Sex-specific life history patterns in bluegill (Lepomis macrochirus): interacting mechanisms influence individual body size

The ultimate body size that an individual fish achieves can be a function both of direct effects of growth or indirect effects associated with the timing of sexual maturation (and associated energetic tradeoffs). These alternatives are often invoked to explain variation in body size within and among fish populations, but have rarely been considered simultaneously. We assessed how resource availability and timing of maturation interact to influence individual body size of bluegill (Lepomis macrochirus). Resource availability (high and low food) and the social structure of the population (presence or absence of large, mature males) were varied in experimental ponds. Food ration affected growth (larger fish in the high food treatments) and the social structure of the population affected timing of maturation (early maturation of males in the absence of large males). Treatment effects, however, were sex-specific; males responded to the social structure of the population and females were more responsive to resource availability. We also found individuals that became sexually mature were smaller than those that remained immature, although results were sex-specific and resource dependent. For males, individuals that matured were smaller when resources were limited; mature and immature females showed no difference in body size regardless of food ration. We show that both resource availability and the processes that control timing of maturation interact in sex-specific ways to influence body size of bluegill. These results suggest that a more robust explanation for variable body size requires consideration of sex-specific interactions between ecological (food and growth) and evolutionary (timing of maturation) mechanisms.     

Reproductive biology and implications for management of the orange-spotted grouper Epinephelus coioides in the southern Arabian Gulf

The reproductive biology of Epinephelus coioides was determined from the examination of 1455 individuals collected between July 2005 and June 2007 in the southern Arabian Gulf. Histological preparations of gonads indicated that males were either derived from a juvenile phase or the transition of postspawning females, confirming a diandric protogynous sexual pattern. The spawning season was well defined, occurring once a year during April and early May. Peaks in spawning occurred after the full and new moons and was completed within a single lunar cycle. The presence of mature males over the entire size and age range and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. While specimens undergoing sexual transition were only observed in size and age ranges of 335–685 mm total length ( L _T) and 5–6 years, patterns in the proportion of males in size and age classes suggested that sex change occurred at a relatively constant rate after female maturation up to the maximum size (1002 mm L _T) and age (11 years). Relationships between reproductive output and capacity with size and age indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of E. coioides . The maximum age, small size and young age at sexual maturation ( L _min= 320 mm L _T, 2 years, for females and 242 mm L _T, 1 year, for males) conflict with the general pattern for large epinepheline groupers and may be a direct result of the intensive demersal fishery in the southern Arabian Gulf.     

Age and size-dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed,Lepomis gibbosus

Synopsis In this study, we used pumpkinseed,Lepomis gibbosus, populations in two east-central Ontario lakes to test for age and size-dependent effects on the probability and timing of reproduction of mature females. Pumpkinseed body size characteristics differed in the two lakes; Little Round Lake harbours a stunted population and Beloporine Lake does not. Age-class distribution of mature females was determined by a maturity assessment on fish collected just prior to spawning and at the mid-spawning period, combined with an early-season capture-mark-recapture survey. Spawning females were collected throughout the breeding season to assess age and size-related temporal trends, and to compare their age and size distribution with that of mature females at large. The proportion of age 2 females in Little Round Lake that actually spawned was considerably lower than the proportion of mature age 2 females at large. Furthermore, age 2 females that spawned in this lake did not do so until late in the breeding season. In contrast, the proportion of young/small females spawning in Beloporine Lake was comparable to the proportion of young/small mature females at large, and both small and large females spawned throughout the breeding season. Small mature females in Little Round Lake may have had no other option but to spawn late in the season because of their poor body condition. In Beloporine Lake, condition factor early in the breeding season in age 2–4 females was higher than that of Little Round Lake females, suggesting that limited energy reserves in the spring may have prevented young Little Round Lake females from early spawning. Our results show that the likelihood and timing of reproduction are both age and size-dependent in some populations. Small individuals that delay seasonal maturation and spawn late in the summer probably contribute little to the population due to the restricted growth and reduced overwinter survival of their progeny.To whom correspondence should be addressed     

Maturation of female common snook Centropomus undecimalis: implications for managing protandrous fishes

