Population dynamics and reproductive cycle of Pholoë minuta (Polychaeta: Sigalionidae) in Galway Bay

Pholoë minuta (Fabricius) is a prominent member of the Amphiura filiformis (O.F. Müller) subcommunity in Galway Bay, Ireland. The aim of the present investigation was to study the population dynamics and reproductive cycle of P. minuta in this area. Monthly sampling over a 2 yr period (November 1978–October 1980), at a permanent station (18 m) with a fine sand substrate, revealed a mean density of 96 individuals per 0.1 m². The width of the fifth elytron was correlated with total body length and was used as a basis for analysing population structure. Four distinct year classes were evident and estimates of recruitment levels and growth rates were calculated. P. minuta is polytelic and reproduces for the first time when approaching 3 yr of age. Spawning occurred in March—April and appeared to be associated with a seasonal low in bottom-water temperature.     

The annual reproductive cycle of the holothurian Aslia lefevrei (Dendrochirota: Echinodermata)

The dendrochirote holothurian Aslia lefevrei (Barrois) is locally and densely aggregated (>70 individuals m^-2) in Galway Bay, on the west coast of Ireland. It has a discrete, relatively short breeding period from the end of February to the beginning of April, with peak activity in March. The sexes are separate and the single gonad, consisting of numerous tubules (100 to 800), lies in the anterior coelom. Both gonad index values and histological observations indicate that gametogenesis resumes in June/July after a brief post-spawned resting period. Little oocyte growth and sperm production occurs over the warm summer months when the holothurian is actively feeding. Gonad index values, however, increase rapidly during this period and reach high values by early winter without concomitant increases in maturity indices. This is subsequently followed by slight reductions in the gonad indices, before reaching peak values in February. It is suggested that this is probably due to the initial build-up and subsequent assimilation of storage products in the gonad, the development of gametogenic products and the frequent assimilation thereafter of unwanted ova. Spawning generally takes between 5 and 60 min, with the entire contents of the gonad rarely being shed in a single release. During this process, the holothurian raises its anterior end and, with the extended tentacles close together, emits a stream of gametes from the genital pore in the mid-dorsal line. Laboratory experiments and general observations indicate that increased temperature and low light levels may influence spawning behaviour.Contribution No. 248 to the School of Marine Sciences, University College Galway, Ireland     

Larval abundance and recruitment of the sand dollar Dendraster excentricus in Monterey Bay,California, USA

Seasonal abundance of planktonic larvae of the sand dollar Dendraster excentricus was determined in Monterey Bay, California, USA. Larvae were counted from two offshore stations and also over a coastal sand dollar bed, and these data were compared with settlement in the sand dollar bed, with adult population structure and with adult reproductive condition. These measurements were made during the period July 1978 to October 1980 and in October 1981. Sand dollar larvae were most abundant in the plankton during the summer, a period when phytoplankton productivity tended to be high and currents were relatively slow and variable. In some years, small-scale current variations appeared to prevent many larvae in the open bay from reaching the nearshore sand dollar beds; however, in other years, massive shoreward transport of the larvae evidently did occur, since the adult population in the sand dollar bed exhibited a mode in size, indicating a large settlement. A comparison of settled individuals in 1980 and the adult size-frequency distribution in 1981 gives an estimated mortality rate of 88% yr^-1 for early juveniles of D. excentricus.     

Reproductive cycle,development and recruitment of two geographically separated populations of the dendrochirote holothurian Aslia lefevrei

