CAN VELOCITY AFFECT GROWTH AND FECUNDITY OF FACULTATIVE RIVERINE FISH SPECIES?

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1 POISH JOURNA OF ECOOGY (Pol. J. Ecol.) Regular research paper Tadeusz PENCZAK Department of Ecology and Vertebrate Zoology, University of Łódź, 12/16 Banacha Str., Łódź, Poland, penczakt@biol.uni.lodz.pl CAN VEOCITY AFFECT GROWTH AND FECUNDITY OF FACUTATIVE RIVERINE FISH SPECIES? ABSTRACT: The length and body weight at subsequent annuli of four facultative riverine fish species: roach (Rutilus rutilus.), perch (Perca fluviatilis.), bream (Abramis brama.) and pike (Esox lucius.) living in lakes and rivers were compared on the basis of published data. Pike grew better in lakes (significant differences in length, and some for weight), but the other species in rivers. However, significant differences between medians were observed in all cases only for body length of roach, in many cases for the length of bream, as well as a few for its weight, and no one for perch. Comparable data on relative fecundity in both habitats were available for roach and pike. Pike females produced more eggs in lakes, but roach in rivers, and the differences were significant. KEY WORDS: lakes, rivers, fish, facultative riverine, standard length, body weight, fecundity. 1. INTRODUCTION Growth, fecundity and diet of fish are functions of environmental conditions. In temperate zones, fish growth is mostly correlated with water temperature, but fecundity with food supply (Wootton 1990, Mann 1991). In rivers, discharge and velocity can also strongly affect the population parameters and fish food resources (Statzner et al., 1988, Grossman et al., 1998, Matthews et al. 1996, Matthews 1998). Velocity may act as a concealing parameter for growth, because the energy the fish has to expend maintaining its position in or swimming is not used for growth (East and Magnan 1987). Fish species adapted to life in running waters are named reophilic, and those in stagnant water limnophilic, while in both habitats eurybiontic or facultative riverine (Wolter 2001, Kruk and Penczak 2003); the last name is in the frequent use. Absence of clear environmental preferences of facultative riverine species can indicate that they have large adaptable possibilities in realization of life history strategies (Wootton 1990, Poff and Allan 1995). Facultative riverine species can use flowing and stagnant waters, even alternately, also during their ontogeny (S chreiber 2002). I think that perch (Perca fluviatilis.) and roach (Rutilus rutilus.) began to invade rivers in central Poland not long ago. When in the 60s I began my fish inventory study in the Warta River (biggest tributary of the Oder River) and the Pilica River (biggest left-site tributary of the Vistula) perch was absent or rare at the majority of sample sites and roach was journal 10.indb :38:16

2 358 Tadeusz Penczak not abundant when reophilic species were still present in slightly polluted river sections (Penczak 1968, 1969). At the turn of the 80s and 90s, when water pollution level reached its highest value, perch became the second dominant after roach, in both rivers (Penczak et al. 1996, Kruk and Penczak 2003, Kruk 2006). Roach has achieved its first position in the ichthyofauna when increased sewage inputs freed it from competitors as it was shown already in the Narew River, biggest right-side tributary of the Vistula (Penczak and Koszalińska 1993). Since 1995 till now despite the fact that water quality is improving continuosly roach and perch remained dominants in both rivers (Kruk 2006, and data not published yet). In Europe such species as perch, roach, bream (Abramis brama.) and pike (Esox lucius.) can live in stagnant and running waters, but studies on their response in terms of growth and fecundity to conditions in both these habitats were not comparable. Hence the aim of this study is to test a hypothesis whether growth and fecundity of the four species have been realized on the same level during their ontogeny in running (rivers) and stagnant (lakes) waters. I have not encountered on such comparative study, as well as no information in books on the subject (Ricker 1968, Gerking 1978, Casselman 1983, Wootton 1990, Mann 1991). 2. MATERIA AND METHODS The published data on growth, and fecundity for perch, roach, bream and pike from Poland were used (Appendix), but also from certain research in other European countries (Żukov 1965, Williams 1967, Cragg-Hine and Jones 1969, Mann 1978, Kipling 1983, C owx 1988, 1990). For analyzing growth we used standard length longitudo corporis (l.c.) in centimeters, and body weight in grams, both on time of annulus formations. Fecundity was considered on the basis of relative fecundity, i.e. number of eggs per 1 kg of fish body weight of females (Bagenal 1973). Number of samples from lakes and rivers were not proportional and distribution of the investigated parameters were skewed, hence the Kruskal-Wallis test was adopted for post-hoc non-parametric comparisons (Quinn and Keough 2002). The parameters of length, weight and fecundity for fish from both habitats were compared with the U Mann-Whitney test (Z ar 1998). 3. RESUTS Medians of body length and weight on time of annuli formations calculated for roach, were higher in rivers than in lakes (Figs 1 and 2). However, compared results display significant differences for the third, fourth and fifth age length groups (Table 1), despite sample size (n = 10 33). The median values calculated for perch were slightly higher for the body weight in rivers, smaller or equal for length (Figs 1 and 2), but no compared pair results (both length and weight) displays significant difference. Table 1. Post-hoc comparisons of medians of roach body lengths for subsequent age groups (1 9) between lakes () and rivers (R). Explanations: n is number of observations (with the U Mann-Whitney test). Significant P values are bolded. Age groups U statistics P n n R 1-R R R R R R R R R journal 10.indb :38:16

