Reproductive biology of the threatened golden galaxias Galaxias auratus Johnston and the influence of lake hydrology

Golden galaxias Galaxias auratus (31–235 mm fork length, L _F) were collected monthly from littoral habitats in Lakes Crescent and Sorell, Tasmania, Australia, between July 2000 and December 2002. Spawning habitats were identified and monitored in both lakes, and surveyed in Lake Crescent. Trends in gonado-somatic indices and reproductive stages of development indicated that gonad development in both sexes begins in midsummer and peaks in late autumn to early winter. Males mature at smaller sizes (50% at 52 mm L _F) than females (50% at 76 mm L _F), larger individuals are predominately females (95% of fish ≥138 mm L _F), and overall male to female ratios are female biased ( c . 1:2). Spawning occurs late autumn to early spring (water temperatures = 1·4–9·7° C) with peaks in spawning activity in winter (mean water temperatures <5° C). Demersal adhesive eggs ( c. 1·5 mm diameter) were found on cobble substrata ( c. 20–250 mm diameter) in littoral areas ( c. 0·2–0·6 m deep) and fecundity of fish 71–181 mm L _F ranged from 619 to 14 478 eggs. The rate of change in water level over the 20 days prior to monthly sampling was important in explaining the occurrence of spent fish and this accounted for temporal differences in spawning between the populations. Lake hydrology influences the reproductive cycle of G. auratus by possibly providing a stimulus for spawning and it controls the availability of spawning habitat in Lake Crescent. Seasonal hydrological cycles ( i.e. rises during late autumn to winter) and a minimum water level of 802·20 m Australian Height Datum in Lake Crescent during autumn (above which littoral areas of cobble substratum are inundated) are critical to G. auratus populations.     

Serum calcium concentrations in white sucker, Catostomus commersoni Lacepède, and bluegill, Lepomis macrochirus Rafinesque, in northern Wisconsin lakes: relation to pH and waterborne calcium

We measured serum calcium concentrations in mature white suckers and bluegills collected from clear-water Wisconsin lakes before spawning. The mean serum calcium concentration in female white suckers was greater in a circumneutral lake than in two naturally acidic lakes, whereas mean concentrations in fish from the two acidic lakes did not differ. Serum calcium concentrations in female bluegills from four acidic and four circumneutral lakes varied among lakes, but no relation was apparent between serum calcium level and either lake pH or waterborne calcium. In contrast, a strong negative correlation between mean serum calcium concentration and relative density of bluegill populations in the eight lakes was observed, suggesting that serum calcium levels in this species are affected by density-dependent factors. Mean serum calcium concentrations were substantially lower in mature male bluegills than in females, which is the norm prior to spawning. We hypothesized that serum calcium levels in female white suckers are depressed by exposure to pH 6.0 or less in calcium-poor waters. The utility of serum calcium concentrations, measured before spawning, as a measure of pH-related stress on populations of white suckers and other synchronous spawners merits further study. However, this measure is probably not useful for evaluating acid stress on fish species such as the bluegill, which have asynchronous reproductive cycles.     

Prevalence and intensity ofCapillaria catostomi (Nematoda: Trichuroidea) in white sucker (Catostomus commersoni) in southern Lake Huron,Canada

Synopsis A total of 314 white suckers (Catostomus commersoni) taken monthly during May to December 1975 and in April and May 1976 from southern Lake Huron, Ontario, Canada, were examined forCapillaria catostomi.Prevalence determined from fresh white suckers (66%) was significantly lower than from frozen white suckers (81%). However, there was no significant difference in prevalence ofC. catostomi in samples taken at monthly intervals; in male and female white suckers; or in the age categories.Mean intensity of infection was not significantly different in fresh (9.6 [± 13.41]) or frozen (10.4 [± 13.11]) white suckers. There was no significant difference in intensities between monthly samples in male and female white suckers or in the age categories. Suckers were infected with 1–172 worms. The percentage of male, female and immature worms did not change with sex or age of host or by month. The frequency distribution showed the parasites were overdispersed.A possible life cycle forC. catostomi involving an oligochaete intermediate host is discussed and an attempt made to explain the almost constant levels of prevalence and intensity.     

