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Homoptera, Delphacidae) Biological and taxonomie differentiation inth e Ribautodelphaxcollinus complex (Homoptera, Delphacidae) CENTRALE LANDBOUWCATALOGUS 0000 0234 7124 Promotor: dr. ir. R. H. Cobben hoogleraar in de diertaxonomie \W C. F. M. den Bieman Biological and taxonomie differentiation in the Ribautodelphax collinus complex (Homoptera, Delphacidae) Proefschrift ter verkrijging van de graad van doctor in de landbouwwetenschappen, op gezag van de rector magnificus, dr. C. C. Oosterlee, in het openbaar te verdedigen op woensdag 28 oktober 1987 des namiddags te vier uur in de aula van de Landbouwuniversiteit te Wageningen. BIBLIOTHEEK •^IR/UNIVERSITEIT svo-- ^bcM ±( Omslagontwerp: F.J . J. von Planta Druk: Grafisch bedrijf Ponsen &Looije n - Wageningen fS^Jofl^, \R\ Stellingen 1. De redenering dat onderzoek van parthenogenetische dieren leidt tot inzicht in de evolutie van seksuele voortplanting is discutabel, omdat, voor zover bekend, alle parthenogeneti­ sche dieren ontstaan zijn uit seksuele vormen. Lynch, M. (1984). Quart. Rev. Biol. 59:257-290 . 2. Selectieve partnerkeuze speelt een belangrijke rol bij de handhaving van populaties, die bestaan uit biseksuele en pseudogame vormen. Dit proefschrift. 3. Akoestische signalen van vrouwtjes krijgen ten onrechte weinig aandacht als soortspecifiek kenmerk bij Auchenorrhyncha. Dit proefschrift. 4. De grote intraspecifieke variatie in akoestische signalen van vrouwtjes van Ribautodelphax pungens en R. imitantoides is in tegenspraak met Patersons stelling dat 'soortherkennings- signalen' gebufferd zijn tegen snelle veranderingen. Paterson, H. E. H. (1985). Transvaal Mus. Mon. 4:21-29 . 5. Pseudogamie is een vorm van parthénogenèse, die voorkomt dat de parthenogenetische vorm de seksuele oudersoort(en) verdringt. 6. Het belang van interspecifieke concurrentie tussen fytofagen wordt vaak overschat. Jermy, T. (1985). Z. zool. Syst. Evolut.-forsch. 23: 275-285. 7. De bepaling van de volgorde van kenmerkverandering van een 'multistate' electroforetisch locus mag niet gebaseerd worden op de relatieve verandering in electroforetische mobiliteit. Green, D. M. (1986). Syst. Zool. 35:283-296 . 8. Discussies over het ontstaan van polyploide dieren zijn gefixeerd op een hybride oorsprong en houden te weinig rekening met een mogelijke autopolyploide oorsprong. Bullini, L. (1985). Boll. Zool. 52: 121-137. 9. Seksuele voortplanting is ook van belang voor het voortbestaan van parthenogenetische vormen, omdat hierdoor de kansen op genetische variabiliteit van polyfyletische parthe- nogeneten vergroot worden. Vrijenhoek, R. C. (1984). In: Population biology and evolution, 217-231. Springer Verlag. Dit proefschrift. 10. De lange duur van taxonomische promotieonderzoeken en het grote aantal publikaties per onderzoek illusteren de complexiteit van de onderzochte materie. Storms, J. J. H. (1986). Vakbl. Biol. 66:405-407 . 11. De geringe hoeveelheid geld zowel nationaal als internationaal beschikbaar voor de taxon­ omie én de omvang van haar taak én haar economisch belang. Wilson, E. O. (1986). Science 230: 1227. Andersen, A. (1987). Nature 329:6 . 12. De neiging om het verzamelen van insekten aan banden te leggen via de nieuwe Natuur­ beschermingswet leidt tot een verminderende belangstelling voor deze diergroep en schaadt daardoor hun bescherming. 13. Bij de discussie over de kosten van kinderopvang wordt te weinig rekening gehouden met de kosten van de traditionele kinderopvang. 