Reproductive biology of the guitarfish Rhinobatos percellens (Chondrichthyes,Rhinobatidae) from the São Paulo Coast,Brazil, western South Atlantic Ocean

The reproductive biology of the guitarfish Rhinobatos percellens was studied from 751 specimens caught by bottom pair trawlers off the coast of São Paulo, Brazil, between c. 24° 00′ S; 45° 15′ W and c. 25° 10′ S; 47° 52′ W, from September 2007 to August 2009. The total length (L_T) and total mass (M_T) relationship for males and females combined was M_T = 1·29E‐06 L_T^3·15 (r = 0·99, n = 751). The mean L_T of sexually mature specimens was 548 mm for males and 583 mm for females. Clasper growth was allometric and showed three distinct phases. Most claspers were calcified in specimens of c. 550 mm L_T. The mean diameter of the largest oocyte was 29·8 mm, the mean ovarian fecundity was seven oocytes and ovulation occurred between August and November. Uterine fecundity ranged from two to 13 embryos (mean of five embryos). Larger females had higher litter sizes and larger embryos; the size‐at‐birth was c. 200 mm L_T. The hepato‐somatic index oscillated seasonally for males and females; the gonado‐somatic index had little variation in males, but varied seasonally in females. The presence of many non‐pregnant adult females and of encapsulated eggs during two consecutive seasons suggests a resting period between gestations and the possibility of diapause.     

Reproductive biology of the greeneye spurdog Squalus chloroculus (Squaliformes,Squalidae)

The reproduction of the greeneye spurdog Squalus chloroculus was studied based on animals caught in the multispecies and multi‐gear southern and eastern scalefish and shark fishery on the upper continental slope off southern Australia. One hundred and ninety‐nine females (502–990 mm, total length, L_T) and 189 males (515–810 mm L_T) were examined. The female reproductive cycle, based on 41 breeding animals, is continuous and triennial, with the pregnancy period estimated to be 31–34 months, seasonal and synchronous with the ovarian cycle; a third of the breeding female population is estimated to give birth between September and December each year. The estimated L_T at which 50% of females are mature is 799 mm (95% c.i. : 794, 804), whereas the L_T at which 50% are maternal is 825 mm (95% c.i. : 817–833), but these estimates are probably biased by the phenomenon of apparent change of L_T at maternity and L_T at maturity following severe length‐selective fishing mortality. Litters ranged from four to 15 embryos with a 1:1 sex ratio, and litter size increased with maternal length. The breeding cycle of males is neither seasonal nor synchronous with the female cycle. The estimated L_T of males where 50% are mature was 629 mm (95% c.i. : 603, 645).     

Geographical variability in life‐history traits of a midslope dogfish: the brier shark Deania calcea

Deania calcea (n = 420) were collected from the catch of deep‐water trawlers in the southern and eastern scalefish and shark fishery in southern Australia during the years 2008–2011. The total length (L_T) range varied between sexes, females being larger (n = 264; 280–1530 mm) than males (n = 156; 310–921 mm). The reproductive cycle in this population is non‐continuous and asynchronous. The estimated L_T at which 50% of males are mature is 807 mm and is 914 mm for females. Populations of D. calcea in higher latitudes appear to mature at a larger size than conspecifics in lower latitudes, in both the northern and southern hemispheres. Litters ranged from three to 10 embryos with a 1:1 sex ratio, but litter size does not increase with maternal L_T. Deania calcea shows geographical variability in its biological parameters and gathering information on life‐history traits of populations is vital to understand the trade‐offs made by this species in response to environmental conditions and to predict intraspecific spatial differences. Such information is a basis for specific spatial management to protect populations from excessive fishing.     

Biological observations on the broadfin shark Lamiopsis temminckii (Carcharhiniformes: Carcharhinidae)

Biological information was collected from 214 individuals of the broadfin shark Lamiopsis temminckii measuring 418 to 1782 mm total length, L_T. Size at maturity (L₅₀) for females and males was estimated at 1430 and 1368 mm L_T, respectively, while mature and gravid females were observed from 1350 mm L_T with litter sizes 2–8 and size at birth 418–650 mm L_T. Analysis of stomach contents revealed a variety of prey, primarily crustaceans (54·0%), teleosts (42·7%) and cephalopods.     

