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Coleoptera, Staphylinidae) Ökomorphologische Diversität und Funktion des Klebfangapparates mitteleuropäischer Stenus-Arten (Coleoptera, Staphylinidae) Dissertation der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) vorgelegt von Lars Koerner aus Berlin Tübingen 2020 Gedruckt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen. Tag der mündlichen Qualifikation: 15.07.2020 Dekan: Prof. Dr. Wolfgang Rosenstiel 1. Berichterstatter: Prof. Dr. O. Betz 2. Berichterstatter: PD Dr. Michael Heethoff Inhalt Übersicht über Publikationen sowie Anteil der Autoren i-ii Inhalt 1 Zusammenfassung 3 Abstract 7 1 Einleitung 12 1.1 Stand der Wissenschaft bezüglich des Klebfangapparates der 12 Gattung Stenus Latreille, 1797 1.2 Fragestellung und Zielsetzung der Dissertation 15 1.2.1 Untersuchung des Klebfangapparates der Gattung Stenus 15 1.2.2 Phylogenie 17 2 Ergebnisse und Diskussion 20 2.1 Vergleichende Untersuchungen zum Mechanismus des 20 Klebfangapparates der Kurzflügelkäfergattung Stenus Latreille, 1797 (Coleoptera, Staphylinidae) 2.1.1 Bestimmung der beim Klebfangmechanismus erzeugten Kräfte 21 2.1.2 Zusammenhang zwischen der Morphologie der Haftpolster, der 22 resultierenden Haftkraft und des Beutefangerfolges 2.1.2.1 Morphologie der Haftpolster 22 2.1.2.2 Vergleich von Druck- und Adhäsionskräften 23 2.1.2.3 Vergleich mit anderen biologischen Haftsystemen 28 2.1.2.4 Beutefang 30 2.1.2.5 Ökomorphologie - Zusammenhang zwischen Morphologie, 31 Beutefangverhalten und Leistungsfähigkeit des Fangapparates 2.1.3 Einfluss der Oberflächenrauheit und -energie auf die 36 resultierende Haftkraft 2.1.4 Funktionelle Grundprinzipien des Stenus-Haftsystems 42 2.1.5 Die Haftpolster der Gattung Stenus als Träger von Bakterien 46 2.2 Molekulare Phylogenie der Kurzflügelkäfergattungen Stenus 48 Latreille, 1797 und Dianous Leach, 1819 (Coleoptera, Staphylinidae, Steninae) Literatur 57 1 Inhalt Danksagung 66 Lebenslauf 67 Publikationen 70 PUBLIKATION I Functional morphology and adhesive 70 performance of the stick-capture apparatus of the rove beetles Stenus spp. (Coleoptera, Staphylinidae) PUBLIKATION II Adhesive performance of the stick-capture 82 apparatus of rove beetles of the genus Stenus (Coleoptera, Staphylinidae) toward various surfaces PUBLIKATION III Divergent morphologies of adhesive predatory 92 mouthparts of Stenus species (Coleoptera, Staphylinidae) explain differences in adhesive performance and resulting prey-capture success PUBLIKATION IV The labial adhesive pads of rove beetles of the 112 genus Stenus (Coleoptera: Staphylinidae) as carriers of bacteria PUBLIKATION VI Loss of the sticky harpoon—COI sequences 122 indicate paraphyly of Stenus with respect to Dianous (Staphylinidae, Steninae). 2 Übersicht über Publikationen (vollständige Publikationsliste siehe Kapitel 10 „Lebenslauf“) sowie Anteil der Autoren: Publikationen als Erstautor Publikation I: Functional morphology and adhesive performance of the stick-capture apparatus of the rove beetles Stenus spp. (Coleoptera, Staphylinidae) Lars Koerner, Stanislav N Gorb, Oliver Betz (2012). Zoology 115: 117–127. Publikation II Adhesive performance of the stick-capture apparatus of rove beetles of the genus Stenus (Coleoptera, Staphylinidae) toward various surfaces Lars Koerner, Stanislav N Gorb, Oliver Betz (2012). Journal of Insect Physiology 58: 155– 163. Publikation III Divergent morphologies of adhesive predatory mouthparts of Stenus species (Coleoptera, Staphylinidae) explain differences in adhesive performance and resulting prey-capture success Lars Koerner, László Zsolt Garamszegi, Michael Heethoff, Oliver Betz (2017). Zoological Journal of the Linnean Society, 181: 500–518. Publikation IV The labial adhesive pads of rove beetles of the genus Stenus (Coleoptera: Staphylinidae) as carriers of bacteria Lars Koerner, Volkmar Braun, Oliver Betz (2016). Entomologia Generalis 36: 33–41. Publikation V Loss of the sticky harpoon—COI sequences indicate paraphyly of Stenus with respect to Dianous (Staphylinidae, Steninae) Lars Koerner, Michael Laumann, Oliver Betz, Michael Heethoff (2013). Zoologischer Anzeiger 252: 337–347. i Anteil der Autoren an den vorgelegten Publikationen: Lars Koerner: Design und Durchführung der Experimente und Studien, Feldprobenahme, Datenerfassung und -analyse, Vorbereitung und Verfassen der Manuskripte zur Veröffentlichung Oliver Betz: Betreuung der gesamten Arbeit Stanislav N. Gorb: Mitarbeit bei der Versuchsplanung, methodische Unterstützung, Diskussion der Manuskripte (Publikationen I und II) Michael Heethoff: Methodische Unterstützung und Beteiligung an der Versuchsplanung und Datenanalyse, Diskussion der Manuskripte (Publikation III und V) Michael Laumann: Methodische Unterstützung