Spawning Dynamics and Size Related Trends in Reproductive Parameters of Southern Bluefin Tuna,Thunnus maccoyii

Knowledge of spawning behaviour and fecundity of fish is important for estimating the reproductive potential of a stock and for constructing appropriate statistical models for assessing sustainable catch levels. Estimates of length-based reproductive parameters are particularly important for determining potential annual fecundity as a function of fish size, but they are often difficult to estimate reliably. Here we provide new information on the reproductive dynamics of southern bluefin tuna (SBT) Thunnus maccoyii through the analysis of fish size and ovary histology collected on the spawning ground in 1993–1995 and 1999–2002. These are used to refine previous parameter estimates of spawning dynamics and investigate size related trends in these parameters. Our results suggest that the small SBT tend to arrive on the spawning ground slightly later and depart earlier in the spawning season relative to large fish. All females were mature and the majority were classed as spawning capable (actively spawning or non-spawning) with a very small proportion classed as regressing. The fraction of females spawning per day decreased with fish size, but once females start a spawning episode, they spawned daily irrespective of size. Mean batch fecundity was estimated directly at 6.5 million oocytes. Analysis of ovary histology and ovary weight data indicated that relative batch fecundity, and the duration of spawning and non-spawning episodes, increased with fish size. These reproductive parameter estimates could be used with estimates of residency time on the spawning ground as a function of fish size (if known) and demographic data for the spawning population to provide a time series of relative annual fecundity for SBT.     

Reproductive guilds (maturation,spawning frequency and fecundity) in the black pomfret,Parastromateus niger (Carangidae) in the Kuwaiti waters of the Arabian Gulf

Oogenesis, oocyte maturation pattern, spawning rhythm, spawning frequency, batch fecundity and oocyte diameter–frequency distribution of the black pomfret, Parastromateus niger (Bloch, 1795) in Kuwaiti waters were investigated from October 2003 to September 2005, using histological and morphological methods. The process of development is divided into four major phases: (i) primary growth phase; (ii) secondary growth phase; (iii) maturation phase; and (iv) spawning phase, followed by the regressed phase. Development of the yolky oocyte is an asynchronous process resulting, by the time of oocyte maturation, in a clear differentiation between a ready batch of oocytes (ready for spawning) and a reserve pool. Consequently, P. niger is capable of spawning multiple times throughout the reproductive season. Spawning frequency estimates, based on final oocyte maturation (FOM) method indicated that the species spawns once every 2.8 days during an 8‐month spawning season lasting from February to September, with a potential annual number of 22.4 spawns. Batch fecundity (BF) (2132–2001 648, mean 406 010 eggs), was significantly positively related to both standard length (SL) (P < 0.05) and ovary‐free body weight (OFBW) (P < 0.05), both parameters being good predictors of BF (r² = 30.8% for SL, from 22 cm onwards, and r² = 29.6% for OFBW, from 129.5 g onwards). No significant differences in monthly BF were found throughout the spawning season. Relative batch fecundity was 336 eggs/g OFBW; thus, estimate for potential annual relative batch fecundity was 7526 eggs g^?1 OFBW. The oocyte diameter–frequency distribution analysis revealed a multimodal distribution (at 100–200, 300–400 and 500–700 μm), confirming the evidence of multiple spawning.     

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.     

Success and failure assessing gonad maturity in sequentially hermaphroditic fishes: comparisons between macroscopic and microscopic methods

