The spermatogenetic process in Barbus longiceps, Capoeta damascina and their natural sterile hybrid (Teleostei, Cyprinidae)

Testicular ultrastructure was studied in Barbus longiceps, Capoeta damascina and their natural hybrid. The testes of these teleosts belong to the unrestricted or lobular type. Germ cell morphology is similar in the parental males. In the hybrid, spermatogenesis does not extend beyond the pachytene of the first meiotic division, probably due to the unsuccessful pairing of the homologous chromosomes. Hybrid testes are occupied mainly by degenerating primary spermatocytes, at the leptotene and pachytene stages. In both parents and the hybrid, Sertoli and Leydig cells are characterized by the presence of granular endoplasmic reticulum and of mitochondria with tubular cristae. Due to the arrest of spermatogenesis, the male germ cell protective barrier is absent in the hybrid. Germ cell nuclear size was measured by a computerized analysis system, using light-microscopy images. In the parents and the hybrid, germ cells attain a uniform inter-individual nuclear size when they reach the first meiotic prophase. The nuclear size of primary spermatocytes is similar among the three groups of fish, possibly reflecting their close genetic relationship.     

A new natural intergeneric cyprinid hybrid from the Jordan River drainage, with a key to the large barbine cyprinids of the southern Levant

In Ain al-Qunaiya. an isolated source within the Jordan River drainage basin of Jordan, several fishes were found to have characters intermediate between two cyprinid species. Barbus canis Valenciennes in Cuvier & Valenciennes. 1842 and Capoeta damascina (Valenciennes in Cuvier & Valenciennes. 1842). The hypothesis of their hybrid origin is advanced. A key to the large barbine cyprinids of the Jordan River system and their hybrids is given.     

Seasonal changes in condition, nutrition, gonad maturation and energy content in barbel, Barbus sclateri, inhabiting a fluctuating river

The objective of this study was to investigate the seasonal cycle of condition, nutrition and gonad development, as well as the magnitude of seasonal variations in energy content of somatic and gonad tissues in juveniles, males and females of Sclater's barbel, Barbus sclateri, from the upper Guadalete River (south Spain). The influence of reproductive cycle on somatic changes was also investigated and discussed. Measurements of condition factor (K), somatic index (SI), gonadosomatic index (GSI) and somatic and gonad energy content (J g^-1 dw) were made in individual specimens taken from the Guadalete River monthly for 12 months. This freshwater ecosystem is characterised by strong seasonal fluctuations in water and flow level, temperature and food supply. It was found that in general juveniles, males and females of barbel exhibited a similar condition, nutrition and somatic energy cycle throughout the year, with the highest values in spring and the lowest in summer. Depletion of K, SI and somatic energy storage from spring until mid-summer seems to be associated with high metabolic demands during this period, and in adult fish also with spawning-related activity. The gonad index and energy content of the gonad were the highest in April and the lowest in summer for both sexes. Spawning took place during late spring – early summer, with fish quiescent by mid summer. The energy required for ovarian development (3970 J g^-1 dw) was greater than the one for testes development (2763 J g^-1 dw). Data on gonad energy content indicated a period (March to April) of intense energy accumulation (64% males and 37% females) which was related to the decline in the average somatic energy content in males and females. The somatic energy content was linearly related to K and SI. In the same way, GSI correlated positively with gonad energy. Linear trends were found between somatic parameters (K, SI and energy content) and gonad parameters (GSI and energy content). This revised version was published online in July 2006 with corrections to the Cover Date.     

Plasma sex steroids and gonadosomatic index variations during ovarian development of female wild yellowfin seabream (Acanthopagrus latus)

The yellowfin seabream (Acanthopagrus latus) has a good potential for mariculture in Iran, so studies on various aspects of its culture has been carried out. In this study, the characteristic of reproduction in this species was investigated. The involvement of steroid hormones in the development of gonads was assayed by sampling blood and gonad tissue in 85 females of A. latus in fish caught from the Musa Creek, north-west Persian Gulf from October 2010 to May 2011. Gonadosomatic index (GSI) and hepatosomatic index (HSI) were recorded. Developmental stages were associated with changes in sex steroids, GSI and HSI. GSI and HSI began to increase in November (immature stage) and reached the highest value in March (ripe stage), decreasing sharply in April (spent stage). Plasma values of 17β-estradiol (E2) levels reached their highest levels in spawning females in March and then decreased in April. Progesterone levels were low through ovarian development, reached a maximum simultaneously with testosterone in February (mature stage) and then decreased in March (ripe stage). The seasonal changes observed in plasma steroid levels and GSI reflected the pattern of oocyte development and the spawning behavior of yellowfin seabream and were typical of the patterns described in most multiple spawners studied to date.     

