Hydrography and distribution dynamics of larval and juvenile fishes in the coastal waters of the Tanshui River estuary,Taiwan, with reference to estuarine larval transport

Distribution dynamics of fish larvae and juveniles in the coastal waters of the Tanshui River, Taiwan was studied fortnightly using surface horizontal tows with a larval net in daytime during the period from early April through early June 1991. Environmental factors, including water temperature, salinity, dissolved oxygen, pH, transparency and depth at sampling stations, were also monitored. A total of 10737 fish eggs and 1387 individuals, representing 43 families and 93 species, was collected during five cruises from 12 stations in the coastal waters. Most fish were estuarine-dependent marine species. Liza macrolepis, Ambassis gymnocephalus, Terapon jarbua, Mullidae and Gobiidae were the most dominant, making up 64.7% of the total catch. Early life stages, including egg, preflexion, flexion and postflexion larvae were abundant in surface samples. However, yolk-sac larvae were absent in the surface water, probably due to an ontogenetic behavioral shift as a consequence of a change in specific weight during early development. The species composition of fish larvae and juveniles was related to the microhabitats found in the coastal waters. The physico-chemical conditions, along with ontogenetic behavior, played an important role in larval fish distribution in the coastal waters.     

Composition and seasonal fluctuations of meiofauna in a Southern African mangrove estuary

Fluctuations in population structure and abundance of mangrove meiofauna were monitored over a period of 14 months. The greatest mean abundance was directly correlated with mean redox potential (Eh), but showed poor relationships with pH and temperature. Maximum density occurred at the mid to high tide levels and nematodes accounted for 80% of the total numbers. Large ciliates were the next most abundant group accounting for 6.4%. These were followed by oligochaetes (4.5%), turbellarians (3%) and kinorhynchs (2.7%). The remainder consisted of low numbers of copepods, polychaetes, gastrotrichs, the larvae of crustacea (prawns and crabs) and insects. Although there was a tendency for density to increase in summer, the numbers were variable and seasonal correlations with physical parameters were obscure. The estimated production from the standing crop is 4.34 gC m^-2 vr^-1.     

Temporal succession and spatial segregation of clupeoid larvae in the coastal waters off the Tanshui River Estuary, northern Taiwan

In the coastal waters off the Tanshui River Estuary in northern Taiwan, eight species of clupeoid larvae were observed. They exhibited a distinct temporal succession in association with seasonal temperature changing. The time of peak abundance of Etrumeus teres was in January, Engraulis japonicus in late April, Sardinella spp. in mid-May, Dussumieria elopsoides in early June, Thryssa dussumieri in late June, Stolephorus insularis in mid-September, Encrasicholina heteroloba in early October and E. punctifer in mid-November. The time intervals of the temporal succession of the fishes were approximately 15 to 25 d in the spring/summer and 25 to 35 d in the autumn/winter fishing seasons. Also, they showed spatial segregation by distributing in areas with different water depths: Sardinella spp. at a water depth of 10 to 20 m, T. dussumieri at less than 10 m, E. heteroloba at 20 to 50 m and E. punctifer at 10 to 40 m. The larvae of these sympatric clupeoid species segregated their nursing periods and areas apparently to reduce competition for habitat and thus to maximize the utilization of resources. Received: 3 March 1997 / Accepted: 19 March 1997     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Statistical modeling of seasonal and environmental influences on the population dynamics of an estuarine fish community

A statistical modeling study was performed on the population fluctuations of the 15 commonest fish species frequenting the tidal Scheldt estuary in Belgium. These included marine juvenile and seasonal visitors, estuarine residents and diadromous fish species that were recorded on the filter screens of a power plant cooling-water intake between September 1991 and April 2001. The species population abundance was regressed against a candidate set of 6 environmental variables and 13 instrumental variables, accounting for seasonality and long-term trends present in the data. Population abundances of the different species were, in general, best described by seasonal variables. Seasonal components contributed, on average, up to 63.8% of the variance explained by the models. Ten species were found to show a slightly negative, though significant, trend over the period of the survey. Most models also included at least one environmental variable, and 25.4% of the explained variance could be attributed to environmental fluctuations. Of all physico-chemical variables, dissolved oxygen was the most important predictor of fish abundance, suggesting that the estuary suffered from poor water quality during the survey. Temperature, salinity, freshwater flow, suspended solids and chlorophyll a concentrations were minor determinants of fish abundance.Communicated by O. Kinne, Oldendorf/Luhe     

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     

Diel and tidal cycles regulate larval dynamics in salt marshes and mangrove forests

