Reproductive biology of the brown smoothhound shark Mustelus henlei,in the northern Gulf of California,México

Female brown smoothhound sharks Mustelus henlei were found to reproduce annually. A mature female carried both developing oocytes in the ovary and developing embryos in the uteri concurrently for c. 1 year. A great variability in the size of embryos was recorded each month, and the maximum embryo sizes were found from late January to mid‐March. The largest oocytes in mature females were observed in mid‐March. Gestation lasted c. 10 months. A linear relationship between maternal total length (L_T) and the number of pups per litter (litter size one to 21) was estimated. Birth L_T was reached in c. 280 mm. Females and males matured at 570–660 and 550–560 mm L_T, respectively. Difference in the litter size among Californian coast (one to 10) and northern Gulf of California (one to 21) populations existed for this smoothhound shark.     

Life‐history traits of the long‐nosed skate Dipturus oxyrinchus

This work investigates life‐history traits of the long‐nosed skate Dipturus oxyrinchus, which is a common by‐catch in Sardinian waters. The reproductive variables were analysed from 979 specimens sampled during scientific and commercial hauls. Females (10·4–117·5 cm total length, L_T) attained larger sizes than males (14·5–99·5 cm L_T). To evaluate age and growth, a sub‐sample of 130 individuals (76 females and 54 males) were used. The age was estimated by annuli counts of sectioned vertebral centra. Four models were used for the length‐at‐age data: the von Bertalanffy, the exponential, the Gompertz and the logistic functions. According to the Akaike's information criterion, the Gompertz model seemed to provide the best fitting curve (L_∞ mean ± s.e. : 127·55 ± 4·90 cm, k: 0·14 ± 0·09, IP: 3·97 ± 0·90 years). The oldest female and male were aged 17 (115·5 cm L_T) and 15 years (96·0 cm L_T), respectively. Lengths at maturity were 103·5 cm for females and 91·0 cm for males, corresponding to 90% of the maximum observed length in both sexes. The monthly distribution of maturity stages highlighted an extended reproductive cycle, with spawning females and active males being present almost throughout the year, as confirmed by the gonado‐somatic index. Ovarian fecundity reached a maximum of 26 yolked follicles with a mean ± s.e. size of 19·7 ± 6·5 mm.     

Living on the edge: latitudinal variations in the reproductive biology of two coastal species of sharks

Differences in the reproductive biology of both the Australian weasel shark Hemigaleus australiensis and the Australian sharpnose shark Rhizoprionodon taylori were apparent between individuals from the southern‐most extent of their range in eastern Australia (Moreton Bay) and those from northern Australia. For H. australiensis from Moreton Bay the total length (L_T) at which 50% of individuals were mature (L_T50) was 759 mm for females and 756 mm for males, values that were respectively 17–26% larger than reported for the species in northern Australia. The relatively low percentage (63%) of pregnant mature females and presence of small, similar‐sized, embryos in utero in both May and November suggested a semi‐synchronous, annual reproductive cycle in Moreton Bay, whereas a synchronous, biannual reproductive cycle occurred in northern Australia. It is likely that H. australiensis has a resting phase between gestation cycles at the southern‐most extent of its range. For R. taylori from Moreton Bay the L_T50s were 588 and 579 mm for females and males, respectively, values 2–3% larger than for individuals from the mid‐Queensland coast and 31–35% larger than for individuals from northern Australia. The length at which 50% of the females were maternal (611 mm L_T) in Moreton Bay was greater than the L_T50, indicating that not all sharks mate immediately after maturing. Rhizoprionodon taylori in the south had an annual reproductive cycle incorporating a 7–8 month embryonic diapause, with pups probably born in February. A mean fecundity of 7·5 was almost double that reported from northern Australia. Regional variations in the reproductive characteristics of H. australiensis and R. taylori may influence their resilience to fishing and other anthropogenic pressures. The substantial differences reported here highlight the importance of region‐specific life‐history parameters to successful management and conservation.     

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.     

Life histories of two deep‐water Australian endemic elasmobranchs: Argus skate Dipturus polyommata and eastern spotted gummy shark Mustelus walkeri

