The influence of food competition and host specificity on the transmission of Triaenophorus crassus (Cestoda) and Cystidicola farionis (Nematoda) to Coregonus lavaretus and Coregonus albula (Pisces:Coregonidae) in Finland

As a superior competitor for planktonic food, vendace (Coregonus albula), when abundant, is expected to displace whitefish (Coregonus lavaretus) from feeding on plankton and to force it to rely more on benthic food. The predicted result would be a reduced abundance of the copepod-transmitted cestode Triaenophorus crassus in whitefish, but an increase in the abundance of the nematode Cystidicola farionis transmitted via benthic amphipods. We studied the occurrence of both parasites in whitefish during 1991-1996 in three interconnected areas at Lake Saimaa, Finland, where the densities of the vendace stocks varied due to natural fluctuation in year-class strengths. In accordance with our hypothesis, some indication of the effect of the density of the vendace population on abundance of C. farionis infection in whitefish was found, but not in the case of T. crassus. Only 0.2% of vendace were infected with T. crassus, while up to 100% of the whitefish in the yearly samples harboured the parasite. In further experiments we clarified which copepod species in Lake Saimaa act as first intermediate hosts of T. crassus, and verified from stomach samples the exposure of both whitefish and vendace to those species. Experimental infections indicated that the infectivity of T. crassus is lower for vendace than for whitefish. We suggest that the reason why vendace stock density does not affect T. crassus infection in whitefish is that T. crassus is transmitted in littoral areas during a short period in spring. At that time of the year copepods are abundant and available to both whitefish and vendace, but since the parasite is less infective to vendace. they do not become infected. At other times of the year, dense vendace stocks may force whitefish to shift to benthic food, which includes amphipods transmitting C. farionis. During vendace stock decline, whitefish may, however, continue to feed on plankton and avoid exposure to C. farionis.     

Selective predation on parasitized prey--a comparison between two helminth species with different life-history strategies.

In Lake Fjellfr?svatn, northern Norway, the larval helminths Cyathocephalus truncatus and Cystidicola farionis use Gammarus lacustris as intermediate hosts and Arctic charr (Salvelinus alpinus) as final hosts. There was sampled 1,433 live G. lacustris from the lake and 1,964 G. lacustris from stomach contents of the charr. Prevalence of infection were, respectively, 0.49% and 3.72% for C. truncatus, and 0.21% and 0.20% for C. farionis. Usually, only 1 parasite was present in each host, and the 2 parasite species never co-occurred. Gammarus lacustris amphipods parasitized by C. truncatus were positively selected by the Arctic charr and were consumed approximately 8 times as often as were the unparasitized amphipods or the amphipods infected with C. farionis. This suggests that G. lacustris amphipods infected with C. truncatus larvae are more susceptible to predation than noninfected specimens, probably because of parasite-induced alterations in behavior or visibility. Alternatively, this could also be explained by selection toward the largest G. lacustris specimens observed, which are also the most frequently parasitized amphipods. However, the data show clearly that this was not a result of size-selective predation by the charr. In contrast, the presence of C. farionis did not increase the susceptibility to predation of its intermediate host. The discrepancy between the 2 helminth species supports the hypothesis that parasite-increased susceptibility to predation is related to the life history strategies of the parasites.     

Parasites as indicators of individual feeding specialization in Arctic charr during winter in northern Norway

The relationship between infection with the food-transmitted parasites Diphyllobothrium dendriticum, D. ditremum (Cestoda) and Cystidicola farionis (Nematoda) and prey selection was studied in individual Arctic charr from Lake Takvatn, northern Norway. There was no correlation between parasites transmitted throughout prey organisms from benthic habitats (amphipods) and pelagic habitats (copepods). A strong relationship between infection with a parasite species and the corresponding intermediate host from the stomach content of individual charr, indicated an individual feeding specialization. Independent of size, charr specialized on the intermediate hosts of all three parasite species. Some charr maintained this specialization on specific prey items throughout the winter period. These parasite species are considered to be useful indicators of past prey selection.     