und Beteiligung an der Versuchsplanung und Datenanalyse, Diskussion des Manuskriptes (Publikation V) László Zsolt Garamszegi: Methodische Unterstützung und Beteiligung an der Datenanalyse und Diskussion des Manuskriptes (Publikation III) Volkmar Braun: Methodische Unterstützung und Beteiligung an der Datenanalyse und Diskussion des Manuskriptes (Publikation IV) ii Zusammenfassung Zusammenfassung Mit mehr als 3000 beschriebenen Arten bildet die Gattung Stenus die artenreichste Tiergattung überhaupt (Puthz, pers. Mitt.). Innerhalb der Familie der Staphylinidae gelten sie als die einzigen Vertreter, die ein stabförmig verlängertes Fanglabium aufweisen, das durch Hämolymphdruck innerhalb weniger Millisekunden hervorgeschnellt werden kann. Bleibt ein Beutetier an den zu Haftpolstern umgebildeten Paraglossen haften, wird die Beute durch Retraktion des Labiums zwischen die Mandibeln gebracht und kann gefressen werden. Die beim Beutefang auftretenden Haftkräfte wurden bislang nur theoretisch bestimmt (Kölsch 2000), während die beim Ausschleudern des Labiums auftretenden Druckkräfte noch völlig unbekannt sind. Zentraler Teil der vorliegenden Arbeit ist die Bestimmung der über den Klebfangmechanismus erzeugten Druck- und Zugkraft. Dazu wurde eine Messmethode, mit der es erstmals möglich ist, den genauen zeitlichen Verlauf der entwickelten Kräfte zu verfolgen und statistisch auszuwerten. Die Tiere sind visuell orientiert und "schießen" mit ihren Klebzungen auf einen sich bewegenden Insektennadelkopf, der mit einem Kraftmessgerät verbunden ist (Publikationen I-III). Umfangreiche Studien von Betz (1996) an verschiedenen Stenus-Arten zeigten, dass größere und strukturell komplexere Haftpolster zu einem erhöhtem Fangerfolg führen. Diese Schlussfolgerung wird durch die neuen Experimente unterstützt und mit der Integration der Performanzmessungen kann nun erstmalig ein direkter Zusammenhang zwischen der Morphologie der Haftpolster, deren Haftperformanz und dem Beutefangerfolg hergestellt werden. Die während des Klebfanges entstehenden Druck- (beim Auftreffen der Klebpolster auf eine Beuteattrappe) und Adhäsionskräfte (beim Rückzug der Klebpolster von der Attrappe) wurden bei 14 mitteleuropäischen Vertretern der Gattung Stenus bestimmt (Publikation III). Mit Werten zwischen 0.28 mN bei S. morio und 1.08 mN bei S. bimaculatus zeigen vor allem die Adhäsionskräfte signifikante interspezifische Unterschiede, welche auf die unterschiedliche Größe und die Komplexität der Haftpolster (d.h. der Anzahl an Hafthaaren und Haftkontakten) zurückzuführen sind (Publikation III). Die durchschnittlichen Druckkräfte während des Fangschlages erreichten Werte zwischen 0.05 mN bei S. biguttatus und 0.18 mN bei S. juno und waren somit um eine Größenordnung niedriger als die Adhäsionskräfte. Außerdem konnte gezeigt werden, dass ein signifikanter Zusammenhang zwischen Druckkraft und Zug- (Adhäsions-)kraft besteht, d.h., je größer die Kraft mit der die Klebpolster auf ein Beuteobjekt auftreffen, desto größer ist auch die Adhäsion zwischen Klebpolstern und Beute. Dieses Ergebnis weist auf einen druckempfindlichen Haftmechanismus bei Stenus hin. Die phylogenetisch-vergleichenden Analysen zeigten weiterhin, dass sich die Zugfestigkeit (Haftkraft pro Haftposterfläche) mit zunehmender Haftpolsterfläche verringert, sodass Stenus-Arten mit 3 Zusammenfassung kleineren Haftpolstern zwar geringere absolute Adhäsionskräfte erzeugen, aber insgesamt höhere Zugfestigkeiten erreichen und somit adhäsiv effizientere Haftpolster besitzen. Die berechneten Werte für die Zugfestigkeit lagen dabei zwischen 51.9 kPa bei S. bimaculatus und 153.2 kPa bei S. humilis (Publikation I, III). Die negative Korrelation von Zugfestigkeit und Haftpolsterfläche zeigt, dass das Haftsekret, welches während des Fangschlages über die Fläche des Haftpolsters verteilt wird, einen sehr großen Beitrag zur Haftfähigkeit des Fangapparates leistet. Dadurch sind auch Arten mit kleineren Haftpolstern befähigt, eine relativ gute Haftperformanz zu erreichen. Die während des Fangschlages auftretenden Kräfte können in Bezug mit Daten über den Fangerfolg bei verschiedenen Beutegrößen gesetzt werden. Bei diesen Fangversuchen wurden verschiedene einheimische Stenus-Arten mit Springschwänzen (Heteromurus nitidus) verschiedener Größen- (bzw. Gewichts-)klassen konfrontiert und die Anwendungshäufigkeiten beider Fangtechniken (Labium/Mandibel) sowie der Fangerfolg analysiert. Die Analysen zeigten, dass eine höhere Adhäsionskraft der Klebpolster zu einem höheren Fangerfolg gegenüber Collembolen der Art H. nitidus führt. Dieser Effekt war beim Fang größerer Collembolen wesentlich stärker ausgeprägt (p < 0.001) als beim Fang kleiner Collembolen
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