The boreo-mediterranean holothurian Aslia lefevrei (Barrois) occupies cryptic habitats from low-water mark to ca. 20 m. During the present work specimens were collected from the west coast of Ireland (February 1979 to August 1980), where A. lefevreiis a prominent element in two discrete epibenthic assemblages, and from the north-west of Spain (September 1981 to August 1982), where this holothurian is densely aggregated in association with raft cultivation of the mussel Mytilus edulis. Its reproductive cycle, spawning process, embryological development, settlement, and early growth rates have been documented by means of laboratory and field studies. Depending on geographic location, spawning in the wild occurs between January and May. Development to the gastrula takes some 3 to 4 d. Uniformly ciliated dipleurula/auricularia hatch 12 to 24 h after gastrulation. The pentactula stage is attained approximately 7 d after fertilization, following which the larva settles. Newly settled individuals utilize the five primary tentacles for both feeding and locomotion. With the advent of small numbers of podia in the ambulacral regions, two of the primary tentacles prolong their locomotory and anchorage capabilities while the remaining tentacles develop adult characteristics and function solely in feeding. Growth was followed for an estimated 30 mo period using length-frequency analyses. In the first year of life, A. lefevrei attain an average length of 3.4 cm; individuals on the sea floor can attain up to 18 cm, indicating a possible life expectancy of 10 yr or more.Contribution No. 301 from the School of Marine Sciences, University College Galway, Ireland     

Spectral properties and photosynthetic action in red-tide populations of Prorocentrum micans and Gonyaulax polyedra

Three sets of experiments were carried out between April and September 1987 to assess reproductive energetics in a laboratory population of the capitellid polychaete Capitella capitata type I. In the first experiment, the population was sampled intensively over a short period of 3 wk. Weight-specific reproductive output did not differ significantly between females and hermaphrodites. However, because hermaphrodites were significantly bigger than females, their fecundity was significantly higher. In the second experiment, fecundity and weight-specific reproductive output were monitored weekly over the oscillation cycle of the population (30 wk). Both fecundity and weight-specific reproductive output varied drastically (40 to 80 eggs per female and 17 to 57%, respectively). In the third experiment, non-sexed juveniles were reared for 15 d on six ration levels (normalized on the basis of organic nitrogen) of four different food sources. The maximum weight-specific reproductive output was 70% (for females). Both fecundity and weight-specific reproductive output were positively correlated with food ration and varied with food type. The data on weight-specific reproductive output recorded in the present study have been compared with those of other iteroparous polychaetes in the literature. Reproductive output in C. Capitata type I varies more widely and is greater than in other iteroparous polychaetes. In this species, reproductive output and population dynamics vary concomitently with the level of organic matter in the habitat.     

Genetic heterogeneity in a single year-class from a panmictic population of adult blue rockfish (<Emphasis Type="Italic">Sebastes mystinus</Emphasis>)

We used microsatellite genetic markers to investigate adult population structure and the formation of a new year-class in Sebastes mystinus (blue rockfish). Since S. mystinus may live as long as 45 years and reach reproductive age at approximately 5 years, the adult population may contain as many as eight generations of reproductive adults. We investigated whether the juveniles of the 2000 year-class and the adult population were genetically homogeneous along the California coast. We sampled approximately 100 juveniles from three sites, two sites along the Monterey Peninsula (Carmel and Monterey) in central California and one at Fort Ross in northern California, and approximately 50 adult S. mystinus from five sites throughout the population center. The adult sampling spanned approximately 700 km from the northern Channel Islands to Fort Bragg. The juveniles showed significant heterogeneity in allele frequencies among distant locations and genetic homogeneity among adjacent locations. In contrast, the adults showed genetic homogeneity over large distances (San Miguel Island to Fort Bragg), indicating little limitation of gene flow in this region. Allele frequencies of juveniles differed from adult samples and in some cases reduced genetic diversity indicative of sweepstakes recruitment (small sample of the adult reproductive potential). The genetic structure of the 2000 year-class suggests that despite a genetically homogenous adult population, settled juveniles can be genetically heterogeneous along the California coast. The results also suggest that the adults, with several year-classes, are capable of maintaining a panmictic population despite the genetic distinctiveness of individual year-classes.     

