Sex and reproductive aspects in Serranus cabrilla (Osteichthyes: Serranidae): macroscopic and histological approaches

The gonads of Serranus cabrilla (Linnaeus, 1758) from the Canary Islands were studied between September 1992 and July 1993. Macroscopic classification of the maturity stages was compared with histolog ical classification of gonad characteristics, and the effect␣of the classification method on the determination of the spawning season and size at maturity was investigated. Analysis of gonad organization and development confirmed that this species is synchronously hermaphroditic. Ovarian-tissue development is asynchronous, and testicular tissue consists of continuous spermatogonia. Six stages of oogenesis and five of spermatogenesis are described, based on differences in staining and in size and on differences in the nucleus and cytoplasm structure as viewed through a light microscope. Two types of atretic degeneration of eggs are also described. Agreement between macroscopic and histological staging was low, although both methods provided similar results in spawning-season determination and size at maturity. The spawning season of S. cabrilla is from February to July, with a peak in May. Fish size at first maturity (size at which 50% of all fish sampled are in relevant maturity stage) is 152 mm standard length (SL), and size at mass maturity (size at which 95% of all fish sampled are in relevant maturity stage) is 167 mm SL. Received: 21 May 1996 / Accepted: 9 August 1996     

Movements of albacore tuna (Thunnus alalunga) in the South Pacific: Evidence from parasites

Parasites were collected from over 400 albacore (Thunnus alalunga) caught by surface trolling and longlining in the south-west Pacific between 1985 and 1988. Parasites found included 1 apicomplexan, 3 nematode species, 4 cestode species, 1 acanthocephalan, 12 digenean species and 3 copepod species. Twelve of these parasite species which could be accurately recognised and counted were used in the subsequent analyses. Parasite data from albacore caught around New Zealand show a decrease in prevalence of three didymozoid parasites with increasing fish length up to a fork length of 70 to 79 cm. The subsequent increase in prevalence of these didymozoids in large longline-caught fish is consistent with fish returning from spawning in tropical waters where re-infection is presumed to occur. Albacore collected at widely separate locations in the south-west Pacific have differences in parasite prevalence, supporting an hypothesis that juvenile albacore move south to New Zealand from the tropics and do not return until the onset of sexual maturity. Albacore appear to move along the subtropical convergence zone, as indicated by a decline in prevalence and abundance ofAnisakis simplex andHepatoxylon trichiuri from New Zealand to the central South Pacific. This is supported by tagging and seasonal movements of the fishery.     

Between-area differences in age and length at first maturity of the orange roughy Hoplostethus atlanticus

Mean age and length at onset of maturity were estimated for orange roughy (Hoplostethus atlanticus) populations off New Zealand, Namibia, and Great Britain. Significant between-area differences were apparent in both these parameters. Implications of the between-area differences on stock structure of orange roughy in New Zealand waters are discussed. A proportional relationship between age at onset of maturity and modal size of fish in the mature population is demonstrated. This is consistent with later-maturing fish experiencing a longer period at a pre-maturity growth rate that is relatively faster than the rate of somatic growth after maturity. Received: 22 December 1997 / Accepted: 5 May 1998     

Reproductive biology of the commercial sea cucumber Holothuria fuscogilva in the Solomon Islands

 Reproduction of Holothuria fuscogilva (Selenka, 1867) in the Solomon Islands was investigated over a 4 yr period (1994 to 1998) by macroscopic and microscopic examination of the gonad tubules, the gonad index (GI) method, histological examination of gametogenesis, and spawning-induction trials. The gonad consisted of numerous tubules that dominated the coelom of gravid specimens. New tubules appeared in March, and grew in size and extent of branching until they reached their maximum size and maturity in August. Spawning occurred from August to October, with the majority of gametes released during October, although it was only partial in many individuals. After spawning, the tubules appeared wrinkled and resorbed into the gonad basis. A five-stage gonad maturity scale based on the macroscopic appearance of the gonad tubules corresponded with discrete stages of gametogenesis identified by histology. Gametogenesis was initiated in mid-March, with oogenesis and spermatogenesis occurring in parallel, followed by the growing stage (May to July) which was marked by active gamete development. Successful induction of spawning during the breeding period corroborated the GI and histological data. The uniform growth of gonad tubules indicated that H. fuscogilva in the Solomon Islands does not conform to the progressive tubule recruitment model described for other holothurians. An important application of this study is that the appearance of gonad tubules, removed by biopsy, can be used to determine the gonad condition of wild adults or captive broodstock. Received: 28 June 1999 / Accepted: 8 March 2000     

