Life-history biology of Peniagone azorica and P. diaphana (Echinodermata: Holothurioidea) from the north-east Atlantic Ocean

Aspects of the reproduction and population biology of two elasipodid holothurians collected during the period 1973–1984 from the north-east Atlantic Ocean were examined. The reproductive biology of both the benthic species Peniagone azorica and the benthopelagic P. diaphana are similar. A primary oocyte forms from an oogonium and grows to about 100m diameter, whereupon it undergoes vitellogenesis and increases to 300m before being spawned. A high proportion of primary oocytes are not spawned and undergo a complicated breakdown process resulting in the formation of an amorphous sac in the ovary wall. It is possible that when an ovarian tubule is full of these sacs it atrophies and drops off, allowing other tubules to develop. The maximum egg size for both species suggests abbreviated larval development. In P. azorica, juveniles reach sexual maturity for the first time at about 30 mm length. The population structure suggests that the adults grow slowly, although it is possible that the observed unimodal distribution in size-frequencies results from one or several recruitments. Recruitment to the population is probably infrequent and may occur irregularly. Comparison of the population structure of closely spaced samples suggests a patchy distribution on the bottom. The benthopelagic lifestyle of adult P. diaphana suggests that the larvais also planktonic.     

Reproductive cycle of two populations of Ophionereis schayeri (Ophiuroidea) in New South Wales

Reproduction of two populations of Ophionereis schayeri (Müller and Troschel) in New South Wales was investigated from November 1991 through January 1993. The reproductive cycle was documented by histological examination of the gonads and by image analysis to determine the oocyte size-frequency distributions. Both populations of O. schayeri were mature in December and January and had a major summer spawning period between January and February. Thereafter, the condition of the gonads exhibited interindividual variability, with asynchronous low-intensity gamete release through August. O. schayeri has the potential to spawn for up to 8 mo of the year. This breeding pattern with synchronous spawning in summer and asynchronous gamete release through winter was similar in both populations. During autumn and winter, the gonads contained developing gametes and advanced gametes ready for spawning. The eggs spawned by O. schayeri during these seasons started their growth during the previous spring vitellogenic period, and continued to grow through summer. Upon reaching full size, they were stored for spawning outside the main breeding period. Spring is marked by increased spermatogenic and vitellogenic activity followed by maturation and spawning. Throughout their development, the oocytes of O. schayeri are surrounded by a follicle consisting of periodic acid-Schiff-positive (PAS+) haemal fluid and PAS+ yolk by the oocytes. In the testes, the haemal sinus projects into the centre of the spermatocyte columns, thus appearing to play a nutritive role in support of both oogenesis and spermatogenesis. Late vitellogenic oocytes of O. schayeri are firmly anchored to the germinal epithelium by an attachment complex consisting of specialised attachment cells and basophilic strands that radiate between the oolemma and the nucleus. Spawning is associated with rupture of the follicles, which remain as prominent, empty U-shaped profiles in the ovaries. O. schayeri produces copious numbers of 200 m-diam oocytes, suggesting that this species is a broadcast spawner and that it has a modified ophiopluteus or vitellaria larva.     

Reproductive cycle of Golfingia margaritacea,a bipolar sipunculan,in subantarctic water

The reproductive cycle of the bipolar sipunculan Golfingia margaritacea collected from Ushuaia, Argentina (54°49 S; 68°13W) in 1986 and 1987 is reported. The population sampled showed a sex ratio of 1:1. Males, however, frequently outnumbered females. Measurements of coelomic oocytes delineated an annual reproductive cycle. Small oocytes were present in the coelom all year: a subpopulation began to increase size about the middle of spring and reached its maximum size in late summer. Spawns may occur from the end February to April. May is characterized by resorption of large oocytes. Monthy monitoring of the gonads revealed mitotic activity and a continuous release of gametocytes to the coelom. Parasites were detected in the gonads. Large coelomic oocytes were also present in the coelom during the same season in the Northern and Southern Hemispheres, but egg sizes were significantly larger in boreal populations.     