Bounded hybrid superiority in an avian hybrid zone: effects of mate, diet, and habitat choice

There has been considerable debate in the study of hybrid zones as to whether hybrids may be superior to parental types within the area of contact (bounded hybrid superiority). In birds, naturally occurring hybridization is relatively common, and hybridization within this group always involves mate choice. If hybrids are superior, females choosing heterospecific mates should be expected to show higher fitness under the conditions prevalent in the hybrid zone. Hybrid superiority under these circumstances would reduce reinforcement and thereby help to maintain the hybrid zone. To examine this issue, we studied reproductive performances of hybrids and parental species of gulls (Larus occidentalis and Larus glaucescens) at two colonies within a linear hybrid zone along the west coast of the United States. This hybrid zone contains predominantly gulls of intermediate phenotype. Previous studies indicated that hybrids were superior to one or both parental types, but provided no data on possible mechanisms that underlie this hybrid superiority. Using a hybrid index designed specifically for these species, we identified to phenotype more than 300 individuals associated with nests, including both individual males and females within 73 pairs in the central portion of the hybrid zone and 74 pairs in the northern portion of the hybrid zone. There was little evidence of assortative mating, and what little there was resulted solely because of pairings within intergrades. In the central hybrid zone, females paired with hybrid males produced larger clutches and hatched and fledged more chicks compared with females paired to western gull males. This was a result of heavy predation on eggs in sand habitat, where male western gulls established territories. In contrast, many hybrid males established territories in vegetated cover that was less vulnerable to predation. In the northern part of the hybrid zone, clutch size did not differ among pair categories, however, there were differences in hatching and fledging success, with females paired to hybrid males showing better success compared to females paired to glaucous-winged gull males. Hybrids showed better hatching and fledging success in the north because hybrids are more like western gulls than glaucous-winged gulls in foraging behavior, taking a higher percentage of fish in their diet, which enhances chick growth and survival. This is believed to be the first documentation of bounded hybrid superiority that delineates the mechanisms that underlie hybrid superiority.     

Some aspects of the reproductive biology ofBarbus spp.,Capoeta damascina and their hybrids (Cyprinidae, Teleostei) in Israel

The reproductive biology and gonad cycle of three Cyprinid fish species:Barbus canis (Valenciennes, 1842),B. longiceps (Valenciennes, 1842) andCapoeta damascina (Valenciennes, 1842), in the upper Jordan River system of Israel were studied by monthly sampling over a two-year period. The reproductive activity of the three species was found to peak from January to April, mostly involving upstream migration towards spawning grounds on river beds 400–900 m above the Jordan River. Hybrids of the three species were collected in nature: in those ofBarbus canis ×Capoeta damascina, the gonads possessed both types of gametes, spermatogonia and oogonia, all of which became arrested at an early stage of development, and infertile; in hybrids of detected in nature, males had oocytes dispersed in the testis, whereas in females, the ovaries had small islets of spermatogonial tissue. In these female hybrids the oocytes ripened normally and spawning occurred.     

Hybridization studies in the planthopper genus Ribautodelphax (Homoptera,Delphacidae)

Seven known Ribautodelphax species and four recently discovered taxa were crossed. Forty-three percent of the interspecific crosses were successful. Interspecific crosses generally resulted in low numbers of hybrids. In some combinations prolonged larval development of the hybrid males was observed. Backcrosses showed that both male and female hybrids were fertile. Reproductive isolation was mainly maintained by premating barriers. Insemination tests revealed that prolonged confinement of males and females lowered mating thresholds. The taxonomic status of the known species and new taxa was shown to be in need of revision. No indications for a hybrid origin of the pseudogamous triploid Ribautodelphax females were found.     

