Plasma sex steroid profiles and the seasonal reproductive cycle in male and female winter flounder,Pleuronectes americanus

The pattern of seasonal gonadal development and variations in plasma sex steroids were investigated in adult male and female winter flounder, Pleuronectes americanus (Walbaum), from Conception Bay Newfoundland beginning August 1987 to December 1988. The winter flounder reproductive cycle can be divided into five consecutive phases of relative reproductive activity including: (1) rapid gonadal recrudescence in the fall (August–December); (2) continued slow gondadal growth in females, or maintenance of the well developed gonads in males, during the winter (December–February); (3) a prespawning phase of gonadal maintenance in the spring (March–April); (4) spawning early in the summer (May–June) after the female gonads reach peak weight; and (5) the summer postspawned period (June–August) when the gonads remain regressed. Female gonadal recrudescence in August is characterized by small increases in plasma estrogen levels and recruitment of small oocytes (150 m) into yolk accumulation. For the winter months, estradiol-17 levels in the plasma remain stable, approximately 15 ng ml^-1, until rising again together with testosterone to peak hormone levels just prior to spawning in conjunction with the highest seasonal values for the gonadosomatic index (GSI) and oocyte diameter. After spawning, the ovaries are regressed and sex steroids in the plasma fall to very low levels. Rapid seasonal recrudescence of the gonads in males is evident from rising GSI values, which reach a maximum in October, and from substantial early seasonal increases in plasma testosterone and 11-ketotestosterone. Afterwards, although GSI values subsequently decline presumably reflecting the process of spermiogenesis and/or onset of spermiation in some males, the testes remain relatively well developed with the presence of sperm throughout the spawning season. As the proportion of spermiating males increases, the plasma levels of the androgenic steroid hormones rise to peak circulating levels from April to June. At the end of the summer reproductive season, the testes of post-spawned males become regressed and plasma testosterone and 11-ketotestosterone levels fall, reaching the lowest seasonal values.     

A comparative study of the length-weight equation and the condition factor of Tilapia zillii from Lake Mariut,Egypt

The length-weight relationship of Tilapia zillii Gerv., seasonally collected from Lake Mariut (a brackish-water lake), Egypt, has been studied. It was found that mature fishes are fatter than spent individuals, and that this mainly affects the exponent n (W=a L ⁿ). While the females are nearly of equal fatness to the males, a higher value of the exponent n is recorded for the females; this is due to the higher weight of the gonads. The smaller fish are more fecundite than the larger, elder individuals; this is revealed by comparison of the condition factor K and GSI (gonadosomatic index) for ripo and spent fish, respectively.     

Synchronized development of gonad and bioluminescent light organ in a highly sexually dimorphic leiognathid fish, Photoplagios rivulatus

In order to investigate the possible coupling between sexual maturation and the light organ system (LOS) development in leiognathid fish, we determined the seasonal changes in the gonad maturation and the light organ enlargement in Photoplagios rivulatus, one of the leiognathid species possessing highly sexually dimorphic LOS. The fish specimens collected from March 2001 to August 2002 were assessed for standard length (SL), body weight (BW), sex, gonad weight (GW) and the light organ weight (LW). Gonadosomatic index (GSI = 10²GW/BW) and percent weight of the light organ to body weight (PLW = 10² LW/BW) were used to demonstrate the extent of development of gonad and the LOS, respectively. Mean GSIs in both male and female increased in June through August. The mean PLW in males showed a similar trend to that of the GSI, while PLW in females showed no clear seasonal trends. A significant positive correlation was found between the GSI and the PLW in males but not in females. The onset of sexual maturity in males (55 mm SL) coincided with that of the light organ enlargement. These results strongly support the functional coupling between reproduction and bioluminescence in this species of leiognathid. From an evolutionary perspective, sexual dimorphism in the LOS is likely to have evolved through sexual selection for reproductive success in leiognathid fishes.     

