Reproductive aspects of the Atlantic angel shark Squatina dumeril in the southern Caribbean Sea

The maturity and reproduction of the Atlantic angel shark Squatina dumeril were assessed using 77 females (29·2–110·4 cm total length; L_T) and 269 males (58·7–108·2 cm L_T) harvested by artisanal gillnetters off Venezuela. The biased sex ratio implied segregation or sex‐specific gear selectivity. Based on the development of the reproductive tract, 50% L_T at sexual maturity (L_T50, mean ± s.e .) for females and males were estimated at 86·14 ± 0·64 and 81·55 ± 0·12 cm, respectively. Uterine fecundity ranged between one and six and with a maximum embryo size of 25·7 cm L_T. Gravid females were observed from August to December, including those close to parturition and while the gestation period was not confirmed, the size of ovarian follicles among some specimens implied protraction. The low fecundity of the species supports close monitoring of catches.     

Sex identification and PIT‐tagging: tools and prospects for studying intersexual differences in freshwater fishes

This study evaluated a technique to allow the long‐term monitoring of individual fishes of known sex in the wild using sex confirmation in close proximity to the reproductive period combined with individual tagging. Hundreds of partially migratory roach Rutilus rutilus were tagged with passive integrated transponders (PIT) following sex determination in spring and various performance measures were compared with fish tagged outside the reproductive period in autumn. Short‐term survival was >95% for R. rutilus sexed and tagged under natural field conditions. Total length (L_T) did not affect the probability of survival within the size range tagged (119–280 mm), nor were there differences in timing of migration the following season between individuals sexed and tagged in spring and individuals tagged in autumn (i.e. outside the reproductive period). Also, a similar per cent of R. rutilus sexed and tagged in spring and tagged in autumn migrated the following season (34·5 and 34·7%). Moreover, long‐term recapture data revealed no significant differences in body condition between R. rutilus individuals sexed and tagged in spring, individuals tagged in autumn and unmanipulated individuals. The observed sex ratio of recaptured fish did not differ from the expected values of equal recapture rates between males and females. Hence, there is no observable evidence for an adverse effect of tagging close to the reproductive period and therefore this method is suitable for studying intersexual differences and other phenotypic traits temporarily expressed during reproduction at the individual level in fishes.     

Differences in fat accumulation between immature male and female Atlantic salmon Salmo salar after dietary administration of tetradecylthioacetic acid

This study provoked sex‐specific differences in fat metabolism in Atlantic salmon Salmo salar, by dietary administration of tetradecylthioacetic acid (TTA) during their first spring and winter in the sea. The effects of TTA were evaluated in June of the first spring and May of the second spring in the sea, by analysing white muscle‐fat content. Muscle fat in males and females differed significantly as a result of TTA in their diet and diet interacted with the sex of the fish. The fat content during the first spring after dietary TTA was lowered by a greater amount in females than in males, 3·1–4·3%, respectively (P < 0·05). In contrast, during the second spring, fat content was lowered by a greater amount in males than in females, 15·8–16·7%, respectively (P < 0·01). Condition factor followed a similar pattern to the muscle fat. The results indicate that the difference in male and female fat accumulation dynamics is related to sex‐specific reproduction biology of S. salar. In addition, the findings show that it is important to consider the sex of the fish and the season of the year when studying fat dynamics and reproductive biology of S. salar.     

Reproductive strategy and recruitment dynamics of amphidromous goby Sicyopterus japonicus as revealed by otolith microstructure