For two protogynous hermaphrodite fish species, the performance of visual gonad analysis techniques was evaluated to determine when the use of macroscopic methods was sufficient and when microscopic techniques were necessary. Simple macroscopic gonad analysis was found to be a powerful tool for distinguishing sex and whether or not females were spawning capable or ripe for black sea bass Centropristis striata (n = 1443) and red porgy Pagrus pagrus (n = 980), often producing results that were in close agreement with more complex and expensive microscopic techniques. Estimates of key reproductive variables, such as size‐dependent sex‐change ogives, spawning season duration, spawning fraction and batch number, were also very similar or equal between methods. Apparent seasonal spawning activity was also predicted similarly by each method and the patterns were highly correlated with seasonal patterns in gonado‐somatic indices. In contrast, distinguishing between immature females and those that were mature, but inactive, proved difficult when using macroscopic methods and, in these cases, predictions often differed from those produced microscopically. In turn, maturity ogives differed significantly between methods for C. striata (maturity ogives could not be generated for P. pagrus as nearly all fish encountered were mature). Agreement rates among male phases were also very low. Macroscopic methods were able to identify signs of sex transition in very advanced specimens, but early signs were only evident microscopically. While much more detail is visible microscopically, here several population‐scale parameters important for fisheries management were estimated equally well with the unaided eye for C. striata and P. pagrus. For comprehensive, fishery‐independent surveys and long‐term research programmes in particular, determining when microscopic techniques are and are not necessary can greatly improve efficiency and reduce costs without compromising data quality.     

Reproductive biology of Galaxias maculatus (Jenyns 1842) in the Río Ovando,a high‐latitude environment in southernmost Patagonia,Argentina

This study establishes the reproductive cycle, batch fecundity and oocyte diameter of a diadromic population of the puyen, Galaxias maculatus, in the Río Ovando, Tierra del Fuego, (54°S), at the southernmost limit of its distribution. Given the short ‘favourable season’ in relation to other populations of the species (in terms of temperature, photoperiod and food availability), the study also explores the trade‐offs between feeding and oocyte production and between phases of growth and reproduction. The reproductive cycle was analyzed by the monthly evolution of the proportion of gonadal maturity stages (determined from microscopic examination of the gonads). Oocytes were measured using a micrometric ocular scale and classified by diameter and morphological characteristics; absolute fecundity was established as the total number of hydrated oocytes per ovary. The studied population exhibits a repetitive spawning strategy, given the coexistence of post‐ovullatory follicles and hydrated oocytes in histological sections and the presence in ripe ovaries of an intermediate cohort of yolked oocytes ready for hydration. Individual fecundity is lower (1422 ± 422 oocytes/ovary) than in other puyen populations, but the egg production increases through individual repetitive spawnings during the protracted spawning period (from October to February). Females attain larger sizes than those of other South American populations (the largest female reaching 115 mm total length), maximizing its potential fecundity. The present paper contributes to the knowledge of the variability of reproductive traits of G. maculatus in relation to diadromic populations, given that the bulk of information in South America refers to landlocked populations.     

Reproductive biology of squaretail coralgrouper Plectropomus areolatus using age‐based techniques

The squaretail coralgrouper Plectropomus areolatus was identified as a fast‐growing, early maturing and relatively short‐lived aggregation‐spawning epinephelid. Examinations of sectioned otoliths found females and males first maturing at 2 and 3 years, respectively, suggesting protogynous hermaphroditism; however, no transitionals were observed in samples. Age distribution for the two sexes was similar and both were represented in the oldest age class; however, significant sex‐specific differences in size‐at‐age were identified. Both sexes fully recruit into the fishery at age 4 years and reach 90% of asymptotic length by age 3 years. Underwater visual assessments, combined with the gonado‐somatic indices, revealed a 5 month reproductive season, with interannual variability observed in the month of highest density within the spawning aggregation. Catch restrictions on adults during spawning times and at reproductive sites, combined with gear‐based management and enhanced enforcement, are recommended to maintain spawning stocks. Based on the available evidence, the sexual pattern for this species is unresolved.     

Reproductive Biology of Three Commercially Important Hemiramphid Species in South-eastern Australia

