A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

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.     

Age structure and spawning of a Northumberland population of Melinna cristata (Polychaeta: Ampharetidae)

A Northumberland (England) population of Melinna cristata was studied from October, 1967 to February, 1970. M. cristata is a gregarious sedentary polychaete which lives in muddy substrates. It is dioecious, and has a very restricted breeding season. The gametes are shed solely by the two pairs of posterior nephridia, which undergo morphological changes just before the breeding season; this allows them to accommodate the mature gametes. After spawning, these nephridia ternal and, within 2 to 3 weeks of spawning, the benthic larvae metamorphose into juveniles. The juveniles grow from 3 to 10 mm during their first year, and can be distinguished from the rest of the population by their size and the complete absence of coelomic gametes. After about 12 months, the juveniles release their first batch of gametes into the coelom from the germinal epithelium, and they then cannot be distinguished from the rest of the population. The development of coelomic gametes takes 7 to 10 months to complete, so M. cristata is potentially capable of breeding for the first time when 2 years old. The majority of worms survive spawning, and M. cristata probably breeds annually for several years. In this population, not all potential breeders spawn, some resorb their mature gametes and release another batch of gametes into the coelom. The Northumberland population of M. cristata is near the southernmost limit of the species distribution, which suggests that environmental conditions for this population are not optimum. The population appears to maintain itself by producing fewer oocytes and by only part of the population spawning.     

Size-related reproductive biology of the infaunal bivalve<Emphasis Type="Italic"> Abra alba</Emphasis> (Bivalvia) in Kinsale Harbour (south coast of Ireland)

The reproductive behaviour of 248 specimens comprising four size classes of the infaunal bivalve Abra alba (Wood) was investigated between September 1994 and November 1995 in Kinsale Harbour. Small males returned a very small proportion of sexually mature and spawning animals. By contrast, no spawning females were found in this size class, suggesting that males develop and mature at a smaller size than females. Mean gonad maturity indices indicated that, except for the smallest size class, A. alba exhibits two spawning events in 1 year, i.e. a lesser spring and a major autumn spawning event. The spring event was well synchronised between males and females, while the autumn spawning was more broadly synchronised between the sexes. Only one spawning event was recorded for very large males and females, also occurring in spring. Spawning stages were recorded for both sexes over all size classes between February and November, indicating only a short period of sexual repose, if any. Oocyte diameters did not show a discrete size range from developing to spawning stages. All stages were seen to contain varying proportions of smaller, less mature oocytes, suggesting that large A. alba females do not develop and spawn en bloc, but release their gametes periodically. Maturity indices showed only a broad similarity with off-bottom temperatures.Communicated by O. Kinne, Oldendorf/Luhe     

Reproductive biology of the antipatharian black coral Antipathes fiordensis in Doubtful Sound, Fiordland, New Zealand

The reproductive biology of Antipathes fiordensis Grange, a species endemic to south-western New Zealand, was followed from April 1994 to May 1995. Ten colonies were individually tagged in Doubtful Sound and sampled on a monthly basis in order to determine their reproductive activity. The fecundity of each of the five tagged female colonies was determined by estimating the total number of polyps per colony from photographs of each colony and by planimetry, the proportion of gravid polyps per colony, and the mean number of oocytes per gravid polyp. In addition, 56 colonies were sampled in March 1995 to estimate the sex ratio, height at sexual maturity, and mean sizes of females and males. A. fiordensis was found to be a dioecious species which, in the absence of gonads in the polyps, has no obvious external morphological differences between the sexes. Broadcast spawning of gametes is the likely mode of reproduction. Gametogenesis began in November 1994 and was highly synchronous within and between colonies, with spawning occurring in March 1995. The sex ratio in adults was 1:1. Colonies reached sexual maturity between the heights of 70 and 105 cm which, based on existing estimates of growth rate, corresponds to a minimum age for sexual maturity of about 31 yr. The largest oocytes measured ranged from 100 to 140 μm in size. Female colonies produced between 1.3 and 16.9 million oocytes, with the larger colonies dominating the reproductive output of the population. Received: 23 June 1997 / Accepted: 1 August 1997     

