Reproductive cycle and sex inversion in razor fish, a protogynous labrid in the southern Mediterranean Sea

The reproductive biology of the Mediterranean razor fish Xyrichthys novacula was investigated by demographic data and histological analysis of the female, intersexual and male gonads. Specimens were collected by bottom trawl on a monthly basis between June 2000 and July 2001 in a sandy bay in southern Thyrrenian. Gonad histology confirmed that the Mediterranean razor fish is a monandric, protogynous hermaphrodite. Females reached first sexual maturity at 100 mm ( L _T) and the estimated mean L _T at first maturity ( L ₅₀) was 125 mm. Females exhibited asynchronous ovarian development and multiple ovulations occurred over the spawning period. Vitellogenesis started in early May and spawning occurred from late May until late September. Sexual transition involved a large‐scale atresia of all oocyte stages and a massive degeneration of ovarian tissue followed by primordial germ cells proliferation. Sex change began at spawning time (June) but transitional individuals tended to cluster at the end of the reproductive period (September). They accounted for 17·1% of the population sampled and were found in a broad size range (105–150 mm L _T).     

Reproductive cycle and sexual maturity of the anglerfish Lophiomus setigerus in the East China Sea with a note on specialized spermatogenesis

The reproductive cycle and sexual maturity of the anglerfish Lophiomus setigerus were examined. Spermatids were released from the germinal cysts into the lumina of the seminal lobules, and both spermatids and spermatozoa were present in the lumina of the seminal lobules and sperm ducts. Spermatogenesis and vitellogenesis occurred throughout most of the year. The testes of males were full of spermatozoa throughout the year, with spawning from May to November. Males and females reached sexual maturity at a mean total length and age of 178 mm, 3.3 years and 303 mm, 6.1 years, respectively. There were clear seasonal cycles in the gonadosomatic index (I_G) and hepatosomatic index ( I _H) in females. The mean I_G of females increased rapidly with ovarian development while the mean I _H decreased from the middle of vitellogenesis to final maturation. Mean values of I _G and I _H in males increased with testicular development.     

Effects of fish size on spawning time in Norwegian spring-spawning herring

During October-January in a northern Norwegian fjord system and in February-March at the main western Norway spawning grounds, an index I _G of gonad weight of spring-spawning herring Clupea harengus in percentage of expected gonad weight at full maturity for a given total length L _T, tended to increase with L _T in the range 27–31 cm as the proportion of recruit spawners decreased. Insignificant differences in I _G were found between L _T groups in the range 32–37 cm (repeat spawners). This is contrary to that suggested in other studies and signifies that generally the stock spawns in only two waves, repeat spawners first and the recruit spawners second. I _G also increased with somatic and total condition factor, which signifies that spawning time may be influenced by summer feeding conditions.     

Reproductive biology and implications for management of the orange-spotted grouper Epinephelus coioides in the southern Arabian Gulf

The reproductive biology of Epinephelus coioides was determined from the examination of 1455 individuals collected between July 2005 and June 2007 in the southern Arabian Gulf. Histological preparations of gonads indicated that males were either derived from a juvenile phase or the transition of postspawning females, confirming a diandric protogynous sexual pattern. The spawning season was well defined, occurring once a year during April and early May. Peaks in spawning occurred after the full and new moons and was completed within a single lunar cycle. The presence of mature males over the entire size and age range and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. While specimens undergoing sexual transition were only observed in size and age ranges of 335–685 mm total length ( L _T) and 5–6 years, patterns in the proportion of males in size and age classes suggested that sex change occurred at a relatively constant rate after female maturation up to the maximum size (1002 mm L _T) and age (11 years). Relationships between reproductive output and capacity with size and age indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of E. coioides . The maximum age, small size and young age at sexual maturation ( L _min= 320 mm L _T, 2 years, for females and 242 mm L _T, 1 year, for males) conflict with the general pattern for large epinepheline groupers and may be a direct result of the intensive demersal fishery in the southern Arabian Gulf.     

Life-history traits of introduced Iberian pumpkinseed Lepomis gibbosus relative to native populations. Can differences explain colonization success?

