Experimental evidence for trait utility of gill raker number in adaptive radiation of a north temperate fish

North temperate fish in post‐glacial lakes are textbook examples for rapid parallel adaptive radiation into multiple trophic specialists within individual lakes. Speciation repeatedly proceeded along the benthic–limnetic habitat axis, and benthic–limnetic sister species diverge in the number of gill rakers. Yet, the utility of different numbers of gill rakers for consuming benthic vs. limnetic food has only very rarely been experimentally demonstrated. We bred and raised families of a benthic–limnetic species pair of whitefish under common garden conditions to test whether these species (i) show heritable differentiation in feeding efficiency on zooplankton, and (ii) whether variation in feeding efficiency is predicted by variation in gill raker numbers. We used zooplankton of three different size classes to investigate prey size dependency of divergence in feeding efficiency and to investigate the effect strength of variation in the number of gill rakers. Our results show strong interspecific differences in feeding efficiency. These differences are largest when fish were tested with the smallest zooplankton. Importantly, feeding efficiency is significantly positively correlated with the number of gill rakers when using small zooplankton, also when species identity is statistically controlled for. Our results support the hypothesis that a larger number of gill rakers are of adaptive significance for feeding on zooplankton and provide one of the first experimental demonstrations of trait utility of gill raker number when fish feed on zooplankton. These results are consistent with the suggested importance of divergent selection driven feeding adaptation during adaptive radiation of fish in post‐glacial lakes.     

The role of gill raker number variability in adaptive radiation of coregonid fish

Gill raker divergence is a general pattern in adaptive radiations of postglacial fish, but few studies have addressed the adaptive significance of this morphological trait in foraging and eco-evolutionary interactions among predator and prey. Here, a set of subarctic lakes along a diversifying gradient of coregonids was used as the natural setting to explore correlations between gill raker numbers and planktivory as well as the impact of coregonid radiation on zooplankton communities. Results from 19 populations covering most of the total gill raker number gradient of the genus Coregonus, confirm that the number of gill rakers has a central role in determining the foraging ability towards zooplankton prey. Both at the individual and population levels, gill raker number was correlated with pelagic niche use and the size of utilized zooplankton prey. Furthermore, the average body size and the abundance and diversity of the zooplankton community decreased with the increasing diversity of coregonids. We argue that zooplankton feeding leads to an eco-evolutionary feedback loop that may further shape the gill raker morphology since natural selection intensifies under resource competition for depleted prey communities. Eco-evolutionary interactions may thus have a central role creating and maintaining the divergence of coregonid morphs in postglacial lakes.     

Structure,development and function of the branchial sieve of the common bream,Abramis brama,white bream,Blicca bjoerkna and roach,Rutilus rutilus

Synopsis The filter feeding organ of cyprinid fishes is the branchial sieve, which consists of a mesh formed by gill rakers and tiny channels on the gill arches. In order to establish its possible role during growth we measured the following morphological gill raker parameters over a range of sizes in three cyprinid fishes, bream, white bream and roach: inter raker distance, bony raker length, raker width, cushion length and channel width. At any given standard length common bream has the largest inter raker distance, roach the lowest and white bream is intermediate. In the comb model of filter feeding the inter raker distance is considered to be a direct measure of the mesh size and retention ability (= minimal size of prey that can be retained) of a filter. For the three species under study there is a conflict between the comb model and experimental data on particle retention. Lammens et al. (1987) found that common bream has a large retention ability whereas roach and white bream have a much smaller one. A new model, the channel model (Hoogenboezem et al. 1991) has been developed for common bream; in this model the lateral gill rakers can regulate the mesh size of the medial channels on the other side of the gill slit. The present data indicate that this model is not appropriate for white bream and roach. At any given standard length white bream and roach only reach 70% of the raker length of common bream, which means that in this model the gill slits should to be very narrow during filter feeding. The gill rakers consist of a bony raker and a fleshy cushion. The bony rakers have a rather long needle-like part outside the cushion in bream, but not in white bream and roach which have blunt gill rakers. Blunt gill rakers are not suited to reduce the diameter of the medial channels. The comb model seems more appropriate for white bream and roach, but doubts about the validity of this simple model remain. The sum of the areas of the medial channels is an approximation of the area through which water flows in the filter. This channel area therefore gives an impression of the capacity or flow rate of the filter. With this capacity estimation and an estimation of energy consumption we calculated an energy ratio of filter feeding. The energy ratio decreases with increasing standard length with an exponent close to the expected exponent of -0.40. The energy ratio is highest in bream, intermediate in white bream and lowest in roach.     

