School density affects the strength of collective avoidance responses in wild‐caught Atlantic herring Clupea harengus: a simulated predator encounter experiment

An experimental study in a semi‐controlled environment was conducted to examine whether school density in wild‐caught Atlantic herring Clupea harengus affects the strength of their collective escape behaviours. Using acoustics, the anti‐predator diving responses of C. harengus in two schools that differed in density were quantified by exposing them to a simulated threat. Due to logistical restrictions, the first fish was tested in a low‐density school condition (four trials; packing density = 1·5 fish m^?3; c. 6000 fish) followed by fish in a high‐density school condition (five trials; packing density = 16 fish m^?3; c. 60 000 fish). The C. harengus in a high‐density school exhibited stronger collective diving avoidance responses to the simulated predators than fish in the lower‐density school. The findings suggest that the density (and thus the internal organization) of a fish school affects the strength of collective anti‐predatory responses, and the extent to which information about predation risk is transferred through the C. harengus school. Therefore, the results challenge the common notion that information transfer within animal groups may not depend on group size and density.     

Co‐inheritance of sea age at maturity and iteroparity in the Atlantic salmon vgll3 genomic region

Co‐inheritance in life‐history traits may result in unpredictable evolutionary trajectories if not accounted for in life‐history models. Iteroparity (the reproductive strategy of reproducing more than once) in Atlantic salmon (Salmo salar) is a fitness trait with substantial variation within and among populations. In the Teno River in northern Europe, iteroparous individuals constitute an important component of many populations and have experienced a sharp increase in abundance in the last 20 years, partly overlapping with a general decrease in age structure. The physiological basis of iteroparity bears similarities to that of age at first maturity, another life‐history trait with substantial fitness effects in salmon. Sea age at maturity in Atlantic salmon is controlled by a major locus around the vgll3 gene, and we used this opportunity demonstrate that these two traits are co‐inherited around this genome region. The odds ratio of survival until second reproduction was up to 2.4 (1.8–3.5 90% CI) times higher for fish with the early‐maturing vgll3 genotype (EE) compared to fish with the late‐maturing genotype (LL). The L allele was dominant in individuals remaining only one year at sea before maturation, but the dominance was reversed, with the E allele being dominant in individuals maturing after two or more years at sea. Post hoc analysis indicated that iteroparous fish with the EE genotype had accelerated growth prior to first reproduction compared to first‐time spawners, across all age groups, whereas this effect was not detected in fish with the LL genotype. These results broaden the functional link around the vgll3 genome region and help us understand constraints in the evolution of life‐history variation in salmon. Our results further highlight the need to account for genetic correlations between fitness traits when predicting demographic changes in changing environments.     

Probabilistic maturation reaction norms assessed from mark–recaptures of wild fish in their natural habitat

Reaction norms are a valuable tool in evolutionary biology. Lately, the probabilistic maturation reaction norm approach, describing probabilities of maturing at combinations of age and body size, has been much applied for testing whether phenotypic changes in exploited populations of fish are mainly plastic or involving an evolutionary component. However, due to typical field data limitations, with imperfect knowledge about individual life histories, this demographic method still needs to be assessed. Using 13 years of direct mark–recapture observations on individual growth and maturation in an intensively sampled population of brown trout (Salmo trutta), we show that the probabilistic maturation reaction norm approach may perform well even if the assumption of equal survival of juvenile and maturing fish does not hold. Earlier studies have pointed out that growth effects may confound the interpretation of shifts in maturation reaction norms, because this method in its basic form deals with body size rather than growth. In our case, however, we found that juvenile body size, rather than annual growth, was more strongly associated with maturation. Viewed against earlier studies, our results also underscore the challenges of generalizing life‐history patterns among species and populations.     

