Predation of Northern Pike (Esox lucius L.) Eggs: A Possible Cause of Regionally Poor Recruitment in the Baltic Sea

Northern pike (Esox lucius) spawning habitat and egg mortality were studied in three spawning areas in 2001 along the Swedish coast of the Baltic proper: the Blekinge Archipelago, Kalmar Sound and coastal freshwater streams. Spawning peaked during the last week of April in streams, at temperatures ranging from 7.7 to 8.9 °C and during the first week of May in brackish waters, at temperatures ranging from 8.9 to 13.8 °C. Spawning occurred in shallow waters, at depths between 0.2–1.5 m, but generally most of the spawn was found in the shallowest areas. In streams, eggs were mainly attached to emersed vegetation, while in brackish sites pike eggs were well scattered among flooded emersed plants, submersed plants and filamentous algae. Mean egg density varied between 469–1829 m^–2 with the lowest density observed in Kalmar Sound. The calculated egg loss occurring from approximately one day after spawning to one day before hatching ranged from 41±7% in coastal streams to 67±6% in the Blekinge Archipelago and 100% in Kalmar Sound. The significant removal of eggs from spawning sites in Kalmar Sound and Blekinge was most likely due to predation from several fish species. In situ observations and stomach analyses suggested that many pike eggs in Kalmar Sound were lost to the three-spined stickleback (Gasterosteus aculeatus), a species that presently dominate the littoral small fish community. This study therefore suggests that egg predation by sticklebacks and other fishes may be a possible cause of the reported poor recruitment of coastal pike populations in the Kalmar Sound region.     

Multispecies spawning sites for fishes on a low‐latitude coral reef: spatial and temporal patterns

Spawning sites used by one or more species were located by intensively searching nearshore coral reefs of Kimbe Bay (New Britain, Papua New Guinea). Once identified, the spawning sites were surveyed repeatedly within fixed 5 m radius circular areas, for > 2000 h of observations ranging from before dawn to after dusk spanning 190 days between July 2001 and May 2004. A total of 38 spawning sites were identified on the seven study reefs distributed at an average of one site every 60 m of reef edge. Pelagic spawning was observed in 41 fish species from six families. On three intensively studied reefs, all 17 spawning sites identified were used by at least three species, with a maximum of 30 different species observed spawning at a single site. Spawning was observed during every month of the study, on all days of the lunar month, at all states of the tide and at most hours of the day studied. Nevertheless, the majority of species were observed spawning on proportionately more days from December to April, on more days around the new moon and in association with higher tides. The strongest temporal association, however, was with species‐specific diel spawning times spanning < 3 h for most species. While dawn spawning, afternoon spawning and dusk spawning species were differentiated, the time of spawning for the striated surgeonfish Ctenochaetus striatus also differed significantly among sites. The large number of species spawning at the same restricted locations during predictable times suggests that these sites are extremely important on this low‐latitude coral reef.     

Aggregation and spawning of the humphead wrasse Cheilinus undulatus (Pisces: Labridae): general aspects of spawning behaviour

The humphead wrasse Cheilinus undulatus formed resident spawning aggregations daily after high tide at specific locations along the seaward edge of the Palau barrier reef. The location and extent of one aggregation site remained consistent for 6 years with no physical features distinguishing it from adjacent areas. Spawning was documented most months and probably occurred year round with possible seasonal and lunar variation. Spawning males arrived first at the site, followed by females and potentially small primary males. The aggregation female to male sex ratio was estimated to be between 6:1 and 10:1. A maximum of 15 males and 100–150 females were observed at the site. A male courtship posture with the anal fin pointed, the caudal fin folded down and the dorsal fin folded against the body was maintained while swimming a few metres off the bottom in view of females. When ready to spawn females rose up as the posturing male passed and the pair released gametes in a relatively sedate fashion near the surface along the shelf break. No attempted predation on spawning adults was seen. Egg predation after spawning was uncommon. On days with early to mid‐day high tides the spawning period started 2·0–2·5 h after high tide when the speed of lagoon–ocean tidal currents peaked and lasted c. 1 h. On days with later afternoon high tides, spawning occurred sooner after high tide and before current speeds peaked. Other fishes with planktonic eggs spawned at the site as pairs or small groups in a rough succession after high tide with C. undulatus, the last species to spawn.     

