Seasonality,lunar periodicity of settlement and microhabitat association of juvenile humpback red snapper <Emphasis Type="Italic">Lutjanus gibbus</Emphasis> (Lutjanidae) in an Okinawan coral reef

Snappers (Lutjanidae) are one of the important fisheries resources in tropical and subtropical waters. However, there have been few studies clarifying the ecology of newly settled juvenile snappers. The aims of the present study were to clarify the seasonality and lunar periodicity of the larval settlement, and the microhabitat association of newly settled juveniles for Lutjanus gibbus in an Okinawan coral reef. Fifteen coral patches were chosen and underwater visual surveys were conducted during a 2-year period. The larval settlement of the species mainly occurred between May and October during the 2-year survey period. In terms of lunar periodicity of the settlement, the newly settled juveniles were mainly found between the last quarter moon and new moon during the main settlement season. Back calculations of settlement dates based on otolith microstructure showed that most juveniles settled during the last quarter moon and new moon. Stepwise multiple regression analyses revealed that the newly settled juveniles were mainly found on coral patches with a large number of holes on the top of the coral patch and this tendency was mainly consistent during the study period. The volume of holes at the base of coral patches had also a positive effect on the abundance of newly settled juveniles in some cases. The results of the present study suggest that larval settlement has a strong seasonal and lunar periodicity, and newly settled juveniles select coral patches with available refuge spaces.     

Growth rhythms in the shrimp Penaeus vannamei and P. schmitti

Growth increments of the penaeid shrimp Penaeus vannamel (Boone) and P. schmitti (Burkenroad) grown in commercial production ponds in Ecuador and Colombia were measured over a 4 mo period between October 1987 and January 1988. P. vannamel grown from wild caught seed in Ecuadorian ponds exhibited a growth rhythm which significantly coincided with lunar phases. Growth increments of at least 1 gwk^-1 occurred around a new and full moon, while growth increments of less than 1 gwk^-1 occurred during first and last quarters. P. vannamel transported from Ecuador as nauplii, raised in Colombian hatcheries and on grown in Colombian ponds exhibited a cyclic pattern in wet weight increments but not correlation to lunar cycles was observed. P. schmitti showed no cyclic pattern in growth. The two species under consideration are indigenous to two different, and quite unique areas displaying different environmental characteristics. The link between the growth pattern, biological and lunar cycles is discussed.     

Lunar rhythms of vertical migrations coded in otolith microstructure of North Atlantic lanternfishes,genus Hygophum (Myctophidae)

Otoliths of five Hygophum species were examined by means of light and scanning electron microscopy. In otoliths of four species (H. benoiti, H. macrochir, H. reinhardtii and H. taaningi) a strong cyclic pattern of the incremental structure was observed. In the fifth species (H. hygomii) such a pattern did not exist. An analysis of archival data on mesopelagic collections suggested three types of Hygophum spp. migratory behavior in relation to the lunar cycle which corresponded with the otolith microstructure. In H. hygomii only limited influence of moon phase on the uppermost range of night-vertical migration toward the surface was observed. The abundant nighttime occurrence of this species moved from the 0 to 50 m into the 50 to 100 m depth strata at full moon. In H. benoiti, a great part of the population, mainly juveniles, showed a tendency toward cessation of the vertical migrations during the first and fourth quarters of the lunar cycle. H. macrochir and H. taaningi had the strongest correlation of behavior with the lunar cycle. Both species showed arrested vertical migrations at the new moon phase, staying at day depths during the night, i. e., below 400 m. Thus, sequences of clear growth increments in otoliths represented a fast-growth period associated with the night migration to the warm surface layers, while bands without easily distinguishable incremental structure were interpreted as a period of slow growth in deep, cold waters due to limitation of the upward migration range occurring approximately at new moon.     

Sexual reproduction in Acropora (Isopora) (Coelenterata: Scleractinia)

In contrast to the seasonal gamete and planula production of Acropora (Isopora) palifera on Heron Island reef (Lat. 23° S), populations on Lizard Island reef (Lat. 14° S), sampled in 1979, 1981 and 1983, and Salamaua and Busama reefs (Lat. 7° S), sampled from 1980 to 1983, planulated year-round. Intensive sampling of colonies at Salamaua and Busama showed that gametes ripened at two-month intervals and that up to six cycles of gametes and larvae could be produced by an individual colony. Gametes of only a portion of the population — usually close to 50% — ripened each month. The Salamaua population, on average, produced fewer and smaller planulae than the Heron Island population during each two-monthly reproductive cycle. Hypotheses correlating the annual periodicity of breeding in marine animals with latitudinal variation of temperature were tested. In general, the time of breeding in A. palifera at Heron Island reef is much more restricted than theories based on latitudinal variations of water temperature would predict. It is hypothesized that, in the near subtropical environment of Heron Island reef, this species has evolved a life-history strategy that limits the amount of energy allocated to reproduction and allocates more energy to growth.     