To assess the colonization success of pumpkinseed Lepomis gibbosus on the Iberian Peninsula, life-history attributes of pumpkinseed populations from Spanish water bodies were compared to populations in the northern and southern parts of its native range, as well as to those of English water bodies where introduced populations have shown minimal natural range expansion. Discriminate function analysis using five population characteristics [mean age at maturity, mean total length ( L _T) at maturity, gonado-somatic index ( I _G), mean L _T at age 2 years as an indication of juvenile growth rate and relative body condition] strongly differentiated populations from the four regional study areas. Spanish populations were early maturing, showed moderate juvenile growth rate and L _T at maturity and high I _G relative to the other population groups. Spanish populations matured significantly earlier than Canadian and English populations, but not southern U.S.A. populations. Spanish populations, however, had a significantly higher I _G than southern U.S.A. populations, suggesting greater reproductive output. Considering these differences in the context of the Winemiller and Rose triangular life-history strategy model, the pumpkinseed exhibits 'equilibrium' life-history traits that have made other non-native fishes successful invaders in areas where habitat alteration has resulted in more predictable discharge regimes and water levels. Populations in Iberia, however, appear to have achieved their success by adopting a more 'opportunistic' life-history strategy than their native counterparts. High levels of life-history plasticity appear to contribute to the success of this species on the Iberian Peninsula.     

Reproductive biology of the smooth-hound shark Mustelus mustelus (L.) in the Gulf of Gabès (south-central Mediterranean Sea)

The reproductive biology of the smooth-hound shark Mustelus mustelus was studied in the Gulf of Gabès (southern Tunisia). Females were found to mature between 1075 and 1230 mm total length ( L _T) whereas males matured between 880 and 1120 mm L _T. The L _T at which 50% of the population reached maturity was 971 and 1172 mm for males and females, respectively. Male gonads were symmetrical in terms of mass and both functional, whereas in females only the right ovary was functional. The seasonal changes in the oocytes and testes development, embryo length and the occurrence of near-term and post-partum females showed that this species displayed a clearly defined annual reproductive cycle with parturition occurring during late April and early May, after a gestation period of 10–11 months. Mating took place during May and early June and fertilization occurred from early June to early July. The embryo sex ratio was not significantly different from unity. Litter size varied from four to 18 embryos and was positively correlated with maternal L _T. The young were born with a L _T of 340–420 mm.     

Age and size compositions, growth and reproductive biology of the breaksea cod Epinephelides armatus, a gonochoristic serranid

Details of the reproductive biology, size and age compositions and growth of the breaksea cod Epinephelides armatus , the sole representative of Epinephelides , were obtained by collecting monthly samples of a wide total length ( L _T) range of individuals from coastal marine waters at 31–32° S on the lower west coast of Australia. Although the modal L _T class of females (250–299 mm) was markedly less than that of males (400–449 mm), the modal ages of the two sexes were similar, i.e. 4 v. 5 years, respectively. The similarity in the age compositions and the histological demonstration that the gonads of all E. armatus consist solely of either ovarian or testicular tissues demonstrate that this species is gonochoristic, which is highly unusual for an anthiinine serranid. The absence of a central, membrane-lined 'ovarian' lumen in the testes of juveniles would account for adult testes containing neither this ovarian remnant nor the peripherally located sperm sinuses that are found in the mature testes of almost all other serranids. The results demonstrate that E. armatus exhibits a very unusual pattern of sexual development for a serranid. The spawning period of E. armatus lasts for c. 9 months, which is long for a species in temperate Western Australian waters, but comparable with that of many other relatively small serranids elsewhere. Females grow slower than males, attaining L _T at 3, 5 and 10 years of c. 200, 285 and 420 mm, respectively, compared with c. 215, 315 and 450 mm, respectively. Females, however, attain maturity at a greater L _T and older age than males.     