The assumption for hermaphroditic fish species that mature individuals of the terminal sex arise directly from mature individuals of the primary sex has led to the use of sex ratios as a proxy for age at maturity (A₅₀). The timing of transition and deficient energy reserves, however, can result in a delay between transition and spawning. To test the assumption of female maturity and investigate the relationship between maturation and energy reserves, common snook, Centropomus undecimalis, a protandrous hermaphrodite, were collected from rivers, estuaries, inlets and offshore habitats on the east coast of Florida during 2010–2015. Immature females were observed every month, with lowest proportions during the peak spawning months of July and August. When calculated based on sex ratio, A₅₀ (8.1 years) overestimated the age at which 50% of the female population was, in fact, mature (4.1–4.9 years). Best-fit models indicate that mesenteric fat index (I_F) and hepato-somatic index (I_H) were significantly affected by gonad phase, month and size and weakly by habitat. In post hoc analysis, immature female I_F did not differ significantly from developing and regenerating females, but immature female I_H was significantly lower than that for all mature phases except animals in the regressing phase. Although immature females may have sufficient energy in terms of fat, it appears that energy is not allocated to reproductive processes, as evidenced by lower I_H. Nonetheless, approximately 95% of females were spawning-capable during peak spawning months, suggesting that the energy threshold at which immature females reach maturity is met by most females each spawning cycle.     

Reproductive biology and implications for management of the spangled emperor Lethrinus nebulosus in the southern Arabian Gulf

The reproductive biology of the spangled emperor Lethrinus nebulosus in the southern Arabian Gulf was determined from the examination of 781 individuals collected between September 2008 and August 2009. There was no histological evidence of adult sex change, and sex ratios did not differ significantly from unity across all size and age classes. Testes had an ovarian structure and a remnant lumen which was not used for sperm transport; furthermore, residual oocytes were observed in the testes of some immature, resting and developing males. A dramatic change in the sex ratio of the smallest and youngest size and age classes suggested that juvenile female to male sex change occurred between 21·0 cm fork length (L(F)) and 24·0 cm L(F) at 1 year of age. The combination of histological evidence with the sexual composition of the size and age structures suggests a non-functional protogynous hermaphroditic sexual pattern, which is analogous to functional gonochorism. The spawning season was well defined, occurring once a year during March, April and early May. Peaks in spawning occurred after full moons, there was a cessation in spawning activity after new moons and spawning was completed within three lunar cycles. The distribution of males over the entire size and age ranges and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. Size-specific and age-specific reproductive potential indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of L. nebulosus. The maximum recorded age (11 years), small mean size and young age at sexual maturation (L(m50) = 26·7 cm L(F), 2·1 years, for females and 19·4 cm L(F), 0·5 years, for males) may be a direct result of intensive demersal fishing in the southern Arabian Gulf.     

Female-biased operational sex ratio of sexual host fish in a gynogenetic complex of Carassius auratus

To study the coexistence of sexual and gynogenetic forms, we examined the population structure of a gynogenetic complex of the Japanese crucian carp, Carassius auratus Temminck et Schlegel, during the April–June reproductive season by collecting 1225 mature fish that migrated from Lake Suwa to a tributary river for spawning. There were more sexual fish (about 80%) than gynogenetic fish in this complex, and the operational sex ratio in the sexual form was female biased (males were about 20%). Mean standard length and body weight of sexual females were larger than those of sexual males. Sex ratio was male biased in smaller fish (standard length, <8.5 cm) but female biased in larger fish (standard length, ≥8.5 cm). We determined age by scale ring marks; the average age of sexual females was higher than that of males, but there was no significant difference in the average age between sexual and gynogenetic females. Sex ratio in the sexual form was more female biased for old than for young fish, and the mean size of sexual females was larger than that of males of the same age. The clear female-biased sex ratio and age difference between sexual females and males can be explained either by (1) higher mortality of males or by (2) female-biased sex allocation. The latter process reduces the disadvantage of sex and contributes to the coexistence of sexual and gynogenetic forms. Received: November 24, 2000 / Accepted: March 6, 2001     

Sexual maturity cycle and spawning of Greenland halibut Reinhardtius hippoglossoides in the Davis Strait

Female sexual maturation cycle and the main spawning time of Greenland halibut Reinhardtius hippoglossoides in the Davis Strait were studied through regularly collected samples during 1 year starting in spring 2003. Samples were collected from the southern slope of the Davis Strait Ridge between Canada and Greenland in the depth range 1000–1500 m. Female sexual maturation was described using different approaches: gonado‐somatic index, visual macroscopic maturity stage index, histological microscopic maturity index and oocyte diameter measurements. A significant increase in the gonado‐somatic index was seen from September onwards until February with a maximum estimated value of 18%. The proportion of mature fish increased from December until March. At the same time, the proportion of females with a low gonado‐somatic index also increased from February, indicating that spawning had occurred and females were recovering. Oocyte diameter distribution revealed a leading cohort development during autumn through to December to February. A coupling between sexual maturity and fish condition was seen for females in maturing condition indicating a steady build up of stored energy in the liver from June to November.     