Reproduction in the diadematoid sea urchin Centrostephanus rodgersii in contrasting habitats along the coast of New South Wales, Australia

Reproduction in the sea urchin Centrostephanus rodgersii was examined in two types of habitats (“barrens”, i.e. habitats characterised by the high crustose coralline algal cover typical of urchin-barren grounds, and by the absence of macroalgae; and “fringe”, i.e. habitats characterised by a high macroalgal biomass and few C. rodgersii) at four locations in New South Wales. The four locations: the Solitary Islands, Sydney, Ulladulla and Eden, span the distribution of C.␣rodgersii from the subtropics at its northern limit to temperate waters near its southern limit. Histology and estimates of gonad retrieval rate (GRR) from January 1994 to October 1995 indicated that reproduction was synchronous at all locations. An increase in the tempo of gametogenesis in May and onset of spawning in June at all locations is consistent with entrainment in response to exogenous factors. Over the range studied, C. rodgersii experienced relatively similar daylength cycles and contrasting sea-temperature cycles. Short days and lunar conditions coinciding with the solstice appear likely proximate cues for the onset of spawning. The major difference in reproduction among locations was in the duration of spawning. In the southern parts of its range breeding occurred over a 5 to 6 mo period, whereas at the Solitary Islands it lasted ≃1 mo. At most locations the GRRs were significantly higher in the fringe habitat than in the barrens habitat. The lower reproductive output of urchins in the barrens habitat was attributed to the food-poor conditions typical of this habitat. The developing fishery for C. rodgersii is likely to be most effective from March to early May. Urchins from barrens areas may not provide sufficient yield to warrant harvesting. Received: 29 October 1997 / Accepted: 18 May 1998     

Mass spawning by the sea urchin Evechinus chloroticus (Echinodermata: Echinoidea) in a New Zealand fiord

A mass synchronous spawning of the sea urchin Evechinus chloroticus (Valenciennes) was observed in situ in Doubtful Sound, a large New Zealand fiord. Spawning occurred between 17:30 hrs and 18:30 hrs on 27 January 1994 and coincided with a full moon, spring tides and a period of decreasing sea temperatures. During spawning, the sea urchins formed a dense spawning aggregation of both sexes, with >90% of the urchins observed spawning at the time. Spawned gametes clouded the water column, and some were eaten by small labrid fish species. The spawning, which may have been as widespread as 40 km, marked a 42 to 50% decrease in gonad indices and resulted in a widespread, dense cohort of  E. chloroticus larvae within the fiord. Received: 25 September 1997 / Accepted: 6 March 1998     

Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic

Egg data from ichthyoplankton monitoring sites in the western English Channel (1988–2003) and northern Spain (1990–2000) and macroscopic maturity data from biological samples of purse seine landings in western and southern Iberia (1980–2004) are used to describe the spawning seasonality of sardine (Sardina pilchardus) in European waters of the northeast Atlantic using generalised additive models. The fitted models reveal a double peak in spawning activity during early summer and autumn in the western Channel, a wider spring peak off northern Spain and a broad winter season in the western and southern Iberian Peninsula. At all sites, a high probability of spawning activity was observed over at least 3 months of the year, with the duration of the season increasing with both decreasing latitude and increasing fish size. Off western and southern Iberia there are indications that the spawning season has been of longer duration in recent years for all size classes (reaching in some cases 8 months of the year for large fish). These patterns are in general agreement with existing literature and theoretical expectations of sardine spawning being driven locally by the seasonal cycle of water temperature, assuming preferences for spawning at 14 –15°C and avoidance for temperatures below 12°C and above 16°C. Regional quotient plots indicated that spawning tolerance to higher temperatures increases progressively with decreasing latitude. Despite the weak evidence for geographical differences in temperature tolerance that may have some genetic origin, the degree of spatio-temporal overlap in sardine-spawning activity within Atlantic European waters is unlikely to promote any reproductive isolation in that area.     

Do cod form spawning leks? Evidence from a Newfoundland spawning ground

Catch and acoustic telemetry data were used to test three predictions of lekking behaviour in Atlantic cod (Gadus morhua L.) at a small-scale spawning ground (∼25 km²) in Placentia Bay, Newfoundland: (1) that cod form male-skewed aggregations during spawning; (2) spawning arenas occur at consistent sites; and (3) on the arenas, there will be higher proportions of spawning fish. Forty-two fishing sets from 1998 to 2003 indicated male-skewed sets early in the spawning season (April) in 5 of 6 survey years. Male-skewed sets were consistently distributed at depths < 50 m and mostly in one part of the ground. The proportions of spawning females and spent males were significantly higher in male-skewed sets, whereas sets with equal sex ratios had significantly higher proportions of immature males and spent females. In addition, cod of both sexes were significantly larger in male-skewed sets. Telemetric tracking of 25 cod in 2002 and 2003 (12 males, 13 females) indicated that both males and females were highly mobile while present on the spawning ground. Overall, our results add support to the hypothesis that cod employ a lekking mating system.     

