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Lsffiss999&9S! sta^/, ©Zoologische Staatssammlung München;download:i http://www.biodiversitylibrary.org/; www.biologiezentrum.at O # im '^öncVie^^ SPIXIANA W ^ Zeitschrift für Zoologie LSffiSS999&9S! Konvergente Farbmusterentwicklungen bei Tagfaltern Freilanduntersuchungen in Asien, Afrika und Lateinamerika IXIANA • Supplement 2i • München, Ol. November 1996 • ISSN 0177-7424 • ISBN 3-931516-11-3 ©Zoologische Staatssammlung München;download:SPIXIAHA http://www.biodiversitylibrary.org/; www.biologiezentrum.at ZEITSCHRIFT FÜR ZOOLOGIE herausgegeben von der ZOOLOGISCHEN STAATSSAMMLUNG MÜNCHEN SPIXI ANA bringt Originalarbeiten aus dem Gesamtgebiet derZoologischen Systematik mit Schwerpunkten in Morphologie, Phylogenie, Tiergeographie und Ökologie. Manuskripte werden in Deutsch, Englisch oder Französisch angenommen. Pro Jahr erscheint ein Band zu drei Heften. Umfangreiche Beiträge können in Supplementbänden herausgegeben werden. SPIXIANA publishes original papers on Zoological Systematics, with emphasis on Morphology, Phylogeny, Zoogeography and Ecology. Manuscripts will be accepted in German, English or French. A volume of three issues will be published annually. Extensive contributions may be edited in Supplement volumes. Redaktion - Editor-in-chief M.BAEHR Fotoarbeiten: M.MÜLLER Einbandgestaltung: Architekturbüro H.-J. SCHMIDT-SCHICKETANZ und Partner Dipl. Ing. Jan LEWERENZ Grundriß und Schnitt derZoologischen Staatssammlung Manuskripte, Korrekturen und Besprechungsexemplare Manuscripts, galley proofs, commentaries and review sind zu senden an die copies of books should be addressed to Redaktion SPIXIANA ZOOLOGISCHE STAATSSAMMLUNG MÜNCHEN Münchhausenstraße 21, D-81 247 München Tel.(089)8107-0 - Fax (089) 81 07-300 SPIXIANA - Journal of Zoology published by The State Zoological Collections München Die Deutsche Bibliothek - CIP-Einheitsaufnahme [Spixiana / Supplement] Spixiana : Zeitschrift für Zoologie / hrsg. von der Zoologischen Staatssammlung, München. - Supplement. München : Pfeil Früher Schriftenreihe ISSN 0177-7424 21. Winhard, W.: Konvergente Farbmusterentwicklungen bei Tagfaltern. - 1996 Winhard, Walter: Konvergente Farbmusterentwicklungen bei Tagfaltern ; Freilanduntersuchungen - in Asien, Afrika und Südamerika / von Walter Winhard. München : Pfeil, 1996 Zugl.: München, Univ., Diss., 1996 (Spixiana : Supplement ; 21) ISBN 3-931 51 6-1 1-3 Copyright © 1996 by Verlag Dr. Friedrich Pfeil, München Alle Rechte vorbehalten - All rights reserved. No part of this publication may be reproduced, stored in a retrieval System, or transmitted in any form or by any means, electronic, mechanical, photocopying or otherwise, without the prior permission of the Copyright owner. Applications for such permission, with a Statement of the purpose and extent of the reproduction, should be addressed to the Publisher, Verlag Dr. Friedrich Pfeil, P.O. Box 65 00 86, D-81214 München, Germany. ISSN0177-7424 - ISBN3-931516-1 1-3 Printed in Germany Verlag Dr. Friedrich Pfeil, P.O. Box 65 00 86, D-81214 München, Germany Tel.(089)74 28 270 - Fax (089) 72 42 772 - E-Mail: [email protected] ©Zoologische Staatssammlung München;download: http://www.biodiversitylibrary.org/; www.biologiezentrum.at Konvergente Farbmusterentwicklungen bei Tagfaltern Freilanduntersuchungen in Asien, Afrika und Südamerika Walter Winhard Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität Mi^inchen Verlag Dr. Friedrich Pfeil, München ©Zoologische Staatssammlung München;download: http://www.biodiversitylibrary.org/; www.biologiezentrum.at ©Zoologische Staatssammlung München;download: http://www.biodiversitylibrary.org/; www.biologiezentrum.at SPIXIANA Supplement 21 1-192 München, Ol. 11. 14% ISSN 0177-7424 ISBN 3-931516-11-3 Konvergente Farbmusterentwicklungen bei Tagfaltern Freilanduntersuchungen in Asien, Afrika und Südamerika Von Walter Winhard VVinhcird, W. (1996): Konvergente Farbmusterentwicklungen bei Tagfaltern. Freiland- untersuchungen in Asien, Afrika und Südamerika. - Spixiana Suppl. 21: 1-192 Some major aspects of the phenomenon of mimicry remain puzzling. The evaluation of collected specimens clearly shows the well-known forms of mimicry, i.e. Batesian and Müllerian mimicry, but extensive observations in nature give a more complicated picture, which is not so easily explained by the 'classical' forms of mimicry. In many cases, the obvious similarities of butterflies in their natural habitats do not fit the theories of Batesian or Müllerian mimicry either. The frequency of 'modeis" and "mimics" may also not be in accordance with the expectations drawn from theory. During extensive field work in South-East Asia, Africa and South America, an addi- tional and more generalizing theory has been developed to explain patterns of similarity among syntopic