Annual variations in the reproductive traits of Pomatoschistus microps in a Mediterranean lagoon undergoing environmental changes: evidence of phenotypic plasticity

The spawning period of the common goby Pomatoschistus microps from 1993 to 1997 in the Vaccarès lagoon did not vary, except in 1997 when it was longer due to the reproduction of the young-of-the-year. Egg size and number, and reproductive allocation varied greatly with one year to another. Female common gobies increased both their fecundity per spawning act and their egg size from 1993 to 1995. The annual variation in the reproductive effort suggests a high phenotypic plasticity of reproductive traits in P. microps , in the face of environmental perturbations. In winter 1993–1994, a centennial flood of the Rhône River caused major hydrological changes in the lagoon in less than 1 week, affecting many invertebrates and fish for several years. The reproductive investment of the common goby increased, possibly as a consequence of those environmental changes.     

Size Dependent Male Reproductive Tactic in the Two-Spotted Goby (Gobiusculus flavescens)

Male investment in testes and sperm duct gland in the polygamous nest breeding two-spotted goby Gobiusculus flavescens (Fabricius) was investigated in relation to time in reproductive season and individual physical parameters. This small teleost fish is most likely the most abundant species found along the rocky shores of the North East Atlantic. The two-spotted goby has a single reproductive season, during which nest-caring males can raise several clutches of offspring. According to the literature the males are on average larger than the females. Here we report for the first time a population showing a reversal of this trend, with males on average being smaller than females, a difference likely caused by a large proportion of small males. Early in the breeding season these small males have typical sneaker characters, with relatively large testes and small seminal duct glands compared to the larger dominant territorial males. The presence of these two alternative male reproductive tactics is confirmed by histological studies, which shows the presence of sperm in the sperm duct glands (SDG) of smaller males, but not in the SDG of intermediate and larger males. To our knowledge, males with typical sneaker characters have not been reported in earlier studied populations of two-spotted goby. Interestingly we found that testes investment declined significantly over the course of the breeding season, and that this reduction was significantly more pronounced in small compared to the large males. Further, a significant increase in seminal duct gland (SDG) mass was observed for the smaller males over the breeding season. We propose that this indicates a possible shift in mating tactic by smaller males from a parasitic to a nest-holding tactic over the course of the breeding season. Thus, the observed size dependent plasticity in investment in SDG over time suggests that the reproductive tactic of G. flavescens is conditional, and possibly influenced by mate availability and male—male competition.     

Reproductive effort and efficiency in the female common goby, Pomatoschistus microps (Krøyer) (Teleostei: Gobioidei)

Reproductive effort and efficiency are terms which together help to describe the reproductive investment of spawning fish. In this paper they are applied to a small and abundant inshore fish, the common goby, Pomatoschistus microps (Kroyer), based on data collected from the wild, and by modelling likely breeding patterns. Wet calorimetry was used to determine the energy content offish tissue, and this allowed reproductive effort to be expressed as both an energy and a weight ratio of gonad to somatic body. The gonadosomatic index (GSI) calculated as an energy ratio was greater than that calculated as a dry "weight ratio". For ripe females the GSI (dry weight) at the onset of breeding was 67.5%. Theoretical and observed values of the maximum number of egg batches laid by a single female during a 16-week breeding season are 9-10, and this corresponds to a total caloric investment of almost 12 kJ. Estimates of the reproductive efficiency, using the dietary energy content during the breeding season, range from 28.2% to a maximum of 44.0%. These values are among the highest calculated for a fish species, and are in keeping with the long and intensive breeding strategy observed in the wild population.     

Annual changes in brain concentration of arginine vasotocin and isotocin correspond with phases of reproductive cycle in round goby,Neogobius melanostomus

Reproductive cycle of seasonally breeding fish is synchronized with changes of photoperiod and temperature in environment. We hypothesize that arginine vasotocin (AVT) and isotocin (IT) are involved in timing and synchronization of seasonal reproductive activity in the round goby (Neogobius melanostomus). To verify this hypothesis, we examined the annual profiles of brain AVT and IT in round goby males and females in relation to their reproductive cycle. Wild round gobies were exposed to annual environmental changes in their natural habitats from where they were sampled monthly over a year. AVT and IT were measured using HPLC with fluorescence detection preceded by solid-phase extraction. This study shows seasonal variations in brain AVT and IT levels. Profiles of changes were similar in males and females: the peak of AVT was observed before spawning in March-April, whereas that of IT during spawning in May–June. Furthermore, the lowest AVT level was noted out of breeding season from November to January, while the level of IT decreased immediately at the end of the spawning. The results show that high AVT levels correlate with pre-spawning period whereas the highest IT levels correspond to spawning. A significant decline in AVT and IT in non-spawning season coincided with the quiescent phase of gametogenesis in both sexes.     

