Seasonal bottlenecks in diet shifts and growth of smelt in a large eutrophic lake

Seasonal growth of smelt Osmerus eperlanus was studied in the Kiihkelyksenselkä basin in the eutrophic Lake Hiidenvesi, southern Finland. The growth of age 1+ and 2+ year smelts was very low probably due to the discontinuous supply of large invertebrates Chaoborus flavicans and Mysis relicta . Diet shift back to zooplankton in autumn, suggested the existence of a bottleneck in the growth of these age groups. Slow growth rate was further intensified by the onset of maturity at age 2 years, combined with the high diet and habitat overlaps between similar‐sized 1+ and 2+ year smelts. Older 3+ and 4+ year smelts shifted to piscivory, i.e . cannibalism, during the growing season, which was seen as rapid increases in their growth rates.     

First maturity and spawning periodicity of hatchery‐origin pallid sturgeon in the upper Missouri River above Fort Peck Reservoir,Montana

The pallid sturgeon Scaphirhynchus albus conservation propagation program has augmented declining wild populations since the 1990s and the older age classes of hatchery‐origin fish are beginning to reach sexual maturity in the wild. Currently, the majority of the information available on the age and size at first maturity and spawning periodicity for pallid sturgeon in the upper basin is from captive hatchery‐origin pallid sturgeon (i.e. age and size at first maturity and spawning periodicity) or from wild pallid sturgeon artificially spawned in the propagation program (i.e. spawning periodicity). The purpose of this study was to document age and size at first maturity and spawning periodicity of known age hatchery‐origin pallid sturgeon that have reached maturity in the wild. Radio‐tagged pallid sturgeon in the upper Missouri River upstream of Fort Peck Reservoir were serially sampled in the early‐spring over multiple years and assigned to reproductive classifications each year based on sex‐steroid concentrations. The youngest reproductively‐active male hatchery‐origin pallid sturgeon sampled was 14.5 years old and the youngest female was 18. Hatchery‐origin males were observed having annual (N = 3) and biennial (N = 2) reproductive cycles. The observed spawning periodicity was similar to what has been reported elsewhere for the species. The youngest mature fish in this study are older and larger than what has been reported for those retained in captivity, indicating that body size alone is not a reliable predictor of maturity for pallid sturgeon.     

Marbled lungfish growth rates in Lake Baringo, Kenya, estimated by mark‐recapture

Mark‐recapture was used to quantify the relationship between body mass ( M _T) and individual growth rates of sub‐adult marbled lungfish Protopterus aethiopicus in Lake Baringo, Kenya. Specific growth rate (in mass) was found to be a well defined and decreasing function of M _T over the size range of recaptured fish. Growth trajectories based on this function indicated that Lake Baringo African lungfish reached maturity at an age of c . 3 years and had a low reproductive effort.     

Variation in brook and brown trout survival within and among seasons, species, and age classes

The Program MARK was used to generate and test a plausible set of survival models for brook trout Salvelinus fontinalis and brown trout Salmo trutta to determine whether survival differed by season, species or age class. Apparent survival varied with time and age, but not by species. For the older (1999) age class, survival was lowest during the autumn of their age 1+ year whereas survival was lowest for the younger (2000) age class during the early summer of their age 1+ year. Additionally, estimates of survival for the younger age class exceeded those of the older age class in all but one interval (early summer 2001) but significant differences were observed in only one of these intervals (autumn 2000). To determine whether the observed seasonal differences in survival were related to seasonal differences in movement rates, multi‐strata spatial models were used within Program MARK to determine the probability of moving for each interval. In‐site movement rates were found to be low regardless of season (average for all cohorts combined was 5%). The ability of the multi‐strata modelling approach to detect temporal variability in movement, however, was potentially limited by spatial scale of the study reach ( c . 1 km). Differences in survival between different aged fishes could lead to selection acting on age at maturity or reproductive effort at a given age.     

Puberty and recrudescence in cultured male yellowtail flounder: plasma androgens and spermatogenesis

