Sediment-associated cues for larval settlement of Polydora cornuta and Streblospio benedicti (Polychaeta, Spionidae)

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1 Sediment-ssocited cues for lrvl settlement of Polydor cornut nd Strelospio enedicti (Polychet, Spionide) Sediment-ssoziierte Signle für die lrvle Ansiedlung von Polydor cornut und Strelospio enedicti (Polychet, Spionide) Von der Fkultät für Mthemtik und Nturwissenschften der Crl von Ossietzky Universität Oldenurg zur Erlngung des Grdes und Titels einer Doktorin der Nturwissenschften (Dr. rer. nt.) ngenommene Disserttion von Fru Zit Seesvári georen m in Kecskemét, Ungrn

2 Gutchter: Junior Prof. Dr. Tilmnn Hrder Zweitgutchter: Prof. Dr. Krsten Reise Tg der Disputtion: 31. Jnur 27

3 The most exciting phrse to her in science, the one tht herlds new discoveries, is not Eurek! ut Tht s funny... Isc Asimov

4 ERKLÄRUNG ERKLÄRUNG VERÖFFENTLICHUNGEN ERKLÄRUNG Teilergenisse dieser Areit sind ls Beitrge in Fchzeitschriften erschienen (Kpitel 3), ls Mnuskript eingereicht worden (Kpitel 4) oder in der Diplomreit von Justus Lodemnn verwendet worden (Teile von Kpitel 3). Mein Beitrg n der Erstellung der Areiten wird im Folgenden erläutert: SEBESVARI, Z., ESSER, F., NEUMANN R., SCHULTE, A., BRINKHOFF, T., HARDER, T. Monospecies cteril films on sediment induce lrvl settlement of the infunl spionid polychetes Polydor cornut nd Strelospio enedicti. J. Exp. Mr. Biol. Ecol. Sumitted. Entwicklung des multiple-choice Testverfhrens durch Z. S.; Hälterung der Polycheten durch Z. S. und F. E.; Isolierung der Bkterien durch A. S. und Z. S.; Durchführung der Siedlungstests durch Z. S. und R. N.; Bereitstellung ein Teil der Bkterien und phylogenetische Anlysen durch T. B.; Erstellung des Mnuskriptes durch Z. S. SEBESVARI, Z., ESSER, F., HARDER, T. 26. Sediment-ssocited cues for lrvl settlement of the inenthic spionid polychetes Polydor cornut nd Strelospio enedicti. J. Exp. Mr. Biol. Ecol. 337, Etlierung der Polychetenkultur im Lor und Entwicklung des no-choice Testverfhrens durch Z. S.; Kultivierung der Polycheten durch Z. S. nd F. E.; Durchführung der prktischen Areiten durch Z. S. und F. E.; Erstellung des Mnuskriptes durch Z. S.; Üerreitung durch Z. S. nd T. H. LODEMANN, J. 25. Untersuchung zum Einfluss usgewählter iologischer, chemischer und physiklischer Prmeter uf ds lrvle Siedlungsverhlten von Strelospio enedicti und Polydor cornut. Diplomreit, Universität Oldenurg. Betreuung der Areit und Bereitstellung der Polychetenkultur durch Z. S., F. E. und T. H. 1

5 ERKLÄRUNG KONFERENZBEITRÄGE SEBESVARI, Z., ESSER, F., HARDER, T. 26. Bcteril cues for lrvl settlement of the spionid polychete Polydor cornut. VII Interntionl Lrvl Biology Symposium, Coos By, Oregon, USA. Vortrg. ESSER, F., SEBESVARI, Z., HARDER, T. 25. Influence of iogeochemicl sediment prmeters on infunl coloniztion of spionids in tidl flts (Wdden Se, Germny). ASLO, Sntigo de Compostell, Spnien. Poster. SEBESVARI, Z., ESSER, F., HARDER, T. 24. Sediment relted settlement cues for lrve of the polychete Polydor cornut. VI Interntionl Lrvl Biology Conference, Hong Kong, Chin. Vortrg. THEMATISCH FREMDE VERÖFFENTLICHUNGEN WÄHREND DER PROMOTION AZAIZEH, H., SALHANI, N., SEBESVARI, Z., SHARDENDU, S., EMONS, H. 26. The potentil of two wetlnd plnts to phytoremedite selenium in constructed wetlnd. J. Phytoremed. 8, SEBESVARI, Z., ETTWIG, K.F., EMONS, H. 25. Biomonitoring of tin nd rsenic in different comprtments of limnic ecosystem with emphsis on Coricul flumine nd Dikerogmmrus villosus. J. Environ. Mon. 7, EMONS, H., SEBESVARI, Z., FALK, K., KRACHLER, M. 24. Occurrence nd specition of rsenic, ntimony nd tin in specimens used for environmentl iomonitoring of limnic ecosystems. In: Hirner, A.V., Emons, H. (eds.) Orgnic metl nd metlloid species in the environment: Anlysis, distriution, processeses nd toxicologicl evlution. Springer, Berlin, Heidelerg, New York. AZAIZEH, H.A., SALHANI, N., SEBESVARI, Z., EMONS, H. 23. The potentil of rhizosphere microes isolted from constructed wetlnd to iomethylte selenium. J. Environ. Qul. 32,