Cyclical mucous sheet formation on poritid corals in the San Blas Islands,Panama

Mucous sheets are common on poritid corals as well as other reef coelenterates, but the frequency, cause and function of these sheets have rarely been investigated.Porites furcata Lamarck andPorites astreoides Lamarck colonies at three sites in the San Blas Islands, Panama, were monitored for the presence or absence of mucous sheets every 2 d between 11 May and 27 August 1983. Concurrent measurements of polyp expansion, particulate matter concentrations, salinity, water temperature and sedimentation rates were also made at each site. Mucous sheet formation was not correlated with any of the environmental parameters measured. Mucous sheet formation by both species had a lunar periodicity.P. furcata formed mucous sheets around the time of the full moon, whileP. astreoides formed mucous sheets during the first quarter (around a week before full moon). Mucous sheet formation was correlated with polyp contraction behavior and may be a secondary effect of a lunar cycle in polyp activity.     

Patterns of lunar settlement and early recruitment in Caribbean reef fishes at Panamá

Lunar patterns of settlement of 15 Caribbean reef fishes were assessed from daily collections of newly arrived fishes from each of two small (A=63 and B=28 m²) patch reefs at Punta de San Blas (Panamá) during two consecutive 2 yr periods (1984, 1985 and 1986, 1987). In 12 species settlement was lunar-cyclic, with a broad peak of activity around the new moon in 11 cases, and at first quarter in 1 species. However, 3 of these lunar-cyclic species displayed intermittent semilunar periodicity in settlement, with peaks around the quarter moons. One other species had a semilunar cycle of settlement with peaks at both quarter moons. Two species apparently lacked lunar settlement cycles. In some, but not all, paris of congeners: (a) monthly variation in settlement intensity was positively correlated, (b) the duration of settlement pulses in the same month consistently differed, and (c) the timing of settlement pulses in the same month often differed (although one species did not consistently arrive before the other). Recruitment of juvenile fish that survived to the end of the lunar cycle in which they settled was monitored monthly, concurrently with settlement, at a number of large sites scattered up to 2 km apart. The levels of variation in the amount of settlement and recruitment each month differed in only 2 of 17 cases, with settlement variation exceeding recruitment variation in only one of these. The intensity of recruitment was positively correlated with the intensity of settlement in 16 of 17 cases. In 15 of those 16 cases, R ² values for linear regressions of recruitment on settlement exceeded R ² values for curvilinear regressions that would indicate either increasing or decreasing recruitment success with increasing intensity of settlement. Thus it appears that variation in recruitment can be used to estimate variation in settlement, and that recruitment success may be density-independent.     

Reproductive biology and larval development of the temperate soft coral <Emphasis Type="Italic">Dendronephthya gigantea</Emphasis> (Alcyonacea: Nephtheidae)

Dendronephthya gigantea (Verrill, 1864), the dominant species in the waters surrounding Jeju Island, Korea, is a gonochoric internal brooder that releases its planulae from July to September. The ratio of females to males in this azooxanthellate soft coral is 2:1. Oogenesis takes place for 12 months and spermatogenesis for 3–5 months. Gametes mature as seawater temperatures increase, suggesting a seasonal factor in the reproductive cycle. Planulae brooded in the gastrodermal canal were expelled around the times of the full moon and the new moon; a significant difference in numbers expelled was not found between day and night. Ciliated planulae had negative buoyancy after planulation, and showed rapid metamorphosis into a primary polyp stage within 2 days.     

