Population dynamics and secondary production in Melinna palmata (Polychaeta: Ampharetidae)

A Southampton Water (England) population of Melinna palmata Malmgren was studied from July 1978 to March 1980. M. palmata is a gregarious sedentary polychaete living in muddy substratum. The distribution of the polychaete in Southampton Water was related to the silt content of the sediment. The species occurs in sediments of <60% silt, it is dioecious, and the gametes are shed during a prolonged breeding season throughout the year, with peaks in March and July. Mature oocytes measure between 180 and 240 m in diameter at spawning, and fertilization is external. Unshed oocytes are resorbed. Annual secondary ary production was 0.42 gC m^–2yr^–1 and the P:B ratio was 2.19.     

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.     

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.     

Life history and population dynamics ofKinbergonuphis simoni (Polychaeta: Onuphidae)

The relationship between life-history characteristics and population dynamics were investigated in the onuphid polychaeteKinbergonuphis simoni (Santos, Day and Rice) between 1982 and 1987. The studied population is located in Upper Tampa Bay, Florida, USA. This worm attains sexual maturity at 5 to 10 mo of age, depending on temperature. Several consecutive broods are produced during a female's life time, with 7 to 26 young per brood. Adults die in June–July after the breeding season. Generations of breeding individuals do not overlap and the replacement of generations occurs in July–August. Only one extended breeding season is experienced per life time. Life span does not exceed 2 yr. Field population density in 1982 was high in fall and spring (2000 to 9000 individuals/m²), low in summer (1600 individuals/m²), and slightly depressed in winter (2500 to 4000 individuals/m²). Changes in population density may be explained by seasonal, temperature-controlled changes in instantaneous birth and death rates. Birth rates are high in fall and spring, while death rates are high in early summer. Death of juveniles contributes to the winter density decrease.     

Ovary structure and oogenesis in Cirratulus cirratus (Polychaeta: Cirratulidae)

Oogenesis in polychaetes has, in most species, two main phases: a coelomic phase during which vitellogenesis occurs, and an ovarian phase. Details of the latter are known in very few species. Efficient reproduction requires coordination between these two phases of oocyte differentiation. The ovaries of Cirratulus cirratus (O. F. Müller) occur 2/segment throughout the fertile region of the body; 5 main stages of oocyte differentiation can be recognised on them: stem cells, oogonia, premeiotic oocytes, early oocytes and terminal oocytes. A quantitative study of the composition of the ovary throughout the reproductive cycle shows that the ovary size is approximately constant, but that the percentage of cells in the terminal oocyte stage is reduced to zero prior to spawning. This stage in oocyte differentiation is therefore a critical one, at which oogenesis may be arrested. The ovary alternates between a proliferative phase when the terminal oocytes are present, and a non-proliferative phase when they are absent. The overall rate of germ cell production for each ovary is low, less than 1 cell/ovary/day, and the, ovary tissue turnover time is greater than 2 years. A qualitative model describing the production of oocytes is presented, and the implications of the low rate of germ cell production are discussed. A low rate of germ cell proliferation, as in Cirratulus cirratus, implies that the rate of oocyte output into the coelom will be controlled by factors influencing the later stages of ovarian oocyte differentiation.     

Reproduction of a Northumberland population of the polychaete Cirratulus cirratus

A population of Cirratulus cirratus (O. F. Müller) was sampled from September 1966 to September 1968. The population is found intertidally in crevices at Cullercoats Bay, Northumberland (UK). Measurements of the sexual maturity and size of the animals have been made. The breeding season is not restricted, spawning occurs in the population asynchronously throughout the year. Ripe worms and worms that have just spawned are invariably present, but the majority of females are always at intermediate stages of oogenesis. Adult males, however, always contain active spermatozoa. Females, but not males, are devoid of coelomic gametes after spawning; a new generation of coelomic oocytes begin to accumulate in the coelom after 30 days. The growth of the coelomic oocytes is relatively slow, the coelomic phase of oocyte development takes between 1 and 2 years for completion. The coelomoducts which serve for the emission of the gametes are described. An interpretation of the female population structure based upon length and sexual maturity data suggests that most females survive to spawn 2 or 3 times, and some to spawn 4 or 5 times. They grow in length by about 2 cm in the period between successive spawnings. The reproduction of Cirratulus cirratus is compared with that of other annelid species breeding throughout the year, and this unusual mode of reproduction is discussed.     