Effects of varying temperature and food availability on growth and reproduction in first‐time spawning female Atlantic cod

The somatic growth, sexual maturation and fecundity of individually marked first‐time spawning female Atlantic cod Gadus morhua were examined under different varying temperature and feeding regimes over the months preceding spawning. A negative correlation between somatic and oocyte growth was found, reflecting the changing energy allocation pattern. Nevertheless, the somatic growth of mature individuals was at least as high as those of immature fish over the period of vitellogenesis. Potential fecundity was positively correlated with body size, but neither temperature or feeding regime significantly affected this relationship. Consequently, fish with unlimited feeding opportunity invested more energy into somatic growth during vitellogenesis compared to those held under a restricted ration. This indicated that once Atlantic cod had made the decision to invest in first reproduction, they allocated a certain amount of energy relative to their size into egg production and any surplus was invested into somatic growth. Low temperature led to an arrest in the onset of vitellogenesis and significantly affected the number of females that matured.     

Reproductive biology ofAwaous guamensis, an amphidromous Hawaiian goby

Synopsis Spawning season, size at first reproduction, oocyte maturation, and fecundity ofAwaous guamensis, an amphidromous Hawaiian goby, were studied from June 1989 through May 1991 in the Wainiha River, Kau'ai, Hawai'i. Female fish larger than 73 mm standard length (SL) had mature gonads from August through December in 1989 and 1990. Gonadosomatic index (GSI) values for mature females ranged from 0.2 to 14.5 during the spawning season. Male fish larger than 64 mm SL had elevated GSI values from June 1989 through December 1989 and from August 1990 through December 1990. Mature sperm were found in two male fish collected in January and February. GSI values for mature males ranged from less than 0.01 to 4.0 in the spawning season. Size-frequency distributions of measurements of vitellogenic oocyte diameters and microscopic observations of oocytes indicated this species has group-synchronous oocyte development. Ovarian maturation stages examined over a 29-month period suggest that members of the stock spawned at different times within the spawning season, although mass spawning events have been documented for this species. Estimates of clutch sizes from nests measured in situ were comparable to estimates of potential fecundity from in vitro examination of ovaries, and indicated that female fish deposited an entire clutch during a spawning event. No evidence for multiple spawning by an individual fish in a single season was found. However, microscopic observations of brown bodies in some ovaries suggested that individual fish probably spawn more than once in a lifetime.     

Growth of dark chub, Zacco temmincki (Cyprinidae), with a discussion of sexual size differences

I examined the annual and seasonal growth of dark chub, Zacco temmincki, in a Japanese river. Investigation of opercular rings showed that the fish reached a maximum age of 8 years. There was no significant sexual size difference at younger ages (3–5 years), but males were larger than females at older ages (6–7 years). Annual increments of length and weight for males that were recaptured were also larger than those for females. The fish spawn from June to August. Females grew for a short period from April to May, but males on average grew for a longer period from April to August. There was no sexual difference in growth rate except during the spawning period. Annual growth rate was negatively correlated with fish length in each sex. The sexual size differences at older ages of the fish might be due to the polygynous mating system in which most mature males could not obtain females and invested for somatic growth in the spawning period, and a short growing season that was overlapped considerably with the spawning period.     

New data on the reproductive biology of shortspine thornyhead <Emphasis Type="Italic">Sebastolobus alascanus</Emphasis> (Sebastidae) in the northwestern Pacific

New data on fecundity, sex ratio in different size groups, and the rate of sexual maturation of shortspine thornyhead Sebastolobus alascanus in the northwestern part of the Pacific Ocean are provided. Individual absolute fecundity of the studied females varies in the range of 175 000–821000 (on average, 392500) eggs. In Pacific waters off the Kuril Islands and Kamchatka, among individuals with a length to 41 cm, males dominate; in size groups of 42–59 cm, sex ratio is almost equal; among larger fish, females prevail. In the western part of the Bering Sea among analyzed fish with a length to 44 cm, sex ratio is almost equal; among larger individuals, females dominate. Half of males and females of shortspine thornyhead off the eastern coast of the northern Kuril Islands and southeastern Kamchatka mature at a length of approximately 41 cm that can be recommended as a commercial size for the given species in this area.     