Development of skeleton elements in juvenile dwarf and small sympatric forms of <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> complex from Lake Davatchan (Transbaikalia)

The development of skeleton elements was studied in prolarvae, larvae, and fry of two morphologically different forms of the Arctic char Salvelinus alpinus complex—small plankton-eating and dwarf (having a seasonal change of plankton and benthic feeding)—that sympatrically inhabit Lake Davatchan (Transbaikalia). According to genetic data, these forms diverged in this lake from a specialized lacustrine ecotype of char and reached reproductive isolation. Comparative analysis revealed similarity in the shape of anlagen and transitive states of cranial bones. In prolarvae and larvae of the dwarf form, the onset of similar morphological states of skeleton elements occurs at a smaller body length than in fish of the small form. Differences in the development of serial skeleton structures (teeth, gill rakers, fin rays, centra, and scales), heterochronies of timing of appearance and rate of differentiation of skeleton elements were found. It is suggested that these differences resulted from the ecological differentiation of the two forms according to spawning grounds that became local habitats of larvae and fry (sublittoral and hypolimnion). The initial divergence could be caused by the presence in the lake of two (autumn and spring-summer) peaks of numbers of food organisms.     

Diet niche relationships among predator and prey fish species in their early life stages in Lake Võrtsjärv (Estonia)

In Lake Võrtsjärv pikeperch was observed not to shift to piscivory in their first autumn of life, although juvenile stages of a variety of fish species were abundant in the lake. It was hypothesized that the diets of predator and prey fish fry overlap and that coarse fish species are important food competitors for juvenile piscivores and thus, pikeperch and perch fry do not shift to piscivory during their first growing season. To discover the possible linkages in this pattern, in 2009 the feeding relationships of pikeperch, perch, ruffe and roach fry were analysed. The stomach content analyses showed that in the summer period, Mesocyclops leuckarti was the most frequent prey for perch and ruffe, pikeperch consumed Leptodora kindti in large quantities, and roach ate mostly plant material. Towards autumn, M. leuckarti was the most abundant prey for all percids. However, average stomach content weight and the number of prey items eaten by ruffe were considerably higher than for other fish fry. Since the feeding opportunities of fish fry are considered poor in the examined lake, the prey has the potential to restrict the recruitment to piscivory of their predators, as prey fish seem to have better abilities to persist in this ecosystem. Furthermore, supposed competition in the juvenile stage may result in a reduced top‐down effect on coarse fish.     

The influence of light environment on depth of visual feeding by larvae and fry of Coregonus pollan (Thompson) in Lough Neagh

The depth limits for visual feeding by pollan larvae and fry, Coregonus pollan , were determined in Lough Neagh, Northern Ireland, by combining laboratory estimates of the minimum light threshold for feeding activity with field measurements of spectral, diurnal and seasonal changes in light penetration. Although feeding thresholds were lowest in the blue light (400–490 nm), the rapid attenuation of this region in the lake meant that the depth limits of feeding in situ were determined by response to the green region (490–595 nm). Effects of changes in irradiance and light penetration on the depth of feeding are considered during a seasonal study. Increased sensitivity, as the juvenile stages develop, may be offset by reduced light penetration due to algal growth. Over an annual period the maximum feeding depth for pollan varied between 2.6 and 4.1 m. In the juvenile stages of two other predatory species from the lake, perch, Perca fluviatilis , and pike, Esox lucius , the corresponding ranges were 1.1–1.9 m and 2.7–4.6 m respectively.     

Seasonal and ontogenetic dynamics in trophic transmission of parasites

Transmission rates from the intermediate (amphipods) to the definitive hosts (fish) were quantified for two helminth species ( Cyathocephalus truncatus , Cestoda, and Cystidicola farionis , Nematoda) both seasonally and through the ontogeny of the final hosts (arctic charr, Salvelinus alpinus , and brown trout, Salmo trutta ). Amphipods ( Gammarus lacustris ) were important prey for both fish species, especially in the autumn. Both parasite species had low infection levels in amphipods compared to high abundance in fish. The seasonal variations in transmission rate of C. truncatus procercoids from amphipods to fish were in accordance with the observed abundance in fish hosts, being highest in the autumn and lowest during late winter and early summer. During summer, however, the estimated monthly transmission rates of C. truncatus were higher than the observed infection levels in the fish, suggesting restricted establishment success and shorter development time and longevity of the cestode in fish at higher temperatures. The accumulated transmission of C. farionis over the ontogeny of arctic charr was similar to the observed age-specific infection levels, reflecting a high establishment success and longevity of this parasite in charr. In contrast, brown trout exhibited an infection level that was much lower than the estimated transmission rates, suggesting a high resistance against C. farionis in these fish. The magnitudes of the estimated transmission rates were sufficient to explain the paradoxical contrast between low infection levels in the intermediate and high in the final hosts.     