and synchronously occuring tropical butterflies. In general, a consider- able number of species converge in their overall appearance in flight, but if attacked by birds, the different species try to escape with different patterns of evasive behaviour, flight speed and directional changes, thereby creating a 'confusion group'. This reduces the chances of learning on the part of the predators and prevents a more pronounced convergence on the part of the prey, the palatable butterflies. In some cases wing struc- tures, such as elongated tails, serve to deceive predators. Species of very different system- atic Position may thus partake in this kind of mimicry grouping without being forced to develop the high degree of conformity typical of Batesian mimicry, as it is sufficient for the similarity to be apparent during rapid and mostly erratic flight. The new results provide a broader view of the mimicry phenomenon and they make the process of fine tuning in the classical forms of mimicry more plausible. The findings in this paper are supported by an extensive array of examples. (For more detailed information in English, see Summarv on page 1 1 1, in Spanish on page 113.) Dr. Walter Winhard, Austr. 4, D-87666 Pforzen. ©Zoologische Staatssammlung München;download: http://www.biodiversitylibrary.org/; www.biologiezentrum.at Inhalt Einleitung Problemstellung 1.2 Batessche Mimikry 8 1.3 Müllersche Mimikry 9 1.4 Ähnlichkeit genießbarer Arten 10 1.5 Freßfeinde 10 1.6 Chemische Verteitiigung bei Faltern 12 2. Beobachtungsorte 2.1. Auswahl der Beobachtungsorte 13 2.2. Der Khao Phra Thaew-Naturpark in Thailand 14 2.3. Die kenianische Küstenregion 15 2.4. Beobachtungsorte in Südamerika 16 3. Erhebung der Befunde und Methodik der Untersuchungen 3.1. Bestimmung und Präparation der auftretenden Arten 18 3.2. Erfassung des Verhaltens 3.2.1. Ermittlung der Falterhäufigkeit 18 3.2.2. Durchschnittliche Häufigkeit des Wechsels der Flugrichtung pro Sekunde und Fluggeschwindigkeit 21 3.2.3. Sonstiges Verhalten 22 3.3. Attacken der Freßfeinde 3.3.1. Direkte Beobachtung 22 3.3.2. Indirekte Erschließung 22 3.4. Füttertmgsexperimente 23 3.5. Statistische Auswertung der gewonnenen Daten (Meßergebnisse) 3.5.1. Flugrichtungswechsel und Fluggeschwindigkeit 24 3.5.2. Falterhäufigkeit und Verteilung der Attacken von Freßfeinden 24 3.6. Gesichtspunkte für die Zusammenstellung der Farbbilder 25 4. Ergebnisse und Diskussion cier Befunde 4.1. Flugyerhalten der Tagfalter 4.1 .1 Durchschnittliche Häufigkeit des Flugrichtungswechsels pro Sekunde und Fluggeschwindigkeit 25 4.1.2. Ändervmgen des Flugverhaltens bei äußeren Einwirkungen (Störungen) und bei der Balz. 29 4.1.3. Besondere Anpassungen an die Bedrohung durch Freßfeinde (Feindvermeidungsverhalten der Falter) 32 4.1.4. Diskussion 33 4.2. Selektionsdruck auf die Tagfalter 4.2.1. Häufigkeit der Attacken gegen schmackhafte und ungenießbare Falter 34 4.2.2. Wie Freßfeinde Falter jagen (Jagdmethoden der Freßfeinde) 35 4.2.3. Diskussion 36 4.3. Füttenmgsexperimente 4.3.1. Verlauf 37 4.3.2. Diskussion 39 4.4. Typische Fälle Batesscher und Müllerscher Mimikry 4.4.1. Thailändische Faltergemeinschaften 4.4.1.1. Pachlioptn aristolodünc, Atropliniieiim coon, die 9-form poh/tcs von Papilio poli/tcs und die ?-form distaiüiaiuis von Papilio mernnon als Ähnlichkeitskomplex in Thailand (Abb. 8. 16-17; mit Diskussion) 40 ©Zoologische Staatssammlung München;download: http://www.biodiversitylibrary.org/; www.biologiezentrum.at 4.4.1.2. Danainae (vor allem der Gattung Eiiploca) und ihre Nachaiimer im Khao Phra Thaew-Forest-Park (Abb. 8.18-23; mit Diskussion) 45 4.4.2. Südamerikanische Faltergemeinschaften 4.4.2.1. Mimikryringe mit den Pieridenarten Disniorphia aiuphionc bzw. Disinorphia nsti/iioiiic und Disinorphia astyocha 4.4.2.1.1. Hypothyris lycaste antonia und ähnliche Arten in der Westkord illere Ecuadors (Abb. 8.24; mit Diskussion) 50 4A.2A.2. Ein vergleichbarer Mimikryring in den Wäldern um die Wasserfälle des Iguazü (Abb. 8.25; mit Diskussion) 54 4.4.2.2. Heliconius erato und Heliconius melpomene (Abb. 8.27), Hclicoiüits cii/soiiyiiuis und Hcliconiits hccalcsia cxiiiiius (Abb. 8.28) in Ecuador, Heliconius crato phyllis (Abb. 8.26, rechts unten) bei den Wasserfällen des Iguazü (mit Diskussion) 56 4.4.2.3. Heliconius erato und Eresia lansdorfi in den argentinischen und brasilianischen Wäldern um die Wasserfälle des Iguazü (Abb. 8.26, rechte Spalte; mit Diskussion) 59 4.4.2.4. Heliconius erato und Adelplia isis in den argentinischen und brasilianischen Wäldern um die Wasserfälle des Iguazü, Adelpha lara und ähnliche Arten im südlichen Ecuador (Abb. 8.29; mit Diskussion) 61 4.4.3. Afrikanische Faltergemeinschaften 4.4.3.1. Amauris niavius dominicanus und ähnliche Arten an der kenianischen Küste (Papilio
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