Reproductive strategies of Red-bellied Piranha (<Emphasis Type="Italic">Pygocentrus nattereri</Emphasis> Kner, 1858) in the white waters of the Mamirauá flooded forest,central Brazilian Amazon

Despite being an important member of neotropical fish assemblages, as well as a species with an unenviable reputation, little is known about the reproductive ecology of Red-bellied Piranha, Pygocentrus nattereri, in the wild. We tracked the reproductive activity of piranhas in the flooded forests of Mamirauá Reserve, Central Brazilian Amazon, for 2 years, in an investigation that included over 3,000 individual fish. Contrary to expectation piranhas had not one, but rather two annual reproductive seasons, tuned to water level fluctuation and the flooding pulse. Females were found to have up to 30,000 oocytes, little more than one third of which were mature and available for spawning in a single batch. Sexual maturation (of both males and females) occurred at around 160 mm. At this point sexually active individuals (of both sexes) became dark and lost most of their red coloration. Ontogenetic changes in habitat selection were strongly associated with the spawning behavior of mature adults. We also found that flooded marginal vegetation and marginal grasses inside lakes were the preferred spawning areas. In contrast, non-reproductive individuals were found in open water and under floating meadows.     

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.     

Terminal spawning in the deepwater squid Moroteuthis ingens (Cephalopoda: Onychoteuthidae)

Maturation in the onychoteuthid squid Moroteuthis ingens was found to be irreversible, with death following shortly after sexual maturation and spawning. Both males and females were found with spent gonads. The ovary reaches very large sizes in mature females and probably prevents feeding by constricting the caecum. There was also a marked difference in the tissue integrity between immature females and females which had reached sexual maturity. Mature and spent females showed advanced tissue breakdown with individuals having a thin mantle wall with an inelastic, gelatinous appearance. Histological examination of the mantle wall revealed that the tissue breakdown was due to a drastic histolysis of muscle tissue and, to a lesser extent, collagen fibres. Mature males also showed some tissue breakdown and loss of muscle fibres but this was not as dramatic as in the females. These features are considered in relation to processes contributing to terminal maturation in M. ingens.     

Reproduction in the piranha Serrasalmus spilopleura, a neotropical fish with an unusual pattern of sexual maturity

Synopsis Reproduction in female Serrasalmus spilopleura (Characiformes, Characidae) from the Itumbiara reservoir (18°28 S, 48°36 W), Paranaíba River, Brazil, was examined with emphasis on the relationship between condition factors and coelomic fat, the annual reproductive cycle and the unusual dynamics of its sexual maturity. The condition factor and coelomic fat index follow a similar pattern along the stages of reproductive cycle. Variations in the condition factor along the reproductive cycle were assigned to variations in the fat contents of the carcass. Serrasalmus spilopleura exhibits characteristics of partial spawning fish and reproduces throughout the year. Sexual maturity occurs at 17.8 cm standard length. Resting and totally spent females were not captured. Females cycle from partially spent to intermediate/advanced maturation without going into the totally spent or resting stages. The initial maturation stage is restricted to fish of intermediate size. Once the initial maturation stage has started the fish remains there for a considerable period of its lifetime before entering in the intermediate maturation stage, an uncommon sexual maturity pattern. The long duration of the initial maturation stage was responsible for the females not returning to the resting stage after spawning.     

Reproductive biology of brown trout, Salmo trutta macrostigma Dumeril 1858, in a tributary of the Ceyhan River which flows into the eastern Mediterranean Sea

The study describes some key elements of the reproductive biology, including spawning season, age at sexual maturity, fecundity and egg diameter of the native brown trout, Salmo trutta macrostigma, in a tributary of the Ceyhan River. A total of 197 brown trout (118 females and 79 males) were captured in 2000–2001 by electric fishing. In observations on monthly changes, the gonadosomatic index (GSI) and the monthly frequency distribution of egg diameter confirmed that spawning lasted from November to January. Some 27.7% of the females and 62.5% of the males attained sexual maturity in their second year. The smallest fork length (FL) of brown trout attaining sexual maturity was 17.4 cm for males and 17.8 cm for females. Mean fecundity in age groups II, III, IV and V were 360, 452, 693 and 1283 eggs per female, respectively. One 9‐year‐old female had a unique 3232 egg count. The mean fecundity of the sampled population was 554 eggs per fish, positively correlated with the FL (mm) (R = 0.8227 ) and body weight (R = 0.8130). The diameter of mature eggs in the spawning season ranged from 3.250 to 5.930 mm, with a 4.146 mm average. Mean egg diameter in age groups II, III, IV and V in the spawning season were 0.813, 3.799, 4.663 and 5.243 mm, respectively. Fecundity, egg weight and diameter were statistically different in all age groups.     