Factors affecting estimates of size at age and growth in grey triggerfish Balistes capriscus from the northern Gulf of Mexico

Growth zones in dorsal spines of grey triggerfish Balistes capriscus from the northern Gulf of Mexico were utilized to estimate growth and examine factors that may affect estimates of size at age. Age was estimated from dorsal‐spine sections of 4687 individuals sampled from U.S. waters during 2003–2013, including both fishery‐independent (n = 1312) and fishery‐dependent (n = 3375) samples. Ninety‐six per cent (n = 4498) of these sections were deemed suitable for ageing; average per cent error between two independent readers was 10·8%. Fork length (L_F) ranged from 65 to 697 mm and age estimates from 0 to 14 years. Both sex and sample source (fishery‐independent v. recreational) significantly affected estimated size at age for 2–6 year‐old fish. Data were pooled between sources to fit sex‐specific von Bertalanffy growth functions. Results for the female model were L_∞ = 387 mm L_F, k = 0·52 year^?1 and t₀ = 0·01 year, while for males L_∞ = 405 mm L_F, k = 0·55 year^?1 and t₀ = 0·02 year. These results were significantly different between sexes and indicate clear sexual dimorphism. Thus, growth should be modelled separately by sex when examining population parameters or conducting stock assessment modelling. The positive bias in estimates of size at age computed for recreational v. fishery‐independent samples also has clear implications for stock assessment as growth functions computed with fishery‐dependent samples would tend to overestimate stock productivity.     

Validation of scale‐age determination in European grayling Thymallus thymallus using tag‐recapture analysis

To validate age determination from scales in European grayling Thymallus thymallus, the scale‐read age of fish was compared with the true age obtained by tag‐recapture analysis. A total of 3997 individuals were tagged with visible implant tags and passive integrated transponder (PIT) tags in the River Wylye, south‐west England during 1999–2007. Annual repeat surveys were undertaken and collected scales read without prior knowledge of tag‐recapture age. Accuracy of fish ageing by scales was highest in 1 and 2 year‐old fish but decreased in older fish. In later life stages (>4 years old), underestimation of age occurred and the error in reading scales rose to 51·9% in 5 year‐old fish. Age assigned from scales underestimated the tag‐recapture assigned age by as much as 3 years. This study suggests that use of scales is an appropriate method to age a short‐lived population of T. thymallus inhabiting productive lotic systems. The underestimation of age in older fish, however, needs to be considered in the management of fish stocks because it may lead to undesirable exploitation of population.     

Effect of spatial differences in growth on distribution of seasonally co‐occurring herring Clupea harengus stocks

The mechanisms most likely to determine the distribution of the two major herring Clupea harengus stocks in their common early summer feeding ground in the eastern North Sea, Skagerrak and Kattegat were investigated through analysis of acoustic survey data from six consecutive years. No change was detected in biomass of North Sea autumn spawning C. harengus (NSAS) over time, whereas the biomass of western Baltic spring spawning C. harengus (WBSS) declined severely. Analyses of centre of abundance by stock showed no change in NSAS distribution, whereas the WBSS changed to a more western distribution over time. Contrary to previous perception of the juvenile migration, NSAS were found to leave the study area at the age between 1 and 2 years and WBSS 1 year olds were encountered in the Skagerrak. The estimated parameters of von Bertalanffy growth equations showed marked differences between areas with fish in the eastern part of the area having the lowest size at age at all ages. Further, their growth conditions appeared to deteriorate progressively over the period studied. Both NSAS and WBSS showed the highest condition in the North Sea and Skagerrak while condition was substantially lower in age Kattegat. The westward movement of spring spawners over time suggests that growth rate and possibly density of conspecifics influence the migration pattern and distribution of C. harengus in the area. In contrast, there was no evidence to suggest that distribution was constant over time within stocks or that distribution reflected size‐dependent limitations on migration distance.     

Fish length exclusively determines sexual maturation in the European whitefish Coregonus lavaretus species complex

The probability that a fish matures at a certain age and length (the so‐called probabilistic maturation reaction norm, PMRN) was analysed for a European whitefish Coregonus lavaretus species complex population living in the Austrian pre‐alpine Lake Irrsee. Fish length was found to be the only relevant determinant of maturation probability, and females matured at slightly smaller sizes than males.     