Demographic structure and life history traits of the common goby Pomatoschistus microps (Teleostei,Gobiidae) in a Mediterranean coastal lagoon (Rhône River delta,France)

Demographic structure and life history traits of the common goby Pomatoschistus microps were investigated in a brackish water lagoon of the Rhône River delta (Mediterranean Sea, southern France). The size frequency distribution and gonadosomatic index indicated that three different age groups occurred and reproduced successively in the lagoon, resulting in a long spawning period from March to September and a high investment in reproduction. This high investment in reproduction, which contrasts with that found in other mostly northern European populations, probably relates to the unpredictability of the gobyˈs environmental conditions.     

Reproductive biology of an endemic,amphidromous goby Lentipes concolor in Hawaiian streams

Synopsis Egg size, fecundity, nest site selection, and breeding structure of the amphidromous Hawaiian goby Lentipes concolor were studied to determine if there was any relationship between these aspects of the breeding biology of this fish and the very long larval period of Hawaiian stream gobies. To do this, these parameters were compared with those known for other gobioid fishes. While eggs were small and females were found to lay up to four nests in a season, reproductive biology and mating behavior of L. concolor were typical of many gobies in spite of its amphidromous life history pattern. Lentipes concolor nests were found from October to June with a spawning peak in February. Nests tended to be positioned closer to the stream bank, and under larger rocks than if they were situated randomly. Depth and mean water column velocity over nest sites did not differ from that at randomly selected points in the stream.     

Not All Offspring Are Created Equal: Variation in Larval Characteristics in a Serially Spawning Damselfish

The way organisms allocate their resources to growth and reproduction are key attributes differentiating life histories. Many organisms spawn multiple times in a breeding season, but few studies have investigated the impact of serial spawning on reproductive allocation. This study investigated whether resource allocation was influenced by parental characteristics and prior spawning history in a serial spawning tropical damselfish (Pomacentrus amboinensis). The offspring attributes of isolated parents of known characteristics were monitored over a 6-week breeding period in the field. Smaller females produced larvae of longer length and larger energy reserves at hatching. This finding is contrary to several other studies that found larger females produce offspring of greater quality. We found that resource allocation in the form of reproductive output was not influenced by the number of spawning events within the breeding season, with larger females producing the greatest number of offspring. Larval characteristics changed as spawning progressed. There was a general decline in length of larvae produced, with an increase in the size of the larval yolk-sac, for all females regardless of size as spawning progressed. This trend was accentuated by the smallest females. This change in larval characteristics may reflect a parental ability to forecast unfavourable conditions as the season progresses or a mechanism to ensure that some will survive no matter what conditions they encounter. This study highlights the importance of accounting for temporal changes in reproductive allocation in studies of reproductive trade-offs and investigations into the importance of parental effects.     

The seasonal partitioning of energy in an estuarine fish, the common goby, Pomatoschistus microps Krøyer

The common goby, Pomatoschistus microps , is an abundant brackish water species which has an appreciable role, both as predator and prey, in the ecosystems of British estuaries. The partitioning of energy within each sex is described and discussed, and energy values are provided for the gonad, liver and carcass throughout almost the entire life of one year class. Partitioning of resources within the body at different times of the year shows considerable variation, not only with time but also between the sexes. In particular, the energy content of the male gonad during the spawning season is never more than 3.7% of that of the female. Important aspects of the biology of the species which relate to energy partitioning are maturation at an early age, high reproductive investment, high stochastic mortality, and death usually after a single breeding season.     

Reproductive biology of racer goby Neogobius gymnotrachelus in the Włocławski Reservoir (Vistula River, Poland)

Initial studies on the reproductive parameters and spawning behaviour of racer goby Neogobius gymnotrachelus, one of three Ponto‐Caspian gobiid species recently invading Polish waters, were carried out in W?oc?awski Reservoir (lower River Vistula, Poland), where spawning lasted from April to August. Males were on average about 23.3% longer and 53.8% heavier than females. The smallest female with mature gonads was 58 mm TL (estimated age 2+ years), and males were sexually mature at 59 mm TL (age of 1+ years). Gonado‐somatic index at the beginning of spawning was seven times larger in females (mean = 8.4) than that in males (mean = 1.2). Mean absolute fecundity was 952 eggs per female (361–2236). Egg diameter ranged from 0.140 to 1.558 mm, being of bi‐modal distribution, which suggests two or three spawning bouts. Length–frequency analysis of juveniles revealed a multimodal distribution. Specialized reproductive behaviour (nest construction and parental care of eggs) and an extended spawning period increase the probability that racer goby can successfully establish self‐sustaining populations in novel environments.     

