Daily increments on the otoliths of larval and juvenile Coregonus spp., and their modification by environmental factors

The sagittal otoliths of pre- and posthatch embryos, larval, and juvenile coregonids (Coregonus spp.) were examined for growth increments. Under laboratory conditions, a check is formed on the day of hatching and subsequently one growth increment per day is deposited during at least 265 days. Under the experimental conditions of this study, the age of young coregonids can therefore be determined with high accuracy.In starving larvae, both increment width and ring contrast decrease during the first 10 days after hatching until daily increments are no longer recognizable. A change from one diet to another, alteration of the water temperature, or a short starvation period lead to the formation of characteristic ring patterns which appear on the otoliths within 1 to 3 days. These patterns are highly reproducible among all specimens of each treatment group and can therefore be used as intrinsic marks. They could be applied to hatchery-reared coregonids, thus providing a basis for judging the efficacy of stocking operations.     

Synchronized spawning ofanguilla japonica inferred from daily otolith increments in leptocephali

The exact timing ofAnguilla japonica spawning was determined from analyses of daily otolith increments in leptocephali collected near a spawning area west of the Mariana Islands on July 1–18, 1991. The birth dates of the 54 leptocephali examined (10.2 to 30.5 mm in total length) ranged from May 22 to June 24, 1991, the individuals clearly comprising two age groups, May-born fish (mode May 28) and June-born fish (mode June 21). The data showed thatA. japonica spawns intermittently during the spawning season, with fixed synchronized timing. Each group of leptocephali collected along three different north-south transects (131°, 134° and 137°E between 10° and 22°N) comprised both May-born and June-born fish. The latter were dominant along the easternmost and middle transects, whereas the May-born fish were more abundant along the westernmost transect. The modal ages of the June-born and May-born fish collected along 137°E on July 1–3 were 13 d and 35 d, respectively, while those of the two age groups collected along 134°E on July 17 and 18 were 28 d and 50 d, respectively. These data show that the interval between the sampling dates for the two transects (ca. 15 d) corresponded closely to the differences in modal ages of specimens from the two transects (15 d) for both the May- and June-born fish, and further, that the difference in modal age between the two age groups (22 d) was the same at both transects. A similar correspondence in total length was also observed between the two age groups at the above two transects. The findings clearly demonstrated parallel westward transport by the North Equatorial Current for both the May- and June-born eel leptocephali, which originated from a spawning area estimated as being between 141° and 143°E.     

Environmental effects of introducing whitefish,Coregonus lavaretus (L.), in Lake Ring

The impact of whitefish (Coregonus lavaretus (L.)) on the trophic structure of eutrophic lakes was studied in Lake Ring, a small eutrophic Danish lake (22.5 ha, mean depth 2.9 m) in which the natural fish fauna is dominated by pike (Esox lucius L.), perch (Perca fluviatilis L.), and eel (Anguilla anguilla (L.)), roach (Rutilus rutilus (L.)) and burbot (Lota lota (L.)) being the only other fish species present. A total of 10993 0+ whitefish were stocked in the lake from October 1989 to July 1990 and the structure of the fish, zooplankton and benthic invertebrate communities studied during the period 1989–1991. Stomach contents analysis revealed that the whitefish mainly ate Daphnia and copepods in 1990–1991, the proportion of copepods decreasing with increasing size of the fish and Daphnia being the overall most important food source. The density of Daphnia in the lake decreased from 72 ind. 1^-1 in 1989 to 9 ind. 1^-1 in 1991; concomitantly the large species Daphnia magna and D. pulex almost disappeared and the density of cyclopoid copepods increased from 72 to 101 ind. 1^-1, presumably because of improved food conditions, while that of calanoid copepods remained virtually unchanged. As a result chl-a increased from 19 to 47 µg 1^-1 and Secchi depth decreased from 2.4 m to 1.7 m, despite there being no change in total P and total N (0.6 mg P 1^-1 and 1.3 mg N 1^-1, respectively). Changes were also observed in the benthic invertebrates; Chaoborus, oligochaetes, and chironomids all decreased, whereas Pisidium increased. It is concluded that the stocking of whitefish in eutrophic lakes for commercial purposes may delay their recovery, or even lead to enhanced eutrophication.     