Age, growth, and sexual maturity of the yellownose skate Dipturus chilensis in the south‐eastern Pacific

The age and growth parameters of Dipturus chilensis were estimated by counting growth rings from thin sections of vertebral centra from 400 fish (246 females and 154 males), ranging from 23 to 124 cm total length (L_T), and backcalculating fish lengths at previous ages. Marginal increment analysis lent support to the hypothesis of annual deposition of band‐pairs, which formed during the winter months. The oldest female D. chilensis aged in this study was 21 years and 117 cm L_T, whereas the oldest male was 18 years and 93 cm L_T. A 4·7% index of average per cent error (I_APE) suggested that this is a precise method for calculating the age of D. chilensis. Observed L_T were lower than backcalculated L_T, which implies the influence of Lee's phenomenon. The von Bertalanffy growth equations, based on mean length‐at‐age data, were estimated as L_t = 128·3 (1 ? e^{?0·112 (t + 0·514)}) for females and L_t = 107·8 (1 ? e^{?0·134 (t + 0·862)}) for males where t is age (years). Growth was significantly different between sexes: females reached a larger adult size. Ages and lengths at 50% maturity were estimated at 14 years of age and 106 cm L_T for females and 11 years of age and 86 cm L_T for males. At c. 14 years, there was a decline in growth rates in females. The factor most likely responsible for this was sexual maturity, which caused a discontinuity in growth of female fish. These results show that this species is slow‐growing, long‐lived, relatively large and of delayed maturity, characteristics that make it vulnerable to exploitation.     

Comparison of fin ray sampling methods on white sturgeon Acipenser transmontanus growth and swimming performance

Effects of two fin‐ray sampling methods on swimming performance, growth and survival were evaluated for hatchery‐reared sub‐adult white sturgeon Acipenser transmontanus. Fish were subjected to either a notch removal treatment in which a small section was removed from an anterior marginal pectoral‐fin ray, or a full removal treatment in which an entire marginal pectoral‐fin ray was removed. Control fish did not have fin rays removed, but they were subjected to a sham operation. A modified 3230 l Brett‐type swim tunnel was used to evaluate 10 min critical station‐holding speeds (S_CSH) of A. transmontanus, immediately after the fin ray biopsies were obtained with each method. Survival and growth were evaluated over a 6 month period for a separate group of fish subjected to the same biopsy methods. Mean ± s.e . 10 min S_CSH were 108·0 ± 2·3, 110·0 ± 2·6 and 115·0 ± 3·5 cm s^?1 for the notch removal group, full removal group and control group, respectively, and were not significantly different among treatments. Behavioural characteristics including tail‐beat frequency and time spent hunkering were also not significantly different among treatment groups swimming at the same speeds. There were no mortalities and relative growth was similar among treatment groups. Average biopsy time for the notch removal method was lower and the wounds appeared to heal more quickly compared with the full removal method.     

Changes in growth and maturation parameters of Pacific sardine Sardinops sagax collected off California during a period of stock recovery from 1994 to 2010

Whether fluctuation in density influenced the growth and maturation variables of three aggregated cohorts (fish born during the 1986–1993, 1996–2003 and 2004–2008 periods) of Pacific sardine Sardinops sagax caeruleus collected off the Californian coast from 2004 to 2010 was investigated. Using a von Bertalanffy mixed‐effects model with aggregated cohorts as covariates, estimated growth rate significantly covaried with aggregated cohorts. Growth rate (K) was modelled as a fixed effect and estimated to be 0·264 ± 0·015 (±s.e ). Statistical contrasts among aggregated cohorts showed that the 1996–2003 cohorts had a significantly lower growth rate than the other two aggregated cohorts. The theoretical age at length zero (t₀) and the standard length at infinity (L_S∞) were modelled as random effects, and were estimated to be ?2·885 ± 0·259 (±s.e ) and 273·13 ± 6·533 mm (±s.e ). The relation of ovary‐free mass at length was significantly different among the three aggregated cohorts, with the allometric coefficient estimated to be 2·850 ± 0·013 (±s.e ) for the S. sagax population. The age‐at‐length trajectory of S. sagax born between 1986 and 2008 showed strong density dependence effects on somatic growth rates. In contrast to the density‐dependent nature of growth, the probability to be mature at‐size or at‐age was not significantly affected by aggregated cohort density. The size and the age‐at‐50% maturity were estimated to be 150·92 mm and 0·56 years, respectively. Stock migration, natural fluctuations in biomass and removal of older and larger S. sagax by fishing might have been interplaying factors controlling growth parameters during 1986–2010.     