Two year classes of cultured male yellowtail flounder Limanda ferruginea were followed in order to examine puberty and recrudescence at the gonadal level, and to assess the impacts of culture conditions on male reproduction. Cultured males demonstrated a propensity for maturation as 1 year‐old fish. Pubertal activation of testicular steroid production was associated with the appearance of primary spermatocytes, but a single case of a premeiotic endocrine activation was detected. The onset of puberty occurred during September and October, when adults were post‐spawned or in early recrudescence. Once initiated, pubertal development proceeded in an uninterrupted progression to full maturity by June at 22 months of age. Spermatogenetic activity during the autumn led to a peak in the gonado‐somatic index and total testicular mass by December. Milt could be first expressed from some pubertal males at this time, well in advance of the May to August spawning period in captivity. Meiotic activity could be absent from the testes by the spring, but residual areas of meiotic activity were found in pubertal males of one year class which may indicate a slower spermatogenetic rate. Androgen levels peaked during prespawning and early spawning; 11‐ketotestosterone (11‐KT) was the dominant androgen throughout spermatogenesis. Recrudescing 2 year‐old males showed greater testicular development and androgen production. Spermatogenetic patterns were similar to those seen in pubertal males, with the distinction that recrudescing males had a more discernable mitotic phase prior to meiotic activity; mitotic and meiotic activity overlapped considerably during early puberty. Testicular development well in advance of spawning may confer an evolutionary advantage to males of this cold‐water marine flatfish.     

A study of the population biology of Athevina boyeri Risso, 1810 in Aberthaw Lagoon, on the Bristol Channel, in South Wales

The biology of the sixth population of A. boyeri Risso, 1810 to be recorded outside of the Mediterranean basin is described. The 2 year life cycle was similar to that of other populations of the species that have been investigated. A maximum standard length of 93 mm was recorded for 2-year-old fish. Sexual maturity was attained by females at a length of 39 mm. A temporal lag in ovarian development was noted between the two spawning age groups, with the older age group spawning first. Details are provided of batch spawning in A. boyeriand the difficulties of assessing fecundity in batch spawners is discussed. Discrepancies between direct and indirect methods of assessing batch size indicated that the eggs around the largest mode may not represent the next batch to be spawned. Evidence is presented to suggest that in A. boyeri these eggs represent a ' store' from which batches of eggs are subsequently released.     

The role of life history in the relationship between population dynamics and environmental variability in two Mediterranean stream fishes

Chub Squalius torgalensis and nase Chondrostoma lusitanicum , in a Mediterranean stream, showed important differences in life-history traits and population dynamics. Both species reached mean maturity at age 2 years. Chub lived up to age 5 years, spawned in March to June, grew at a maximum rate of 0·59 mm mm⁻¹ year⁻¹ and showed a low reproductive allocation, with fecundity and egg size increasing with body size. Nase lived up to age 4 years, spawned in January to April, grew at a maximum rate of 0·46 mm mm⁻¹ year⁻¹ and showed a high reproductive allocation, with egg size independent of body size. Both chub and nase showed moderate fluctuations in population size during 1991–1998, but differed in factors driving density at age. Density of age 1 year juvenile chub decreased following severe summer droughts and proportionate survival prevailed thereafter. Density of age 2 year adult nase decreased following severe spring floods, but neither environmental nor parental stock effects were detected for juveniles and older fishes. The results illustrated the interplay between life history and environmental variability in driving fish population dynamics, with impacts of both summer droughts and spring floods being contingent on species-specific patterns of spawning and reproductive investment.     

Food Preference of Perch (Perca fluviatilis) Stocked in a Previously Fishless,Acid Bog Lake and Development of the Fish Population in Absence of Piscivory

The almost fishless, acid Lake Fuchskuhle was physically partitioned into 4 sections and stocked with young, 10‐ to 13‐cm perch. Their body growth was normal only up to age 4 years. Although the perch spawned successfully, reproduction was negligible because only few offspring survived the first year of life. Food for young perch was plentiful, benthic invertebrates composing the main source. Because the population included only 2 to 3 year classes, the perch could not shift from benthic feeding to piscivory as they grew. As a result, growth rate decreased in older fishes (over 4–5 years old). The benthic food supply alone could not satisfy the energy requirements of the older perch.     

Age, growth and reproduction of the cyprinid Rutilus lemmingii (Steindachner, 1866) in the River Huebra, Central Spain

Age, growth and reproduction of the R. lemmingii population of the River Huebra, Duero basin are analysed. Females dominated older age classes and lived up to 6 years (5+) while males only reached 5 years (4+). The growing season extended from April to September; growth rates were similar for both sexes. 0+ fish condition increased during their first summer and spring. In older fish, condition cycle was related to gonad development and showed some differences between sexes. The number of females reaching maturity at age 1 + almost doubled that of males; females also matured at smaller size. Both fecundity and egg size increased with female length: mean egg counts varied between 974 for 1 + individuals and an estimated 10491 for 5+ fish. Eggs were produced as a single batch, but were released fractionally during April and May.     