Gonadal development and associated changes in liver size and sexual steroids during the reproductive cycle of captive male and female Atlantic cod (Gadus morhua L.)

Gametogenesis in female and male Atlantic cod (Gadus morhua L.) was investigated by sampling blood plasma and gonadal tissue from 19 to 33-month-old fish. The reproductive cycles of both female and male Atlantic cod are characterized by distinct annual variations in gonadal size and developmental stage and these are associated with changes in sex steroids and liver size. I(H) did not change during early gonadal development, but both spent females and males had lower I(H) than late maturing females and spermiating males, respectively. In females I(G) was correlated to plasma E2 levels and they were highest in spawning females. The lowest levels during the reproductive cycle were observed in spent females. Plasma T levels were low throughout ovarian development, and were at a minimum in spent females. 11-ketotestosterone in plasma of males increased rapidly during spermiation, while T increased at earlier testicular stages and reached maximum during spermiation. High plasma levels of steroids in male and female cod during spawning serve to promote further development and growth of less advanced stages of germ cells.     

Life history traits of Scardinius acarnanicus (Economidis, 1991) (Pisces: Cyprinidae) in two Greek lakes (Lysimachia and Trichonis)

Age, growth, reproduction and mortality of Scardinius acarnanicus, an endemic cyprinid fish from central and western Greece, were investigated in two contrasting environments, the eutrophic Lake Lysimachia and the oligotrophic to mesotrophic Lake Trichonis. Maximum ages observed were 7+ for males and 10+ for females in Lake Lysimachia and 8+ for males and females in Lake Trichonis. Scardinius acarnanicus grew allometrically (slope of fork length–somatic weight regressions >3) and relatively rapidly until age 2 or 3 years, then slowed at a time coincident with maturation. Total instantaneous mortality rates of fish from Lake Trichonis were higher than in Lake Lysimachia. Scardinius acarnanicus is a multiple spawner; monthly values of the gonadosomatic index (GSI) indicated that spawning occurred mainly in April. Mean egg diameter in Lake Lysimachia was 1.54 ± 0.2 and 1.28 ± 0.16 mm in Lake Thrichonis. In Lake Lysimachia, the population sex ratio departed markedly from 1 : 1; females were more abundant at ages >3 and the overall male : female ratio was 1 : 1.47. In Lake Trichonis, females were more abundant at ages >6; the overall male : female ratio was 1 : 1.16, not significantly different from a 1 : 1 ratio.     

Female mate choice and spawning behaviour of Chinook salmon under experimental conditions

After spawning their first nest, female chinock salmon Oncorhynchus tshawytscha paired with small males ( c. 46% of female weight) spent an average of 16·2 h between spawning of successive nests compared with 9·6 h for females paired with large males ( c. 112% of female weight) ( P< 0·05). Neither frequencies of female nest construction behaviours (digging and probing) nor male courtship behaviour (crossovers and quivers) differed between large- and small-male pairs. Male quivering frequencies were correlated significantly with female digging and probing frequencies, whereas the crossover frequencies were not. It is suggested that delayed spawning by females in the presence of relatively small males is a primary mechanism by which females in the genus Oncorhynchus exhibit mate choice.     