6 ABSTRACT ABSTRACT ABSTRACT Ptchy distriution of infunl polychetes my result from ctive site selection of lrve guided y sediment ssocited microil cues. This hypothesis ws tested in still wter lortory settlement ssys with nd without choice option for lrve of two spionid polychetes with indirect development: Polydor cornut nd Strelospio enedicti. Lortory rood cultures of these spionids were estlished nd yielded sufficient numer of lrve with plnktotrophic development for iossys. Lrve were oviously le to ccept or reject ttrctive nd unttrctive sediment qulities in no-choice ssys in species specific mnner. In oth species lrve displyed high settlement (75 to 95 %) in response to nturl sediment nd significntly low settlement (5 to 5 %) in shed sediments, whilst in cse of P. cornut significntly low settlement (25 to 55 %) resulted in sterile sediment. Low settlement in sediments treted y steriliztion or comustion most likely resulted from vriety of fctors such s modified sediment fric, grin size distriution nd quntity of dsored orgnic mtter. To experimentlly ddress the potentil role of microorgnisms nd microil metolites s meditors of lrvl settlement, shed sediment ws inoculted with mixture of vile microorgnisms detched from nturl sediment of dult hitts. The presence of vile microorgnisms significntly incresed lrvl settlement in comprison to the control of shed sediment indicting tht lrvl settlement ws t lest prtilly medited y the presence of microorgnisms ssocited with sediment. Susequently, 15 cteril isoltes otined from nturl sediment of dult hitts elonging to 5 phylogenetic clsses i.e. α-proteocteri (4), γ-proteocteri (3), Bcillles (3), Flvocteri (3) nd Sphingocteri (2), were screened for their ility to induce lrvl settlement in P. cornut nd S. enedicti. Recoloniztion of shed sediment with cteri resulted in low cteril cell densities (< 1 5 cells g -1 sediment). At these densities none of the 15 isoltes triggered settlement of spionid lrve. Recoloniztion of sterilized nturl sediment resulted in cteril densities etween cells g -1 sediment, i.e. one order of mgnitude lower thn in nturl sediment. In no-choice ssys two out of 15 isoltes, i.e. Loktnell sp. DF11 strin (α-proteocteri, Roseocter-clde) nd strin 54 (Flvocteri, closest mtch t GenBnk Psychroflexus tropicus) significntly triggered settlement of P. cornut lrve. There ws no correltion etween the 3

7 ABSTRACT phylogenetic ffilition of cteri nd their effect on lrvl settlement. Both inductive isoltes nd the non-inductive isolte Rhodoctercee cterium strin DF16 (α-proteocteri, Roseocter clde) were further tested in multiplechoice ssys using four prllel experimentl tretments of sediment, i.e. nturl, sterile nd two sediment tretments of different quntity nd qulity of cteri. Multiple-choice ssys reveled cler differences in lrvl settlement of oth spionids. Generlly, nturl sediment triggered high rtes of settlement while sterile sediment evoked significntly lower rtes of settlement. At cteril cell densities of 2. to 9. x 1 8 g -1 sediment strin DF11 nd 54 induced similr rtes of settlement s the control of nturl sediment. DF16 did not trigger lrvl settlement t ny of the densities (2. x 1 6 to 5. x 1 7 g -1 ) under investigtion. Ded or suspended cells s well s wter solule products of DF11 did not induce lrvl settlement either. The results of this study suggest tht the settlement cue of S. enedicti nd P. cornut is of cteril origin ut not relted to unique cteril genus, het lile nd ssocited with the sediment. Furthermore, the settlement of P. cornut is influenced y gregrious ehvior nd the signl is minly hroured in sediments formerly inhited y conspecific dults. In S. enedicti gregrious ehvior does not influence lrvl settlement. 4