Effects of temperature and food supply on the growth of whitefish Coregonus lavaretus larvae in an oligotrophic peri‐alpine lake

The relative roles of temperature and food availability on the seasonal and daily growth of whitefish Coregonus lavaretus larvae were investigated in the oligotrophic peri‐alpine Lake Annecy, France. During the spring from 2004 to 2007, surface water temperature and density of potential zooplankton prey were monitored, and 2688 larvae were caught and measured. In addition, the daily growth of 130 larvae was estimated retrospectively by investigating the microstructure of their otoliths. Temperature played the predominant role in controlling both seasonal and daily growth of early larvae. In contrast, the abundance of Mesocyclops leuckarti and larval density was only slightly correlated to larval growth, suggesting no food limitation nor strong interindividual competition over the study period. Overall, these findings run counter to concerns about potential food limitation, but sound a warning about the potential impact of climate change on fish ecology and fisheries management.     

Post-stocking perturbations in a population of subarctic whitefish,Coregonus lavaretus (L.): effects on growth,condition and cestode infection

Lake Kilpisjärvi was stocked with a total number of 440 000 migratory and benthic whitefish fry during the period of 1959–1964. Thereafter, the Diphyllobothrium ditremum plerocercoid infection became a problem decreasing the feasibility and commercial exploitation of the whitefish stock. Since dense fish populations may cause stunting, all stockings in the study area were stopped in an attempt to enhance the fisheries. The aim of the present study was to explore the effect of the cessation of the stocking on the growth, condition, catches and D. ditremum infection of the whitefish population. A total of 1594 whitefish were sampled with benthic gill nets. Growth was estimated from length-at-age and mass–length relationship was estimated with a non-linear regression in 1982–1983, 1992–1993 and 1997. Parasite infection was studied in 1992–1993 and 1997. Growth and condition were poor in the beginning of the 1980s. In 1992–1993, the growth and condition of the fish improved as the numerical catch per unit effort decreased from 1982 to 1992. The mean catches of other fish species were low. There were no significant changes in whitefish growth or condition from 1992–1993 to 1997. From the 1970s to 1982–1983, during slow growth and poor condition of the fish, the median gillraker count was decreasing to 19 (min–max 14–26). In 1992–1993, the gillraker count had increased significantly to 23 (min–max 19–26). However, the improved growth and condition did not cause decrease in the D. ditremum burden. On the contrary, based on our own results and literature notes, the parasite abundance seems to increase during the observation period of 1964–1997.     

Ontogenetic changes in habitat use by whitefish,Coregonus lavaretus

Synopsis The whitefish, Coregonus lavaretus, in the lake Mj?sa exhibited two niche changes during their life cycle. Juveniles (< 25 cm body length) were confined to the shallow (0–30 m) epibenthic zone. Medium sized whitefish (25–35 cm body length) expanded their habitat use to include the deep (30–90 m) epibenthic zone as well as the pelagic zone. From a body length of 35 cm, habitat use was restricted to the deep epibenthic zone. Small fish in the shallow epibenthic zone ate small and medium-sized prey (zooplankton, insect larvae and surface insects). Medium-sized fish in this zone were in addition feeding on the larger amphipod, Pallasea quadrispinosa. In the pelagic zone, the diet of medium-sized whitefish was dominated by zooplankton, although some larger prey like surface insects and age-0 smelt, Osmerus eperlanus, were also eaten. In the deep epibenthic zone, the diet of both medium-sized and large (< 35 cm) whitefish consisted mainly of the large prey P. quadrispinosa.     

Validation of otolith daily increment formation for two temperate syngnathid fishes: the pipefishes Stigmatopora argus and Stigmatopora nigra

Otoliths were used for the first time to successfully validate the age of members of the family Syngnathidae: the spotted pipefish Stigmatopora argus and the wide-bodied pipefish Stigmatopora nigra. Otolith increments were deposited daily in (1) known-age juveniles ranging in age from 0 to 31 days and (2) adults that had been stained with alizarin complexone, and a hatch mark was found on all otoliths which represented day 0. Otolith increment validation will allow development of growth models for S. argus and S. nigra, essential to understanding and managing these exclusive seagrass species.     

Does water calcium content influence the distinctness of daily growth increments in the otoliths of larval whitefish (Coregonus lavaretus L.)?