Semelparous or iteroparous: resource allocation tactics in the ayu, an osmeroid fish

Ayu Plecoglossus altivelis is an annual osmeroid fish with a single breeding season. Its reproductive mode tends to vary such that large females spawn once, while small females spawn twice with an interval of some 2 weeks. The hypothesis was examined that certain female reproductive modes are governed by adequate resource allocation according to maternal body size, to maximize lifetime reproductive success. A series of rearing experiments revealed that immediately before spawning, larger females had relatively higher diversion from the soma and higher investment into the gonad, in contrast to smaller females which maintained higher protein synthetic activity for further reproduction. It was concluded that the extant plasticity in the reproductive mode that occurred in P. altivelis was an outcome of different tactics on the semelparity-iteroparity continuum rather than maladaptive deviation from the optimum.     

Life-history traits of Amur sleeper, <Emphasis Type="Italic">Perccottus glenii</Emphasis>, in the invaded Vistula River: early investment in reproduction but reduced growth rate

In recent decades, Amur sleeper is one of the most invasive fish species in Eastern and Central Europe. Generally, it is assumed that success of an invasive species can largely depend on the plasticity of its life-history traits, e.g., indicated by higher investment in reproduction in the initial stage of its invasion. On the other hand, such energy allocation to production of gonads should negatively impact somatic growth rate. The aim of this article was to explore this phenomenon in a non-native population of Amur sleeper inhabiting artificial reservoir on a large lowland river in Central Europe. Through analysis of the population age structure, sex-dependent growth rate and life-history traits we assessed investment in reproduction and its possible relationship with growth pattern as well as compared the results with literature data from native and introduced range. Samples collected monthly from April 2005 to March 2006 were used to estimate sex ratio, GSI, fecundity, eggs diameter frequency over the year, duration of spawning season, and mode of spawning. Age structure was calculated from scales and the same data were also used to back-calculate standard length (SL) at age. The population in the Włocławski Reservoir was represented by eight age groups, and its life span was one of the longest recorded both in its natural range and introduced areas. The weight–length relationship showed that the Amur sleeper grew isometrically and there was no difference between females and males. Although the Amur sleeper growth rate was relatively slow in the Włocławski Reservoir, its increment in the first year of life was comparable to that in other introduced areas and higher than in its natural range. The female age of maturation (1+) found in the reservoir was earlier than observed in its natural range as well as in most introduced areas (2+, 3+). Female length at maturation was similar to that reported elsewhere. The back-calculated standard length (SL) of females and males showed that in the first two age classes (1 and 2) males achieved larger SL than females. Meanwhile, at age 3, 4 and 5, females were larger than males but these differences were insignificant, except for class 5. For both sexes the average annual increment of SL decreased with age but in females the increment was always higher than in males (P < 0.05). The growth rate differences between females and males resulted possibly from their unequal investment in reproduction. Multiple spawning in the Włocławski Reservoir lasted from April to the end of August and was almost 3 months longer than in other regions. Thus, this higher investment in reproduction displayed by, e.g., earlier maturation of females and longer spawning season in the Włocławski Reservoir, may contribute to invasive success of Amur sleeper in newly colonized areas; however, the costs of this strategy result in slower growth in older age classes.     

Reproductive biology of the silver pomfret, Pampus argenteus (Euphrasen), in Kuwait waters

The reproductive activities of the silver pomfret, Pampus argenteus (Euphrasen), in Kuwait waters were investigated from March 1996 to February 1998. Observations on the seasonal distribution of maturity stages and variations in seasonal fluctuations in the gonadosomatic index (GSI) confirmed recent findings that the spawning period begins in May. The species has a prolonged spawning period in the females extending from May to August, whereas the males mature in April and ripe specimens were encountered in monthly samples until September. The recruitment pattern confirmed the one breeding season. There are two spawning peaks, the first in May and the second in August. Variations in GSI relative to fish length indicated that females and males are most fecund at about 24.5–26.4 cm and 20.5–22.4 cm length classes, respectively. The males mature earlier than females, at a minimum size of 12.5–14.4 cm, while the females mature at 20.5–22.4 cm. The oocyte diameter‐frequency distribution suggests a serial rhythm of spawning. Fecundity ranged from 28 965 to 455 661 and correlated positively with: (a) standard length (P < 0.006); (b) ovary weight (P < 0.001); and (c) body weight (P < 0.001), and negatively with egg size (P < 0.003).     