Two Australian endemic elasmobranchs, the Argus skate Dipturus polyommata and the eastern spotted gummy shark Mustelus walkeri, were collected from the by‐catch of a prawn Melicertus plebejus trawl fishery off Queensland. Age and growth parameters were estimated from growth band counts in vertebral sections of 220 D. polyommata and 44 M. walkeri. Dipturus polyommata males and females had an observed maximum age of 10 years and reached maximum sizes of 369 and 371 mm total length (L_T), respectively. Mustelus walkeri lived longer, with the oldest female aged 16 years and measuring 1050 mm stretched total length (L_ST), and oldest male aged 9 years and 805 mm L_ST. Dipturus polyommata grew relatively fast with a von Bertalanffy growth completion parameter of k = 0·208 year^?1 with males reaching maturity at 4·0 years (c. 278 mm L_T) and females at 5·1 years (c. 305 mm L_T). Mustelus walkeri grew more slowly with k = 0·033 year^?1 with males estimated to mature at 7–9 years (670–805 mm L_ST) and females at 10–14 years (833–1012 mm L_ST). Length at birth inferred from neonate D. polyommata was 89–111 mm L_T while for M. walkeri it was estimated to be 273 L_ST based on the value of L₀ from the von Bertalanffy growth model. Both species appeared to have continuous reproductive cycles and low fecundity with an average ovarian fecundity of eight follicles for D. polyommata and a litter size of five to seven pups for M. walkeri. Based on these life‐history traits, D. polyommata is more resilient to fishing pressure than M. walkeri.     

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 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.     

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.     

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.     

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.     

Proximate causes of sexual size dimorphism in Colorado pikeminnow, a long‐lived cyprinid

Evidence for sexual size dimorphism (SSD) and its possible causes were examined in the endangered Colorado pikeminnow Ptychocheilus lucius, a large, piscivorous, cyprinid endemic to the Colorado River system of North America. Individuals representing 18–24% of the upper Colorado River population were captured, measured, sexed and released in 1999 and 2000. Differing male and female total length‐(L_T) frequency distributions revealed SSD with females having greater mean and maximum sizes than males. Although both sexes exhibit indeterminate post‐maturity growth, growth trajectories differed. The point of trajectory divergence was not established, but slowed male growth might coincide with the onset of maturation. Differing growth rate was the dominant proximate cause of SSD, accounting for an estimated 61% of the observed difference in mean adult L_T. The degree of SSD in adults, however, was also related to two other factors. Evidence suggests males become sexually active at a smaller size and earlier age than females; a 2 year difference, suggested here, accounted for an estimated 12% of the between‐sex difference in mean adult L_T. Temporal shifts in gender‐specific survival accounted for an additional 27% of the observed between‐sex difference in mean adult L_T. Estimated age distributions indicated a higher number of older females than older males and more younger males than younger females in the population during the period of sampling. Dissimilarity of age distributions was an unexpected result because the male : female population sex ratio was 1 : 1 and estimates of long‐term annual survival for adult males and females were equal (88%). Future assessments of SSD in this population are apt to vary depending on the prior history of short‐term gender‐specific survival. Without recognizing SSD, non‐gender‐specific growth curves overestimate mean age of adult females and underestimate mean age of adult males of given L_T. Assuming age 8 years for first reproduction in males and age 10 years for females, the adult male : female ratio was estimated as 1·1 : 1 and mean adult age, or generation time, was estimated as 16·4 years for males and 18·4 years for females.     

Reproductive life history of the introduced peacock grouper Cephalopholis argus in Hawaii

This research investigated the reproductive biology (sex ratio, hermaphroditic pattern, size and age at maturity) of Cephalopholis argus, known locally in Hawaii by its Tahitian name roi. The results suggest that C. argus exhibits monandric protogyny (female gonad differentiation with female to male sex change) with females reaching sexual maturity at 1.2 years (95% c.i .: 0.6, 1.6) and 20.0 cm total length (L_T; 95% c.i .: 19.6, 21.2). The female to male sex ratio was 3.9:1. The average age and L_T at sex change was 11.5 years (95% c.i .: 11.1, 12.9) and 39.9 cm (95% c.i .: 39.5, 41.2), respectively. Current information on spawning seasonality of this species is incomplete, but based on the occurrence of spawning capable and actively spawning females, spawning probably takes place from May to October. Evidence of lunar spawning periodicity was found, with an increased proportion of spawning capable and actively spawning females, and an increased female gonado‐somatic index during first quarter and full‐moon phases. This information fills a valuable information gap in Hawaii and across the species' native range.     