Life-history traits of the endangered river blenny Salaria fluviatilis (Asso) and their implications for conservation

The life-history characteristics of the river blenny Salaria fluviatilis in a Mediterranean stream in north-east Spain have been studied. The maximum age observed was 4+ years in both sexes, although only age groups 0+ and 1+ years were well represented. For both sexes, growth was rapid between April and June (before the summer drought) and in October. The instantaneous rate of monthly growth (April to October) was positively correlated with water depth. All fish attained sexual maturity during their first year of life. Females spawned multiple times. Spawning occurred mainly from June to August, although older females (2+ years old) began in May. The relationship between fecundity and total length recorded for older females in the spring was not significantly different from that obtained for 1 year-old females in the summer. The mean diameter of ripe oocytes, however, was greater in older females. Larger eggs that were laid during a period of good food supply (spring) might have a higher probability of survival. The results suggest that female fecundity is affected by environmental conditions. The species is characterized by fast growth, early maturity, multiple spawning and limited longevity. This may be a good life-history strategy for small fish species in unstable environments, where adult mortality is high, variable or unpredictable. Some life-history traits of the river blenny suggest that this species might be vulnerable to more severe drought conditions than those associated with the Mediterranean climate.     

Monthly indices of the post-smolt growth of Atlantic salmon from the Drammen River, Norway

Indices of the monthly post-smolt growth of three maturity groups of Atlantic salmon Salmo salar from the Drammen River in Norway were examined. Scale samples were collected from 1983 to 2003, providing a retrospective growth history for 2854 individuals. Overall post-smolt growth of all three maturity groups decreased throughout the study period. When the post-smolt year was segmented into 8 monthly indices, there was a significant decline in growth during the fourth and fifth month, typically months of maximum growth. The pre-fishery abundance of one and two sea-winter fish from Norway also showed a decline throughout the study period and was positively correlated to the post-smolt growth of fish returning to the Drammen River. Freshwater growth was negatively correlated to the pre-fishery abundance, thus indicating that factors affecting post-smolt growth during the fourth and fifth months at sea may be responsible for declining return numbers in Norway.     

Breeding biology of a non-diadromous galaxiid, Galaxias vulgaris Stokell, in a New Zealand river

In the Glentui River, Canterbury, New Zealand, Galaxias vulgaris Stokell spawned in late winter and early spring (from July to September), the onset of spawning being temperature-dependent. No females matured in their first year, but 95 and 68 % of age 0+ males were mature in consecutive years. With the exception of one female, all fish aged 1 + and older spawned. In mature fish, overies reached nearly 23%, and testes 11%, of somatic weight. Maturity in males depended on size, whereas in females, age appeared to be the major factor affecting maturity. Recruitment eggs for next season's spawning were distinct from primary oocytes soon after the ripe eggs were shed. Egg numbers varied between 284 and 1911 per fish, and weight was the best indicator of fecundity. Sex ratios of whole samples were not significantly different from 1 : 1, but ratios of potential spawners revealed an excess of males, due mainly to the presence of age 0 + fish. Age0+ males probably play an important role in adverse conditions and act as a form of insurance by significantly increasing the proportion of ripe males, thereby ensuring that all eggs are fertilized. The nest, egg mass and behaviour during the spawning period are described. More than one female may lay eggs in each nest and it is probable that each egg mass receives sperm from more than one male. It is suggested that the presence of fewer and larger eggs in G. vulgaris , compared with the diadromous galaxiids, is the result of a more precise fertilization process.     

Sea growth, smolt age and age at sexual maturation in Atlantic salmon

Relationships between growth at sea, smolt size and age at sexual maturation of Atlantic salmon Salmo salar were tested. The fish were offspring of brood stocks sampled in eight Norwegian rivers at latitudes between 59° and 70° N, hatchery reared and released at smolting at the mouth of the River Imsa (59° N). Smolt size influenced the subsequent growth rate of Atlantic salmon. The larger the fish were at release, the slower the yearly length increment at sea. Mean sea age at sexual maturity, measured as proportion of the returning adults attaining sexual maturity at sea age 2 years, was significantly correlated with mean growth rate during the first year at sea and mean smolt size ( r ²= 0·74, P < 0·001). Fish attaining maturity at a relatively high sea age were more fast growing during their first year at sea than those maturing at a younger age. The results indicate that high sea age at sexual maturation is a population-specific characteristic and associated with high early growth rate at sea.     