Spawning migrations in the monogamous butterflyfish,Chaetodon trifasciatus

Spawning and related behavior of a monogamous butterflyfish,Chaetodon trifasciatus, were observed at Kuroshima Island, Okinawa, Japan. Each heterosexual pair defended a feeding territory in the daytime. Spawning occurred at dusk on the days around full or new moon in the daytime. Spawning occurred at dusk on the days around full or new moon in the vicinity of offshore tidal currents. Spawning migration to such sites occurred in pairs, the feeding territories of which were located in areas of inshore currents. In the evening each pair established a small temporary territory, which they spawned adjacent to and slept within until the next morning. The distribution of sleeping sites as well as tidal current directions may determine the spawning sites of this butterflyfish.     

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.     

Evolution of Female Egg Care in Haremic Triggerfish,Rhinecanthus aculeatus

Paternal care is predominant among telcost fishes with external fertilization. This study describes maternal care in a haremic coral-reef fish and discusses the possible factors leading to its evolution. Both sexes of the triggerfish Rhinecanthus aculeatus (Balistidae) maintained territories; some individuals for more than 8 years. Each male's territory overlapped 2-3 female territories. Pair-spawning occurred around sunrise. Only females cared for the demersal eggs until hatching, which occurred just after sunset on the day of the spawning. No predation was observed on eggs under the maternal care, but experimental removal of parental females decreased the hatching rate to nearly zero. Egg-guarding females foraged as frequently as males, but less than half of non-spawning days. Spawning occurred only in the periods of about 1 wk around the new or full moon, and individual females spawned up to three times in each period. Thus, the maternal care did not significantly affect the duration of the females' spawning intervals, while males would suffer mate loss if they performed parental care. In this situation, maternal care should be the evolutionarily stable strategy. Evolutionary transition from no care to maternal care and then to biparental care is suggested in the Balistidae.     

Spawning timing of the cleaner wrasse, Labroides dimidiatus, on a warm temperate rocky shore

We observed spawning behaviors of the haremic cleaner wrasse Labroides dimidiatus at a rocky reef in southern Japan. Females released pelagic eggs daily from June to September after a spawning ascent by pairs. When the high tide occurred between noon and evening (quarter moon to new or full moon), they spawned around the time of the tide providing fast offshore currents. For the rest of the lunar cycle, spawnings occurred during late afternoon independent of the state of the tide at the time of day. This spawning pattern is quite different from that of conspecifics on coral reefs that always spawn around the high tide at all lunar stages to avoid potential egg predators. The wrasse ascended high in the water column, probably because of its predator-immune characters as a cleaner. The steep slope of the study site seemed to contribute to releasing gametes far above aggregating planktivores. Thus, gametes were not subject to heavy predation. Current direction might not always affect survival of spawned pelagic eggs on the temperate reefs so crucially as it does on coral reefs. We conclude that spawning during unfavorable tidal conditions during the daytime may be better than spawning during the best tidal condition in the crepuscular period when predation pressure on adults will be high. Intraspecific variation in the timing and location of spawning of the wrasse may result from different conditions for larval survival. Received: February 6, 2000 / Revised: June 16, 2000 / Accepted: July 17, 2000     

Interspecific variation in spawning time and male mating tactics of the parrotfishes on a fringing coral reef at Iriomote Island,Okinawa

