Deepwater broadcast spawning by Montastraea cavernosa, Montastraea franksi, and Diploria strigosa at the Flower Garden Banks, Gulf of Mexico

Broadcast spawning by corals is a tightly synchronized process characterized by co-ordinated gamete release within 30–60 min time windows once per year. In shallow water corals, annual water temperature cycles set the month, lunar periodicity the day, and sunset time the hour of spawning. This tight temporal regulation is critical for achieving high fertilization rates in a pelagic environment. Given the differences in light and temperature that occur with depth and the importance of these parameters in regulating spawn timing, it has been unclear whether deeper coral can respond to the same environmental cues that regulate spawning behaviour in shallower coral. In this report, a remotely operated vehicle was used to monitor coral spawning activity at the Flower Garden Banks at depths from 33 to 45 m. Three species Montastraea cavernosa, Montastraea franksi, and Diploria strigosa were documented spawning within this depth range. All recorded spawning events were within the same temporal windows as shallower conspecifics. These data indicate that deep corals at this location either sense the same environmental parameters, despite local attenuation, or communicate with shallower colonies that can sense such spawning cues.     

Combined effects of tidal and diurnal cycles on spawning of the puffer,Takifugu niphobles (Tetraodontidae)

Synopsis Spawning in the pufferTakifugu niphobles was observed at Tomioka in western Kyushu, Japan, and the factors affecting the spawning time were estimated using a multiple regression analysis. Spawning in this puffer occurred in the intertidal beach during evening rising tides around the full and new moons. Both the time of commencement and ending of a day's spawning were related mainly to the tidal cycle, and occurred later when the high tides occurred later. However, the termination of a day's spawning was also affected by the diurnal cycle; spawning ended earlier when sunset was earlier. Besides, a day's spawning tended to be concentrated within a shorter time span when the interval between the commencement of spawning and the sunset was shorter. Thus the spawning time of the puffer is regulated not only by the tidal but also by the diel cycle, and it is suggested that the adaptive significance of the spawning reactions to both cycles should be examined separately.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

The Role of Temperature and Embryo Development Time in the Diel Timing of Spawning in a Coral-reef Damselfish with High-frequency Spawning Synchrony

We studied the diel timing of spawning in the demersally spawning Hawaiian damselfish, Dascyllus albisella, from mid-June to late-September 1997 at two small patch reefs in Hawaii. Our objectives were to elucidate daily timing of spawning in relation to water temperature, diel timing of hatching, and short-period spawning synchrony. Spawning occurred every 5–7 days at both reefs, with all spawning on a reef concluded either within a single day (1-day spawning) or within two successive days (2-day spawning). Spawning began in early morning and continued for most of the day. There was a significant, positive linear relationship between mean daily average water temperature (= daily average temperature averaged over the period starting from the day following the last spawning day of the preceding nest cycle till the day before the first spawning day of the current cycle) and peak spawning hour of day, for 1-day spawning, and the first and second days of 2-day spawning at both reefs. The relationship between mean daily average water temperature and peak spawning hour of day was comparable among all spawning-day classes and reefs. Hatching occurred on the fourth day of development throughout the study despite the 26.5–29.1°C change in water temperature during the study period, and hatching was restricted to within two hours after sunset. We propose that D. albisella's peak spawning time is positively correlated with increased water temperature because it maintains the benefits of synchronous spawning within two constraints: the narrow daily period of hatching, and the inverse relationship between water temperature and embryo developmental time.     

Diel and lunar spawning periodicity of the hawkfish Paracirrhites forsteri (Cirrhitidae) on the reefs of Kuchierabu-jima Island, southern Japan

We observed spawning behavior of the hawkfish Paracirrhites forsteri on reefs of southern Japan. Spawning generally occurred after sunset, prior to the full and new moon with semi-lunar spawning peak periodicity. No egg predation was observed in spawning after sunset, and high tide often occurred at dusk in the lunar phases. Thus, diel timing and lunar synchronicity may increase larval survival. We found that the mating activity tended to start earlier in the day during early mating season than during mid-to-late mating season. The advantages of the earlier start of mating activity were also examined in relation to adult biology contexts.     