Litter Overlap in Gambusia hubbsi: Superfetation Revisited

The frequency of litter overlap, the simultaneous presence of two litters at different stages of development within a single female, varies among populations of Gambusia hubbsi in different habitats on Andros, Bahamas. In freshwater bluehole and shallow water populations, less than 2% of the females carried two litters, independent of the difference in developmental stage between the litters. In well field populations, 2% of the females carried two litters composed of immature and mature oocytes, whereas 10% carried developing oocytes and late stage embryos, and 16% carried mature oocytes (or older) and late stage embryos. Among females from populations that we introduced into unoccupied well fields 2% carried two litters composed of immature and mature oocytes, about 14% carried developing oocytes and late stage embryos, and less than 6% carried mature oocytes (or older) and late stage embryos. When two litters composed developing oocytes and late stage embryos were present, the two litters were of equal size, and the total number of offspring was 1.5 times larger than expected on the basis of female body length. When two litters composed mature oocytes (or older stages) and late stage embryos were present, the two litters were also of equal size, and the total number of offspring was as expected on the basis of female body length. Litter overlap was independent of female size. A comparison of the consequences of litter overlap for strictly lecithotrophic and strictly matrotrophic modes of reproduction with the patterns of litter overlap observed in G. hubbsi, suggests that litter overlap in G. hubbsi reduces the cost of reproduction, but does not increase the rate of offspring production.     

Temporal and latitudinal comparisons of reproductive parameters in a heavily exploited shark,the bonnethead,Sphyrna tiburo (L. 1758), in the southern Gulf of Mexico

In the southern Gulf of Mexico, the bonnethead shark, Sphyrna tiburo, is one of the most frequently captured species in landings of small-scale fisheries. Based on the analysis of two fishery-dependent sampling periods (1993–1994 and 2007–2014), this study aimed to determine reproductive parameters and identify temporal differences between the two time periods. In the first sampling period, 776 males and 352 females with a size range of 28.0–120.0 cm total stretched length (L_T) were analysed, and in the second sampling period, 387 males and 432 females with a size range of 28.0–122.0 cm L_T were analysed. The size at 50% maturity in the second sampling period was significantly different between sexes, 82.6 cm L_T for females and 73.8 cm L_T for males (no estimation was possible for the first sampling period). The size at 50% maternity was not different between sampling periods, 97.3 cm L_T for the first sampling period and 99.0 cm L_T for the second sampling period. Litter size varied from 3 to 19 embryos and the average was not statistically different in both periods, 10.1 (S.D. = 3.8) for the first sampling period and 11.3 (S.D. = 3.5) for the second sampling period. The female reproductive cycle is asynchronous, and it seems to be annual, with a gestation period of 5–6 months, and a consecutive ovarian cycle and gestation period. Temporal (between sampling periods) and latitudinal (southern Gulf versus northern regions) variations occur in the synchronicity of the reproductive cycle, temporal variation in the relationship between maternal length and litter size, and latitudinal variation in average size of mature sharks.     

Timecourse of oocyte development in saithe Pollachius virens

Wild caught North Sea saithe Pollachius virens were monitored for growth, sex steroid profiles and oocyte development pre‐spawning and measured for egg size and group fecundity during the spawning season in the laboratory. Vitellogenesis commenced in late October–early November, at a leading cohort size (C_L) of c. 250 µm, after which oocytes grew rapidly in size until spawning started in February. Notably, a distinct cortical alveoli stage was virtually absent with yolk granules observed in developing oocytes at the very beginning of vitellogenesis. Little atresia was observed pre‐spawning, but atretic re‐absorption of remnant oocytes containing yolk granules was found in all females immediately post‐spawning. As expected, concentrations of sex steroids, oestradiol‐17β (females), testosterone (both sexes) and 11‐ketotestosterone (both sexes), increased pre‐spawning before dropping post‐spawning. The present experiment provides the first validation of sex steroid levels in P. virens. Post‐ovulatory follicles were visible in histological sections from female gonads 9–11 months post‐spawning, but then disappeared. Spawning commenced around a C_L of c. 750 µm (700–800 µm). Hydrated oocytes (eggs) measured between 1·04 and 1·31 mm (mean = 1·18 mm) with decreasing sizes towards the end of spawning. The average estimated realized fecundity was c. 0·84 million eggs (median female total length, L_T = 60 cm). Spawning lasted from 13 February to 29 March.     