Spatial and temporal variation in growth rates and maturity in the Indo-Pacific squid <Emphasis Type="Italic">Sepioteuthis lessoniana</Emphasis> (Cephalopoda: Loliginidae)

Age, growth and maturity parameters are described for the Indo-Pacific squid Sepioteuthis lessoniana both temporally and spatially (equatorial, tropical and subtropical). Tropical squid that grew through periods of warming water temperatures grew 9% faster than squid that grew through periods of cool water temperatures. The tropical spring-hatched and equatorial squid had similar growth rates (3.24, 3.18 g/day) and these were significantly faster than the tropical summer/autumn hatched squid (2.89 g/day). The oldest squid aged was 224 days, but the majority of individuals were <200 days. Subtropical squid were larger, older and matured later than equatorial and tropical spring-hatched squid. The mean weight of subtropical squid aged between 100 and 150 days was >400 g, 85% greater than tropical and equatorial squid. Geographical differences revealed that subtropical mature winter females and males had mean ages >150 days, respectively 17% and 23% older than their tropical mature winter counterparts. Temporal differences in age at maturity were also evident with tropical winter females and males having a mean age of ~140 days, respectively 41% and 25% older than their summer counterparts. Cooler subtropical and winter tropical squid had the heaviest gonads (>15 g ovaries, >1.5 g testes) compared to summer tropical and equatorial squid. However, relative gonad investment (GSI) values of the cooler squid were significantly lower with cool subtropical and winter tropical females having GSI values <3, which was about half the value of the warmer water females. This study revealed considerable plasticity in the size-at-age of this species. The tropical population had growth parameters that fluctuated between an equatorial strategy (fast growth, small body size, and small gonads) and a subtropical strategy (large body size, slower growth, and large gonads) depending on season.     

Analysis of the relationship between gonads and hepatopancreas in males and females of the crab Sesarma intermedia,with reference to resource use and reproduction

At the beginning of the reproductive period in May, there was an increment in both the gonad and heaptopancreas. During the peak phase in June, with high ovary production, there was a decrease in the hepatosomatic index (HSI), particularly for the Ariake females. The minimum gonadosomatic index (GSI) of 2% in August and September for the Ariake and in September and October for the Hakata populations was inversely related to the increase of HSI. A second rise in GSI was observed in November at the 1% level as against the June peak of 5% for the Ariake and 3.5% for the Hakata females, and it remained at this level until December, while the HSI decreased to the minimum. The December minima for both GSI and HSI were about the same as the initial February values. The spermathesomatic index (SSI) had a slight increase to 0.8% in August, but remained constant at 0.5% throughout the year. Likewise, GSI for males was constant at the 0.5% level, with minor fluctuations, and did not show any clear relationship with the HSI as for female GSI. Females had higher hepatosomatic indices but the soma weighed less than males throughout the year. Again in May, medium-sized females had the highest GSI followed by large and then small-sized individuals. As the HSI steadily decreased, GSI increased with advancing egg development stages for ovigerous females. The quantitative stomach analyses showed that females had higher stomach quantity indices (SQI) in May than in September, when the breeding period had ended. Contrary results were observed for males. Also, female SQI in May was higher than that for males.     

Length-structured analysis of the reproductive season of anchovy and common sardine off central southern Chile

The anchovy (Engraulis ringens) and common sardine (Strangomera bentincki) are two small pelagic fish which have a similar reproductive strategy off central southern Chile. The seasonal reproductive dynamics of both the species was investigated by taking into account the gonadosomatic index (GSI) and visual maturity data as a function of female size for the period 1993–1999, and also the seasonal pattern in condition factor per size classes. Larger females of common sardine (repeat spawners) have a reproductive peak earlier in the season than younger females at first maturity, while larger anchovy have a reproductive peak delayed as compared with first-time spawners. The condition factor of females exhibited an inverse cycle with the seasonal pattern of GSI, and delayed by approximately 6 months. Although larger females tend to present better and wider condition than smaller females, the condition of females seems to be delayed in larger females probably mediated by the seasonal pattern in food availability. This “energy storage strategy” in spring and summer time seems to be size-dependent and past energetic reserves could also affect the egg production and timing of reproduction. We concluded that the reproductive season of anchovy and common sardine is different and mediated by the length structure in the seasonal upwelling system off central southern Chile.     