Population dynamics, sex ratio and size at sex change in a protandric simultaneous hermaphrodite, the spiny shrimp Exhippolysmata oplophoroides

One of the main goals of sex allocation theory is understanding sex ratio evolution. However, theoretical studies predicting sex ratios in species with unusual sexual systems, such as protandric simultaneous (PS) hermaphroditism, are rare. In PS hermaphrodites, juveniles first develop into functional males that mature into simultaneous hermaphrodites later in life. Here, we report on the sex ratio (males/males + hermaphrodites) in the PS hermaphroditic shrimp Exhippolysmata oplophoroides. A 2-year study demonstrated that hermaphrodites dominated the population in two different bays. This skewed sex ratio may be explained by limited encounter rates among conspecifics. In agreement with this idea, the density of shrimps was extremely low (≤1 shrimp km⁻²) at the two study sites. Size at sex phase change and sex ratios remained relatively stable through time at the two bays. The stability of these parameters might be explained by the rather steady population structure of this species during the study period. A review of sex ratios in PS hermaphroditic shrimps (Lysmata and Exhippolysmata) revealed considerable variation; some species have male- and others hermaphrodite-skewed sex ratios. The conditions explaining inter- and intra-specific sex ratio variation in protandric simultaneous hermaphroditic species remain to be addressed.     

Demography and reproductive biology ofMelinna palmata (Ampharetidae: Polychaeta) in Inner Galway Bay on the west coast of Ireland

A high-density population of the polychaeteMelinna palmata Grube was the focus of a two-year study at a shallow-water location in Inner Galway Bay on the west coast of Ireland. Mean densities were highest in the second year of the study following successful recruitment. Size-frequency analysis of population structure (April 1983 to March 1985) revealed the presence of four cohorts, with the 1983 settlement becoming the dominant modal class. Size comparison of this cohort with the 1982 settlement showed that growth rates were lower and mortality higher in the 1983 settlement, which may be related to density dependent intra-specific competition.M. palmata at this location live for 2 to 2.5 yr with a small number surviving to 3 yr of age.M. palmata in Galway Bay is dioecious and has an even sex ratio. Reproductively mature worms were approximately 2 yr old when they first spawned. Spawning took place between May and July, when mature oocytes had a modal size of ca. 290µm. Proliferation of previtellogenic oocytes (ca. 10µm) from the gonads occurred following spawning. Growth over the winter was slow and was followed by a period of rapid growth between March and May, coincident with rising water temperatures. The presence of mature unspawned oocytes, which were resorbed after commencement of new proliferation, was taken as evidence of the species propensity for polytely. However, post-spawning mortality appeared to be high, so that only a small number of individuals survived to spawn the following year. The highest abundance of mature males was encountered between May and August, but lower levels of mature stages were present throughout the year.     

Reproduction in Amphiura filiformis (Echinodermata: Ophiuroidea): Seasonality in gonad development

The burrowing ophiuroid Amphiura filiformis (O. F. Müller) (Echinodermata: Ophiuroidea) colonises an extensive area and is a numerically dominant member of the macrofauna in Galway Bay on the west coast of Ireland. There, it has a discrete, relatively short annual breeding period, from June to September, with peak activity in the month of August. The breeding period approximates in timing those of other European populations. The oocytes of a particular year are laid down in the autumn of the previous year; they begin to develop in spring, with the period of fastest growth in May/June. Gonad growth corresponds quite closely to the annual rise in water temperature, with spawning taking place during the warmest months of the year. The species is polytelic, and it is suggested that the long-lived Galway Bay individuals may breed every year for several years. The relationship between spawning and population dynamics is discussed in the light of current, available literature concerning A. filiformis from other parts of Europe. The role of so-called ultimate factors in the process of ophiuroid maturation and spawning is discussed.     