8 ZUSAMMENFASSUNG ZUSAMMENFASSUNG ZUSAMMENFASSUNG Aggregiertes Vorkommen von inenthischen Polycheten im Hitt knn durch eine von sediment-ssoziierten mikroiellen Signlen geleitete ktive Sustrtwhl der Lrven erklärt werden. Diese Hypothese wurde durch lorsierten lrvlen Ansiedlungstests für zwei Spioniden mit indirekter Entwicklung, Polydor cornut und Strelospio enedicti, getestet. Für die eiden Polychetenrten wurden Lorkulturen etliert, um einen gnzjährigen Zugng zu plnktonischen Lrven zu gewährleisten. In Testreihen ohne Auswhlmöglichkeit wren die Lrven offensichtlich in der Lge ktiv eine Siedlungsentscheidung zu treffen. Beide Arten zeigten hohe Siedlungsrten in ntürlichem Sediment (zwischen 75% und 95 %) und niedrige im ei 6 C verschten Sediment (zwischen 5% und 5 %). Im durch Autoklvieren sterilisierten Sediment siedelte S. enedicti in gleich hohen Rten wie im ntürlichen Sediment, während die Siedlung von P. cornut im sterilen Sediment signifiknt niedriger wr ls im ntürlichen Sediment (zwischen 25% und 55 %). Niedrige Siedlungsrten im verschten zw. sterilen Sediment können jedoch durch eine Vielzhl von Prmetern wie die veränderte Struktur, Korngrößenverteilung und Qulität zw. Quntität des orgnischen Mterils verurscht werden. Um die potenzielle Rolle der Mikroorgnismen und deren Metoliten in der lrvlen Ansiedlung zu untersuchen, wurde verschtes Sediment mit leendigen Mikroorgnismen eimpft, die durch Aschütteln von ntürlichem Sediment us dem Hitt der dulten Tiere gewonnen wurden. Die Anwesenheit dieser Mikroorgnismen führte zu signifiknt erhöhten Siedlungsrten der Lrven eider Arten im Vergleich zu den uneimpften Negtivkontrollen, ein Hinweis druf, dss die lrvle Ansiedlung dieser Spioniden - zumindest teilweise - von sediment-ssoziierten Mikroorgnismen geleitet wird. Um Auslöser des positiven Siedlungssignls zu identifizieren, wurden 15 kterielle Isolte des ntürlichen Leensrumes eider Spioniden uf die Fähigkeit lrvle Ansiedlungen zu eeinflussen getestet. Die Isolte gehörten zu 5 phylogenetischen Klssen, nmentlich zu α-proteocteri (4), γ- Proteocteri (3), Bcillles (3), Flvocteri (3) und zu Sphingocteri (2). Ds Beimpfen von verschtem Sediment führte regelmäßig zu Zellzhlen von < 1 5 Zellen g -1 Sediment. In dieser Zelldichte vermochte keine der 15 Isolte die lrvle Ansiedlung zu eeinflussen. Ds Beimpfen von sterilem Sediment führte 5

9 ZUSAMMENFASSUNG zu Zellzhlen zwischen 1 7 und 1 8 Zellen g -1 Sediment. Dmit sind diese Zelldichten etw um eine Größenordnung niedriger ls die ntürlicher mriner Sedimente. In lrvlen Ansiedlungstests ohne Auswhlmöglichkeit hen zwei der fünfzehn Isolte, DF11 (α-proteocteri, Roseocter-Stmm) und Isolt 54 (Flvocteri), die Siedlung der Lrven eider Arten signifiknt gefördert. Die eiden siedlungs-induktiven Isolte und ein nicht-induktives Isolt Rhodoctercee cterium strin DF16 (α-proteocteri, Roseocter clde) wurden mittels Ansiedlungstest mit Vierfchuswhl näher untersucht. Dei wurden ntürliches, steriles sowie zwei mit Isolten verschiedener Zelldichte oder Qulität eimpftes Sediment ngeoten. Die Ansiedlungstests mit Vierfchuswhl hen ein klres Siedlungsmuster eider Arten gezeigt. Im Allgemeinen wr ntürliches Sediment sehr ttrktiv für die lrvle Ansiedlung, während steriles Sediment von den Lrven signifiknt seltener gewählt wurde. Die eiden induktiven Isolte DF11 und 54 hen ei Zelldichten von 2. is 9. x 1 8 g -1 Sediment gleich hohe Siedlungrten gezeigt wie ds ntürliche Sediment. DF16 ht die lrvle Ansiedlung ei Dichten, die mit diesem Isolt erzielr wren (2. x 1 6 is 5. x 1 7 g -1 Sediment) nicht eeinflusst. Durch Hitze getötete oder leendige er suspendierte Zellen sowie wsserlösliche Metolite des Isoltes DF11 hen die lrvle Ansiedlung eenflls nicht eeinflusst. Die Ergenisse dieser Areit weisen druf hin, dss die Siedlungssignle für die Spioniden S. enedicti und P. cornut zumindest teilweise kterieller Herkunft sind, er nicht uf eine einzelne kterielle Gttung zurückzuführen sind. Die signlgeenden chemischen Verindungen sind hitzelil und sedimentssoziiert. Desweiteren konnte gezeigt werden, dss die Siedlung von P. cornut durch ds Vorhndensein von Individuen der gleichen Art positiv eeinflusst wird. Dei ist esonders Sediment, in dem ereits vorher dulte P. cornut leten, ttrktiv für die lrvle Ansiedlung, während die Adulten selst und deren leere Röhren keine oder nur geringe Siedlungsinduktion zeigten. Bei der lrvlen Ansiedlung von S. enedicti scheinen die eigenen Artgenossen keinen nchweisren Einfluss zu hen. 6