Temporal variation in feeding morphology and size-structured population dynamics in fishes

Considerable variation in morphology associated with resource use is a classic example of local adaptation to the environment. We demonstrate that a temporal change in feeding morphology might occur within a population. In a 5-year natural field experiment, we estimated gill raker morphology, resource density and population dynamics of the roach and its competitor, the perch. Despite a variation in density of zooplankton resources and perch across years, no change in roach density was observed. However, gill raker morphology in roach covaried with size structure of the zooplankton resource across years. A laboratory experiment confirmed that gill raker morphology has a great effect on roach foraging efficiency on zooplankton and that there is a functional trade-off with regard to zooplankton foraging. We suggest that the response in gill raker structure is an adaptation to deal with the rapid population dynamics of zooplankton, which are in turn mediated by changes in the size structure of the competing perch.     

Genetic differentiation in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka

The genetic and environmental basis for polymorphism in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka, was investigated. Analysis of 30 full sib families involving pure types and reciprocal hybrids revealed that the variation was partitioned significantly among families within cross types and among cross types in both traits. As in the wild, kokanee displayed more gill rakers than sockeye; reciprocal hybrids displayed intermediate counts. Gill raker length also varied markedly among cross types, with pure sockeye displaying 19% longer gill rakers than comparable sized kokanee. This difference was in the opposite direction predicted, given the common positive association between gill raker number and length in sympatric morphs of the same species in fishes. Gill raker number and length were generally not correlated within cross types, suggesting independent divergence of the traits. The results are discussed in relation to genetic and trophic divergence of the morphs and to factors selecting for differentiation in the two gill raker traits.     

Feeding habits and gill raker morphology of three planktivorous pelagic fish species off the coast of northern and western Kyushu in summer

Feeding habits and gill raker morphology were examined for the three major planktivorous pelagic fishes, Japanese anchovy Engraulis japonicus , Pacific round herring Etrumeus teres and Japanese jack mackerel Trachurus japonicus , off the northern and western coasts of Kyushu, in the north‐eastern part of the East China Sea in the summer months of 2001. Using fishes in the same size range (80–140 mm, standard length), the stomach contents of the three fish species were compared. The diet of the Japanese anchovy mainly consisted of Oncaeidae copepods, while the diets of the Pacific round herring and Japanese jack mackerel were dominated by calanoid copepods at all stations. Comparisons between prey size in the stomach, zooplankton size in the water and gill raker morphology suggested that the stomach contents of the three species were characterized mainly by the difference in the feeding behaviour between Japanese anchovy (filter‐feeding) and the other two species (particulate‐feeding), rather than by the difference in the morphology of feeding apparatus only. It was concluded that behavioural adaptations in the feeding of these pelagic fishes brought about trophic partitioning to some degree in this pelagic ecosystem in summer. Although the diets of these three species overlapped to some extent, there was still little likelihood of competition between the Japanese anchovy and the other two species. The potential for competition between the Pacific round herring and the Japanese jack mackerel is discussed.     

Intraspecific phenotypic variation among alewife populations drives parallel phenotypic shifts in bluegill

Evolutionary diversification within consumer species may generate selection on local ecological communities, affecting prey community structure. However, the extent to which this niche construction can propagate across food webs and shape trait variation in competing species is unknown. Here, we tested whether niche construction by different life-history variants of the planktivorous fish alewife (Alosa pseudoharengus) can drive phenotypic divergence and resource use in the competing species bluegill (Lepomis macrochirus). Using a combination of common garden experiments and a comparative field study, we found that bluegill from landlocked alewife lakes grew relatively better when fed small than large zooplankton, had gill rakers better adapted for feeding on small-bodied prey and selected smaller zooplankton compared with bluegill from lakes with anadromous or no alewife. Observed shifts in bluegill foraging traits in lakes with landlocked alewife parallel those in alewife, suggesting interspecific competition leading to parallel phenotypic changes rather than to divergence (which is commonly predicted). Our findings suggest that species may be locally adapted to prey communities structured by different life-history variants of a competing dominant species.     