Fecundity regulation by atresia in turbot Scophthalmus maximus in the Baltic Sea

Down‐regulation of fecundity through oocyte resorption was assessed in Baltic Sea turbot Scophthalmus maximus at three locations in the period from late vitellogenesis in April to spawning during June to July. The mean ± s.d . total length of the sampled fish was 32·7 ± 3·1 cm and mean ± s.d . age was 6·2 ± 1·5 years. Measurements of atresia were performed using the ‘profile method’ with the intensity of atresia adjusted according to the ‘dissector method’ (10·6% adjustment; coefficient of determination was 0·675 between methods). Both prevalence (portion of fish with atresia) and intensity (calculated as the average proportion of atretic cells in fish displaying atresia) of atresia were low in prespawning fish, but high from onset of spawning throughout the spawning period. Atretic oocytes categorized as in early alpha and in late alpha state occurred irrespective of maturity stage from late prespawning individuals up to late spawning fish, showing that oocytes may become atretic throughout the spawning period. Observed prevalence of atresia throughout the spawning period was almost 40% with an intensity of c. 20%. This indicates extensive down‐regulation, i.e. considerably lower realized (number of eggs spawned) v. potential fecundity (number of developing oocytes), suggesting significant variability in reproductive potential. The extent of fecundity regulation in relation to fish condition (Fulton's condition factor) is discussed, suggesting an association between levels of atresia and fish condition.     

Gonad development and size‐at‐maturity of silver therapon Leiopotherapon plumbeus (Kner 1864; Teleostei:Terapontidae) in tropical volcanic lakes in south Luzon,Philippines

Gonad development of the silver therapon Leiopotherapon plumbeus in two volcanic crater  lake  habitats (Sampaloc Lake, Taal Lake) in south Luzon, Philippines was examined during the annual reproductive cycle. The minimum body size‐at‐maturity of fish in these two lake habitats was also compared. Four gonad development stages were characterized as basis for the classification of ovarian (immature, maturing, mature, spawned) and testicular maturation (immature, maturing, mature) phases. The occurrence of all development stages in individual gonads suggest an asynchronous development whereby advanced stages are recruited continuously from a pool of younger stage germ cells to result in elevated female and male GSI throughout the annual cycle due to active gonadogenesis. Together with the increasing occurrence of advanced stage oocytes and spermatozoa from March until October, the elevated GSI of fish may indicate peak gonadal growth during the onset of the dry season (December–January) for eventual spawning from the beginning (May–June) until the end of the wet season (October–November). In both lake habitats, male fish were smaller than females but, regardless of sex, the minimum size‐at‐maturity of fish in Sampaloc Lake was significantly smaller than fish in Taal Lake. Overall, asynchronous development during oogenesis and spermatogenesis allows for year‐round reproduction of silver therapon, with elevated gonad growth in the dry season in preparation for spawning during the wet season. Compared with fish in Taal Lake, a smaller size‐at‐maturity of fish in Sampaloc Lake may be a response of the wild fishery stock to long‐term high fishing mortality and degradation of the lake habitat.     

Reproductive strategy and recruitment dynamics of amphidromous goby Sicyopterus japonicus as revealed by otolith microstructure

The daily ages of 312 of 879 newly recruited postlarvae of Sicyopterus japonicus, collected from the Shuang‐Chi Estuary in north‐eastern Taiwan during February 1996 to April 1997, were determined from daily growth increments in their otoliths. Pelagic larval duration, growth rate of the marine larval stage and hatching dates were estimated, and recruitment timing was linked to environmental factors. The mean ±s .d . total length (L_T) and daily ages of S. japonicus at recruitment to the estuary were estimated to be 33·95 ± 1·31 mm (range 30·7 to 38·1, n = 317) and 163·72 ± 12·79 days (range 130 to 198, n = 312), respectively. The recruitment of S. japonicus larvae is size dependent not age dependent because L_T of the larvae is independent of age at recruitment. Periodic analysis indicated that L_T and growth rate of the larvae were inversely correlated with the age at recruitment, which means that the fast‐growing individuals recruited earlier. The growth rate of S. japonicus in the marine larval stage was synchronous with marine productivity in this subtropical area, i.e. the spring cohort recruited in the autumn had a higher growth rate than the autumn cohort recruited the following spring. The main spawning season of S. japonicus as backcalculated from otolith daily increments was in autumn, a relatively low productivity period compared with spring. During this season, there were fewer competitors and predators than in the more productive spring. The recruitment of 95% of postlarva coincided with low salinity (14) and low water temperature (23° C) in the river mouth that provided a buffer area for the adaptation of the larvae for upstream migration. This unique reproduction strategy and prolonged larval duration facilitated the widespread distribution of the fish along the coasts of East Asia.     