Variability of GnRH secretion in two goby species with socially controlled alternative male mating tactics

Male reproductive phenotypic plasticity related to environmental-social conditions is common among teleost fish. In several species, males adopt different mating tactics depending on their size, monopolizing mates when larger, while parasitizing dominant male spawns when smaller. Males performing alternative mating tactics are often characterized by a strong dimorphism in both primary and secondary reproductive traits. According to studies on sex-changing species and on species where only one male morph is reproductively active, male alternative phenotypes are expected to vary also in gonadotropin-releasing hormone (GnRH) neurons in forebrain preoptic area (POA). Here, we compared the intra- and inter-sexual variations in number and size of GnRH neurons, along with gonads and male accessory structure investment, in two goby species, the grass goby, Zosterisessor ophiocephalus, and the black goby, Gobius niger, characterized by male alternative mating phenotypes. In both species, older and larger males defend nests, court and perform parental care, while younger and smaller ones try to sneak territorial male spawning. We found that grass goby and black goby have different patterns of GnRH expression. Grass goby presents a clear intra-sexual dimorphism in GnRH expression, related to the occurrence of alternative mating tactics, while in the black goby, only inter-sexual differences are observed. The inter- and intra-specific variability in the GnRH neurons in these two goby species is discussed in light of the differences in migratory behavior, nest type, and mating system.     

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.     

The ecology, behavior, and conservation of the tidewater goby, Eucyclogobius newberryi

Understanding the ecology and behavior of endangered species, such as the tidewater goby, Eucyclogobius newberryi, is important for identifying problems and formulating solutions for species recovery. The tidewater goby forms isolated populations in California's coastal lagoons, creeks, and marshes. Rapid declines in the number of populations led to its listing as an endangered species in 1994. This benthic fish prefers stillwater habitats and feeds on small invertebrates. It is an annual species with an extended breeding season. Fish are larger in marsh habitats than in lagoon or creek habitats. The male digs a spawning burrow, preferably in sand, where he provides care for a single clutch. The tidewater goby is sex-role reversed: females compete more intensely than males for access to mates. The tidewater goby is a species at risk, in part due to narrow habitat preferences, isolation of populations, short lifespan, lack of marine dispersal, and vulnerability to introduced predatory fishes. Attributes that favor its recovery include euryhaline tolerances, rapid reproductive rate, its potential for opportunistic feeding, and the possibility of natural recolonization under certain circumstances. Potential conservation measures include protecting coastal marshes that adjoin creeks and lagoons, maintaining natural hydrologic regimes, preventing artificial breaching of the sandbar at the estuary's mouth, and preventing introductions of predatory fishes. Captive breeding and reintroduction of tidewater gobies are potential tools for recovery, provided that underlying problems of habitat availability and suitability and issues of genetic integrity and disease transmission are addressed. Further research into the tidewater goby's utilization of marsh habitats, dispersal mechanisms, response to artificial breaching events, and metapopulation genetics would provide additional information for management.     

Female reproductive biology of the sand smelt in brackish lagoons of southern France

The breeding season of Atherina boyeri living in the brackish lagoons of Mauguio, Pérols and Méjean (Hérault, southern France) was very protracted (February-September). The peak of the reproductive effort occurred in April, May and June. Larger fishes started spawning earlier, and stopped later, than smaller ones. Batch fecundity varied from 4 to 447 oocytes and was positively related to female length and weight. Monthly mean fecundity increased at the beginning of the breeding season and decreased between June and July. Individual mean oocyte diameter ranged from 1.34 to 1.94 mm and was not related to fish length. Monthly mean oocyte size declined throughout the breeding season as water temperature rose. There was no trade-off between number and egg size. The strategy of these sand smelts seemed well adapted to ensure spawning success without endangering their survival. The highest reproductive effort period occurred in such a way that environmental conditions were suitable to the best survival and growth of hatched larvae. When breeding ceased, fishes, particularly the youngest ones, had enough time to improve their condition before overwintering.     