Male Isaza (Gymnogobius isaza, Gobiidae) prefer large mates: a counterstrategy against brood parasitism by conspecific females

Like many other gobies, male Isaza (Gymnogobius isaza) which are endemic to Lake Biwa, Japan, exclusively care for broods in nests. This goby may have an optimal range of brood size (i.e., an average clutch size of about 2000–3000 eggs) within which they may produce larger numbers of hatching young because much larger broods may be destroyed by fungal infection before hatching. This optimal brood size hypothesis (Takahashi et al. in J Ethol 22:153–159, 2004) predicts that (1) after spawning, both males and females will refuse additional spawning by other gravid females (second females) to keep brood sizes within optimal ranges, (2) larger fish will repel second females more successfully than will smaller fish, and thus, (3) both sexes prefer larger mates. To examine these predictions, we first observed Isaza’s aggressive behaviors in aquaria and investigated whether fish attacked and repelled second females that were introduced after spawning, and, if so, what were the sizes of fish that did so. Large fish, regardless of sex, aggressively prevented second females from entering the nest, but second females larger than the pairs displaced the pair females forcibly and spawned eggs into their clutches. Mate choice experiments showed that males preferred large females. Although females’ choice of large mates was not confirmed, many results may largely coincide with the predictions of the optimal brood size hypothesis. Thus, Isaza males’ choice of large mates will be advantageous for defending against brood parasitism by conspecific females and for achieving optimal clutch size.     

Ontogeny of sexual development in the roach (Rutilus rutilus) and its interrelationships with growth and age

The roach (Rutilus rutilus) has become a sentinel species for the study of sexual disruption in wild fish populations as a consequence of exposure to endocrine disrupting chemicals (EDCs). Little is known, however, about the normal ontogeny of sexual development in this species. Here, we analyzed the ontogeny of sexual development in captive‐bred roach and assessed how growth rate and fish size affected the timing of both sexual differentiation and sexual development over a 2‐year period. Ovarian differentiation was first recorded at 68 days post‐fertilization (dpf) and this preceded testicular differentiation (first recorded at 98 dpf). In contrast, sexual maturation occurred at an earlier age in males (300 dpf) compared with females (728 dpf). No differences in body size (length or weight) were recorded between male and female roach until the fish were 415 dpf. Studies on three populations of roach which grew at different rates showed that the timing of sexual differentiation was highly variable and more related to fish size than to fish age. Time to sexual maturation was also variable among populations but, subsequent to their first year of life, gonadal status was less well associated with fish size. Interestingly, the sex ratio of the population was biased towards females in populations that grew more rapidly during early life. The findings presented here provide a valuable foundation of work to support both field‐ and laboratory‐based assessments on the effects of EDCs, and other stressors, on sexual differentiation and development in the roach. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

‘Good-genes’ and ‘compatible-genes’ effects in an Alpine whitefish and the information content of breeding tubercles over the course of the spawning season

Some models of sexual selection predict that individuals vary in their genetic quality and reveal some of this variation in their secondary sexual characteristics. Alpine whitefish (Coregonus sp.) develop breeding tubercles shortly before their spawning season. These tubercles are epidermal structures that are distributed regularly along the body sides of both males and females. There is still much unexplained variation in the size of breeding tubercles within both sexes and with much overlap between the sexes. It has been suggested that breeding tubercles function to maintain body contact between the mating partners during spawning, act as weapons for defence of spawning territories, or are sexual signals that reveal aspects of genetic quality. We took two samples of whitefish from their spawning place, one at the beginning and one around the peak of spawning season. We found that females have on average smaller breeding tubercles than males, and that tubercle size partly reveals the stage of gonad maturation. Two independent full-factorial breeding experiments revealed that embryo mortality was significantly influenced by male and female effects. This finding demonstrates that the males differed in their genetic quality (because offspring get nothing but genes from their fathers). Tubercle size was negatively linked to some aspects of embryo mortality in the first breeding experiment but not significantly so in the second. This lack of consistency adds to inconsistent results that were reported before and suggests that (i) some aspects of genetic quality are not revealed in breeding tubercles while others are, or (ii) individuals vary in their signaling strategies and the information content of breeding tubercles is not always reliable. Moreover, the fact that female whitefish have breeding tubercles of significant size while males seem to have few reasons to be choosy suggests that the tubercles might also serve some functions that are not linked to sexual signaling.     

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.