Population dynamics and secondary production in an estuarine population of Caulleriella caputesocis (Polychaeta: Cirratulidae)

Between July 1978 and March 1980, the distribution, population dynamics and secondary production of Caulleriella caputesocis St. Joseph, 1864, in Southampton Water, South England, were investigated. The distribution of the polychaete was related to amount of silt and copper-content of the sediment, the highest densities occurring in sediment containing 60 to 100% silt and less than 50 ppm copper. Breeding occurred at a low level throughout the year, usually when the worms were over one year old, with peak breeding between May and September. C. caputesocis in Southampton Water is monotelic, and oocytes measured 120 to 160 m in diameter at spawning. Unshed oocytes were resorbed. Annual secondary production varied between 0.01 and 5.90 g C m^-2 yr^-1 and P:B ratio of the species ranged between 0.84 and 7.09.     

Population dynamics and secondary production in an estuarine population of Nephtys hombergii (Polychaeta: Nephtyidae)

From July 1978 to March 1980, a study was made on the distribution, population dynamics and secondary production of Nephtys hombergii Audouin et Edw. occurring in the sublittoral industrialised region of Southampton Water in south England. The distribution of the worm was related to the silt content and copper level of the sediment, the greatest densities of N. hombergii being found in sediment containing 60 to 100% silt. Breeding occurred at a low level throughout the year, with a maximum in July to September and November to January in the second year of growth. Spawning occurred when the oocytes measured 200m in diameter, and unshed gametes were resorbed. Annual production varied between 0.092 and 4.32 g C m^-2 yr^-1 (ash-free dry weight) and amounted to 1.9–39.4% of the total macrofaunal production at the sampling stations. The production:biomass (P:B) ratio of the species varied between 1.6 and 2.9.     

Gametogenesis in Harmothoe imbricata (Polychaeta: Polynoidae)

The processes of gametogenesis in Harmothoe imbricata are described and illustrated. For most of their development the gametes are attached to the walls of segmental blood vessels. In males, only meiosis and sperm differentiation occur while the cells are free in the coelom. In females, oocytes are only released into the coelom briefly, after vitellogenesis has been completed. H. imbricata thus differs from most polychaete species so far described — where in males the spermatogonia divide mitotically to form rosettes after they have been released into the coelom, and in females the whole of vitellogenesis occurs after release of oocytes into the coelom. The oocytes of H. imbricata enter the first metaphase of meiosis before they are shed at spawning. The elongated head of the sperm in H. imbricata is atypical of polychaetes in general.     

Tube-worm-sediment relationships of Diopatra cuprea (Polychaeta: Onuphidae)

Diopatra cuprea (Bosc) builds a long, nearly vertical parchment tube in sediments, which is reinforced with bits of shell, sediment or debris. The tube can be divided on the basis of structure into reinforced sections built at or above the sediment surface, and unreinforced sections built below the sediment surface. If sediment accumulates faster than the worm can build reinforced tube, an unreinforced section results. When this occurs, D. cuprea builds reinforced tube at the new sediment surface, and then pulls it down into the substratum until it joins the next lower reinforced section, or until the worm can no longer overcome sediment resistance. In the latter case, the unreinforced section occurs between two reinforced sections. The tube cap is trimmed as sediment is eroded from around it. Selection for reinforcing materials occurs only in the transitional zone from unreinforced to reinforced tube. The exposed portion of the tube (the tube cap) extends from 1 to 6 cm above the sediment surface as an inverted J. The tube mouth is oriented perpendicularly to the direction of current flow. Tube maintenance continues at temperatures below feeding-response blockage (5° to 10°C), but ceases below about 1.8°C.     

Reproductive behaviour ofNereis diversicolor (Annelida: Polychaeta)

Nereis diversicolor O. F. Müller, collected on the tidal flats of the Jadebusen (North Sea, FRG) in autumn 1987 and 1988, was exposed to different temperatures in the laboratory. Results indicated that maturation was induced by temperatures above 6°C; spawning in early spring was synchronized by raising temperatures after a period of low temperatures in winter, and occurred mainly at new and full moon. During reproduction the female stays inside the burrow; the male releases sperm in front of it; the resultant larvae remain in the tube for 10 to 14 d protected by the female.     

Secondary production ofPrionospio caspersi (Annelida: Polychaeta: Spionidae)

The annual somatic production ofPrionospio caspersi Laubier was estimated, between July 1986 and 1987, at a sandy shallow-bottom station in the area facing the Po River Delta, Italy, where this species is one of the most abundant. Primary recruitment occurs in summer, and maturation of gametes the following spring. Body sizes were estimated from the surface area of the anterior end of worms, projected onto a monitor and recorded using a digitizer tablet. Measurements, to the tenth setiger, were correlated with total length, total body surface area and AFDW. The annual production was 8.06 g m^–2 yr^–1 (AFDW) and the production:biomass (P:B) ratio 4.09. The results are discussed in relation to some physical and chemical features of the environment, stressing differences with the life cycle of other coexisting populations.     