Reproduction of the commercial sea cucumber Holothuria whitmaei [Holothuroidea: Aspidochirotida] in the Indian and Pacific Ocean regions of Australia

Seasonal trends in the gonad index (GI) of two widely separated populations of black teatfish, Holothuria whitmaei (formerly included in Holothuria nobilis), were investigated between Pacific (Great Barrier Reef) and Indian Ocean (Ningaloo Reef) coral reefs of Australia. Reproductive activity followed a similar annual trend, with the GI of both populations peaking typically between April and June. Macroscopic and histological analysis of Ningaloo Reef specimens revealed that large germinal tubules, positioned centrally on the gonad basis, progressed through four maturity stages: growing (II); mature (III); partly spawned (IV); and spent (V). Growing tubules dominated the central region of the gonad basis between January and March, followed by an increase in the number of mature tubules throughout the GI peak (April–June). The progressive appearance of partly spawned and spent tubules between June and October suggests that spawning in H. whitmaei continues intermittently over an extended period throughout the austral winter. The examination of the gonad structure of sexually mature male and female specimens identified five tubule size classes in total (C₁ to C₅), each of differing physical and gametogenic status. In females, smaller C₁ tubules located at the anterior edge of the gonad basis contained pre- to early vitellogenic oocytes. Larger C₂ and C₃ tubule cohorts, positioned centrally on the gonad basis, contained mid- to late-stage vitellogenic oocytes. Smaller C₄ and C₅ tubules, located at the posterior edge, contained only relict oocytes. Similar physical and gametogenic differences were apparent between tubule cohorts in male specimens. We propose that these results, together with evidence of incomplete gonad resorption over the austral summer, indicate that gonad development in H. whitmaei conforms to the predictions of the Tubule Recruitment Model (TRM). The TRM is reported rarely among tropical aspidochirotes, and results presented here (1) provide the first direct evidence of this model in H. whitmaei, and (2) confirm that this species is one of the few winter-spawning tropical invertebrates.     

Reproduction of the Antarctic fish Nototheniops nudifrons

The reproductive biology of the Antarctic fish Nototheniops nudifrons (Lönnberg, 1905) was analyzed by examination of the gonads of fish collected in March and April 1985 in trawls near Low Island, Antarctic Peninsula, and compared to direct observations of reproduction and early larval development in the laboratory from March to October of the same year. Males and females reached sexual maturity at an age of 4 to 5 yr. Mature males differed in coloration from females and immature males. Ovaries of sexually mature females contained two distinct size modes of vitellogenic oocytes, representing two separate clutches of developing eggs. Females spawned 100 to 3 500 demersal eggs, which were laid in a nest in crevices or under rocks, and guarded by the male for about 4 mo. Females did not assist in nest defense or egg care. Most spawning in the field and in the laboratory occurred in late fall and early winter (May to June). A second clutch may be spawned in spring (November and December). Eggs hatched after 124 d, and larvae were raised for 38 d. Otoliths of larvae contain internal microincrements, which are deposited in a daily fashion, and are visible by light microscopy. The otoliths of 32 adult fish were examined by scanning electron microscope, and counts of microincrements in these otoliths allowed the backcalculation of hatching dates. Estimated hatching dates were between September and May.     

Relationships between the movements,growth, age structures,and reproductive biology of the teleosts Sillago burrus and S. vittata in temperate marine waters