Reproductive biology of Haemulon plumierii in the south‐western Atlantic Ocean's most extensive reefs: implications for fisheries management

The reproductive biology of the white grunt Haemulon plumierii was studied from 360 individuals obtained from artisanal fisheries landings in the Abrolhos Bank, Brazil, between August 2010 and March 2012. The overall sex‐ratio did not differ significantly from 1:1, although males predominated in larger size classes. β‐Binomial modelling of historical sex‐ratio data indicated that the catch rate of females has increased in recent years. Females reached maturity at a smaller total length (L_T; 214 mm) than males (235 mm L_T) and the L_T at which 50% of all individuals are mature (L₅₀) was 220 mm, corresponding to 41·5% of the maximum recorded L_T. Variation in the gonado‐somatic index and in the relative frequency of reproductive stages indicates that reproduction occurs year round, with increased activity during the austral spring and summer. Fecundity was not size dependent. The reproductive parameters provided here can support management measures focussed on seasonal closures during spawning peaks (September to November and February to March) and minimum sizes (>L₅₀) for the capture of this important artisanal fisheries resource in Abrolhos, the region with the largest and most biodiverse coralline reefs in the South Atlantic Ocean.     

Life history of the river goby Glossogobius callidus (Teleostei: Gobiidae)

The river goby Glossogobius callidus is native to freshwater and estuarine habitats in South Africa. Individuals [21.1–144.4 mm total length (L_T)] were sampled from impoundments in the Sundays River Valley, Eastern Cape, from February 2014 to March 2015. The largest female was 137.2 mm L_T, and the largest male was 144.4 mm L_T. Length-at-50% maturity was 75.2 ± 2.1 mm L_T for males and 76.2 ± 2.0 mm L_T for females. Absolute fecundity was 1028.2 ± 131.7 oocytes per fish, and relative fecundity was 50.1 ± 18.1 oocytes per gram. The spawning season extended from October to December. Fish were aged using sectioned sagittal otoliths. The growth zone periodicity was validated using edge analysis. Longevity was more than 7 years for females and more than 6 years for males. Length-at-age was similar for the two sexes and was best described using the von Bertalanffy growth model as L_t = 74.7(1 − e^{–1.0(t + 0.1)}) mm L_T for the entire population. Using the population age structure, the mortality rate was estimated at 1.3 per year.     

Age and size at sexual maturity of the smooth skate Malacoraja senta from the western Gulf of Maine

Age and size at sexual maturity was determined for 185 male and 96 female smooth skates Malacoraja senta (ranging in size from 370 to 680 mm total length L_T), collected from the western Gulf of Maine. Maturity ogives for males, based on clasper length, testis mass and the proportion of mature spermatocysts in the testes, suggest that 50% maturity occurs between 9 and 10 years and 560 mm L_T. Maturity ogives for females, based on ovary mass, shell‐gland mass and maximum follicle size, suggest that 50% maturity occurs at age 9 years and 540 mm L_T.     

Reproductive biology of skates, Bathyraja(Ishiyama), along the eastern Bering Sea continental slope

Eight Bathyraja skate species, Bathyraja aleutica, Bathyraja interrupta, Bathyraja lindbergi, Bathyraja maculata, Bathyraja minispinosa, Bathyraja parmifera, Bathyraja taranetzi and Bathyraja trachura, collected along the eastern Bering Sea continental slope were examined for reproductive features. The sex ratio for six of eight Bathyraja skate species studied was c. 1 : 1 with males dominating for B. interrupta and B. parmifera. Five of eight species examined, B. aleutica, B. lindbergi, B. parmifera, B. taranetzi and B. trachura approximately equalled or exceeded the maximum reported total length, L_T. Maximum L_T for three species (B. interrupta, B. minispinosa and B. parmifera) revealed that the largest individuals were males and that in three other species (B. lindbergi, B. maculata and B. trachura) both females and males grew to approximately the same size. Both females and males matured at approximately the same L_T for seven of eight species. Only female B. aleutica matured at a much larger size than males. All species matured within a narrow size range following a protracted juvenile stage and once mature these skates appeared to grow very little. First maturity for all species occurred at ≥80% of their maximum L_T. The number of oocytes did not significantly differ between the left and right ovary for any species. The total number of mature oocytes increased with L_T for six of eight species examined. The number of oocytes for one species, B. aleutica, increased in number until 145 cm L_T, then declined in those individuals greater in length. The egg cases of all species are described, of which four (B. lindbergi, B. parmifera, B. taranetzi and B. trachura) have never been previously reported. Evidence is presented that suggest skates may be limited to as little as one and as many as 13 spawning seasons, depending on the species, and that senescence may occur in older individuals.     