Gonad development and sperm motility of the diving beetle Cybister brevis Aubé, 1838 (Coleoptera: Dytiscidae) in response to seasonal changes in Japan

Gonad maturation in wild Cybister brevis Aubé, 1838 from Japan was investigated to determine the physiological mating season. Breeding experiments showed that hatching occurred from May to September, and the peak was observed in July. The gonadosomatic index (GSI) in collected females (ovary development) showed the highest value in May. The GSIs were low in other seasons. On the other hand, the GSIs in collected males (testes and accessory glands) remained at constant values during the experimental period. Interestingly, high sperm motility was exhibited in May and September, whereas it was low in December and March. This indicates the maturation difference between females and males. Females mature only at the beginning of the breeding season (May), but males do so from May to September. This result suggests that males have a wide-ranging maturation period and earlier sexual maturation than females, ensuring a successful fertilization.     

A comparative study on morphology, growth rate and reproduction of Clarias gariepinus (Burchell, 1822), Heterobranchus longifilis Valenciennes, 1840, and their reciprocal hybrids (Pisces, Clariidae)

A detailed study on reproduction indicated that female C. gariepinus mature earlier (5–7 months) than female H. longifilis (12–14 months). In the reciprocal hybrids, female first sexual maturity was attained at 20–21 months. Both reciprocal hybrids and parental species displayed an equilibrated sex ratio. Macroscopical and microscopical observations revealed numerous abnormalities in gonadal development of the hybrids. GSI and fecundity in female hybrids were considerably lower than those found in the parental species. Moreover, intra-ovarian tumors occurred in 20% of the hybrids. In hybrid males, GSIs were generally higher than those found in the parental species, but the spermatozoa concentration in the semen was about 100 times less. Despite these abnormalities, viable larvae, resulting from F2 and backcross fertilizations, were obtained.     

Temporal variation in condition index and meat quality of Lunella undulata (Turbinidae), in relation to the reproductive cycle

ABSTRACT

Lunella undulata is a subtropical-temperate marine gastropod which is harvested commercially from temperate reefs in Australia. This paper aimed to evaluate monthly variation in the condition index (CI) and biochemical composition of the foot tissue of L. undulata in relation to the spawning cycle. The gonadosomatic index (GSI) varied significantly between males and females and across months (P?=?0.0001), ranging from 43% to 70% in males and 41% to 71% for females. The highest GSI, as well as the highest CI for both sexes was recorded in December and April, whilst meat yield peaked in January, September and October. Protein, lipid, fatty acid and heavy metals in the foot tissue also significantly varied between months (P?<?0.05). Throughout all 14 months of sampling, the foot tissue of L. undulata showed good nutritional quality, with high levels of protein and the polyunsaturated fatty acids, docosapentaenoic acid, eicosapentaenoic acid and arachidonic acid. Overall, despite some temporal variability in the biochemical composition, L. undulata could be harvested at any time of the year for human consumption, although it may be best to avoid peak spawning times, which occur around January in northern NSW.     

Behavioral reproductive isolation inDrosophila silvestris,D. heteroneura,and their F1 hybrids (Diptera: Drosophilidae)

We investigated the role that courtship and aggressive interactions may have for the maintenance of reproductive isolation betweenDrosophila silvestris andD. heteroneura. We examined the behavioral bases of reproductive isolation between the parental species and we examined the courtship success of each sex of both reciprocal F₁ hybrids when paired with the parental species. We found reduced copulation success among heterotypic parental pairs compared to homotypic pairs, which was primarily due to the lack of courtship initiation betweenD. silvestris males andD. heteroneura females. When hybrid males from both reciprocal crosses were paired with parental females their copulation successes were not significantly different from that of parental males. In contrast, hybrid females from both crosses had reduced copulation success withD. silvestris males, which in turn was primarily due to a reduced success of reaching later stages of courtship. The time spent in copulation by hybrid males was intermediate between the two parental males. We studied aggression by observing the interactions of males of heterotypic pairs, both between the parental species and between the hybrids and parental males. A lack of aggressive interactions betweenD. silvestris males andD. heteroneura males in addition to the lack of courtship suggests thatD. silvestris males do not respond toD. heteroneura individuals of either sex. Hybrid males were equally successful in winning fights with bothD. silvestris andD. heteroneura males. These results indicate that the behavioral isolation betweenD. silvestris andD. heteroneura may be largely a consequence of the earliest stages of interactions. The two species may differ either in activity levels or in morphological or chemical traits that are important for species and mate recognition. The relatively high copulation and aggressive success of hybrids indicates that sexual selection against hybrids alone is unlikely to be a sufficient force to reduce gene flow and maintain species distinctions.     