In larval and juvenile whitefish ( Coregonus lavaretus L.) from Lake Constance, Germany, the otolith increments are deposited daily, whereas daily deposition could not be confirmed in larval whitefish from Lake Pyhäselkä, Finland. The calcium concentration in Lake Constance is high (around 1.3 m m ), while calcium deficiency is typical for Finnish lakes (around 0.15 m m ). Therefore, the hypothesis that the distinctness of daily otolith increments in whitefish is related to water calcium content was tested by rearing three groups of Lake Constance whitefish in water of 0.2, 1.3 and 4.7 m m Ca. The eggs were incubated in lake water (1.3 m m Ca), and the larvae were acclimated to the experimental calcium concentrations on the day of hatching. After 39 days of ad libitum- feeding with Artemia nauplii, the three groups did not differ significantly in total length, wet and dry weight, and otolith length and width. The daily increments were easily recognizable, and contrast between dark (D)- and light (L)-zones was the same in the fish of all test groups. For the experimental set-up of this study, and particularly the range of calcium concentrations tested, the hypothesis that water calcium content influences the distinctness of daily otolith increments was rejected.     

Preparing sagittae for examination of daily growth increments of young‐of‐the‐year fishes: a modification of the embed method

A modification (termed the slide‐glass‐embed‐method, SGEM) of the embed method for preparing fish sagittae is described. The SGEM is based on a very simple principle: a dome of mixed resin containing the embedded sagittae loses hardness after being heated and can be easily cut with dissecting scissors.     

Daily ration of juvenile Coregonus lavaretus (L.) fed on living zooplankton

Juvenile whitefish, Coregonus lavaretus (L.), wet weight 0.04 to 5.2 g, from Lake Constance were kept at 10, 12 and 16° C water temperature, respectively and fed with living zooplankton. The experimental duration lasted 72 to 120 h. Daily rations were defined as the amount of zooplankton remaining subtracted from the amount of zooplankton added after a 24 h interval. The mortality of the zooplankton was determined in parallel experiments without fish. Relative daily ration (zooplankton weight/fish weight) v. fish weight increased up to a fish dry weight of approximately 0.12 g and then decreased steadily. The maximum daily ration was about 270% of fish body wet weight (wet/wet) corresponding to 75% of body dry weight (dry/dry), respectively. In fishes of a dry weight higher than 0.12 g (wet weight 0.65 g) a significant difference in food intake was found between 12 and 16° C. The specific growth rate ranged from nearly 0 up to 33% per day. No correlation was found between daily ration and specific growth rate.     

Reactions in individual fish to strobe light. Field and aquarium experiments performed on whitefish (Coregonus lavaretus)

This study describes how individual whitefish Coregonus lavaretusreact to strobe light. Field experiments were performed in a net enclosure on fish tagged with ultrasonic transmitters. A strobe light array was switched on near the tagged fish. The fish moved away from the light and increased their swimming speed. Aquarium experiments under controlled conditions were carried out in rearing tanks at Saimaa Fisheries and Aquaculture Station in Finland. A strobe light was directed from the side of the basin just ahead of, directly at, and behind the fish at a close range. In the first two cases fish responded by a distinct turn and a change in swimming direction away from the light. The fish did not change its swimming direction when light was aimed from behind. It is concluded that strobe light may be used to prevent fish from swimming into a specific area. Implications for development of new fishing equipment and research concerning fishes in areas with water power stations is briefly discussed.     

The effect of dietary phosphorus deficiency on the immune responses of European whitefish (Coregonus lavaretus L.)

Low phosphorous (P) feeds in aquaculture are recommended to reduce eutrophication of water systems. However, the feed should be adequate for normal growth and intact immune defence. Influence of low dietary P supply on non-specific and specific immune defence of European whitefish (Coregonus lavaretus) was studied in two trials. For Trial 1, a semi-purified, low-P diet was formulated and supplied with 0, 2.5, 5.0, 7.5, or 10.0 g P kg(-1)to obtain a P gradient of 4.4-14.9 g kg(-1)diet. Diets were fed to four replicate groups of fingerling whitefish for 42 days in a flow-through, freshwater system maintained at 15 degrees C. Fish fed with the P-unsupplemented diet had significantly lower plasma immunoglobulin M (IgM) levels than fish fed with the P-fortified diets (means 2.53 vs. 3.19 mg ml(-1); P=0.047). Plasma lysozyme activity did not differ between fish fed with the P-unsupplemented diet and fish fed with the P-fortified diets (means 21.8 vs. 25.5 U ml(-1); P=0.107). For Trial 2, fish were acclimated for 49 days by feeding the lowest or highest dietary P contents, and thereafter immunised with a single intraperitoneal injection of a novel antigen, bovine gamma globulin (BGG). A clear antibody response against BGG was elicited but the antibody levels did not differ between fish fed low-P or high-P contents. Similar to Trial 1, plasma IgM was lower in fish fed low-P diets, and low-P diet did not influence plasma lysozyme activity. In both trials the growth of fish, having received low-P diet, was significantly lowered. It is concluded that P deficiency has only minor effects on the immune parameters in whitefish, and the practical aquafeed with P contents sufficient for normal growth does not compromise immune functions of this species.     