10 CONTENTS CONTENTS CONTENTS 1 GENERAL INTRODUCTION DISPERSAL OF PLANKTONIC LARVAE CHARACTER AND PERCEPTION OF SETTLEMENT SIGNALS SETTLEMENT, METAMORPHOSIS AND RECRUITMENT POST-SETTLEMENT EVENTS OBJECTIVES GETTING STARTED: SELECTION AND CULTURING OF TEST POLYCHAETES SELECTION OF SUITABLE TEST POLYCHAETES POLYDORA CORNUTA STREBLOSPIO BENEDICTI CULTURING OF ADULT POLYCHAETES LARVAL CULTURE TECHNIQUES LARVAL DEVELOPMENT AND SETTLEMENT BEHAVIOR DEVELOPMENT OF SETTLEMENT ASSAYS TO STUDY ACTIVE SETTLEMENT CHOICE IN POLYCHAETE LARVAE INTRODUCTION Lrvl competence Lrvl selectivity t settlement versus delyed metmorphosis Influence of light on lrvl settlement MATERIAL AND METHODS Development of settlement ssys Collection nd tretment of nturl sediment smples Setup of settlement ssys Assessment of potentil experimentl rtifcts Sttisticl nlyses RESULTS Settlement in different sediment qulities (no-choice ssys) Settlement in different sediment qulities (multiple-choice ssys) Lrvl competence Lrvl selectivity t settlement versus delyed metmorphosis Testing the influence of light on lrvl settlement Assessment of potentil experimentl rtifcts DISCUSSION Lrvl settlement in different sediment qulities

11 CONTENTS Lrvl competence nd selectivity issues Suitility of different ssy designs to test lrvl settlement THE ROLE OF SEDIMENT-ASSOCIATED BACTERIA IN THE SETTLEMENT OF INFAUNAL POLYCHAETES INTRODUCTION MATERIAL AND METHODS Testing the influence of nturl cteril communities on lrvl settlement Testing the influence of cteril isoltes on lrvl settlement Monitoring the tretment efficiencies RESULTS Isoltion of cteri nd phylogenetic nlysis Lrvl response to nturl cteril community Lrvl response to sediment inoculted with cteril isoltes Effect of cteril cell density nd viility on lrvl settlement The effect of wter solule cteril products on lrvl settlement The effect of suspended cteri on lrvl settlement DISCUSSION DO CONSPECIFICS TRIGGER LARVAL SETTLEMENT? INTRODUCTION MATERIAL AND METHODS RESULTS Lrvl response to conspecific dults Lrvl response to different constituents of the dult hitt Lrvl response to conspecific dults nd juveniles DISCUSSION CONCLUSION REFERENCES ACKNOWLEDGEMENT CURRICULUM VITAE

12 GENERAL INTRODUCTION 1 GENERAL INTRODUCTION GENERAL INTRODUCTION Mny sessile enthic mrine invertertes, such s oysters, rncles nd polychetes, develop vi pelgic lrvl phse followed y enthic juvenile nd dult existence. Advntges of lrvl dispersl re genetic exchnge etween different popultions, voidnce of competition for resources with dults, fst (re)coloniztion of new hitts nd incresed ility to withstnd locl extinction (Levin 1984, Pwlik 1992, Pechenik 1999). Possile disdvntges include dispersl wy from fvorle hitts or conspecifics, greter vulnerility to environmentl stresses nd predtion (Thorson 195, Pechenik 1999). The erly life-history of mrine orgnisms is commonly divided into series of functionl or ehviorl stges, such s pssive dispersl, development, chievement of competence for settlement nd metmorphosis, ctive sustrte explortion, initil settlement, metmorphosis nd survivl of the newly settled individuls (Rodríguez et l. 1993). This thesis focuses on ctive lrvl choice t initil settlement in two spionid polychetes with indirect development, Polydor cornut nd Strelospio enedicti. Since there re mny inconsistencies in the literture out the reltive importnce of pssive nd ctive dispersl, the existence nd role of ctive lrvl choice nd the terminology of competence, settlement, metmorphosis nd recruitment; these issues re riefly summrized in the following. 1.1 DISPERSAL OF PLANKTONIC LARVAE During the pelgic phse, which lsts in species-specific pttern from few hours to severl months, inverterte lrve disperse over vrious sptil scles. On lrge scle (< 1s of kilometers) lrvl dispersion is driven y hydrodynmic processes such s currents, tides nd stgntions. Mximl dispersion cpcity of lrve is relted to the length of their plnktonic life nd to the rte nd direction of previling currents (Scheltem 1986). Wheres lrge scle distriution is commonly ccepted to e driven y hydrodynmic processes, lrvl fte shortly prior to settlement is discussed controversively. The hypotheses tht lrve redy to settle sink pssively through the wter column like "seeds in the wind" ws formulted very erly (e.g. Yonge 1937) nd is still discussed in the literture. In well-lnced study Butmn investigted the reltionship etween smll scle hydrodynmic processes nd ptterns of initil lrvl settlement (Butmn 1989). 9