Night irradiance and synchronization of lunar release of planula larvae in the reef coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus), which is known to release planula larvae on a monthly cycle, was grown in full daytime solar irradiance, but with four treatments of night irradiance: (1) natural night irradiance, (2) shifted-phase (total darkness during nights of full moon with artificial irradiance at lunar intensity on nights of new moon), (3) constant full moon (full lunar irradiance every night), and (4) constant new moon (total darkness every night). The reproductive cycle of the corals held in the shifted-phase treatment moved out of synchrony with the cycle of corals exposed to a natural lunar cycle of night irradiance. Two previously described types of P. damicornis were tested. The Type Y normally start releasing larvae at full moon, with peak production at third quarter. In the shifted-phase treatment they began releasing planulae at new moon (artificial full moon), with peak production at first quarter. The Type B corals, that normally start releasing planulae at new moon with peak production at first quarter, began to release planulae at full moon (artificial new moon), with peak production at third quarter. Populations of corals grown either in the constant full moon or constant new moon treatment quickly lost synchronization of monthly larva production, although production of planulae continued. Thus spawning is synchronized by night irradiance.Contribution No. 702 of the Hawaii Institute of Marine Biology     

Reproductive cycles and larval morphology of three Recent species of Argyrotheca (Terebratellacea: Brachiopoda) from Mediterranean submarine caves

Although the three examined species of Argyrotheca are quite common in the Mediterranean, little is known about their reproductive cycles. This study shows that Argyrotheca cordata and Argyrotheca cistellula have continuous breeding activity, indicated by the occurrence of ripe eggs and larvae in their brood pouches throughout the year. The absolute number of larvae found in the brood pouches of both species corresponds to the adult's size. The third species examined, Argyrotheca cuneata, tends to breed in autumn, with larvae only present in September and November, but the sample-size was small. Larval development is very similar in all three Argyrotheca species, each embryo going through a gastrula stage, a two-lobed and a three-lobed stage. Before leaving the brood pouch, the larvae have an apical lobe with a girdle of elongated cilia, a mantle lobe with a midventral band of cilia and a pedicle lobe without ciliation. The larvae of A. cordata and A. cuneata possess four bundles of larval setae, whereas A. cistellula shows no setae at all. Received: 6 November 1998 / Accepted: 15 March 1999     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panama, and Galapagos Islands (Ecuador)

Reproductive ecology of two major reef-building corals in the eastern Pacific [Pocillopora damicornis (Linnaeus) andPocillopora elegans Dana] was investigated between 1984 and 1990 in Costa Rica, Panama (Gulf of Chiriqui and Gulf of Panama), and the Galapagos Islands (Ecuador) following the 1982–83 El Niño disturbance. Mature spermatocytes and oocytes were found in both species and were usually present in the same polyp in nearly equal ratios. Numerous zooxanthellae were observed in mature, unfertilized oocytes. Although spawning in field populations was not observed, it is likely that both pocilloporids are simultaneous hermaphroditic spawners, as evidenced by the disappearance of mature gametes after full moon. This is in contrast with most known pocilloporid corals that brood and release planula larvae. Corals were reproductively most active in the thermally stable environments of Costa Rica and the Gulf of Chiriqui (Panama) where 32 to 90% of all colonies contained gametes. In the moderately varying thermal conditions in the Galapagos Islands, 16 to 40% of colonies contained gametes, and in the pronounced seasonal upwelling environment of the Pearl Islands (Panama) only 6 to 18% of colonies contained gametes. Year-round reproduction occurred in Costa Rica and the Gulf of Chiriqui, whereas reproduction was confined to warm periods in the seasonally varying environments of the Galapagos Islands and the Gulf of Panama. Pocilloporid corals in Costa Rica and the Gulf of Chiriqui demonstrated lunar spawning activity, with mature gametes present a few days before and after full moon. Some limited spawning may have occurred also at new moon. While frequent gamete maturation has been demonstrated in this study, the relatively low rates of larval recruitment occurring on eastern Pacific coral reefs disturbed by the 1982–83 El Niño suggest that the recovery of important frame-building corals could be greatly prolonged.     