Genetic, spatial, and temporal components of precise spawning synchrony in reef building corals of the Montastraea annularis species complex

When organisms release gametes into the sea, synchrony must be precise to increase fertilization and decrease hybridization. We tagged and genotyped over 400 spawning corals from the three species in the Montastraea annularis species complex. We report on the influence of species, individuals, and genotypes on timing of spawning from 2002 through 2009. During their annual spawning event M. franksi spawns on average 2 h after sunset, whereas M. annularis and M. faveolata spawn 3.5 h after sunset. Only M. franksi and M. annularis have compatible gametes. Individual colonies of the same genotype spawn at approximately the same time after sunset within and across years (within minutes), but different genotypes have significantly different spawning times. Neighboring colonies, regardless of genotype, spawn more synchronously than individuals spaced further apart. At a given distance, clone-mates spawn more synchronously than nonclone-mates. A transplant experiment indicates a genetic and environmental influence on spawn time. There is strong, but not absolute, concordance between spawn time, morphology, and genetics. Tight precision in spawning is achieved via a combination of external cues, genetic precision, and perhaps conspecific signaling. These mechanisms are likely to influence reproductive success and reproductive isolation in a density-dependent manner.     

Semilunar spawning cycle of the humbug damselfish Dascyllus aruanus

A very distinct semilunar spawning cycle was found in a population of the damselfish Dascyllus aruanus on the coral reefs of Sesoko Island, Okinawa. Spawning occurred from June to September, only in the early morning, during a period of 2–4 days immediately before or around the time of the new and full moon. Males cared for the eggs deposited on the substrate for 2.5 days until hatching. Hatching occurred just after sunset, i.e., at the high tide of spring tide; the strong ebb current then would rapidly disperse the newly hatched larvae offshore. Females tended to synchronize spawning in a male's nest, also because multiple clutches in a nest would be more likely to survive until hatching. Thus, the distinct semilunar spawning cycle may favor females in reducing mortality of both eggs and larvae. Received: July 9, 1999 / Accepted: January 29, 2000     

A tropical bird can use the equatorial change in sunrise and sunset times to synchronize its circannual clock

At higher latitudes, most organisms use the periodic changes in day length to time their annual life cycle. At the equator, changes in day length are minimal, and it is unknown which cues organisms use to synchronize their underlying circannual rhythms to environmental conditions. Here, we demonstrate that the African stonechat (Saxicola torquatus axillaris)-an equatorial songbird-can use subtle solar cues for the annual timing of postnuptial moult, a reliable marker of the circannual cycle. We compared four groups that were kept over more than 3 years: (i) a control group maintained under constant equatorial day length, (ii) a 12-month solar time group maintained under equatorial day length, but including a simulation of the annual periodic change in sunrise and sunset times (solar time), (iii) a 14-month solar time group similar to the previous group but with an extended solar time cycle and (iv) a group maintained under a European temperate photoperiod. Within all 3 years, 12-month solar time birds were significantly more synchronized than controls and 14-month solar time birds. Furthermore, the moult of 12-month solar time birds occurred during the same time of the year as that of free-living Kenyan conspecifics. Thus, our data indicate that stonechats may use the subtle periodic pattern of sunrise and sunset at the equator to synchronize their circannual clock.     