Sillago burrus and S. vittata both use sheltered, nearshore shallow waters ( 1.5 m) as nursery areas. However, the juveniles of the former species remain there for only a few months, before migrating into deeper waters (5 to 15 m) as they increase in size, whereas some juvenile S. vittata do not undergo a similar migration until considerably later. S. burrus rarely exceeded 2 yr of age and was never found beyond 4 yr of age. Although only a small number of S. vittata exceeded 2 yr of age, a few individuals of this species were caught between 4 and 7 yr old. The maximum and asymptotic lengths of S. burrus (251 and 180 mm, respectively) were far lower than those of S. vittata (325 and 320 mm, respectively), whereas the growth coefficients (K) were much higher for the former species, i.e. 2.4 vs 0.4. Virtually all S. burrus, and also those S. vittata that moved into deeper waters early in life, spawned at the end of their first year of life. Since relatively few S. burrus reached 2 yr of age, the attainment of almost full size by the end of their first year of life enables a relatively large number of eggs to be produced by fish at the end of their first year-for many, their only spawning period. Those individuals of S. vittata that remained in their shallow nursery areas until the end of their first year of life, did not reach maturity until the end of their second year of life. The proportions of mature gonads and the numbers of yolk-vesicle and yolk-granule oocytes and post-ovulatory follicles in ovaries were far higher in both S. burrus and S. vittata during December to February than in any other month, demonstrating that these two species spawn largely in these summer months. During this period, the ovaries of individual S. burrus and S. vittata often contained post-ovulatory follicles, as well as yolkvesicle and yolk-granule oocytes that ranged widely in size, strongly suggesting that both species are multiple spawners. During the spawning period of S. burrus, the ovaries possessed large numbers of hydrated oocytes and no post-ovulatory follicles or the reverse situation, and the oocytes tended to form several relatively discrete size groups. This indicates that S. burrus produces eggs in batches and that the spawning of the members of this species is synchronised. The presence of large numbers of yolk-granule oocytes with migrating nuclei in the ovaries of many S. vittata at certain times suggests that this species is also a batch- and synchronised spawner. Comparisons between the results of the present study and past work emphasise that the relationships between the timing of offshore movements and the sizes and ages at first maturity vary considerably amongst whiting species.     

Reproduction and reproductive variability in the coral Pocillopora verrucosa from the Republic of Maldives

The sexuality, reproductive mode, and timing of reproduction of Pocillopora verrucosa from the Republic of Maldives, Indian Ocean, were assessed using serial histological sections. These showed that P. verrucosa is an annual simultaneous hermaphrodite, with gonads arranged in two opposing arcs of alternating testes and ovaries, six gonads in each arc. No planulae were observed in any dissection or histological analysis carried out, therefore making brooding unlikely. Broadcast spawning is inferred from the disappearance of mature gametes from samples collected between late March and April 1991. Mean oocyte size at spawning was 53.5 m and mean potential fecundity was 7300 ooctyes cm^-2yr^-1. The reproductive pattern of P. verrucosa in the Maldives is compared to that of the same species in different locations, the short breeding season in March to April occurring earlier in the Maldives than in Red Sea populations. The maximum mean oocyte diameter found in Maldivian specimens was much smaller than elsewhere. The year-to-year variation in numbers of oocytes/polyp within single colonies, and the variation between colonies within the population was significant (p<0.01). Therefore, it is possible that fecundity is not a good variable to use when monitoring stress on coral reefs unless larger numbers of estimates can be routinely made. Some colonies contained immature oocytes at spawning that were not released and continued to grow to approximately twice the size of the spawned oocytes that were presumed to be mature. These unspawned oocytes were oosorbing, and were characterised by the presence of zooxanthellae and large numbers of vacuoles in their cytoplasm. This gradual expansion and oosorption of unspawned oocytes has not been documented previously for corals.     