Diversification, Sympatric Speciation, and Trophic Polymorphism of Arctic Charr, Salvelinus Alpinus Complex, in Transbaikalia

In northern Transbaikalia, independently evolving landlocked populations of Arctic charr are found in mountain lakes. To assess the diversity of charr in this region, speciation modes involved in the evolution of charr forms, and the role of trophic polymorphism in their divergence, we studied the morphology and feeding of dwarf, small, and large forms of Arctic charr from a number of Transbaikalian lakes. Meristic data on charr from five lakes support the earlier conclusion that the three forms do not represent separate lineages but have independently diverged in sympatry in each of the lakes. In 10 lakes, the dwarf form showed varying degrees of differentiation from normal (small and large) charr in meristic characters (up to morphologically distinct and presumably reproductively isolated groupings), which is viewed as various levels of sympatric divergence. Indexes of gill raker length in fish from 20 lakes vary among populations of both dwarf and normal charr, with forms having short and long rakers being sympatric in some of these lakes. However, the index can be used only for comparing charr of different forms up to about 32cm fork length (FL) because it is strongly negatively correlated with size in larger fish. The study of charr diets in 21 lakes indicates that large charr are piscivorous whereas dwarf and small charr feed on a wide range of invertebrates, partitioning these resources in different ways. Planktivores, including very specialized ones, and non-planktivores (benthic feeders, insectivores) can be identified within the small and dwarf forms. The proportion of plankton in the diets of dwarf and small charr is positively correlated with the number and length of gill rakers while the proportion of benthos is negatively correlated. Allopatric planktivorous and non-planktivorous small charr differ in body proportions; parallel emergence of such morphotypes in different parts of the range is a characteristic feature of the Salvelinus alpinus complex.     

Differences in gill raker morphology between two local populations of a benthophagous filter-feeding fish, Goniistius zonatus (Cheilodactylidae)

Gill raker morphology of a benthophagous fish Goniistius zonatus (Cheilodactylidae) (10.9–29.2 cm SL), using a filter-feeding mode, was compared between two locations (Morode and Arakashi) in southern Japan. Although gill raker number and gill raker length at the two locations did not differ, gill raker spacing was narrower relative to overall fish size at Morode than at Arakashi, mainly because gill raker width was greater at Morode. The difference of gill raker spacing is unlikely to have a genetic or physiochemical explanation. Small invertebrates (≤1.0 mm) were dominant on the substrate at Morode but were less common at Arakashi. Such small animals were consumed by many fish at Morode but were rarely exploited at Arakashi. At Morode, the narrow gill raker spacing would be effective in retaining small prey, which should be an important energy resource in this population. The difference of interraker spacing at the two locations seems to be related to available prey size at each location. Received: November 14, 2000 / Revised: February 13, 2001 / Accepted: February 28, 2001     

Trophic polymorphism, habitat and diet segregation in Percichthys trucha (Pisces: Percichthyidae) in the Andes

Divergent natural selection affecting specific trait combinations that lead to greater efficiency in resource exploitation is believed to be a major mechanism leading to trophic polymorphism and adaptive radiation. We present evidence of trophic polymorphism involving two benthic morphs within Percichthys trucha , a fish endemic to temperate South America. In a series of lakes located in the southern Andes, we found two morphs of P. trucha that could be distinguished on the basis of gill raker length and five other morphological measures, most of which are likely associated with the use of food resources. The differences were consistent across all lakes examined, and were correlated with habitat use and diet. Individuals with longer gill rakers were more abundant in the littoral zone (littoral morph) while the short gill-raker morph was more abundant at 10 m depth and deeper (deep benthic morph). Both morphs fed primarily on benthic invertebrates, but the littoral morph fed more on larval Anisoptera than did the deep benthic morph. Phenotypic correlations among traits were high for the littoral morph, but low and non-significant for the deep-benthic morph. We suggest that gill raker length may influence the relative efficiency of suction feeding for the two morphs. This is the first evidence of trophic polymorphism in fishes from temperate South America.     