Synopsis The reproductive biology of eastern sea garfish Hyporhamphus australis, eastern river garfish H. regularis ardelio, and snub-nosed garfish Arrhamphus sclerolepis were described throughout their respective ranges in the coastal waters of New South Wales (NSW), Australia. Peaks in gonadosomatic indices indicated that spawning of eastern sea garfish occurred in late spring and early summer (November–December) on the south coast of NSW, and in winter and early spring (June–September) on the north coast. Eastern river garfish spawned between July and December in NSW estuaries and snub-nosed garfish spawned between October and January in the Clarence River. The sex ratios in commercial catches of eastern sea garfish from the north coast of NSW were biased toward male fish, but approached equality for fish caught from the south coast. Sex ratios were significantly biased toward female snub-nosed garfish, and female eastern river garfish from all estuaries sampled. Mean (±SE) batch fecundity was 1498 ± 110 (range: 98 – 3449) ripe oocytes per female for eastern sea garfish, 917 ± 36 (range: 102 – 2268) ripe oocytes per female for eastern river garfish, and 716 ± 104 (range: 20 – 2956) ripe oocytes per female for snub-nosed garfish across the range of mature sizes examined. There was a linear relationship between batch fecundity and fish size for all three species of garfish. Eastern sea garfish approached 50% maturity at a larger size than snub-nosed, or eastern river garfish. Size at 50% maturity was similar for male and female eastern river and snub-nosed garfish, but male eastern sea garfish matured at a significantly smaller size than females. All three species appear capable of spawning in the spawning season immediately following the one in which they were born. Mature female fish of all three species had distributions of oocyte diameters consisting of three or four modes, which strongly suggests asynchronous oocyte development and a multiple spawning strategy during the spawning season. Implications for the management of garfish fisheries in NSW are also discussed.     

Intra‐population trends in the maturation and reproduction of a temperate marine herbivore Girella elevata across latitudinal clines

Latitudinal variation in the reproductive characteristics of a temperate marine herbivore, rock blackfish Girella elevata, was examined from three regions of the south‐eastern Australian coast. Biological sampling covered 780 km of coastline, including the majority of the species distribution. The sampling range incorporated three distinct oceanographic regions of the East Australian Current, a poleward‐flowing western boundary current of the Southern Pacific Gyre and climate‐change hotspot. Girella elevata are a highly fecund, group synchronous (multiple batch)‐spawner. Mean fork length (L_F) and age at maturity were greater for females than males within all regions, with both male and female G. elevata of the southern region maturing at a greater size and age than those from the central region. Estimates of batch fecundity (F_B) were greatest in the northern and southern regions, relative to the central region where growth rates were greatest. Significant positive relationships were observed between F_B and L_F, and F_B and total fish mass. Gonado‐somatic indices indicated latitudinal synchrony in spawning seasonality between G. elevata at higher latitudes, spawning in the late austral spring and summer. A late or prolonged spawning period is evident for G. elevata from the northern region. Juvenile recruitment to intertidal rock pools within the central and southern regions was synchronous with the spawning season, however, no juveniles were found within the northern region. The implications of latitudinal variation in reproductive characteristics are discussed in the context of climate and oceanographic conditions of south‐east Australia.     

The tempo of reproduction in Hyphessobrycon pulchripinnis(Characidae), with a discussion on the biology of ‘multiple spawning’ in fishes

Synopsis We describe the short-term patterns of egg production and release in the lemon tetra, Hyphessobrycon pulchripinnis (Characidae) as observed over a six month aquarium study and then use our results and those of others to both describe general patterns and derive comparative predictions. Female lemon tetras ovulated about once every four days; differences among individuals were small and inconsistent. As in other species, the probability of ovulation depended strongly on time since last ovulation, indicative of an ovarian cycle; furthermore we found no obvious long-term patterns of ovulation within females and batch fecundity was independent of the length of the two previous interovulation intervals and of the one following. Each batch of ovulated eggs was released over an average of 23.1 spawning acts, beginning as soon as the lights went on in the morning. The number of spawning acts increased with ovulation fecundity but decreased with the number of other females spawning in the tank on that day. The mass of eggs produced every four days was greater than that of the remaining ovary, and the mass of eggs that could be produced in a season was greater than that of the female. This latter observation highlights the most important consequence of repeated reproduction within a season - increased reproductive output - and leads to predictions associating it with less seasonal environments (e.g. low latitudes), as well as with small ovaries and small body size. The significance of releasing a batch of eggs over many spawning acts remains unknown.     