Spawning time and male mating tactics of parrotfishes (family Scaridae) were investigated on a fringing coral reef at Iriomote Island, Okinawa. Spawning was observed in 14 species, and more frequently in more abundant species such as Chlorurus sordidus, Scarus rivulatus and Chlorurus bowersi. At the reef-edge spawning site, C. bowersi spawned at high tide, C. sordidus spawned both at high tide and in the early morning, whereas Calotomus carolinus and most of the Scarus species such as S. rivulatus spawned only in the early morning, mostly 0630–0830 h. Spawning only in the early morning irrespective of tide phase and moon age has seldom been reported from the scarid species of other localities. It is suggested that spawning in the early morning would be adaptive in species such as S. rivulatus, which migrated considerable distances (ca. 500 m) to the inshore feeding sites, in order to minimize feeding losses due to migration. For male mating tactics, pair spawning by territorial TP (terminal phase) males occurred in all 14 species, and streaking and group spawning by nonterritorial small IP (initial phase) males were seen more frequently in more abundant species. Moreover, group spawning by nonterritorial TP males, which were larger than the IP males but smaller than the territorial TP males, frequently occurred in S. rivulatus. Such mating tactics of TP males have not been reported from Scaridae.     

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.     

Predation on fish eggs by white perch,Morone americana,in western Lake Erie

Synopsis The white perch,Morone americana, is an east coast estuarine species that invaded Lake Erie in the 1950's, but did not increase in abundance until the mid 1970's. We studied its distribution and feeding during spawning in the Sandusky River, Ohio in 1981–1983. White perch were present in the area from early April through May, but abundance was highest on bedrock riffles about 45 km upstream from Lake Erie. Spawning activity peaked in the last week of April when temperatures approached 18°C. White perch collected in early April had eaten walleye,Stizostedion vitreum vitreum, eggs. As spawning activity of white perch increased, feeding activity declined, and most fish collected during late April contained no food. Egg predation increased again in May, but the eggs eaten then were those of white bass,Morone chrysops, white perch, and possibly other species. We have no evidence that egg predation by white perch has affected walleye or white bass recruitment, but it could become a problem if white perch continue to increase in abundance.     

Female spawning migrations of the protogynous wrasse,Halichoeres marginatus

Spawning sites and spawning migration paths of tagged females of the protogynous wrasse,Halichoeres marginatus, were studied on the shallow reefs at Kuchierabu-jima Island, Japan. Males set up mating territories above prominent rocks on the offshore reef slope in the late afternoon, and pair-spawned with females, which had migrated there from their home ranges located in inshore areas. Small females migrated to the spawning sites near their home ranges, whereas large females migrated to various spawning sites located within a wide area, including downcurrent sites. Spawning at the downcurrent sites favors transport of eggs offshore, thereby increasing the female’s fitness. The spawning sites where an individual had spawned as a female were subsequently used for mating after it had changed sex. It is suggested that the wide migration of females to various spawning sites, enables the storing up of information on those sites, which later helps in the acquisition of mating territories after changing sex.     

Features of the reproductive ecology of the rock catfish Silurus lithophilus

Some aspects of the reproductive ecology of the rock catfish Silurus lithophilus (Siluriformes: Siluridae), a species endemic to the Lake Biwa system, were investigated from April to July in 1989–1994. Field observations were conducted at a rocky shoal, which was also a spawning ground of S. biwaensis, along the shore of the Seta River, the lake's outlet. Spawning of S. lithophilus occurred around midnight at shallow, rocky places on the shoal (5–70 cm in water depth) from early May to mid-July, starting earlier than S. biwaensis (mid-May to mid-July). Spawning tended to occur at low water temperatures irrespective of high-water events, unlike S. biwaensis, and in low densities (fewer than 8 fish per night) compared to S. biwaensis (1–45 fish per night). Although the rock catfish tended to appear and spawn at higher frequencies on nights when S. biwaensis spawned or appeared, it always avoided conflict with the latter species by spawning at sites far from those used by S. biwaensis or after S. biwaensis had left the vicinity. The survival of eggs of S. lithophilus might be favored by a strategy of diverting predator attention from them by synchronizing spawning nights with S. biwaensis. Received: February 27, 2001 / Revised: October 17, 2001 / Accepted: November 8, 2001     

A survey of the spawning of perch (Perca fluviatilis), pike (Esox lucius), and roach (Rutilus rutilus), using artificial spawning substrates in lakes