Ecological consequences of food partitioning for the fish population structure in a eutrophic lake

In the highly eutrophic lake, Frederiksborg Slotssø, the diet composition of the bream (Abramis brama L.) and roach (Rutilus rutilus L.) populations was examined during three periods with different food availability. The length range of bream and roach was 9–34 cm (TL) and 5–18 cm (TL), respectively. The relative food composition was examined for 2 cm and 1 cm length intervals of bream and roach, respectively. During all three periods, bream shifted from benthic cladocerans (Alona sp.) to zooplankton and chironomids within a transitional length of 15.0–20.0 cm. These foodshifts were coupled with a change in feeding behaviour from particulate to filter feeding. The biomass of chironomids was too low to sustain the consumption of larger bream (>20.0 cm) which initiated feeding in the pelagic zone even in periods when the mean length and biomass of the preferred zooplankton, Daphnia cucullata, were low. In contrast to bream, roach fed mainly on zooplankton. With increasing size, roach progressively shifted to larger zooplankton species due to the increasing mesh size of their branchial system. The importance of benthic animals in the diet of roach was minor due to low feeding efficiency on prey buried in the sediment. Detritus appeared in the diet of bream and roach in periods of low availability of animal food items. Feeding on detritus may provide an energetic advantage to bream and roach and increase the carrying capacity for these species in lakes, where detritus is highly abundant. Especially for the larger fish due to the decrease in their relative metabolic demands. However, the ability of bream to filter feed and with increasing size to retain food items smaller than those retained by roach may be the main mechanism for the dominance of bream over roach in highly eutrophic lakes.     

The Impact of Fish Predation and Cyanobacteria on Zooplankton Size Structure in 96 Subtropical Lakes

Zooplankton are relatively small in size in the subtropical regions. This characteristic has been attributed to intense predation pressure, high nutrient loading and cyanobacterial biomass. To provide further information on the effect of predation and cyanobacteria on zooplankton size structure, we analyzed data from 96 shallow aquaculture lakes along the Yangtze River. Contrary to former studies, both principal components analysis and multiple regression analysis showed that the mean zooplankton size was positively related to fish yield. The studied lakes were grouped into three types, namely, natural fishing lakes with low nutrient loading (Type1), planktivorous fish-dominated lakes (Type 2), and eutrophic lakes with high cyanobacterial biomass (Type 3). A marked difference in zooplankton size structure was found among these groups. The greatest mean zooplankton size was observed in Type 2 lakes, but zooplankton density was the lowest. Zooplankton abundance was highest in Type 3 lakes and increased with increasing cyanobacterial biomass. Zooplankton mean size was negatively correlated with cyanobacterial biomass. No obvious trends were found in Type 1 lakes. These results were reflected by the normalized biomass size spectrum, which showed a unimodal shape with a peak at medium sizes in Type 2 lakes and a peak at small sizes in Type 3 lakes. These results indicated a relative increase in medium-sized and small-sized species in Types 2 and 3 lakes, respectively. Our results suggested that fish predation might have a negative effect on zooplankton abundance but a positive effect on zooplankton size structure. High cyanobacterial biomass most likely caused a decline in the zooplankton size and encouraged the proliferation of small zooplankton. We suggest that both planktivorous fish and cyanobacteria have substantial effects on the shaping of zooplankton community, particularly in the lakes in the eastern plain along the Yangtze River where aquaculture is widespread and nutrient loading is high.     