Shifting dynamic forces in fish stock fluctuations triggered by age truncation?

Accumulating evidence shows that environmental fluctuations and exploitation jointly affect marine fish populations, and understanding their interaction is a key issue for fisheries ecology. In particular, it has been proposed that age truncation induced by fisheries exploitation may increase the population's sensitivity to climate. In this study, we use unique long‐term abundance data for the Northeast Arctic stock of cod (Gadus morhua) and the Norwegian Spring‐Spawning stock of herring (Clupea harengus), which we analyze using techniques based on age‐structured population matrices. After identifying time periods with different age distributions in the spawning stock, we use linear models to quantify the relative effect of exploitation and temperature on the population growth rates. For the two populations, age truncation was found to be associated with an increasing importance of temperature and a relatively decreasing importance of exploitation, while the population growth rate became increasingly sensitive to recruitment variations. The results suggested that the removal of older age classes reduced the buffering capacity of the population, thereby making the population growth rate more dependent on recruitment than adult survival and increasing the effect of environmental fluctuations. Age structure appeared as a key characteristic that can affect the response of fish stocks to climate variations and its consequences may be of key importance for conservation and management.     

Growth and mortality of larval Myctophum affine (Myctophidae,Teleostei)

The growth and mortality rates of Myctophum affine larvae were analysed based on samples collected during the austral summer and winter of 2002 from south‐eastern Brazilian waters. The larvae ranged in size from 2·75 to 14·00 mm standard length (L_S). Daily increment counts from 82 sagittal otoliths showed that the age of M. affine ranged from 2 to 28 days. Three models were applied to estimate the growth rate: linear regression, exponential model and Laird–Gompertz model. The exponential model best fitted the data, and L₀ values from exponential and Laird–Gompertz models were close to the smallest larva reported in the literature (c. 2·5 mm L_S). The average growth rate (0·33 mm day^?1) was intermediate among lanternfishes. The mortality rate (12%) during the larval period was below average compared with other marine fish species but similar to some epipelagic fishes that occur in the area.     

Female Baltic herring Clupea harengus allocate resources from growth to reproduction in poor feeding conditions

The trade‐off between somatic growth and reproduction in the female Baltic herring Clupea harengus was investigated from 1984 to 2002. During the study period, growth decreased, as a consequence of decreasing salinity and weakening of feeding conditions. Production of muscle and ovarian tissue decreased in repeat spawners, but investment in reproduction took an increasing amount of the total production of new tissues. This suggested that a shift in allocation to reproduction takes precedence over body growth in the reproductive strategy of C. harengus. The process also indicated one possible mechanism leading to dwarf forms in fish populations.     

Climate effects on size‐at‐age: growth in warming waters compensates for earlier maturity in an exploited marine fish

Over the past 3 decades, North Sea Atlantic cod (Gadus morhua) have exhibited variable length‐at‐age along with declines in spawning stock biomass and timing of maturity. Multiple factors affecting growth and development in fish acted on this economically important stock over the same period including warming waters and an intensive fishery. Here, we employ North Sea cod as a model population, exploring how a physiologically relevant temperature metric (the growing degree‐day, GDD; °C day) can be used to compare year‐classes on a physiologically relevant time‐scale, disentangling influences of climate (thermal history) on observed length‐at‐age trends. We conclude that the trends in North Sea cod length‐at‐age observed during the last three decades can be explained by a combination of temperature‐dependent growth increases and a trend toward earlier maturation, the latter likely induced by the intensive fishing pressure, and possibly evidence of fisheries‐induced evolution.     