Seasonal growth of wild Atlantic salmon juveniles and implications on age at smoltification

The seasonal growth trajectories of wild Atlantic salmon Salmo salar juveniles by age group within the Margaree River, Canada, are described. Circuli counts from scales were used to infer growth rates at different ages and these were used to predict the proportions of age 2‐ and 3‐year old smolts from different portions of the watershed. In the wild Atlantic salmon juveniles from the Margaree River, there was no bimodality in fork length frequencies and no 1 year old smolts were produced. Water temperature differences during the growing season were insufficient to explain the differences in growth rates and size at age among the sites sampled. There was a positive association between the growth rate in the first year and the subsequent age at smoltification. In the Margaree River, differences in tributary specific growth rates and size at age were expected to produce important differences in the relative ages at smoltification.     

Reproductive biology of the milk shark Rhizoprionodon acutus and the bigeye houndshark Iago omanensis in the coastal waters of Oman

Female Rhizoprionodon acutus were found to mature between 62 and 74 cm total length ( L _T) whereas males matured between 63 and 71 cm L _T. The L _T at 50% maturity was 64·3 cm for females and 64·7 cm for males. Litter size varied from one to six embryos, and was positively correlated with maternal L _T. Female embryos outnumbered males by a ratio of 2·3:1. The size at birth was c . 37 cm L _T. Full‐term embryos and post‐partum females were observed during all seasons although their occurrence was highest in spring. Spermatozoa were rarely recorded in the oviducal gland, indicating that this species does not store sperm. It was not possible to generate maturity curves for Iago omanensis but it was evident that females matured by the time they reached 35 cm and males were mature by 31 cm L _T. This species displayed a clearly defined reproductive cycle with parturition occurring primarily in spring, after a gestation period of c . 1 year. Maximal embryo size was 19 cm L _T while maximal litter size was 24 embryos. The oviducal gland appeared to act as a seminal receptacle and it appeared that females may utilize these stores by not mating every year.     

Countergradient variation and secondary sexual color: phenotypic convergence promotes genetic divergence in carotenoid use between sympatric anadromous and nonanadromous morphs of sockeye salmon (Oncorhynchus nerka)

Genetically distinct anadromous (sockeye) and nonanadromous (kokanee) morphs of the Pacific salmon, Oncorhynchus nerka, develop identical, brilliant red color at maturity during sympatric breeding in freshwater streams. The marine and lacustrine environments they occupy prior to maturity, however, appear to differ in the availability of dietary carotenoid pigments necessary to produce red coloration. We tested the hypothesis that kokanee, which occupy carotenoid-poor lakes, are more efficient at using the dietary pigments than are sockeye, which occupy the more productive North Pacific Ocean. In a 2-year controlled breeding study, flesh and skin color of mature and immature crosses fed a low-carotenoid diet were quantified with both a chromameter and by chemical extraction of carotenoid pigments. Results revealed striking countergradient variation in carotenoid use, with kokanee approximately three times more efficient at sequestering the pigments to the flesh musculature than similar age sockeye. This difference translated into virtually nonoverlapping differences between pure crosses in secondary sexual color at maturity, when the pigments are mobilized and transported to the skin. Kokanee crosses turned pinkish red over most of their body, whereas sockeye turned olive green. The olive green was similar to the breeding color of residuals in the wild, the progeny of anadromous sockeye that remain in fresh water and are believed to have given rise to kokanee on numerous independent occasions. Reciprocal hybrids were similar to each other and intermediate to the pure crosses, indicating additive genetic inheritance. Mate choice trials with sockeye males in the wild showed the ancestral morph strongly preferred red over green models. These results suggest a preference for red mates maintained in nonanadromous breeding populations drove the reevolution of the red phenotype in kokanee via more efficient use of dietary carotenoid pigments. This is a novel, yet hidden, mechanism by which sexual selection promotes the genetic differentiation of these sympatric populations.     

Fitness consequences of the timing of metamorphosis in a freshwater crustacean

Metamorphosis is a common life-cycle transition in organisms as diverse as amphibians, insects, fishes and crustaceans, and the timing of this transition often affects an individual's fitness. Here, we measured age and size at metamorphosis in laboratory-reared individuals of the freshwater copepod, Diaptomus leptopus , and then followed individuals over their entire life cycle to assess the fitness consequences of variation in age and size at metamorphosis. In 3 separate experiments, individuals were raised in different food conditions: low food (0.2 μg C/ml) switched to high food (0.7 μg C/ml), or high food switched to low food, at several different larval and juvenile stages. Control individuals were reared on high or low food concentrations over their entire life cycles. For each individual, we measured age and size at metamorphosis and age and size at maturity; for females, we also measured total lifetime egg production, longevity, and calculated a composite fitness measure, λ. Statistical analyses showed no significant effects of age or size at metamorphosis on these same traits measured at maturity, or on the fitness components we estimated. The first individuals to mature had the highest total egg production and individual fitness; differences in body size at maturation explained none of the variation observed in fitness components. Our results show that metamorphosis was uncoupled from maturity and from fitness components by growth and development achieved during the juvenile phase of the life cycle, and support the conclusion that fitness consequences of metamorphosis depend fundamentally on the organization of an organism's life cycle. They also suggest that body size plays a different life-history role in these organisms than is recognized in most poikilotherms, and suggest the hypothesis, based on laboratory experiments, that selection may act primarily on juvenile developmental rates in field populations.     

