Effect of variable food levels on reproductive performance of breeding female three-spined sticklebacks

The study examined the response to variation in food ration within a breeding season on components of the reproductive performance of mature female three-spined sticklebacks Gasterosteus aculeatus , a batch spawner. The main effect of food availability was on the duration of the inter-spawning interval (ISI). Low ration resulted in an increase in the number of days until the next spawning. A low ration during one inter-spawning interval increased the length of the next ISI, even if there was a high ration in the latter. Females that experienced high rations for two successive ISIs showed an increase in batch fecundity over the successive spawnings. There was no tendency for fish on low rations to produce bigger eggs or eggs with higher lipid concentration. Females on low ration had a lower carcass dry matter concentration and a lower liver dry weight immediately after their third spawning. Female sticklebacks responded to a low ration with little change in egg number or size, but a slight reduction in physiological condition. Egg characteristics and, to a lesser extent, batch fecundity are insensitive to current rate of food consumption, but the interval between spawnings is sensitive to both current and previous rates of consumption.     

Effect of spawning number and ration on reproductive performance of the batch-spawning three-spined stickleback Gasterosteus aculeatus

An experiment quantified the effect of food ration and spawning number on the breeding season reproductive performance of batch-spawning, female three-spined sticklebacks Gasterosteus aculeatus . Individually housed females were maintained on fixed rations of 2, 4, 8 or 16% of their initial postspawning mass of live enchytraeid worms from immediately after their first spawning until they ceased spawning. Number of spawnings correlated positively with ration. Total breeding season reproductive investment (total wet mass of eggs produced as a percentage of initial postspawning female mass) ranged from 38% at the 2% ration to 147% at the 16% ration. At the lower rations, postspawning mass, batch fecundity and wet and dry masses of the batch declined over successive spawnings, with the rate of decline inversely related to ration. At the highest ration, there was no decline in batch fecundity and postspawning mass increased over successive spawnings. Mean reproductive investment per inter-spawning interval was higher at the highest ration, but at all rations declined over successive spawnings. Mean reproductive effort per inter-spawning interval (wet mass of eggs spawned as a percentage of the wet mass of food consumed over the inter-spawning interval) was inversely related to ration. At the higher rations, reproductive effort showed no trend over successive spawnings. Neither spawning number nor ration had a systematic effect on egg diameter, wet mass per egg, dry mass per egg or total lipid content of the eggs. If the rate of food intake was insufficient, although batch fecundity declined, the main adjustment was a reduction in the number of spawnings in the breeding season.     

The effects of rainfall on different components of seasonal fecundity in a tropical forest passerine

Seasonal fecundity is a composite metric that is determined by component parameters such as clutch size, nest survival and re‐nesting probability. Many of these component parameters are known to vary with environmental conditions, in particular rainfall prior to or during the breeding season. In some species, seasonal fecundity is positively related to rainfall, but little is known about which component parameters of seasonal fecundity respond most strongly to rainfall. We used intensive nest monitoring of a multi‐brooded tropical forest passerine, the Montserrat Oriole Icterus oberi, to examine the effects of rainfall during the pre‐breeding season on component parameters of annual fecundity. We monitored all nests of a total of 42 pairs over 5 years in which rainfall varied substantially. We then related clutch size, nest survival, onset and length of the breeding season, re‐nesting probability and re‐nesting interval to pre‐breeding season rainfall using generalized linear mixed models that accounted for random variation across sites and individual pairs, and incorporated other variables known to affect the response. Higher pre‐breeding season rainfall led to an increase in clutch size and a decrease in re‐nesting interval, but nest survival, re‐nesting probability and length of the breeding season were not affected by variation in rainfall. The onset of the breeding season was delayed in very dry years. We conclude that higher rainfall is likely to increase food availability and thus body condition of female Montserrat Orioles, leading to an increase in fecundity due to larger clutch sizes.     

Fecundity of the three-spined stickleback, Gasterosteus aculeatus (L.)