Reproduction and larval development of Marenzelleria viridis (Polychaeta: Spionidae)

The pelagic larvae of the polychaete Marenzelleria viridis (Verrill, 1873) are newcomers among the meroplankton of the Baltic Sea, where they sometimes achieve an abundance of up to 21x10⁶ ind m^-3 near the coast, especially in late autumn and even in winter. Benthos samples were collected in the Darss-Zingst bodden chain from April 1992 to 1993 and used to ascertain the reproductive stages of individuals. Development from fertilized egg to benthic juvenile is described on the basis of field material. Details of the morphology of various pelagic stages and the young benthic worm are presented as drawings and scanning electron microscope photographs. In 1992, development of the gametes started in mid-May. The individuals reached maturity in late September after about 20 wk. The first planktonic stage was the fertilized egg. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Gastrotrochs are present on the third and thereafter every alternate segment. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage, and neuropodial bidentate hooks from the 10th to 11th setiger on. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. The reproductive seasons of various populations of M. viridis and the developmental patterns of their larvae are discussed, and differences are compared with relevant findings for the spionids.     

Functional morphology and feeding behavior of Paraprionospio pinnata (Polychaeta: Spionidae)

The functional morphology and feeding behavior of Paraprionospio pinnata (Ehlers) were studied. The tentacular feeding palps of P. pinnata possess four groups of cilia: laterals, latero-frontal cirri, frontals and basal transverse rows. The lateral cilia beat in metachronal waves creating in current that flows toward the frontal surface of the palp. The latero-frontal cirri deflect suspended particles onto the frontal surface and potentially resuspend deposited particles. The frontal cilia line the groove of the palp and transport particles to the ciliated pharynx. The basal transverse cilia also beat in metachronal waves and together with the frontal cilia of the branchiae produce a U-shaped current that removes particles rejected by the pharynx from the burrow. The significance of particle selection at the site of the pharynx in spionid polychaetes is hypothesized to be constrained by the interaction of specific morphologies and behaviors that reduce the effects of epifaunal predation and browsing. The relationship between sediment permeability and type of respiratory or ventilation current is hypothesized to be a potentially important factor in understanding the distribution of spionid polychaetes.     

Sublethal predation on Polydora cornuta (Polychaeta: Spionidae): patterns of tissue loss in a field population,predator functional response and potential demographic impacts

Patterns of tissue loss due to sublethal predation and potential effects on population dynamics were investigated in the polychaete Polydora cornuta Bosc, 1802. Field observations in southeastern Connecticut showed that the percentage of adult worms regenerating segments and/or feeding palps varied temporally from May to November 1982, but the general pattern suggests a constant level of sublethal encounters. Over all sampling dates, a mean of 14.9 and 7.0% of the population was found regenerating posterior segments and palps, respectively. Worms lost a mean of 19.1% of their segments. There was a weak, but statistically non-significant, size-specific difference in the number of regenerating segments per individual. Worms with 20 to 39 segments and 60 to 80 + segments lost an average of 13.8 and 17.2% of their segments, respectively, but worms with 40 to 59 segments lost an average of 23.4% of their segments. The incidence of regeneration (palps and segments) increased linearly with increasing adult density in the population. There was also a strong positive linear relationship between the number of segments available and the number of segments lost at the population level. This suggests that the constant level of sublethal predation can be explained by predators taking prey in proportion to their density and/or the number of segments available. This pattern is supported by results from a laboratory functional response experiment which indicated that at high prey density, partial consumption of P. cornuta by the predatory polychaete Eteone heteropoda was greater than lethal consumption. Lethal predation of P. cornuta by E. heteropoda did not vary across experimental density treatments. Demographic modelling suggested that sublethal predation can reduce the population growth rate of P. cornuta, but the reduction is less than if the added predation pressure was solely lethal. It was estimated that up to 25% of the population could be preyed upon in a sublethal manner before the potential for population growth fell below population maintenance levels.     