3 Growth and fecundity of facultative riverine fish 359 Fig. 1. Increase in standard length in populations of roach, perch, bream and pike at the time of annuli formations in lakes () and rivers (R). Statistical symbols explanation is presented using pike as an example. Table 2. Post-hoc comparisons of medians of bream body lengths for subsequent age groups (1 11). Explanations as in Table 1. Age groups U statistics P n n R 1-R R R R R R R R R R R journal 10.indb :57:22

4 360 Tadeusz Penczak Fig. 2. Increase in weight in populations of roach, perch, bream and pike at the time of annuli formations. Symbols explanation on Fig. 1. Table 3. Post-hoc comparisons of medians of bream body weights for subsequent age groups (1 11). Explanations as in Table 1. Age groups U statistics P n n R 1-R R R R R R R R R R R journal 10.indb :57:22

5 Growth and fecundity of facultative riverine fish 361 Table 4. Post-hoc comparisons of medians of pike body lengths for subsequent age groups (1 7). Explanations as in Table 1. Age groups U statistics P n n R 1-R R R R R R R Table 5. Post-hoc comparisons of medians of pike body weights for subsequent age groups (1 4). Explanations as in Table 1. Age groups U statistics P n n R 1-R R R R Fig. 3. Relative fecundity (number of eggs per kg of fish body weight of females) of roach and pike in lakes and rivers. Symbols explanation on Fig. 1. journal 10.indb :57:23

6 362 Tadeusz Penczak Medians for length and weight of bream, on successive annuli formation, attained higher values in rivers than in lakes (Fig. 1 and 2). Differences in length values between fish from both environments were significant with exception of two age groups (Table 2), while in weight values only three age groups differed significantly (Table 3). Pike grew better in lakes than in rivers without exception, and this concerns medians of both length and weight (Figs 1 and 2). Highly significant differences in these parameters were recorded, with exception of the seventh annulus for length (Table 4), and the first and fourth annuli for weight (Table 5). Comparable data on relative fecundity were available for roach and pike. Pike females produced more eggs in lakes, but roach in rivers (Fig. 3). Both differences were significant on the level P < DISCUSSION Variability of specific morphological and behavioral traits clearly indicates that natural selection has increased surplus power on them (Ware 1982). ife history theory has explained how energy is allocated by fish into various vital functions, and we know at present that only 0 18% of ingested food is invested into growth, and calculated averages are rather very low (Ware 1982). Fish growth is essential and frequently critical element of fish population assessment that can be directed by many biotic and abiotic factors (Wootton 1990, Weerd and Komen 1998, Pegg and Pierce 2001). Better growth is a good indicator of conditions in given habitat because larger bodied animals can maintain homeostasis over a wider range of conditions (Gaston 1990), and it results in more rapid colonization of empty or poorly settled sites (Gaston 1988, Taylor and Gotelli 1994). Greater breadth of food resources, which is undoubtedly provided by meandering rivers, should be positively correlated with better growth ratio and it was confirmed in this paper. Among the four investigated species greater body length and weight values on successive annuli were obtained for roach, perch (exception for body weight) and bream living in rivers, and only the pike was growing significantly better in lakes. Unfortunately, the majority of these differences between fish from rivers and lakes were not significant and this concerns roach and perch mainly. Velocity may act as a masking factor for growth, because when fish maintain their position in the water column by swimming against a current, much less energy is available for growth (Wootton 1990). Each fish shows significant preference for a convenient velocity value ( amouroux et al. 1998), which is not easy to estimate. Individuals experiencing moderate velocity are in a better physiological condition than others living in still water, which is a result of training effect. In other cases competitive interactions with conspecifics may be more intense in still water (Wootton 1990). While comparing growth ratio of pike in river and lakes Kapczyńska and Penczak (1969) recorded difference not only in growth but also in body shape; this concerns perch in this research as well. Fish from lakes had a more elongated body than fish caught in rivers. Coming back to insignificant differences common to both roach and perch we have to underline that truly objective age determination strongly depends on applied techniques, tools and the type of chosen calcified structure (Casselman 1983, Mann and Steinmetz 1985). In papers selected for this study the variability of aging methods was unfortunately rather high. Our study can also suffer from the fact that we used data from a large part of Europe. There are many papers showing that growth rate is dependent on geographical variation (Pyron 1999), but some of them provide evidence that some fish attain larger size at lower latitudes (Modde and Scalet 1985) while others at higher latitudes (Isley et al. 1987, C onover and Present 1990). The impact of velocity on fecundity was positive for roach, but negative for pike. Roach in the Pilica River (Vistula system) matures at age 3+, and produces more eggs at subsequent annuli than in lakes (Penczak et al. 1977). Perhaps this is caused by the fact that fecundity success is inversely related to increase in velocity, and the species is not journal 10.indb :38:17