The number of eggs spawned by female three-spined sticklebacks Gasterosteus aculeatus (L.) was highly correlated with the size of the fish expressed either as total length or as weight after spawning. There were no significant correlations between the size of the eggs measured either as wet weight per egg or as dry weight per egg, and the size of the fish. Nor were there significant correlations between the calorific value of the eggs and the length or weight of the fish. Regressions relating egg production to the length and the weight of the fish are given. These results and an analysis of previous studies on the fecundity of the stickleback suggest that variations in fecundity are primarily a reflection of variation in the size of the fish at maturity, and that this size is related to the race of the stickleback and to environmental conditions such as food supply which influence the growth of sticklebacks.     

Life-history variation within a three-spined stickleback population in the Camargue

Among individuals of female three-spined sticklebacks Gasterosteus aculeatus from a population in the Camargue, southern France, studied in 12 successive years, adult L _T ranged from 31–64 mm, clutch size ranged from 33–660 eggs, and mean egg diameter per clutch ranged from 1.15–1.67 mm. Because the population was strictly annual, inter-annual variation corresponded to variation among generations having experienced different environmental conditions. Body mass varied significantly among years, suggesting an effect of varying environmental conditions. Gonad mass and clutch size increased with body mass, but mean egg diameter was not correlated to body mass. Body mass-adjusted gonad mass, interpreted as reproductive effort per clutch, did not vary significantly among years, suggesting that this trait was not influenced by environmental conditions. Body mass-adjusted clutch size and egg size varied significantly among years. Inter-annual variation in body length at breeding, clutch size and egg size was of the same order of magnitude as inter-population variation reported by other authors for this species. During the breeding season, reproductive effort and clutch size tended to increase. Egg size tended to decrease during the breeding season but this seasonal pattern varied among years. Observed life-history variation is discussed both in terms of its evolutionary significance and methodological implications in the study of life-history variation among populations.     

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.     

Reproductive regulation in the female Japanese minnow, Pseudorasbora parva (Cyprinidae)

The Japanese minnow, Pseudorasbora parva, is a multiple spawner that lays eggs repeatedly during the spawning season. In laboratory aquaria, 19 of 63 females did not spawn, whereas 44 other females laid 167 to 6285 eggs in 1 to 14 mating sequences. Neither fish density nor sex ratio (number of males per female) affected fecundity, but the increase in fish density significantly decreased the growth rate of females. The increase in male length significantly increased fecundity in aquaria which contained only a single male, whereas it decreased female growth rate. These relationships were not evident in aquaria housing three or ten males. Since larger males were more dominant and had larger testes than smaller males, the correlation between fecundity and male size strongly suggests that individual females regulated fecundity to increase their reproductive success.     

Empirical evidence of a trade-off between reproductive effort and expectation of future reproduction in female three-spined sticklebacks

Optimal life-history models generally predict that the reproductive effort of iteroparous organisms may increase with age, as their expectation of future reproduction decreases. The population of three-spined sticklebacks (Gasterosteus aculeatus) in the Camargue (Rhone River Delta, France) is annual, all adults dying after their first breeding season. As the three-spined stickleback is a multiple spawner, we tested the hypothesis that reproductive effort may increase during the breeding season on field data. From 1987 to 1998, 653 female sticklebacks were collected in the field during the breeding seasons. The body size, body weight and weights of the liver, gonads and carcass were measured for these individuals. Only gravid females with mature eggs (176 fish) were included in the analysis. Considering the female three-spined stickleback as a capital breeder, the energetic resources available for allocation between soma and gonads were estimated by its body weight. Somatic condition decreased during the breeding season and reproductive effort (gonad weight relative to body weight) increased. These patterns did not vary significantly between years. These observed variations in reproductive effort during the breeding season can be interpreted as empirical evidence of a trade-off between reproductive effort and expectation of future reproduction.     

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.     

Maternal investment in reproduction and its consequences in leatherback turtles

Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001–2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.     