Diel and lunar patterns of reproduction in the Caribbean and Pacific sergeant major damselfishes Abudefduf saxatilis and A. troschelii

Diel and lunar reproductive patterns of the Caribbean and Pacific sergeant major damselfishes Abudefduf saxatilis and A. troschelii respectively, were studied i the San Blas and Northern Perls Islands of Panama in 1981 and 1982. These are sister-species that have diverged since the uplift of the isthmus of Panama. Caribbean A. saxatilis females typically complete spawning in the nests of males earlier in the morning than do Pacific A. troschelii females. Spawning is rarely observed during the afternoon. Hatching occurs four or five days after spawning in the Caribbean species, and after four days in the Pacific species. In both, fry hatch during the hour following sunset. There also are differences in lunar patterns of spawning between the two species. The Caribbean A. saxatilis spawns throughout the month, exhibiting no lunar periodicity. In the Pacific A. troschelii spawns within nine days on either side of new moon. During the wet season, reproductive activity at a site spans three or four days, but is extended to eight or more days during dry season upwelling in the Bay of Panama. Thus, reproductive synchrony is much greater within than between colonies of the Pacific sergeant major, particularly in the wet season. Reproductive synchrony in the Pacific sergeant major might be beneficial because fry hatching at dusk encounter favorable tides for transport away from reef-based predators around new moon, or because reproductive synchrony reduces the risk of predation on unhatched embryos in nests guarded by males. Tidal amplitude and the rate of predation on embryos in nests are greater in the Pacific.     

Sexual reproduction of the reef coral Montastreacavernosa (Scleractinia: Faviidae) in the Santa Marta area, Caribbean coast of Colombia

Sexual reproduction of the reef-building coral Montastreacavernosa was studied in the Santa Marta area, Caribbean coast of Colombia, from October 1990 to October 1991. The area is subjected to a seasonal upwelling–outwelling regime. From microscopical analysis of tissue sections sampled each lunar month around the full moon, it was confirmed that this is a gonochoric, broadcasting species, with a single gametogenic cycle per year, and a 1:1 sex ratio. Oogenesis began a few weeks after spawning, and extended for 11 mo. The onset of spermatogenesis occurred just before the full moon of June, later than at other localities where sea-water temperatures during the first half of the year are not as low. The amount of reproductive tissue strongly increased in both sexes after July, in association with an increase in sea-water temperature and a decrease in photoperiod. The gonad index of fully mature female colonies was about four times lower than at other localities, perhaps due to the stressful seasonal regime. The male gonad index was lower than that of the female, indicating sex-related differences in the rates of biomass allocation to reproduction. As in other Caribbean localities, spawning occurred after the full moons of August and September. However, there was some evidence of a third spawning episode after the full moon of October, possibly associated with a delay in the occurrence of maximum sea-water temperatures in near-equatorial localities compared to higher latitudes. The probability of cross-fertilization in this species with a gonochoric breeding system and a broadcasting mode of reproduction is favored by its balanced sex ratio, its usually high within-reef zone abundance, and by localized and repeated spawning episodes, synchronized by lunar phase. Received: 19 April 1996 / Accepted: 24 September 1996     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). I. Gametogenesis and larval brooding

Anthelia glauca Lamarck, 1816 is a gonochoric, external-brooding soft coral found in KwaZulu-Natal. It is reproductively active in the summer months. The development of gametes produced in late summer is arrested in winter. Several stages of gametes are found at the base of the polyps, and female polyps produce several cycles of larvae over an extended breeding period of 4 to 5 months. Larvae are brooded in a unique pharyngeal brooding pouch not yet described in other coral species. The brood pouch consists of an expansion of the pharynx with constrictions proximal and distal to the embryos and larvae. Our data suggest that egg transfer and fertilization occur at full moon and the mature larvae are released after new moon. Zooxanthellae are absent in A. glauca oocytes, but zooxanthella infestation commences at the immature larval stage. Received: 15 July 1997 / Accepted: 12 March 1998     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Short-term temporal patterns of early recruitment of coral reef fishes in the tropical eastern Pacific

Short-term temporal patterns of recruitment have been described in a variety of coral reef fishes and have often been related with lunar and tidal cycles. While the relative importance of lunar and tidal factors in determining recruitment patterns has been difficult to assess, most studies have been done in the Caribbean and Indo-Pacific, where tidal amplitudes are small. We studied the short-term temporal dynamics of fish recruitment at Gorgona Island (tropical eastern Pacific), where there is a large tidal amplitude (~4.4 m). Every other day during three consecutive months in 1998, we directly measured the magnitude of reef fish recruitment to standardized coral units (SCUs) isolated from natural reefs. A total of 40 species from 21 families settled on the SCUs. Of 11 species with sufficient numbers for meaningful statistical analyses, two (Lutjanus guttatus and Pomacanthus zonipectus) had lunar recruitment with peaks near the new moon; three combined species of antennariids showed semilunar recruitment with peaks near moon quarters; and eight other species showed sporadic and aperiodic recruitment pulses. The contribution of lunar (moonlight intensity) and tidal factors (tidal amplitude and net tidal flow) to recruitment dynamics varied among species, although it was generally low (<18%) even among species with periodic patterns, except perhaps in L. guttatus. In this species, recruitment magnitude correlated negatively with moonlight intensity, accounting for 34.5% of the variance. Post-settlement predation by roving predators may be one cause of this relationship. In the remaining species, particularly those with sporadic and aperiodic recruitment pulses, stochastically varying weather and oceanographic events may be more important in determining temporal variation in recruitment.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Spawning periodicity and habitat of the palolo worm Eunice viridis (Polychaeta: Eunicidae) in the Samoan Islands

