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Einnischungsmechanismen Bei Rotwidderchen (Lepidoptera: Zygaenidae) Auf Kalkmagerrasen Der Schwäbischen Alb (Baden-Württemberg)

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Einnischungsmechanismen Bei Rotwidderchen (Lepidoptera: Zygaenidae) Auf Kalkmagerrasen Der Schwäbischen Alb (Baden-Württemberg) EINNISCHUNGSMECHANISMEN BEI ROTWIDDERCHEN (LEPIDOPTERA: ZYGAENIDAE) AUF KALKMAGERRASEN DER SCHWÄBISCHEN ALB (BADEN-WÜRTTEMBERG) Dissertation zur Erlangung des Doktorgrades Dr. rer. nat. der Fakultät für Naturwissenschaften der Universität Ulm vorgelegt von WOLFGANG WAGNER aus MEMMINGEN Ulm 2002 Die vorliegende Arbeit wurde an der Abteilung für Experimentelle Ökologie (Biologie III) der Universität Ulm angefertigt. Referentin: Prof. Dr. E. KALKO Abteilung für Experimentelle Ökologie (Bio III) Universität Ulm Koreferent: Prof. Dr. G. GOTTSBERGER Abteilung für Systematische Botanik und Ökologie (Bio V) Universität Ulm Dekan: Prof. Dr. WITSCHEL Inhaltsverzeichnis 1. Einleitung - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 1 2. Material und Methoden 2.1 Untersuchungsgebiet 2.1.1 Geologie und Klima - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2 2.1.2 Lage und Fläche der Untersuchungsgebiete - - - - - - - - - - - - - - - - - - - - - - - 3 2.2 Untersuchte Arten - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 7 2.3 Erfassungsmethodik und Untersuchungszeitraum 2.3.1 Raupen - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 8 2.3.2 Falter - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 9 2.3.3 Nektarpflanzen - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 10 2.3.4 Gebietsanalyse - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 11 2.3.5 Vegetation - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 12 2.3.6 Markierungsexperiment - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 12 2.3.7 Andere Tiergruppen - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 13 2.3.8 Parasitoide - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 13 2.4 Zuchtmethoden - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 13 2.5 Nomenklatur und Determination - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 14 2.6 Begleit-CD - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 15 3. Ergebnisse 3.1 Witterungsverlauf im Untersuchungszeitraum - - - - - - - - - - - - - - - - - - - - - 15 3.2 Datengrundlage - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 17 3.3 Gesamtphänologie 3.3.1 Zygaena minos und Z. purpuralis - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 18 3.3.2 Zygaena loti - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 22 3.3.3 Zygaena viciae - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 23 3.3.4 Zygaena lonicerae und Z. ephialtes - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 24 3.3.5 Zygaena carniolica - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 24 3.3.6 Zygaena transalpina - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 26 3.3.7 Zygaena filipendulae - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 27 3.4 Abundanz und Dominanz in den einzelnen Gebieten 3.4.1 Untersuchungsgebiete 1998-2001 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 28 3.4.2 Untersuchungsgebiete 2000-2001 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 33 3.5 Gebietsanalyse 3.5.1 Allgemeine Faktoren - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 36 3.5.2 Raupennahrungspflanzen - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 42 3.6 Weitere Aspekte zur Larvalökologie 3.6.1 Verhalten - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 47 3.6.2 Larvalhabitat - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 48 3.7 Zuchtergebnisse 3.7.1 Zygaena filipendulae - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 50 3.7.2 Zygaena transalpina - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 52 3.7.3 Zygaena loti - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 52 3.7.4 Zygaena viciae, Z. minos, Z. purpuralis und Z. carniolica - - - - - - - - - - - - - - 53 3.7.5 Kreuzungsversuch von Z. minos und Z. purpuralis - - - - - - - - - - - - - - - - - - 54 3.8. Blütenökologie 3.8.1 Nektarpflanzenspektrum- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 