Trade-off between current reproductive effort and delay to next reproduction in the leatherback sea turtle

The trade-off between current and future reproduction plays an important role in demographic analyses. This can be revealed by the relationship between the number of years without reproduction and reproductive investment within a reproductive year. However, estimating both the duration between two successive breeding season and reproductive effort is often limited by variable recapture or resighting effort. Moreover, a supplementary difficulty is raised when nonbreeder individuals are not present sampling breeding grounds, and are therefore unobservable. We used capture–recapture (CR) models to investigate intermittent breeding and reproductive effort to test a putative physiological trade-off in a long-lived species with intermittent breeding, the leatherback sea turtle. We used CR data collected on breeding females on Awa:la-Ya:lima:po beach (French Guiana, South America) from 1995 to 2002. By adding specific constraints in multistate (MS) CR models incorporating several nonobservable states, we modelled the breeding cycle in leatherbacks and then estimated the reproductive effort according to the number of years elapsed since the last nesting season. Using this MS CR framework, the mean survival rate was estimated to 0.91 and the average resighting probability to 0.58 (ranged from 0.30 to 0.99). The breeding cycle was found to be limited to 3 years. These results therefore suggested that animals whose observed breeding intervals are greater than 3 years were most likely animals that escaped detection during their previous nesting season(s). CR data collected in 2001 and 2002 allowed us to compare the individual reproductive effort between females that skipped one breeding season and females that skipped two breeding seasons. These inferences led us to conclude that a trade-off between current and future reproduction exists in leatherbacks nesting in French Guiana, likely linked to the resource provisioning required to invest in reproduction.     

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.     

Investment in testes,sperm‐duct glands and lipid reserves differs between male morphs but not between early and late breeding season in Pomatoschistus minutus

This study of the sand goby Pomatoschistus minutus, a nest‐holding fish with paternal care, focused on gonadal investment among males of different sizes collected early and late in the breeding season. All males caught at the nest had breeding colour, whereas trawl‐caught fish consisted of males both with and without colour. The absence or presence of breeding colour was a good predictor of testes investment. Compared to males with breeding colour, males without colour were smaller in body size but had extraordinarily large testes. In absolute terms, testes mass of males without breeding colour was on average 3·4 times greater than those of males with breeding colour. Since small colourless males are known to reproduce as sneaker males, this heavy investment in testes probably reflects that they are forced to spawn under sperm competition. Contrary to testes size, sperm‐duct glands were largest among males with breeding colour. These glands produce mucins used for making sperm‐containing mucous trails that males place in the nest before and during spawning. Since both sneakers and nest‐holders potentially could benefit from having large glands, this result is intriguing. Yet, high mucus production may be more important for nest‐holders, because it also protects developing embryos from infections. There was no significant effect of season on body size, testes or sperm‐duct glands size, but colourless males tended to be less common late in the season. Possibly this may indicate that individual small colourless males develop into their more colourful counterparts within the breeding season.     

Age-specific variation in reproduction is largely explained by the timing of territory establishment in the New Zealand stitchbird Notiomystis cincta

1. Using data from 327 nests over a consecutive 8-year period we examined age-specific variation in reproduction in a population of stitchbirds (or hihi) Notiomystis cincta and related how differences in reproductive performance were linked to the timing of territory establishment and breeding. 2. Across the population all reproductive parameters showed a quadratic relationship with an increase mainly between the first and second breeding season and a decline after the fourth year. A longitudinal analysis showed evidence of senescence by the sixth year in the numbers of chicks fledged and recruited. 3. Reproductive increases between years 1 and 2 were the result of poor-quality females dying after their first breeding season (differential selection hypothesis) in combination with surviving females showing improvements in reproduction in their second year (individual improvement/constraint hypothesis). 4. There was no effect of mate experience or territory quality on improvements in breeding between years. 5. The key variable influencing reproductive output was the timing of breeding. Birds that started laying earlier were more likely to lay multiple clutches in any given season. This was the main difference between first-year and older birds; generally first-year birds initiated egg laying later and consequently laid fewer clutches. 6. Approximately half of all first-year birds did not establish their territory until after the breeding season had begun. This delay in territory establishment resulted in these birds delaying breeding, which resulted in them having a lower reproductive output relative to all other birds. First-year birds that managed to establish their territory before breeding commenced, had similar rates of reproduction as older birds. 7. There was a positive relationship between the timing of territory establishment during a female's first year and her hatching date in the previous breeding season. We hypothesize that this was because late-hatched females were less able to effectively compete for territories against earlier-hatched members of their cohort, and this delayed their establishment and breeding in their first year. Thus, this social constraint is likely to be a major factor driving age-specific reproductive variation in this population.