Influence of seasonal changes in relative liver size, condition, relative gonad size and variability in ovarian development in multiple spawning fish species used in environmental monitoring programmes

Basic biological information was collected to assist in understanding the biology of selected small-bodied fishes for use in environmental monitoring programmes. Gonado-somatic index ( I _G) and hepato-somatic index ( I _H) profiles of female blacknose dace Rhinichthys atratulus , mummichog Fundulus heteroclitus and golden shiner Notemigonus crysoleucas were similar, decreasing gradually during the spawning season. I _G and I _H profiles of female northern redbelly dace Phoxinus eos peaked twice suggesting oocyte recruitment was continuous throughout the spawning season. Regressions of ovary mass to adjusted body mass ( M _A) were more variable in dace relative to the other fish species. For female blacknose dace, variability in the relationship between ovary mass and M _A was minimized by selecting either 2 year-old fish and fish that weighed 2–4 g.     

Ovarian morphology of the Japanese eel in Mikawa Bay

Annual changes in gonadal maturation of female Japanese eel Anguilla japonica in sea water were investigated histologically over 5 years in the Mikawa Bay, Japan, where they occurred throughout the year except in March. Almost all immature Japanese eels (yellow eels) occurred mainly from April to September, and they were rare after November. In contrast, maturing Japanese eels (silver eels) occurred from October to February. The gonado‐somatic index ( I _G) and oocyte diameters of yellow eels were <1·0 and 150 μm, respectively, and oocytes were at the peri‐nucleolus or the oil droplet stages. The I _G and oocyte diameters of silver eels were greater than those of yellow eels and most oocytes developed to the primary yolk globule stage. The numbers of silver eels lacking oocytes at the primary yolk globule stage increased after January in Mikawa Bay, although I _G and oocyte diameters remained unchanged. In contrast, silver eels caught at the mouth of the bay in January possessed oocytes that had advanced to the secondary yolk globule stage. These observations indicate that oocyte development changes seasonally, especially after winter in Mikawa Bay.     

Influence of female age, body size and environmental conditions on annual egg production of the bullhead

A population of bullhead Cottus gobio inhabiting a mountainous river network, was examined for the influence of maternal condition (total length, L _T, and age class in years) and environmental factors (water temperature and fish density) on fecundity, egg size and net annual reproductive effort ( E _R, defined as the ratio of total egg production mass to body mass) within several stocks. Regardless of location within the drainage network, bullhead reproduction was synchronous, with only a single batch and fecundity ranging from 69 to 896 eggs per female. As with L _T fecundity increased with age, although the difference between age classes 5 and 6 years was smaller than between other age classes. Egg diameter also increased with age from age classes 2 to 4 years, but then tended to decrease for age classes 5–6 years. Multiple stepwise regression was used to examine the effect of female L _T, water temperature and fish densities on bullhead egg production for each of the four age classes (2, 3, 4 and 5–6 years) separately. When statistically significant, water temperature always had a positive effect and fish densities a negative effect, on fecundity and egg size. The L _T always had a positive effect on fecundity for all the age classes. In contrast, L _T had a positive effect on egg size and E _R only for female reaching maturity (2 years), whereas intermediate aged females (3–4 years) were insensitive and the oldest fish showed a negative response.     

Using the composite variables of reproductive morphology, histology and steroid hormones to determine age and size at sexual maturity for the thorny skate Amblyraja radiata in the western Gulf of Maine

Age at size was linked to sexual maturity in 77 male and 93 female thorny skates Amblyraja radiata from the western Gulf of Maine using three criteria: 1) gross reproductive morphology, 2) histology and 3) steroid hormone concentrations. Age-bias plots and the coefficient of variation from vertebral band counts suggested that the ageing method represented a non-biased and precise approach to the age assessment of A. radiata . Maturity ogives for males, based on data gathered for clasper length, circulating testosterone concentrations and proportion of mature spermatocysts within the testes, predicted that 50% maturity occurred at a total length ( L _T) of 865 mm and c. 10·90 years of age. For females, maturity ogives, based on data gathered for ovary mass, shell gland mass, follicle size and circulating oestradiol concentrations, predicted that 50% maturity occurred at 875 mm L _T and c. 11·00 years of age. Collectively, the results suggest that analysis of several contemporaneous reproductive variables offers an accurate determination of sexual maturity in the thorny skate.     