Variation in the release and recruitment of larvae of estuarine invertebrates affects the distribution and abundance of adults, as well as trophic interactions in both the plankton and the benthos. Larval release and supply are often timed to environmental cycles such as the diel and tidal cycles. Here, we determined using plankton tows whether the abundance of larvae spanning salt marsh and mangrove habitats across the intertidal landscape varied with diel and tidal cycles. Using three different sampling designs across two sites and within each of two estuaries over a 12-month period, we covered a range of spatial and temporal scales. This allowed us to test the general prediction that densities of meroplankton in the water column would be greater during nocturnal ebb tides than during other phases of the diel or tidal cycle. As predicted, nocturnal ebb tides yielded the highest densities of meroplanktonic larvae and were dominated by first-stage crab zoeae and this finding was most pronounced in the salt marsh. Throughout the course of the year, greater numbers of meroplankters consistently occurred during the ebb tide compared with the flood tide. The densities of other taxa (e.g. gastropods and polychaetes) showed no clear trends with diel or tidal cycles. This study highlights the effects of these pervasive physical cycles on the timing of larval release and supply in the salt marsh–mangrove complex, and emphasises their contribution to the trophic interactions and the dynamics of benthic populations within estuaries.     

Spatial and seasonal differences in the fish fauna in the shallows of a large Australian estuary

Samples of fish were collected by beach seine throughout the shallow waters of the large Peel-Harvey estuarine system (south-western Australia) in the wet (June to November) and dry periods (December to May) between August 1979 and July 1981. The number of species, density and biomass declined with distance from the estuary mouth and rose with increasing temperature and salinity. Both classification and ordination distinguished the faunal composition of the saline reaches of the rivers from that of the narrow Entrance Channel and two large basins (Peel Inlet and Harvey Estuary). Classification also separated the fauna of the riverine group into wet- and dry-period components, and divided samples taken in the Entrance Channel from those in the basins. Differences between the faunal composition of the Peel Inlet and its tributary rivers were related to differences in salinity regime. The riverine fauna was subjected to much more variable and lower minimum salinities. Species characteristic of the rivers included teleosts such as Atherinosoma wallacei and Amniataba caudavittatus, which are estuarine sensu stricto in southwestern Australia, the semi-anadromous Nematalosa vlaminghi and juveniles of the marine Mugil cephalus. The species diagnostic of the wet periods in the rivers were the estuarine species A. wallacei and Favonigobius suppositus, while the dry periods were characterised by the marine species Atherinomorus ogilbyi and Sillago schomburgkii. Marine species also characterised the Entrance Channel (Favonigobius lateralis, Sillago bassensis), whereas the indicators in Peel Inlet and Harvey Estuary were Hyporhamphus regularis and Apogon rueppellii, both of which can pass through the whole of their life cycle in estuarine as well as marine environments.     

Species composition,community structure and zoogeography of fishes of mangrove estuaries in the Solomon Islands

Mangrove estuaries in the Solomon Islands are well developed, but are small and isolated from each other by extensive fringing coral reef lagoons. A total of 136 species of fish were recorded from 13 estuaries (6 estuaries in Kolombangara, 3 in New Georgia, 3 in Rendova and 1 in the Florida Group); none contained more than 50 species. Sampling took place during five 3 wk expeditions from 1986 to 1988. The mean biomass of 11.60 g m^–2 is comparable with that of similar estuaries in northern Australia. Cluster-analysis revealed two patterns of fish species composition. The first group, in which Gobiidae were the most numerous taxon, inhabited soft, muddy-bottom estuaries. The second group, dominated by Pomacentridae, lived in hard-bottom, log-strewn estuaries. The role of the estuaries as nursery grounds for coral reef species was assessed and found to be insignificant, but they are used as feeding grounds by mobile piscivorous species. The species composition of Solomon Island estuaries was compared with that of other Indo-Pacific estuaries. No endemic species were found and the fauna is typical of such mangrove systems throughout the region. However, several taxa that are common in Australia or New Guinea were not found, notably Ariidae, Centropomidae,Pomadasys, and Sciaenidae. These absentees, and the fish fauna as a whole, are discussed in relation to the position of the Solomon Islands at the western edge of the Pacific Plate, the effects of deep-ocean trenches, the recent geological origin of the islands, and possible methods of colonisation from nearby mangroves in Australia and New Guinea. The importance of larval durations and dispersal to colonisation are discussed in relation to oceanic circulation patterns in the Solomon Sea.     