Aspects of reproductive biology of Labeo horie Heckel (Pisces: Cyprinidae) in Lake Chamo, Ethiopia

Labeo horie Heckel is increasingly becoming commercially important in Lake Chamo but its reproductive biology in the lake is not well enough to guide its management. Sex ratio, breeding season, length at first maturity and fecundity of L. horie were studied from 1197 fish samples collected between August 1998 and October 1999 in Lake Chamo, Ethiopia. The sex ratio was significantly different (χ ² = 12.12, P < 0.001). The peak‐breeding period was during the rainy months of March to May, during which time more than 90% of both sexes had ripe gonads. The size at maturity ( L _m50 ) of males was 52 cm while the L _m50 of females was 62 cm. The smallest ripe male in the catch was 46.7 cm and weighed 890 g while the smallest ripe female caught was 49.5 cm and weighed 1145 g. The weight of ripe ovaries ranged from 54.3 g to 991.8 g and contained between 68,400 and 1,320,400 eggs. Relative fecundity ranged between 60 and 290 eggs per g of body weight. The relationships between fecundity and total length and between fecundity and total weight were curvilinear, while the relationship between fecundity and ovary weight was linear. L. horie conformed to the general pattern of reproduction in a tropical environment where peak‐breeding activity occurred during the rainy season. In order to protect the spawning population, fishing pressure should be minimized during breeding time at the shallower littoral regions.     

宁波滑蜥两性异形和雌性繁殖

蜥蜴的雌性繁殖特征对理解两性异形的进化原因起着重要作用。于2011年4月在安徽滁州采集宁波滑蜥(Scincella modesta),定量研究该种形态特征的两性异形和雌性繁殖特征,检验成体形态特征两性异形与雌性繁殖的相关性。研究共采集43条(17♀♀,26♂♂)宁波滑蜥,雄性和雌性个体的最大体长分别为47.4 mm和46.6 mm。雌雄两性在体长和头宽上没有差异,而在腹长和头长上差异显著,雄性有较大的头长,雌性有较大的腹长。宁波滑蜥年产单窝卵。窝卵数和窝卵重与雌体体长及腹长呈正相关,卵重与雌体体长无相关性。窝卵数及卵重的变异系数分别为0.20和0.12。卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数来增加繁殖输出。这些结果表明,宁波滑蜥是雌雄个体大小同形的两性异形模式,性选择使得雄性有着较大的头长,以具有较高的交配成功率,生育力选择使得雌性有着较大的腹长,以具有较大的生育力和繁殖输出。     

Maturation and Reproductive Biology of Female Wild Carp, Cyprinus carpio, in Victoria, Australia

The reproduction of carp Cyprinus carpio L. (Family: Cyprinidae), from Victorian waters in Australia, is studied with detailed analysis of gonad maturation, spawning season, fecundity and oocyte diameter. Results show that carp has a high annual fecundity (AF) (0.12–1.54 million oocytes per fish), which is positively correlated with caudal fork-length (L, mm) and total weight (W, kg) but not age. The relationships between length or weight and AF were statistically significant and best described with the simple linear or quadratic regressions. Mean relative fecundity was 0.163 million eggs kg^–1 whole weight. Egg size was estimated from oocyte diameter in carp from eight stocks. Egg size was proportional to maternal size but not age. Seasonal trends in gonadosomatic indices, together with the changes in the macroscopic and microscopic condition of ovaries, demonstrated that spawning generally peaks during spring–early summer, but also occurs through until autumn and can even start in late winter at some sites. In Victoria, this species is a multiple spawner with asynchronous oocyte development and a protracted spawning season. Stocks generally contain both females that spawn once, and females that spawn repeatedly, within a spawning season. Implications for management for the control of feral carp stocks are discussed.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

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

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

Is body size a good indicator of fecundity in the genus Thaumetopoea? A story told by two intrageneric Mediterranean forest defoliators