13 GENERAL INTRODUCTION She concluded tht in most species pssive deposition determines where lrve initilly encounter the se ed. However, she pointed out tht fter encountering the surfce ctive or pssive redistriution of lrve tkes plce. Redistriution fter sediment contct my hppen through crwling or hopping (Butmn 1989), ctive swimming ove the ottom or pssive redistriution (Eckmn 1979, Plmer 1988). Butmn (1989) hypothesized tht "ctive selection mechnisms proly operte t the time the lrv first reches the ottom, such tht the orgnisms cn opt to remin or leve the depositionl locle. The lrv is not, however, "free to choose" in the sense of freely inspecting ll the options." On this note, ctive site selection of lrve is commonly oserved on smll sptil scles (Keough nd Downes 1982, Woodin 1986, Butmn et l. 1992, Desroy et l. 1997, Hrvey nd Bourget 1997, Pernet et l. 23). 1.2 CHARACTER AND PERCEPTION OF SETTLEMENT SIGNALS A criticl phse in the developmentl mode vi pelgic lrve is the detection of suitle sustrtum for susequent juvenile nd/or dult life. Hitt recognition y lrve is medited y either positive or negtive settlement cues in the form of single or mixed cues (Woodin 1991, Snelgrove nd Butmn 1994). Some lrve metmorphose in response to very specific cues, while others re generlists nd respond to cues of vrious origins (Qin 1999). For orgnisms settling on hrd sustrtum lortory experiments hve confirmed tht exploring lrve respond to physicl fctors (Mullineux nd Butmn 1991, Mid et l. 1994, Thiygrjn et l. 23) nd chemicl cues of iotic nd iotic origin (reviewed y Pwlik 1992, Qin 1999, Steinerg et l. 22). A frequently studied iotic source of cues ws iofilms (gglomertes of ttched cteri, enthic ditoms, fungi nd protozo). Biofilms hve een shown to elicit highly specific lrvl response with respect to their different origin nd/or growth phse in hrd sustrte settlers (ZoBell et l. 1935, Wieczorek nd Todd 1997, Qin et l. 23, Lu et l. 25). Both, smll orgnic metolites nd mcromoleculr extrcellulr polymers hve een identified nd prtilly purified s lrvl settlement cues in mjor iofilm components, i.e. cteri (Fitt et l. 199, Zimmer-Fust nd Tmurri 1994, Hrder et l. 22) cteril extrcellulr polymers (EPS) (Kirchmn et l. 1982, Mki et l. 199, Szewzyk et l. 1991, Lu et l. 23), ditoms (LeTourneux nd Bourget 1988, Hrder et l. 22) nd ditom EPS (Lm et l. 23/25). 1

14 GENERAL INTRODUCTION Although in soft sediment hitts less effort ws spent to identify the cues influencing lrvl settlement thn in hrd sustrt (Woodin 1986), severl prmeters were discussed or identified in this regrd, such s the orgnic content of the sediment (Butmn nd Grssle 1992, Grssle et l. 1992, Zettler 1996), the presence of elevted sulfide concentrtions (Cuomo 1985), sediment disturnce (Woodin et l. 1998, Mrinelli nd Woodin 24), sediment grin size distriution (Pinedo et l. 2), presence of conspecific juveniles or dults in the hitt (clled gregrious settlement) (Highsmith 1982, Olivier et l. 1996, Hrdege et l. 1998) nd hloromtic metolites of symptric orgnisms (Woodin et l. 1993, Hrdege et l. 1998). Similr to hrd sustrte settling orgnisms, the role of microorgnisms hs een suggested for infunl orgnisms (Wilson 1955, Gry 1966, 1967) ut only few follow-up works identifying the role of sediment-ssocited microorgnisms on infunl recruitment ptterns hve een performed since (Hdl et. l 197, Hermnn 1975 nd 1995). The perception of settlement cues is determined y their nture nd chemicl properties. Whilst physicl prmeters sich s sediment grin size or surfce texture hve to e explored y direct surfce contct of lrve chemicl cues my lso e detected in the wter column or in close proximity to surfces, such s the surfce oundry lyer (SBL, few hundred μm thin lyer of wter ner the surfce in which the velocity chnges from zero t the surfce to the free strem vlue wy from the surfce). Under flow conditions wter-solule chemicl cues re likeky perceived t the SBL t est, where flow-speed is low nd chemicl sustnces my ccumulte (Pwlik 1992). The perception of wter solule cues ove the SBL y the lrve is limited due to fst dilution. Under flow conditions relistic model of cue perception might thus function y signl detection ner the surfce ccompnied y controlled ehviorl surfce explortion like crwling, wlking (e.g. rncle cyprids, once ttched, use their ntenne to wlk long the sustrte surfce), drifting with the wter like "lloonist" (pssive edlod trnsport) or "ping pong ll" (ctive chnge of the verticl position) (Aelson 1997). Furthermore, Hdfield (24) reported tht lrve of the nudirnch Phestill sioge rpidly responded to dissolved settlement cues y sinking which enhnced their trnsport to the sustrtum, even in wve-driven turulent flows. In res or time slots of low flow-velocities wter solule cues my e perceivedin the wter column nd induce lrvl settlement (Coon et l. 199, Highsmith 1982). However, in most cses lrve 11

