ALBERT

Notes: Abstract  Evolutionary change in developmental mode in sea urchins is closely tied to an increase in maternal provisioning. We examined the oogenic modifications involved in production of a large egg by comparison of oogenesis in congeneric sea urchins with markedly different sized oocytes and divergent modes of development. Heliocidaris tuberculata has small eggs (95 µm diameter) and the ancestral mode of development through feeding larvae, whereas H. erythrogramma has large eggs (430 µm diameter) and highly modified non-feeding lecithotrophic larvae. Production of a large egg in H. erythrogramma involved both conserved and divergent mechanisms. The pattern and level of vitellogenin gene expression is similar in the two species. Vitellogenin processing is also similar with the gonads of both species incorporating yolk protein from coelomic and hemal stores into nutritive cells with subsequent transfer of this protein into yolk granules in the developing vitellogenic oocyte. Immunocytology of the eggs of both Heliocidaris species indicates they incorporate similar levels of yolk protein. However, H. erythrogramma has evolved a highly divergent second phase of oogenesis characterised by massive deposition of non-vitellogenic material including additional maternal protein and lipid. Maternal provisioning in H. erythrogramma exhibits recapitulation of the ancestral vitellogenic program followed by a novel oogenic phase with hypertrophy of the lipogenic program being a major contributor to the increase in egg size.

Notes: Abstract Reproduction of the commercial sea urchinParacentrotus lividus (Lamarck) from contrasting habitats on the west coast of Ireland was examined from May 1986 through August 1988. Urchins were collected intertidally from an exposed rocky shore and subtidally from a protected bay. Monthly measurements of the gonad index and histological examination of the gonads demonstrated thatP. lividus has an annual reproductive cycle. Although the two populations exhibited similar reproductive patterns over three breeding seasons, there were inter-annual and inter-population differences in the amplitude of gonadal growth and in the time when spawning started. The subtidal urchins had significantly larger gonads and exhibited a longer period of reproductive maturity compared with the intertidal urchins. This difference was also evident in the histology of the ovaries at the beginning of breeding, when most of the subtidal females contained mature ovaries, whereas most of the intertidal females contained premature ovaries. An inter-annual difference in the onset of spawning was observed, with the start of gamete release differing by as much as four weeks between years. It appears that inter-annual differences in sea temperature influence the temporal variation in spawning byP. lividus and that increasing temperature may serve as an exogenous cue for gamete release. The inter-annual variability in the onset of spawning suggests that photoperiod does not cue gamete release. Gonadal growth, on the other hand, occurs during the coldest months of the year and during the period of shortest days, suggesting that temperature and photoperiod may both influence gonadal growth during the winter. Oogenesis and spermatogenesis ofP. lividus were examined histologically. During the post-spawning recovery and growth stages, the gonads gained weight through growth of the nutritive phagocytes and the accumulation of periodic acid Schiff (PAS) + droplets by these cells. The PAS + material appears to play a nutritive role in gametogenesis. For the females, the frequencies of six ovarian maturity stages was assessed at approximately monthly intervals. Small oocytes were present throughout the year and clusters of early oocytes were most apparent during the spent and recovery stages. With the onset of vitellogenesis and subsequent accumulation of ova, the nutritive phagocytes and their PAS + droplets became depleted. The ovarian condition at the onset of breeding was variable, due to differences in the number of vitellogenic oocytes, differences in the number of ova in storage, and differences in the amount of PAS + material. In general, the nutritive phagocyte tissue is reduced by the onset of spawning and is exhausted by the termination of breeding. A similar series of events occurs during spermatogenesis. The relevance of this study for the use ofP. lividus as a brood-stock organism for mariculture is discussed.