Maturity classification and group maturity of the red sea bream Pagrus major

The criteria of ovarian maturity during the spawning season and maturity rates according to age are described for the female red sea bream Pagrus major collected from the Chikuzenkai between April and June, 1984. The degree of maturity of the ovary is categorized into four stages based on the maturation stage of the most advanced oocytes in the ovary, rate of atresia and occurrence of post-ovulatory follicles. The ovarian maturity rates in fish aged 2, 3, 4 and 5 years were shown to be 5.8, 46.2, 75.0 and 97.3%, respectively, assuming that yolked ovaries with low levels of atresia (<50% yolked oocytes affected) attain maturity. The fork length at which 50% of the fish attained maturity ranged from 280 to 300 mm.     

Reproductive biology of the holothurians from the major communities of the New Caledonian Lagoon

The reproductive biology of nine species of aspidochirote holothurians from the major communities of the New Caledonian lagoon has been analysed. The data were obtained from monthly sampling from 1978 to 1982 of their populations on reef slopes and passes for Holothuria nobilis, H. fuscogilva, H. fuscopunctata and Thelenota ananas, in the inner lagoon for Stichopus variegatus, H. scabra and H. scabra versicolor and on reef flats for Actinopyga mauritiana, A. echinites and H. atra. The morphology and anatomy of the gonads and their annual cycle are described. The reproductive cycles, composed of the five phases of gonad growth, maturing, spawning, post-spawning and resting, were determined from the changes in the sexual stages, the gonad indices and the percentage of indeterminate sex. Three main reproductive patterns are recognized, annual cycles with spawning occurring during the warm season or during the cool season and a semiannual cycle. Other population parameters, such as the size of the mature oocytes, the mean size of the individuals at first sexual maturity, and absolute and relative fecundities were calculated and related to the mean size of the species, the taxonomy and the ecology. The reproductive strategies of the species show a gradient that is probably related to the stability of the biotopes, with smaller species living in the more intrable outer reef flats and larger species in deeper lagoon floors.     

Validation of age determination from otoliths of the King George whiting Sillaginodes punctata (Perciformes)

The otoliths of the King George whiting Sillaginodes punctata were assessed for their usefulness in adult-ageing. Whole, transverse-sectioned and broken/burnt sagittae displayed alternating opaque and translucent zones. Some small whole otoliths contained accessory areas of pigment, whilst large otoliths were too thick to transmit light, resulting in ambiguous counts in each case. Both marginal-increment analysis and treatment of fish with tetracycline demonstrated that the first three opaque zones formed annually, the first during the spring/summer of the second year of life, and the second and third in each subsequent spring. Otolith thickness increased linearly with the completion of each new opaque/translucent zone. An ageing protocol based on whole otoliths, supplemented with the breaking/burning of ambiguous otoliths is recommended. An algorithm for calculating age in months from otolith counts based on a fixed birth-date of the mid-point of the spawning season is presented. Received: 18 August 1997 / Accepted: 15 October 1997     

Reproduction and growth of Isostichopus fuscus (Echinodermata: Holothuroidea) in the southern Gulf of California, México

The reproduction and growth of Isostichopus fuscus (Ludwig, 1875) at Isla Espíritu Santo, Baja California Sur, México from July 1992 to September 1993 and from August 1996 to July 1997 were analyzed. The reproductive cycle was monitored by using histological analysis. Five gonadal stages were described according to cell types present in the gonad; undetermined, gametogenesis, maturity, spawning and post-spawning. This holothurian reproduces annually during summer (July to September), when sea surface temperature reaches 27 °C. The gonad index is related to the gonadal stages. The first sexual maturity for I. fuscus was calculated at 367 g (4 to 5 years old) according to the von Bertalanffy equation, and nearly all the specimens at Isla Espíritu Santo were 5 to 7 years old (including some individuals over 35 years old). Data of the length–weight relationship showed that I. fuscus grows allometrically at the studied site (b = 1.83). The mortality rate was relatively high (M = 0.51), indicating that in a typical cohort, 40% of the recruits does not survive the first year, and less than 20% reaches maturity. Received: 15 July 1998 / Accepted: 19 August 1999     