Comparative oocyte morphology and fecundity of five characid species from São Francisco River basin,Brazil

The morphology of vitellogenic oocytes and the batch fecundity of five tropical forage species belonging to the family Characidae, were studied in 104 late‐maturing ovaries. Significant morphological differences between vitellogenic oocytes and ovarian follicles were found. The lowest batch fecundity values were recorded in Hemigrammus marginatus (480 ± 163) and Orthospinus franciscensis (1701 ± 562), which were the smaller species in terms of total length, body weight and oocyte diameter. The highest batch fecundity value was observed in Tetragonopterus chalceus (8384 ± 3944) having the highest GSI and oocyte diameter. Batch fecundity and gonad weight was highly correlated followed by body weight and total length. Relative fecundity was estimated per unit total length, body weight and gonad weight. The wide variation in fecundity observed between the specimens and the species analysed is possibly related to the multiple spawning reproductive strategy of these fishes. Since T. chalceus have higher values of batch and relative fecundity, it is concluded that this species has a higher reproductive potential than the other forage species studied.     

Gonad maturation and spawning of cobia,Rachycentron canadum (Linnaeus, 1766) off the Dungun coast,Malaysia

The reproductive biology of cobia, Rachycentron canadum, from the coastal waters of Dungun, Malaysia was studied from June 2014 to May 2015. From commercial trawls, a total of 201 samples (combined sexes) were collected (fork lengths [FL] 37.5–124.0 cm; body weights 0.5–20.4 g). The overall sex ratio of females to males was 1:0.9, which was not significantly different (χ² = 2.12, df = 1; p < .05). Estimations of length at 50% maturity (L₅₀) showed that both sexes matured at approximately 75 cm FL; estimated spawning frequency was 6 days. Mean batch fecundity (BF) ranged from 0.55 to 4.32 million eggs. The average number of eggs per gram of ovary was from 2,100 to 5,400 eggs, and relative fecundity 147 eggs/g. There was a weak positive correlation (r² = .48) between BF and female FL as well as BF with an ovary‐free body weight (r² = .56), possibly due to females being in a continuous spawning condition and some possibly half‐spent, based on the histological examination of the female gonads. Despite cobia being asynchronous spawners, the gonadosomatic index in both males and females showed peaks in June, November, and particularly March. Based on histological examination, spawning‐capable males were encountered throughout the study period, whereas spawning‐capable females in the late developing subphase were found mostly in March and April. This is the first study on the reproductive aspects of cobia in Malaysian waters.     

Spawning frequency in the eastern Australian river garfish, Hyporhamphus regularis ardelio (Hemiramphidae)

Patterns in the timing of spawning of the eastern Australian river garfish Hyporhamphus regularis ardelio were investigated by sampling fish throughout 24?h during their peak spawning period. Patterns in gonadosomatic indices, the presence of hydrated oocytes and post-ovulatory follicles in histological ovary sections, together with previously demonstrated polymodal distributions of oocytes undergoing final maturation, indicated that river garfish were capable of spawning at any time during the 24?h period. Spawning had either recently occurred or was imminent in all females sampled suggesting that river garfish could spawn multiple times in a given season. Although energy resources are used in the production of low numbers (<2500) of large (~2.5?mm diameter) eggs per batch, total fecundity during the reproductive season may be elevated through multiple spawning events. Implications of these findings on current fishery management regulations are discussed.     

Spawning frequency, fecundity, egg weight and spawning type of silver pomfret, Pampus argenteus (Euphrasen) (Stromateidae), in Kuwait waters

Silver pomfret, Pampus argenteus, were collected by fishing with drift gillnets on one spawning ground in Kuwait waters during 1998–2000. Fish size frequency, sex ratio, maturation cycle, spawning frequency, fecundity and egg weight were assessed. The length–weight relationship differed between sexes whereby females were significantly bigger than males. Spawning started in mid‐May and continued until early October. During this time the water temperature ranged from 26.0 to 32.8°C, salinity was ? 39.0‰ and water depth ranged between 5 and 12 m. Large females spawned earlier than young spawners and the overall percentage of males during the spawning period was 70.3%. Spawning occurred after 13.00 h, with peak spawning between 15.00 and 18.00 hours during outgoing tide. Mean daily spawning frequency amounted to 63.2%. Spawning activity was found to be associated with the lunar cycle and spawnings were concentrated during the first and third quarters of the moon period, indicating a semilunar reproduction cycle. It was concluded that a female would spawn at least six times during the season. No change was observed in relative fecundity during the peak spawning season (June–August). Average relative batch fecundity was 176.3 eggs g^?1 somatic weight (SW), corresponding to a relative total fecundity of 1058 eggs g^?1 SW, which is 1.5 times higher than estimates obtained from counting the standing stock of oocytes. Bigger fish produced heavier eggs and the egg weight decreased as the spawning season progressed. Based on gonadal cycles, oocyte size frequency distribution and total fecundity, we concluded that silver pomfret is a multiple batch spawner with indeterminate fecundity.     