Reproductive life history of the introduced peacock grouper Cephalopholis argus in Hawaii

This research investigated the reproductive biology (sex ratio, hermaphroditic pattern, size and age at maturity) of Cephalopholis argus, known locally in Hawaii by its Tahitian name roi. The results suggest that C. argus exhibits monandric protogyny (female gonad differentiation with female to male sex change) with females reaching sexual maturity at 1.2 years (95% c.i .: 0.6, 1.6) and 20.0 cm total length (L_T; 95% c.i .: 19.6, 21.2). The female to male sex ratio was 3.9:1. The average age and L_T at sex change was 11.5 years (95% c.i .: 11.1, 12.9) and 39.9 cm (95% c.i .: 39.5, 41.2), respectively. Current information on spawning seasonality of this species is incomplete, but based on the occurrence of spawning capable and actively spawning females, spawning probably takes place from May to October. Evidence of lunar spawning periodicity was found, with an increased proportion of spawning capable and actively spawning females, and an increased female gonado‐somatic index during first quarter and full‐moon phases. This information fills a valuable information gap in Hawaii and across the species' native range.     

Reproductive aspects of the yellowtail snapper Ocyurus chrysurus from the southern Gulf of Mexico

In this study, sex ratio, spawning season, fork length (L(F)) at maturity (L(F50)), batch fecundity and spawning frequency were characterized for the continental population of Ocyurus chrysurus from the Campeche Bank, in the southern Gulf of Mexico. A total of 1657 specimens were collected from February 2008 to January 2009. The overall sex ratio (male:female) and sex ratios by size-class showed no significant differences from an expected 1:1 ratio. The Campeche Bank population did not conform to the reproductive seasonality pattern characteristic of a continental population. A protracted spawning season that extended from January to September with peaks occurring mainly between April and May and additionally in September was observed. The population conformed, however, to the sexual maturity pattern observed for populations and species associated with a continental margin. Fish of both sexes reached the onset of sexual maturity at a similar and small L(F) of c. 14 cm, and L(F50) (L(F) at which 50% of females and males become mature) was 21·3 and 19·4 cm. Asynchronous-type ovarian development was observed for this species and batch fecundity estimates ranged from 14,102 to 164,756 oocytes (mean ±S.D. = 43,852 ± 32,684 oocytes). The overall spawning frequency estimate was once every 8·3 days or 26 times during the 9 month spawning season.     

Steroid hormone profiles and correlative gonadal histological changes during natural sex reversal of sobaity kept in tanks and sea-cages

The gonadosomatic index of sobaity Sparidentex hasta fits well with the natural spawning season of February-March. At 12 and 24 months all the population matured as males. However, during this time, between April-September intersex gonads were seen. From October of the third year of life, females were seen for the first time. These changes in the histology of the gonads were correlated with the hormonal profile. Testosterone showed no significant correlation with sex but 11-ketotestosterone correlated very well with the mature male gonads and thus can be used for sexing. Oestradiol-17β higher in sex reversed individuals (females) in the spawning season than in males. The sequence of events in the protandrous sex change of this fish during the first two spawning seasons is male → intersex → male. However, during the third spawning season (age 36–37 months) this sequence is male →intersex → male or female (10–17%).     

Reproductive biology of the East African lungfish (Protopterus aethiopicus) in Mwanza Gulf, Lake Victoria

The population of lungfish (Protopterus aethiopicus (Heckel)) was sampled by bottom trawling for 12 months from June 1984 to May 1985. A total of 576 lungfish were investigated. The sex ratio in the population was 1 male : 1 -98 females. Average size at maturity was found to be 960cm total length (TL) but the smallest matured male measured 72-9 cm and the female 840 cm TL. It was not possible to establish spawning times and reproductive activity trends either from the variation in relative condition factor ‘kn’ or from gonadosomatic index (GSI). Whereas male lungfish seem to show no further increase in GSI after gonadal maturity stage III, females exhibit GSI values that vary directly with oocyte maturation and ovarian weights. Fat deposited along the gonads was found to be a better basis for assessing maturation and spawning in lungfish. The potential spawning period was found to be from September to May. Absence of lungfish with spent gonads over the 12-month period and the presence of different egg sizes in ovaries, suggest that the fish is a multiple spawner. Fecundity was found to range from 705 eggs (in 84-cm TL fish) to 14,922 eggs (in 130-cm TL Fish).     