Maturation and diel reproductive periodicity of round scad (Carangidae: Decapterus punctatus)

The diel reproductive periodicity of Carangidae is poorly known but appears to be highly variable between species. Some species spawn during the day, others are believed to spawn at night, and it is demonstrated here that round scad, Decapterus punctatus, spawn at dusk. We collected D. punctatus in the eastern Gulf of Mexico during three April cruises (1995, 1996, and 1997). Based on histological criteria, size at 50% maturity was 113 mm fork length (FL) for males and 128 mm FL for females. The gonad-somatic index (GSI) of mature males was significantly different between hours and appeared to show diel periodicity. Diel periodicity was also observed in changes in female GSIs, whole oocyte diameters, and ovarian histology. The average GSI of mature females fluctuated two-fold between day and night, and the size distribution of whole oocytes in some fish was bimodal (at 0.3-0.4 and 0.7-0.8 mm diameter) at dusk rather than unimodal during most of the diel cycle. Histological preparations revealed that these rapid changes in ovarian GSIs and oocyte size distributions were the result of final oocyte maturation. Germinal vesicle migration was observed from 0900 to 1400 hours eastern standard time (EST), germinal vesicle breakdown was evident as early as 1100 hours EST, and ovulation occurred as early as 1800 hours EST. Spawning frequency (approximately every 5 days) was similar whether calculated from the proportion of females with hydrated oocytes during the afternoon or from the proportion of females with postovulatory follicles during the morning. Batch fecundity correlated with fish size and ranged from 5,500 to 34,700 hydrated eggs per individual. These findings do not support published hypotheses that young-of-the-year D. punctatus reproduce before their first winter or that D. punctatus reproductive output is bimodal within a year.     

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.     

Reproductive biology and ecology of a deep-demersal fish, alfonsino Beryx splendens, over the seamounts off New Caledonia

The reproductive biology of the alfonsino Beryx splendens was studied by histological examinations, gonadosomatic index and macroscopic scales of maturation of a large sample of gonads. Alfonsino is a gonochoric species. The size-frequency distribution of the sex ratio was bimodal and considered to be due to size dimorphism. In New Caledonia, the breeding period of this species occurs during the southern summer, with a peak in December to January. The spawning stage is attained at a minimum fork length of 28 cm for females and 33 cm for males. The size at which 50% of the population attain sexual maturity (FL₅₀) is 33.2 cm for females and 34.5 cm for males. Maximum potential fecundity is estimated to lie between 270 000 to 675 000 eggs for fish between 34 and 40 cm in fork length. It was possible to differentiate vegetative zones, in which juvenile alfonsino grow until they reach maturity, from reproductive zones (fishing grounds) which are inhabited by mature individuals. The larvae and juveniles could be carried from the reproductive zone to the vegetative zone by currents in an oceanic eddy system. Received: 26 April 1996 / Accepted: 20 September 1996     

Spatial–temporal pattern of bluefin tuna (Thunnus thynnus L. 1758) gonad maturation across the Mediterranean Sea

Atlantic bluefin tuna (BFT; Thunnus thynnus) is a migrating species straddling the North Atlantic Ocean and Mediterranean Sea. It is assumed that this species is divided into a western and an eastern stock, which spawn in the Gulf of Mexico and the Mediterranean Sea, respectively. To learn more about the reproductive behavior of the eastern BFT stock, we tracked gonadal development in adult fish that were sampled between April and July during three consecutive years (2003–2005). Sampling campaigns were carried out using common fishing methods at selected locations within the Mediterranean Sea, namely Levantine Sea, Malta, and Balearic Islands. An additional sampling point, Barbate, was situated northwest of the Straits of Gibraltar along the Atlantic coast. Morphometric parameters such as the total body mass (M _B) and the weights of the gonads (M _G) were recorded, and the respective gonadosomatic index (GSI) values were calculated. The data collected revealed two important trends: (1) GSI values are higher in fish caught in the eastern rather than the western locations across the Mediterranean Sea, and (2) the GSI reaches maximum values between late May and early June in Levantine Sea (eastern Mediterranean Sea), and only 2 and 4 weeks later in the central (Malta) and western (Balearic Islands) locations, respectively. The advanced gonadal development in BFT correlates well with higher sea surface temperatures. Our findings also distinguish the northern Levantine Sea BFT population (mean M _B 78.41 ± 4.13 kg), and the Barbate BFT population with the greatest M _B (all fish sampled > 100 kg). These data reflect a situation in which the eastern Mediterranean basin may function as a habitat for young BFT, until they gain a larger M _B and are able to move to the Atlantic Ocean. However, the existence of genetically discrete BFT populations in the Mediterranean Sea cannot be ruled out.     

