Reproductive Dynamics and Potential Annual Fecundity of South Pacific Albacore Tuna (Thunnus alalunga)

The reproductive biology of albacore tuna, Thunnus alalunga, in the South Pacific Ocean was investigated with samples collected during broad-scale sampling between 2006 and 2011. Histology was done in a single laboratory according to standard protocols and the data analysed using generalized linear mixed-effects models. The sex ratio of albacore was female biased for fish smaller than approximately 60 cm FL and between 85 and 95 cm, and progressively more male biased above 95 cm FL. Spawning activity was synchronised across the region between 10°S and 25°S during the austral spring and summer where sea surface temperatures were ≥24 °C. The average gonad index varied among regions, with fish in easterly longitudes having heavier gonads for their size than fish in westerly longitudes. Albacore, while capable of spawning daily, on average spawn every 1.3 days during the peak spawning months of October to December. Spawning occurs around midnight and the early hours of the morning. Regional variation in spawning frequency and batch fecundity were not significant. The proportion of active females and the spawning fraction increased with length and age, and mature small and young fish were less active at either end of the spawning season than larger, older fish. Batch fecundity estimates ranged from 0.26 to 2.83 million oocytes with a mean relative batch fecundity of 64.4 oocytes per gram of body weight. Predicted batch fecundity and potential annual fecundity increased with both length and age. This extensive set of reproductive parameter estimates provides many of the first quantitative estimates for this population and will substantially improve the quality of biological inputs to the stock assessment for South Pacific albacore.     

Batch fecundity and spawning frequency of invasive Hemiculter leucisculus (Basilewsky, 1855) in Erhai Lake,China

Hemiculter leucisculus are multiple spawners with indeterminate fecundity, although previous fecundity estimates have assumed a determinate spawning pattern. Batch fecundity and spawning frequency of H. leucisculus were studied in Erhai Lake, China, in which the fish, as an exotic species, has become a successful colonizer. Spawning frequency was 16.05 %, as determined from the percent of females with postovulatory follicles 12 to 36 h old during the peak reproductive period (from May 27 to August 8). The average interval between spawning was 6.2 days and there were more than 16 total spawning batches during the peak reproductive period. The average batch fecundity (mean ± SD) was 11,934?±?5,921 hydrated oocytes in 40 females (standard length or SL: 9.1–14.4 cm), while relative batch fecundity was 560?±?137 eggs g^?1 wet weight. The total potential annual fecundity was approximately 190,944 oocytes over the whole spawning season and was much higher than the estimated standing stock (31,585 oocytes) of yolked oocytes (36 females, SL: 9.5–16.2 cm) at the beginning of the spawning season in April 2010. This high annual fecundity is likely to have contributed to successful invasion of Erhai Lake by H. leucisculus.     

Reproduction in an invasive exotic catfish Pterygoplichthys disjunctivus in Volusia Blue Spring,Florida, U.S.A.

Morphology and reproductive strategies of a population of invasive loricariid catfish, Pterygoplichthys disjunctivus, in Volusia Blue Spring, Florida, were examined using morphological characters, gonado‐somatic indices and batch fecundity. This species has cystovarian ovaries, a left ovary that is significantly larger than the right ovary, and is highly fecund. Ovaries contained multiple oocyte size classes and completely spent females were not found, both of which indicate that this species is a multiple spawner. Reproductive patterns changed significantly between the first (2005) and the last (2007) years of this study; female fish became capable of spawning at a smaller size, a greater proportion of the captured population were capable of spawning each year and female batch fecundity increased, as did male and female gonado‐somatic indices. Plasticity of P. disjunctivus reproductive patterns is discussed in relation to resource availability and environmental stability.     

Spawning frequency and batch fecundity of horse mackerel, Trachurus trachurus (L.), in the Saronikos Gulf (Greece)

Horse mackerel, Trachurus trachurus (L.), is a multiple spawning fish with most probably an indeterminate fecundity. Histological sections of gonads were used to identify the hydrated oocytes, the migratory-nucleus stage oocytes, and the new and old post-ovulatory follicles. Spawning frequency determination, based on the mean percentage estimation of the females that occurred in different spawning states such as migratory-nucleus stage oocytes and post-ovulatory follicles during two successive reproductive periods, was found to average once every 5.8 and 4.8 days, respectively. High spawning frequency was observed at the peak of spawning. Relative batch fecundity of 205 oocytes/g fish weight was estimated by the hydrated and migratory-nucleus method. Since potential annual spawnings were found to be equal to 16, potential annual relative fecundity could be 3280 oocytes/g fish weight.     