7 Growth and fecundity of facultative riverine fish 363 well adapted to flowing water yet, hence lack of the suitable adaptation is compensated by higher fecundity. ACKNOWEDGEMENTS: I thank Łukasz Głowacki for correction of English. Thanks are directed to A. Januszkiewicz for help in collecting published data. 5. REFERENCES Antosiak B Wzrost szczupaka (Esox lucius.) w jeziorach okolic Węgorzewa [Pike growth in the lakes of the vicinities of the Węgorzewo Town] Roczn. Nauk Rol. 77, B, 2: (in Polish, Eng. Summ.) Bagenal T.B Fish fecundity and its relations with stock and recruitment Rap. Rapp. P.-v. Réun. Cons. perm. int. Explor. Mer, 164: Bi ałokoz W Płodność wybranych gatunków ryb z kilku jezior mazurskich przygotowanych do doświadczalnego nawożenia [Fecundity of selected fish species from several Masurian lakes prepared for artificial fertilizing] Roczn. Nauk Rol. 95, H, 4: (in Polish) Brylińska M. (Ed.), 1986 Ryby słodkowodne Polski [Freshwater fishes of Poland] Wyd.I. PWN, Warszawa, 429 pp. (in Polish) Brylińska M. (Ed.), 1991 Ryby słodkowodne Polski [Freshwater fishes of Poland] Wyd.II. PWN, Warszawa, 429 pp. (in Polish) Brylińska M. (Ed.), 2000 Ryby słodkowodne Polski [Freshwater fishes of Poland] Wyd. III. PWN, Warszawa, 521 pp. (in Polish) Br ylińska M., Br yliński E eszcz [Bream] PWRi, Warszawa, 223 pp. (in Polish) Casselman J.M Age and growth assessment of fish from their calcified structures techniques and tools (In: Proceedings of the international workshop on age determination of oceanic pelagic fishes: tunas, billfishes, and sharks, Eds: E.D. Prince and.m. Pulos) NOAA Tech. Rep., NMFS 8, p Conover D.S.O., Present T.M Counter-gradient variation in growth rate: compensation for length of the grooving season among Atlantic silversides from different latitudes. Oecologia, 83: Cowx I.G Distribution and variation in the growth of roach, Rutilus rutilus (.), and dace, euciscus leuciscus (.), in a river catchment in south-west England J. Fish Biol. 33: C owx I.G The reproductive tactics of roach, Rutilus rutilus (.) and dace, euciscus leuciscus (.) populations in the Rivers Exe and Culm, England Pol. Arch. Hydrobiol. 37: Cragg-Hine D., Jones J.W The growth of Dace euciscus leuciscus (.), Roach Rutilus rutilus (.) and Chub Squalius cephalus (.) in Willow Brook, Northamptonshire J. Fish Biol. 1: East P., Magnan P The effect of locomotion activity on the growth of the brook charr, Salvelinus fontinalis Mitchill. Can. J. Zool. 65: Gaston K.J The intrinsic rates of increase of different body size Ecol. Entom. 14: Gaston K.J Patterns in the geographical ranges of species Biol. Rev. 65: Gerking S.D Ecology of freshwater fish production Blackwell Scientific Publications, Oxford, 520 pp. Grossman G.D., Ratajczak R.E., Crawford M., Fre eman M. C Assemblage organization in stream fishes: effects of environmental variation and interspecific interactions Ecol. Monogr., 68: Isley J.J., Noble R.., Koppelman J.B., Phillip D.P Spawning period and first-year growth of northern, Florida, and intergrade stocks of largemouth bass Trans. Am. Fish. Soc. 11: Kapczyńska A., Penczak T Wzrost szczupaka, Esox lucius., w rzekach Wyżyny Łódzkiej i terenów przyległych [Growth of pike, Esox lucius. In the rivers of the Łódź Upland and adjoining areas] Prz. Zool. XIII, 1: (in Polsh, Eng. Summ.) Karpińska-Waluś B Wzrost płoci (Rutilus rutilus.) w jeziorach okolic Węgorzewa [Growth of roach (Rutilus rutilus.) in the lakes of the vicinities of the Węgorzewo Town] Roczn. Nauk Rol. 77, B, 2: (in Polsh, Eng. Summ.) Kipling C Changes in the growth of pike (Esox lucius) in Windermere J. Anim. Ecol., 52: Korkuć M., Frankiewicz P., Zalewski M Wzrost dominujących gatunków ryb w rzece Grabi [Growth of dominant fish species in the Grabia River] Acta Univ. odz., Folia limnol. 5: (in Polsh, Eng. Summ.) Korwin-Kossakowski M., 1976 Płodność i wzrost szczupaka (Esox lucius innaeus 1758) z jeziora Warniak [Fecundity and growth of pike (Esox lucius innaeus 1758) from ake Warniak] Roczn. Nauk Rol. 97, H, 2: (in Polsh, Eng. Summ.) journal 10.indb :38:17