Reproductive characteristics of pike‐characids Boulengerella cuvieri (Ctenoluciidae) in the middle Xingu River,Eastern Amazon

This study evaluated the reproductive characteristics of pike‐characids, Boulengerella cuvieri, during the hydrological cycle in the Xingu River, eastern Amazon. The sex ratio was 1:1 and the gonado‐somatic index and relative frequency of maturation stages indicate a short breeding season of single phase spawning that coincides with a filling and flood period. Mean standard length at first sexual maturity for female B. cuvieri was estimated to be 22·9 cm.     

Reproductive strategies of a temperate Australian sciaenid (teraglin,Atractoscion atelodus)

Worldwide there have been declines in stocks of exploited sciaenids. Despite a long history of exploitation, there has been no study of the life history characteristics of the fishery-important sciaenid, teraglin, Atractoscion atelodus. This study describes the reproductive characteristics of A. atelodus within the major area of its distribution and fishery, New South Wales (NSW) Australia. Characteristics studied included the size and age at maturity, fecundity, spawning mode and season, and these are compared to congeneric species. Atractoscion atelodus displays year-round batch spawning behaviour with asynchronous oocyte development and indeterminate fecundity. This is unusual for sciaenids, which typically display spring-summer spawning and only over a few months. The length and age at which 50% of the population matures for both males and females is 36 cm fork length (FL) and 1 year, a smaller size compared to other closely related sciaenids. While the species displays resilient reproductive characteristics such as protracted year-round spawning, constant supply of vitellogenic oocytes and relatively small/young age at maturity, there are also characteristics that make it susceptible to over-exploitation. These include highly female skewed sex ratios, as well as their schooling and voracious feeding behaviour leading to the potential for high catch rates.     

Energy allocation strategy of skipjack tuna Katsuwonus pelamis during their reproductive cycle

The lipid composition of somatic and reproductive tissues was determined for female skipjack tuna Katsuwonus pelamis caught in the western Indian Ocean between latitude 10° N and 20° S and longitude 40° and 70° E. The highest total lipid (TL) contents were in the liver and gonads, with white muscle levels approximately three‐fold lower. Three lipid classes dominated: triacylglycerols (TAG), sterol esters and wax esters (SE‐WE) and phospholipids (PL). Collectively, these accounted for between 70 and 80% of TLs. Changes in lipid concentrations were evaluated over the maturation cycle. Immature fish had the lowest gonad and liver TL levels; concentrations of TL, TAG, SE‐WE and PL accumulated from immature to mature (spawning‐capable) phase, reflecting sustained vitellogenic activity of the liver and a transfer of lipids to developing oocytes from the onset of vitellogenesis. Gonado‐somatic and hepato‐somatic indices were positively correlated with each other and positively related to TL in the gonads and liver. Fulton's condition index and lipid concentrations in muscle did not vary significantly over the maturation cycle; fat content in the main storage tissues was undepleted as the ovary developed. Hence, K. pelamis apparently supports reproduction directly from food intake over the breeding season. In the gonads, reserve lipids (SE‐WE and TAG) and sterols were related to batch fecundity but this was not the case for somatic and hepatic tissues. These results suggest that K. pelamis utilizes an income breeding strategy.     

