Methods for investigating patterns of mortality and quantifying cause-specific mortality in sea-farmed Atlantic salmon Salmo salar

Methods for investigating patterns of mortality and quantifying cause-specific mortality in Atlantic salmon Salmo salar farming were developed. The methods were further used to investigate mortality and patterns of mortality for the first 3 mo after sea transfer in the 2006 year-class autumn smolts (SO) of Norwegian farmed Atlantic salmon. In the study population, which consisted of 20 pens at 10 sites, cause-specific mortality was examined by 11 fish health professionals during 8 visits to each pen. Cause-specific mortality proportions were used to convert crude mortality into cause-specific mortality. Cumulative mortality in the study period was 2.1% in the study population compared with 3.7% for the 2006 year-class SOs in the national database. Of this cumulative mortality, 73 and 59% took place in 20% of the pens in the study and the reference population, respectively. Daily mortality rates in the study population showed a variation from 0 to 2376 per 100,000 fish where the majority of mortality was observed during disease outbreaks. All study pens had periods of low baseline mortality and some pens had no increased mortality during the study period. Of 2088 dead fish examined, 92% (1929 fish) were assigned a specific cause of death, and in 97% of these 1929 fish the investigators reported the given cause of death to be likely or very likely. Ulcers were the main cause of death, accounting for 43% of the assigned mortality, and infectious agents were involved in 64% of the total mortality. The study shows that probable causes of death can be established in Atlantic salmon farming and their contribution to total mortality measured.     

A fish-eye view of riverine hydropower systems: the current understanding of the biological response to turbine passage

One-way connectivity maintained by fish passing through hydropower turbines in fragmented rivers can be important to population dynamics, but can also introduce a new and significant source of mortality. Sources of mortality during turbine passage can come from several sources including blade strike, shear forces, cavitation, or pressure decreases and parsing the contributions of these individual forces is important for advancing and deploying turbines that minimize these impacts to fishes. We used a national hydropower database and conducted a systematic review of the literature to accomplish three goals: (1) report on the spatial distribution of turbine types and generation capacities in the USA, (2) determine fish mortality rates among turbine types and fish species and (3) examine relationships between physical forces similar to those encountered during fish turbine passage and fish injury and mortality. We found that while Francis turbines generate 56 % of all US hydropower and have the highest associated fish mortality of any turbine type, these turbines are proportionally understudied compared to less-common and less injury-associated Kaplan turbines, particularly in the Pacific Northwest. While juvenile salmonid species in actual or simulated Kaplan turbine conditions were the most commonly studied, the highest mortality rates were reported from percid fishes passing through Francis turbines. Future studies should focus on understanding which species are most at-risk to turbine passage injury and mortality and, subsequently, increasing the diversity of taxonomy and turbine types in evaluations of turbine injury and mortality.     

Size-related mortality due to gnathiid isopod micropredation correlates with settlement size in coral reef fishes

The transition between the planktonic and the benthic habitat is a critical period for the larvae of many demersal marine organisms. Understanding the potential constraints on the timing of this habitat transition, called settlement, is important to understanding their biology. Size-specific mortality can set the limits on lifestyle and help explain ontogenetic habitat shifts. We examined whether size-based mortality risks after settlement may include micropredation by ectoparasites by testing whether survival of settlement-stage fish varies with fish size when exposed to a reef-associated micropredator. Fish (14 species) were exposed to one blood-sucking gnathiid isopod overnight, with appropriate controls; gnathiid feeding success and survival, and fish mortality were recorded relative to fish size. After adjusting for fish relatedness, we found the relationship between fish mortality and size differed with gnathiid exposure: for gnathiid-exposed fish, the mean mortality of the smallest fish was much higher (57%) than unexposed controls (10%), and decreased to ~0% for fish >12 mm standard length (SL); mortality was almost nil in controls. Thus, a predicted optimal size to switch habitat and reduce mortality risk from micropredation should be >12 mm SL. We then asked what species might be at greater risk and if the steep increase in survival at ~12 mm SL might coincide with settlement at larger sizes among fishes. Across 102 other species (32 families), 61% settled at ≥12 mm SL. After adjusting for relatedness, mean fish settlement size was 15.0 mm and this was not significantly different from 12 mm. Thus, settlement size clusters around the minimum fish size threshold our gnathiid experiment predicted would be large enough to survive a gnathiid encounter. These results suggest micropredators may contribute to size-selective mortality during settlement processes and are consistent with the hypothesis that the pelagic phase provides fish an escape from certain micropredators.     