The reproductive biology of a new species of sea cucumber, Holothuria (Mertensiothuria) arenacava in a Kenyan marine protected area: the possible role of light and temperature on gametogenesis and spawning

The sea cucumber Holothuria arenacava was discovered in the Mombasa marine reserve in 1997 and described by Samyn et al. (2001). The reproductive biology of this holothurian was investigated in order to (1) characterize the reproductive pattern, (2) examine the relationship among environmental parameters including temperature, light and lunar period, and (3) examine the relationship between the reproductive pattern and feeding of this new species. The gonad index method and microscopic examination of gonads was used to analyze samples collected for a period of 13 months. H. arenacava displayed an annual reproductive cycle with gametogenesis commencing in July during the south-east monsoons, when temperature and light intensity are lowest along the Kenyan coast. Gonad growth peaked in February–March at the end of the north-east monsoons when temperatures and light reach their annual maxima along the Kenyan coast. The higher correlation between light intensity and gonad growth (r=93) than temperature (r=0.71), coupled with the fact that temperatures continued to drop for a month after gametogenesis had already commenced, suggests that light intensity and not temperature is the cue for the onset of gametogenesis in this species. Spawning was synchronized between females and males and occurred during a short period between March and May (inter-monsoonal period) when both temperature and light intensity decrease along the Kenyan coast. Male and female gonad indices showed significant variation with lunar day and no lunar periodicity was observed in this sea cucumber. The sex ratio of the population of H. arenacava was skewed towards significantly more females than males, and females were significantly larger and had larger gonads and gonad indices than males. These life history strategies including spawning during a short discrete period, more and larger females that have larger gonads (i.e., typically more fecund), and spawning just prior to the peak in phytoplankton concentrations, a time that is probably more favorable for larval development, may serve to increase the reproductive success of this sea cucumber.     

Ovarian allometry and the use of the gonosomatic index: a case study in the Mediterranean sardine,<Emphasis Type="Italic"> Sardina pilchardus</Emphasis>

The gonosomatic or gonadosomatic index [GSI=(ovary weight/fish weight)×100] has been widely used in fisheries science and experimental reproductive studies as a simple, low-cost measure of reproductive condition. However, its properties have not been fully evaluated, and several pitfalls, such as size-dependence and changes in ovarian allometry, may invalidate its use. In the present study, we examined ovarian allometry and the appropriateness of GSI for assessing ovarian activity in the Mediterranean sardine Sardina pilchardus. The analysis was based on a large sample of histologically scored females collected over an annual cycle in the eastern Mediterranean (Aegean and Ionian Seas). First, we examined GSI dependence on fish size by comparing ovary weight–on–fish weight relationships in different stages of oocyte development. The effects of recent spawning (incidence of postovulatory follicles) and intensity of follicular atresia were also addressed. In a subsequent step of the analysis, we applied generalized linear models (GLMs) to analyze the relationship between GSI and histological stage, taking into account the parallel effects of additional factors related to geographical region, month of capture, fish length and energetic reserves (fat stage, hepatosomatic index—HSI). Ovarian growth was isometric in all stages of oocyte development and states of atresia, but altered to positive allometric at the stage of oocyte hydration. Oocyte growth and intensity of atresia significantly affected GSI. Fish length, geographical region, month of capture, fat stage and HSI did not substantially affect GSI, further strengthening the appropriateness of the index for the Mediterranean sardine. Finally, we provide first evidence, based mainly on available (but yet limited) published information for other fish species, that: (1) the pattern of ovarian allometry may not be altered by fecundity variations and (2) the pattern of ovarian allometry may reflect the pattern of oocyte growth (i.e. isometric vs. allometric ovarian growth could reflect group-synchronous vs. -asynchronous oocyte development). Alternatively, the pattern of ovarian allometry could reflect the presence or absence of size dependency in the relative fecundity (eggs per gram of body weight) and/or egg size. The isometric ovarian growth in the Mediterranean sardine could be attributed to the lack of such size-dependent effects.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Parental care and the cost of reproduction in a Mediterranean fish

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

Sexual reproduction and external brooding by the Caribbean gorgonianBriareum asbestinum