Experimental assessment of the probabilistic maturation reaction norm: condition matters

The probabilistic maturation reaction norm (PMRN) describes an individual''s probability of maturing at a given age as a function of size and other relevant phenotypic traits. Population-level shifts in the PMRN are often interpreted to indicate genetic as opposed to phenotypic changes in maturation in fish. Inferences derived from trends in the PMRN have been challenged, warranting an experimental assessment of the method. This was accomplished in a laboratory experiment using zebrafish (Danio rerio). Fish were reared under different food levels to induce variation in growth and maturation. Plasticity in maturation was not entirely captured by the demographic age- and length-based PMRN. Adding condition to the PMRN captured a greater amount of environmental variation in maturation probability. Nevertheless, significant differences in the PMRNs among the food levels remained after accounting for the influences of age, size and condition on maturation probability indicating plasticity of the PMRN. This was particularly pronounced between fish held on low food levels as compared with fish experiencing abundant resources, with the latter experiencing higher size-specific maturation probabilities. Our analysis emphasizes the need for incorporating salient physiological traits influencing maturation, such as condition, to make accurate inferences about documented shifts observed in the position of PMRNs on maturation trends in wild fish stocks.     

Variation in growth,mortality, length and age compositions of harvested populations of the herbivorous fish Girella tricuspidata

Commercial gillnet and beach‐seine catches of Girella tricuspidata from seven estuaries in eastern Australia were examined for differences in fork length (L_F), sex and age composition, and populations were assessed for growth and mortality. Fish 220–350 mm L_F dominated landings across all estuaries sampled, regardless of gear type. Few fish >10 years of age were observed in the catches, with fish aged 3–5 years, and 4–7 years, being most abundant in the catches in the four most northern estuaries and three southern estuaries, respectively. There was considerable variation in the L_F of G. tricuspidata at any given age and the oldest male and female were 21 and 24 years, respectively. There were no consistent differences between sexes or latitudinal regions in the growth and mean L_F at age of fish in each individual age class between 3 and 8 years. Growth of females was greater than males in the northern region, but not elsewhere. Estimates of the instantaneous rate of total mortality (Z) were dependent on estuary and year, ranging from 0·30 to 1·01, whereas the corresponding estimates of fishing mortality (F) ranged from 0·12 to 0·90. Populations of G. tricuspidata appear to have been heavily exploited, primarily relying on young fish recruiting to the fishery. The ecosystem‐wide effects of harvesting this dominant mobile teleost herbivore need to be assessed further.     

Effects of the sea louse Lepeophtheirus salmonis on temporal changes in cortisol,sex steroids,growth and reproductive investment in Arctic charr Salvelinus alpinus

Groups of mature (5+ year old) Arctic charr Salvelinus alpinus held in sea water were exposed for 34 days to either a high (mean ±s.e . 0·15 ± 0·01 sea lice Lepeophtheirus salmonis g^?1 fish mass) (HI), medium (0·07 ± 0·00 sea lice g^?1 fish mass) (MI) or no [control (C)] sea‐lice infection during early stages of gonad development (June to July). Infection with sea lice resulted in increased plasma cortisol concentrations and this was related to intensity of infection; females tended to have higher cortisol concentrations than males at high infection intensities (HI group: female c. 130 ng ml^?1; male c. 80 ng ml^?1). Plasma osmolality (C c. 330, MI c. 350 and HI c. 415 mOsm) and chloride concentrations (C c. 135, MI c. 155 and HI c. 190 mM) increased significantly with infection intensity, indicating osmoregulatory problems in infected fish. A strong positive relationship between plasma osmolality and cortisol concentration was recorded. Plasma sex‐steroid concentrations were influenced negatively by sea‐lice infection, particularly in the HI group, and were inversely related to plasma cortisol concentrations. The most heavily infected fish postponed the initiation of reproductive development until exposed to fresh water and timing of ovulation tended to be delayed in these fish. Growth rate and condition were negatively influenced by sea‐lice infection and growth rate was inversely related to plasma cortisol concentrations. Sea‐lice infection resulted in mortality among females in the HI group, and the proportion of maturing females was lower in the MI group (46%) than in the controls (85%). Egg production in the MI and HI groups was c. 50 and 30% of the C group. Egg size, embryonic survival and fry mass did not differ across groups. Sea lice influence reproductive development and egg production in S. alpinus, and consequently these parasites may influence populations via sublethal effects on broodfish, affecting growth and condition, and their reproductive output.