Cystidicola farionis as a swimbladder parasite of the whitefish in the Bothnian Bay

Material comprising 3167 specimens of the parasite Cystidicola farionis was assembled from 1741 individuals of the sea-spawning whitefish, Coregonus nasus in the shallow northeastern and deeper central areas of the Bothnian Bay. Occasional parasites were found in the immature fish (≦ 175 mm), but infection increased in the two larger classes as the fish aged, reaching 27% in the northeastern area, and as much as 67% in the southern area. This difference is explained partly by the more frequent feeding on Pontoporeia affinis , the intermediate host for the parasite, the deeper waters extending nearer the coast in the majority of the central and southern Bothnian Bay. The increased intensity of infection with age does not vary significantly between the areas studied. No clear seasonal fluctuations in the prevalence or intensity of infection could be found.     

ECOLOGICAL SEPARATION BETWEEN WHITE AND PINK-BACKED PELICANS IN THE RUWENZOFU NATIONAL PARK, UGANDA

The number and distribution of the White Pelican Pelecanus onocrotalus roseus and the Pink-backed Pelican P. rufescens were studied in the Ruwenzori National Park, Uganda, by means of aerial surveys carried out at approximately weekly intervals between May 1968 and December 1969. The White Pelican is migratory and was present in the Park at highest numbers in August 1968 and May 1969 when 2510 and 3927 respectively were recorded. It does not breed within the Park. The Pink-backed Pelican is resident, with a population averaging about 1350 birds. Pelicans are distributed in discrete flocks on loafing grounds. Flocks of the White Pelican tend to be bigger than those of the Pink-backed. Forty-three loafing grounds were recognized in the Park, and it is shown that there is no difference between the species in the frequentation of these grounds. The areas favoured by both species are characterized by flat, open stretches close to water; steep slopes and heavily vegetated shore-lines are avoided. The feeding behaviour was studied by direct observation and through cine-photography. Only the White Pelican feeds communally in coordinated groups; this species also often feeds singly. The feeding sequence of both species is described, and is divided into four phases:searching, trapping, catching and swallowing. The searching phase lasts about six times longer in the Pink-backed Pelican than in the White. The White Pelican feeds further from the shore than does the Pink-backed. There is little difference between the species in the daily feeding pattern. Feeding takes place throughout the day but mostly from dawn to about 10.00 hrs and from 17.00 hrs until dusk. The rest of the day is spent mainly in preening, resting or bathing. The food of pelicans is exclusively fish. The White Pelican takes mainly large Tilupia, and Haplochromis and fish fry to a lesser extent. The Pink-backed Pelican feeds largely on fish fry, but Tilapia and Haplochromis are frequently taken and by weight are more important than the fry. However, the Tilapiu are smaller than those taken by the White Pelican. Ecological separation between the two species is achieved through these differences in their feeding behaviour. It is estimated that the White Pelican takes 1201 g and the Pink-backed Pelican 776 g of fish each day. The total amount of fish eaten by both species from Lake George during 1969 was calculated to be 591 709 kg. This is small (c. 3%) compared with the estimated total fish production of the lake but quite appreciable (12.7%) as a proportion of the total caught by man. It is concluded that the pelican populations are probably not important as a link in the chain whereby organic matter transferred from land to water by the Hippopotamus is returned to the land.     

Diel feeding rhythms and daily food consumption of juvenile Atlantic salmon in the River Alta, northern Norway

Atlantic salmon fry (0+) sampled from the River Alta exhibited only minor differences in stomach content weights and feeding rates throughout diel periods, but feeding rates were generally lowest at night. In contrast, salmon parr (1+ to 3+) had large diel fluctuations in stomach content weight, with the largest weights usually being recorded during the night and early morning. Accordingly, their feeding rates were highest at night. This nocturnal feeding pattern was consistent throughout all sampling occasions, and appeared to reflect a persistent feeding periodicity in the salmon parr. The daily food consumption rates of both fry and parr were highest during midsummer and decreased towards autumn.     