The reproductive biology of the gonochoricBriareum asbestinum (Pallas), a common Caribbean gorgonian, was studied in the San Blas Islands (Panama), La Paguera (Puerto Rico) and the Bahamas. Of 721 colonies examined at the height of the reproductive season, during the summers of 1985 to 1988, all were either male or female. No hermaphroditic colonies or polyps were observed. The sexratio of Panama populations was significantly biased in favor of males (2.1:1,n=454) over all four years of the study. A significant male bias was also found in populations sampled in Puerto Rico (2.3:1,n=143) and in the Bahamas (2.5:1,n=124). The consistency of this geographically widespread skewed sex-ratio is unique among octocorals and scleractinians. Ooyctes appear annually in September and reach maximum diameter (600 to 900µm) in June and July of the following year. The 3 to 5 mo developmental cycle of spermaries is shorter than that of females and begins in March. Spawning in males is synchronous and occurs following the full moons of June and July. Fertilization is internal. Embryos are released from polyps very early in development, often whilst still in the process of cleaving. Embryos remain attached to the outside of the colony for 3 to 5 d, many becoming entrapped in mucous sheets secreted by the colonies. Larvae are negatively buoyant and settle rapidly once displaced from the colony. Males become reproductive at smaller branch sizes than females. Reproductive effort (gonads polyp^–1 and total gonad volume polyp^–1) for both males and females increased with increasing branch size and in polyps away from the tip and base of the branch.Please address all correspondence and requests for reprints to Dr Brazeau at his present address: Department of Biology, University of Houston, Houston, Texas 77204-5513, USA     

Influence of habitat on the reproductive biology of the deep-sea hydrothermal vent limpet <Emphasis Type="Italic">Lepetodrilus fucensis</Emphasis> (Vetigastropoda: Mollusca) from the Northeast Pacific

Habitat selection by the hydrothermal vent limpet, Lepetodrilus fucensis, in Northeast Pacific hydrothermal vent ecosystems, may influence its reproductive output, as it occupies habitats with varying physico-chemical conditions that reflect the availability of nutritional resources. Histological techniques were used to determine size at first reproduction, gametogenesis, reproductive output, and fecundity in relation to shell length (SL), through examination of the gonads of male and female L. fucensis, collected from five different hydrothermal vent habitat types with different temperature anomalies and hydrothermal fluid flow vigour: vigorous (VIG), diffuse (DIF), tubeworm bushes (TWB), peripheral (PER), and senescent areas (SEN). Both male and female L. fucensis exhibited early maturity, with the first reproductive event occurring at 3.8 and 3.9 mm shell length, respectively. All stages of gamete development were present in the gonads of males and females, suggesting continuous gametogenesis and asynchronous reproduction in this species. Gametogenic maturity of limpets did not vary among actively venting habitats (VIG, DIF, TWB, and PER), but was significantly lower in males and females from SEN habitats. Mean oocyte diameter was largest in females from VIG habitats, and smallest in females from SEN habitats, than in those from the other habitats (DIF, TWB, and PER). Females from actively venting habitats also had greater actual fecundity than those from senescent habitats. While the gametogenic pattern of L. fucensis appears phylogenetically constrained, selection of actively venting habitats by L. fucensis maximizes its reproductive output. The multiple feeding strategies of L. fucensis may allow for a constant supply of energy to be allocated to reproduction in any habitat except senescent vents. Early maturity, high fecundity, and continuous production of gametes suggest a reproductive strategy characteristic of an opportunistic species, and may be contributing to the extremely abundant populations of L. fucensis observed in the Northeast Pacific vent ecosystem.     

Manipulation of the seasonal reproductive cycle in winter flounder,Pleuronectes americanus,using a gonadotropic hormone releasing hormone

A gonadotropic hormone-releasing hormone analogue ([D-Ala⁶, Pro⁹-NHEt]LHRH, GnRH-A) was administered to the winter flounder, Pleuronectes americanus (Walbaum) at different stages of the reproductive cycle from September 1988 to July 1989. Both males and females responded to releasing hormone treatment throughout the year except for a brief summer period in post-spawned fish when the gonads were regressed. In females, GnRH-A markedly increased plasma estradiol-17 and testosterone levels and accelerated growth of the ovaries during the period of early gonadal recrudescence beginning in August. In prespawning females, GnRH-A treatment increased plasma testosterone levels in association with a stimulation of germinal vesicle migration and ultimately ovulation. During the fall, GnRH-A treatment of males increased plasma androgen levels (testosterone and 11-ketotestosterone) and stimulated growth of the testes. In February, a time when spermiation is initiated in only half of the males, the proportion of spermiating males was increased by treatment with releasing hormone, indicating that spermiation can be advanced in some males by ca. 2 mo.A portion of this work was given as a poster presentation at the XIth International Symposium on Comparative Endocrinology, Malaga, Spain, May 14–20, 1989     