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9 Growth and fecundity of facultative riverine fish 365 Ricker W.E Methods for assessment of fish production in fresh waters IBP Handbook no. 3, Blackwell Scientific Publications, Oxford, 313 pp. Rolik H., Rembiszewski J.M Ryby i krągłouste (Pisces et Cyclostomata) [Fishes and cyclostomes] PWN, Warszawa, 252 pp. (in Polish) S chreiber A Diferences in levels of heterozygosity in populations of the common gudgeon (Gobio gobio, Cyprinidae) among adjacent drainages in Central Europe: an effect of postglacial range dynamics? Heredity, 89: Statzner B., Gore J.A., Resh V.H Hydraulic stream ecology: observed patterns and potential applications J. North Am. Benthol. Soc., 7: Szczepanek E Wzrost płoci, jelca i szczu paka w rzece Widawce [Growth of roach, dace and pike in the Widawka River] Acta Univ. odz., Folia limnol., 3: (in Polish) Szczerbowski J.A., Martyniak A., Terlecki J Wzrost szczupaka, krąpia, jelca, klenia i płoci w rzekach zlewiska Łyny [Growth of pike, white bream, dace, chub and roach in the rivers of the Łyna River system] Roczn. Nauk Rol., 97, H, 2: (in Polish, Eng. Summ.) Staff F., 1950 Ryby słodkowodne Polski i krajów ościennych [Freshwater fishes of Poland and adjoining countries] Trzaska, Evert, Michalski, Warszawa: 286 pp. (in Polish) Tay lor C.M., G otel li N.J The macroecology of Cyprinella: correlates of phylogeny, body size, and geofraphic range Am. Nat. 144: Terle ck i J., 1973 Płodność szczupaka [Fecundity of pike] Esox lucius (innaeus 1758) z jeziora Śniardwy Roczn. Nauk Rol. 95, H, 3: (in Polish, Eng. Summ.) Ware D.M Power and evolutionary fitness of Teleosts Can. J. Fish. Aquat. Sci., 39: Weerd J.H., Komen J The effects of chronic stress on growth in fish: a critical appraisal Comp. Bioch. Physiol. Part A, 120: Wilkońska H Wzrost płoci (Rutilus rutilus.) w podgrzanym jeziorze icheńskim [Growth of roach (Rutilus rutilus.) in heated ake icheń] Roczn. Nauk Rol., 97, H, 4: (in Polish, Eng. Summ.) (Received after revising Noveber 2006) Williams W.P The growth and mortality of four species of fish in the river Thames at Reading J. Anim. Ecol., 36: Włoszczyński B Tempo wzrostu ryb dolnego biegu rzeki Wełny [Growth rate of fish in the lower course of the Wełna River] Rocz. WSR w Poznaniu XXII: (in Polish, Eng. Summ.) Wojno T Sezonowe zmiany wzrostu leszcza (Abramis brama.) z jeziora Wdzydze [Seasonal changes of bream (Aramis brama.) growth in ake Wdzydze] Roczn. Nauk Rol., 84, B, 2: (in Polish, Eng. Summ.) Wolter C Conservation of fish species diversity in navigable waterways andsc. Urban. Plann., 53: Wootton R.J Ecology of Teleost Fishes Chapman and Hall, ondon. Załachowski W Szczupak [Pike] PWRi, Warszawa, 251 pp. (in Polish) Zawisza J Szybkość wzrostu leszcza, certy, brzany i krąpia w środkowym biegu Wisły w okolicach Warszawy [Growth rate of bream, vimba, barbel and white bream in the middle course of the Vistula River in the vicinities of Warsaw] Roczn. Nauk Rol., 57: (in Polish, Eng. Summ.) Zawisza J Wzrost ryb w jeziorach okolic Węgorzewa. Próba ustalenia niektórych prawidłowości [Growth of fish in the lakes of the vicinities of the Węgorzewo Town. An attempt at determining certain regularities] Roczn. Nauk Rol., 77, B, 2: (in Polish, Eng. Summ.) Zar J.H Biostatistical analysis Prentice-Hall, Englewood Cliffs, New Jersey. Z ieliński P Wzrost, pokarm i zagęszczenie dominujących gatunków ryb karpiowatych w rzece Warcie po utworzeniu zbiornika zaporowego [Growth, food and density of dominant cyprinid fish species in the Warta River after the creation of a dam reservoir] Acta Univ. odz., Folia limnol., 5: (in Polish, Eng. Summ.) Żukov P.J., 1965 Ryby Białorusi [Fishes of Byelorussia] Nauka i Technika, Mińsk: (in Belarussian) Żuromska H Wzrost okonia (Perca fluviatilis.) w jeziorach okolic Węgorzewa [Growth of perch (Perca fluviatilis.) in the lakes of the vicinities of the Węgorzewo Town] Roczn. Nauk Rol. 77, B, 2: (in Polish, Eng. Summ.) journal 10.indb :38:18