The daily ages of 312 of 879 newly recruited postlarvae of Sicyopterus japonicus, collected from the Shuang‐Chi Estuary in north‐eastern Taiwan during February 1996 to April 1997, were determined from daily growth increments in their otoliths. Pelagic larval duration, growth rate of the marine larval stage and hatching dates were estimated, and recruitment timing was linked to environmental factors. The mean ±s .d . total length (L_T) and daily ages of S. japonicus at recruitment to the estuary were estimated to be 33·95 ± 1·31 mm (range 30·7 to 38·1, n = 317) and 163·72 ± 12·79 days (range 130 to 198, n = 312), respectively. The recruitment of S. japonicus larvae is size dependent not age dependent because L_T of the larvae is independent of age at recruitment. Periodic analysis indicated that L_T and growth rate of the larvae were inversely correlated with the age at recruitment, which means that the fast‐growing individuals recruited earlier. The growth rate of S. japonicus in the marine larval stage was synchronous with marine productivity in this subtropical area, i.e. the spring cohort recruited in the autumn had a higher growth rate than the autumn cohort recruited the following spring. The main spawning season of S. japonicus as backcalculated from otolith daily increments was in autumn, a relatively low productivity period compared with spring. During this season, there were fewer competitors and predators than in the more productive spring. The recruitment of 95% of postlarva coincided with low salinity (14) and low water temperature (23° C) in the river mouth that provided a buffer area for the adaptation of the larvae for upstream migration. This unique reproduction strategy and prolonged larval duration facilitated the widespread distribution of the fish along the coasts of East Asia.     

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.     

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.     

Manipulation of the energetic state of Atlantic salmon Salmo salar juveniles and the effect on migration speed

Atlantic salmon Salmo salar smolts were produced with similar energetic states as wild S. salar and the effect of low energetic state on smolt migration was tested. The total energetic state of the fish (body lipids and proteins) in the spring was correlated with Fulton's condition factor (K). Fish at a low energetic state swam slower but migrated further than fish at a higher energetic state when tested in two experimental streams. During a period of starvation throughout the winter and spring, fish conserved their body‐lipid reserves at 1·5% by using more protein as an energy source and the metabolic shift occurred between 3·5 and 1·5% body lipids. An energetic state of approximately 3·5 kJ g^?1 (K ≈ 0·65) appeared to be the critical limit for survival.     

Time and size at seaward migration influence the sea survival of Salmo salar

Whether time of seaward migration of young Atlantic salmon Salmo salar influences their subsequent survival and growth was investigated in the River Imsa, south‐western Norway. Salmo salar were tagged when moving downstream through a trap near the outlet between 1976 and 2010 and recaptured on their adult return. Most descended as smolts in April and May, but some descended during the other months of the year. Annual variation in timing of the smolt migration was significantly correlated with variation in water temperature during spring. Mean total body length of the descending S. salar varied with month of seaward migration. The sea survival of S. salar emigrating from the River Imsa between January and May was 2·8 times higher than for those descending between June and December. The sea survival of the various cohorts decreased with increasing river temperature in April to May, prior to the smolt migration, and decreasing day number when the smolts moved to sea. The size of smolts descending the river between April and May did not affect the survival at sea as much as it affected the survival of migrants descending in any other month of the year. The majority of the downstream migrating S. salar were 2 years old, but proportionally, more 1 year olds moved downstream in the autumn than in the rest of the year. Mean duration between downstream migration of the young and the return migration of the grilse was shortest (12·7 months) for those descending in July and August and longest for those descending in October (21 months). Mean monthly specific growth rate was highest for those migrating downstream between May and July and lowest for those emigrating in September. Based on the present results, it was hypothesized that S. salar emigrating between April and August migrated directly out into the ocean, while those that emigrated between October and March stayed in the estuary until the subsequent spring.     

Long‐term effects of photoperiod,temperature and their interaction on growth,gill Na+, K+‐ATPase activity,seawater tolerance and plasma growth‐hormone levels in Atlantic salmon Salmo salar

