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Ecological and Life History Characteristics of Ruffe (Gymnocephalus Cernuus) in Relation to Other Freshwater Fish Species

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Ecological and Life History Characteristics of Ruffe (Gymnocephalus Cernuus) in Relation to Other Freshwater Fish Species J. Great Lakes Res. 24(2):228-234 Internat. Assoc. Great Lakes Res., 1998 Ecological and Life History Characteristics of Ruffe (Gymnocephalus cernuus) in Relation to Other Freshwater Fish Species Jyrki Lappalainen* and Jakob Kjellman Department of Limnology and Environmental Protection P.O. Box 27, FIN-00014 University of Helsinki, Finland ABSTRACT. Ecological and life history characteristics of ruffe (Gymnocephalus cer- nuus) were studied in relation to 33 other native freshwater fish species in Finland. The descriptive ecological characteristics included 1) trophic guild, 2) species interaction, 3) adult habitat, 4) spawning habitat, 5) reproductive behavior, and 6) reproductive guild. The life history variables included 1) age at maturity, 2) length at maturity, and 3) fecundity as a number of eggs. Three different clusters of fish species were found in cluster analysis based on the ecological characteristics. The most distinct cluster of 7 species was comprised of predators. Ruffe was grouped into the largest cluster together with 20 other species and was closest to the cyprinid, gudgeon (Gobio gobio). In multidimensional scaling species were placed in two hypothetical dimensions firstly by their interaction with other species, trophic guild and adult habitat and secondly by their spawning habitat. Ruffe was placed near the center of the plot, suggesting that it has no special ecological requirement among the characteristics and species studied. Subsequent analysis of the life history characteristics revealed that ruffe was similar to the most typical r-selected species in Finland, showing high fecundity and low length at maturity, but with slightly slower growth rate. We, therefore, concluded that adult ruffe is a potential invader species showing no special ecological requirements and with life history characteristics typical for r-selected species. INDEX WORDS: Ruffe, life history, ecological characteristics, Finland, r-selected. INTRODUCTION Ruffe (Gymnocephalus cernuus) is a common benthic fish species in Finnish freshwaters. Other frequently occurring species in addition to ruffe include Eurasian perch (Perca fluviatilis), northern pike (Esox lucius), roach (Rutilus rutilus), and burbot (Lota lota) (Koli 1981, Tonn et al. 1990). As with many other small-sized fish, ruffe has received little attention due to its low economic value. However, in recent years it has colonized new wa- ters outside its former distribution range. In these newly colonized waters the abundance of ruffe has increased to high levels, a circumstance that has raised questions concerning its possible negative effects on indigenous fish species (e.g., Pratt et al. 1992, Rösch and Schmid 1996, Savino and Kolar 1996, Winfield et al 1996). The purpose of the present paper was to examine the ecological similarities of ruffe in relation to other native freshwater fish species in Finland, based on six ecological characteristics. We were interested in how various species are placed in hypothetical dimen- sions and especially in relation to ruffe, based on these six ecological characteristics. We hypothesised that species near each other, thus showing similarities in the ecological characteristics examined, should compete for niche dimensions when co-occurring. *Corresponding author. E-mail: [email protected] The species studied were also classified according to their age, fecundity, and length at maturity. Mandrak (1989) has shown that successful invaders have broad niche dimensions and life history characteristics typical for r-selected species. In general, r-selected species have short generation time, small size, high fecundity, and rapid growth. Hence we sugges- ted that ruffe should be closer to r-selected than K-selected species among the native freshwater fish species in Finland. MATERIALS AND METHODS Ecological and Life History Characteristics Ecological and life history characteristics were estimated for the 34 freshwater fish species native to Finland. This classification was based mostly on Finnish literature (Pitkänen 1961; Seppovaara 1982; Koli 1984, 1990; Mikelsaar 1984; Kangur and Kangur 1996). Ecological characteristics were estimated for adult fish at an age corresponding to maturity in about 50% of the fish and included 1) trophic guild, 2) species interaction, 3) adult habitat, 4) spawning habitat, 5) reproductive behavior, and 6) reproductive guild. These characteristics were the same as those used by Mandrak (1989) with the exception that the adult and spawning habitats were further divided into lentic and lotic water habitats and the repro- ductive guild was classified