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     

Effects of climate and demography on reproductive phenology of a harvested marine fish population

Shifts in phenology are a well‐documented ecological response to changes in climate, which may or may not be adaptive for a species depending on the climate sensitivity of other ecosystem processes. Furthermore, phenology may be affected by factors in addition to climate, which may accentuate or dampen climate‐driven phenological responses. In this study, we investigate how climate and population demographic structure jointly affect spawning phenology of a fish species of major commercial importance: walleye pollock (Gadus chalcogrammus). We use 32 years of data from ichthyoplankton surveys to reconstruct timing of pollock reproduction in the Gulf of Alaska and find that the mean date of spawning has varied by over 3 weeks throughout the last >3 decades. Climate clearly drives variation in spawn timing, with warmer temperatures leading to an earlier and more protracted spawning period, consistent with expectations of advanced spring phenology under warming. However, the effects of temperature were nonlinear, such that additional warming above a threshold value had no additional effect on phenology. Population demographics were equally as important as temperature: An older and more age‐diverse spawning stock tended to spawn earlier and over a longer duration than a younger stock. Our models suggest that demographic shifts associated with sustainable harvest rates could shift the mean spawning date 7 days later and shorten the spawning season by 9 days relative to an unfished population, independent of thermal conditions. Projections under climate change suggest that spawn timing will become more stable for walleye pollock in the future, but it is unknown what the consequences of this stabilization will be for the synchrony of first‐feeding larvae with production of zooplankton prey in spring. With ongoing warming in the world’s oceans, knowledge of the mechanisms underlying reproductive phenology can improve our ability to monitor and manage species under changing climate conditions.     

Natural spawning of a Hawaiian sea urchin,Tripneustes gratilla

Observations of natural spawning events are rare but critically important for the field of fertilization ecology. For broadcast spawners, knowing the natural behavior of spawning, including proximity of animals and the timing of spawning, is essential for informing spawning experiments and directing future study. Here we describe a natural spawning event that took place on the island of Maui in the winter of 2010. Groups of Tripnuestes gratilla were observed to spawn in the late afternoon shortly after a local high tide that was a lower high of a mixed, semidiurnal tide cycle. Urchins appeared to increase the probability of fertilization by ascending to the local high points of the reefs and spawning within groups of 2–5. Although the majority of the urchins that spawned were T. gratilla, two individuals of Echinothrix diadema were observed to spawn at the same time, raising questions about potential hybridization in the wild.     

Diel spawning periodicity of red snapper Lutjanus campechanus in the northern Gulf of Mexico

Ovaries of red snapper Lutjanus campechanus were examined histologically to determine rates of oocyte maturation, diel spawning periodicity and whether lunar cycle influenced spawning rhythm. Hydration of red snapper oocytes began during the mid‐morning hours; c . 5 h was necessary for oocytes to become fully hydrated and ovulation occurred no more than 5 h after oocytes attained full hydration. Appearance of fresh postovulatory follicles after 1330 hours and the absence of hydrated oocytes after 1830 hours signified that red snapper spawning occurred during this 5 h period. In addition, evidence of a peak in spawning was seen near 1600 hours. Postovulatory follicles degenerated within a 24 h time period. A lunar spawning cycle was not evident.     

The timing of spawning in capelin (Mallotus villosus Müller) at a coastal location in eastern Newfoundland

We tested three hypotheses concerning the timing of spawning for a circumpolar species, capelin (Mallotus villosus), for which timing of larval emergence is known to be synchronized by physical conditions. The first hypothesis, developed from previous studies, was that spawning would be synchronized by upwelling events. Initial results from Middle Cove Beach in eastern Newfoundland indicated that spawning was not synchronized with upwelling. We next hypothesized that spawning was a function of several environmental variables. Results from logistic regression indicated that neither single-factor nor multi-factor models could explain the timing of spawning. Single variables could predict spawning in some years but no variable could reliably predict the time of spawning year after year. Finally, we hypothesized that the probability of spawning increased as a set of significant variables approached preferred levels. For capelin at Middle Cove, the set of variables that influence capelin spawning were identified as wave height, sea surface roughness and capelin abundance in the water. Thus only a combination of variables explained the timing of spawning for capelin. Preferred conditions for capelin spawning were wave heights less than 20 cm at the beach, a sea surface with a slight ripple, and an intermediate rank abundance of capelin in the water corresponding to hundreds to thousands of individuals. Capelin abundance alone was not a useful predictor. During the course of the study we observed a shift in the dates that capelin arrived and spawned at the beach. During 1987–1990 capelin spawned at Middle Cove Beach during June, but in more recent years (1991–1993) capelin did not arrive or spawn until July.     