Estimates of daily mortality from a neascus trematode in age-0 shortnose sucker (Chasmistes brevirostris) and the potential impact of avian predation

Parasites may be an important component of early life mortality in fishes, but assigning part of total mortality to parasites is difficult. The Chapman-Robson mortality estimator is a robust and potentially valuable way to quantify the added mortality of parasites when age data are available. We used daily age data and the Chapman-Robson catch-curve procedure to estimate daily mortality for 15 years in juvenile age-0 shortnose suckers (Chasmistes brevirostris), and for 6 years, the daily mortality of fish with and without black spot infection, a trematode whose final host is a piscivorous bird. Infected fish always had higher mortality rates than uninfected fish, and for 3 years when those differences were significant, the added daily mortality for infected fish was 3.6–3.7 %. Based on the proportion infected each year, and for durations of 15–50 d, juvenile populations were 18.3–38.6 % lower than they would have been without black spot infections. There were no significant differences in growth between infected and uninfected fish in most years and little indication of a direct metabolic impact of infections. Thus, this added mortality, primarily in July and August, seemed unlikely to be an indirect result of infection and was most likely due to predation. The source of that predation is unknown but the parasite’s final hosts, piscivorous birds, seemed the most obvious candidate for this added mortality.     

Many Vulnerable or a Few Resilient Specimens? Finding the Optimal for Reintroduction/Restocking Programs

Most reintroduction and restocking programs consist of releasing captive-raised juveniles. The usefulness of these programs has been questioned, and therefore, quality control is advisable. However, evaluating restocking effectiveness is challenging because mortality estimation is required. Most methods for estimating mortality are based on tag recovery. In the case of fish, juveniles are tagged before release, and fishermen typically recover tags when fish are captured. The statistical models currently available for analyzing these data assume either constant mortality rates, fixed tag non-reporting rates, or both. Here, instead, we proposed a method that considers the mortality rate variability as a function of age/size of the released juveniles. Furthermore, the proposed method can disentangle natural from fishing mortality, analyzing the temporal distribution of the captures reported by fishermen from multiple release events. This method is demonstrated with a restocking program of a top-predator marine fish, the meagre (Argyrosomus regius), in the Balearic Islands. The estimated natural mortality just after release was very high for young fish (m ₀ = 0.126 day^-1 for fish 180 days old), but it was close to zero for large/old fish. These large/old fish were more resilient to wild conditions, although a long time was needed to achieve a relevant reduction in natural mortality. Conversely, these large/old fish were more vulnerable to fishing, creating a trade-off in survival. The release age that maximizes the number of survivors after, for example, one year at liberty was estimated to be 1,173 days. However, the production cost of relatively old fish is high, and only a few fish can be produced and released within a realistic budget. Therefore, in the case of the meagre, increasing the number of released fish will have no or scarce effects on restocking success. Conversely, it is advisable implement measures to reduce the high natural mortality of young juveniles and/or the length of time needed to improve fish resilience.     