1. Body size correlates with several factors such as reproductive fitness, environmental changes, the quality and quantity of food during critical development stages, and the feeding season. For the Palearctic moths of the genus Thaumetopoea (Lepidoptera; Notodontidae), the larval development is crucial and differs between species according to their feeding season; larvae of the pine processionary moth Thaumetopoea pityocampa (Denis & Schiffermüller 1775) feed during winter while larvae of its congeneric cedar processionary moth Thaumetopoea bonjeani (Powell 1922) develop during summer in North Africa.
2. This discrepancy in lifecycles leads to different reproductive traits such as egg batch length, number of eggs per batch, eggs protection mechanisms and female body size.
3. According to Darwin's fecundity advantage hypothesis (1871), female body size should have a positive influence on reproductive fitness, since larger females supposedly have higher fecundity. The universal allometric scaling phenomenon rule proposed by Rensch (1950) predicts that the degree of sexual size dimorphism tends to decrease with the increase of female body size.
4. Here, two morphometrical parameters that is, body size and scale size, estimated from body measurements of individuals of both species, feeding on the same host Atlantic cedar Cedrus atlantica (Manetti & Carrière 1855) (Pinales; Pinaceae) in Algeria were proposed. The aim was to find out traits that might rule the competition for food and space, in particularly fecundity and body size.
5. Results of the present study highlight a female-biased sexual size dimorphism in both species. The positive correlation between female body size and fecundity shown in this study weakly supports Darwin's hypothesis. Finally, the intrageneric test performed leads to conclude that Rensch's rule does not hold in the considered species.

     

Measuring temporal variation in reproductive output reveals optimal resource allocation to reproduction in the northern grass lizard, Takydromus septentrionalis

We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005 to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a particular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producing more, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annual reproductive output were all positively related to female size (snout–vent length). Females switched from producing more, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutch mass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequent clutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, but clutch mass of either the first or the second clutch remained unchanged across the sampling years. The regression line describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither the line for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more markedly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs and between reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis ; (2) females allocate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shifts in reproductive output and egg size are determined ultimately by natural selection.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 315–324.     

A comparative study of egg mass and clutch size in the Anseriformes

The factors explaining interspecific differences in clutch investment in precocial birds are poorly understood. We investigated how variations in clutch characteristics are related to environmental factors in a comparative study of 151 extant species of ducks, geese and swans (Anseriformes). Egg mass was negatively related to clutch size in a phylogenetic regression, a relationship that was much stronger when controlling for female mass. Nest placement was related to both egg size and clutch size, with cavity-nesting species laying more but smaller eggs. Egg size was positively correlated with incubation period and with female mass, and also with sexual size dimorphism (i.e. male mass relative to that of the female). Clutch size was not related to female mass. Species with long term pair bonds laid smaller clutches and larger eggs. The size of the breeding range was strongly positively correlated with clutch size and clutch mass, and its inclusion in multivariate models made other biogeographical variables (hemisphere, breeding latitude or insularity) non-significant. The small clutches in insular species appear to be a product of small range size rather than insularity per se. Our results suggest there is an evolutionary trade-off between clutch and egg size, and lend support to Lack’s resource-limitation hypothesis for the waterfowl.     

Repeated Mating and Female Fecundity in the Simultaneously Hermaphroditic Land Snail Arianta arbustorum

Summary

To evaluate the influence of repeated mating on female fecundity in the simultaneously hermaphroditic, self-incompatible land snail Arianta arbus-torum, the number and size of clutches, egg size and hatching success of individuals from 3 populations were determined under conditions of isolation and grouping during one breeding season in a field cage experiment. Only adult snails which had mated and oviposited in the preceding year were used.

Sperm storage enabled isolated individuals to continue with the production of fertilized eggs. Snails kept singly or in groups differed neither in number of clutches laid nor in egg size. But isolated snails laid smaller clutches than did grouped snails. As a result isolated snails produced fewer eggs per breeding season. Hatching success varied greatly between populations and rearing conditions. In general, isolated snails had fewer hatchlings than grouped snails, indicating that individuals prevented from remating suffered a reduced fitness.