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 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.     

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 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 variation among local populations of Atlantic cod from the Norwegian Skagerrak coast

Small‐scale spatial variation in life history was found among genetically distinct local populations of Atlantic cod Gadus morhua from the Norwegian Skagerrak coast. Among populations, age at 50% maturity varied from 2·6 to 3·8 years, total ( L _T) length at 50% maturity from 35 to 60 cm, annual survival from 33 to 64%, mean L _T at age 4 years from 43 to 63 cm, and mean backcalculated L _T at age 1 year from 8 to 12 cm.     

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.     

Reproductive characteristics and population decline of four species of skate (Rajidae) off the eastern coast of Canada

Four of the most common species of skate (Rajidae) were studied off eastern Canada to determine if their reproductive characteristics were linked to their population trajectories. The fecundity of the winter skate Leucoraja ocellata , the little skate Leucoraja erinacea , the thorny skate Amblyraja radiata and the smooth skate Malacoraja senta averaged between 41 and 56 egg cases per year for each species. For all species but L. ocellata , males matured at larger sizes and at later ages than females. Theoretical rates of population increase for non-equilibrium populations of L. ocellata ( c . 0·07), M. senta ( c . 0·14) and L. erinacea and A. radiata ( c . 0·20) were low compared to most fishes, indicating that north-west Atlantic skates are intrinsically unproductive, yet are theoretically capable of supporting low-level fisheries. Nevertheless, the results of 36 years of research surveys indicate that the abundance of mature L. ocellata , A. radiata and M. senta all decreased by >90% since 1970, indicating that past fishing mortality (both directed and undirected) has outstripped the net productivity of the skate populations on the eastern Scotian Shelf. The relationship between maximum age ( t _max) and age of maturity ( t _mat) was a better predictor of population growth rate than was body size, with the species exhibiting the highest ratios of t _mat : t _max ( L. ocellata = 0·68, M. senta = 0·66) having the lowest predicted population growth rates. L. ocellata appears to have the lowest productivity and has experienced the greatest population decline, thus raising concerns over its future status.     

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.     

The reproductive biology of Scyliorhinus canicula in the Bristol Channel, U.K.

Among 972 Scyliorhinus canicula , the overall sex ratio (males:females) was 1:1·23. Males matured between 490 and 540 mm L _T with 50% maturity at 520 mm. Females matured between 520 and 640 mm L _T with 50% maturity at 550 mm. Female gonadosomatic index peaked in May. Fecundity increased with fish length. The egg laying season lasted 10 months with a peak in June and July. The importance of erect, sessile invertebrates such as bryozoans, poriferans and hydroids as egg-case deposition sites has been established. Recently hatched fish were born in an equal sex ratio and at a mean length of 100·5 mm and mean weight of 3·07 g. One intersexual fish is described.     

Osteological development of the vertebral column and of the caudal complex in Dentex dentex

Osteological development of the vertebral column and caudal complex in common dentex was described under extensive larval rearing conditions. Generally, the cartilaginous bones developed prior to the membranous bones. The development of the axial skeleton began with the formation of the hypural 1, the neural arches 2 and 3, as well as the haemal arches 1–8 at 4·8, 4·9 and 5·0 mm total length ( L _T, measured in vivo ), respectively. By 7·5 mm L _T, all the cartilaginous elements were formed, except for the ventral ribs, which formed between the range of 8·4–18·0 mm L _T. The caudal lepidotrichia were the first membranous bones to appear (5·3 mm L _T) and attain their full meristic count (7·4 mm L _T), followed by the vertebral centra, which formed between 6·6 and 9·7 mm L _T. By 25·0 mm L _T, all the elements were fully ossified except for the ventral ribs. The developmental direction and order of all the elements were studied with respect to their formation and ossification. The results were discussed in the contexts of ichthyoplankton, ecology and aquaculture. Compared with other Sparidae species, common dentex followed a pattern of relatively rapid rate of osteological development.     

Sexual and geographical variation in life history parameters of the shorthorn sculpin

A total of 293 shorthorn sculpins Myoxocephalus scorpius from Tromsø, northern Norway, were sampled between November 1998 and April 1999 to determine sex, total length, age, growth, maturity and mortality. Females grew to larger sizes ( L _∞=26·9 v. 18·5 cm), matured later (2 v. 1 year of age) at larger size (maturation length=16 v. 14 cm L _T), and had lower instantaneous mortality rates (0·93 v. 1·20 year⁻¹) than males. The life history parameters of shorthorn sculpins in northern Norway were more similar to the parameters of short-lived central European populations than to the parameters of the long-lived population of Newfoundland. This study confirms that northern Norwegian shorthorn sculpins exhibit sexual dimorphism as in other shorthorn sculpin populations. The relationships between growth pattern, age at maturity and mortality rates observed in the Tromsø population and in other shorthorn sculpin populations, correspond well with the predictions from a published life history model.     