Seasonal and spatial changes in the larval fish fauna within a large temperate Australian estuary

A total of 66814 fish larvae, representing 37 families and 74 species, were collected in samples taken monthly between January 1986 and April 1987 from 13 sites located at frequent intervals throughout the large Swan Estuary in south-western Australia. The Gobiidae was the most abundant family, comprising 88.2% of the total number of larvae, followed by the Clupeidae (3.4%), Engraulididae (2.9%) and Blenniidae (1.0%). The most abundant species were Pseudogobius olorum (53.3%), Arenigobius bifrenatus (31.2%) and Engraulis australis (2.9%). Abundance of fish larvae in the lower, middle and upper regions of the estuary each reached a maximum between mid-spring and early summer, 2 to 4 mo before the attainment of maximum temperatures. Larvae of species such as Nematalosa vlaminghi and Apogon rueppellii were collected only between November and February, whereas those of others such as P. olorum, E. australis and Leptatherina wallacei were present over many months. The times and locations of capture of larvae have been related to the distribution and breeding periods of the adults of these species. The mean monthly number of species was far greater in the lower than upper estuary (14.7 vs 2.7), whereas the reverse was true for mean monthly concentration (42 vs 197 larvae per 100 m³). Classification, using the abundance of each of the 74 species recorded at the different sites, showed that the composition of the larval fish fauna in the lower, middle and upper estuary differed markedly from each other. Most larvae caught in the lower estuary belonged to marine species, whereas those in the upper estuary almost exclusively represented species that spawn within the estuary. The fact that the larvae of the 59 species of marine teleosts recorded during this study were restricted mainly to the lower estuary, and yet contributed only 6.2% to the total numbers for the whole estuary, helps to account for the relatively high species diversity in this region. The lack of penetration of many of these larvae beyond the first 12.5 km of the estuary presumably reflects the weak tidal effect in the wide basins of the middle estuary and saline regions of the tributary rivers. The larvae of the 13 teleosts that typically spawn within the estuary contributed 93.8% to the total numbers of larvae. Most of these estuarine-spawned larvae belong to teleosts that deposit demersal eggs and/or exhibit parental care (egg-guarding and oral and pouch-brooding), characteristics which would maximize their chances of retention within the estuary.     

Spatio-temporal structure of pelagic larval and juvenile fish assemblages in coastal areas of New Caledonia,southwest Pacific

This study aims at describing the diversity and composition of larval and juvenile fish assemblages in coastal areas of New Caledonia, southwest Pacific, and identifying the environmental factors that influence the seasonal and spatial patterns of these assemblages. A total of 97 taxa belonging to 7 orders and 26 familis were captured in three bays near Nouméa by light trapping every month between January 2002 and June 2003. The assemblages were dominated by Clupeiform larvae and juveniles (96.4% of total abundance) and followed by Perciform larvae (3%). The number of taxa per sample varied from less than five in July–August to more than ten in October–November and abundances followed the same seasonal pattern. Analyses of similarity showed significant differences in the assemblages caught in the three bays and analyses of contribution to the dissimilarity revealed that these differences were due to the most abundant families. The constant part of the relationship between environmental variables and the composition of assemblages was assessed by the partial triadic analysis STATICO, a statistical approach that takes into account the strong seasonality of the data. Rainfall, wind direction and thermal stratification of the water column were found to play a major role in the structure of the assemblages, although tidal amplitude and wind speed became important when Clupeidae and Engraulididae were excluded from the analyses. The richness, relative abundances and seasonal variations of the assemblages caught in three bays under study are close to what has been observed elsewhere in the tropics. This study shows the efficiency of the STATICO analysis for identifying the environmental factors that have a permanent effect on assemblages and sorting them out from those which act temporally or on specific locations. The high abundances and diversity of coral-reef fish larvae observed in coastal zones of New Caledonia suggest that further studies are needed to fully explore the role of the coastal zones of New Caledonia as nurseries.     