14. Knarsentanden zal verstommen als de tandarts uit het ziekenfondspakket verdwijnt. Proefschrift van C. F. M. den Bieman. Biological and taxonomie differentiation in the Ribautodelphax collinus complex (Homoptera, Delphacidae). Wageningen, 28 oktober 1987. Aan Marianne Aan mijn ouders VOORWOORD Ik wil graag allen danken, die bijgedragen hebben aan het onderzoek en de totstandkoming van dit proefschrift. Op de eerste plaats dank ik mijn promotor René Cobben, die het initiatief nam voor het projekt. Ik dank je voor de vrijheid, die ik kreeg bij de uitvoering van het onderzoek. Jouw hulp en interesse hebben het resultaat mede bepaald. Kees Booij introduceerde mij in de Delphacidae en ik ben je dankbaar voor de vele diskussies en je hulp bij de start van het onderzoek. Peter de Vrijer dank ik voor zijn bijdrage aan de opzet van de experimenten en zijn kommentaar op de artikelen. Ik dank Theodoor Heijerman, Ton de Winter, Christiaan Kooyman en Adriaan Guldemond voor hun waardevolle bijdragen en suggesties en de goede werksfeer. Ebbe Schumacher taught me the eleetrophoretic technique and our cooperation resulted in chapter 6. Maarten Drost, Margreet Bloemhof en Ton de Winter hebben met veel inzet en kreativiteit hun doktoraalonderzoek verricht en jullie resultaten hebben in belangrijke mate bijgedragen aan dit proefschrift. Wouter Tigges was een steunpilaar want zonder jouw inzet voor de kweken zouden vele experimenten niet mogelijk zijn geweest. Julius Pattiapon dank ik voor zijn hulp bij het uitwerken van de eindeloze reeksen geluidssignalen. The animated discussions with Sakis Drosopoulos on the origin of triploid planthoppers stimulated my research. You made our excursion to Greece a success. The investigations of the Mediterranean Delphaaid fauna of Manfred Asche, Reinhard Remane and Hannelore Hoch have been very important to my study. You generously shared your experience and materials with me. Veel medewerkers van de vakgroep Entomologie en van de Gekombineerde Diensten hebben mij geholpen. In het bizonder wil ik Truus de Vries danken voor het typen van het volledige proefschrift en Jack Brangert en Hein Visser voor de vele foto's. De stichting "Het Nationale Park De Hoge Veluwe" en Staatsbosbeheer Limburg verleenden toestemming voor het doen van onderzoek in hun terreinen. Mijn onderzoek is mede mogelijk gemaakt door een subsidie van de Stichting voor Biologisch Onderzoek in "Nederland (BION), die gesubsidieerd wordt door de Nederlandse Organisatie voor Zuiver Wetenschappelijk Onderzoek (ZWO). De Uyttenboogaart-Eliasen Stichting verleende een reiesubsidie. Ik ben Marianne Hamdankbaar voor de steun en stimulans tijdens dit onderzoek. Zonder jouw hulp zou het veldwerk, de basis van mijn onderzoek, nooit zo suksesvol hebben kunnen verlopen. INHOUD- CONTENT S Inleiding 1 Host-plant relations in the planthopper genus Ribautodelphax (Homoptera, Delphacidae). Ecol.Ent . 12:163-172 . 3 2 Acoustic differentiation and variation in planthoppers of the genus Ribautodelphax (Homoptera,Delphacidae) .Neth . J. Zool.36 :461-480 . 13 3 Variability in female calling signals inmixe d populations ofpseudoga - mous forms andbisexua l Ribautodelphax species (Homoptera,Delphacidae) . Neth. J. Zool. (accepted). 35 4 Hybridization studies in the planthopper genus Ribautodelphax (Homoptera Delphacidae). Genetica (accepted). 53 5 Karyotypic variation inbisexua l species and pseudogamous forms of the planthopper genus Ribautodelphax (Homoptera,Delphacidae) . Genetica (accepted). 73 6 Allozyme polymorphism in planthoppers of the genus Ribautodelphax (Homoptera,Delphacidae) ,an d the origin of the pseudogamous triploid forms, (submitted). 89 7 Coexistence of pseudogamous and bisexual planthoppers of the genus Ribautodelphax (Homoptera,Delphacidae) .(submitted) . 107 8 Taxonomie evaluation of the Ribautodelphax collinus complex (Homoptera, Delphacidae). (manuscript). 