Synchronous female spawning and male mating behavior in a land-locked population of Japanese charr, Salvelinus leucomaenis japonicus

In resource-based promiscuous mating systems, synchronous spawning of females affects competition among males and variation in the reproductive success of males. We documented the mating behavior of Japanese charr (Salvelinus leucomaenis japonicus) through an annual breeding season to examine the relationship between female spawning synchrony and male mating behavior. Females spawned highly synchronously in the population studied, i.e., approximately half the spawning was finished within the first three days of the entire spawning season (11 days). The daily operational sex ratio (OSR) was nearly 1:1 through the spawning period. The number of males around a spawning female was very small (1.21+/-0.49 males per female) over the spawning ground and period, suggesting that a competitive male could effectively chase subordinate males away from a spawning female. A few males attempted to sneak near the oviposition site of females (16%; 9 of 57 breeding groups), while some males adopted sneaking tactics in the initial phase of females' spawning (24%). We did not observe any males to succeed in sneak fertilizations. We conclude that in this Japanese charr population, the synchronous spawning of females was related to the unbiased daily OSR, male aggregation around females, and consequently whether and how efficiently males engaged in sneak mating behavior.     

Characteristics of the spawning migrations of brown trout, Salmo trutta L., and rainbow trout, S. gairdneri Richardson, in Great Lake, Tasmania

Upstream spawning migrations of mature brown trout, S. trutta , and rainbow trout, S. gairdneri , were studied in Liawenee Canal, Great Lake from 1949 to 1985. Brown trout migrations normally occurred from early April to mid-May and rainbow trout from late August to early November. In 1983, 16 425 brown trout and 1338 rainbow trout passed through a fixed upstream diversion trap. Brown trout spawning migrations occurred predominantly over the temperature range 6–10° C, while rainbow trout migrated predominantly over the range 5–11° C. Migrations peaked at water temperatures of 7.6°C (males) and 7.8°C (females) for brown trout, and 8.3°C (males) and 9.6°C (females) for rainbow trout. Rainbow trout migrations occurred at high flow conditions and were positively correlated with canal flow increases, while brown trout migrated under low canal flow. Mean length, weight and condition of rainbow trout of both sexes decreased significantly during migrations. Female brown trout decreased in weight and condition but not in length; male brown trout did not change in condition despite decreases in both length and weight during migrations. Overall sex ratio was 2:1 (female:male) for both species, with the relative proportion of male fish decreasing as migrations progressed. Age composition changed during migrations; dominant age classes were 3 < 4 < 5 + years for both species. Comparison of length, weight, condition and age revealed minor changes during the 37-year period 1949–1985.     

Effect of altitude on the sex ratio in populations of Silene acaulis (Caryophyllaceae)

Alatalo, J. M. & Molau, U. 1995. Effect of altitude on the sex ratio in populations of Silene acaulis. — Nord. J. Bot. 15: 251–256. Copenhagen. ISSN 0107–055X.
Predicted increase of female frequencies in gynodioecious Silene acaulis was tested along an altitude gradient in northern Sweden. Average female frequencies for the four sites increased with altitude from 42% to 59% within a short geographical distance. This follows the outcrossing hypothesis, that female frequencies should be positively correlated with selfing rates of hermaphrodites in populations. More adverse environmental conditions should favour gynodioecy in areas where reproduction to a greater part relies on vegetative reproduction or selfing. Further, a significant difference in corolla width was found between females and the larger hermaphrodites, but not between sites. Cushion size and the number of flowers per cushion decreased with attitude.     

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     

Effective size of subpopulation of the early run sockeye salmon Oncorhynchus nerka from Azabach's Lake (Kamchatka): effect of density on variance of reproductive success

Using Parensky's approach for estimating the number of breeding pairs, we determined effective subpopulation size Ne in early-run sockeye salmon Oncorhynchus nerka from Azabach'e Lake (Kamchatka) in 1977 through 1981. On average (over years and populations), biased sex ratio decreased Ne by 7% as compared to the number of fish on the spawning sites (Ni). High density reduced the Ne/Ni ratio by 62-66% because some fish were excluded from spawning. Dominance polygyny as compared to monogamy and random union of gametes could reduce Ne by about 17%.     