Comparative fertilization biology of gymnolaemate bryozoans

Locations and oogenic stages of eggs at the time of sperm-egg fusion were determined for nine gymnolaemate species exhibiting different reproductive strategies with respect to site of embryonic development (e.g., water column, introvert, embryo sac, or ovicell) larval type, and zooid sexuality. Ovarian, coelomic, and recently spawned oocytes obtained from freshly collected colonies between 1988 and 1995 were stained with aceto-orcein and examined for the presence of a sperm nucleus. All nine species are shown to have a similar fertilization biology in which sperm fuse with eggs in or near ovaries and egg activation is delayed until eggs are spawned. In species such as the etenostomeAlcyonidium sp. and the cheilostomeElectra pilosa (Linnaeus, 1767), which produce marierons oocytes that are spawned into the water column and have cyphonautes larvae, sperm fuse with late growth stage ovarian oocytes after germinal vesicle breakdown (GVBD) at or near ovulation. In species that produce few oocytes, brood embryos, and release coronate larvae, sperm fuse with late growth stage ovarian primary oocytes before GVBD, as in the ctenostomeBowerbankia gracilis Leidy, 1855, or with very early growth stage ovarian primary oocytes, as in the cheilostomesCribrilina corbicula (O'Donoghue, 1923),Dendrobeania lichenoides (Robertson, 1900),Hippodiplosia insculpta (Hincks, 1882),Schizoporella serialis (Heller, 1867),T ricellaria gracilis (Smitt, 1867), andWatersipora arcuata Banta, 1969. Fertilization success was high in all nine species, with 83 to 100% of oocytes at an appropriate oogenic stage containing a sperm nucleus or early male pronucleus. Gymnolaemate bryozoans may maintain high levels of fertilization success by: (1) concentrating spawned sperm from the water column using colony feeding currents; (2) localizing sperm-egg encounters in or near the ovary, (3) using sperm when available by permitting sperm to fuse with early growth stage oocytes; and (4) selfing.     

Sexual reproduction in the compound ascidian Diplosoma listerianum (Tunicata). I. Metamorphosis,storage and phagocytosis of sperm in female duct

In the colonial ascidian Diplosoma listerianum (collected in the Lagoon of Venice in 1986 and 1991), ovulating eggs detach themselves from the ovary wall and segregate in the tunic without exposure to seawater. With the aim of finding evidence of the pathway followed by spermatozoa to approach the oocyte, histological and ultrastructural observations were made. The results showed that the hollow ovary elongates in a fertilization canal which flanks the sperm duct and opens externally, next to the anus. Intercellular extended tight junctions isolate the lumen of this canal from the blood. Sperm were found in the fertilization canal, both free and in intracellular vacuoles of wandering cells (phagocytes). These sperm showed differences in comparison to those from the sperm duct, which are possible signs of a sperm reaction. The main modifications were shown by the dense groove, a narrow invagination of the plasmalemma bound to the nuclear envelope by dense material. In non-modified sperm, the groove runs spirally all along the head, while in those in the fertilization canal, it coils in the anterior half of the head, pressing back the long mitochondrion and endoplasmic tubules. These modifications were interpreted to be result of the release of the dense groove, considered to be a sort of stretched, contractile spring. The presence of sperm in the fertilization canal and particularly in phagocytes is discussed in relation to the capacity for exogenous sperm storage by zooids and to the necessity of waste clearance in order for successive waves of fresh sperm to be able to move towards fully grown oocytes. The origin of the female fertilization canal, the role of its tight junctions, and the complex sperm morphology are discussed as adaptations to internal fertilization in D. listerianum.     

Spawning and development in <Emphasis Type="Italic">Osedax</Emphasis> boneworms (Siboglinidae,Annelida)

We report observations on spawning and early development in bone-eating worms of the genus Osedax. Individual females of Osedax rubiplumus were observed at 1820 m depth freely spawning hundreds of oocytes, and females of an undescribed species, Osedax “orange collar”, were observed spawning in laboratory aquaria. Cytological and molecular analysis of the spawned oocytes of two Osedax species revealed no evidence for the bacterial endosymbionts that the female worms require for their nutrition, suggesting that the bacteria must be acquired later from the environment, as they are in other siboglinids. Individual O. “orange collar” females released an average of 335 (±130) eggs per day, but the number of oocytes spawned per day varied greatly, suggesting that not all the females spawned daily. Fertilization rates of the spawned oocytes varied from 0 to 100%, though most females showed nearly 100% fertilization rates. Oocytes spawned in the laboratory at 4–6°C were negatively buoyant. If fertilized, these oocytes extruded polar bodies and then after at least four hours cleaved unequally. Subsequent cleavages occurred in a spiral pattern at roughly 2-h intervals, resulting in free-swimming trochophore larvae after 24 h. These lecithotrophic trochophores swam for 9–16 days before settling with several hooked chaetae, similar to those of dwarf Osedax males. The larval life span of the Osedax species studied in the laboratory appears to be shorter than in closely related Vestimentifera. Osedax rubiplumus, on the other hand, has much larger oocytes and so may have greater dispersal potential than these other Osedax species. The high fecundity and apparently continuous reproduction of Osedax boneworms permits the opportunistic exploitation of sunken vertebrate bones. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal cycle of lysosomal enzyme activities in the mantle tissue and isolated cells from the musselMytilus edulis

