Seasonal changes in biochemical composition of body walls,gonads and pyloric caeca in two populations ofSclerasterias mollis (Echinodermata: Asteroidea) during the annual reproductive cycle

Seasonal changes in biochemical composition of tissues are compared in a shallow inshore and deeper offshore population of the seastarSclerasterias mollis (Hutton, 1872) on the Otago shelf, New Zealand, from 1985 to 1986. The biochemical composition of the body wall, gonads and pyloric caeca of the two populations did not differ greatly, even though reproductive output and nutrient storage of reserves were consistently higher in the offshore population. The biochemical composition of the ovaries and testes differed. High levels of carbohydrate and lipid were maintained in the ovaries from the commencement of vitellogenesis until spawning occurred. The testes had higher ash levels during spermatogenesis. During the annual reproductive cycle, an inverse relationship in the carbohydrate levels between the ovary and the pyloric caeca suggests nutrient transfer from the pyloric caeca to the gonads during gametogenesis. Annual changes in biochemical composition indicate that the body wall is also used for nutrient storage.     

Seasonal changes in pyloric caecum and gonad indices during the annual reproductive cycle in the seastar Asterias forbesi

Seasonal changes in pyloric caecum and gonad indices were studied in a population of the seastar Asterias forbesi (Desor) from East Rockaway Inlet (Long Island, New York) during two annual cycles (1980/81, 1982/83). Pyloric caeca indices increased during fall, reached a maximum about April, and declined sharply to a minimum in mid-summer. Gonad indices increased during fall and winter and reached a maximum about May. Judging from gonad size analysis, spawning occurred in late June to early July, at bottom temperatures of 16° to 18°C. There was no long-term inverse relationship between pyloric caecum and gonad indices. Since nutrients and energy ingested during the important fall feeding period are utilized simultaneously for body growth and gonad development, it is unnecessary for the pyloric caeca to store nutrients over long periods as occurs in A. rubens and many other species. The specific caeca-gonad relationship is probably an adaptation to the extreme seasonal thermal fluctuations of the NW Atlantic, which constrain the feeding activity of A. forbesi and limit the possibility of long-term storage. In this population, most individuals grow rapidly in their first year (including first two summers) and spawn in their third summer. Few seastars survive to spawn again in their fourth summer. However, gametogenesis may take place in some individuals in their second summer. Year-to-year variability in mean size at spawning may reflect temporal variation in environmental conditions (weather, food availability) at this site.     

Energy investment in the annual reproduction cycle of female red porgy, <Emphasis Type="Italic">Pagrus pagrus</Emphasis> (L.)

Seasonal cycle in energy content of gonad, perivisceral fat, liver, and muscle was used to estimate the energy cost of spawning of female red porgy (Pagrus pagrus) from Buenos Aires coasts, Argentina. Spawning was observed primarily between November and January, although a low-intensity spawning was also observed at the beginning of this period. The energy required for gonad development of female red porgy (the difference between the maximum and minimum energy content of ovaries along the year) was 2,652 J g⁻¹ dry weight or 558 kJ. Female red porgy expend 22.8% of their total body energy (3,481 kJ) during the spawning period. Of this energy, relative contributions from different sources were as follows: ovaries 62.5% (496 kJ), perivisceral fat 17.8% (142 kJ), liver 10.3% (82 kJ), and muscles 9.4% (75 kJ). Female spawners lost about 7.2% (119 g) of their total body weight (1,654 g) in the spawning period. From the 7.2% of the total weight lost, female gonad represented a relative reduction of 73.8% (87.6 g), the perivisceral fat declined by 5.7% (6.8 g), the liver declined by 10.6% (12.5 g), and muscle tissue exhibited a 10% loss in total weight (11.9 g). Reproductive effort represented 7.6% of food energy, while resting metabolism accounted for almost 38% of food energy. Despite the fact that red porgy females feed throughout the entire spawning period, it is shown that between pre-spawning (September, water temperature 15°C) and mid-spawning (December, water temperature 19°C) they lost 45 and 38% of their muscle and liver lipids, respectively, and 89% of the perivisceral fat associated with the gut and body cavity. Red porgy used both their energy reserves and food energy during ovarian maturation. This asynchronous species have a strategy of a lower energy allocation and a higher dependence of food during the spawning time (late-spring and summer) for successful reproduction.     