This study was undertaken to examine the long‐term effects of photoperiod, temperature and their interaction on growth, gill Na⁺,K⁺‐ATPase (NKA) activity, seawater tolerance and plasma growth‐hormone levels in Atlantic salmon Salmo salar pre‐smolts and smolts. The fish (mean ± s.e . initial body mass = 15·9 ± 0·4 g) were reared on two photoperiods (continuous light, LL, and simulated natural photoperiod, LDN, 60° 25′ N) and two temperatures (8·3 and 12·7° C) from June to May of the following year. Mean body mass was affected by photoperiod, temperature and their interactions. Both temperature groups on LL developed peak levels in gill NKA activity from October to November, 4–5 months prior to the natural season for the parr–smolt transformation. Fish at 12° C showed peak levels in NKA activity 4–6 weeks before the fish at 8° C. Fish in all four experimental groups showed maximum NKA activity within a similar size range (113–162 g). The present findings further indicate that smoltification in S. salar is to some extent driven by size, and that S. salar will develop smolt characteristics, e.g. a marked increase in NKA activity, within a similar size range. Faster‐growing S. salar will, thus, reach this size threshold at a relatively younger age.     

Identifying reproductive events using archival tags: egg‐laying behaviour of the small spotted catshark Scyliorhinus canicula

The use of archival depth telemetry as a means of remotely assessing the reproductive rates of free‐ranging fishes is explored. This is achieved by electronically tracking the vertical movements of individual female small spotted catsharks Scyliorhinus canicula in the natural environment, whilst simultaneously evaluating the temporal and vertical distributions of egg‐laying in this species. Distinctive patterns of short‐term (0·3–3·7 h), shallow‐water activity are documented in the time–depth profiles of female S. canicula that occur at an appropriate depth (1·0–2·3 m) and periodicity (every 10–12 days) to represent egg‐laying behaviour. Putative egg‐laying behaviour was exhibited simultaneously by two individually tracked female S. canicula during late‐spring and early‐summer. The results highlight that, provided species behaviour is suitable and complementary methods such as previous data, laboratory experiments and field surveys can be used to validate the patterns observed, archival depth telemetry offers an unobtrusive means by which egg production and egg‐laying behaviour of free‐living fishes can be estimated. As precise information regarding life‐history parameters is difficult to obtain for free‐ranging fish species, this technique could be used to improve the parameterization of species demographic models that are relevant to the management of wild fish populations.     

FIELD AND CULTURE STUDIES OF A POPULATION OF ENDARACHNE BINGHAMIAE (PHAEOPHYTA) FROM SOUTHERN CALIFORNIA1

Abundances of the erect, blade phase of Endarachne binghamiae J. Ag. (Scytosiphonales, Phaeophyta) varied seasonally at a southern California rocky intertidal site. Blade cover and density were much greater in the fall through early spring; blades were mostly absent from quadrats during the summer. Blade abundances were negatively correlated with both seasonal variations in seawater temperature and photoperiod. Laboratory culture studies failed to provide evidence for sexual reproduction. The life history appears to be of the “direct” type with plurangia-produced zooids germinating into crustose disks. Most disks developed erect blade clusters under spring/fall (17° C) and winter (13° C) temperatures over the range of natural photoperiods employed (14:10, 12:12, 10:14 h LD). In contrast, cultures held under the summer temperature (21° C) produced almost entirely crustose growths regardless of photoperiod. Similar results were obtained for cultures grown at 100 and 200 μE · m^?2· s^?1. E. binghamiae blades were fertile throughout the year and produced viable zooids indicating that reproductive seasonality did not influence the seasonal pattern of blade abundance. Culture and field studies suggest that the initiation of new erect blade clusters from crustose disks is confined to the cooler months of the year (winter and spring). The summer reduction or absence of E. binghamiae blades appears to be due to increased mortality rates and temperature constraints on the development of new erect bladed thalli. Hypothetical causes of mortality are desiccation stress, sand burial, increased grazing activity and a genetically-based short life span.     