A histological study on the development of the digestive system of Pseudosciaena crocea larvae and juveniles

The ontogenetic development of the gut and accessory organs in large yellow croaker Pseudosciaena crocea was investigated using light microscopy from hatching up to the juvenile stage (40 days post hatch, dph). At 3 dph (mean ±  s . d ., 4·1 ± 0·1 mm total length, L _T), coinciding with the buccopharynx opening, larvae started to feed exogenously, and the gut consisted of a well‐developed buccopharynx, a partially‐differentiated oesophagus and an intestine divided in three regions (anterior intestine, intermediate intestine and rectum). Yolk reserves were not completely depleted at the onset of exogenous feeding, and a period of mixed nutrition was observed up to 6 dph (4·3 ± 0·1 mm L _T), when yolk was definitively exhausted. Important morphological changes occurred at the end of the larval period, coinciding with metamorphosis. At 17 dph (6·8 ± 0·6 mm L _T), pyloric caeca differentiated at the junction of the pyloric stomach and the anterior intestine. Gastric glands were first observed at 21 dph (9·2 ± 1·2 mm L _T), coinciding with the morphological development of the stomach in three different regions (cardiac, fundic and pyloric) according to the histological characteristics of their mucosa. At this age, large longitudinal folds appeared in the median and posterior oesophageal mucosa. These morphological and histological features suggested the achievement of a digestive system characteristic of large yellow croaker juveniles and adults.     

A comparison of the maturation and growth of female flathead sole in the central Gulf of Alaska and south‐eastern Bering Sea

Female flathead sole Hippoglossoides elassodon maturity appears not to be area dependent since the total length ( L _T) at which 50% were mature ( L _T50) was similar for the central Gulf of Alaska (333 mm) and south‐eastern Bering Sea (320 mm) areas. Likewise the age at which 50% were mature ( A ₅₀) was similar in the south‐eastern Bering Sea (9·7 years) and central Gulf of Alaska (8·7 years). The timing of female flathead sole spawning may also be similar between areas. Batch or serial spawning was indicated for flathead sole. Female flathead sole grew at a similar rate in both the Gulf of Alaska and Bering Sea. In contrast, males grew faster in the Bering Sea than in the Gulf of Alaska. Males grew more slowly than females in both areas after 5 years of age, and reached a smaller maximum L _T. The growth of both sexes was similar during 1993 and 1996 in the Gulf of Alaska.     

The reproductive biology and ecology of the Port Jackson shark Heterodontus portusjacksoni in the coastal waters of eastern Australia

The reproductive biology and ecology of the Port Jackson shark Heterodontus portusjacksoni was investigated at three locations on the central and southern coast of New South Wales (NSW), Australia from January 2002 to December 2005 using underwater visual census surveys and samples obtained from a commercial fishery. Adults displayed sexual dimorphism in total length ( L _T) at sexual maturity, with males maturing between 762 and 772 mm L _T and females between 902 and 905 mm L _T. The mean ovarian fecundity was estimated at 16 offspring per female but was unrelated to female L _T. Male gonado-somatic ( I _G) and hepato-somatic ( I _H) indices and female I _G declined from July to November as did maximum ovarian follicle diameter and the diameter of the three largest follicles. Adults were absent from inshore reefs between December and July. Hence, H. portusjacksoni has a synchronous annual breeding season in NSW, which occurs between July and November (the austral winter to spring), with a peak in oviposition from August to October. Heterodontus portusjacksoni copulatory and ovipository behaviour are reported for the first time. Copulation was observed and involved oral grasping of the female's pectoral fin by a single male, which wrapped his body around hers to insert one clasper. Ovipositing females appeared to search crevices in the reef prior to delivering a single capsule, which was washed into the crevice by water movement, with the female departing very soon after oviposition. This study represents the first rigorously quantitative analysis of H. portusjacksoni reproductive biology and ecology in NSW waters.     