15 GENERAL INTRODUCTION hve een reported to detect chemicl settlement signls through direct contct with the surfce or in the SBL (Pwlik 1992, Morse nd Morse 1984, Perce und Scheiling 199). 1.3 SETTLEMENT, METAMORPHOSIS AND RECRUITMENT Burke (1983) defined settlement s the termintion of the pelgic life nd considered it s ehviorl, reversile trit, wheres metmorphosis is developmentl process expressed in morphologicl nd physiologicl chnges, which is irreversile nd occurs only once during the life cycle. Unfortuntely the distinction etween settlement nd metmorphosis hs not een mde this clerly in the relevnt literture. Rodríguez et l. (1993) considered settlement s "the pssge from pelgic wy of life to enthic wy of life" nd distinguished two phses of the settlement process: ehviorl phse of explortion for suitle sustrte nd phse of permnent residence or ttchment to the sustrte, which triggers metmorphosis. Although oth, settlement nd metmorphosis re often interlinked, in some cses they cn e triggered independently (Coon et l. 199). Moreover, in soft sediment hitts Rodriguez's requirement of "permnent residence" in the sustrte cn not pplied since individuls often leve the sediment fter initil settlement s lrve, post-lrve or juveniles (Woodin 1991, Duchêne 24). Additionlly, mny species metmorphose grdully cusing difficulties in ccurtely determing the "event" of metmorphosis in settlement ssys. For exmple, Pectinri koreni (Polychet, Pectinriide) displys n intermedite stge etween mettrochophore lrve nd juveniles, clled ulophore lrve or post-lrve. Post-lrve hve lost mettrochophore ttriutes ut miss some juvenile trits (Thiéut et l. 1998). The term recruitment is usully defined s the entry into the enthic popultion nd lso includes the survivl of the individul up to specific size fter settlement (Frschetti et l. 23). Recruitment is lso used s n umrell term including the relese of lrve into the wter column, their trnsport, plnktonic mortlity, settlement nd post-settlement survivl (Jenkins et l. 1999). In this thesis settlement ssys were crried out to evlute initil lrvl settlement. In this regrd settlement ws considered nd defined s ehviorl, repetle trit. 12

16 GENERAL INTRODUCTION 1.4 POST-SETTLEMENT EVENTS Once settled nd metmorphosed locl disturnces, predtion or differences in the hitt requirements of juvenile nd dults my cuse disloction of juveniles nd dults. These post-settlement processes my susequently chnge initilly estlished species-specific distriution ptterns. The reltive importnce of prend post-settlement processes t the formtion of sptil ptterns differs etween hrd nd soft-sustrtes nd mong species (Stoner 199, Ólfsson et l. 1994, Frschetti et l. 23). Generlly, in soft sediment settlers the limittion of moility is y fr not s strict s in hrd sustrte settlers (Cummings et l. 1995, Whitltch et l. 1998, Snelgrove et l. 1999, Srd et l. 2, Norkko et l. 21, Stocks 22, de Montudouin et l. 23, Hernndez Guevr 24). Studies investigting the ility of juveniles nd dults for ctive settlement choice hve een rre so fr. In settlement experiments with juveniles insted of lrve Woodin (1993, 1995) oserved high selectivity t settlement. Adult polychetes used y Gry (1967) to test the ttrctiveness of different sediments displyed ctive sustrte choice. 1.5 OBJECTIVES The nlysis of sediment cores long trnsects in tidl flts of the Wdden Se (Germny) reveled distinctive ptterns of undnce in spionid polychetes with lrvl development (Stmm 2). The oserved ptterns were neither correlted with sediment chrcteristics such s the silt content nor the orgnic cron nd nitrogen content (Stmm 2). Therefore, I hypothezised tht the distriution ptterns my hve resulted from ctive site selection of lrve triggered y fctors others thn silt, orgnic cron nd nitrogen content. In prticulr, I focused on the role of cteri ssocited with the sediment surfce s meditors in the settlement process. To test this hypothesis, I chose two spionid species with pelgic lrvl development, Polydor cornut Bosc, 182 nd Strelospio enedicti Wester, 1879 nd rised the following questions: Do lrve of P. cornut nd S. enedicti ctively ccept or reject different sediment types? Is there ny ehviorl evidence tht ptchy distriution in the field is the result of lrvl choice? 13