InMytilus edulis L., gametogenesis takes place in the mantle at the expense of the connective storage tissue. There are two main types of storage cells: vesicular (VC) cells storing large amounts of glycogen and adipogranular (ADG) cells containing large numbers of protein granules, lipid droplets and lesser amounts of glycogen. One of the ways in which stored reserves can be mobilized for gamete formation is by controlled autophagy, in which the cellular constituents are degraded by lysosomes. Mussels were collected from the Menai Strait, North Wales, and monthly measurements made, over two years (1984–1986), of the activities of lysosomal acid hydrolases (acid phosphatase,-N-acetyl-D-glucosaminidase and-glucuronidase) and Cathepsins B and L in the mantle tissue, isolated ADG cells, low-density cells and, during spawning, in the mature oocytes of female mussels. The lysosomal proteinases, Cathepsins B, L and H, were further characterised by activation with thiol compounds and inhibition with thiol blockers and by leupeptin. Because of the low activity in the mantle tissue ofM. edulis, Cathepsin H was not assayed on a seasonal basis. There was a general increase in lysosomal enzyme activity during the winter, which can be related to increased autolysis in the storage cells and to the process of maturation in the developing oocytes. The activity of Cathepsin B was highest in the ADG and low-density cells, implying an important role in proteolysis within the ADG cells. By contrast, Cathepsin L displayed the highest activity in the mature oocytes, suggesting a major function of Cathepsin L in the development and maturation of the oocytes. Two different-glucosidase activities were measured in the monthly assays, one with a pH optimum of 4.5 (acid) and the other at pH 7.5 (neutral). Highest activities of the acid-glycosidase were found in the low-density cells, but there were no significant seasonal changes in the mantle tissue as a whole. Activities of the neutral-glucosidase were low in the ADG cells and mature oocytes, but showed high activities in the mantle tissue, with marked seasonal changes that corresponded to the mobilization of glycogen reserves in the VC cells.     

Long-term changes in reproductive patterns of the holothurian<Emphasis Type="Italic"> Oneirophanta mutabilis</Emphasis> from the Porcupine Abyssal Plain

Time-series studies (1989–2002) in the NE Atlantic have shown large-scale changes in the composition and structure of the benthic community on the Porcupine Abyssal Plain. Radical changes in the abundance of some species in 1996 led to a significant shift in the way in which organic matter was reprocessed at the seabed. This article examines the reproductive processes of the holothurian Oneirophanta mutabilis collected during the time series. The reproductive biology of O. mutabilis is reviewed. No males were evident in any of the samples. The sex ratio (females: no gender) was biased significantly towards no-gender individuals. The maximum egg size was 650 m and there was no evidence of synchrony in reproduction. Significant changes in the oocyte-size distribution and the fecundity of O. mutabilis were noted through time, coinciding with the time of greatest faunal change in the benthic community. There was an increase in the proportion of previtellogenic oocytes and a decrease in the proportion of mature vitellogenic oocytes in 1997 and 1998, in parallel with a significant decrease in fecundity of the post-1996 samples. Samples from 2002 showed a reversal in the reproductive trends, with an increase in the proportion of mature vitellogenic oocytes and fecundity. The results are discussed in relation to large changes in abundance of the epibenthos on the Porcupine Abyssal Plain. It is suggested that the superabundance of certain megafaunal species on the abyssal seafloor affected the availability of trophic resources for O. mutabilis, leading to the changes in its reproductive effort.     