Relationships of reproductive traits and body size with attainment of sexual maturity and age in Blanding's turtles (Emydoidea blandingi)

To place associations among body size, age at maturity, age, and reproductive traits of a long-lived organism in the context of current life history models based on the concept of norms of reaction, we examined data from a mark-recapture study of Blanding's turtles (Emydoidea blandingi) in southeastern Michigan during 24 of the years between 1953 and 1988. Females matured between 14 and 20 years of age. Both the smallest and largest adult females in the population were reproducing for the first time in their lives. This result suggests that a combination of differences in juvenile growth rates and ages at maturity, and not indeterminate growth, are the primary cause of variation in body size among adults. Body size variation among individuals was not related to age at sexual maturity. Females that had slower growth rates as juveniles matured later at similar mean body size compared to those with more rapid growth that matured at an earlier age. As a result, a linear model of age at sexual maturity with growth rates of primiparous females between hatching and maturity was significant and negative (R² = 0.76). Frequency of reproduction of the largest and smallest females was not significantly different. Clutch size did not vary significantly with age among either primiparous or multiparous females. Clutch sizes of primiparous females and multiparous females were not significantly different. However, older females (>55 years minimum age) reproduced more frequently than did younger females (minimum age <36 y).     

The biology of the bigeye grenadier at South Georgia

The biology of the bigeye grenadier Macrourus holotrachys caught as by‐catch in the Patagonian toothfish Dissostichus eleginoides longline fishery conducted around South Georgia was investigated to improve data available for fisheries management. Age estimates suggest that M. holotrachys is a moderately slow growing species ( K  = 0·10), reaching ages of >30 years and attaining total lengths ( L _T) >80 cm ( L _∞ = 33). The size at which 50% of females had started to mature ( L _int50) for M. holotrachys was 21 cm pre‐anal length ( L _PA) and occurred at c . 9 years old. Estimates of natural mortality and Pauly's growth performance index were found to be low ( M  = 0·09 and Φ = 2·82 respectively). Gonad maturity stage was described from macroscopic and histological investigation. Mature ovaries had oocytes at all developmental stages with between 22 and 55% likely to be spawned each year. Absolute fecundity ranged from 22 000 to 260 000 eggs and was positively correlated with both pre‐anal length and mass. A highly skewed sex ratio of 32 : 1, females : males, was found for specimens caught by longlines but not for a small sample of shallower trawl‐caught specimens. It is suggested that females are far more susceptible to longline capture than males. Macrourus holotrachys is a bentho‐pelagic predator and scavenger that feeds on a wide range of fishes and invertebrates. The fish are long lived, slow‐growing species typical of deep‐water grenadiers; fisheries management strategies should reflect their probable susceptibility to overfishing.     

Gonadal maturity in vimba (Vimba vimba L.) raised in carp ponds

The aim of the study was to determine gonadal maturity in male and female vimba after induced spawning in the years 2001 throughout 2004, based on spawners derived from an anadromous population obtained from the Barycz River. For these 4 years, fry derived from each successive spawning were raised in carp ponds at the Experimental Hatchery Centre of the Polish Angling Association in Szczodre, Poland. In February 2005, gonads were collected from males and females and their maturity determined after histological preparations. Additionally, in females the diameters of oocytes undergoing trophoplasmic growth were determined and compared. Histological analysis showed that all females had gonads at maturity stage IV, except the youngest fish, in which maturity stage I was determined. It was also shown that the size of oocytes undergoing trophoplasmic growth was related to age: the older the females, the greater the proportion of large‐diameter oocytes. In addition, some 2‐year‐old females had a considerable proportion of oocytes undergoing trophoplasmic growth despite being at stage IV of ovarian maturity. Among males, 2‐ to 4‐year‐olds had gonads at maturity stage III, in which the spermatogenesis process was active. One‐year‐old males had testes at maturity stage II, with visible spermatogonia entering meiotic divisions. When keeping vimba under pond conditions, only 2 years are needed to obtain a mature spawner from a spawning school. The older the school, the higher the probability that a greater number of spawners will enter spawning.