Life histories of the powan,Coregonus lavaretus (L.) (Salmonidae,Coregoninae) of Loch Lomond and Loch Eck

Four aspects of the life histories of the two populations of powan Coregonus lavaretus (L.) in Scotland are described: growth (Eck powan are shorter and with greater year to year variance than Lomond); sexual maturation (Eck powan mature younger, but at similar weight to Lomond); spawning (timing in Eck varies, but is consistent in Lomond); and recruitment/mortality (fecundity, sex ratios, and mortality also vary in the short term). Short term differences between the physiological ecology of the populations can be ascribed to the size and topography of the lochs. Long term differences are more difficult to account for, and are more important in that they may signal changes in sustainability. Conservation of powan must be considered in terms of their synecological relationships, not in isolation.     

Parallel evolution of ecomorphological traits in the European whitefish Coregonus lavaretus (L.) species complex during postglacial times

The extensive phenotypic polymorphism in the European whitefish has triggered evolutionary research in order to disentangle mechanisms underlying diversification. To illuminate the ecological distinctiveness in polymorphic whitefish, and evaluate taxonomic designations, we studied nine Norwegian lakes in three watercourses, which each harboured pairs of divergent whitefish morphs. We compared the morphology and life history of these morphs, documented the extent of genetic differentiation between them, and contrasted the niche use of sympatric morphs along both the habitat and resource axes. In all cases, sympatric morphs differed in the number of gill rakers, a highly heritable trait related to trophic utilization. Individual growth rate, age and size at maturity, diet and habitat use also differed between morphs within lakes, but were remarkably similar across lakes within the same morph. Microsatellite analyses confirmed for all but one pair that sympatric morphs were significantly genetically different, and that similar morphs from different lakes likely have a polyphyletic origin. These results are most compatible with the process of parallel evolution through recurrent postglacial divergence into pelagic and benthic niches in each of these lakes. We propose that sparsely and densely rakered whitefish sympatric pairs may be a likely case of ecological speciation, mediated in oligotrophic lakes with few trophic competitors.     

Effects of freshwater to seawater transfer on osmoregulation,acid-base balance and respiration in river migrating whitefish (Coregonus lavaretus)

Osmoregulation, acid-base balance and respiratory parameters were investigated in whitefish following transfer from freshwater to salt water. Whitefish acclimated successfully to 25 ppt brackish water but died after direct transfer to 32 ppt sea water. Transfer to brackish water induced rapid (<6 h) and permanent increases in plasma [Na⁺], [Cl^–], total [Ca] and [Mg]. The extracellular hyperosmolality effected a transient (<3 days) muscle tissue dehydration and red blood cell shrinkage. Exposure to brackish water decreased both the arterial O₂ tension and whole body O₂ uptake. The extracellular acid-base status changed from an initial respiratory acidosis at 1 h towards a pronounced metabolic acidosis at 48 h of brackish water exposure. Red cell pH_i decreased in parallel with extracellular pH_e, but the in vivo pH_i/pH_e was only 0.26, suggesting some selective protection of red cell pH_i. Plasma cortisol concentration and gill Na⁺, K⁺-ATPase activity increased after exposure to high ambient salinity, reflecting the induction of hypo-osmoregulatory mechanisms. The physiological changes in whitefish are discussed in relation to salinity-induced effects in other salmonid fishes.Abbreviations CO₂ solubility in plasma - water O₂ capacitance coefficient - BW brackish water - C _T total CO₂ content in plasma - FW fresh water - Hb hemoglobin - Hct hematocrit - M _b body mass of fish - MCHC mean cellular hemoglobin concentration - PCO₂ carbon dioxide tension - pH _e extracellular pH - pH _i intracellular pH - PO_{2 in} oxygen tension in water flowing in - PO_{2 out} oxygen tension in water flowing out - ppt parts per thousand - RBC red blood cell(s) - SW sea water - V _m water flows through chamber - OV ₂ ml O₂ consumed per kg per hour     

Effects of starvation on the formation of daily growth increments in the otoliths of milkfish, Chanos chanos (Forsskål), larvae

The effects of starvation on daily growth and increment formation in the otolith were examined using a double oxytetracycline-labelling method on larval milkfish, Chanos chanos (Forsskål), reared under different feeding regimes. The results indicated that the differences in body and otolith growth between the larvae fed once and three times a day were not significant, and that the otolith growth increment was deposited daily in both groups of fed larvae. In contrast, the starved larvae grew at a slower rate than fed larvae in body length and otolith dimensions, and the otolith growth increment in the starved larvae was not deposited on a daily basis. After undergoing starvation, the larvae were unable to recover their normal growth either in otolith increment deposition or in body and otolith growth even though they were fed. Therefore, the application of ageing techniques based on counting otolith growth increments seems to be inaccurate for starved larvae.     