The utility of a long‐term acoustic recording system for detecting white seabass Atractoscion nobilis spawning sounds

This study reports the use of a long‐term acoustic recording system (LARS) to remotely monitor white seabass Atractoscion nobilis spawning sounds at three sites along the southern California coastline, adjacent to Camp Pendleton. On the basis of previous studies of A. nobilis sound production relative to periods of known spawning activity, LARS were set to continuously record ambient sounds for a 2 h period around sunset from April to June 2009. Acoustic analyses identified A. nobilis courtship sounds on 89, 28 and 45% of the days at the three locations, respectively. From 474 h of acoustic data, spawning‐related sounds (chants) were detected on 19 occasions in 2009 with an additional 11 spawning chants recorded during a 2007 validation period. Most spawning chants occurred within 30 min of sunset during the months of May and June at a mean ±s.d . surface temperature of 18·2 ± 1·2° C. Consecutive daily spawning activity was not apparent at any sites in 2009. Atractoscion nobilis spawning chants were recorded at all three sites, suggesting that shallow rocky reefs which support kelp forests provide suitable A. nobilis spawning habitat. Results confirm the utility of passive acoustic recorders for identifying A. nobilis spawning periods and locations.     

Spawning aggregations of Lutjanus cyanopterus(Cuvier) on the Belize Barrier Reef over a 6 year period

Cubera snapper Lutjanus cyanopterus aggregated to spawn at Gladden Spit, a salient sub‐surface reef promontory seaward of the emergent reef and near the continental shelf edge of Belize. Their spawning aggregations typically formed 2 days before to 12 days after full moon from March to September 1998–2003 within a 45 000 m² reef area. Peak abundance of 4000 to 10 000 individuals was observed between April and July each year, while actual spawning was most frequently observed in May. Spawning was observed consistently from 40 min before to 10 min after sunset within a confined area ≤1000 m². Data suggested that cubera snapper consistently formed seasonal spawning aggregations in relation to location, photoperiod, water temperature and lunar cycle, and that spawning was cued by time of day but not tides. The cubera snapper aggregation site was included within the Gladden Spit Marine Reserve, a conditional no‐take fishing zone.     

Reproduction of the introduced oyster Crassostrea gigas (Bivalvia: Ostreidae) cultured on rafts in Spain

The oyster Crassostrea gigas was introduced in Spain for aquaculture purposes; however, until now, it is not known whether populations are established in the wild, being necessary to define whether this species is spawning and which environmental variables trigger this process. The influence of environmental parameters on the reproduction of C. gigas was evaluated from January 2008 to October 2009 with oysters grown on a raft in the Ría de Arousa (Galicia, NW Spain). Temperature and chlorophyll a are directly correlated to sexual maturation. Oysters can mature at temperatures below 14°C. The temperature necessary for spawning differs between the sexes, requiring a temperature above 15°C for males and 18°C for females. Females had a single massive spawn between June and September, while males had partial spawning from May to December with two peaks, one in May–September and another in October–December, with the second peak more pronounced. The first spawning peak is related to high temperatures and concentrations of chlorophyll a, and the second spawning peak is mainly related to the food availability in the water. The spawning asynchrony may be impeding establishment of wild C. gigas populations in Spain.     