A Genetic Basis for the Inviability of Hybrids between Sibling Species of Drosophila

A mutation of Drosophila melanogaster whose only known effect is the rescue of otherwise lethal interspecific hybrids has been characterized. This mutation, Hmr, maps to 1-31.84 (9D1-9E4). Hmr may be the consequence of a P element insertion. It rescues hybrid males from the cross of D. melanogaster females to males of its three sibling species, D. simulans, D. mauritiana and D. sechellia. This rescue is recessive, since hybrid males that carry both Hmr and a duplication expected to be Hmr+ are not rescued. Hmr also rescues the otherwise inviable female hybrids from the cross of compound-X D. melanogaster females to males of its sibling species. This rescue is also recessive, since a compound-X heterozygous for Hmr does not rescue. Another mutation, discovered on the In(1)AB chromosome of D. melanogaster, is also found to rescue normally inviable species hybrids: unlike Hmr, however, In(1)AB rescues hybrid females from the cross of In(1)AB/Y males to sibling females, as well as hybrid males from the cross of In(1)AB females to sibling males. These data are interpreted on the basis of a model for the genetic basis of hybrid inviability of complementary genes.     

The evolution of gonad expenditure and gonadosomatic index (GSI) in male and female broadcast‐spawning invertebrates

Sedentary broadcast‐spawning marine invertebrates, which release both eggs and sperm into the water for fertilization, are of special interest for sexual selection studies. They provide unique insight into the early stages of the evolutionary succession leading to the often‐intense operation of both pre‐ and post‐mating sexual selection in mobile gonochorists. Since they are sessile or only weakly mobile, adults can interact only to a limited extent with other adults and with their own fertilized offspring. They are consequently subject mainly to selection on gamete production and gamete success, and so high gonad expenditure is expected in both sexes. We review literature on gonadosomatic index (GSI; the proportion of body tissue devoted to gamete production) of gonochoristic broadcast spawners, which we use as a proxy for gonad expenditure. We show that such taxa most often have a high GSI that is approximately equal in both sexes. When GSI is asymmetric, female GSI usually exceeds male GSI, at least in echinoderms (the majority of species recorded). Intriguingly, though, higher male GSI also occurs in some species and appears more common than female‐biased GSI in certain orders of gastropod molluscs. Our limited data also suggest that higher male GSI may be the prevalent pattern in sperm casters (where only males release gametes). We explore how selection might have shaped these patterns using game theoretic models for gonad expenditure that consider possible trade‐offs with (i) somatic maintenance or (ii) growth, while also considering sperm competition, sperm limitation, and polyspermy. Our models of the trade‐off between somatic tissue (which increases survival) and gonad (which increases reproductive success) predict that GSI should be equal for the two sexes when sperm competition is intense, as is probably common in broadcast spawners due to synchronous spawning in aggregations. Higher female GSI occurs under low sperm competition. Sperm limitation appears unlikely to alter these conclusions qualitatively, but can also act as a force to keep male GSI high, and close to that of females. Polyspermy can act to reduce male GSI. Higher male than female GSI is predicted to be less common (as observed in the data), but can occur when ova/ovaries are sufficiently more resource‐intensive to produce than sperm/testes, for which some evidence exists. We also show that sex‐specific trade‐offs between gonads and growth can generate different life‐history strategies for males and females, with males beginning reproduction earlier. This could lead to apparently higher male GSI in empirical studies if immature females are included in calculations of mean GSI. The existence of higher male GSI nonetheless remains somewhat problematic and requires further investigation. When sperm limitation is low, we suggest that the natural logarithm of the male/female GSI ratio may be a suitable index for sperm competition level in broadcast spawners, and that this may also be considered as an index for internally fertilizing taxa.     