The development of spawning in perch, pike and roach in Lake Geneva has been studied by means of artificial spawning substrates, laid at different depths, from 1984 to 1993. In Lake Geneva, perch spawned in May. A rise of surface water temperature up to 14 °C stimulated spawning activity while bad weather (surface temperature at 10 °C) induced a spread of the spawning period over more than one month. The spawning period was delayed in years when the mean width of perch egg-ribbons was the largest; this corresponded to the biggest females. At the beginning of the spawning period (early in May), perch preferentially chose a depth of 4 m to spawn. In contrast, at the end of the spawning period, maximum spawning intensity was observed at a depth of 12 m. This phenomenon was more pronounced when water temperature rose above 14 °C in the top 4 m while it remained below 12 °C at a depth of 12 m. Pike spawned at the end of April and at the beginning of May in Lake Geneva. They preferred spruce branches among the different spawning substrates that we tested in Lake Vouglans. When the water temperature increased at the surface of Lake Geneva, pike preferred to lay their eggs on substrates set at 3 m depth where the temperature was cooler than in the surface layer (10.5 °C vs 14 °C). Roach spawned during the last two weeks of May or during the first two weeks of June in Lake Geneva, depending on water temperature. Spawning had generally been spread over a week, but a sudden decrease of water temperature could slow spawning intensity. Roach were able to lay their eggs on natural or synthetic substrates, located 0.5 m below the surface near the shore as well as, at a distance of several hundred metres from the shore. The survival of eggs was always above 90% for perch and roach and generally above 70% for pike.     

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 of the sea ravenHemitripterus villosus in Peter the Great Bay, Sea of Japan

The spawning habits of the sea ravenHemitripterus villosus were investigated by scuba diving at shallow rocky bottom sites around Bolshoi Pelis Island (Peter the Great Bay, Sea of Japan). Spawning occurs in September, when the surface water temperature drops. The optimum spawning temperature is 17–18°C. The fish density in nest sites reaches 120/300 m². The mean individual fecundity is about 10000 eggs. Over 35% of the egg masses ofH. villosus are eaten by echinoderms, primarilyPatiria pectinifera andStrongylocentrotus nudus. Since fecundity is relatively low and parents do not take care of the egg masses, predation can strongly affect the abundance of this species.     

Mass spawning of Caesio teres (Pisces: Caesionidae) at Enewetak Atoll,Marshall Islands

Synopsis Mass spawning of Caesio teres occurred between March and August, 1983 on a reef emerging from deep water just inside the East Channel of Enewetak Atoll, Marshall Islands. Aggregations and spawning were observed from one day before until three days after the full moon. Spawning aggregations were not observed during the new or quarter moons. Spawning commenced after high tide, when the current began to flow out the pass from lagoon to ocean. The spawning aggregation of close to 1000 individuals migrated to the spawning site. Spawning occurred when the aggregate ascended to near the water's surface. Subgroups dashed horizontally within the aggregate, releasing a highly visible gamete cloud. Predation on spawning adults was not observed. Predation on spawned eggs was noted. The eggs of C. teres are described. Mid-Pacific Research Laboratory, Enewetak Atoll, Republic of the Marshall Islands Present address: Motupore Island Research Station, P.O. Box 320, University of Papua New Guinea, Papua New Guinea     

Location and timing of Asian carp spawning in the Lower Missouri River

We sampled for eggs of Asian carps, (bighead carp Hypophthalmichthys nobilis, silver carp H. molitrix, and grass carp Ctenopharyngodon idella) in 12 sites on the Lower Missouri River and in six tributaries from the months of May through July 2005 and May through June 2006 to examine the spatial and temporal dynamics of spawning activity. We categorized eggs into thirty developmental stages, but usually they could not be identified to species. We estimated spawning times and locations based on developmental stage, temperature dependent rate of development and water velocity. Spawning rate was higher in the daytime between 05:00 and 21:00 h than at night. Spawning was not limited to a few sites, as has been reported for the Yangtze River, where these fishes are native, but more eggs were spawned in areas of high sinuosity. We employ a sediment transport model to estimate vertical egg concentration profiles and total egg fluxes during spawning periods on the Missouri River. We did not identify substantial spawning activity within tributaries or at tributary confluences examined in this study.     