Reproductive biology of Psammobatis extenta

Psammobatis extenta collected from the commercial bottom trawl fishery operating in Puerto Quequén, Argentina, between 2000 and 2001, had a sex ratio of 1:1 (535 specimens). Both the largest male and female measured 313 mm total length ( L _T). The relationship between total body mass and L _T was significantly different between sexes; mature females weighed more than mature males for a given L _T. The size at which 50% of males were sexually mature was 262 mm L _T (83·6% of the maximum size), whereas 50% of females were mature at 249 mm L _T (80·0% of the maximum size). The gonadosomatic index in mature males and females was slightly higher during summer, but did not show significant variation over the sampling period. The hepatosomatic index of males and females was higher during the autumn and winter and followed a distinct seasonal pattern. Evidence suggested a continuous reproductive cycle during the year, with a maximum number of females carrying egg-cases in summer.     

Morphological and histological changes in the swim bladder during maturation of the Japanese eel

Silver eels Anguilla japonica ( I _G 1·5–3·5) had more developed rete mirabile, gas gland and submucosa than yellow eels ( I _G 0·4–1·5) and the development of swim bladder components increased with sexual maturity only in the early maturation process ( I _G 3·5). These observations indicate that the Japanese eel develops its swim bladder in either the river or shallow sea water and leaves for the open sea after the swim bladder has become adapted to a deeper-sea environment.     

Reproductive biology of two sympatric skates in the south‐west Atlantic: Psammobatis rudis and Psammobatis normani

The total lengths ( L _T) at which 50% were mature of Psammobatis rudis and Psammobatis normani , in the south‐west Atlantic were: P. rudis , 428 mm for males and 414 mm for females and P. normani , 443 mm for males and 403 mm for females. Clasper length in mature males was greater in P. normani than in P. rudis , whereas oviducal gland width was not different between species. Females of P. normani with egg cases were found in every month sampled, and in January, March, April and July in P. rudis , although insufficient samples were available to identify peak oviposition times. Geographic variation in size frequency and maturity were found. The effects of oceanographic conditions and fishing pressure are discussed. Size at 50% maturity in both species was >74% of the maximum L _T, indicating late sexual maturity and low potential stock recovery rate.     

Age, growth, and reproductive biology of the blackbelly rosefish from the Carolinas, U.S.A.

Otoliths and gonads of blackbelly rosefish Helicolenus dactylopterus dactylopterus were collected from the commercial fishery off the Carolinas in 1994–1997. Opaque bands on transverse sections of otoliths were determined to be annuli by analysis of marginal increments. Opaque zone formation occurs between July and January. Ages ranged from 7 to 30 years. Blackbelly rosefish have intraovarian gestation. Fertilization is internal, as free spermatozoa were found primarily in resting ovaries from July through early December with peak occurrence in September through November. There was a delay of 1–3 months before fertilization, as oocyte development did not begin until December. Occurrence during January through April of early-celled embryos, the most advanced stage observed, and postovulatory follicles indicated that oocyte development was rapid. Egg development occurs in a clear gelatinous matrix secreted into the ovarian cavity. The reproductive mode is a zygoparous form of oviparity, intermediate between oviparity and viviparity. Population sex ratio departed markedly from 1: 1 for most length intervals. Males were more abundant at lengths >250 mm L _T and the overall male: female ratio was 1 : 0·60.     

Distribution, reproduction and feeding of the Greenland shark Somniosus (Somniosus) microcephalus, with notes on two other sleeper sharks, Somniosus (Somniosus) pacificus and Somniosus (Somniosus) antarcticus

The size and depth distribution, stomach contents and reproductive status of Somniosus ( Somniosus ) microcephalus, Somniosus ( Somniosus ) pacificus and Somniosus ( Somniosus ) antarcticus were examined from specimens collected from the North and South Atlantic, North and South Pacific, and the Southern Ocean. Specimens ranged in size from 42 to 480 cm total length, L _T, and were taken from depths of 35–1280 m. Stomach contents included coelenterates, gastropods, cephalopods, echinoderms, crustaceans, elasmobranchs, teleosts, penguins, marine mammals and human waste. Female S. ( S. ) microcephalus mature at c . 450 cm and S. ( S. ) antarcticus at c . 435 cm L _T; a female S. ( S. ) pacificus of 430 cm was mature. Male S. ( S. ) microcephalus mature at c . 300 cm, but male S. ( S. ) antarcticus may not mature until c . 400 cm L _T. The size at birth in these three species of Somniosus is c . 40 cm L _T.     

