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 biology of the endangered Oxleyan pygmy perch Nannoperca oxleyana Whitley

The reproductive biology of the Oxleyan pygmy perch Nannoperca oxleyana is described from simultaneous studies of wild populations in north-eastern New South Wales and mature fish held in aquaria. In the wild, 50% of males and females matured at total lengths of 24·0–25·9 and 28·0–29·9 mm, respectively. The species displays sexual dichromatism during the spawning season, with males developing more intense red and brown fin and body colouration, and black pelvic fins. Captive male broodfish displayed territoriality during the breeding season, closely guarding sites within artificial, plant-like substrata in which pairs of fish spawned adhesive eggs. Protracted serial spawning of wild and captive fish occurred from September to April and May at mean water temperatures ≥16·6° C and day length ≥10·7 h. Captive broodfish spawned on an average of 57% of days during the 256 day spawning period. Gonado-somatic indices averaged 0·7% for all ripe males and 4·1–4·2% for all ripe females collected. Mean total and batch fecundities of captive females were 1323 eggs per fish and 7·8 eggs per fish per day, respectively, and relative fecundity was 587 eggs g⁻¹ of body mass. Batch fecundity of wild females was estimated at 7·8 eggs per fish. The adaptive significance of this reproductive strategy in a harsh, variable environment is discussed.     

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.     

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.     

Reproductive biology of gag in the southern Gulf of Mexico

Aspects of the reproductive biology of gag Mycteroperca microlepis in the southern Gulf of Mexico were studied by following seasonal variations in the gonado‐somatic index and through histological examination of gonads. Gag were collected from inshore and offshore waters of the Campeche Bank, Yucatan, Mexico, between April 1996 and December 2001. This species is a protogynous hermaphrodite, and appeared to be depth‐size distributed. The smallest gag (9–49 cm L _F) collected were all juvenile females, and were caught in inshore waters (1–10 m depth), while the largest (49–116 cm L _F), mainly adult females, males and transitionals, were captured in offshore waters (33–167 m depth). Overall the offshore male to female ratio was female‐biased (1 : 3·3) and differed significantly from unity. The species spawns at depths of c . 50–53 m, from early winter to mid‐spring, with peak spawning activity occurring between January and March. Fifty per cent of females reached first maturity at 72·1 cm L _F. At 103 cm L _F, 50% of sampled females had changed into males. Gag can be considered a monandric species, and sexual transition for this grouper seemed to occur in fish distributed within a narrow size range (85–111 cm L _F). The results are compared with those of other authors for gag stocks from the south‐east Atlantic coast of the U.S.A. and the north‐east Gulf of Mexico.     

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.     

Reproductive cycles in a reared strain of the mummichog, a daily spawner

Annual, lunar, and diel samplings were taken from a strain of mummichog (Arasaki strain) reared in outdoor tanks under natural conditions, to examine gonadal maturity. Gonads of yearling fish were quite immature in September. During late autumn and winter, a gradual increase in the GSI of both sexes was observed, and the growth of cortical alveolus phase oocytes in females and basal spermatogenesis in males progressed. In late February, a rapid increase in the GSI of both sexes, vitellogenesis in females, and active spermatogenesis in males, occurred. The spawning period of the yearling fish was from late March to August judging from the presence of milt-producing males and ovulated females. The spawning period of the underyearling fish started in the same month as the yearlings, but terminated 1 month earlier. Plasma levels of oestradiol-17 β (E₂) in females and testosterone in males were high during the spawning period in the yearlings. In the underyearlings, however, E₂ levels peaked in early spring, and declined in the latter part of the spawning period. Neither a lunar nor semilunar cycle was evident in the reproductive activity of this fish, which proved to be a typical daily spawner. Females showed an apparent daily reproductive cycle; oocyte maturation commenced at about 1200 hours, germinal vesicle breakdown (GVBD) occurred at about 2400 hours, and ovulation was completed by 2400 hours, 24 h after GVBD. Such clear annual and daily reproductive cycles make this strain of mummichog a suitable model for the study of environmental and endocrine regulation of reproductive cycles in marine and estuarine teleosts.     

Plasma non-esterified fatty acid profiles in wild Atlantic salmon during their freshwater migration and spawning

Total plasma non-esterified fatty acid (NEFA) levels increased significantly in adult Atlantic salmon during the first months of their upstream migration and spawning in the Exploits River, Newfoundland, Canada. The highest levels occurred in May and were 5467±270·43 nmol ml⁻¹ for females and 4617±334·70 nmol ml⁻¹ for males. Significantly higher levels were maintained by females compared with males for most of the upstream migration. Between August and October, total plasma NEFA levels declined by 61% in females but only 23% in males. The decline in plasma monounsaturated and polyunsaturated fatty acid levels accounted for 74% of the loss of NEFAs in females. Specific plasma NEFAs such as 16: 0 (palmitic), 16: 1 (palmitoleic), 18: 1n9 (oleic) and 20: 5n3 (eicosapentaenoic acid) differed significantly between males and females during migration and spawning. The mean gonadosomatic index ( I _G) values of females in May and just prior to spawning were 0·37±0·01 and 10·25±0·32, respectively. The rapid decline in the plasma NEFA content of females coincided with the largest increase in their I _G (1·85±0·02–10·25±0·32). Corresponding I _G values for males were 0·34±0·01 in May and 3·33±0·78 prior to spawning. Plasma NEFA levels of spent salmon did not differ between sexes and were significantly lower than those of salmon preparing to spawn.     