Dynamics and biological cycle of a Melinna palmata (Ampharetidae) population during the recolonisation of a dredged area in the vicinity of the harbour of Brest (France)

The reproductive cycle and growth parameters were studied in a Melinna palmata population, for 3 yr after dredging, in the vicinity of the harbour of Brest, France. This species first settled in May 1979, 8 mo after dredging had ceased. Measurements of the diameter of coelomic oocytes indicated that spawning took place from August to October and settlement in autumn and early winter. The first settled cohort displayed a high growth rate (0.25 mm width mo^-1 in summer) and a low longevity (22 mo); the growth rate of the second settled cohort was not significantly different from the first, but its longevity was higher (28 mo); the third settled cohort had a lower growth rate (0.15 mm width per month), but lower mortality and therefore a longer life expectancy. A fourth cohort appeared in August 1981 but was not followed because of the termination of the study. M. palmata colonised the dredged area in two ways: by settlement of juveniles in autumn and by probable immigration of young and adults in May. The population increased in number and biomass from the end of dredging (August, 1978) to June 1981 and decreased after this period. The demographic strategy of this species changed from r- to K-selection during this recolonization as a result of intraspecific and interspecific competition.     

Quantitative and ultrastructural studies on the reproductive biology of the polychaete Pholoe minuta in Galway Bay

Previous works by the present authors have identified, in a qualitative fashion, the reproductive cycle of Pholoe minuta (Fabricius) in inner Galway (53°13.5N; 9°7W). This quantitative, histological study, carried out over the period late May 1981 to late April 1982, shows that P. minuta is polytelic (two to three annual spawnings in an individual's lifespan). Individuals become sexually mature for the first time when approaching 3 yr of age. Estimates were made of the size of the reproductively active population (47 to 49%), sex ratio (1:1), duration (10 to 11 mo) and pattern (unimodal) of gametogenesis and fecundity levels (210 198 eggs m^-2). Maturation (mean oocyte size=85 m) and spawning (for 1 wk during late March through early April) were shown to be synchronised. Throughout oogenesis, follicle cells play an important role in binding together oocyte clusters, and desmosomal contacts are evident between oocyte and follicle cell plasmalemmas. No evidence for the synthesis of yolk precursors in the follicle cell cytoplasm was discovered. Oocytes are rich in Golgi apparatuses (yolk synthesising) and nuclear pores are numerous. Rough endoplasmic reticulum is distended in nature. Egg envelope development does not appear to commence until after the onset of vitellogenesis. Mature spermatozoa are of the primitive bullet-shaped type, with an unmodified acrosome. A possible method of elevation membrane formation is described.     

Analysis of individual year-classes of a marine fish reveals little evidence of first-generation hybrids between cryptic species in sympatric regions

As settled juveniles and adults, blue rockfish (Sebastes mystinus) are nonmigratory inhabitants of kelp and rocky reef habitats along the California coast, USA, and prior to settlement, they possess a pelagic larval and juvenile stage lasting 3–5 months. A previous study of adults revealed two cryptic species within S. mystinus and evidence of reproductive isolation in a region where both cryptic adults co-occur. Given this pattern of reproductive isolation, we investigated the degree of hybridization or introgression in individual year-classes shortly after juvenile settlement in two different years (2001 and 2002). Using microsatellite markers, we found little indication of hybridization in new juvenile year-classes despite an adult population that comprised both cryptic species. However, we found an average of two percent of hybrid or introgressed individuals in regions with a low frequency of one of the two species. Therefore, while the lack of hybrids or introgression supports the hypothesis of reproductive isolation between the cryptic species within S. mystinus, the age-structured analysis also revealed a spatial pattern of low-frequency differences in the number of introgressed individuals. These results suggest that reproductive barriers may breakdown when one of the two species predominates the regional adult gene pool.     

Population dynamics and production of euphausiids

Seasonal changes in abundance, size and aspects of the population structure of Meganyctiphanes norvegica (M. Sars) and Nyctiphanes couchi (Bell) are described from samples taken with the Continuous Plankton Recorder at 10 m depth over a 2 yr period (1966 and 1967) in the North Atlantic Ocean and the North Sea. M. norvegica lived for a maximum of just over 2 yr, and adults of both year-classes spawned during a limited breeding season in the spring or summer. N. couchi spawned over a prolonged breeding season, giving rise to a complex of cohorts with overlapping size ranges. It was concluded that 3 or 4 cohorts were spawned in each year and that the maximum life span was probably greater than 1 yr, although maturity may be attained in less than a year. Estimated annual production at 10 m depth for M. norvegica ranged from 0.80 to 18.74 mg m^-3yr^-1 and for N. couchi from 0.67 to 8.23 mg m^-3yr^-1. P:B ratios ranged from 1.3:1 to 6.3:1 for M. norvegica and 4.0:1 to 5.5:1 for N. couchi.     