Physiological effects of the lunar cycle on the spawning of a coral reef fish,Abudefduf Vaigiensis: in vivo and in vitro trait

The aim of the present study was to investigate the lunar cycle effects of the spawning of Audefduf vaigiensis through in vivo and in vitro analysis. For this purpose, the indices of GSI, serum levels of sex steroids, including testosterone (T), 17α-hydroxyprogesterone (OHP), 17α, 20β-dihydroxyprogesterone (DHP), and 11-keto-testosterone (11-KT) as well as the germinal vesicle breakdown (GVBD) were measured. The sampling pattern was weekly, based on the moon cycles as the new moon (NM), the first quarter (FQ), the full moon (FM), and the last quarter (LQ). In females, the highest in vivo values of the GSI index were obtained in FQ and LQ, and in males, this value was significantly higher in LQ than NM. The highest in vivo level of OHP in females was observed in FQ, whereas in males was obtained in FM. In both sexes, the in vivo serum levels of DHP were obtained in LQ. In males, the level of 11-KT were at the peak in NM. In vitro analysis showed the highest rate of GVBD in LQ. Moreover, the in vitro levels of T, OHP, and DHP were significantly higher in LQ compare to NM in both sexes. However, in males, the in vitro levels of 11-KT was significantly higher in NM than LQ. These cyclical changes obtained from in vivo plasma steroid hormones and in vitro data on GVBD suggested that lunar periodicity is a major external regulator that synchronized ovarian and testicular activity of A. vaigiensis with emphasis on spawning phenomenon.

     

Production of fertile unreduced sperm by hybrid males of the Rutilus alburnoides complex (Teleostei,cyprinidae). An alternative route to genome tetraploidization in unisexuals.

The hybrid minnow Rutilus alburnoides comprises diploid and polyploid females and males. Previous studies revealed that diploid and triploid females exhibit altered oogenesis that does not involve random segregation and recombination of the genomes of the two ancestors, constituting unisexual lineages. In the present study, we investigated the reproductive mode of hybrid males from the Tejo basin, using experimental crosses and flow cytometric analysis of blood and sperm. The results suggest that diploid hybrids produced fertile unreduced sperm, transmitting their hybrid genome intact to offspring. Triploid hybrids also produced unreduced sperm, but it was not possible to obtain data concerning their fertility. Finally, tetraploid hybrids produced fertile diploid sperm, which exhibited Mendelian segregation. Tetraploid R. alburnoides may reestablish biparental reproduction, as individuals of both sexes with the appropriate constitution for normal meiosis (two haploid genomes from each parental species) are likely to occur in natural populations. Tetraploids probably have arisen from syngamy of diploid eggs and diploid sperm produced by diploid hybrid males. Diploid hybrid males may therefore play a significant role in the dynamics of the complex, starting the evolutionary process that may ultimately lead to a new sexually reproducing species.     

Reproduction characteristics of a stunted population of rudd, <Emphasis Type="Italic">Scardinius erythrophthalmus</Emphasis> (Linnaeus, 1758) living in the Anzali Lagoon (the southwest Caspian Sea,Iran)

A total of 486 specimens of Scardinius erythrophthalmus caught in the Anzali lagoon, a large lagoon located in the southwest Caspian Sea (Iran), between March and June 2007 were examined. Age determination showed that the maximum ages observed were 4+ in males and 5+ in females. Isometric growth was identified from length-weight relationship in males, females and pooled data. There was no significant difference from parity in the overall sex ratio of 252 males to 234 females. The fish spawn from mid April to late May, with peak spawning in mid May with the highest average GSI value of 7.12 and 13.52 for males and females respectively. The absolute fecundity ranged between 1482–59620 eggs with the mean of 9287.87 eggs while relative fecundity ranged from 127.8 to 1737.6 eggs/g with the average of 709 eggs/g of body weight. Egg diameter ranged from 0.43 to 1.23 mm with a mean of 0.73 mm. The characteristics of rudd in the stunted population from the Anzali lagoon differ markedly from those of other localities of its range.     

Post‐zygotic isolation in cactophilic Drosophila: larval viability and adult life‐history traits of D. mojavensis/D. arizonae hybrids

Drosophila mojavensis and Drosophila arizonae are cactophilic flies that have been used extensively in speciation studies. Incomplete premating isolation, evidence of reinforcement, and a lack of recent introgression between these species point to a potentially important role for post‐zygotic isolating barriers in this system. Other than hybrid male sterility, however, post‐zygotic isolation between D. mojavensis and D. arizonae has received little attention. In this study, we examined viability and life‐history traits of D. mojavensis/D. arizonae F₁ hybrids from sympatric crosses. Specifically, we reared hybrids and purebreds on the natural host cacti of each parental species and compared viability, development time, thorax length, and desiccation resistance between hybrids and purebreds. Interestingly, hybrid females from both crosses performed similarly or even better than purebred females. In contrast, hybrid sons of D. arizonae mothers, in addition to being sterile, had shorter average thorax length than males of both parental species, and hybrid males from both crosses had substantially lower desiccation resistance than D. mojavensis males. The probable cost to hybridization for D. mojavensis females resulting from reduced desiccation resistance of hybrid sons may have been an important selective factor in the history of reinforcement for crosses involving these females.     