Synchronous aggregated pseudo-copulation of the sea star <Emphasis Type="Italic">Archaster angulatus</Emphasis> Müller &; Troschel, 1842 (Echinodermata: Asteroidea) and its reproductive cycle in south-western Australia

Mass individual pairing or pseudo-copulation of the sea star Archaster angulatus was observed in November and December 2009, suggesting a late-spring/summer spawning period for this species on the west coast of Australia. Detailed measurements were made on the second of these occasions. Density of sea stars was 1.11 per m² and 68.5% were in mating pairs. Copulating pairs were mostly male on female, occasionally male on male. There was no difference in size between males and females in mating pairs. No evidence was found to indicate mating and spawning is coincident with lunar or tidal cycles. Females outnumbered males by more than 20%, but the difference in sex ratio was not statistically significant. Analysis of the reproductive cycle revealed that gonad indices reached their peak in October and declined from then until January. Histological sections of gonads confirmed that sea stars are in peak reproductive condition in October and November and are fully spent by January. Males have a much lower (ca. 1/3rd) gonad index than females when each are in peak reproductive condition, the second lowest recorded for any sea star. Pyloric caecae indices showed little annual variation and monthly averages of just 3–4% are among the lowest ever recorded for an asteroid. It is suggested that these characteristics are associated with the copulatory behaviour of the deposit feeding A. angulatus, enabling the species to maintain a high level of fertilisation success while also minimising the allocation of energy to gonad development in habitats with low or variable food availability. However, it is still difficult to explain why a species which ensures a high level of fertilisation by pseudo-copulation also does this en masse and synchronously. One hypothesis is that competition for males and the benefits of having eggs fertilised by multiple males favours both synchrony and aggregation.     

Effects of seasonality on the reproductive cycle of <Emphasis Type="Italic">Diadema</Emphasis> aff. <Emphasis Type="Italic">antillarum</Emphasis> in two contrasting habitats: implications for the establishment of a sea urchin fishery

Reproduction of Diadema aff. antillarum was examined between 2002 and 2005 at subtidal rocky bottoms around the Canary Islands. Two contrasting habitats (urchin barrens and grazing fronts) characterized by different levels of food availability were chosen, and factors thought to influence reproductive periodicity were monitored, including temperature, photoperiod, phytoplankton abundance and benthic food availability. Histological analyses showed that D. aff. antillarum had an annual reproductive cycle that was relatively synchronous across the studied sites and habitats. Photoperiod was the most significant factor that correlated with gonad periodicity; benthic food availability of 2 month lag was also correlated. However, some differences were detected between males and females in the timing of the onset of gametogenesis. Spawning was synchronized between both sexes from June to August. Results suggest that the optimum time of year to harvest urchin gonads would be between May and June when gonads were maximal in size but not full of gametes. This species may not provide optimal conditions for an industrial scale fishery, as sea urchins occurring in high density had small gonads and those producing larger volume or more marketable gonad tissue occurred in low densities where harvesting costs would exceed profit.     

Male limitation of female reproductive success in a pipefish: effects of body-size differences

Summary In the pipefish Syngnathus typhle, a species with exclusive male parental care, males limit female reproductive success because of their limited brood pouch space and long pregnancy. Sexual size dimorphism is absent in these 1-year-old animals but increases with age so that older females are larger than similarly aged males. Because fecundity is related to size in both sexes and increases more rapidly with body size in females than in males, the difference in growth increases female fecundity more, relative to male fecundity, as the fish get older. We therefore predicted that male limitation of female reproductive success is even more severe when all age classes are considered. To measure a female's maximum reproductive rate, she was provided with three males. Small 1-year-old females produced as many eggs, or produced eggs at the same rate, as a male of similar size could care for. Small females filled on average 1.06 males within the time span of one male pregnancy and actually produced on average 10 eggs fewer than needed to fill a similarly sized male. Large 2-year-old females, in contrast, produced on average a surplus of 149 eggs and filled 2.7 similarly sized males within the course of one pregnancy. The difference between females of the two size classes was highly significant. Males prefer to mate with larger females if given a choice. In nature sex ratios are equal, and males limit female reproductive success in the whole population. Therefore, small females are more severely constrained by mate availability than are larger females because males choose to mate with larger females. Offprint requests to: A Berglund