Morphological differentiation and resource polymorphism in three sympatric whitefish Coregonus lavaretus(L.) forms in a subarctic lake

Three field‐identified whitefish Coregonus lavaretus forms in Lake Muddusjärvi, Finland, were compared in morphology, diet and prey size. First, these forms were studied with univariate and multivariate analysis to assess morphological divergence at a higher resolution level than in the field. Second, stomach contents were analysed to estimate diet‐overlap among forms. Finally, the relationship between prey size and morphology was examined. The whitefish were assigned to the initial field‐classification with 99·2% and 98·8% accuracy for morphologic and meristic traits, respectively. The small sparsely‐rakered form (SSR) had the shortest rakers and largest gillraker space, followed by the large sparsely‐rakered form (LSR) with intermediate gillraker length and gillraker space, while densely‐rakered whitefish (DR) had the longest rakers and smallest gillraker space. The two sparsely‐rakered whitefish forms (LSR and SSR), consumed mainly benthic macroinvertebrates, while densely‐rakered whitefish (DR), utilized pelagic food items. Average diet‐overlaps between whitefish forms were low in June‐September (Schoener's α = 0·02 − 0·23). Gillraker number and length were negatively correlated to prey length in the diet ( r  = −0·73, and r  = −0·60), while gillraker space was positively correlated with prey length ( r  = 0·81). The fact that these whitefish forms were morphologically and ecologically segregated, and that gillraker traits probably have a functional value in food selection, further suggests that natural selection has been important in structuring life‐history trajectories into divergent niche use.     

Morphological divergence and origin of sympatric populations of European whitefish (Coregonus lavaretus L.) in Lake Femund, Norway

Combining morphological and genetic analysis, we compared patterns of diversification within and between morphs among sympatric European whitefish (Coregonus lavaretus L.) populations in Lake Femund, Norway. Seven external populations, from potential colonization routes into Lake Femund were included. We found that deep-, shallow-, river- and bay spawning populations are distinct morphs in Lake Femund. Within morphs, populations range from being similar genetically (Fst=0-0.005) among deep-spawning populations to being highly differentiated (Fst=0.153) between bay-spawning populations. Between morphs, genetic differences ranged from a low (Fst=0.008-0.022) between deep- and shallow-spawning populations to high difference (Fst=0.125-0.143) between shallow- and bay-spawning populations. A higher proportion of molecular variance was seen among (3.9%) than within morphs (2.8%). The adaptive gene combinations behind the four morphs seem to have originated within the lake, although the lake could have been colonized from more than one source population.     

Validation of daily otolith increments in larval and juvenile Chinese sucker, <Emphasis Type="Italic">Myxocyprinus asiaticus</Emphasis>

Otolith development was observed and the formation of daily growth increments in otoliths of Chinese sucker, Myxocyprinus asiaticus, was validated by monitoring known-age larvae and juveniles in the laboratory from 2003 to 2005. Otolith shape changed with larval and juvenile development, and there was an exponential relationship until a body length of 16 mm or so, and a linear relationship after a body length of 16 mm between otolith size and fish size. The first increment was identified in larvae 1 day after hatching. The regressed equations between daily age (D) and increment number in otoliths (N) were N = −0.64 + 0.96D in lapillus, and N = −0.31 + 0.98D in sagitta. The slopes were not significantly different than 1.0. This demonstrated that otolith increments in this species were formed daily and can be used for daily age determination.     

Daily growth increments in the otoliths of the three-spined stickleback, Gasterosteus aculeatus L.

Daily increment formation was demonstrated in the sagittae of the three-spined stickleback, Gasterosieus aculeatus L., under both laboratory and field conditions. The onset of daily increment formation coincided with the day of hatching. The main sources of inaccuracy in age determination resulted from difficulties in resolving narrow increments in the sagittae of fish held at low temperatures and the increasing incidence of checks with age. The advantage of daily increments for investigating growth in this species was demonstrated from a comparison between increment-derived age-at-length data and length-at-capture data, since the latter significantly under-estimated early growth rate within a population.