17 GENERAL INTRODUCTION To test this hypothesis, I investigted different lrvl developmentl stges of P. cornut nd S. enedicti for their ility to ctively ccept or reject sediments in still wter lortory ssys. Sediments under investigtion were untreted or sterilized, shed or cid wshed sediment. If lrve, s hypothesized, did ctively select sediment qulity, different preferences would e displyed in the presence of clerly differentited sediment types (Chpter 3). Do lrve respond to signls derived from cteri ssocited with sediment? The min emphsis of this pproch ws to focus on the role of sedimentssocited microil cues s potentil triggers of lrvl settlement in P. cornut nd S. enedicti. Microorgnisms from nturl sediment nd different cteril isoltes from the hitt of dult polychetes were reestlished on sterile sediment to test if different cteril isoltes nd their undnce on the sediment influenced lrvl settlement (Chpter 4). Do lrve detect settlement signls in the wter column or on the sediment surfce? The signl triggering lrvl settlement my e detected y the lrv either on the surfce or in the overlying wter. A series of screening experiments were performed to loclize the settlement cue (Chpter 4). Does the presence of conspecifics influence settlement of lrve? Do the lrve ehve gregriously t settlement? Species specific ptterns my e cused y gregrious lrvl settlement, i.e. lrve prefer to settle in sustrtes with lredy settled conspecifics (Knight- Jones 1953). Therefore the settlement of P. cornut lrve in presence of conspecific dults, juveniles, empty tues nd culture sediment ws investigted nd compred with lrvl settlement in sediment inhited y S. enedicti nd culture sediment of S. enedicti, respectively (Chpter 5). To investigte lrvl settlement in spionid polychetes it ws necessry to develop nd set up spionid cultures under lortory conditions, design nd compre different settlement iossys, study the competence-progression of lrve y temporl correltion of lrvl morphology s well s their ility to settle. These works re outlined in Chpter 2 nd 3. 14

18 GETTING STARTED 2 GETTING STARTED: SELECTION AND CULTURING OF TEST POLYCHAETES GETTING STARTED 2.1 SELECTION OF SUITABLE TEST POLYCHAETES To test lrvl settlement, the mintennce of lortory sed rood cultures is dvntgeous. So fr, lortory studies with lrve of infunl invertertes hve minly een crried out with ivlves (Bchelet et l. 1992, Snelgrove et l. 1993, Turner et l. 1994, Snelgrove et l. 1998, Dunn et l. 1999, Cummings nd Thrush 24) nd polychetes (Jägersten 194, Wilson 1937, Gry 1967, Ch nd Bhud 2). Among the polychetes, specil emphsis hs een devoted to Cpitell sp. I, which in turn hs ecome stndrd ssy species for lortory studies ll over the world (Grssle 198, Cuomo 1985, Duilier 1988, Pechenik nd Cerulli 1991, Biggers nd Lufer 1992, Butmn nd Grssle 1992, Snelgrove et l. 1993, Cohen nd Pechenik 1999, Snelgrove et l. 21, Mrinelli nd Woodin 24, Thiygrjn et l. 25). Spionids, one of the lrgest polychete fmilies, hve een rely investigted concerning lrvl settlement so fr. Among spionid polychetes only Strelospio enedicti ws used in lrvl settlement ssys. Here, the impct of pollutnts (endosulfn nd PAHs) on lrvl of S. enedicti ws tested (Chndler nd Scott 1991, Chndler et l. 1997). Spionids seemed to e suitle orgnisms to study lrvl sustrte selection, ecuse mny of them disperse vi plnktonic lrve (Rouse nd Pleijel 21) nd species specific distriution ptterns of spionids commonly occur in the field (e.g. Stmm 2). Mny spionid polychetes re prticulrly esy to otin due to their high undnce in ech nd esturine hitts. They live in urrows or tues (Rouse nd Pleijel 21) with typicl undnces of severl thousnd individuls per squre meter (Zjc 1991). Spionids re recognized y pir of elongted grooved plps extending from the hed. Spionid lrve re generlly rooded in the mternl tue until they develop numer of segments nd re relesed into the plnkton (Blke nd Arnovsky 1999). The following criteri were used to select suitle test species mong spionids: 1) occurrence in the Wdden Se (logistics), 2) development vi plnktotrophic lrve to ensure nturl dispersion potentil nd ecologicl relevnce of the ssy, 15