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.     

Differences in size and growth rates of male and female migrating Japanese eels in Pearl River, China

The Sr/Ca ratios in otoliths of silver Japanese eels Anguilla japonica , in Pearl River, China, indicated that both sexes did not stay in brackish water and grew in fresh water from the glass eel stage until spawning migration. This did not support the hypothesis that females tended to distribute upstream and males might be restricted to estuaries. The back-calculated total length of males at glass eel stage was not significantly different from that of females, indicating that the hypothesis that small glass eels became males and larger ones became females may not be true. The mean (±S.D.) age and total length of males at migration were 6·4±1·6 years and 48·3±4·5 cm, which were significantly smaller than for females, 8·3±1·6 years and 61·4±4·1 cm. The age of migration was related inversely to growth rate for both sexes. Growth parameters of the von Bertalanffy growth equation were K =0·21 cm year°1, L _∞=55·7 cm and t _o=-0·55 year for males and K =0·14 cm year⁻¹, L _∞=77·5 cm and t _o=-0·60 year for females. The difference in asymptotic length ( L _∞) between males and females may be because females postpone migration to achieve larger size for maximizing reproductive success.     

On the life history of the lesser gurnard (Scorpaeniformes: Triglidae) inhabiting the Agulhas Bank, South Africa

Growth analysis based on sectioned sagittal otoliths revealed the lesser gurnard Chelidonichthys queketti on the Agulhas Bank to be relatively fast growing and long lived, with ages of up to 7 years being recorded. Total length at age (mm) was described best by the specialized von Bertalanffy growth model as L _T=306·1 (1 − e^{0·53(t+0·18)}). First approximations of total, natural and fishing mortality rates were determined at 0·73, 0·38 and 0·35 year⁻¹ respectively. The adult population was male dominated with a sex ratio of 1 female: 1·2 males with the mean size of males and females being similar. The lesser gurnard is an iteroparous species with females maturing by the end of the first year of life (195 mm L _T), thereafter spawning throughout the year with reproductive activity peaking over spring and late summer. The lesser gurnard appears to exhibit similar life-history patterns to other triglid species in that it can be classified as a generalist. Generalistic life-history characteristics such as a fast growth rate, early sexual maturity at a relatively large size, a non-seasonal spawning pattern, feeding on a variety of prey organisms and the ability to inhabit various substrata could all contribute to it maintaining a high biomass on the Agulhas Bank.     

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.     

The reproduction of anglerfish Lophius piscatorius Linnaeus from the north-west coast of Scotland

The reproductive cycle of anglerfish was studied by examining fresh material obtained from commercial landings and research vessel surveys. The general morphology and histology of the gonads are described. This species has unusual gonads. The testes are tubular, and beanshaped in transverse section. The maturation process of the testis is similar to that of most other teleosts. The ovaries are confluent and form a single, flattened tube. During maturation, the width and, in particular, the length of the ovaries increases dramatically. A gelatinous matrix is produced during the maturation process, which contributes to the enormous size and weight of ripe ovaries. Ripe testes can be found throughout the year, whereas ripened ovaries were found only between November and May. Females outnumbered males in length classes greater than 70 cm and no males larger than 90 cm were found. The estimated mean length at maturity (1_50%) was 73 cm for females and 49 cm for males. Because only the large female anglerfish are capable of spawning, and because virtually all large anglerfish are female, trends in their abundance resulting from commercial exploitation should be monitored with care.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

Reproductive biology of Carcharhinus acronotus in the coastal waters of South Carolina

The reproductive biology of blacknose sharks Carcharhinus acronotus in the western North Atlantic Ocean was studied by examining specimens collected in the coastal waters of South Carolina. Males begin the maturation process between 875 and 910 mm fork length ( L _F), as indicated by the presence of functional claspers and siphon sacs. The presence of vitellogenic oocytes and developing oviducal glands and uteri indicated that females begin to mature at c . 870 mm L _F. Length at which 50% of the population reached maturity was 896 and 964 mm L _F, equivalent to 4·3 and 4·5 years, for males and females, respectively. Gonado‐somatic indices suggested that spermatogenesis and vitellogenesis began after December. Mating took place during the end of May and the beginning of June. Fertilization occurred during late June and early July, suggesting that female blacknose sharks were capable of sperm storage. Based on the timing of fertilization and occurrence of females carrying near‐term pups in late May and early June, the gestation period for blacknose sharks was c . 11 months. Female blacknose sharks reproduced biennially based on the absence of vitellogenic oocytes in near‐term females and there being no indication of vitellogenesis in postpartum females. Male blacknose sharks were capable of reproducing annually as indicated by turgid genital ducts, which were observed in all mature males collected during late May and early June.     