长江口南北支浮游动物群落生态学研究

长江口特殊的生态环境使得浮游动物的种类组成、密度和时空分布等生态特征有别于其它水体。2005年秋季（丰水期）和2006年春季（枯水期）对长江口南北支浮游动物进行了2个航次的调查,沿南北支由西向东设立3个采样断面共20个采样点。共采集到浮游动物227种,其中轮虫65种,原生动物107种,枝角类15种,桡足类40种。优势种包括龟甲轮虫（Keratella）、刺簇多肢轮虫（Polyarthra trigla）、萼花臂尾轮虫（Brachionus calyciflorus）、砂壳虫（Difflugia）、表壳虫（Arcella）、砂壳纤毛虫（Tintinnid）、中华哲水蚤（Calanus sinicus）、汤匙华哲水蚤（Sinocalanus dorrii）、广布中剑水蚤（Mesocyclops leuckarti）、象鼻溞（Bosmina）等。文章对南北两支浮游动物的种类组成和结构进行了比较,南支浮游动物的种类、数量及生物多样都高于北支,南北两支的浮游动物群落相似度很低。南北两支水域环境的不同造成其浮游动物群落结构的较大差异,文章分析了环境因子与浮游动物群落结构的相关性,其中水温是重要的影响因素,水温的季节性决定了浮游动物在时间分布上的变化。盐度对浮游动物群落结构的影响主要决定了其物种生态类型及空间分布。从整体上看,南北支主要受长江径流控制,浮游动物的种类组成以淡水种为主,咸水种在断面3出现较多。     

Heterotrophic bacterial dynamics in the lower Fraser River,its estuary and Georgia Strait,British Columbia,Canada

Annual cycles in the glucose heterotrophic potentials of the bacterioplankton of the lower Fraser River, its estuary and Georgia Strait surface waters were observed. Maximal microbial heterotrophic potentials were noted during the winter and early spring months, whereas maximal heterotrophic productivities were observed to occur during the summer and fall months. Temperature and soluble organic carbon levels appear to be the most important of the assayed physico-chemical parameters in regulating activities and productivities of the heterotrophic microbes of these waters.     

长江口南北支浮游动物群落生态学研究

长江口特殊的生态环境使得浮游动物的种类组成、密度和时空分布等生态特征有别于其它水体。2005年秋季(丰水期)和2006年春季(枯水期)对长江口南北支浮游动物进行了2个航次的调查,沿南北支由西向东设立3个采样断面共20个采样点。共采集到浮游动物227种,其中轮虫65种,原生动物107种,枝角类15种,桡足类40种。优势种包括龟甲轮虫(Keratella)、刺簇多肢轮虫(Polyarthra trigla)、萼花臂尾轮虫(Brachionus calyciflorus)、砂壳虫(Difflugia)、表壳虫(Arcella)、砂壳纤毛虫(Tintinnid)、中华哲水蚤(Calanus sinicus)、汤匙华哲水蚤(Sinocalanus dorrii)、广布中剑水蚤(Mesocyclops leuckarti)、象鼻溞(Bosmina)等。文章对南北两支浮游动物的种类组成和结构进行了比较,南支浮游动物的种类、数量及生物多样都高于北支,南北两支的浮游动物群落相似度很低。南北两支水域环境的不同造成其浮游动物群落结构的较大差异,文章分析了环境因子与浮游动物群落结构的相关性,其中水温是重要的影响因素,水温的季节性决定...     

Sewage fluxes and seasonal dynamics of physicochemical characteristics of the Bhagirathi-Hooghly River from the lower stretch of River Ganges,India

ABSTRACT

To understand the temporal variations of the physicochemical characteristics of the Bhagirathi-Hooghly River (BHR), three locations representing three districts of West Bengal were selected. The material fluxes from 34 drains during pre-monsoon season was quantified. The analysis of variance (ANOVA) revealed that no significant spatial variations were observed for the physicochemical parameters, whereas seasonal variations were significant. The mean discharge was found to be highest (247.2?×?10³?m³?d^?1) in the midstream drains. Highest mean concentrations of dissolved oxygen (DO) (7.35?mg?L^?1) and nitrate (0.81?mg?L ^?1) were observed during the post-monsoon season followed by the monsoon and pre-monsoon. According to the BIS, WHO and the European standard of water quality (pH, 6.5–8.5; Nitrate, 0–2.5?mg?L^?1;DO, ≥5?mg?L^?1), the results of the respective parameters revealed the BHR system is maintained at high to good water quality, meaning that the BHR system is slightly altered from its pristine environment. The mean concentrations of biological and chemical oxygen demands were found to be high during the monsoon season, revealing that a large quantity of refractory organic matter is transported to the eastern Bay of Bengal from the Ganges.     