Temporal constraints on the potential role of fry odors as cues of past reproductive success for spawning lake trout

Deciding where to reproduce is a major challenge for most animals. Many select habitats based upon cues of successful reproduction by conspecifics, such as the presence of offspring from past reproductive events. For example, some fishes select spawning habitat following odors released by juveniles whose rearing habitat overlaps with spawning habitat. However, juveniles may emigrate before adults begin to search for spawning habitat; hence, the efficacy of juvenile cues could be constrained by degradation or dissipation rates. In lake trout (Salvelinus namaycush), odors deposited by the previous year's offspring have been hypothesized to guide adults to spawning reefs. However, in most extant populations, lake trout fry emigrate from spawning reefs during the spring and adults spawn during the fall. Therefore, we postulated that the role of fry odors in guiding habitat selection might be constrained by the time between fry emigration and adult spawning. Time course chemical, physiological, and behavioral assays indicated that the odors deposited by fry likely degrade or dissipate before adults select spawning habitats. Furthermore, fry feces did not attract wild lake trout to constructed spawning reefs in Lake Huron. Taken together, our results indicate fry odors are unlikely to act as cues for lake trout searching for spawning reefs in populations whose juveniles emigrate before the spawning season, and underscore the importance of environmental constraints on social cues.     

Feeding pattern and diet of first feeding brown trout fry under natural conditions

The feeding habits of brown trout Salmo trutta fry were studied during the critical first feeding period in a natural spawning and nursery stream. A low proportion of the fry initiated exogenous feeding before emergence from the gravel, and while having nearly 30% of the yolk sac remaining. This probably reflected low feeding motivation or limited feeding opportunities within the gravel environment. The majority of the fry started feeding after emergence, and after most or all yolk was absorbed. Some fry emerged with large amounts of yolk remaining, while others emerged after yolk exhaustion. The degree of stomach fullness revealed that feeding was more efficient after a territory had been acquired. The diets of the young fry were dominated by chironomid larvae, followed by zooplankton and Plecoptera larvae. Fry dispersing downstream and out of the nursery area were significantly smaller than resident fry, indicating displacement due to competition for territories. The majority of the downstream dispersing fry had initiated feeding, and there was at this point no evidence of starvation in any of the fry. It therefore appeared that the later emerging fry actively migrated out of the overpopulated nursery area to find available territories further downstream.     

Aspects of the biology of Tilapia mossambica Peters (Pisces: Cichlidae) in a natural freshwater lake (Lake Sibaya, South Africa)

An account is given of some aspects of the biology of Tilapia mossambica Peters in Lake Sibaya, South Africa. Lake Sibaya is a warm shallow coastal lake with extensive shallow terraces in the littoral which shelve abruptly into underwater valleys. The substrate is predominantly sandy. Aquatic macrophytes are generally restricted to water 1–7 m deep. Adult fishes (over 8 cm standard length) are usually absent from water deeper than 12m and shallower than 0–5 m, whereas juveniles may occur at all depths, and fry only in very shallow water.
T. mossambica inhabits the littoral and sublittoral in the warm and transition periods (August–April) but moves into deep water in the cool season (May–July). Exposed and sheltered areas are utilized for different purposes by adult fishes, the former for nesting, and latter for feeding and mouth–brooding. Habitat selection by males is governed by the availability of suitable nest sites. Nests are most common in sparsely–vegetated sheltered areas. Brooding females preferred sheltered littoral areas but ventured onto the terrace to release the fry. Juveniles and fry inhabited exposed shores with a temperature gradient which reversed diurnally. The breeding, shoaling and feeding behaviour is described. The biology of T. mossambica in Lake Sibaya is compared to data on the same, and similar species, in other systems. The utilization of available resources in the lake by T. mossambica is discussed and reference is made to the importance of the retention of generalized characters for the successful habitation of the cyclically–renewed habitat of the littoral.     