Reproductive biology of <Emphasis Type="Italic">Cosmasterias lurida</Emphasis> (Echinodermata: Asteroidea) an anthropogenically influenced substratum from Golfo Nuevo,Northern Patagonia (Argentina)

The starfish Cosmasterias lurida was sampled from the pier piles of Almirante Storni in the Golfo Nuevo, northern Patagonia, from November 2001 to November 2003. In this anthropogenically modified site there is an unusually abundant food supply of mussels and large aggregations of starfishes. The reproductive cycle was studied by organ indices and histological examination of gonads. The highest gonad index (GI) values were found in the summer (February–March), followed by a drop in GI values, indicating spawning. Histological analyses indicated that gametogenesis takes place from June to January, sexual maturity occurred during the summer and spawning occurred in April. The GI and pyloric caeca index (PCI) tended to have an inverse relationship, but this was only statistically significant in males. However, biochemical analyses of the starfish indicated no transfer of nutrients between pyloric caeca and gonads in either sex. In fact, simultaneous accumulation of carbohydrates was found in the pyloric caeca and gonads of both sexes.     

Population differences in gonad and pyloric caeca cycles of the New Zealand seastarSclerasterias mollis (Echinodermata: Asteroidea)

Inshore and offshore populations ofSclerasterias mollis (Hutton, 1872) were sampled on the outer continental shelf off the Otago Peninsula, New Zealand, and their nutritional and reproductive cycles are described from 1985–1986. Histological changes in the gonads are generally typical of other asteroids. The gametogenic cycle takes 12 mo. The gonad and pyloric caeca indices in both females and males had an inverse relationship.S. mollis accumulates nutrients in the pyloric caeca during summer and early autumn. The gonads develop in the autumn and winter. Offshore seastars were much larger and had significantly higher gonad and pyloric caeca indices than inshore individuals. These differences in body sizes and organ indices arise from differences in food availability and/or population density.     

Changes in the biochemical composition of the ovaries of the seastar Asterias rubens during its annual reproductive cycle

The biochemical composition of the ovaries of the asteroid echinoderm Asterias rubens was studied, with emphasis on the changes which are manifest during the annual reproductive cycle. Ovarian dry weight displays only minor changes during ovarian development, constituting about 20% of the organ fresh weight. Levels of all constituents show some variation, but in general increase with ovarian growth. Only the free amino acid level is rather constant. Except for glycogen, the contents of other constituents (total lipids, other reducing sugars, free amino acids and proteins) show a rectilinear increase with ovarian growth. Glycogen content increases rectilinearly with time during early vitellogenesis, but remains constant during further development until ovarian maturity. By plotting constituent levels and contents against a non-linear gonad index axis rather than against sampling date, results can be correlated more easily with ovarian growth stages. The possibility of endocytosis of a vitellogenin-like yolk precursor by the ovary is discussed.     

Sexual and asexual reproduction of<Emphasis Type="Italic"> Allostichaster capensis</Emphasis> (Echinodermata: Asteroidea) in Golfo Nuevo

Sexual and asexual reproduction of the fissiparous starfish Allostichaster capensis were examined for 24 months at Bahía Kaiser, Golfo Nuevo, Chubut, Argentina. Gonad indices revealed an annual reproductive cycle, with peak development occurring in late winter (August). Spawning occurred in spring (September). Pyloric caeca indices displayed a clear annual cycle and a reciprocal relationship with gonad indices, showing the storage function of pyloric caeca. The starfish population appears to be largely maintained by fission, as only two females were found. A. capensis also showed an annual cycle of fission, with the highest frequency in spring and summer (from November to January), just after spawning. Incidence of fission was correlated with environmental factors such as photoperiod, seawater temperature, and salinity.Communicated by P.W. Sammarco, Chauvin     

Reproductive cycle of the sea urchin Holopneustes purpurascens (Temnopleuridae: Echinodermata)