Multiple determinants of community structure in shallow marsh habitats,Cape Fear River estuary,North Carolina,USA

The nekton of tidal creeks was studied at 17 sampling localities from September 1977 through August 1978, in the Cape Fear River estuary, North Carolina, USA. Prior to these dates, collections were made at 9 stations beginning in January 1977; these data were used to supplement conclusions drawn from the larger effort. Species recruited from the ocean utilized marsh habitats only temporarily and dominated the catches with over 70% of the total abundance. Their distribution was influenced by salinity gradients and to a lesser extent by substrate characteristics. In addition, temporal habitat partitioning with associated size differences of related species played an important role in structuring marsh nekton communities. A clearly defined ecotone was associated with the mesohaline-polyhaline transition zone, in slainities between 14 and 21 S. Numerous marine stenohaline forms were restricted to salinities above 16 S, thus increasing species richness in high salinity marshes. Despite differences in freshwater flows in 1977 and 1978, major features of the various marsh communities (species associations and relative abundances) exhibited little change throughout the Cape Fear estuary, indicating that these communities were relatively persistent in time. Standing crops for ocean-spawned species at the end of the growing season indicated that considerable annual export in the form of living biomass of fish and shellfish takes place from the marshes. Since most individuals of these species return to the ocean in the fall, an important energy link between the marshes and nearshore marine environment is demonstrated.     

Survival of communally reared larval and juvenile lobsters,Homarus americanus

Survival of individually reared larval and juvenile stage lobsters, Homarus americanus (Milne-Edwards), was significantly higher than in corresponding groups of communally reared individuals. Among communally reared lobsters, the mortality rate was highest in the second-stage larvae and then progressively decreased in the later stages. The relationship between survival and duration of molt period of each life-cycle stage indicates that asynchronous molting in the groups of communally reared lobsters is a contributing factor to the higher mortality rate. The molting and mortality curves of communally held lobsters reared from the first larval to first or second juvenile stage showed best cross correlation at 0- or 1-day time lag. The decreased mortality rate observed in the later larval and juvenile stages appears to have resulted from the establishment of new behavior patterns. Group interactions which are influenced by numerous extrinsic and intrinsic factors lead to higher mortality rate (cannibalism) among communally reared lobsters.     

Sources and major biogeochemical pathways of organic matter in the mangrove system of Rufiji estuary

Fatty acids were extracted from the surface sediments (10 cm) of three sampling sites of Rufiji estuary to infer their sources and biogeochemical pathways. The fatty acids ranging from C₈ to C₂₄ were distinguished from this study, and were broadly classified into saturated (SFAs), monounsaturated (MUFAs) and polyunsaturated fatty acids (PUFAs). SFAs were found to be the major fraction at station 1 and 3 where as at station 2, MUFAs dominated. A total of 19 fatty acids with a total concentration of 64.91 μg/g dry weight were characterised at station 1. C_16:0 was the most abundant fatty acid contributing 21.94% of total fatty acids (TFAs). C_22:2 was the second most abundant, which accounted for 9.46% of TFAs. Fatty acids ranging from C₁₂ to C₂₄ were identified at station 2. C₂₀ fatty acid was the most abundant fatty acid contributing 21.94%, followed by C_16:0. At station 3, fatty acids ranging from C₈ to C₂₄ were obtained. The PUFA C_20:5n-3 was the most abundant fatty acid contributing 21.65%, followed by C_24:0 (15.00% of TFAs). The ratio of lower molecular weight (LMW) to higher molecular weight (HMW) biomarkers was used as an indicators to distinguish higher plants organic matter from algae-derived fatty acids.     

Laboratory study of effects of temperature and salinity on survival and larval development of a population of Rhithropanopeus harrisii from the Mondego River estuary,Portugal

Rhithropanopeus harrisii (Gould) is an introduced species in the estuary of the Mondego River (Portugal): it was first recorded from the Iberian Peninsula in 1989. The larval development of this population was studied under laboratory conditions at different temperatures and salinities, and showed a larval development pattern very similar to that reported for American estuarine populations of this species. Larval development was negatively correlated with temperature. Time to megalopa varied between 7 and 35 d; the first crab (C₁) was reached after a maximum of 11 to 43 d. Larval development was optimum at 25°C and 15S. Larval survival was maximum at 10, 15 and 20S at all three temperatures studied (20, 25 and 30°C). The percentage of abnormal megalopae increased with increasing salinity to a maximum (100%) at 30S; incidence of abnormality was not affected by temperature.     

Spawning behaviour,egg and larval development of the myctophid fish Benthosema pterotum

The distribution of eggs and early larval stages of the myctophid fish Benthosema pterotum was studied during three cruises to the Gulf of Oman during January to February 1981 and February 1983. The eggs and yolk sac larvae were identified and described after artificial fertilization on board the Research Vessel Dr. Fridtjof Nansen. The mature fish spawn in the Gulf at 300 to 100 m depths during early night. The eggs and smallest larvae were found at 300 to 200 m depths. The eggs hatch within 12 h at 21°C. Hatching occurs before the slightly bouyant eggs reach the upper 50 m of the water column.