Changes in the fish and zooplankton communities of Ringsjön,a Swedish lake undergoing man-made eutrophication

During the 20th century Lake Ringsjön has developed from a mesotrophic to a eutrophic lake, and the phytoplankton community has changed from a rather diverse community to a monoculture of blue-green algae. The eutrophication process has accelerated during the last decade. The most important external nutrient loading of today comes from agriculture.Although phosphorus has been shown to be the primary nutrient leading to excessive algal growth in fresh water, several biotic factors — such as interactions between nutrients, phytoplankton, zooplankton and planktivorous fish — may play a decisive role in the occurrence and maintenance of large algal blooms.The aim of this investigation was to study the changes in the fish community of Lake Ringsjön, especially the most dominant planktivores, and the state of the zooplankton community during the seventies. The fish fauna is dominated by cyprinids, especially roach, and there are relatively few predatory fish. During the seventies the mean size of roach decreased, and measurements of the zooplankton community indicated that the predation pressure on zooplankton had increased. The mean sizes of cladocerans such as Daphnia and Bosmina, which were selected for by the planktivorous fish, decreased; the size of the calanoid Diaptomus, which was not preyed upon by the dominating fish species, did not change. The growth of zooplankton-feeding stages of several fish species was retarded, which meant that the growth of young perch decreased, while older roach were mainly affected. In the prevailing situation, planktivorous roach can maintain a numerous population of small individuals, whereas the predatory perch is at a disadvantage, and predation on zooplankton is intense.     

Food resource partitioning and trophic morphology of Brevoortia gunteri and B. patronus

The menhadens Brevoortia gunteri and B. patronus are sympatric and morphologically similar. The two species are planktophagous and exhibited significant ( P < 0·0001) food resource partitioning, with phytoplankton predominating in B. patronus stomachs and zooplankton in B. gunteri. The branchial apparatus of both species is of the typical alosine form. Brevoortia patronus has significantly more branchiospinules per mm (=19·47) and significantly longer gill rakers (=13·35 mm) than B. gunteri (=14·11, =12·01 mm respectively). B. patronus is characterized by a gill raker system forming a fine-meshed filter capable of retaining smaller food items. The results support the hypothesis that food resource partitioning is related to different morphological features of the branchial apparatus.     

Coexistence and origin of trophic ecotypes of pygmy whitefish,Prosopium coulterii,in a south‐western Alaskan lake

Ecologically, morphologically and genetically distinct populations within single taxa often coexist in postglacial lakes and have provided important model systems with which to investigate ecological and evolutionary processes such as niche partitioning and ecological speciation. Within the Salmonidae, these species complexes have been well studied, particularly within the Coregonus clupeaformis–C. laveratus (lake and European whitefish, respectively) group, but the phenomenon has been less well documented in the other whitefish genera, Prosopium and Stenodus. Here, we examined the morphology, feeding biology and genetic structure of three putative forms of the pygmy whitefish, Prosopium coulterii (Eigenmann & Eigenmann, 1892), first reported from Chignik Lake, south‐western Alaska, over 40 years ago. Field collections and morphological analyses resolved a shallow water (< 5 m depth) low gill raker count form (< 15 first arch gill rakers), a deepwater (> 30 m), low gill raker form and a deepwater, high gill raker count (> 15 gill rakers) form. The two low gill raker count forms fed almost exclusively on benthic invertebrates (mostly chironomids), while the deepwater, high gill raker count form fed almost exclusively on zooplankton; differences in diet were also reflected in differences both in δ¹³C and δ¹⁵N stable isotopes. All three forms were characterized by the same major mitochondrial DNA clade that has been associated with persistence in, and postglacial dispersal from, a Beringian glacial refugium. Analysis of variation at nine microsatellite DNA loci indicated low, but significant differentiation among forms, especially between the two low gill raker count forms and the high gill raker count form. The extent of differentiation along phenotypic (considerable) and genetic (subtle) axes among the Chignik Lake forms is similar to that found among distinct taxa of Prosopium found in pre‐glacial Bear Lake (Utah–Idaho, USA) which is probably at least ten times older than Chignik Lake. Our analyses illustrate the potential for the postglacial differentiation in traits subject to divergent natural selection across variable environments.     

Patterns of morphological changes and hybridization between sympatric whitefish morphs (Coregonus spp.) in a Swiss lake: a role for eutrophication?