The reproductive biology of some serolid isopods from the Antarctic

Summary Several species of serolid isopod from Signy Island, South Orkney Islands and also from South Georgia were studied, and aspects of their reproductive behaviour and associated morphology are discussed. InSerolis polita Pfeffer egg-laying took place at a mean age of 28 months, and the eggs were then incubated in a ventral marsupium for a period of 20 months before the release of juveniles in the spring. The mean egg dry weight at spawning was 0.68 mg forS. polita and 3.35 mg forSerolis cornuta Studer. The egg weight increased during marsupial development due to the uptake of minerals, but the total calorific content fell. Within each species brood size was linearly related to female size and the mean number of eggs varied from 40 forS. polita to 232 forSerolis pagenstecheri Pfeffer. There was no detectable marsupial mortality.S. polita appeared to breed repeatedly at two-year intervals, butS. cornuta probably only breeds once. The reproductive biology of Antarctic serolids is compared with that of temperate isopods, and it is shown that they produce larger young which is related to their larger adult size. The extremely protracted period of marsupial incubation is associated with the need to synchronize reproduction with the seasonal nature of Antarctic primary production.     

The reproductive ecology of the invasive Asian paddle crab,Charybdis japonica (Brachyura: Portunidae), in northeastern New Zealand

The Asian paddle crab, Charybdis japonica, native to the northwest Pacific, is an invasive species that has established populations within northeastern New Zealand. Here, we provide a detailed examination of C. japonica's reproductive seasonality and gametogenesis, sizes at physiological maturity, and sex ratios outside its native range. Trapping in the Weiti River Estuary, Whangaparoa (36°38.4′S, 174°43.6′E) from February 2010 to May 2012 indicated a male‐biased population. Reproductive seasonality largely depended on female condition, since males contained ripe spermatozoa year‐round. Female gametogenesis began during late autumn, with oocyte development occurring throughout winter to early spring. Based on the presence of mature gametes during late winter and the appearance of gravid females in early November, mating began during early spring, with spawning in mid‐spring. Spawning coincided with a large drop in the gonad index, and occurred between sea surface temperatures of 17–22°C and 11.75–13.5 h day length. Females remained reproductive for up to 5 months of the year. Histological observation suggested that CW50 (the carapace width at which 50% of the population is mature) for females was 46.39 mm, while CW50 for males could not be determined due to a paucity of immature males. Histological analysis also suggested that females of C. japonica could produce store sperm and multiple broods annually.     

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.     

Pair formation in the herbivorous rabbitfish Siganus doliatus

This study investigated the basis of pair formation in the abundant herbivorous rabbitfish Siganus doliatus on Orpheus Island, Great Barrier Reef. Pair formation was the most common social system in S. doliatus, with 67·4% of all individuals occurring in pairs. Pairs were stable (i.e. individuals remained with the same partner throughout the study) and pair members were found within 5 m of each other 82·9% of the time. Of the examined pairs, 25% were homosexual resulting in a proportion of heterosexual pairs (75%) that was significantly lower than expected if pairs were formed solely for reproductive reasons. Therefore, although reproduction appears to be the main driver of pair formation in S. doliatus, other factors are likely to influence this behaviour. The high density of individuals on the reef crest (5·7 ± 0·9 individuals 200 m^?2; mean ± s.e .) and extensively overlapping home ranges of pairs indicated that the defence of territories plays no role in pair formation. Instead, it appears that pair formation in S. doliatus is driven, in part, by other, non‐reproductive, ecological factors. It is suggested that pair formation allows for increased vigilance against predation and enables S. doliatus to execute a novel feeding behaviour.     

Using life‐history information to assess potential effects of shrimp trawling on small fishes