Growth and ovarian development of Maurolicus muelleri during spring

The size, fecundity and gonad development of Maurplicus muelleri (Gmelin) were studied in Herdlefjorden, western Norway, from January to June 1994. In January, the two age groups formed separate sound-scattering layers (SSL). Juvenile 1-group fish were found to form the upper SSL, while adult 2+ individuals inhabited the lower, these distributions reflecting the different feeding to fitness functions of these two age groups. In June there was only one SSL, and in March and May some 2+ were also found in the upper SSL. Growth of the younger fish to 2+ fish size was observed over the period, and it was found that M. muelleri mature after 1 yr, although size at maturity was lower than found in previous studies. Growth of the 2+ fish changed as the period progressed, to allow gonadal development after a winter of low feeding. Increased investment in development of the gonads was also observed in the 1-group fish over the study period, but substantial maturation only occurred from May to June. Absolute fecundity was size dependent, although high variability was found in the larger size ranges. Oocyte development occurred in discrete batches, although lack of gonadal synchrony between individuals means that spawning is continuous in the population as a whole, once the spawning period begins. Batch fecundity was found to be similar to previous studies, and 2+ fish were found to start spawning earlier, and with an increased intensity, than the 1-group fish.     

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.     

Social inhibition of maturation in females of the temperate wrasse Pseudolabrus celidotus and a comparison with the blennioid Tripterygion varium

In two temperate reef fish, Pseudolabrus celidotus (Bloch and Schneider) and Tripterygion varium (Bloch and Schneider) studied near Leigh, New Zealand, most spawning activity was concentrated during the first 2 months of spawning seasons which lasted about 5 months. In P. celidotus, maturation and spawning of first-year females (0+) was delayed with respect to older (1+yr) females. Within the 0+ age class, the minimum size of maturation declined as the spawning season proceeded. However, in all females of T. varium, ripening and spawning took place over the same time ranges. Observations on the seasonal patterns of female aggression suggested that larger female P. celidotus were socially inhibiting the maturation of small females. The level of interaction was high compared to that in T. varium, and reached a peak over the onset of the spawning season. This hypothesis was tested by the removal of larger (1+yr) female P. celidotus from a field population. The remaining 0+ females increased in ovary weight and matured earlier than those in an undisturbed area. We argue from this experiment that it is of advantage for P. celidotus females to spawn early in the season and of advantage to inhibit other females from doing so. Two possible reasons are suggested, the first applying to sexchanging species such as P. celidotus, the second to temperate-water fishes in general.     

The dual functions of sea urchin gonads are reflected in the temporal variations of their biochemistry

Fatty acid analyses are emerging as a powerful technique to probe trophic interactions between organisms. In this paper, the application of both this procedure and gonad index (GI) determination on two populations (intertidal and subtidal) of the echinoid Psammechinus miliaris is reported. The investigation spanned the 3-month spawning period of Scottish west coast populations. In both populations a progressive decrease in the GI was found, coupled with an increasing maturity stage (from mature to spent). Sexual maturation and decrease in GI was synchronous between the two populations. In conjunction, there were distinct changes in gonad biochemistry. Differences in the fatty acid composition of the gonad reflected the changes in sexual maturation. Mature males and females had significant differences in the fatty acid composition of their gonads, whereas post-spawned individuals showed no gender differences. Male urchins had higher levels of polyunsaturated fatty acids (PUFAs) compared to females, and there was a dramatic reduction in the fatty acids 22:6(n−3) and 20:5(n−3) with increasing maturity stage. Using multivariate statistical techniques, these changes in the fatty acid composition of the sea urchin gonad were linked to habitat related diet differences combined with gender differences. These changes in the fatty acid signatures clearly reflect the dual function of the gonad as both a nutrient store and a reproductive organ.     

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     

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.     

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

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

Reproductive cycle and gametogenesis in the black abalone Haliotis cracheroidii (Gastropoda: Prosobranchiata)

Seasonal changes in gonald size and stages of gametogenesis in the black abalone Haliotis cracheroidii were related to changes in environmental parameters. H. cracheroidii showed an annual reproductive cycle terminating in a synchronized spawning in late summer. Gametogenesis was initiated immediately after spawning. Gametes were present in the gonad through the winter months. Gametogenesis was initiated a second time in the spring months. Maximal gonad growth (to a gonad index of 20%) occurred during summer months prior to spawning. Changes in gonad size andperiods of initiation of gametogenesis revealed no apparent correlation with changes in seasonal water temperature. Changes in gonad size showed no apparent relation to change in day length. Total polysaccharide levels in foot tissue changed seasonally, indicating that food availability is probably not a factor in directly regulating gonad growth. Gonad index data for the chiton Katharina tunicata (collected over a 10 year period) showed no apparent correlation to seasonal change in water temperature.