Histology of the gonads and observations on the social behavior of the Caribbean angelfish Holacanthus tricolor

Male and female specimens of Holacanthus tricolor were collected in May 1981 and September 1982 off the Island of St. Croix. U.S. Virgin Islands, and their gonads were examined histologically. All testes examined showed vasa deferentia which were characteristic of secondary males, suggesting protogynous sex change. A transitional individual supported this interpretation. Ovarian histology and size-specific egg counts revealed seasonal differences in spawning frequency and clutch size. Females appeared to spawn every day in May. The timing of final maturation and hydration of the oocytes indicated that spawning occurred in the late afternoon or early evening. Not all females spawned every day in September, and clutch sizes were smaller than those of May. Behavioral observations of H. tricolor in the field revealed that males defended harems of two to four smaller females. Within each harem, females of different sizes had overlapping home ranges, while the home ranges of females of similar size did not overlap. Besides size dimorphism between the sexes, H. tricolor exhibited a previously unreported sexual dichromatism.     

Reproductive pattern in the brooding and broadcasting sea star Pteraster militaris

Pteraster militaris (O. F. Müller, 1776) broods young in an aboral chamber and broadcasts gametes and offspring as well. The reproductive cycle of a population of P. militaris from the northwest Atlantic Ocean was examined over a 23 mo period from September 1985 to July 1987. The species is gonochoric (sex ratio 1:1), with only one incidence of hermaphroditism observed (n=261). Gonad indices were lower in females than in males, possibly an adaptation to the limited space available in the brood chamber. Body component and maturity indices, gonad histology and response to 1-methyladenine indicate that females in the population have a continuous breeding season, with a slight decrease in intensity in the spring. Males have a more pronounced decrease in the intensity of reproduction in the spring. Gonad development in males in synchronous both within and between individuals. Gonad development in females is asynchronous.     

Reproduction of the commercial sea cucumber Holothuria scabra (Echinodermata: Holothuroidea) in the Solomon Islands

Over a 3-year period (1996-1998), reproduction of the commercial sea cucumber Holothuria scabra (Jaeger, 1833) was investigated in the Solomon Islands to determine the spawning pattern and whether gametogenesis is continuous or seasonal. The gonad consisted of a single cohort of tubules that developed uniformly. Macroscopic examination of the gonads revealed that mature gametes were present throughout the year. Individuals with gonads at different stages of maturity were present in most samples. Partly spawned gonads were prevalent in females, whereas mature gonads were prevalent in males. The time at which the peak gonad index was recorded differed among years. Although gametogenesis was continuous, with a potential for prolonged gamete release, a period of enhanced spawning occurred during the dry season, from September to December. Maximum gonad indices were reached prior to and during this period of enhanced spawning. Histology revealed that gametogenesis reinitiated in partly spawned gonads, resulting in the presence of gametes at different stages of development in the gonad. The uniform growth of gonad tubules indicated that H. scabra does not conform to the progressive tubule recruitment model described for holothurian oogenesis. Continuous reproduction in H. scabra and prolonged availability of mature gametes would facilitate use of this species for aquaculture.     

Parental care and the cost of reproduction in a Mediterranean fish

Summary Symphodus tinca is a common near-shore Mediterranean labrid fish in which females may sometimes spawn their eggs over hundreds of square meters, or alternatively spawn into well-defined algal nests. Eggs spawned in either manner are fertilized, but widely scattered eggs receive no parental care, whereas eggs spawned into nests are usually guarded by the male until they hatch. Here, I report weight changes of individual marked fish that engaged in a variety of different reproductive behaviors during three breeding seasons. Males gained weight at 0.15% per day outside the spawning season, and added 29–78% to their overall body weight between reproductive seasons, even following substantive weight losses in a spawning season (up to 20% among nesting males). Nesting and nest-guarding males lost an average of 0.32% and 0.41% of their body weight per day in 1986 and 1987. This cost is four times greater than reproduction for nonnesting males, which registered a 0.03% daily weight gain. Actively spawning females lost 0.06% of their body weight daily during the spawning season. While long-term growth rates did not appear to be substantially affected by reproduction in either sex or by parental care in males, present work does not exclude the possibility that such long-term effects may exist.     