Lipid and fatty acid content in wild white seabream (Diplodus sargus) broodstock at different stages of the reproductive cycle

The lipid and fatty acid content of the gonads, liver and muscle of wild white seabream males and females was studied at different stages of the reproductive cycle. Samples were taken from mature white seabream at pre-spawning (November), mid-spawning (March) and post-spawning (June) stages. The results showed that lipid accumulates in gonads and muscle from November to March. The gonadosomatic index (GSI) was also increased during this period. Male gonads showed a greater increase in polar lipid (PL) than neutral (NL), while female gonads displayed the reverse. The increase in both neutral and polar lipid was higher in the muscle of males than in females. In the same period, male livers showed no changes either in lipid content or the hepatosomatic index (HSI), while female livers registered an increase in both lipid content and HSI. Between March and June, in both males and females, total, neutral and polar lipid decreased sharply in the gonads and muscle. Muscular lipid content reduction was more pronounced in males than females. On the other hand, the lipid content of the liver in males and females remained relatively constant. In general terms, the amounts of major fatty acids (16:0, 18:1n-9, 20:5n-3 and 22:6n-3) in gonadal and muscular polar and neutral lipid in both males and females increased from November to March and declined thereafter. Variations of the liver fatty acid content were less extreme. In the period from mid-spawning to post-spawning, the presence of 20:4n-6 in polar and neutral lipid increased to a notable extent in all organs studied.     

Behavioural isolation may facilitate homoploid hybrid speciation in cichlid fish

Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental species. In the absence of post‐mating isolation due to structural changes in the genome, or temporal or spatial premating isolation, another form of reproductive isolation would be needed for homoploid hybrid speciation to occur. Here, we investigate the potential of behavioural mate choice to generate assortative mating among hybrids and parental species. We made three‐first‐generation hybrid crosses between different species of African cichlid fish. In three‐way mate‐choice experiments, we allowed hybrid and nonhybrid females to mate with either hybrid or nonhybrid males. We found that hybrids generally mated nonrandomly and that hybridization can lead to the expression of new combinations of traits and preferences that behaviourally isolate hybrids from both parental species. Specifically, we find that the phenotypic distinctiveness of hybrids predicts the symmetry and extent of their reproductive isolation. Our data suggest that behavioural mate choice among hybrids may facilitate the establishment of isolated hybrid populations, even in proximity to one or both parental species.     

Gonadal development and spawning cycle in the digynic hermaphrodite sharpsnout seabream Diplodus puntazzo (Sparidae) off the Canary Islands, northwest of Africa

Diplodus puntazzo is a species characterized by a rudimentary hermaphroditism, with a low level of protandry. The study found that the overall sex ratio was one male to 2.03 females, diverging significantly from 1 : 1, and that there was a ratio of one male to 0.11 transitional fish. The sex ratio varied with season: during the reproductive period the ratio was 1 : 1.86 (male : female), significantly lower than in the non‐reproductive period in which the sex ratio was approximately 1 : 2.26. Both sexes had similar size distributions; however, females were predominant in all size intervals, especially in the larger ones. No significant difference in mean total length (TL) between females (330 mm), males (319 mm) or transitionals (321 mm) was found. The smallest functional female and male analysed were 159 and 157 mm TL, and the largest were 520 and 510 mm TL, respectively. No significant difference in length at first level of maturity, attained at the second year of life, was found between males (281 mm) and females (292 mm), respectively. A protracted spawning season was identified for the species: from September to February, with a peak in November. GSI values followed the same pattern for males and females, with values of females being similar to those of males; the highest values occurred between September and December, with a maximum in November. Occurrence of transitional fish was significantly higher during the month with highest values of GSI than in the resting period. The study found that vitellogenesis did not begin synchronously for all females, and the simultaneous presence of oocytes in all stages of development indicated an asynchronous mode of ovarian development. The presence of postovulatory, together with tertiary yolk vesicle oocytes, indicated that the species is a multiple spawner.