Occurrence and significance of nitrogen transport in the brown alga Laminaria digitata

Nitrogen transport in a Laminaria digitata (Huds.) Lamour population growing at Arbroath, Scotland has been investigated (during the period 3 June 1981 to 3 July 1981), using ¹⁵N as tracer. NO ₃ ^- was assimilated both by the blade meristem and by the mature blade. NO ₃ ^- uptake by the blade meristem alone was insufficient to supply the nitrogen demand for growth. This additional demand was met by nitrogen transport from the mature blade to the meristem. It was estimated that 70% of the nitrogen demand of the meristematic region was supplied by nitrogen transport from the mature blade. Although transport occurred, the size of the endogenous pool of stored nitrogen in the mature blade did not change, the quantity of nitrogen transported being equivalent to the amount of NO ₃ ^- assimilated during the experimental period. The possible co-transport of nitrogen and carbon is discussed.     

Tidal influences on the photoperiodic induction of tetrasporogenesis in Bonnemaisonia hamifera (Rhodophyta)

Photoperiodic control of tetrasporangium formation was studied in two low intertidal populations of Bonnemaisonia hamifera (Trailliella-phase) in Galway Bay, Ireland between September 1985 and February 1986. Photoperiodic induction was affected by tides, principally by the time of day at which high and low water occurred through the spring/neap cycle. High water of spring tides falling at the beginning and end of the day reduced the effective daylength for plants, and caused an early onset of reproduction. The threshold irradiance signalling light-dark transition was estimated to lie between (1-) 5 and 10 mol photons m^-2 s^-1 (400 to 700 nm). Low water of spring tides falling in the middle of the day exposed plants to high air temperatures during periods when seawater temperatures were below the lower limit for sporangium induction. This caused reproduction to be resumed. Quantitative differences between the two populations were caused by microclimatic effects (temperature, light reduction during tidal flooding) resulting from their different vertical heights on the shore.     

Reproductive success,relative abundance and population structure of two species of Nephtys in an estuarine beach

During 1978 and 1979 specimens of Nephtys hombergii in the River Tyne estuary (North East England) became gravid during the winter but did not spawn in the spring breeding season; instead their gametes were resorbed. The drastic effects of these spawning failures were apparent in the population structure in 1980, when the 1978 and 1979 year-classes were shown to be virtually absent, by the size of the individuals in the population and the numbers of growth lines in their jaws. The population structure in 1975, by contrast, indicated that recruitment during the preceding 4 to 5 yr had been good. Spawning was also effective in 1980, and an 0 group was detected in August 1980. Comparable data has been obtained for N. caeca, which occurs in the same beach. It shows that recruitment of this species during 1976–1980 was consistently good, whereas analysis of the age-class frequencies suggests that recruitment of N. caeca was poor in 1973 and 1975. The dominance relationship of the two species in the beach has been reversed during the period 1975–1980. It is suggested that the sympatric distribution of the two species is maintained in part by periodic reproductive failure. The causes of this are not yet known; possible reasons, including the extremely cold conditions experienced in 1978–1979 are discussed. The difference in frequency of the 1978, 1979 and 1980 year classes will now permit a rigorous evaluation of the relationship between the number of growth lines in the jaws and age in N. hombergii.     