Fecundity and reproductive effort in Ophryotrocha labronica (Polychaeta: Dorvilleidae)

Reproductive effort in terms of fecundity and energy allocation was studied in Ophryotrocha labronica La Greca and Bacci, 1962 a small, semicontinuous iteroparous species. In O. labronica fecundity is more or less constant throughout life, and the total fecundity of the 64 couples examined in this experiment only declined by 12% from the first to the last spawning. No linear relationship was found between fecundity and body size.␣The energy content of germinal and somatic tissues was determined by differential scanning calorimeter. The␣reproductive effort and a reproductive index based on the fifth spawning were evaluated to compare the␣patterns of energy allocation of a semicontinuous iteroparous species with semelparous and annual iteroparous species. The reproductive index ranged from 0.20 to 0.77, with an average of 0.480. The average reproductive effort was 0.840, ranging from 0.60 to 0.96. These figures highlight the enormous amount of energy that O. labronica allocates to reproductive tissues. Received: 23 July 1997 / Accepted: 20 October 1998     

Anchorage of embryos in Spirorbinae (Polychaeta)

Methods of incubation are taxonomically important in Spirorbinae. About half of the known species incubate in the operculum, but the remainder are usually described simply as incubating within the tube. Studies of tube incubation show that the embryos are anchored in various ways. They form an egg string, which is attached to the tube by a posterior filament in Spirorbis and Spirorbella; they adhere more generally to the tube wall in Circies and Paradexiospira; they are attached to specialised thoracic stalks in Protolaeospira, Helicosiphon and Romanchella; they adhere to the body surface in Metalaeospira and probably in Eulaeospira; they are not attached to either body or tube in Paralaeospira. Tube incubation is found predominantly in temperate or cold waters. In most species from the southern hemisphere, embryos are anchored to the bodies of the parents, but in most species from the northern hemisphere, embryos are fixed to the walls of the parent tubes.     

Functional morphology and feeding behavior of Scolelepis squamata (Polychaeta: Spionidae)

Functional morphology and feeding behavior of Scolelepis squamata (Müller) were studied. Gut contents consisted of unconsolidated sedimentary particles, fecal pellets of other species, and a wide variety of embryos, larvae, and juveniles. Unlike other spionid polychaetes the palps of S. squamata lack a median, ciliated groove. Particles captured by the palps were brought to the pharynx by a complete contraction of the palp. In the presence of a current, S. squamata helically coiled their palps, and fed almost exclusively on suspended and resuspended particles. In contrast to most spionid polychaetes the palps of S. squamata are not deciduous. Various aspects of the morphology and feeding behavior of S. squamata are discussed in terms of their functional and ecological significance.     

Zur sexuellen Differenzierung von Ophryotrocha puerilis (Polychaeta: Eunicidae)

Intact young males of Ophryotrocha puerilis Clap. Mecz. which are about to reach the female phase are capable of producing oocytes even under conditions of fasting. Intact isolated females may change sex under fasting conditions; their oocytes are not shed but resorbed. Decerebrated males which are about to reach the female phase at the moment of decerebration are unable to develop oocytes. Decerebrated females begin earlier with the production of spermatozoa than females under fasting conditions. The oocytes of decapitated females are either resorbed (young females with oocytes ranging in diameter up to 60 μm), or shed in egg masses capable of development after fertilization (older females with oocytes of 90 μm diameter or more). The effect of decerebration on females is compensated completely by implantation of isolated prostomia from female donors. Decerebrated males with implanted isolated prostomia from female donors immediately start oocyte production; these oocytes are not shed. In the majority of cases, decerebrated females change to the male phase after implantation of prostomia isolated from male donors. The mutual influence of a pair depends on direct contact of the partners. In isolated females, anterior fragments, consisting of prostomia and some segments, may induce change of sex and shedding of oocytes. On the other hand, intact females may induce sex reversal in anterior fragments. Even isolated prostomia from female donors may induce change of sex in intact females. A possible influence of intact females upon isolated prostomia was not found. Intact females and decerebrated females with implanted prostomia from female donors exhibit no mutual influence when placed together.     

Population structure and reproductive biology of Ceratonereis limnetica (Polychaeta: Nereididae) at Lower Portland,Hawkesbury River,Australia

Ceratonereis limnetica Hutchings and Glasby was collected from Lower Portland, Hawkesbury River, New South Wales, Australia, from March 1982 to September 1983. C. limnetica is one of a relatively few polychaete species in which the adults normally tolerate reverine conditions. Reproduction in this species has been modified accordingly, differing from the common pelagic spawning of many nereidids. Adults of both sexes remain essentially unmodified and spawn on the bottom. Embryos and larvae are brooded in specially constructed tubes in the sediment by the female. Both sexes have a life span of about 1 yr, dying soon after spawning. The morphological and reproductive behavioural similarities between C. limnetica and C. aequisetis are considered to indicate a close phylogenetic affinity.