Age- and weight-dependent susceptibility of rainbow trout Oncorhynchus mykiss to isolates of infectious haematopoietic necrosis virus (IHNV) of varying virulence

The virulence of 5 European and 1 North American isolate of infectious haematopoietic necrosis virus (IHNV) was compared by infecting female sibling rainbow trout ('Isle of Man' strain) of different weights and ages (2, 20 and 50 g). The fish were exposed to 10(4) TCID50 IHNV per ml of water by immersion, and the mortality was recorded for 28 d. Two new IHNV isolates from Germany were included in the investigation. One was isolated from European eels kept at 23 degrees C (+/- 2 degrees C) and the other was not detectable by immunofluorescence with commercially available monoclonal antibodies recognising the viral G protein. The results showed that IHNV isolates of high or low virulence persisted in rainbow trout of all ages/weights for 28 d, with the exception of fish over 15 g in the eel IHNV (DF [diagnostic fish] 13/98)-infected groups from which the virus could not be reisolated on Day 28. The smallest fish were most susceptible to an infection with any of the IHNV isolates. The lowest cumulative mortality (18%) was observed in fingerlings infected with the North American isolate HAG (obtained from Hagerman Valley), and the highest mortality (100%) in DF 04/99 infected fish. The DF 04/99 and O-13/95 viruses caused mortality in fish independent of their weight or age. The isolates FR-32/87 and I-4008 were virulent in fish up to a weight of 20 g and caused no mortality in larger fish. In the IHNV HAG- and DF 13/98 (eel)-infected rainbow trout, no signs of disease were observed in fish weighing between 15 and 50 g. An age/weight related susceptibility of rainbow trout was demonstrated under the defined conditions for all IHNV isolates tested.     

Effects of entrapment and cooling water discharge by the Bergum Power Station on 0+ fish in the Bergumermeer

The numbers, distribution and size of 0+ fish in Bergumermeer and in the cooling water system of the Bergum Power Station were registered from 1978 to 1981. An exponentially decreased vulnerability to entrapment by the power station was found with increasing size of fish. Because of the large water intake in relation to the lake volume, entrainment mortality of fish larvae in May and June is large and probably of the same order of magnitude as the natural mortality. In the same period the Bergumermeer can be supplied with large amounts of fish larvae, immigrating passively with the water current. Impingement mortality later on in the season is insignificant compared with natural mortality. The heated effluents from the power station influence the distribution and the size of 0+ fish in the lake. Year-class strength of pikeperch may be improved due to the larger size of 0+ pikeperch. Less water intake as a possibility to minimize entrainment is discussed.     

A 27-year study of brown trout population dynamics and exploitation in Lake Songsjøen, central Norway

From 1968–1984 (period I), a brown trout Salmo trutta , population in a 70-ha oligotrophic lake in central Norway was exploited using larger mesh gill-nets selectively removing the larger fish. From 1985–1994 (period II), intermediate sized fish were removed using smaller-mesh sizes gill-nets. Fishing mortality and CPUE were correlated positively with effort and numbers of fish >3 years old for period II. The gill-net catchability was correlated negatively with spawner biomass and number of trout >3 years old. The significant positive correlation between natural mortality and stock biomass and spawning stock biomass indicated density-dependent mortality. The significant correlation between spawning stock and recruitment described by the Ricker model, indicated density-dependent recruitment of 1-year-old trout. The fishing regimes in the two periods affected the population dynamics and density differently. Selective removal of smaller fish permitted the larger fish to survive, and was beneficial in reducing fish density and maintaining stocks at low levels, consequently, achieving the expected increase in fish growth rates.     

Injury and short term mortality of benthic stream fishes – a comparison of collection techniques

Small benthic fish such as darters are frequently collected for stream inventory purposes or to document habitat use, with the intent of releasing the fish unharmed following enumeration. The purpose of this study was to examine the injury and short term mortality (8 d) of greenside darters captured by live wire pot trapping and electrofishing, using two different settings (80Hz, 6ms and 60Hz, 6ms). Two different electrofishing techniques were used, spot electrofishing and sweep electrofishing. Short term mortality was highest for fish collected in live pot traps. Abrasion from the wire traps appeared to remove scales and irritate the skin. By the conclusion of the study, 74% of the fish caught in live pot traps were dead from fungal lesions. Greenside darters captured by all electrofishing methods exhibited low short term mortality (< 10%). The only initial mortality, hemorrhaging and spinal damage, occurred for fish collected using 80Hz, 6ms sweep technique, although the short term mortality was still far less than that observed among trapped fish. The spot electrofishing technique resulted in no injury, with either of the settings. Live trapping produces little initial mortality, and thus may be wrongly viewed as a safe alternative for the collection of threatened benthic stream fishes, compared to electrofishing. We suggest that researchers studying small fish in warmwater systems use caution when collecting and handling fish for subsequent release. This revised version was published online in July 2006 with corrections to the Cover Date.     