Feeding ecology of silver hake larvae on the Western Bank, Scotian Shelf, and comparison with Atlantic Cod

The feeding ecology of the larvae of silver hake Merluccius bilinearis was examined during two time periods (October 1998 and December 1992) on the Western Bank, Scotian Shelf, north‐west Atlantic, and compared with the feeding ecology of Atlantic cod Gadus morhua larvae collected in the same samples in December 1992. During both time periods silver hake exhibited strong selection for late stage copepodids and adult copepods at a small size (>3·5 mm total length, L _T). The niche width measured as the diet breadth index ( I _DB) of silver hake declined rapidly as they increased in size and remained relatively constant from 3 to 11 mm L _T, during each time period. Atlantic cod larvae exhibited a broader niche width that was curvilinear over the same L _T. Atlantic cod were also less selective than silver hake, incorporating both naupliar and early stage copepodids in their diets throughout the length classes examined. Simple isometric relationships did not explain the differences in diet, as Atlantic cod larvae continued to feed on early stages of copepods at large size, while silver hake larvae quickly switched to large prey items. The strong selection and narrow I _DB observed for silver hake probably reflects adaptation to spawning during the periods between major secondary production peaks in temperate waters.     

Reproductive biology of the milk shark Rhizoprionodon acutus and the bigeye houndshark Iago omanensis in the coastal waters of Oman

Female Rhizoprionodon acutus were found to mature between 62 and 74 cm total length ( L _T) whereas males matured between 63 and 71 cm L _T. The L _T at 50% maturity was 64·3 cm for females and 64·7 cm for males. Litter size varied from one to six embryos, and was positively correlated with maternal L _T. Female embryos outnumbered males by a ratio of 2·3:1. The size at birth was c . 37 cm L _T. Full‐term embryos and post‐partum females were observed during all seasons although their occurrence was highest in spring. Spermatozoa were rarely recorded in the oviducal gland, indicating that this species does not store sperm. It was not possible to generate maturity curves for Iago omanensis but it was evident that females matured by the time they reached 35 cm and males were mature by 31 cm L _T. This species displayed a clearly defined reproductive cycle with parturition occurring primarily in spring, after a gestation period of c . 1 year. Maximal embryo size was 19 cm L _T while maximal litter size was 24 embryos. The oviducal gland appeared to act as a seminal receptacle and it appeared that females may utilize these stores by not mating every year.     

Gape morphology of cod Gadus morhua L., haddock Melanogrammus aeglefinus (L.) and whiting Merlangius merlangus (L.) through metamorphosis from larvae to juveniles in the western Irish Sea

Variations in standard length ( L _S), gape size ( S _G) and jaw length ( L _J) were studied in larval and juvenile gadoids (cod Gadus morhua , haddock Melanogrammus aeglefinus and whiting Merlangius merlangus ) from 4 to 70 mm. The increase in S _G and L _J was not linear with respect to L _S. The relationship was best described by segmented regression lines in all three species, with an inflection point at c . 10·5 mm. The S _G and L _J increased more rapidly in relation to larval L _S for individuals smaller than this inflection point size. The rates of increase slowed significantly post-inflection, an effect more noticeable in S _G data compared to L _J data. In each case, the inflection point fell in the intermediate period of development between the larval and juvenile stages, which could be considered as metamorphosis. Published equations that have been used to predict S _G from L _J lead to the overestimation of gape. New relationships are presented, which may be used to predict S _G from measurements of either L _S or upper jaw length in cod, haddock and whiting.