Reproductive and population parameters of spiny dogfish Squalus acanthias in the south‐western Atlantic Ocean

The objective of this study was to estimate reproductive and population parameters of the spiny dogfish Squalus acanthias for the south‐western Atlantic Ocean. In total, 2714 specimens (1616 males and 1098 females) were collected from surveys carried out using research vessels. Males ranged from 225 to 861 mm total length (L_T) and females from 235 to 925 mm L_T. The size at maturity of females (651 mm) was significantly greater than that of males (565 mm). The maximum proportion of mature individuals (P_max) of the gestation ogive was <1, which indicates that a proportion of mature females was not in gestation. This inactivity may be explained by the occurrence of resting periods between cycles or by the asynchrony of the reproductive cycle. The estimated P_max for the maternity ogive suggested that about one third of mature females were in the maternity stage (i.e. with embryos >156 mm). The temporal and spatial co‐occurrence of non‐gravid adult females at different stages of ovarian development, as well as gravid females at all embryonic development stages would indicate that the female reproductive cycle in the south‐western Atlantic Ocean is asynchronous. The results indicate that S. acanthias is susceptible to fishing pressure on account of its length at maturity, extended reproductive cycles and low fecundity.     

Reproductive biology of the southern thorny skate Amblyraja doellojuradoi (Chondrichthyes,Rajidae)

The total lengths (L_T) of 193 males (209–556 mm) and 130 females (275–515 mm) of Amblyraja doellojuradoi, a commercial by‐catch species on the Argentinean continental shelf, which are increasingly retained, were analysed. No sexual dimorphism was observed in the L_T at which 50% of individuals were sexually mature; males matured at 448 mm and females at 411 mm, c. 80 and 82% of maximum L_T. The hepato‐somatic index was similar among sexes, but significantly different between maturity stages, being lower in mature than immature specimens. Males had no seasonal difference in the hepato‐somatic index and females had the lowest index in autumn. The gonado‐somatic index was lower in males than in females and significantly higher in mature than immature specimens of both sexes. Males had the highest index in autumn and females had no seasonal difference. Collectively, these results would indicate that A. doellojuradoi breeds in autumn.     

Catch composition and reproductive biology of Sphyrna lewini (Griffith & Smith) (Carcharhiniformes, Sphyrnidae) in Indonesian waters

Data on the length compositions and reproductive biology have been collected for Sphyrna lewini in Indonesian waters, in which this species makes an important contribution to the biomass of its artisanal and small‐scale fisheries. These data were obtained by recording relevant body measurements and counts for this species while visiting Indonesian fish landing sites. The fish, which had been caught by gillnetting and longlining, covered a wide length range and included substantial numbers of immature, maturing and mature individuals. The vast majority of S. lewini <1100 mm total length (L_T) had been caught by gillnetting, whereas most of those above this length had been obtained through longlining. The number of embryos in pregnant females, which ranged from 14 to 41, with a mean of 25, was positively correlated with the L_T of those females. Birth occurred at c. 400 mm L_T and predominantly in October and November. The number of females was similar to that of males among smaller fish, but far greater than that of males among larger fish. This presumably reflects a faster growth and greater longevity of females, selectivity of longlining for females and a tendency for males to move outside the area fished. For males, the relationships between clasper length and the extent of its calcification, and also with sexual maturation and L_T have been determined. Females attained maturity (L_T50) at a far larger size (2285 mm) than males (1756 mm). The L_T of virtually all females and all males taken by gillnetting were less than their respective L_T50 at maturity, and c. 67 and 51% of the longline catches of females and males, respectively, were likewise immature. This feature, together with the substantial catches of S. lewini and the life cycle traits of elasmobranchs, suggest that this species is likely to be prone to overfishing in Indonesian waters. Furthermore, the removal of large numbers of this apex predator will presumably be affecting the trophic structure in the waters in which it is fished.     