The current study presents information on size distributions, size at recruitment to the fishery, size at maturity and patterns of reproduction for several small benthic fishes caught as by‐catch in the southern Gulf of California (Mexico) shrimp trawl fishery: sand perch Diplectrum spp., lumptail searobin Prionotus stephanophrys, bigscale goatfish Pseudupeneus grandisquamis and silver stardrum Stellifer illecebrosus. Pseudupeneus grandisquamis, P. stephanophrys and S. illecebrosus populations were all sexually dimorphic in size. Total‐length (L_T)‐based analyses did not provide reliable information on survival and growth. The majority of sampled P. grandisquamis and S. illecebrosus were caught before reproductive maturity, whereas the majority of Diplectrum spp. and almost all P. stephanophrys were mature when caught. L_T at 50% gear retention (L_Tc, mm) v. 50% maturity (L_Tm, mm): Diplectrum spp. 124·53 v. 131·43; P. grandisquamis 90·98 v. 135·20; S. illecebrosus 82·55 v. 137·30. L_Tc for P. stephanophrys was 104·73, but L_Tm could not be modelled for this species as almost all captured individuals were mature. Diplectrum spp., P. grandisquamis and S. illecebrosus were indeterminate spawners, whereas P. stephanophrys appeared to be a determinate spawner. Sex ratios were equal for each of the gonochoristic species. In general, the gonado‐somatic index (I_G) increased with increasing L_T for all except P. stephanophrys, where I_G decreased with increasing L_T for both males and females. Mature individuals of all taxa were found throughout the sampling period (September to March), and I_G increased with sample day for all except females of P. grandisquamis. The current data suggest the potential for fishery effects on sampled populations of P. grandisquamis and S. illecebrosus.     

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.     

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.     

Changes in growth and maturation parameters of Pacific sardine Sardinops sagax collected off California during a period of stock recovery from 1994 to 2010

Whether fluctuation in density influenced the growth and maturation variables of three aggregated cohorts (fish born during the 1986–1993, 1996–2003 and 2004–2008 periods) of Pacific sardine Sardinops sagax caeruleus collected off the Californian coast from 2004 to 2010 was investigated. Using a von Bertalanffy mixed‐effects model with aggregated cohorts as covariates, estimated growth rate significantly covaried with aggregated cohorts. Growth rate (K) was modelled as a fixed effect and estimated to be 0·264 ± 0·015 (±s.e ). Statistical contrasts among aggregated cohorts showed that the 1996–2003 cohorts had a significantly lower growth rate than the other two aggregated cohorts. The theoretical age at length zero (t₀) and the standard length at infinity (L_S∞) were modelled as random effects, and were estimated to be ?2·885 ± 0·259 (±s.e ) and 273·13 ± 6·533 mm (±s.e ). The relation of ovary‐free mass at length was significantly different among the three aggregated cohorts, with the allometric coefficient estimated to be 2·850 ± 0·013 (±s.e ) for the S. sagax population. The age‐at‐length trajectory of S. sagax born between 1986 and 2008 showed strong density dependence effects on somatic growth rates. In contrast to the density‐dependent nature of growth, the probability to be mature at‐size or at‐age was not significantly affected by aggregated cohort density. The size and the age‐at‐50% maturity were estimated to be 150·92 mm and 0·56 years, respectively. Stock migration, natural fluctuations in biomass and removal of older and larger S. sagax by fishing might have been interplaying factors controlling growth parameters during 1986–2010.     

Life history and initial assessment of fishing impacts on the by‐catch species Dules auriga (Teleostei: Serranidae) in southern Brazil

The life history of Dules auriga, a small hermaphrodite serranid species inhabiting deep waters and a frequent component of the discarded catch of bottom trawling in southern Brazil, was studied to assess the fishery effects on the stock through the estimation of the remaining spawning‐potential ratio. Sampling was conducted throughout a year and included specimens to determine sex, maturity and age. Age was validated by the edge type and marginal‐increment analysis. The oldest and the largest individuals were 9 years and 195 mm total length. Growth parameters fitted to the von Bertalanffy equation were L_∞ = 178·34 mm, k = 0·641 year^?1 and t₀ = ?0·341 years. Length and age at first maturity were 140·72 mm and 2 years, respectively. The reproductive season was throughout the austral spring and summer. The assessment of the effects of fishing showed that it may have resulted in a loss of 50% of the spawning potential. This loss may be higher when taking into account the uncertainty in the life‐history parameters and could be considered of concern for the population. Fast growth, moderate longevity, long spawning season, small size and age at maturity make D. auriga relatively resilient to the removal of biomass by fishing. When considering the uncertainty, however, the losses of the spawning potential have been severely reducing the population resilience in the face of ecosystem changes.