55 3.8.2 Blühphänologie und Widderchenabundanz - - - - - - - - - - - - - - - - - - - - - - - 58 3.8.3 Konkurrenz mit anderen Blütenbesuchern - - - - - - - - - - - - - - - - - - - - - - - - 63 3.9 Weitere Aspekte zur Imaginalökologie - - - - - - - - - - - - - - - - - - - - - - - - - - - 64 3.10 Markierungsexperiment im Lindletal 3.10.1 Populationsdynamik bei Z. minos und Z. purpuralis - - - - - - - - - - - - - - - - - 65 3.10.2 Mobilität bei Z. minos und Z. purpuralis - - - - - - - - - - - - - - - - - - - - - - - - - 68 3.10.3 Zygaena filipendulae - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 69 3.11 Parasitoide und Feinde - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 70 3.12 Morphologische Besonderheiten - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 71 3.13 Einnischung der Grünzygaene Adscita geryon - - - - - - - - - - - - - - - - - - - - - 72 3.14 Abschätzung der Genauigkeit der eingesetzten Transektmethode - - - - - - - 74 4. Diskussion 4.1 Methoden 4.1.1 Erfassungsmethoden - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 75 4.1.2 Zuchtmethoden - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 77 4.1.3 Markierungsexperiment - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 78 4.1.4 Gebietsanalyse - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 78 4.2 Einnischungsphänomene 4.2.1 Phänologie - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 79 4.2.2 Nahrungsbiologie - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 85 4.2.3 Habitatansprüche - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 87 4.3 Mechanismus der zeitlichen und räumlichen Einnischungsphänomene 4.3.1 Notwendige Entwicklungszeit der Arten - - - - - - - - - - - - - - - - - - - - - - - - - - 91 4.3.2 Phänologische Folgen der unterschiedlichen Entwicklungszeit - - - - - - - - - - 94 4.3.3 Auswirkungen der Entwicklungszeit auf die räumliche Verteilung - - - - - - - 96 4.4 Dichteregulatoren 4.4.1 Nektar- und Raupennahrungspflanzen - - - - - - - - - - - - - - - - - - - - - - - - - - 97 4.4.2 Biotopfaktoren - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 99 4.4.3 Witterung - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 101 4.4.4 Prädatoren und Parasitoide - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 102 4.4.5 Konkurrenz - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 103 4.4.6 Folgen für die Abundanz - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 106 4.5 Populationsdynamik und Mobilität - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 107 4.6 Synoptische Betrachtung - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 110 4.7 Folgen für den Naturschutz 4.7.1 Gefährdung - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 111 4.7.2 Schutzerfordernisse - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 113 5. Zusammenfassung/Summary - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 115 6. Farbtafeln - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 120 7. Literaturverzeichnis - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 131 8. Anhang 8.1 Beschreibung der Einzelgebiete mit Flächenunterteilung - - - - - - - - - - - - - - 136 8.2 Tabellen (Faktoren der Gebietsanalyse, Vegetationsaufnahmen Z. minos) - - 141 Abkürzungen in der PDF-Datei M: Männchen (Singular) MM: Männchen (Plural) W, WW: Weibchen 1. Einleitung Die Rotwidderchen sind eine phänotypisch und verhaltensbiologisch weitgehend einheitliche Schmetterlingsgruppe aus der Familie Zygaenidae. Ihr tagaktives Verhalten, ihre auffällige schwarz-rote Warnfärbung (aposematisches Muster) und ihr auf Magerrasen meist abundantes Auftreten machen sie zu auffälligen Erscheinungen der mitteleuropäischen Fauna. Alle Arten können aus mit der Nahrung aufgenommenen Substanzen oder de novo Cyanoglukoside synthetisieren (FRANZL et al. 1986), die unter anderem in Wehrsekretbehältern gespeichert werden und den Tieren in jedem Stadium einen Schutz vor zumindest vertebraten Prädatoren wie beispielsweise Vögeln bieten (RAMMERT 1992). Aufgrund ihrer leichten Erfaßbarkeit eignen sich die Rotwidderchen gut für popula- tionsökologische Studien zur Erforschung von Populationsdynamik und Ausbreitungsme- chanismen. Darüber hinaus
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