Migration and reproductive biology of Mugil liza (Teleostei: Mugilidae) in south Brazil

The mullet Mugil liza occurs along the Atlantic coast of South America from Venezuela to Argentina, but 95% of the commercial catch is collected from south Brazil between São Paulo and Argentina. Mugil liza is a single spawner with oocyte development occurring synchronously in two groups. Spawning happens in marine areas and occurs after migration. The reproductive migration occurs from Argentina (38° S) to the southern Brazilian states (24–26° S) from April to July, with peak spawning in June between northern Santa Catarina and Paraná. The presence of hyaline oocytes was associated with high salinity and sea surface temperatures of 19–21° C, and followed the seasonal northward displacement of these oceanographic conditions. The average size at first maturity (L_m) for both sexes was 408·3 mm total length, L_T. Males (L_m = 400·1) matured earlier than females (L_m = 421·9 mm). Fecundity ranged from 818 992 to 2 869 767 oocytes (mean = 1 624 551) in fish that were between 426 and 660 mm L_T.     

Primary male development of two sequentially hermaphroditic groupers,Epinephelus akaara and Epinephelus awoara (Perciformes: Epinephelidae)

Gonad ontogeny of the Hong Kong grouper Epinephelus akaara (a bi‐directional sex changer) and the yellow grouper Epinephelus awoara (a protogynous hermaphrodite) was examined for the first time from post‐larval phase until first sexual maturation, by histology. Approximately 20 specimens of each species were collected randomly every 2–7 weeks from rearing tanks with natural sea water and temperature between June 2013 and June 2014. The paired gonadal primordia (GP) were observed at 6 weeks after hatching (wah) for both species; however, gonia were first observed in GP at 16 wah for E. akaara and at 8 wah for E. awoara. The timings for the appearance of primary‐growth stage oocytes (O1) and the completion of ovarian lumen (OL) varied; both at 27 wah for E. akaara, and at 18 and 23 wah for E. awoara respectively. A bisexual‐phase gonad with an OL, O1 and scattered spermatogenic cysts (SC) was observed at 27–29 wah for both E. akaara and E. awoara. Sexual differentiation was subsequently observed from the bisexual‐phase gonad at 34 wah for E. akaara, and 41 wah for E. awoara, with the appearance of cortical‐alveolus stage oocytes (O2) for developing female and the proliferation of SC for developing primary male (i.e. from juvenile directly). Ovaries of mature females contained the vitellogenic stage oocytes (O3) and scattered SC; testes of mature primary males had sperm in sperm sinuses within the gonadal wall and remained O1. Minimum age of first sexual maturation for both female and primary male of E. akaara was at 41 wah; minimum total length (L_T) of female (143 mm) was larger than that of primary male (137 mm L_T). Minimum age and size of first sexual maturation for female of E. awoara (47 wah and 149 mm L_T, respectively) were larger than those of E. akaara. Developing primary males of E. awoara were found at 41–58 wah, however, mature males were not observed, indicating inconsistency in first sexual maturation for E. awoara. This study provided strong evidences of primary male pathway in E. akaara and E. awoara; the latter is confirmed to be diandric.     

Maturation and sexual ontogeny in the spangled emperor Lethrinus nebulosus

The reproductive development and sexual ontogeny of spangled emperor Lethrinus nebulosus populations in the Ningaloo Marine Park (NMP) were investigated to obtain an improved understanding of its evolved reproductive strategy and data for fisheries management. Evidence derived from (1) analyses of histological data and sampled sex ratios with size and age, (2) the identification of residual previtellogenic oocytes in immature and mature testes sampled during the spawning season and (3) observed changes in testis internal structure with increasing fish size and age, demonstrated a non‐functional protogynous hermaphroditic strategy (or functional gonochorism). All the smallest and youngest fish sampled were female until they either changed sex to male at a mean 277·5 mm total length (L_T) and 2·3 years old or remained female and matured at a larger mean L_T (392·1 mm) and older age (3·5 years). Gonad masses were similar for males and females over the size range sampled and throughout long reproductive lives (up to a maximum estimated age of c. 31 years), which was another correlate of functional gonochorism. That the mean L_T at sex change and female maturity were below the current minimum legal size (MLS) limit (410 mm) demonstrated that the current MLS limit is effective for preventing recreational fishers in the NMP retaining at least half of the juvenile males and females in their landed catches.     