Reproductive cycle of Centrostephanus rodgersii (Echinoidea), with recommendations for the establishment of a sea urchin fishery in New South Wales

The reproductive cycle of the sea urchin Centrostephanus rodgersii (Agassiz) was investigated in two populations, at Clovelly and Little Bay, in Sydney, New South Wales, Australia. C. rodgersii were collected at monthly intervals from February 1992 through January 1993. The reproductive cycle was determined by histological examination of oogenesis and spermatogenesis, monthly measurements of gonad index (GI), and induction of spawning by KCl injection. C. rodgersii has an annual reproductive cycle that was highly synchronous in both populations. From February to June, gametogenesis was accompanied by a decline in the amount of nutritive tissue in the gonads. The urchins were mature from June to September, with peak spawning between July and August, as indicated by a significant drop in GI. The breeding season of C. rodgersii therefore coincides with the lowest sea temperatures and the shortest days of the year. The gonads returned to the recovering condition within a month of spawning, with a substantial thickening of the nutritive layer along the gonad wall, and the GI returned to near pre-spawning levels. As a result, the spent phase was rarely found in C. rodgersii. With the exception of a significant decrease in the GI following spawning of urchins from the Clovelly population, the GI measurements did not show any distinct pattern through time. Specimens spawned in response to KCl injection from mid-May to early October, with the maximum response in July. Although all individuals sampled were at a similar stage of maturity at any one time, inter-site differences were seen with all of the methods used. Gonad indices from Little Bay were consistently higher and less variable than those from Clovelly for most of the year. The Little Bay population could also be induced to spawn for a longer period of time than could the Clovelly population. The breeding season of the Little Bay population appears to be longer than that of the Clovelly population. The relationship between size and sexual maturity was also examined. All C. rodgersii with a test diameter of >60 mm could be induced to spawn and produced viable gametes. C. rodgersii has been nominated for commercial exploitation in New South Wales, and the results of this investigation are used to make recommendations on the timing and size limits for a fishery utilising this species.     

Reproductive biology of the spiny lobster, <Emphasis Type="Italic">Panulirus Penicillatus</Emphasis>, in the southeastern coastal waters off Taiwan

The reproductive biology of spiny lobster, Panulirus penicillatus, was studied based on 2,068 lobsters, ranging from 34.28 to 131.60 mm carapace length (CL), sampled in Taitung coastal waters from September 2003 to December 2004. The overall sex ratio approximated 1:1 (χ² = 0.02, P > 0.05), but the monthly sex ratios in 2004 showed significant differences and males were predominant in sizes larger than 80 mm CL. Reproductive activity, assessed using histology, a gonadosomatic index and percentage of ovigerous females, indicated that the mature females could be found in every month and that the major spawning occurred from May to September. The presence of re-developing/re-ripe ovaries by month and size-specific spawning time suggest that larger mature females (>60 mm CL) spawn at least three times a year while smaller new mature females spawn at least once a year. For females, the estimated sizes at 50% physiological and functional maturity were (mean ± SE) 56.46 ± 0.56 mm CL and 66.63 ± 1.07 mm CL. The estimated sizes at functional maturity were between 72 and 74 mm CL for males. The number of eggs per spawning event (brood size, BS) was related to CL by the equation Y _BS = 2.4 × 10^-3CL^4.18 (r ² = 0.902, n = 12). Female lobsters with CL ranging from 60 to 80 mm made the greatest contributions to egg production because of their high brood size and active reproductive activity. A minimum legal size should be established for the fishery to protect egg production potential of lobster population in the southeastern coastal waters off Taiwan.     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Seasonality of coral reproduction in the Dampier Archipelago,northern Western Australia