Reactivation of koi herpesvirus infections in common carp Cyprinus carpio

Two co-habitation studies with common carp were conducted to determine whether latent infections of koi herpesvirus (KHV) exist. Fish were exposed to KHV using 2 different temperature profiles, which induced low and high initial mortality. Subsequently, certain groups of fish were co-habited with naive fish while others were not. Koi herpesvirus was reactivated in fish from 3 of the 5 experimental tanks. Reactivation of the virus occurred regardless of the initial mortality associated with the virus or whether fish were co-habited with naive fish. The reactivation of the virus in our experiments occurred several months after the initial exposure to KHV and appeared to be temperature dependent.     

Stress and survival of small pike-perch Sander lucioperca (L.) after trawling and chilling

The mortality and stress responses (plasma cortisol concentration) of undersized (total length, L _T, <370 mm) pike-perch Sander lucioperca caught as by-catch in a trawl fishery were assessed. The effects of three different holding methods on pike-perch caught by trawl were compared: (1) fish that were transferred to ambient-temperature water (15·0–21·4° C) immediately after capture and not exposed to chilling water, (2) fish held in chilling water for 10 min and (3) fish exposed to chilling water for 2 h. The sample fish were held and monitored in individual plastic restrainers at ambient-water temperature for 48 h after each of the handling procedures. Total mortality was similar for fish held in chilling tanks for 10 min (27·2% total mortality) compared to fish that were only held in ambient-temperature water (28·2% mortality). Total mortality reached 91·3% when fish were held in chilled water for 2 h. Mortality and plasma cortisol concentrations correlated inversely with size (96–368 mm L _T) and directly with lake-water temperature (15·0–21·4° C) after treatment. In addition, the effect of chilling was observed in elevated cortisol concentration and delayed start of recovery. The results show that quickly removing (<10 min) undersized pike-perch from chilled water could markedly improve the survival of released fish. Due to delayed recovery, however, fish should be allowed to recover (30–40 min) in a deck tank before release.     

Relative contribution of exotoxin and micropredation to icthyotoxicity of two strains of Pfiesteria shumwayae (Dinophyceae)

The mechanism by which Pfiesteria shumwayae (Glasgow and Burkholder) kills fish is controversial. Several studies have implicated a Pfiesteria-associated exotoxin in fish mortality while other studies indicate that physical attack of dinoflagellates on fish (micropredation) and not exotoxin is responsible. We examined the ichthyotoxicity of two strains of P. shumwayae (CAAE 101272 and CCMP 2089) in a bioassay system that exposed test fish to the dinoflagellates both with and without direct contact in the same aquarium at the same time. Dinoflagellate-free supernatants from both strains were also tested for toxicity. The results showed that direct contact between P. shumwayae and fish significantly enhanced fish mortality with both strains (P < 0.001). About 87.5% and 100% of fish died when exposed directly to CAAE 101272 and CCMP 2089, respectively. When protected from direct contact with Pfiesteria cells, 19% of the fish exposed to CAAE 101272 and 6% of those exposed to CCMP 2089 died. No deaths were observed in controls. Supernatant killed fish when obtained from cultures of CAAE 101272 but not when obtained from CCMP 2089.Analysis of variance showed that, for both strains, fish mortality in Pfiesteria-inoculated bioassays was significantly higher than control bioassays both with and without direct contact (P < 0.001). Differences between strains were not significant (P = 0.3). These results indicate that both strains are associated with exotoxin production. However, the dominant and most consistent mechanism of fish mortality observed in this study required physical contact between fish and Pfiesteria cells.     