Population biology of hyperbenthic crustaceans in a cold water environment (Conception Bay,Newfoundland). II.<Emphasis Type="Italic"> Acanthostepheia malmgreni</Emphasis> (Amphipoda)

Seasonal population dynamics of the gammarid Acanthostepheia malmgreni Goës in Conception Bay, Newfoundland, were examined from October 1998 to November 2000. This species exhibited a 2.5-year life span, with the reproductive cycle correlating with seasonal phytoplankton flux. Females were semelparous and died following a 5-month brooding period and the subsequent release of juveniles in April and May. The biennial life cycle of this population should result in the presence of two cohorts in the hyperbenthos at any given time. However, the cohorts alternated in strength from year to year, which affected annual density, biomass and production during the study period. Densities were 64±87 ind. per 100 m³ in 1999 and 491±492 ind. per 100 m³ (mean±SD) in 2000. Secondary production was estimated at 18–44 mg C m^–2 in 1999 and 180–311 mg C m^–2 in 2000. The annual P/B ratios were 0.89 and 2.27 in 1999 and 2000, respectively. Growth varied both among and within cohorts, with different life-history stages exhibiting variable growth rates ranging from 0 to 12 mg dry mass month^–1.Communicated by J.P. Grassle, New Brunswick     

Determinants of reproductive potential and population size in open populations of Patella vulgata

The importance of external and internal population processes in determining variation in reproductive output and variation in population size were quantified with model simulations for open populations of the sequentially hermaphroditic limpet Patella vulgata using field data from the Isle of Man and South West Ireland. Cross-correlation analyses of model outputs and elasticity analyses show that population dynamics are dominated by the effects of large females, and that recruitment adds little to reproductive output. However, populations experiencing low but highly variable recruitment appear male limited and recruitment pulses carrying young males into the population are correlated to reproductive output with a 2–5-year lag. We conclude that pulses in recruitment can be a major structuring force in these limpet populations, but site-specific post-recruitment processes will determine the relative importance of recruitment to population dynamics and the lag between recruitment and reproductive output.     

Nitrogen uptake kinetics in three year-classes of Laminaria groenlandica (Laminariales: Phaeophyta)

Nitrate and ammonium uptake rates were measured for three year-classes of the perennial macrophyte Laminaria groenlandica Rosenvinge, collected from nitrogen-depleted waters in Barkley Sound, British Columbia, Canada, in summer 1981. A time course of uptake rate revealed that ammonium uptake was high during the first hour and then decreased for all three year-classes; the opposite pattern was exhibited for the time course of nitrate uptake rate. Nitrate uptake rate increased linearly with nitrate concentration up to the highest level tested (60 M). The nitrate uptake rate of first-year plants was three times higher than second- and third-year plants; ammonium uptake rates showed similar patterns to those for nitrate. The interaction between nitrate and ammonium was examined for first-year plants. Nitrate and ammonium were taken up simultaneously and uptake rates were identical and equal to uptake rates when only nitrate or ammonium was present in the medium. Therefore, first-year plants are able to take up twice as much inorganic nitrogen per unit time when both nitrate and ammonium are present. First-year plants showed significant diel periodicity in ammonium uptake rates, whereas second- and third-year plants showed no periodicity in nitrate or ammonium uptake rates.     

Annual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus

Interannual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus (De Haan) was studied for a 7-yr period (1981 to 1987) in an intertidal mudflat in the estuary of the Tatara-Umi River, Fukuoka, Japan. Density of juveniles fluctuated greatly, and maximum annual density was low in 1981 to 1984 (22.0 to 44.0 m^-2) and very high in 1985 to 1987 (98.5 to 188.6 m^-2). In 1980, recruits may have been very few, though there is no data on or before 1980, because no crab of the 1980 cohort was found during the study period. Density of adult crabs was almost stable (11.0 to 32.6 m^-2), perhaps because of density-dependent mortality. Under the high density conditions prevailing from 1985 to 1987 crabs grew slowly and longevity was shorter. Male chela and female abdomen were dwarfish, and females produced few and small broods. The changes of life-history traits under high density conditions were thought to result from strong intraspecific competition for space.