Coral spawning in Western Australia (WA) occurs predominantly in the austral autumn in contrast to the Great Barrier Reef (GBR) on Australia’s east coast where most spawning occurs in spring. Recent work, however, suggests a second spawning period in northern WA with at least 16 Acropora spp spawning in spring or early summer. This discovery has initiated a re-examination of reproductive seasonality in northern WA, particularly on inshore reefs adjacent to large development projects, such as the site of this study in Mermaid Sound, in the Dampier Archipelago. Three locally abundant taxa, Porites spp, Pavona decussata and Turbinaria mesenterina were sampled monthly from September 2006 to May 2007 to determine sexuality, the mode of reproduction and the time of gamete maturity. All three taxa were gonochoric broadcast spawners. Porites spp. colonies were mature in November and December, P. decussata in March and April. In contrast, most colonies of T. mesenterina contained mature gametes for up to 5 months beginning in November, suggesting either individuals are releasing gametes on multiple occasions, or they retain mature gametes for more than 1 month. Field surveys to determine the reproductive status of the remaining coral assemblage were conducted prior to the full moon in October 2006 and March 2007. Only four species contained mature gametes in October 2006. In contrast, 55 species contained mature gametes in March 2007. We conclude that the major spawning season of corals on shallow-inshore reefs in the Dampier Archipelago is autumn, although taxa that spawn in spring and summer include Porites spp., Acropora spp. and possibly T. mesenterina that are numerically dominant at many of these sites. Consequently, management initiatives to limit the exposure of coral spawn to stressors associated with coastal development may be required in up to five months per year.     

Courtship and spawning in the emperor angelfish Pomacanthus imperator,with comments on reproduction by other pomacanthid fishes

Courtship and spawning are described for Pomacanthus imperator and 4 Centropyge spp. at Enewetak Atoll, Marshall Islands, observed from 15 August to 15 September, 1981. In all species, courtship and spawning occur at dusk, continue throughout most of the lunar month, are preceded by male display to the female, and culminate in the pair's shedding gametes into the water column. The species differ in ascent height, details of courtship behavior, spawning location relative to prominent reef topography and degree and type of sexual dichromatism. Social organization in all species appears based on male defense of harems of 2 to 5 females. Observed behavior is compared to that of confamilials, and discussed relative to proposed theories concerning lunar periodicity of spawning activity, sexual dichromatism and the adaptive significance of spawning ascent behavior.     

Evidence for broadcast spawning as well as brooding in the scleractinian coral Pocillopora damicornis

The reproductive activity of 88 colonies of Pocillopora damicornis at Rottnest Island, Western Australia, was examined over 12 mo (20 August 1988 to 20 August 1989). Larvae were found close to the three new moons of February, March and April, 1989, but in large numbers only during March; 67% of all corals produced larvae. There was no obvious connection between gametogenesis and the production of brooded larvae; however, those corals which did not produce, larvae also did not produce ova but did produce, sperm, and must be considered male colonies. All colonies which produced ova went on to produce planulae. Spermatogenesis occurred throughout the year, but oogenesis occurred between September 1988 and April 1989. Dramatic declines in the percentage of polyps containing eggs and sperm occurred in early February and early April 1989. On these dates the amount of lipid in the tissue of the corals decreased, which suggests that the decline in the percentage of polyps with sperm and eggs was not due to the gametes being resorbed by the polyps. Therefore, P. damicornis at Rottnest Island appears to be both a brooder and a broadcaster with asexual reproduction through brooded larvae and sexual reproduction through spawning of gametes. This accounts for earlier evidence in the literature that some input from sexual reproduction had been occurring in the populations of P. damicornis at Rottnest Island. There is no direct evidence for spawning, as broadcasting of gametes has not been observed, but the indirect evidence provides a strong argument for the existence of this dual reproduction.     

Evidence for multiple spawning in the tropical oceanic squid Stenoteuthis oualaniensis (Teuthoidea: Ommastrephidae)

Indirect evidence suggests that Stenoteuthis oualaniensis is a multiple spawner, a reproductive strategy not reported previously for squid. After spawning once, S. oualaniensis apparently continues to feed, grow, and mature additional oocytes before spawning again. Squid were caught by jigging at night in waters around the main Hawaiian Islands during four cruises from 1981 to 1987. All females from 200 mm mantle length (ML) to at least 318 mm ML were sexually mature, indicating that considerable growth takes place after reaching sexual maturity. Size-frequency analyses of ovarian oocytes revealed continuous egg production throughout adult life. Mature ova are stored in the oviducts, but oviduct fullness is unrelated to body size, indicating repeat spawning.     