Maturity Ogives for South Pacific Albacore Tuna (Thunnus alalunga) That Account for Spatial and Seasonal Variation in the Distributions of Mature and Immature Fish

Length and age at maturity are important life history parameters for estimating spawning stock biomass and reproductive potential of fish stocks. Bias in estimates of size and age at maturity can arise when disparate distributions of mature and immature fish within a population are not accounted for in the analysis. Here we investigate the spatial and temporal variability in observed size and age at maturity of female albacore tuna, Thunnus alalunga, using samples collected across the South Pacific. Maturity status was identified using consistent histological criteria that were precise enough to allow for mature but regenerating females to be distinguished from immature females during the non-spawning season, permitting year-round sampling for maturity estimation in albacore. Using generalised linear mixed models, we found that the proportion of mature females at length varied significantly with latitude and time of year. Specifically, females at northern latitudes (∼10–20°S, where spawning occurs) were mature at significantly smaller lengths and ages than females at southern latitudes (∼20–40°S), particularly during the spawning season (October–March). This variation was due to different geographic distributions of mature and immature fish during the year. We present a method for estimating an unbiased maturity ogive that takes into account the latitudinal variation in proportion mature at length during a given season (spawning or non-spawning). Applying this method to albacore samples from the western region of the South Pacific gave a predicted length at 50% mature of ∼87 cm fork length (4.5 years).     

Demographic Structure,Sex Ratio and Growth Rates of Southern Bluefin Tuna (Thunnus maccoyii) on the Spawning Ground

The demographics of the southern bluefin tuna (SBT) Thunnus maccoyii spawning stock were examined through a large-scale monitoring program of the Indonesian longline catch on the spawning ground between 1995 and 2012. The size and age structure of the spawning population has undergone significant changes since monitoring began. There has been a reduction in the relative abundance of larger/older SBT in the catch since the early 2000s, and a corresponding decrease in mean length and age, but there was no evidence of a significant truncation of the age distribution. Pulses of young SBT appear in the catches in the early- and mid-2000s and may be the first evidence of increased recruitment into the spawning stock since 1995. Fish in these two recruitment pulses were spawned around 1991 and 1997. Size-related variations in sex ratio were also observed with female bias for fish less than 170 cm FL and male bias for fish greater than 170 cm FL. This trend of increasing proportion of males with size above 170 cm FL is likely to be related to sexual dimorphism in growth rates as male length-at-age is greater than that for females after age 10 years. Mean length-at-age of fish aged 8–10 years was greater for both males and females on the spawning ground than off the spawning ground, suggesting that size may be the dominant factor determining timing of maturation in SBT. In addition to these direct results, the data and samples from this program have been central to the assessment and management of this internationally harvested stock.     

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.     

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.     

Age, reproduction and fecundity of the spined loach Cobitis taenia L. (Pisces, Cobitidae) from Lake Klawój (Poland)

This is the first study concerning the features of the reproduction process of the karyologically identified spined loach C. taenia (2n=48). The histology of 71 ovaries, and gonadosomatic index (GSI) of karyologically identified spined loach Cobitis taenia L. from Lake Klawój (Northern Poland) were examined. The absolute and relative fecundity of 25 females was estimated by gravimetric method. The age of fish was determined according to the annual increments of otholits. The spawning of C. taenia from Lake Klawój took place from May to July, at a water temperature exceeding 18.5 degrees C. The GSI values at the beginning of the reproduction period ranged from 7 to 19%. The average absolute fecundity of females was 2078 eggs, with the number ranging from 869 to 3371 eggs. High individual variability in the gonad histology and the GSI values during the reproductive period was observed. Such variability could be the result of beginning the reproduction process in the fish at various times and, probably, due to the various numbers of batches laid and various numbers of eggs per batch.     