This study documents the annual reproductive cycle of the Temnopleurid echinoid Holopneustes purpurascens near Sydney. H. purpurascens reached sexual maturity between 11 and 26 mm diameter and was gonochoristic. The sequence of oogenesis was similar to previously described echinoids. However, the proportion of eggs at each stage showed no temporal pattern in abundance, suggesting that vitellogenesis is continuous throughout the year. Spermatogenesis was also similar to other echinoids and showed a significant increase in the proportion of partially spawned testes in spring, followed by a similar increase in the proportion of partially spawned testes in late spring and summer. Gonad indices showed a peak in weight from late winter to mid-spring (9.10ǃ.47 for females; 6.28ǂ.43 for males; mean-SE), then a decline for both sexes. Although spawning was variable over time, there was a peak in the proportion of spawning from late winter to early summer that correlated with the observed peaks in gonad indices and gametogenic cycles. The demographic implications of this reproductive pattern are that new recruits can enter the population through synchronous spawning during the breeding period, and smaller haphazard spawning events at other times of the year.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Storage and depletion of lipid components in the pyloric caeca and ovaries of the seastarAsterias rubens during its annual reproductive cycle

Detailed information is presented on the changes in lipid-class profiles (polar lipids, total neutral lipids, free sterols, steryl esters, triacylglycerols, neutral ether lipids and free fatty acids), which are manifest during the annual reproductive cycle of the seastarAsterias rubens for both the storage organs (pyloric caeca) and utilizing organs (ovaries). The ovaries appear to accumulate all individual lipid components proportional to progression of vitellogenesis. Pyloric caeca, in contrast, show variable lipid-class levels, probably due to more complex processes in these organs, operating simultaneously: digestion of food, accumulation of storage lipids, mobilization of stored lipids and their release. The results are discussed in terms of current knowledge on the role of lipids in echinoderm reproduction.     

Reproduction of sympatric populations of Patiriella gunnii,P. calcar and P. exigua in New South Wales,asterinid seastars with direct development

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.     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.     

Seasonality of reproduction in the cushion star Odontaster validus at Signy Island, Antarctica

The reproductive and larval biology of the common Antarctic cushion-star Odontaster validus Koehler (Echinodermata: Asteroidea) were studied in a shallow-water population at Signy Island in the maritime Antarctic. The gonad index peaked in April/May, and injection with 1-methyladenine indicated that competence to spawn peaked in May/June. Gastrulae were first detected in May, and maximal water-column larval densities were measured in July. Reproduction was therefore highly seasonal, although not all individuals appeared to spawn each year. The pyloric caeca index increased in summer (typically November to March, but variable between years) and decreased during winter (June to October), suggesting that feeding activity was seasonal. All aspects of biology investigated showed significant interannual variability. Comparison with previous studies in the high Antarctic at McMurdo Sound indicated a similar if more pronounced seasonality at Signy, but the warmer seawater temperatures in the maritime Antarctic (particularly in late winter/early spring from September onwards) resulted in faster larval development at Signy. Received: 21 April 1997 / Accepted: 28 January 1998     

Annual reproductive cycles of the commercial sea urchinParacentrotus lividus from an exposed intertidal and a sheltered subtidal habitat on the west coast of Ireland

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.     

Seasonal changes in biochemical composition of the bivalve Lima hians from the Clyde Sea Area

Seasonal changes in the tissue weight and biochemical composition of the bivalve Lima hians (Gmelin) from a station in the Skelmorlie Bank, Clyde Sea Area, were studied during 1969–1971. The composition of an animal of standard size was calculated for each sampling date to give information on seasonal changes independent of shell growth. Increases in tissue weight took place between April and July as the gonad developed and reserves were built up. Between July and March tissue weight fell, at first rapidly as a result of spawning in the population, and later more slowly as reserves were utilized. Differentiation of the gonad began during the winter, at the expense of stored reserves, but was greatly accelerated during the spring due to increased food from the plankton. The lipid and nitrogen content of a standard animal rose to a pre-spawning maximum in July and then fell. Carbohydrate content also rose during this period, but continued to rise to reach maximum values in September. Carbohydrate and nitrogen both declined during the winter, but there was little indication of any net utilization of lipid during the winter months. The tissues of the ripe female contained more lipid than the male, and the tissues of the ripe male contained more nitrogen than those of the female. The total calorific content of the tissues of the male was greater than that of the female.     