Reproductive biology of Hemiramphus brasiliensis and H. balao (hemiramphidae): maturation,spawning frequency,and fecundity

Analyses of life-history data show that both the size-specific batch fecundities and the age-specific spawning frequencies differ for two halfbeak species, Hemiramphus brasiliensis, the ballyhoo, and H. balao, the balao. Halfbeak ages were determined from sectioned otoliths; histological data was used to describe oocyte development and estimate spawning frequency; and batch fecundity was measured from counts of whole oocytes in final maturation. Hemiramphus brasiliensis lived longer (4 versus 2 years) and had a higher survival rate (14.9% versus 7.5% annually) than H. balao did. Of the two species the larger and longer-lived congener, H. brasiliensis, reached sexual maturity at a larger size (fork length 198 versus 160 mm). The spawning period of age-0 females was strongly related to season, whereas spawning by older females occurred throughout the year. Reproduction by both species peaked during late spring or early summer, and all mature females were spawning daily during April (H. brasiliensis) or June (H. balao). This is the first demonstration of iteroparity for the family Hemiramphidae. H. brasiliensis had a lower batch fecundity (about 1164 versus 3743 hydrated oocytes for a 100-g female) than H. balao did. Such low batch fecundities are typical of the order Beloniformes, but quite different from those of other fishes that live in association with coral reef habitats. H. balao's higher batch fecundity is consistent with the life-history theory that predicts higher numbers of eggs for shorter-lived species; this is possible because H. balao produces smaller hydrated oocytes than H. brasiliensis (modal diameter about 1.6 versus 2.4 mm). The high spawning frequency of Hemiramphus species compensates for their low batch fecundity. The annual fecundity of both species is similar to that of other reef fish species, after adjusting for body size and spawning frequency. The lifetime fecundity of H. balao was very similar to that of H. brasiliensis, after accounting for the differences in survival for each species. This suggests a fine tuning of different reproductive traits over the entire life cycle that results in roughly equivalent lifetime fecundity for both species.     

Reproduction mode of European Bitterling (Rhodeus amarus,Bloch, 1782) determined through rapid oocyte counts and size determination using digital imaging

This study focuses on assessing the reproduction mode of an important model species for evolutionary and behavioural ecology by using digital image analysis: the European bitterling Rhodeus amarus (Bloch, 1782). Specifications of mode of reproduction were determined using oocyte size distribution, seasonal dynamics in mean oocyte diameter and total number of oocytes in ovary samples gained from April to July 2007. The rapid oocyte count was enabled by using lucia image analysis software, which also provided measurement and colour estimations of oocytes. Bitterling ovaries showed features typical for indeterminate spawners, i.e. a continuous distribution of oocyte size over the reproductive season and recruitment of new pre‐vitelogenic oocytes in the second half of the reproductive season. These results are consistent with the view that the European bitterling is a batch spawning fish with indeterminate fecundity.     

Analysis of the cycle of reproduction of Sardina pilchardus (Walbaum, 1792) off the Moroccan Atlantic coast

This work focuses on aspects of reproductive biology of Sardina pilchardus from the Atlantic coast of Morocco. The mean values of batch fecundity estimated for the species is 23150(+/-1301) oocytes for a mean size of 19.5(+/-0.49) cm, the mean relative fecundity being 346(+/-7.34) oocytes per gram of female without ovary. Batch fecundity increases with total length and body weight without ovary. Sizes at first sexual maturity (L50) are reached for males and females at 15.8(+/-0.29) cm and 15.8(+/-0.35) cm, respectively. The spawning period for the population extends between October and July and the spawning peak occurs from October to February. However, the small sardines (14.5-17 cm) in their first reproduction spawn between November and June, whereas larger fish (17.5-25 cm) spawn between October and July. The factor of condition (K) increased in summer during the sexual resting phase. It is weak in winter during the period of reproduction. Regarding, the sex ratio, there was no significant difference in the number of males and females.     

The reproductive cycle and its control; frequency of spawning and fecundity in Blennius pholis L.

Spawning in Blennius pholis L. is seasonal occurring from March to July in the population of the South Gower beaches. There is evidence that both temperature and daylength regulate the reproductive cycle. Cyclical changes in the ovary during the reproductive cycle are studied by analysis of the changing size frequency distribution of the oocytes. Adults are capable of spawning a minimum of eight times in any one spawning season, and females > 11 cm in length have an estimated fecundity of 8,000 eggs.