Whitefish, genus Coregonus, show exceptional levels of phenotypic diversity with sympatric morphs occurring in numerous postglacial lakes in the northern hemisphere. Here, we studied the effects of human‐induced eutrophication on sympatric whitefish morphs in the Swiss lake, Lake Thun. In particular, we addressed the questions whether eutrophication (i) induced hybridization between two ecologically divergent summer‐spawning morphs through a loss of environmental heterogeneity, and (ii) induced rapid adaptive morphological changes through changes in the food web structure. Genetic analysis based on 11 microsatellite loci of 282 spawners revealed that the pelagic and the benthic morph represent highly distinct gene pools occurring at different relative proportions on all seven known spawning sites. Gill raker counts, a highly heritable trait, showed nearly discrete distributions for the two morphs. Multilocus genotypes characteristic of the pelagic morph had more gill rakers than genotypes characteristic of benthic morph. Using Bayesian methods, we found indications of recent but limited introgressive hybridization. Comparisons with historical gill raker data yielded median evolutionary rates of 0.24 haldanes and median selection intensities of 0.27 for this trait in both morphs for 1948–2004 suggesting rapid evolution through directional selection at this trait. However, phenotypic plasticity as an alternative explanation for this phenotypic change cannot be discarded. We hypothesize that both the temporal shifts in mean gill raker counts and the recent hybridization reflect responses to changes in the trophic state of the lake induced by pollution in the 1960s, which created novel selection pressures with respect to feeding niches and spawning site preferences.     

Reconstructing the past density of planktivorous fish and trophic structure from sedimentary zooplankton fossils: a surface sediment calibration data set from shallow lakes

1. As quantitative information on historical changes in fish community structure is difficult to obtain directly from fish remains in lake sediments, transfer function for planktivorous fish abundance has been developed based on zooplankton remains in surface sediment (upper 1 cm). The transfer function was derived using weighted average regression and calibration against contemporary data on planktivorous fish catch per unit effort (PF-CPUE) in multiple mesh size gill nets. Zooplankton remains were chosen because zooplankton community structure in lakes is highly sensitive to changes in fish predation pressure. The calibration data set consisted of thirty lakes differing in PF-CPUE (range 18–369 fish net^–1), epilimnion total phosphorus (range 0.025–1.28 mg P l^–1) and submerged macrophyte coverage (0–57%). 2. Correlation of log-transformed PF-CPUE, total phosphorus and submerged macrophyte coverage v the percentage abundance in the sediment of the dominant cladocerans and rotifers revealed that the typical pelagic species correlated most highly to PF-CPUE, while the littoral species correlated most highly to submerged macrophyte coverage. Consequently, only pelagic species were taken into consideration when establishing the fish transfer function. 3. Canonical correspondence analysis (CCA) revealed that the pelagic zooplankton assemblage was highly significantly related to PF-CPUE (axis 1), whereas the influence of total phosphorus and submerged macrophyte coverage was insignificant. Predicted PF-CPUE based on weighted average regression without (WA) and with (WA(tol)) downweighting of zooplankton species tolerance correlated significantly with the observed values (r² = 0.64 and 0.60 and RMSE = 0.54 and 0.56, respectively). A marginally better relationship (r² = 0.67) was obtained using WA maximum likelihood estimated optima and tolerance. 4. It is now possible, quantitatively, to reconstruct the historical development in planktivorous fish abundance based on zooplankton fossil records. As good relationships exist between contemporary PF-CPUE data and indicators such as the zooplankton/phytoplankton biomass ratio, Secchi depth and the maximum depth distribution of submerged macrophytes, it is now also possible to derive information on past changes in lake water quality and trophic structure. It will probably prove possible further to improve the transfer function by including other invertebrate remains, e.g. chironomids, Chaoborus, snails, etc., and its scope could be widened by including deeper lakes, more oligotrophic lakes, more acidic lakes and lakes with extensive submerged macrophyte coverage (in the latter case to enable use of the information in the fossil record on plant-associated cladocerans).     