Sex ratio and maternal investment in the multivoltine large carpenter bee Xylocopa sulcatipes (Apoideh: Anthophoridae)

Abstract. 1. Females of the multivoltine carpenter bee Xylocopa sulcutipes (Maa) (Hymenoptera: Anthophoridae) usually excavate a straight tunnel in dead twigs and mass provision a linear array of up to ten brood cells with pollen and nectar. An egg is deposited upon each food mass within one cell.
2. Female offspring generally receive a higher provisioning mass (0.180 ± 0.048 g) than males, a significant difference ( P > 0.001). There are, however, male larvae that receive as much food or more as their sisters or female larvae reared in another nest.
3. There is a close positive association between the size of a mother and the weight of provisions for individual daughters, but not for sons.
4. Female offspring are positioned in the innermost brood cells (Gositions 1, 2 and 3). The sex ratio of the outer cells is either significantly male biased (positions 4–6) or skewed towards males (positions 8 and 9). Positions 7 and 10 are in equilibrium.
5. Solitary females produce a significantly female biased sex ratio ( P < 0.01). Sex ratio in social nests is skewed toward females, but not significantly so ( P < 0.2). There is no significant difference between the sex ratio of solitary and social nests ( P = 0.361). The population sex ratio (pooled sex ratio of all broods produced) is significantly female biased ( P = 0.003).
6. Females kept in the laboratory produced female biased sex ratios whilst unmated females produced all-male broods indicating that insemination and ovarian development are not causally related.
7. The expected sex ratio (ESR) under equal investment, calculated as 1/CR (CR = mean male provision weight/mean female provision weight), is 137.5:117.5 (males:females), and differs significantly from that observed, 104:151 (males:females) ( P < 0.001). The 'Local Resource Enhlancement' hypothesis best explains the female biased sex ratio found in X.sulcatipes and its maintenance in the population.     

Evolutionary consequences of cytoplasmic sex ratio distorters

Summary Computer simulations of diploid genetic models were used to examine the consequences of the spread of a cytoplasmic sex ratio distorter on the frequencies of nuclear sex-determination alleles and the spread of nuclear resistance alleles in female biased populations. The cytoplsmic elements considered here override the expression of the nuclear sex-determination genes, turning genetic males into females. When homozygous male genotypes are viable, a cytoplasmic sex ratio historter spreads in a population if the proportion of daughters produced by infected females exceeds the proportion of daughters produced by uninfected females. The equilibrium frequency of male phenotypes is the proportion of uninfected progeny produced by infected females. When homozygous male genotypes are lethal, the conditions for the spread of the cytoplasmic element are more stringent. The spread of a cytoplasmic sex ratio distorter causes an increase in the frequency of nuclear male sex-determination alleles as a result of the unusual combinations of genotypes which mate in infected populations. Eventually, a cytoplasmic element may replace the nuclear gene as the sex-determination mechanism. This occurs without selection. Nuclear genes conferring resistance to cytoplasmic sex ratio distorters generally increase in female biased populations and often restore a 11 sex ratio despite continual selection on the cytoplasmic element to increase its transmission efficiency.     

The reproductive cycle of yellowfin bream, Acanthopagms australis (Günther), with particular reference to protandrous sex inversion