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.     

Reproductive investment in the Silurus meridionalis

A comparison of pre- and postspawning Silurus meridionalis showed that 20·7% of body stored energy was utilized during spawning for a standard male (74·5 cm) and 23·8% for a standard female (85·3 cm). About one-third of the loss of the stored energy was released as eggs by females, and almost all of the energy loss for males and about two-thirds for females were expended in metabolism. Stored lipid as fuel for metabolism supplied 90·0% of energy in males and 95·2% in females, and protein supplied the rest of the energy. Models for predicting energy in released gametes ( G _g), deposited in the body as somatic growth ( G _s), utilized in spawning activity ( S _a), expended in maintenance ( M , including metabolism, faeces and excretion), and food energy ( C ) were developed, and annual energy budgets were compiled. The balanced budget for a male aged 4 was: 100 C =0·06 G _g+11·17 S _a+19·5 G _s+69·2 M , and for a female aged 5: 100 C =5·48 G _g+8·51 S _a+15·8 G _s+70·2 M .     

Age and growth of the round stingray Urobatis halleri at Seal Beach, California

The age and growth of the round stingray Urobatis halleri was determined using vertebral sections from animals collected at Seal Beach, California from 2002 to 2005. Annual periodicity was validated from U. halleri injected with oxytetracycline and maintained in captivity over a 2 year period ( n = 7). The coefficients estimated by the von Bertalanffy growth model were the disc width asymptote ( W _D∞) (286 mm for males and 224 mm for females) and K (0·09 year⁻¹ for males and 0·15 year⁻¹ for females). The age structure of the population consisted of mostly older, mature males and females. Age at maturity was estimated at 3·80 years for females and 3·75 years for males, and the maximum assessed age was 14 years old. Males were more numerous than females throughout the year; however, from May to September, females outnumbered males. The U. halleri age and growth coefficients were comparable to other species in the family Urolophidae. Based on the seasonality and age structure of this population, Seal Beach offers warm-water refuge for U. halleri of reproductive maturity, and the U. halleri at Seal Beach may garner some behavioural thermoregulation benefit.     

Reproductive strategy of a primitive temperate notothenioid Eleginops maclovinus

The reproductive biology of Eleginops maclovinus was examined in the Falkland Islands between October 2000 and December 2002. Males predominated at total lengths ( L _T) of 10 to 52 cm and females at >53 cm L _T. Length frequency analysis showed a bimodal distribution with females representing the larger mode for every month during the study period. Gonad histology revealed that 19% of the histological samples studied were considered to be those from hermaphrodites: morphologically as male gonads but containing protoplasmic oocytes. It was therefore concluded that E. maclovinus is a protandrous hermaphrodite. The size of first (male) maturation of E. maclovinus was 30·73 cm L _T. Males and females matured from August onwards and spawning occurred between September and December at depths of >40 m. Eleginops maclovinus has the smallest eggs and highest fecundity among the notothenioids. The highest potential fecundity was attained at maturity stage III with c . 48 million eggs. Because of further oocyte resorption, this value gradually decreased until the final fecundity ranged from 1·1 to 7·3 million eggs. Oocyte length frequencies in ovaries suggested that E. maclovinus was a batch spawner. Hydrated oocytes were found to contain a large perivitelline space indicating that the egg had a pelagic development.     

Age, growth and reproduction of the barrelfish Hyperoglyphe perciformis (Mitchill) in the western North Atlantic

Otoliths ( n = 847) and gonads ( n = 817) were collected from barrelfish Hyperoglyphe perciformis that were captured by commercial fishermen in the waters off South Carolina and Georgia in 1995, 1997 and 2001–2006. Of the otoliths collected, 97% were aged successfully, and specimens sampled ranged from 5 to 85 years, with a median age of 12 years. The von Bertalanffy growth parameters yielded the equation: L_t = 857·8{1 − e^{−0·0985[ t −(−8·95)]}}, where L_t is fork length ( L _F) at time t . Through histological examination, 94% of the gonads assessed were assigned to a sex and reproductive class. Females spawned from September to May with a peak from November to January. Males spawned year round, but had a peak from September to April. The sex ratio (M:F) for this population was 1:1·34. The smallest mature female was 605 mm L _F and the youngest immature female was 697 mm L _F. Estimates of L _F and age at 50% maturity ( L ₅₀ and A ₅₀) for females were 660 mm L _F (95% CI = 633–667 mm L _F) and 6·08 years (95% CI = 3·50–7·27 years), respectively. The youngest mature male was 575 mm L _F and the oldest immature male was 762 mm L _F, and no estimates of L ₅₀ or A ₅₀ were made for males. It was determined that barrelfish exhibit the typical characteristics of long life span, slow growth and high age at maturity seen in other deepwater fishes, and that care should be taken to manage this species accordingly.