in more detail. The trophic guild was determined by the type of adult food comprising at least 75% of the diet. The categories were herbivory, planktivory, insectivory, piscivory, and omnivory. Species interactions were classified into three cate- gories: predator, prey, or neither. Further, the adult and spawning habitats were characte- rized as lentic and lotic waters by the type of depth continuum: wetland, nearshore, benthic, and pelagic. Reproductive behavior and guild were classified according to Balon (1975). Thus species were classified based on their reproductive behavior as nonguarders or guar- ders. The nonguarders included open-substrate spawners or brood hiders, and the guarders were substrate choosers, nest builders, or internal bearers. Based on the reproductive guild, species were either litho-pelagophils, lithophils, phyto-lithophils, phytophils, psammophils, speleophils, or ariadnophils. All three life history characteristics of age, fecundity, and length were classified according to the age corresponding to maturity in about 50% of the fish. Fecundity, as the number of eggs, was divided into five classes based on a logarithmic scale: 1) 1 to 100, 2) 101 to 1,000, 3) 1,001 to 10,000, 4) 10,001 to 100,000 and 5) 100,001 to 1,000,000. The length at maturity (cm) was classified into the following groups: 1) 1 to 10, 2) 11 to 20, 3) 21 to 40, 4) 41 to 80, and 5) > 80. The ecological and life history characteristics of the species studied were analyzed inde- pendently of their possible occurrence in lakes. Species having many ecological forms such as whitefish (Coregonus lavaretus) and brown trout (Salmo trutta) characteristics typical of lake forms were included. The characteristics thus used for whitefish were of the form having a mean of 42 gillrakers, while those for brown trout were typical of Salmo trutta m. lacustris. Statistical Analysis Cluster analysis (SYSTAT 1992) was used to classify the freshwater fish species according to their ecological characteristics. Since these characteristics were categorical, cluster analysis was based on disagreement percentage among the species. Thus, if all six characteristics were the same, the disagreement percentage would be 0%, as noted for three-spined stickleback (Gasterosteus aculeatus) and nine-spined stickleback (Pungitius pungitius). On the other hand, if no categorical characteristics were the same, the disagree- ment percentage would be 100%, as noted for ruffe and Arctic char (Salvelinus alpinus). Multidimensional scaling was also based on disagreement percentages scaled between 1 and 0. Species were partitioned into clusters with K-means clustering based on their life history characteristics (SYSTAT 1992). The clustering was started with three partitionings, because we expected that at least three groups of species could be found showing characteristics typical for either r- or K-selected species or falling between these two limits. Preliminary analysis showed that four clusters with only one or two species were already formed with six partitions and the results were difficult to interpret. Therefore, no further partitionings were analysed. RESULTS Ecological Characteristics Cluster analysis formed three large groups of species based on ecological characteristics (Fig. 1). All seven predators were grouped into cluster I. Ruffe was grouped into the largest cluster with 20 other fish species. The species in this cluster were all open-substrate spawners except the three- and nine-spined sticklebacks. Cluster II could be further divided into three smaller clusters based on trophic guild. All species in cluster IIa were plankti- vorous, in cluster IIb insectivorous except rudd (Scardinius erythrophthalmus), and species in cluster IIc were omnivorous. In cluster IIb ruffe was grouped into the same cluster with a cyprinid gudgeon (Gobio gobio). Four of the ecological characteristics studied were the same for ruffe and gudgeon: trophic guild, species interaction, spawning habitat, and reproductive behavior (Appendix). The fish species studied were ordinated based on interaction with other species, trophic guild, adult, and spawning habitats in two hypothetical dimensions in multidimensional scali- ng (Fig. 2). In dimension 1 species were placed by their interaction with other species and trophic guild. Predators, i.e., piscivorous species, were placed on the left side of the plot, while nonpiscivorous species were placed on the right side. Furthermore, dimension 1 indi- cated changes in adult habitat from pelagic on the left to benthic and nearshore habitats on the right side of the plot. In dimension 2 species were scaled by their spawning habitat. In the upper part of the graph were lentic-wetland or nearshore spawners, while in the lower part of the plot the species were mainly benthic spawners. Ruffe was placed near the center of the plot in multidimensional scaling (Fig. 2). Subsequent
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