Notes: Abstract The northern Pacific asteroid Asterias amurensis (Lütken) has been recently introduced to Tasmania and is now well-established in east and southeast Tasmania. This sea star is conspicuous throughout the Derwent River estuary and is particularly abundant in the Hobart port area. Reproduction of two populations of A. amurensis at Sullivans Cove and at Sandy Bay was investigated from August 1993 to October 1994 by gonadal histology and measurement of the gonad index (GI). An intense period of vitellogenic and spermatogenic growth started in April, with the breeding condition reached by June. In 1994, peak GI was recorded at Sullivans Cove in July and at Sandy Bay in August. Although these data indicated that the Sullivans Cove population spawned before the Sandy Bay population, histological examination revealed that major spawning activity occurred in both populations from July onwards. In the early part of the breeding season, gametogenesis proceeded in parallel with spawning, with released gametes being replaced by continual gametogenesis. Maintenance of a higher GI during the early part of the breeding season at Sandy Bay was due to prolonged gametogenic replacement at this site. Spawning continued to October 1994, resulting in a sharp decline in the GI. The cyclic expansion and regression of the genital haemal sinus coincident with germinal proliferation and growth, respectively, support the contention that haemal fluid provides nutrients for gametogenesis. In Tasmania, A. amurensis experiences temperature and photoperiod regimes similar to those experienced by endemic populations of this species in the north Pacific. Comparison of reproduction of A. amurensis in Japan and Tasmania at similar latitudes shows that gametogenesis of the northern and southern populations is 6 mo out of phase. This phase shift provides evidence for photoperiodic regulation of gametogenesis in A. amurensis. The similar trend in sea-temperature regimes associated with the stages of gametogenesis in northern and southern populations suggests that temperature also plays a modulatory role. Based on the periodicity of reproduction and the ontogeny of A. amurensis, the planktotrophic larvae of this species would be expected to be in the plankton for several months from the end of winter through summer. Considering the highly fecund, dispersive life-history of A. amurensis, the southeast Tasmanian populations have considerable potential to serve as a seed source for establishment of new populations of this asteroid elsewhere in Tasmania and in mainland Australia.

Notes: Abstract 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.

Notes: Abstract Reproduction and development of the ophiuroidOphionereis olivacea H. L. Clark, a species that broods its young, were investigated in Florida from January through December 1985 and in Belize in June 1985 and April 1986.O. olivacea has a maximum disc diameter of 5.2 mm and is a protandric hermaphrodite. Sex reversal occurs at disc diameters ranging between 2.2 and 4.0 mm; it has 400 µm diameter oocytes and is ovoviviparous. As expected from an obligate out-crosser, this ophiuroid has a similar allocation to male and female reproduction, in terms of gonad volume. Reproduction of the Florida population was monitored for one year.O. olivacea is a sequential brooder and incubates its young during the coldest months of the year. The females produce a single clutch of juveniles each year. Spawning starts in November and peaks in January and February. The ova are spawned synchronously from all the ovaries of each female into the bursae and are retained there. Fertilization is accomplished by intake of sperm into the female bursae.O. olivacea develops through a ciliated embryonic stage that appears to be a modified vitellaria larva lacking ciliary bands. These modified larvae swim in seawater in the bursae. Direct development in this species has resulted from heterochrony in the loss of ophiopluteus features. The number of brooding females in the population increases in November and reaches a maximum in March. JuvenileO. olivacea emerge from the bursae at a disc diameter of 480 µm and each arm has three segments. Recruitment was first detected in April, and juveniles were particularly abundant in June. Small size inO. olivacea is associated with paedomorphic heterochrony with precocious maturation of the males at a disc diameter of 1.5 mm, ~2 mo after taking up their free existence. Brooding as a life history mode inO. olivacea is examined and the adaptations of this species to its warm-water habitats are discussed.

Notes: Abstract Three asterinid seastars, Patiriella gunnii (Gray), P. calcar (Lamarck) and P. exigua (Lamarck) are sympatric in southeast Australia and form part of a series of Patiriella species endemic to Australian shores. Reproduction of co-occurring populations of P. gunnii, P. calcar and P. exigua was investigated in New South Wales from December 1988 to February 1991. Their reproductive cycles were documented by the gonad index (GI) method and by histological examination of the gonads. The annual cycle of the pyloric caecae was also documented and the relationship between the two indices was examined. Oogenesis and spermatogenesis are described in histological detail. The three species produce large yolky eggs and have direct development. Vitellogenesis and weight-gain by the ovaries results from accumulation of periodic acid Schiff (PAS +) material by the oocytes. P. gunnii and P. calcar are gonochoric and have well-defined reproductive cycles, spawning in spring and summer. There was a sharp decline in the GI of P. gunnii in December due to potentially synchronous spawning. Histological examination revealed that initiation of gamete release occurred in August. Gamete release by P. calcar also started in August, with episodic spawning through December. P. gunnii and P. calcar have coincident 3 to 4 mo breeding seasons. In general, P. exigua is protandrous with a broad size range over which sex change occurs. Some P. exigua, however are simultaneous hermaphrodites at the outset of gonad formation and become increasingly female with growth. This species has continuous gametogenesis and oviposits its eggs on the undersides of intertidal boulders, with enhanced oviposition during winter and spring. Newly metamorphosed P. exigua occupy intertidal microhabitats and were located in the field from August to October. It is suggested that increasing sea temperature during spring may serve to cue gamete release by P. gunnii and P. calcar and that enhanced oviposition of P. exigua in winter may be in response to conditions optimal for development of the egg masses at this time. The GI of male and female P. gunnii and P. calcar did not differ, while male P. exigua had a significantly lower GI than the females. Small males are characteristic of this latter species, and it is suggested that the decreased male investment in P. exigua is associated with its non-planktonic development. The gonad and pyloric caeca (PCI) indices of P. gunnii exhibited a reciprocal relationship, indicating that the transfer of nutrients from the caeca to the gonads supports gametogenesis. These indices were partly reciprocal in P. calcar. It is suggested that gonadal growth in P. calcar may be less dependent on caecal reserves than that in P. gunnii. The PCI of P. exigua maintained a similar level throughout the year and was higher than the GI. Although the Patiriella species have coincident breeding periods, temporal differences in the intensity of spawning appears to reproductively isolate them in the field.