The ontogeny of feeding behaviour in four species of centrarchid fish

Relatively little work has been done on the ontogeny of feeding behaviour in fish. One of the aims of this study was to compare the ontogeny of feeding behaviour in four species of centrarchid fish of diverse feeding guilds. A second aim was to test Mayr's hypothesis that the locomotory acts involved in the ontogeny of feeding behaviour would be stereotyped. All four species initially displayed five feeding acts, with three of the acts disappearing from the repertoire at various intervals. The disappearance of the acts appeared to be related to the size and not to the age of the fry. The acts decreased in frequency as the fry aged and differences among the species in frequencies were minimal. The data supported Mayr's hypothesis but a modifying corollary was added to account for changes due to maturation.     

Host-constrained epidemiology of the fish tapeworm Ligula intestinalis (L.)

Investigations into the biology of the roach, Rutilus rutilus (L.), and Ligula intestinalis (L.) populations at Slapton Ley, Devon were carried out between October 1982 and December 1984, and additional data collected from the lake since 1977 have been re-analysed. The Ligula population exhibited some unusual features: a limited specificity, a persistently low abundance and a scarcity of multiple infections. The population dynamics of the roach were also unusual in that the individual growth rate was one of the fastest in Britain, the survival was very poor and the year classes tended to alternate in strength. Whether the unusual epidemiology of Ligula could be explained by the unusual population dynamics of the roach is investigated. The transmission period of Ligula to the roach was limited to their first few months of life, probably due to a limited period of feeding on copepods by the fry. This narrow transmission window was almost certainly the major factor that has prevented the abundance of Ligula plerocercoids attaining high levels in the fry or of increasing in subsequent years, and has resulted in their abundance being determined entirely by transmission events to the fry. The limited transmission period has also caused the life-cycles of the roach and Ligula populations to become synchronized. At other localities, the abundance of Ligula is typically high, and can have a significant effect on the mortality of the host population, but, with the low abundance at Slapton, Ligula-induced roach mortality was insignificant. The rapid growth of the roach fry was the most likely explanation for the limited period of feeding on copepods, and the large size, short life-span and low abundance of the roach probably also constrained the build-up of the Ligula population in the lake. The random frequency distribution and scarcity of multiple infections was not considered unusual in view of the very low plerocercoid abundance. The very low abundance of Ligula, the separation of the spawning of roach and rudd in time and space, and the scarcity of rudd may account for the absence of infection in this species. It was concluded that the Ligula population in Slapton Ley was being constrained by the atypical population dynamics of the roach, and not vice versa as theory predicts.     

Role of diet and experience in the development of feeding behaviour in largemouth bass, Micropterus salmoides

Two groups of largemouth bass, Micropterus salmoides , were reared in the laboratory. One group was reared on an artificial, passive diet (frozen brine shrimp) whereas the second was reared on a natural, active diet (cultured zooplankton). Observations on the development of feeding behaviour indicated that the motor patterns and duration (number of weeks in the behavioural repertoire) of the feeding acts did not differ between fry reared on the two diets. While feeding on their respective diets, natural-diet fry performed significantly more orientations and bites, the two major early feeding acts, than did the artificial-diet fry. When tested with live fish prey, fish reared on the natural diet performed fewer orientations and strikes and captured more prey per fry than did the artificial-diet fry. Natural-diet fry had a significantly better net efficiency (captures minus strikes minus orientations) than did artificial-diet fry. Diet, experience, and length (T.L.) of fry affected their predator efficiency significantly. We argue that providing hatchery-reared bass fry with an opportunity to prey on live forage fish once or twice before their release would enhance their survival and eventual recruitment into natural populations.     

Aggression Level in Different Water Velocities Depends on Population Origin in Grayling, Thymallus thymallus

Phenotypic plasticity is recognized as an important mechanism of adaptation. However, because of its potential costs and limits, it has been hypothesized to be reduced and ultimately become lost when there is no selection for its maintenance. Heterogeneous environments in particular are expected to favour and thus maintain plastic phenotypes. Lakes and rivers differ in their flow regimes. In addition to higher average water velocity, rivers are characterized by distinct spatial and temporal variation in water flow, whereas lakes can be regarded as quite uniform in this respect. We studied whether behaviour, which is generally considered to be highly plastic, shows differences in the degree of plasticity between lake and river populations of the European grayling, Thymallus thymallus, fry in response to different water velocities. Given that aggressive behaviour in fish has been shown to relate to ambient flow regime, we compared aggressiveness of hatchery‐reared grayling originating from hatchery stocks of two lake and two river populations in still and flowing water. River fish showed higher aggressiveness in flowing water compared with still water, whereas aggressiveness of lake fish did not appear to vary according to water velocity. The higher plasticity of aggressiveness evoked in river fish by different water velocities may thus represent an adaptation to more variable flow and presumably related feeding conditions in their natural environment.     