Reproduction in the dusky grouper from the southern Mediterranean

Demographic data and gonad histology confirmed that the dusky grouper Epinephelus marginatus is a protogynous hermaphrodite that follows a monandric pathway to sexual development. Females reached first sexual maturity at 36·7 cm L_s and estimated mean length at first maturity (L₅₀) was 43·8 cm L_s for females and 81·3 cm L_s for males. Adult sex ratios during the reproductive period were c. 3·5: 1 females to males. Females exhibited group-synchronous ovarian development and multiple ovulation occurred over the spawning period. Gonads were ripe from early May and spawning occurred from June until early September. The size of ripe testes (0·6% W )indicated strong oligospermy and suggested a mating system with no sperm competition. Sexual transition was protogynous involving regression of ovarian tissue and proliferation of testicular tissue in the gonads. Transitional individuals occurred from May through November and accounted for 9% of sampled adult population. Sex change occurred in fish 69–93 cm (L_s) long and the size distributions of males and females overlapped over 27% of the L_s range. Special zones were recognized as gathering areas for sexually mature dusky groupers during the reproductive period.     

Maturation, spawning and growth of rock soles off Kodiak Island in the Gulf of Alaska

Northern rock sole Lepidopsetta polyxystra females from the Kodiak Island area, Alaska, reached 50% maturity at 328 mm L _T and an average age of 7 years. In contrast, southern rock sole Lepidopsetta bilineata females reached 50% maturity at 347 mm L _T and an average age of 9 years. Spawning started in midwinter for northern rock sole and peaked during the spring, while spawning for southern rock sole occurred during the summer. The bottom depth for spawning northern rock sole ranged from 43 to 61 m and averaged 45 m; spawning depth for southern rock sole ranged from 35 to 120m and averaged 78m. Both species appeared to develop a single stock of oocytes and to ovulate them in a single spawning. Northern rock sole females grew faster overall ( K =0.24) than southern rock sole females ( K =0.12) but reached a smaller maximum length ( L _∞=430 mm) than southern rock sole ( L _∞=520mm). Males of both species grew more slowly than females after 5 years of age and reached a smaller maximum length.     

A hydro-mechanical approach to the scaling of swimming performance in the sand flathead Platycephalus bassensis Cuvier: effects of changes in morphological features based on fish size

The swimming performance of Platycephalus bassensis at steady speed was assessed with an emphasis on hydrodynamics. The minimum swimming speed to maintain hydrostatic equilibrium for P. bassensis of 0·271 m total length ( L _T) was calculated to be 1·06 L _T s⁻¹. At this speed, the required lift to support the mass of the fish was equivalent to 6·6% of the fish mass; 82·7% of which was created by the body as a hydrofoil, and the rest of which was created by the pelvic fins as hydrofoils. The minimum swimming speed decreased with the L _T of the fish and ranged from 1·15 L _T s⁻¹ for a fish of 0·209 m to 0·89 L _T s⁻¹ for a fish of 0·407 m. The forward movement per tail-beat cycle ( i.e. stride length) was described with an equation including quantities of morphological and hydro-mechanical relevance. This equation explained that stride length was increased by the effect of turbulence characterized by the Reynolds number and demonstrated the morphological and hydro-mechanical functional design of the fish for maximizing thrust and minimizing drag. The larger span of the caudal fin and caudal tail-beat amplitude was associated with larger stride length, whereas greater frictional drag was associated with smaller stride length.     

Constraints of body size and swimming velocity on the ability of juvenile rainbow trout to endure periods without food

The hypothesis that body size and swimming velocity affect proximate body composition, wet mass and size‐selective mortality of fasted fish was evaluated using small (107 mm mean total length, L _T) and medium (168 mm mean L _T) juvenile rainbow trout Oncorhynchus mykiss that were sedentary or swimming ( c . 1 or 2 body length s⁻¹) and fasted for 147 days. The initial amount of energy reserves in the bodies of fish varied with L _T. Initially having less lipid mass and relatively higher mass‐specific metabolic rates caused small rainbow trout that were sedentary to die of starvation sooner and more frequently than medium‐length fish that were sedentary. Swimming at 2 body length s⁻¹ slightly increased the rate of lipid catabolism relative to 1 body length s⁻¹, but did not increase the occurrence of mortality among medium fish. Death from starvation occurred when fish had <3·2% lipid remaining in their bodies. Juvenile rainbow trout endured long periods without food, but their ability to resist death from starvation was limited by their length and initial lipid reserves.     

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.