Reproductive biology of the endangered Oxleyan pygmy perch Nannoperca oxleyana Whitley

The reproductive biology of the Oxleyan pygmy perch Nannoperca oxleyana is described from simultaneous studies of wild populations in north-eastern New South Wales and mature fish held in aquaria. In the wild, 50% of males and females matured at total lengths of 24·0–25·9 and 28·0–29·9 mm, respectively. The species displays sexual dichromatism during the spawning season, with males developing more intense red and brown fin and body colouration, and black pelvic fins. Captive male broodfish displayed territoriality during the breeding season, closely guarding sites within artificial, plant-like substrata in which pairs of fish spawned adhesive eggs. Protracted serial spawning of wild and captive fish occurred from September to April and May at mean water temperatures ≥16·6° C and day length ≥10·7 h. Captive broodfish spawned on an average of 57% of days during the 256 day spawning period. Gonado-somatic indices averaged 0·7% for all ripe males and 4·1–4·2% for all ripe females collected. Mean total and batch fecundities of captive females were 1323 eggs per fish and 7·8 eggs per fish per day, respectively, and relative fecundity was 587 eggs g⁻¹ of body mass. Batch fecundity of wild females was estimated at 7·8 eggs per fish. The adaptive significance of this reproductive strategy in a harsh, variable environment is discussed.     

Growth and reproduction in Canthigaster valentini (Pisces,Tetraodontidae): a comparison of a toxic reef fish with other reef fishes

Synopsis The growth and reproduction of Cantigaster valentini were studied in two sites at Lizard Island, Australia. C. valentini was found to be a gonochore, with a sex ratio very close to 1:1; sexes could be distinguished externally. The growth (in length) of known individuals from both sites was measured at least every two months over two years. Growth rates of males and females decrease as their sizes increase. Growth rates differ between sexes and between sites: males generally grow faster than females and individuals at Mermaid Cove generally grow faster than individuals at Palfrey Island. Spawning is demersal, it occurs daily between 0800 and 1600h, and continues year-round. For females the interval between successive spawnings varies from about 4 days in the warm-water season to about 10 days in the cool-water season. From a comparison of local reproductive output and local recruitment survivorship of larvae in the plankton was estimated to be much higher than in another species (Pomacentrus wardi) for which a similar estimate was available. We suggest that some aspects of the reproductive strategy of C. valentini differ from other, non-toxic reef fishes in ways consistent with a reduced threat of predation upon adults, eggs, and larvae: courtship and spawning are unhurried and occur throughout most of the day; spawning is unrelated to lunar cycles; there is no parental care or defense of fertilized eggs; and embryos often hatch on rising tides.Centre for Environmental and Urban StudiesSenior author's present address: School of Biological Sciences F07, University of Sydney, Sydney 2006, N.S.W., Australia     

Sexual reproduction and early development of the solitary coral Fungia scutaria (Anthozoa: Scleractinia)

Fungia scutaria spawned vigorously with a lunar beriodicity during the summer months of 1981 and 1982. Spawning activity declined in the fall of both years and was absent in winter and spring (1983). There was only one short spawning event per lunar cycle. Each event occurred in the evening between 1700 and 1900 hours 1 to 4 days following the full moon. Fungia scutaria exhibits gonochorism. Females ejected eggs through their mouths into the seawater above. Many of these negatively buoyant eggs settled onto the oral discs and were moved off the edge by ciliary-mucoid activity. Spermatoza from males were similarly expelled in a jet of gastrovascular fluid. Spawned eggs were small and lacked endosymbiotic zooxanthellae. Rapid development led to ciliated solid planulae by the next morning. Within 24 h a mouth had begun to develop. Planulae may have been able to feed within 39 h. Infection with zooxanthellae occurred 4–5 days following spawning. Planulae may have become competent for settlement by 7 days, but attempts to docment settlement produced ambiguous results.