Gill anomalies of perch and roach from four lakes differing in water quality

This study examined gill anomalies of two common freshwater fish species, roach Rutilus rutilus and perch Perca fluviatilis , collected over five seasons in 1989–1990 from four lakes in central Finland. The lakes differed both in water quality and in fish parasite species composition. Particular attention was paid to the differences in chloride cell proliferation. Gill anomalies were generally more common and abundant in roach than in perch. Chloride cell proliferation was the most frequent histological change in roach but was that least often found in perch. Most of the changes were systemic and light in severity. In perch an unidentified lesion of noncellular eosinophilic inclusions in an enlarged epithelium was also recorded. There was a marked pattern in the severity of histological changes in perch when comparing those from the natural Lake Peurunka to those from three 'altered' lakes. Gills of perch from Lake Peurunka possessed more alterations in all the recorded parameters. In roach, a significant differences were detected in chloride cell and epithelial proliferation: these were greater in Lake Peurunka and 'polluted' Lake Vatia. In perch, all histological changes were significantly more frequent at the end of autumn 1989 when compared to other seasons. In roach, increased frequencies of all gill changes were noted in winter and spring samples. Tissue reactions to parasites were either only localized or absent. The systemic gill alterations in freshwater fish may reflect the soft nature and low winter temperature of Finnish fresh waters, in which chemical imbalances, such as acid peaks, may induce more dramatic changes in fish tissues than in harder waters.     

Comparison of losses of planktivorous fish by predation and seine-fishing in a lake undergoing long-term biomanipulation

SUMMARY 1. Piscivore stocking at artificially high densities and fishing are the two common approaches to reduce the amount of planktivorous and benthivorous fish in lake biomanipulation programmes. Both measures have advantages and disadvantages, but their relative efficacy has not previously been directly compared.
2. We calculated the average annual catch of roach and bream in a lake undergoing long-term biomanipulation (Feldberger Haussee, Germany) by seining each year between 1992 and 1998. We compared this value with a bioenergetics estimate of annual consumption rates of the dominant cohorts of piscivores, pikeperch and pike, in 1997 and 1998. We also determined species composition and length distribution of prey fish in stomachs of the piscivores.
3. Roach was the dominant prey species of both pikeperch and pike, whereas bream was rarely taken by either piscivorous species. Seining removed on average larger specimens of roach than were found in the stomachs of the piscivores.
4. Based on stocking densities of the piscivores, published mortality rates, and individual consumption rates, feeding of pikeperch and pike on roach exceeded the manual removal of roach by seining by a factor of 4–15 (biomass) in 1997 and 1998.
5. Based on these results, a combination of fishing and piscivore enhancement is recommended. Whereas the stocks of adult roach and bream have to be reduced mainly by fishing, the predation of piscivores should be directed predominantly towards the juvenile zooplanktivorous fish. Therefore, small size-classes of piscivorous fish should be promoted by fisheries management, including stocking and harvest regulations.     

Field and experimental evidence of the effect of Jenynsia multidentata, a small omnivorous–planktivorous fish, on the size distribution of zooplankton in subtropical lakes

1. Small cladocerans, copepod nauplii and rotifers often dominate the zooplankton community in tropical and subtropical lakes. This is probably because of high predation pressure by small omnivorous–planktivorous fish, but experimental evidence is scarce.
2. This study used two approaches to test the effect of the small omnivorous–planktivorous fish species Jenynsia multidentata , which is frequently abundant in (sub)tropical eutrophic lakes in South America, on the size distribution of zooplankton. In Lake Blanca (Uruguay), which lacks any piscivores, we sampled seasonally for both fish and zooplankton. We also conducted an outdoor mesocosm experiment with treatments containing or lacking J. multidentata .
3. Together, the empirical and experimental data suggest that J. multidentata predation plays an important role in modulating the size structure of the zooplankton community in subtropical lakes. In the absence of J. multidentata , stocked large-sized zooplankters like Daphnia obtusa were abundant in the experiments, while small-sized zooplankton dominated in the presence of fish, as they did in the lake itself from spring to the end of the season.