The gametogenic cycle of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Mollusca: Cephalopoda) as observed on the Atlantic coast of Andalusia (south of Spain)

The present study reports the gametogenic cycle of the common octopus, Octopus vulgaris Cuvier, 1797 in the Atlantic coast of Andalusia between May 2001 and May 2002. A total of 673 specimens of total weight ranging from 30 to 3,732 g were analysed. The gonad development was defined by means of histological methods and gonadosomatic indexes. Four gametogenic stages were identified in males (I immature, II maturing, III mature, IV spawning) and in females (I immature, II maturing, III pre-spawning, IV spawning). The sex ratio obtained was not significantly different from 1:1 (P>0.01). Mature males were found during the whole year, September being the month when the highest rate of sexually active individuals occurred (48%). In females, the gametogenic cycle commenced in December and ended in November, and spawning occurred between April and October, with two maximum peaks in August and September (48% in each). Size at sexual maturity was 850 g of total body weight for males and 1,250 g of total body weight for females. Males mature earlier than females.Communicated by S.A. Poulet, Roscoff     

Multiple spawning in the tropical squid Photololigo sp.: what is the cost in somatic growth?

This study assesses the potential of the tropical loliginid squid Photololigo sp. to lay multiple batches of eggs and examines changes in somatic growth during reproduction. Histological analysis of the ovary and the relative size of the oviduct to mantle weight and ovary weight were used to determine the potential for multiple spawning. Ovaries of mature females always had immature and mature oocytes present, suggesting that not all the oocytes were maturing simultaneously and that multiple batches of eggs were being produced. Furthermore, poor correlations of oviduct weight with body size and ovary weight indicated that mature oocytes were not accumulating in the oviduct for a single spawning event. Both these observations supported the hypothesis that Photololigo sp. has the potential to lay multiple batches of eggs throughout its life. Specific growth rates, length-weight relationships, relative growth of somatic and reproductive tissue and microscopic assessment of muscle tissue were compared between immature and mature females. Growth rates of immature females were almost twice as fast as those of mature females. Mature females also had no large muscle fibres present, suggesting that energy for reproduction was mobilised from the muscle tissue.     

Population biology and larval development of the terebellid polychaete Nicolea zostericola

A 3-year study was conducted on a population of Nicolea zostericola (Oersted: Grube, 1860) at Nahant, Massachusetts (USA), which indicated that this species has a life span of about 1 year. Spawing occurred from mid-April to early July, after a rise in sea-water temperature to about 6°C. Pair formation occurred between sexually mature adults prior to and during spawning. Eggs were deposited in cocoons, where development occurred in 14 to 22 days. Larval develomment is described along with juvenile and adult substrate preferences and behavior. The entire intertidal population disappears following the spawning season in late June, and reappears in September as a result of migration of worms from the sub-littoral zone. Oogenesis consists of 3 phases: a short gonadal and follicular phase, followed by a longer vitellogenic phase during which oocytes are free-floating in the coelomic fluid.Contribution no. 31 from the Marine Science Institute, Northeastern University, Nahant, Massachusetts, USA.     

Reproductive biology of the lanternfish Benthosema pterotum from the Indian Ocean

Samples of Benthosema pterotum (Alcock) were obtained from the Arabian Sea (1976, 1979, 1981, 1983, 1984), off Mozambique (1978) and from the Bay of Bengal (1979). The sexual maturity stages are described. For females, the size of the largest oocytes was found to be most useful for size of the largest oocytes was found to be most useful for assessing maturity. Mature and spawning fish were found in all seasons. Samples from November 1983 differed from all others in having very few mature females. A few apparently spent fish were observed in July to August 1979 and November 1983. Some females mature at 25 mm and males at 20 mm, whereas others can still be immature at 45 and 40 mm, respectively. Of the total number of stations, four had significantly more males than females, twelve significantly more females, and forty-seven displayed no difference in the sex ratios. The vertical migration of the mature and spawning fish did not differ significantly from that of other individuals. Batch fecundity ranged from ca. 200 to 3 000 eggs fish ranging from 27 to 52 mm, relative fecundity from ca. 2 000 to 7 000 eggs g^-1 maternal dry weight. There are indications that B. pterotum spawns only once, but no firm conclusions could be drawn.