Yellowfin bream, Acanthopagrus australis , of all age classes were collected from Moreton Bay, Australia. The species possessed typical sparid ovotestes in which the testis and ovary occur in separate zones. During the spawning period (June-August) juveniles, functional males and functional females could be distinguished by the macroscopic appearance of the gonad. The sex ratio of males to females decreases with age, indicating protandrous sex inversion.
Histological and structural study of the ovotestis showed all fish have previtellogenic cells in the ovarian zone but only juvenile and male fish have developing spermatogenic cells in the testis. Most juveniles become functional males by the age of two years but a small proportion of juveniles develop directly into functional females (primary females). Protandrous sex inversion commences after the spawning period when male fish appear with spermatozoa and no other spermatogenic cells in the testis. During the period November-January male fish with no spermatogenic cells are common and a reduction in size of the testis occurs so that by March-April the ovotestis becomes structurally and histologically similar to the female ovotestis. Some fish remain functional males during their whole life-history (primary males). In functional females vitellogenic cells are present in the ovary only during the spawning period and the testis remains very small in size.     

Gonadal development and spawning cycle in the digynic hermaphrodite sharpsnout seabream Diplodus puntazzo (Sparidae) off the Canary Islands, northwest of Africa

Diplodus puntazzo is a species characterized by a rudimentary hermaphroditism, with a low level of protandry. The study found that the overall sex ratio was one male to 2.03 females, diverging significantly from 1 : 1, and that there was a ratio of one male to 0.11 transitional fish. The sex ratio varied with season: during the reproductive period the ratio was 1 : 1.86 (male : female), significantly lower than in the non‐reproductive period in which the sex ratio was approximately 1 : 2.26. Both sexes had similar size distributions; however, females were predominant in all size intervals, especially in the larger ones. No significant difference in mean total length (TL) between females (330 mm), males (319 mm) or transitionals (321 mm) was found. The smallest functional female and male analysed were 159 and 157 mm TL, and the largest were 520 and 510 mm TL, respectively. No significant difference in length at first level of maturity, attained at the second year of life, was found between males (281 mm) and females (292 mm), respectively. A protracted spawning season was identified for the species: from September to February, with a peak in November. GSI values followed the same pattern for males and females, with values of females being similar to those of males; the highest values occurred between September and December, with a maximum in November. Occurrence of transitional fish was significantly higher during the month with highest values of GSI than in the resting period. The study found that vitellogenesis did not begin synchronously for all females, and the simultaneous presence of oocytes in all stages of development indicated an asynchronous mode of ovarian development. The presence of postovulatory, together with tertiary yolk vesicle oocytes, indicated that the species is a multiple spawner.     

The reproductive success of pike, Esox lucius: aspects of fecundity, egg density and survival

Power-law relationships have been estimated between fecundity and fish length, weight and age for pike from two gravel pit lakes (Main Lake and St Peter's Lake) in Buckinghamshire, England. Relative fecundity, estimated at 17.6 eggs g⁻¹ for Main Lake pike and 19.4 g⁻¹ for St Peter's Lake pike, did not differ significantly between the lakes. Population fecundity for the Main Lake was estimated at 10.6 million eggs and 6.4 million eggs in 1986 and 1987, respectively.
Experimental work involving samples of eggs from 18 artificially fertilized Main Lake female pike showed a significant relationship between egg diameter and female length, and a significant difference between mean egg diameters for 41-cm and 101-cm females. No significant relationships were found between mean egg size and mean fry size, nor between fry length and adult length. Mean size differences between newly hatched, 18-day-old and 41-day-old fry from the 41-cm and 101-cm females were not significant.
Samples of eggs taken at 2-day intervals from 6 April to 5 May in a pike spawning area of the Main Lake revealed clumped distributions of eggs, with average egg densities ranging from 1 to 47 eggs per 0.071 m² (14–671 m ²) on sand and silt substrata. The maximum egg density of 51 per 0.071 m² (729 m⁻²) was found on flooded grass. The stages of development of the eggs were identified over the sampling period. Survival from stage 1 to fry was estimated at 3–6%. Egg losses were estimated at 9–10% day⁻¹.
Egg survival experiments gave overall hatching success rates of 11 % on sand/silt compared with 2.5% on aquatic plants (1986), and 18.9% on undisturbed sand/silt compared with 7.9% on disturbed sand/silt (1987). The effect of siltation on egg hatching is discussed.