Notes: Abstract 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 μm-diam oocytes, suggesting that this species is a broadcast spawner and that it has a modified ophiopluteus or vitellaria larva.

Notes: Abstract 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.

Notes: Abstract The Australian asterinid sea stars Patiriella vivipara and P. parvivipara have the most derived lifehistory pattern seen in the Asteroidea. They are simultaneous hermaphrodites, have intragonadal fertilisation, and incubate their young in the gonads to an advanced juvenile stage. As characteristic of brooding echinoderms, P. vivipara and P. parvivipara are diminutive, with P. parvivipara being the smallest known sea star. These species have the most restricted distribution known in the Asteroidea. Reproduction of two populations P. vivipara in Tasmania, Midway Point and Tesselated Pavement, was examined in specimens collected in 1991, 1992 and 1995. Reproduction of P. parvivipara in South Australia was examined in specimens collected from one population in 1991 and 1994. The gonads are ovotestes. Both species typically contain 6 to 8 predominantly female gonads and 1 predominantly male gonad. In the predominantly female gonads, only a few eggs and a small amount of sperm are produced at a time. Detection of sperm in these gonads requires histological examination. The amount of sperm in the predominantly female gonads appears sufficient to fertilise all the ova produced. The low allocation to male function and the simultaneous presence of mature eggs and sperm suggests that both species may be self-fertile. The amount of sperm in the predominantly male gonads however, would not be expected in exclusively self-fertilisers, suggesting that some outcrossing occurs. The significantly higher reproductive output of P. vivipara at Midway Point than at the Tesselated Pavement indicates that the Midway Point population is particularly important for the conservation of this species. P. vivipara and P. parvivipara have small, 140 to 150 μm-diam eggs. Embryogenesis is asynchronous, with progeny at different stages of development in the gonads. The end-point of brooding is variable in P. vivipara, with juveniles emerging from the parent at sizes ranging between 1.5 and 5.0 mm diam. Juvenile P. parvivipara are born at a diameter of up to 2.5 mm. At birth the juveniles of both species are up to 25–30% of the parent's diameter. Birth involves distension of the gonopore, with a marked separation of the ossicles and softening of the connective tissue around the pore. The juveniles are considerably larger than the ova and depend on extraembryonic nutrition to support their growth. Once the mouth opens, the juveniles prey on their intragonadal siblings. Cannibalism accounts for the substantial post-metamorphic growth. Viviparity lies at the extreme end of the broadcasting-brooding continuum of life histories in Patiriella, and the life-history traits of P. vivipara and P. par-vivipara are compared with those of other Patiriella spp. to assess the changes associated with evolution of viviparity.

Notes: Abstract Reproduction ofHeliocidaris erythrogramma (Valenciennes) andH. tuberculata (Lamarck) was compared through examination of oogenesis, spermatogenesis and monthly measurement of the gonad index. These species occur sympatrically in the Sydney region. Their reproduction was examined at two sites near Botany Bay, New South Wales, Australia, from February 1989 through January 1990.H. erythrogramma produces buoyant, 450µm-diam eggs and the sperm have a head region 10µm in length. By contrast,H. tuberculata produces negatively-buoyant, 95µm-diam eggs and the sperm have a head region 4µm in length. Histochemical examination of the gonads revealed that periodic acid Schiff-positive (PAS +) material stored in the nutritive phagocytes appears to support vitellogenesis in both species. InH. tuberculata this material is utilized in the formation of PAS + yolk oligolecithal eggs, whereas inH. erythrogramma the PAS + material appears to be converted to lipid yolk in macrolecithal eggs.H. erythrogramma had a seasonal reproductive pattern with a 3 mo summer spawning period, whereas both populations ofH. tuberculata had a 9 mo breeding period characterized by the continual presence of nutrient reserves and vitellogenic oocytes which rapidly replaced spawned ova. Spawning ceased only for 3 mo over the summer. Due to the 9 mo spawning ofH. tuberculata it is not clear what factors serve to cue reproduction in this species.