121 Samenvatting 157 Curriculum vitae 163 INLEIDING Dit proefschrift maakt deel uit van het biosystema tisc h onderzoek, dat verricht wordt door de sektie Diertaxonomie van de vakgroep Entomologie, Landbouwuniversiteit Wageningen. Belangrijke aandachtspunten in dit onderzoek zijn o.a. de karakterisering van nauwverwante soorten op basis van biologisch zinvolle kriteria en de rekonstruktie van fylogenetische relaties binnen en tussen soortskomplexen. De resultaten van dit onderzoek vormen een bijdrage aan diskussies over het begrip "soort"e n over het verloopva n soortsvormingsprocessen. Uit de familie van de spoorcikaden (Delphacidae) zijn voor dit biosystematisch onderzoek enkele genera gekozen, die een reeks met een toenemende mate van interspecifieke differentiatie vormen. Het Ribautodelphax collinus komplex vertegenwoordigt daarbijee n basaal nivo van differentiatie. Bij de aanvang van dit onderzoek omvatte het R^_ collinus komplex in Europa eenvierta l beschreven soorten, die slechts een geringe mate van morfologische divergentie vertonen.Variabilitei t inbepaald e diagnostische kenmerken leidde tot twijfel over de taxonomische status van deze soorten. Faunistisch onderzoek in vnl. het Mediterrane gebied resulteerde ind e ontdekking van een aantal nieuwe vormen, waarvan de taxonomische status onduidelijk was. Dit onderzoek draagt bij aan een oplossing van deze taxonomische verwarring. De biosystematiek baseert zich op het "biologisch soortsconcept": soorten zijn eenheden, die in het veld geheel of in belangrijke mate reproduktief geisoleerd zijn van elkaar. Dankzij deze reproduktieve isolatie kunnen soorten verschillen ontwikkelen en blijven de verschillen gehandhaafd. Diverse mechanismen kunnen bijdragen aan het ontstaan van isolatie tussen soorten bv.verschille n inherkenning ssignalen , in habitats en in genitaalbouw. Om de mate van differentiatie tussen de taxa uit het R^ collinus komplex te kunnen bepalen zijn deze taxavergeleke n wat betreft: ekologie m.n. waardplantrelaties; geluidssignalen; kruisbaarheid; karyologie; allozymen en morfologie. Voor een evaluatie van de verschillen is het 1 belangrijk ook de variatie binnen en tussen conspecifieke populaties te bestuderen en daarom is dit onderzoek
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    Incipient non-adaptive radiation by founder effect? Oliarus polyphemus Fennah, 1973 – a subterranean model case. (Hemiptera: Fulgoromorpha: Cixiidae) Dissertation zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) im Fach Biologie eingereicht an der Mathematisch-Naturwissenschaftlichen Fakultät I der Humboldt-Universität zu Berlin von Diplom-Biologe Andreas Wessel geb. 30.11.1973 in Berlin Präsident der Humboldt-Universität zu Berlin Prof. Dr. Christoph Markschies Dekan der Mathematisch-Naturwissenschaftlichen Fakultät I Prof. Dr. Lutz-Helmut Schön Gutachter/innen: 1. Prof. Dr. Hannelore Hoch 2. Prof. Dr. Dr. h.c. mult. Günter Tembrock 3. Prof. Dr. Kenneth Y. Kaneshiro Tag der mündlichen Prüfung: 20. Februar 2009 Incipient non-adaptive radiation by founder effect? Oliarus polyphemus Fennah, 1973 – a subterranean model case. (Hemiptera: Fulgoromorpha: Cixiidae) Doctoral Thesis by Andreas Wessel Humboldt University Berlin 2008 Dedicated to Francis G. Howarth, godfather of Hawai'ian cave ecosystems, and to the late Hampton L. Carson, who inspired modern population thinking. Ua mau ke ea o ka aina i ka pono. Zusammenfassung Die vorliegende Arbeit hat sich zum Ziel gesetzt, den Populationskomplex der hawai’ischen Höhlenzikade Oliarus polyphemus als Modellsystem für das Stu- dium schneller Artenbildungsprozesse zu erschließen. Dazu wurde ein theoretischer Rahmen aus Konzepten und daraus abgeleiteten Hypothesen zur Interpretation be- kannter Fakten und Erhebung neuer Daten entwickelt. Im Laufe der Studie wurde zur Erfassung geografischer Muster ein GIS (Geographical Information System) erstellt, das durch Einbeziehung der historischen Geologie eine präzise zeitliche Einordnung von Prozessen der Habitatsukzession erlaubt. Die Muster der biologi- schen Differenzierung der Populationen wurden durch morphometrische, etho- metrische (bioakustische) und molekulargenetische Methoden erfasst.
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