Responses of succinate dehydrogenase and non-specific alkaline phosphatases and mortality of tilapia to ambient pH stress in a sewage-fed aquaculture pond

The fish, tilapia (Oreochromis mossambicus) of 50-60 g body weight was experimentally exposed to effluent gradients of highly alkaline pH in a sewage-fed aquaculture farm for examining the pH stress-induced responses of mortality and the stress marker enzyme succinate dehydrogenase and the non-specific alkaline phosphatases of fish prior to death at different hours of intoxication. A second trial was performed after two months when water quality changed along the sewage effluent gradient. An in situ experiment was also performed for better understanding of the responses of enzymatic activities attributable to different levels of pH conditions. Time required for 100% mortality of fish tended to increase from 30 min in pH 11.6 to 22 hr in pH 10.2. There was no mortality of fish when water quality improved significantly (with pH ranging between 9.6 to 8.0) after two months. The activities of succinate dehydrogenase and intracellular alkaline phosphatases assayed in gills and liver prior to death of fish tended to reduce with increase in survival hour, following a pattern of decay curve. On the other hand, percent of enzymatic inhibition of the exposed fish over the control increased as the survival hour increased following a pattern of exponential curve. It appears that the highest water pH of 11.6. maximum ratio for ammonium to ammonium hydroxide (1: 21) and reduced level of dissolved oxygen (2.62 mg/l) were perhaps responsible for the 100% mortality of fish within 30 min of their exposure and the enzymatic activities in the gills and liver assayed prior to death of fish tended to reduce as the acclimatization period of fish increased and vice-versa.     

Two classes of deleterious recessive alleles in a natural population of zebrafish, Danio rerio

Natural populations carry deleterious recessive alleles which cause inbreeding depression. We compared mortality and growth of inbred and outbred zebrafish, Danio rerio, between 6 and 48 days of age. Grandparents of the studied fish were caught in the wild. Inbred fish were generated by brother-sister mating. Mortality was 9% in outbred fish, and 42% in inbred fish, which implies at least 3.6 lethal equivalents of deleterious recessive alleles per zygote. There was no significant inbreeding depression in the growth, perhaps because the surviving inbred fish lived under less crowded conditions. In contrast to alleles that cause embryonic and early larval mortality in the same population, alleles responsible for late larval and early juvenile mortality did not result in any gross morphological abnormalities. Thus, deleterious recessive alleles that segregate in a wild zebrafish population belong to two sharply distinct classes: early-acting, morphologically overt, unconditional lethals; and later-acting, morphologically cryptic, and presumably milder alleles.     

The importance of shading by riparian vegetation and wetlands in fish survival in stagnant water holes,Great Ruaha River,Tanzania

Mass fish mortality occurred in November 2006 in a water hole in the Great Ruaha River, Ruaha National Park, and was due to the lack of shade by riparian vegetation or fringing wetlands. Nearby water holes with shade or fringing wetlands suffered no fish mortality. Despite being vulnerable to intense birds’ predation, fish stayed in the top few centimeters due to hypoxia in deeper water.     

Theoretical consideration of effect of fishing mortality on growth and reproduction of fish populations

An increase in fish mortality due to fishing can theoretically change the growth and reproduction of fish populations from the viewpoint of adaptation. We address the issue of how an iteroparous fish should convert surplus energy into somatic growth and reproduction at each age under given conditions of mortality. A model of life history, which maximizes the net reproductive rate using the discrete maximum principle, is improved employing a new relationship between body weight and surplus energy which we have recently proposed. The model is applied to the North Sea plaice Pleuronectes platessa, for which it has been reported that the average length of young fish had increased whereas that of old ones had decreased for some decades. Although the model cannot directly explain the former phenomenon, the two phenomena can be interpreted as a change in the optimal life history due mainly to an increase in mortality.     