The reproductive cycle of Asterina stellifera (Möbius) (Echinodermata: Asteroidea) in the Cabo Frio region, southeastern Brazil

Reproduction of Asterina stellifera was studied in two populations from the Cabo Frio region, southeastern Brazil. One was located in an intertidal seagrass flat at Japonês Island (1994–1995); the second was on a sublittoral rocky shore at Pontal beaches (1999–2000). Reproductive cycle was studied at monthly intervals by organ indices and histological examinations of gonads. Although these two populations were studied in different habitats and at different times, the periodicity of the annual reproductive cycle was very similar for the two populations. The highest gonad index (GI) values were found in August/September (late winter), followed by a drop in GI values, indicating spawning. The GI and pyloric caeca index (PCI) tended to have an inverse relationship in both populations, but it was only statistically significant in the rocky-shore population. Gametogenesis in the two populations was described and roughly confirmed the reproductive cycles determined by GIs. Published online: 9 August 2002     

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.     

Seasonal changes in biochemical composition of the bivalve Abra alba from the Clyde Sea Area

Seasonal changes in the body weight and composition of Abra alba (W. Wood) from south of the Little Cumbrae in the Clyde Sea Area (UK) were studied during 1969–1971. The composition of a specimen of standard size was calculated for each sampling data to provide information on seasonal changes independent of shell growth. Increase in body weight took place rapidly in early summer as the gonad developed and reserves were built up. Spawning in July resulted in a fall in body weight, and this was followed by a slower fall during the autumn and winter as reserves were utilized. The changes are related to seasonal fluctuations in food availability and temperature, which are reflected in the amount of pigment (phaeophytin) retained in the tissues.     

Seasonal changes in the physiology,reproductive condition and carbohydrate content of the cockle Cardium (=Cerastoderma) edule (Bivalvia: Cardiidae)

Pronounced seasonal cycles in the rates of oxygen consumption and feeding were found for Cardium (=Cerastoderma) edule L. measured in the field under ambient conditions. The cockles had a maximum rate of oxygen consumption (0.89 ml O₂ g^-1 h^-1) in April which declined to a minimum of 0.35 ml O₂ g^-1 h^-1 in March. Their feeding rate was variable but had a maximum value (3.91 l g^-1 h^-1) in April and a minimum value (0.73 l g^-1 h^-1) in October. There was no apparent seasonal variation in absorption efficiency, with a mean value of 67.6%. Gametogenesis was initiated in January and the population reached a peak in reproductive condition in April/May, followed by a 3 month spawning period. Carbohydrate reserves were synthesised during spawning, and were then utilised during the winter and early spring. An adaptive function for a reduction in time spent feeding is postulated, and correlations between the rates of certain physiological processes and some exogenous and endogenous variables are discussed.     

Growth and spawning of Urechis caupo (Echiura) in Bodega Harbor,California

Growth and spawning of the large, infaunal echiuran worm Urechis caupo Fisher and MacGinitie were studied at Bodega Harbor on the coast of central California, USA, from 1978 through 1981. In situ growth rates of marked worms were negatively related to initial size. Short-term, summer growth rates (volmo^–1) of small worms (<80 ml) were greater than longer-term growth rates measured over several seasons (asesonal). Size-frequency distributions of worms sampled from two sites also suggested a seasonal growth pattern with relatively fast spring-summer growth and slower winter growth. However, larger worms sometimes lost volume during in situ growth experiments, and the loss was most pronounced during short-term, summer growth periods. It is suggested that energy used in burrow construction may have contributed to volume loss during short-term growth experiments. In contrast, longer-term, aseasonal growth rates were nearly always positive, and indicated that reproductive size (about 56 ml) could be reached within about 1.5 yr of recruitment, and a large size (about 158 ml) could be reached within about 6 yr. A seasonal pattern of spawning was observed during three consecutive years, as indicated by ripeness indices (storage organ dry weight ÷ body wall dry weight). At least two spawning episodes occurred annually: ripe gametes that accumulated in the storage organs during the summer and fall were spawned during the winter; gametes that accumulated during late winter and early spring were spawned during the spring or early summer. Worms were spawned-out by mid-summer.