Reproductive biology of wahoo, Acanthocybium solandri, off eastern Australia

A dearth of basic biological information for wahoo, Acanthocybium solandri, currently hinders the ability of scientists and managers to assess population sustainability and appropriately manage the dramatically increasing global catch. This study examined the gonads of 382 wahoo collected off eastern Australia during 2008–2011 to quantify their reproductive biology in the region. The overall sex ratio of the sample was 3.2:1 (females:males), however this differed significantly among fishing sectors and areas. The estimated fork length at which 50 % of female wahoo reach maturity was 1,046 mm. Similar to the Atlantic Ocean, female wahoo have a protracted summer spawning season during October-February. The mean spawning frequency of female wahoo was uncertain but may be approximately 2–3 days, with evidence of fish actively spawning on consecutive days. Batch fecundity of females was positively correlated with fish size and estimates ranged between 0.65 and 5.12 million oocytes. Relative fecundity was estimated at 122.0 (±9.7) oocytes per gram of ovary free body weight and did not differ with fish size or throughout the spawning season. Estimation of reproductive parameters such as size- and age-at-maturity may facilitate the construction of per-recruit stock assessments of wahoo in the region.     

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.     

Reproductive ecology of Japanese anchovy off the Pacific coast of eastern Honshu, Japan

Spawning frequency of offshore migrant populations of Japanese anchovy Engraulis japonicus ranged from 0·34 to 0·90, and was higher than that of inshore Japanese anchovy. In addition, spawning frequency of offshore populations varied in response to the sex ratio. They spawned at water temperatures of 5.0, 8.6 and 12.6° C, indicating that the minimum critical water temperature of offshore Japanese anchovy was considerably lower than that of inshore populations. A significant positive relationship was observed between water temperature and relative batch fecundity of offshore Japanese anchovy. Furthermore, water temperature was negatively related to egg size, suggesting a trade-o. between relative batch fecundity and egg size for offshore fish. At the same water temperature, relative batch fecundity of offshore populations was higher than that of inshore ones. The comparison of spawning frequency and relative batch fecundity suggests that offshore Japanese anchovy represent a reproductive ecology by which they spawn more eggs than inshore Japanese anchovy within a certain time period.     

Reproductive biology of hardhead catfish Ariopsis felis: evidence for overwintering oocytes

Hardhead catfish Ariopsis felis are a common marine catfish in the coastal waters of the Gulf of Mexico (GOM). The low economic value of this species has depressed interest and research, and although the species is known for its extremely low fecundity and large oocytes, little else is known about this catfish species. A total of 1230 samples across all months of the year from 2016 to 2018 resulted in 681 females, and analysis of gonado-somatic index (I_G) revealed 1% to be a clear cut-off indicating maturity. Females are considered capable of spawning from April to June when I_G averaged 4–8%. Both atresia and post-ovulatory follicles were present in July, suggesting that spawning ends in July in the northern GOM. The 1% I_G cut-off was used to designate maturity, and from that an L₅₀ of 253 mm was estimated. Batch fecundity from 41 females estimated a mean batch size of 36 oocytes. Perhaps the most interesting finding was the presence of secondary growth stage oocytes (e.g., cortical alveoli) from July through November, well outside the spawning capable period. Furthermore, 78% of females had some early vitellogenic oocytes present during the non-spawning season, and the distribution of these relatively large (2–5 mm) oocytes did not change over time. The results here are not only important as reproductive biology information for a common and abundant species, but also present interesting and unusual patterns of non-spawning season oocyte development that is not commonly seen in Western Hemisphere subtropical fish species.     

Reproductive Schedules in Southern Bluefin Tuna: Are Current Assumptions Appropriate?

Southern bluefin tuna (SBT) appear to comprise a single stock that is assumed to be both mixed across its distribution and having reproductive adults that are obligate, annual spawners. The putative annual migration cycle of mature SBT consists of dispersed foraging at temperate latitudes with migration to a single spawning ground in the tropical eastern Indian Ocean. Spawning migrations have been assumed to target two peaks in spawning activity; one in September-October and a second in February-March. SBT of sizes comparable to that of individuals observed on the spawning ground were satellite tagged in the Tasman Sea region (2003-2008) and demonstrated both migrations to the spawning grounds and residency in the Tasman Sea region throughout the whole year. All individuals undertaking apparent spawning migrations timed their movements to coincide with the second recognised spawning peak or even later. These observations suggest that SBT may demonstrate substantial flexibility in the scheduling of reproductive events and may even not spawn annually as currently assumed. Further, the population on the spawning grounds may be temporally structured in association with foraging regions. These findings provide new perspectives on bluefin population and spatial dynamics and warrant further investigation and consideration of reproductive schedules in this 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.     