The relationship between the phase of the moon and the emergence of the epitokous segments of the palolo worm Eunice viridis Gray has been known to the natives of the Samoan Islands for centuries. They predict the date and time of day when the emergence occurs so that they can be ready to catch the worms. This phenomenon is one of the best known examples of lunar periodicity. It was first described scientifically at the end of the last century. My own investigations concern the occurrence of the worms in the reef, in which they gnaw long tunnels through the massive blocks of coral limestone at levels characterized by the occurrence of symbiotic algae. Apparently the algae are the main sources of nutrition for the worms. The casting off of the epitokous segments occurs at the third quarter of the moon in October or November. An analysis of known dates on which the swarms of worms have appeared permitted a precise method of prediction to be formulated. The causality of this periodicity is discussed.     

Reproductive strategies and diet in deep-sea nuculanid protobranchs (Bivalvia: Nuculoidea) from the Rockall Trough

Although the taxonomy of deep-sea protobranch bivalves is becoming better known, relatively little information is available on their reproductive biology and whether or not populations show reproductive periodicities. We have examined the reproductive biology of three common sympatric species as part of a long-term time-series of samples taken from 2900 m in the Rockall Trough from 1973 to 1983. Malletia cuneata Jeffreys, 1876 produces a maximum of 30 oocytes at any one time and these grow to a maximum size of 240 m. Maximum fecundity of Ledella pustulosa (Jeffreys, 1876) and Yoldiella jeffreysi (Hidalgo, 1877) is 174 and 360, respectively, and both species produce an egg of 120 m in diameter. These data indicate lecithotrophic early development in L. pustulosa and Y. jeffreysi, but direct development in M. cuneata; however, evidence from the prodissoconch length of M. cuneata suggests lecithotrophic development. L. pustulosa and Y. jeffreysi also differ from M. cuneata in having a distinct reproductive cycle with spawnout in the winter months. Both the continuously breeding M. cuneata and the seasonally breeding L. pustulosa ingest diatoms, coccoliths and foraminiferans, but whereas the diet of M. cuneata appears to be constant throughout the year there is an apparent reduction in the feeding activity of L. pustulosa concomitant with the deposition of phytodetritus on the deep-sea bed.     

Variation in the genetic composition of coral (Pocillopora damicornis and Acropora palifera) populations from different reef habitats

The genetic structure of populations of the corals Pocillopora damicornis and Acropora palifera was examined in three habitats at One Tree Island during March and April 1993, using electrophoretically detectable variation at six allozyme loci. There were significant genetic differences among populations of P. damicornis within each of the reef crest, lagoon and microatoll habitats. The level of differentiation among populations was similar in each of the habitats. Differences between populations of P. damicornis from lagoon and microatolls were no greater than that within habitats, but genetic differentiation of these from crest populations was much higher. There was no difference in the genetic composition of A. palifera populations within or between the lagoon and microatolls, the only habitats where this species was found. Both coral species had observed:expected (G _O:G_E) genotypic diversity rations >0.80, indicating predominantly sexual reproduction. These data, the high genotype diversity and general conformance of genotype frequencies to those expected under conditions of Hardy-Weinberg, suggested panmixis at each site. The high degree of sexual reproduction in the P. damicornis populations is unusual for a species where asexual reproduction has been the dominant mode of reproduction reported to date. Gene flow in both species was considerable between the lagoon and the closed microatolls. The genetic differences between populations of P. damicornis in these habitats and the reef crest may reflect the relative isolation of all populations within the closed One Tree Lagoon from those outside. However, local currents appear to offer effective means of dispersal between the habitats, suggesting that the genetic differences result from natural selection in the different environments within One Tree Lagoon and the reef crest.