Reproductive biology of two co‐occurring mugilids,Liza argentea and Myxus elongatus,in south‐eastern Australia

The reproductive biology of Liza argentea and Myxus elongatus occurring in two estuaries (Lake Macquarie and St Georges Basin) was found to differ. Gonado‐somatic index values and macroscopic staging of gonads identified the peak spawning period of L. argentea occurred between March and November in Lake Macquarie and January and April in St Georges Basin. In contrast, peak spawning of M. elongatus was concentrated between January and March in both estuaries. Spawning of L. argentea probably occurred in the lower reaches of estuaries as well as in nearshore coastal waters, whereas evidence indicated M. elongatus spawned only in ocean waters. The mean fork length at maturity (L_F50) was greater for females than males in both species, and it also occurred at a larger mean L_F in M. elongatus (males = 230 mm and females = 255 mm) than L. argentea (males = 180 mm and females = 207 mm). Estimates of total potential fecundity were also greater for M. elongatus (425 484–1 157 029) compared to L. argentea (159 933–548 954). Both species had determinate fecundity and displayed a group synchronous pattern of oocyte development, with two distinct size classes of oocytes present in mature ovaries. Liza argentea probably release the larger class of oocytes in one spawning event, but this could not be established for M. elongatus.     

Reproductive biology of the crocodile shark Pseudocarcharias kamoharai

From February 2005 to September 2007, a total of 490 crocodile sharks Pseudocarcharias kamoharai, caught as by‐catch in the swordfish and tuna longline fishery that operates in the tropical western Atlantic Ocean, was studied in regard to their reproductive biology. Maximum observed total lengths (L_T) were 1220 and 1090 mm for females and males respectively, with a high proportion of the catch being composed of mature specimens. Sexual maturity was attained at 760–810 mm L_T for males (L_T50 = 800 mm) and 870–980 mm L_T for females (L_T50 = 916 mm). The size at birth was estimated at 415 mm L_T. Temporal variation in gonad morphology and mass suggests that in this region P. kamoharai, an aplacental viviparous species with oophagy, does not show a well‐defined reproductive seasonality, with mating and parturition occurring possibly over an extended period of the year. Mean ±s.d . fecundity was estimated to be 3·9 (± 0·6) pups per reproductive cycle.     

Goosefish Lophius americanus fecundity and spawning frequency,with implications for population reproductive potential

To improve knowledge of goosefish Lophius americanus' reproductive biology, females were collected during 2009–2012 from the Mid‐Atlantic Bight shelf region of the U.S. east coast. Batch fecundity increased with total length (L_T), from 229 100 to 2 243 300 mature oocytes per female (L_T range: 55·5–112 cm; n = 54). This estimate of fecundity at L_T is lower than one derived from a sample collected during 1982–1985. Examination of whole oocyte diameters in different months indicated that L. americanus is a serial spawner, releasing more than one egg veil per spawning season, as suspected or observed for other Lophius species. Seasonality of spawning was evident from whole oocytes and gonad histology, and from larval fish surveys spanning the U.S. north‐east shelf, and confirmed a protracted (c. 6 months) spawning period. Peak spawning activity progressed northward from spring to autumn. The population‐level implications of these results were explored by estimating population reproductive potential (P_RP), which considered the value of both current and future per capita reproduction using decade‐specific age structure and fecundity at length. P_RP is now more than 50% lower compared with the historical period (1982–1985), a result of the lower proportions of large females and reduced fecundity across all sizes. Mechanisms that could explain this loss of stock productivity are fishing‐induced size–age truncation or regime shifts in egg production caused by changes in energy density of common forage species.