Telemetry and genetics reveal asymmetric dispersal of a lake‐feeding salmonid between inflow and outflow spawning streams at a microgeographic scale

The degree of natal philopatry relative to natal dispersal in animal populations has important demographic and genetic consequences and often varies substantially within species. In salmonid fishes, lakes have been shown to have a strong influence on dispersal and gene flow within catchments; for example, populations spawning in inflow streams are often reproductively isolated and genetically distinct from those spawning in relatively distant outflow streams. Less is known, however, regarding the level of philopatry and genetic differentiation occurring at microgeographic scales, for example, where inflow and outflow streams are separated by very small expanses of lake habitat. Here, we investigated the interplay between genetic differentiation and fine‐scale spawning movements of brown trout between their lake‐feeding habitat and two spawning streams (one inflow, one outflow, separated by <100 m of lake habitat). Most (69.2%) of the lake‐tagged trout subsequently detected during the spawning period were recorded in just one of the two streams, consistent with natal fidelity, while the remainder were detected in both streams, creating an opportunity for these individuals to spawn in both natal and non‐natal streams. The latter behavior was supported by genetic sibship analysis, which revealed several half‐sibling dyads containing one individual that was sampled as a fry in the outflow and another that was sampled as fry in the inflow. Genetic clustering analyses in conjunction with telemetry data suggested that asymmetrical dispersal patterns were occurring, with natal fidelity being more common among individuals originating from the outflow than the inflow stream. This was corroborated by Bayesian analysis of gene flow, which indicated significantly higher rates of gene flow from the inflow into the outflow than vice versa. Collectively, these results reveal how a combination of telemetry and genetics can identify distinct reproductive behaviors and associated asymmetries in natal dispersal that produce subtle, but nonetheless biologically relevant, population structuring at microgeographic scales.     

Within-stream variation in early life-history traits in brown trout

Significant additive genetic variance for most early life-history traits was found in brown trout Salmo trutta living in both allopatry above an impassable waterfall and sympatry (below the waterfall in the same stream) with alpine bullhead Cottus poecilopus. These traits included length, mass and yolk sac volume at hatching, and size at'button-up' (the time when yolk is enclosed within the body cavity). There were small differences in size at hatching and size at button-up among populations (adjusted for egg size). However, sympatric fry grew more rapidly and experienced lower mortality rates during the period of first feeding than allopatric fry. This might indicate behavioural differences between brown trout from the two populations. It is suggested that these phenotypic differences may be a result of adaptation to living in sympatry with alpine bullhead.     

Biological evaluation of frozen zooplankton as food for milkfish (Cbanoschanos) fry

Milkfish fry with an average standard length and weight of 13.88 mm and 3.95 mg, respectively, were reared for 30 days using live and frozen Moina macrocopa and Brachionusplicatilis at feeding densities of 10–20 individuals per ml. Growth, survival and yield were used as indicators of the overall performances of the various treatment groups. Fry fed live M. macrocopa showed gains (both length and weight), growth and survival rates and yields significantly higher than fry fed with other treatment groups (P < 0.05). However, significant reductions in growth and survival rates resulted when fry were fed frozen M. macrocopa. On the other hand, there were no significant differences in growth and survival rates (P < 0.05) in fry fed live or frozen B. plicatilis. The results of the current study showed that although milkfish fry could be grown successfully using B. plicatilis, feeding with live Moina significantly improved growth, survival rate and yield (P lt; 0.05). Frozen Moina was found to be unsuitable as a feed for rearing milkfish fry because it reduced growth rates and increased mortality. Comparisons between live and frozen rotifers have proven the suitability of frozen rotifers as feed for rearing milkfish fry. By freezing surplus rotifers this would permit short term storage in anticipation of high hatchery demand and overcome any unpredictable failures with live cultures.