10 366 Tadeusz Penczak APPENDIX Sources of analyzed materials. Explanations: standard body length, W body weight, F relative fecundity. Reference Species Bream Perch Roach Pike Antosiak 1961 W Białokoz 1973 F F F F Brylińska 1986 W F Brylińska 1991 W F W W W Brylińska 2000 Brylińska and Bryliński 1968 W Brylińska and Bryliński 1973 F Cowx 1988 Cowx 1990 F Cragg-Hine and Jones 1969 Kapczyńska and Penczak 1969 Karpińska-Waluś, 1961 Kipling 1983 Korkuć et al W W W Korwin-Kossakowski, 1976 W F ewandowska-jarzynowa 1966 W Mann 1978 W Mann and Beaumont 1990 Mann and Penczak 1984 W W Marciak 1977 Penczak et al W Penczak et al W W Penczak et al W Penczak et al F Rolik and Rembiszewski 1987 W W W Szczepanek 1988 W W Szczerbowski et al Staff 1950 W Terlecki 1973 F Wilkońska 1977 Williams 1967 W W Włoszczyński 1964 W W W Wojno 1964 W Załachowski 1973 F Zawisza 1951 Zawisza 1961 Zieliński 1991 W Żukov 1965 W F W F F W F Żuromska 1961 W journal 10.indb :38:18