Gonadal development in the coral reef damselfish<Emphasis Type="Italic"> Dascyllus flavicaudus</Emphasis> from Moorea,French Polynesia

The present study investigated the sexual pattern of the yellow-tailed dascyllus, Dascyllus flavicaudus (Randall and Allen), through histological examination of the process of gonadal differentiation and maturation and through male-removal experiments in the laboratory. The study was conducted in Moorea, French Polynesia (17°32S; 149°50W), from 31 December 1994 to 11 March 1995. For gonadal histology, two populations were sampled, one consisting of large aggregations (n=54) and the other consisting of small- to medium-sized groups around isolated corals (n=55). An additional small sample (n=21) was also collected from a population that consisted of small groups around isolated corals. After an initially undifferentiated state, gonads of D. flavicaudus developed an ovarian lumen, followed by oocytes in the primary growth stage. From this ovarian state or from more developed ovaries with cortical-alveolus stage oocytes, some gonads developed into testes through degeneration of oocytes and development of spermatogenic tissue. In all three populations, spermatogenic tissue developed only in gonads that contained pre-vitellogenic oocytes (termed as mixed stage 1–2 gonads). The two main populations did not differ in their expression of sex change despite differences in their social organization. In both populations, size of individuals with mixed stage 1–2 gonads overlapped mainly with the size range of immature females, which suggests that functional female-to-male sex change was rare. The hypothesis that D. flavicaudus is primarily gonochoristic is further supported by removal experiments in laboratory groups, in which removal of a dominant male(s) failed to induce sex change in any of the high-ranking females. Yet, in all three field populations, some fish with mixed stage 1–2 gonads were found within the size ranges of mature females, which indicates possible occurrences of functional sex change. These conflicting results indicate that it may be premature to draw any definitive conclusions about the sexual pattern of D. flavicaudus.Communicated by T. Ikeda, Hakodate     

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.     

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.     

Comparative study of sympatric populations of two hyperiid amphipods,Primno johnsoni and P. evansi,from the eastern North Atlantic Ocean

Hyperiid amphipods are common components of oceanic plankton communities, and yet information is lacking on the ecology and biology of the majority of species. This study compares sympatric populations of two phrosinids, Primno johnsoni Bowman and P. evansi Sheader, using material collected on R.R.S. Discovery Cruises 121 and 140 in the eastern North Atlantic Ocean. The two species were essentially very similar in terms of their vertical distribution and diurnal migration, and in the timing and sequence of reproductive events (onset of maturity, ovary/oocyte development, marsupial development). Mature female and male size ranges were comparable for the two species, males maturing at a small body size, with little subsequent growth, and females producing a succession of 5 (P. evansi) or 6 (P. johnsoni) broods maximum. The species differed significantly in egg size and in total egg output. These dissimilarities were accentuated by differences in the patterns of female mortality in the two species, such that actual egg output per female was 1.8 times greater (2.4 times in terms of total egg volume) in P. johnsoni than in P. evansi. These differences were reflected in the relative size of populations in the field. Although both species are capable of consuming a wide range of planktonic prey, it is assumed that gelatinous species are important, especially as hosts for the parasitoid juvenile stages, and it is suggested that host-specificity might act to separate the niches.     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin     

The reproductive biology of echinothuriid and cidarid sea urchins from the deep sea (Rockall Trough,North-East Atlantic Ocean)