Regulation of local populations of a coral reef fish via joint effects of density- and number-dependent mortality

Density-dependent mortality can regulate local populations - effectively minimizing the likelihood of local extinctions and unchecked population growth. It is considered particularly important for many marine reef organisms with demographically open populations that lack potential regulatory mechanisms tied to local reproduction. While density-dependent mortality has been documented frequently for reef fishes, few studies have explored how the strength of density-dependence varies with density, or how density-dependence may be modified by numerical effects (i.e., number-dependent mortality). Both issues can have profound effects on spatial patterns of abundance and the regulation of local populations. I address these issues through empirical studies in Moorea, French Polynesia, of the six bar wrasse (Thalassoma hardwicke), a reef fish that settles to isolated patch reefs. Per capita mortality rates of newly settled wrasse increased as a function of density and were well approximated by the Beverton-Holt function for both naturally formed and experimentally generated juvenile cohorts. Average instantaneous mortality rates were a decelerating function of initial densities, indicating the per capita strength of density-dependence decreased with density. Results of a factorial manipulation of density and group size indicate that per capita mortality rates were simultaneously density- and number-dependent; fish at higher densities and/or in groups had higher probabilities of disappearing from patch reefs compared with fish that were solitary and/or at lower densities. Mortality rates were ~30% higher for fish at densities of 0.5 fish/m² than at 0.25 fish/m². Similarly, mortality rates increased by ~45% when group size was increased from 1 to 2 individuals per patch, even when density was kept constant. These observations suggest that the number of interacting individuals, independent of patch size (i.e., density-independent effects) can contribute to regulation of local populations. Overall, this work highlights a greater need to consider numerical effects in addition to density effects when exploring sources of population regulation.     

Isolation of the North American strain of viral hemorrhagic septicemia virus (VHSV) associated with epizootic mortality in two new host species of Alaskan marine fish

Thousands of dead Pacific herring Clupea pallasi, Pacific hake Merluccius productus and walleye pollock Theragra chalcogramma were reported in Lisianski Inlet near Pelican, Alaska, USA, on August 1, 1998. The Pacific hake and pollock continued to die through the end of September. Virological examinations of dead fish identified the North American strain of viral hemorrhagic septicemia virus (VHSV) from all 3 species of fish as well as associated high virus titers and possible histopathological lesions. No other primary fish pathogens were detected and there were no apparent environmental causes for fish mortality. This is the first report of VHSV in 2 new Alaskan fish host species and of a natural epizootic associated with VHSV in which progressive mass mortality was observed simultaneously in herring and 2 other species of free-ranging marine fish.     

Overwinter body condition,mortality and parasite infection in two size classes of 0+ year juvenile European bitterling Rhodeus amarus

Body condition and parasite abundance were examined in two size classes of European bitterling Rhodeus amarus during the first overwintering period in two seasons (2007–2008 and 2009–2010). Body condition of large fish did not change during winter, and increased significantly in March. From November to February, small fish showed a decreasing trend in condition. Despite a significant increase in March condition of small fish only reached the same level as before winter. Total parasite abundance increased significantly in winter in both fish size classes, reflecting a seasonal increase in monogenean infection. Large fish were parasitized significantly more than small fish during winter, but only in small fish was a negative correlation between parasite infection and condition found and a significant decrease in parasite abundance recorded after wintering, indicating mortality of heavily infected individuals with low condition during the winter. A trend for higher overwinter mortality in small fish was found under semi‐experimental conditions. The decrease in condition during the winter period in small fish may reflect faster energy depletion generally expected in smaller individuals. The results indicate that parasite infection may contribute to the overwinter mortality of 0+ year R. amarus, with a stronger effect in smaller individuals.     

Effect of epidermal papillomatosis on survival of the freshwater fish Rutilus rutilus

Epidermal papillomatosis occurs in several marine and freshwater fish species. Previously, papillomatosis has been shown to induce mortality in juvenile carp. We studied the effect of epidermal papillomatosis on the survival of adult male roach Rutilus rutilus by caging naturally diseased, marked (by us) fish in the field. Within the constraints of the experimental design, there was no difference in survival between healthy, slightly diseased and heavily diseased fish. Therefore, we conclude that the possible effect of epidermal papillomatosis on the mortality of wild roach is relatively minor.