Variability of key biological parameters of round sardinella Sardinella aurita and the effects of environmental changes

We examined growth rates and reproductive characteristics of Sardinella aurita off Senegal and other coastal areas over a 20 year period (1995–2014) to determine how they relate to variations in environmental characteristics of coastal waters. Based on fish length-frequency data and a coastal upwelling index, we found that S. aurita recruitment tends to occur during the periods of most intensive upwelling (March–April off Senegal). Peak reproduction corresponds to periods of low sea-surface temperature (in February or March). The sex ratio was remarkably consistent during the 30 year study period and so was not affected by environmental changes. We hypothesise that S. aurita takes advantage of the higher zooplankton productivity that occurs in coastal waters when upwelling brings nutrient-rich water to the surface (i.e., it increases its growth rate and accumulates energy reserves for spawning). Growth performance appears to be strongly dependent on environmental conditions. The timing of spawning seems to occur when food (zooplankton) is most available for supplying the energy requirements needed by adults for spawning and early development of larvae. Environmental changes seem to have a significant effect on S. aurita growth and reproduction, which endorses their high phenotypic plasticity.     

Phenological and distributional shifts in ichthyoplankton associated with recent warming in the northeast Pacific Ocean

Understanding changes in the migratory and reproductive phenology of fish stocks in relation to climate change is critical for accurate ecosystem‐based fisheries management. Relocation and changes in timing of reproduction can have dramatic effects upon the success of fish populations and throughout the food web. During anomalously warm conditions (1–4°C above normal) in the northeast Pacific Ocean during 2015–2016, we documented shifts in timing and spawning location of several pelagic fish stocks based on larval fish samples. Total larval concentrations in the northern California Current (NCC) during winter (January–March) 2015 and 2016 were the highest observed since annual collections first occurred in 1998, primarily due to increased abundances of Engraulis mordax (northern anchovy) and Sardinops sagax (Pacific sardine) larvae, which are normally summer spawning species in this region. Sardinops sagax and Merluccius productus (Pacific hake) exhibited an unprecedented early and northward spawning expansion during 2015–16. In addition, spawning duration was greatly increased for E. mordax, as the presence of larvae was observed throughout the majority of 2015–16, indicating prolonged and nearly continuous spawning of adults throughout the warm period. Larvae from all three of these species have never before been collected in the NCC as early in the year. In addition, other southern species were collected in the NCC during this period. This suggests that the spawning phenology and distribution of several ecologically and commercially important fish species dramatically and rapidly changed in response to the warming conditions occurring in 2014–2016, and could be an indication of future conditions under projected climate change. Changes in spawning timing and poleward migration of fish populations due to warmer ocean conditions or global climate change will negatively impact areas that were historically dependent on these fish, and change the food web structure of the areas that the fish move into with unforeseen consequences.     

Aspects of the spawning of western Atlantic butterflyfishes (Pisces: Chaetodontidae)

Synopsis The status of knowledge of spawning among the five shallow waterChaetodon species in the western Atlantic is reviewed. Spawning has been observed for three species in Puerto Rico, St. Croix and the Bahamas, with possible courtship in a fourth.Chaetodon aculeatus spawned near the time of sunset over objects on the reef as single female/male pairs or as two females and one male, with pair spawning in rapid succession. Spawning occurred during much of the lunar month from February to April and it is uncertain whether any lunar periodicity to spawning exists. Male-male aggression was noted. Spawning sites (coral heads) were alternated daily and it is likely that females spawn only once every two days. A single female produced as many as 2090 eggs in a single spawning.Chaetodon capistratus spawned during much of the lunar month from February to April. It spawned about 5 min afterC. aculeatus, occasionally using the same sites, and alternated sites daily. A female produced as many as 3710 eggs in one spawning.Chaetodon striatus spawned from February to April but it is unknown if it has any lunar spawning cycle. No predation attempts by piscivores on spawning adults were seen. Predation byMelichthys niger on eggs ofC. striatus occurred. No egg predation was observed forC. aculeatus andC. capistratus. With an assumed four month reproductive season, alternate day spawning and observed egg production values,C. aculeatus andC. capistratus produce respectively about 100 000 and 200 000 eggs per large female per year. The reproductive strategy of smaller species may be to produce moderate numbers of eggs per day over a spawning season of at least a few months while larger species may produce more eggs per day for a shorter period.