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Ecology and Impact of the Exotic Amphipod,Corophium Curvispinum

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Ecology and Impact of the Exotic Amphipod,Corophium Curvispinum Ecology and impact of the exotic amphipod, Corophium curvispinum Sars, 1895 (Crustacea: Amphipoda), in the River Rhine and Meuse S. Rajagopal, G. van der Velde, B.G.P. Paffen and A. bij de Vaate Reports ofth e project "Ecological Rehabilitation of Rivers Rhine and Meuse" No.75-1998 Institute for Inland Water Management and Waste Water Treatment (RIZA), P.O. Box 17, 8200 AA Lelystad, The Netherlands To be referred to as: Rajagopal, S., G. van der Velde, B.G.P. Paffen & A. bij de Vaate, 1997. Ecology and impact of exotic amphipod, Corophiumcurvispinum Sars , 1895 (Crustacea: Amphipoda), in the River Rhine and Meuse. Report (No. ) of the project "EcologicalRehabilitation of RiversRhine and Meuse" {with abstracts in Dutch, French and German). Institute for Inland Water Management and Waste Water Treatment (RIZA), P.O. Box 17, 8200 AA Lelystad, The Netherlands. Contents Preface I Summary III Samenvatting VII Résumé XI Zusammenfassung XV Listo f figures XIX List oftable s XXIII 1. Introduction 1 1.1. Distribution and range extensiono f Corophium curvispinum 1 1.2.Reason sfo r the present study 1 1.3. Objectives 3 2. Materials andmethod s 3 2.1. Study area 3 2.2. Methods 5 2.2.1. Life history andreproductiv e biology 5 2.2.2. Growth rates 6 2.2.3. Production 7 2.2.4. Distribution andimpact s of C. curvispinum onothe r macroinvertebrates 7 2.2.5. Mud-fixation 7 2.2.6. Filtration capacity 9 2.2.7. Hydrographie parameters 10 2.2.8. Statistical analysis 10 3. Results 10 3.1. Population density 10 3.2. Population structure 13 3.3.Se x ratio 13 3.4. Brooddevelopmen t 13 3.5. Breeding season 18 3.6. Broodsiz e 18 3.7. Losso fembryo sfro mth ebroo dpouc h 24 3.8. Relationship betweenbod y lengthan dweigh t 25 3.9. Growthrat e 25 3.10.Productio n 28 3.11. Mud-fixation 34 3.11.1. Onth e stones 34 3.11.2.O nexperimenta ltile s 38 3.12. Filtrationcapacit y 51 3.13. Seasonalvariation s inpopulatio n densities of macroinvertebrates inth e Lower Rhine 53 3.14. Distribution and impacto f C. curvispinum onothe r macroinvertebrates 62 4. Discussion 70 5. Conclusions and Recommendations 81 References 83 Appendix 89 Preface This project was financially supported by the Ministry of Transport and Public Works, the Ministry of Housing, Physical Planning and the Environment, Netherlands Organisation of the Advancemento fPur eResearc h(BION )an dBeijerinck-Poppin g Foundation.Th eresearc hcommitte e ofthi sprojec tconsiste do f Dr.Ir .G.M .va nDij k(RIVM) ,Mr .A .bi jd eVaat e(RWS/RIZA ) (Chairman), Drs. F.M.J. Oosterbroek (RWS), Dr. F.W.B, van den Brink (KUN) and Prof. Dr. G. van der Velde (KUN).W e are grateful to Prof.J.M .va n Groenendael and Mr. B. Kelleher for useful discussions. Thanks are due to M.J.E. Orbons, M.G. Versteeg and students for their assistance inth efiel d and laboratory. Summary Exoticspecie sinvasion sca nb eviewe da sspecia lcase so frang eextensio n(Hengeveld ,1989) . Thesespecie sar eofte n ecologically andphyleticall yver ydifferen tfro mnativ etaxa .Therefore ,the y may establish or spread due to different life history characteristics and exploiting resources differently from nativeorganism s ina disturbe decosystem s (e.g. humanactivities) .Th e invasion of exotic speciesca n cause drastic changes in ecosystems (Drake eta/. , 1989;Pinkste r etai, 1992; Dick etal., 1993 ; Nichols & Hopkins, 1993; Stewart & Haynes, 1994; Fahnenstiel etat., 1995). In recent years, large numbers of exotic species have invaded the River Rhine (Den Hartog etal., 1992;Va nde nBrin k etai, 1993),however , Corophium curvispinumSars ,ha sbee nsuccessfu lan d dominatesth eepilithi c communities. Inrespons et othi ssuccess ,a stud y wascarrie d outt o assess ecology and impacto f C.curvispinum inth e River Rhinean d Meuse,fro m March 1992t o February 1994. Inth eLowe r Rhine,th elife-cycl eo f C. curvispinumi sbase do nthre egeneration spe ryear .Th e breedingseaso no f C.curvispinum occurre dbetwee nMa yan dOctobe ran dwa sstrongl y correlated with watertemperature . Reproduction generally began inMarc h andwa swel lestablishe d by May. The overwintering generation died during June/July, but reproduction continued until October as a resulto fbreedin gb ysumme rgeneratio nindividuals .Dat ao nth ese xrati oo fC. curvispinumindicat e thatfemale s(6 0t o80% )wer emor eabundan ttha nmales .A lo wpercentag eo fmale swa sobserve d during May/June.Th e meanbroo dsiz e (25± 5egg s(mea n ± SD),rang e= 12-38a t Nijmegen and 18± 3 eggs, range = 10-24 at Lobith) of C.curvispinum inth e Lower Rhine, is one of the highest ever recorded and showed a positive correlation with chlorophyll-a. Like many other crustaceans, a linear relationship exists between the body length and brood size of C. curvispinum. The percentage egg loss from the brood pouch of C. curvispinumi s high when compared with other Corophiumspecies . On stone surfaces, the amount of fixed muddy material including all macroinvertebrates was 38 -1044 g m"2 (dry weight) and 7 -138 g m'2 (ash-free dry weight). On experimental tiles, the amount of mud material,excludin g macroinvertebrates, was found to be 0.3 -16.3 g m"2 (ash-free dryweigh t onmonthl y exposedtiles ) and0. 