The reproductive biology of 5 species of echinothuriid (Phormosoma placenta, Calveriosoma hystrix, Araeosoma fenestrum, Sperosoma grimaldii and Hygrosoma petersii) and 2 species of cidarid (Cidaris cidaris and Poriocidaris purpurata) sea urchins from the deep sea (Rockall Trough) has been examined from samples collected during 1973–1983. In all species the gonads lie within the interambulacrum attached to aboral gonopores and when fully developed occupy most of the test not occupied by the gut or Aristotle's lantern. In all the species, initial oocyte development takes place along the germinal epithelium embedded in nutritive tissue. In all the echinothuriids and in Poriocidaris purpurata, the oocyte grows to ca. 200 to 450 m, at which stage vitellogenesis begins. Oocyte growth continues until a maximum egg size of 1 100 to 1 500 m is attained. In the echinothuriids, two types of nutritive tissue are found. In the carly stages of gametogenesis the oocyte is surrounded by well-structured periodic acid Schiff (PAS)-positive tissue. As the oocyte grows this tissue becomes vacuolated, suggesting that there is a transfer of nutriment to the developing oocyte. In Phormosoma placenta, unspawned oocytes are phagocytosed. There is no evidence of seasonality in any of the echinothuriid species or in Poriocidaris purpurata. Extrapolation with shallow-water echinothuriids suggests that larval development is lecithotrophic, omitting any planktotrophic phase. Of the species examined, only Cidaris cidaris has a reproductive strategy which produces a known larva, although the limited samples did not permit any determination of seasonality in this deep-sea population.     

Reproductive cycle of the endemic cassiduloid<Emphasis Type="Italic"> Cassidulus mitis</Emphasis> (Echinoidea: Cassiduloida) on the Brazilian coast

Cassidulus mitis Krau, 1954 is an endemic species from Rio de Janeiro, Brazil. To study the reproductive cycle of this species, samples were collected at Praia Vermelha, in Guanabara Bay, from October 1998 to April 2000. The sex ratio for C. mitis was 1:1, and reproduction occurred throughout the year. The gonadal index of males was greater than that of females. Five gametogenic stages were identified in males: early growth, premature, mature, partially spawned and early growth with a partially spawned stage. There were no spent or recovery stages such as those seen in females. Females had six gametogenic stages: recovery, early growth, premature, mature, partially spawned and spent stage; there was no early growth with a partially spawned stage as there was in males. The mean oocyte diameter was 382 m (SD=49 m), and the mean number of juveniles per female was 100 individuals (SD=108). The anterior gonads were frequently smaller than the posterior ones, and there were differences in the gametogenic stages between the anterior and posterior gonads. First sexual maturation occurred in individuals with a diameter of 17 mm. The intestinal wet weight was two- to threefold higher than the test wet weight. This may indicate that C. mitis uses its intestine as a weight belt to avoid displacement from the substratum.Communicated by P.W. Sammarco, Chauvin     

Functional morphology of the gonads of the articulate brachiopodTerebratulina retusa

The reproductive anatomy, and ultrastructural features of the gonads of the articulate brachiopodTerebratulina retusa (Linnaeus), are documented based on collections made between October 1985 and October 1986 from the Firth of Lorn, west coast of Scotland. This species is dioecious, and maturity is achieved in both sexes at shell lengths greater than ~5.5 mm. There is no obvious external sexual dimorphism except for slight differences in the coloration of the gonads; testes are white/cream, ovaries are yellow/orange. The gonads occur as four palmate lobes, a pair in each valve. Gonads are formed within a mantle sinus (vascula genitalia), which is an anterior extension of the coelom, that opens posteriorly into the visceral cavity and to the exterior via a pair of metanephridia. The latter serve as gonoducts during spawning. Gametes are borne on genital lamellae formed from a reticulate lattice of connective tissue. The lamellae are an extension of the ileoparietal band and are fused along one margin to the inner mantle membrane. Developing oocytes are closely affixed to the genital lamellae and originate from a pool of proliferating germ cells at its base. Vitellogenic oocytes that are at an advanced stage are released from the genital lamellae, but are retained within thevascula genitalia. Liberated oocytes continue to accumulate yolk and eventually occlude thevascula genitalia, before being spawned. Coelomocytes were identified within the gonads. In spent females these cells appear to be phagocytic and involved in the resorption of necrotic material, while in the male they may serve as nutritive cells.