19 GETTING STARTED 3) cultivle dults nd lrve in the lortory, with relistic purpose of yerround lrvl supply in the lortory culture, 4) known nd descried distriution ptterns in the field, 5) worldwide occurrence to ensure dptility of the ssy. In the North Se, c. 6 spionid species elonging to 15 different gener were reported (listed e.g. in the dtse of the North Se Benthos Project 2 ). Initilly criteri 2 nd 3 were pplied to nrrow down the numer of possile cndidtes followed y evlution of criteri 4 nd 5. A memer of the Polydor group (the species-richest spionid genus in the North Se), Polydor cornut hs een previously cultured in the lortory y Anger et l. (1986) nd Irvine nd Mrtindle (1999). In oth studies continuous supply of plnktotrophic lrve ws chieved t optiml culture conditions. P. cornut is widely undnt in muddy snds of tidl flts in the temperte zone (Hrtmnn-Schröder 1996) nd occurs in the Wdden Se (Dnkers 1981). Stmm (2) reported distinctive ptterns of undnce of P. cornut. Similr oservtion of ptchy distriution of P. cornut ws lso reported in the Atlntic t Estern Pssge, Hlifx, Cnd (Snelgrove et l. 1999). P. cornut ws collected during low tide in mudflts ner Hooksiel (53 38' 31" N, 8 4' 55" E, Wdden Se, Germny) in August 23. The chrcteristic verticl living tues constructed y P. cornut helped to loclize dult smpling sites. Suitle lortory culture methods were developed or dopted from Irvine nd Mrtindle (1999), respectively. Following estlishment of the culture y settlement of ll newly htched lrve continuous lrvl supply ws chieved t the eginning of 24. Detiled informtion on species chrcteristics nd culture conditions re given in Chpter 2.4. Strelospio enedicti (Wester 1879), the only spionid lredy used in lrvl settlement ssys prior to this study (Chndler nd Scott 1991, Chndler et l. 1997) ws reported to e cultivle in the lortory (Lewin 1986, Bridgess nd Heppell 1996, Chndler et l. 1997). Erlier cultivtion ttempts of this species were minly motivted y the intrspecific vrition in its developmentl mode (poecilogony), which predestines S. enedicti for studies of different developmentl nd mternl effects. Although S. enedicti occurs in the Wdden Se, it is less undnt thn Polydor cornut. In the first yer of my study, S. enedicti could not e found t the smpling site in Hooksiel. However in summer 24, the species occurred in Hooksiel nd polychete culture ws 16

20 GETTING STARTED successfully estlished in the lortory. Only femles with plnktotrophic lrvl development mode fulfilled selection criterion 2. Therefore, femles of S. enedicti were preliminry cultured to scertin plnktotrophic development of lrvl roods. Due to the experiences in culture techniques of P. cornut, cultures of S. enedicti were estlished quickly. Sufficient lrvl supply for iossy purposes ws chieved in Septemer 24. Detiled species nd culture informtion is given in Chpter POLYDORA CORNUTA Polydor cornut Bosc, 182 (formerly lso Polydor ligni Wester, 188) is memer of the polydorid species complex (Blke 1969). Adults re up to 32 mm long, 1.5 mm wide nd hve up to 9 setigers (Rdshevsky 25). It colonizes sndy nd muddy hitts. P. cornut tolertes high orgnic contents nd is n erly colonizer of defunted sites (Duer 1984). It uilds U-shped tues up to 4 mm in length nd 1 mm width (Hrtmnn-Schröder 1996). The tues protrude from the snd nd form funnel-like vents esily visile y the nked eye. P. cornut feeds with plps s long s setigers on plnktonic orgnisms nd detritus either from wter or from the sediment surfce (Stmm 2). While recently settled juveniles of P. cornut lmost exclusively show suspension feeding, lrger juveniles nd dults minly feed on deposit unless the flow exceeds ~1 cm/s (Hentschel 1999). P. cornut reproduces y internl fertiliztion. Femles deposit sphericl eggs ( 1-11 μm) in cpsules. Egg cpsules re ttched to the inside wll of the tue nd contin up to 3 eggs. The eggs in roods develop synchronously into lrve, which enter the plnkton with 3-4 setigers nd pproximtely μm length (Blke 1969, Hrtmnn-Schröder 1996, Rdshevsky 25). Relesed lrve spend 2-3 weeks in the wter (Anger et l. 1986) feeding on plnkton nd swimming ctively. A mximum of 85, lrve per cuic meter se wter ws reported y Orth (1971). New segments develop one y one in the growth zone in front of the disclike pygidium. 1,2 1,3 μm long setiger lrve re le to settle nd undergo metmorphosis. Mximum length of lrve found in plnkton ws out 1,7 μm long with 25 setigers (Rdshevsky 25). Additionlly, Mcky nd Gison (1999) reported delphophgic (feeding on the nurse eggs) lrvl development in P. cornut. Adelphophgic femles produced roods in which 17

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