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.     

Estimating Intermittent Individual Spawning Behavior via Disaggregating Group Data

In order to understand fish biology and reproduction, it is important to know the fecundity patterns of individual fish, as frequently established by recording the output of mixed-sex groups of fish in a laboratory setting. However, for understanding individual reproductive health and modeling purposes it is important to estimate individual fecundity from group fecundity. We created a multistage method that disaggregates group-level data into estimates for individual-level clutch size and spawning interval distributions. The first stage of the method develops estimates of the daily spawning probability of fish. Daily spawning probabilities are then used to calculate the log likelihood of candidate distributions of clutch size. Selecting the best candidate distribution for clutch size allows for a Monte Carlo resampling of annotations of the original data which state how many fish spawned on which day. We verify this disaggregation technique by combining data from fathead minnow pairs, and checking that the disaggregation method reproduced the original clutch sizes and spawning intervals. This method will allow scientists to estimate individual clutch size and spawning interval distributions from group spawning data without specialized or elaborate experimental designs.     

Reproductive biology of the endangered Oxleyan pygmy perch Nannoperca oxleyana Whitley

The reproductive biology of the Oxleyan pygmy perch Nannoperca oxleyana is described from simultaneous studies of wild populations in north-eastern New South Wales and mature fish held in aquaria. In the wild, 50% of males and females matured at total lengths of 24·0–25·9 and 28·0–29·9 mm, respectively. The species displays sexual dichromatism during the spawning season, with males developing more intense red and brown fin and body colouration, and black pelvic fins. Captive male broodfish displayed territoriality during the breeding season, closely guarding sites within artificial, plant-like substrata in which pairs of fish spawned adhesive eggs. Protracted serial spawning of wild and captive fish occurred from September to April and May at mean water temperatures ≥16·6° C and day length ≥10·7 h. Captive broodfish spawned on an average of 57% of days during the 256 day spawning period. Gonado-somatic indices averaged 0·7% for all ripe males and 4·1–4·2% for all ripe females collected. Mean total and batch fecundities of captive females were 1323 eggs per fish and 7·8 eggs per fish per day, respectively, and relative fecundity was 587 eggs g⁻¹ of body mass. Batch fecundity of wild females was estimated at 7·8 eggs per fish. The adaptive significance of this reproductive strategy in a harsh, variable environment is discussed.     

Reproductive biology of Oxygymnocypris stewartii in the Yarlung Zangbo River in Tibet, China

We determined the gonadal histology, reproductive characteristics and implications for management of Oxygymnocypris stewartii from individuals collected in the Yarlung Zangbo River. Based on the analyses of gonad development and the size distribution of oocytes, O. stewartii spawns just once each year. Peak spawning occurred between late March and early April, when water temperature exceeded 5°C. The standard lengths and the ages at 50 % maturity for males and females were 273 mm and 357 mm, and 5.1 year and 7.3 years, respectively. The mean fecundity of 34211 and mean relative fecundity of 25.4 eggs per gram of fish body weight suggest that O. stewartii might be especially vulnerable to exploitation.     

The influence of feeding regime on sexual maturation, fecundity and atresia in first-time spawning turbot

Exposure of adolescent turbot Scophthalmus maximus to low rations during vitellogenesis, covering the 4 months immediately prior to spawning, led to a drop of 70% in mean ovary weight, and was associated with poor growth of the vitellogenic oocytes or, in a third of cases, the absence of vitellogenic oocytes. Exposure to low rations during the recruitment of vitellogenic oocytes, 4–8 months prior to spawning, produced a more variable response. A few of these females produced well-developed vitellogenic oocytes, but in most, vitellogenic oocytes were absent or only poorly developed and in some cases there was a high incidence of atresia. Exposure to intermediate rations throughout oocyte recritment and vitellogenesis also depressed oocyte development. In maturing females the number of non-atretic vitellogenic oocytes (relative potential fecundity) averaged 998 per g of somatic body weight of fish at the start of the spawning season in mid-June. Maturation of the males was not significantly influenced by the dietary regime. The small size of the testes and the low levels of milt production, even in fish fed high rations immediately prior to spawning, showed that reproductive investment in males was much lower than in females.