3 -101.8g m" 2(ash-fre e dryweigh to ncumulativ etiles) . A seasonal pattern was observed inth e amount of mud material fixedo n the stones of groins and experimental tiles.Th e amounto f mudmateria lo nth estone s ofgroin san dexperimenta l tiles inth e Lower Rhine was highest during the summer period. A correlation was established between IV population densities of C. curvispinuman d muddy material fixed on the stones and on tiles (dry weight and ash-free dry weight). For example, at Lobith the correlation formula was y = 228.27 + 0.003x (r = 0.74, P < 0.001) for the dry weight of mud in gram per square metre (x is the number of individuals per square metre). For the ash-free dry weight of the muddy tubes, the correlation formulawa sy = 30.5 6+ 0.0006 x (r = 0.84 , P< 0.001) .Th epercentag eo forgani c matter calculated from the dry weight and ash-free dry weight of mud material including macroinvertebrates on the stones ranged from 9%t o 23%(mea n± SD; 15.9± 3.7%) at Lobith andfro m 13%t o 20%(16. 0 ± 2.4%) at Nijmegen. No significant variation inthes e values existed between Lobith andNijmegen . Incomparison ,th e percentage of organic matter of the mud material excluding macroinvertebrates ontile s at Weurt (River Waal) ranged from 11% to 29%(21. 4± 4.3%). Mean values of the rateo f mud fixation per individual calculated from the cumulative density of C.curvispinum an d amounts of mudfixe dcumulativel yo nth estone sa tLobit han dNijmege n indicatea th econstan t rateo fabou t 2.5 u,gindividual' 1day" 1( n= 1,700,000)afte r 400days .Almos t the samevalu e wasfoun d atWeur t (2.3 u,gindividual" 1 day"1; n = 500,000) after 350days . A tentative estimation of filtration by C. curvispinum indicates a possible mean filtration capacity of 5 x 106cm 3 m"2day' 1 in the River Rhine.Th e mean filtration rate of C. curvispinumi n River Meusewa s estimated as0. 3 x 106cm 3m" 2day" 1.Th e variation inth efilterin g potential of C. curvispinumi nth e River Rhine and Meuse isdu e to the much higher population densities in River Rhine compared to the Meuse. However, since there are no data available on the variation in filtration rate of C.curvispinum i n the literature, a thorough investigation is necessary to calculate the actual filtering impact of this filter feeder. The population densities ofvariou s macroinvertebrates were studiedb ycollectin gstone sa t2 1 locations during September 1992 and September 1993 in the Rivers Rhine and Meuse. C. curvispinumwa s the most dominant macroinvertebrate, outnumbering all other species by many orders of magnitude. A maximum C.curvispinum densit y of 642,000 individuals m"2wa s recorded at De Steeg inth e River Ussel. C.curvispinum densities inth e River Rhine branches (River Waal, R. Nederrijn/Lek and R. Ijssel) showeda positiv e correlationwit haverag e stream velocities. Higher population densities of C.curvispinum were recorded in the River Rhine than in the River Meuse. The relatively higher salinity, water temperatures, stream velocities andioni c content (e.g.Sodium ) inth e River Rhine,resultin gfro m industrialdischarge s and mining activities,hav econtribute dt oth e enormous success of C.curvispinum inth e Rhine.Th e success of C.curvispinum isals o relatedt o its strategy in competing for settlement space by means of muddy tubes. The very high densities V of this opportunistic filter feeder had shown an enormous impact on other macroinvertebrates, especially Dreissenapolymorpha (Pallas) ;whic hha sdrasticall ydecline dsinc eth emas scolonisatio n of C. curvispinum.Th e possible reasons are discussed by comparing population densities of D. polymorphabefor e (September 1989) andafte r (September 1993)th e explosive population growth of C.curvispinum alon g the River Rhine. Despite the importance of the population explosion of C.curvispinum inth e River Rhine,ver y little is known about the fundamental features of filtration rate and tube building activity of this species. The success of invasive species is mostly due to their plasticity in response to the characteristics of invaded ecosystem.
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