Reproductive parameters of rhinobatid and urolophid batoids taken as by‐catch in the Queensland (Australia) east coast otter‐trawl fishery

Reproductive variables are provided for batoids regularly taken as by‐catch in the east coast otter‐trawl fishery on the inner‐mid continental shelf off the south‐east and central coasts of Queensland, Australia. Total length at maturity (L_T50 and 95% c.i .) for the eastern shovelnose ray Aptychotrema rostrata was 639·5 mm (617·6–663·4 mm) for females and 597·3 mm (551·4–648·6 mm) for males. Litter size (n = 9) ranged from nine to 20 (mean ± s.e. = 15·1 ± 1·2). This species exhibited a positive litter size–maternal size relationship. Disc width at maturity (W_D50 and 95% c.i .) for the common stingaree Trygonoptera testacea was 162·7 mm (155·8–168·5 mm) for females and 145·9 mm (140·2–150·2 mm) for males. Gravid T. testacea (n = 6) each carried a single egg in the one functional (left) uterus. Disc width at maturity (W_D50 and 95% c.i .) for the Kapala stingaree Urolophus kapalensis was 153·7 mm (145·1–160·4 mm) for females and 155·2 mm (149·1–159·1 mm) for males. Gravid U. kapalensis (n = 16) each carried a single egg or embryo in the one functional (left) uterus. A single female yellowback stingaree Urolophus sufflavus carried an embryo in each uterus. A global review of the litter sizes of shovelnose rays (Rhinobatidae) and stingarees (Urolophidae) is provided.     

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.     

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.     

New records of angular rough sharks Oxynotus centrina in the coastal waters of Malta,with observations on post‐capture resilience and release behaviour

One male and one female angular rough sharks Oxynotus centrina were caught south‐east of Malta between May and June at a depth of 60–100 m < 5 km from shore. The immature female (total length, L_T 565 mm) was landed dead but the male (535 mm L_T) was found alive. This communication presents important biological observations on post‐capture recovery and release behaviour of this species.     

Size and age estimates at sexual maturity for the little skate Leucoraja erinacea from the western Gulf of Maine,U.S.A.

Size and age estimates at sexual maturity were determined for 162 male and 273 female little skates Leucoraja erinacea collected from the western Gulf of Maine. Maturity ogives suggest that 50% maturity in females occurs at age 9·5 years and 480 mm total length (L_T), whereas 50% maturity in males occurs at a slightly younger age of 7·7 years and smaller size of 460 mm L_T. Age estimates were made from 389 L. erinacea ranging in size from 93 to 570 mm L_T. The index of average per cent error and age‐bias plots indicated that the ageing methods were precise and non‐biased. Additionally, annual periodicity of band formation was validated with oxytetracycline in eight individuals (three males and five females) ranging in age from 3 to 12 years. In conclusion, results from this study indicate that L. erinacea exhibits characteristics that make other elasmobranch populations highly susceptible to overexploitation.     

Age, growth, and sexual maturity of the yellownose skate Dipturus chilensis in the south‐eastern Pacific

The age and growth parameters of Dipturus chilensis were estimated by counting growth rings from thin sections of vertebral centra from 400 fish (246 females and 154 males), ranging from 23 to 124 cm total length (L_T), and backcalculating fish lengths at previous ages. Marginal increment analysis lent support to the hypothesis of annual deposition of band‐pairs, which formed during the winter months. The oldest female D. chilensis aged in this study was 21 years and 117 cm L_T, whereas the oldest male was 18 years and 93 cm L_T. A 4·7% index of average per cent error (I_APE) suggested that this is a precise method for calculating the age of D. chilensis. Observed L_T were lower than backcalculated L_T, which implies the influence of Lee's phenomenon. The von Bertalanffy growth equations, based on mean length‐at‐age data, were estimated as L_t = 128·3 (1 ? e^{?0·112 (t + 0·514)}) for females and L_t = 107·8 (1 ? e^{?0·134 (t + 0·862)}) for males where t is age (years). Growth was significantly different between sexes: females reached a larger adult size. Ages and lengths at 50% maturity were estimated at 14 years of age and 106 cm L_T for females and 11 years of age and 86 cm L_T for males. At c. 14 years, there was a decline in growth rates in females. The factor most likely responsible for this was sexual maturity, which caused a discontinuity in growth of female fish. These results show that this species is slow‐growing, long‐lived, relatively large and of delayed maturity, characteristics that make it vulnerable to exploitation.     

Biological observations on the bristly catshark Bythaelurus hispidus from deep waters off the south‐west coast of India

Biological data are presented for the poorly known bristly catshark Bythaelurus hispidus based on specimens collected from the by‐catch of the commercial deep‐sea shrimp trawl fishery operating in the Arabian Sea at depths of 200–500 m off the south‐west coast of India. One hundred and sixty‐two individuals, which ranged from 120 to 366 mm total length (L_T), were collected for this study. Size‐at‐maturity (L₅₀) for females and males was estimated at 252 and 235 mm L_T, respectively. The reproductive mode of B. hispidus was aplacental viviparity, which is the rarest reproductive mode within the Scyliorhinidae and is considered to be the most advanced of the three reproductive modes occurring within this family. Dietary analysis of stomach contents revealed B. hispidus feeds on a variety of prey, primarily fishes.