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© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Systematic, evolutionary, and ecological implications of myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papilionoidea) by KONRAD FIEDLER BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 31 1991 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Die Serie wird vom Zoologischen Forschungsinstitut und Museum Alexander Koenig herausgegeben und bringt Originalarbeiten, die für eine Unterbringung in den „Bonner zoologischen Beiträgen" zu lang sind und eine Veröffentlichung als Monographie rechtfertigen. Anfragen bezüghch der Vorlage von Manuskripten sind an die Schriftleitung zu richten; Bestellungen und Tauschangebote bitte an die Bibhothek des Instituts. This series of monographs, published by the Zoological Research Institute and Museum Alexander Koenig, has been established for original contributions too long for inclu- sion in „Bonner zoologische Beiträge". Correspondence concerning manuscripts for pubHcation should be addressed to the editor. Purchase orders and requests for exchange please address to the library of the institute. LTnstitut de Recherches Zoologiques et Museum Alexander Koenig a etabh cette serie de monographies pour pouvoir pubher des travaux zoologiques trop longs pour etre in- clus dans les „Bonner zoologische Beiträge". Toute correspondance concernante des manuscrits pour cette serie doit etre adressee ä I'editeur. Commandes et demandes pour echanges adresser ä la bibhotheque de I'insti- tut, s. V. p. BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 31, 1991 Preis: 40,— DM Schriftleitung/Editor: G. Rheinwald Zoologisches Forschungsinstitut und Museum Alexander Koenig Adenauerallee 150—164, D-5300 Bonn 1, Germany Druck: Rheinischer Landwirtschafts-Verlag G.m.b.H., 5300 Bonn 1 ISBN 3-925382-33-X ISSN 0302-671X © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Systematic, evolutionary, and ecological implications of myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papihonoidea) by KONRAD FIEDLER BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 31 1991 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CIP-Kurztitelaufnahme der Deutschen Bibliothek Fiedler, Konrad: Systematic, evolutionary, and ecological impHcations of myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papihonoidea) / by Konrad Fiedler. Hrsg.: Zoolog. Forschungsinstitut und Museum Alexander Koenig, Bonn. — Bonn: Zoologisches Forschungsinst. und Museum Alexander Koenig. (Bonner zoologische Monographien; Nr. 31) ISBN 3-925382-33-X NE: GT © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CONTENTS Page Abstract 5 Introduction 6 Lycaenid myrmecophily: the general framework 6 Morphology and function of the myrmecophilous organs of lycaenid caterpillars 11 Communication between lycaenid caterpillars and ants 17 Ecology of lycaenid-ant interactions 19 Aims of the present study 21 Behavioural interactions between lycaenid larvae and ants: a comparative experi- mental study 24 The use of quantitative methods in the investigation of myrmecophily. ... 24 Material and methods 25 Results 26 Myrmecophilous species {Polyommatus coridon - Polyommatus icarus - Polyommatus escheri) 26 Myrmecoxenous species {Lycaena phlaeas - Lycaena tityrus - Lycaena hippothoe - Callophrys rubi - Hamearis lucina) 30 Discussion 32 The function of the myrmecophilous organs 32 Pore cupola organs (PCOs) 32 Dorsal nectary organ (DNO) 34 Tentacle organs (TOs) 35 Comparative aspects 36 Lycaenid systematics and Myrmecophily 39 The system of the Lycaenidae 39 Systematic distribution of myrmecophilous organs within the Lycaenidae 43 The outgroups: NymphaHdae and Riodinidae 43 The subfamihes Poritiinae, Miletinae, and Curetinae 45 Myrmecophilous organs of the Lycaeninae 46 The higher lycaenid taxa and their ant-associations 48 Poritiinae 48 Miletinae 48 Curetinae 49 Lycaeninae 49 Specificity of lycaenid-ant interactions 54 Which ants do visit lycaenid caterpillars? 54 Mechanisms of host-specificity 58 The obligatory and specific myrmecophiles within the Lycaenidae 62 Poritiinae 63 Miletinae 63 Lycaeninae 63 Lycaenid hostplant relationships and myrmecophily 67 Hostplants as selective agents in the Lycaenidae 67 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Database and analytical procedure 69 The hostplant relationships of the higher lycaenid taxa 71 Aberrant feeders: Poritiinae and Miletinae 71 Curetinae 72 Lycaeninae 72 Aphnaeini 72 Lycaenini 73 Thechni (Luciiti - Ogyriti - Zesiiti - Arhopaliti - Thecliti) 74 Eumaeini (Catapaecilmatiti - Amblypodiiti - Loxuriti - lolaiti - Reme- laniti - Hypolycaeniti - Deudorigiti - Eumaeiti) 78 Polyommatini (Candahditi - Lycaenesthiti - Niphanditi - Polyomma- titi) 83 Conclusions 84 General patterns of hostplant use within the Lycaenidae 84 Lycaenid hostplants in comparison to other butterflies 87 Are there trade-offs with myrmecophily? 89 Zoogeography of lycaenid-ant interactions 93 Zoogeography of the Lycaenidae 93 Poritiinae 94 Miletinae 94 Curetinae 95 Lycaeninae 95 Zoogeographie patterns in lycaenid myrmecophily 97 Introductory remarks 97 Europe and North West Africa 99 Japan 102 Australia 104 Thailand and West Malaysia 105 India 106 South Africa 108 Neotropical region 110 Nearctic region Ill Conclusions 113 Evolution of interactions between lycaenids and ants 116 Ants as selective agents for lepidopterous capterpillars 116 Evolution of myrmecophily and its related organs 117 Specializations and reductions 123 Obligatory myrmecophily 123 Secondary myrmecoxeny 128 Hostplants 129 Feeding habits 129 Habitat 129 Species diversity of the Lycaenidae: is myrmecophily a part of the answer? 130 Concluding remarks 132 Acknowledgements 134 Literature cited 135 Appendix: Tables 157 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 5 ABSTRACT 1. ) A quantitative study of the behavioural interactions of 7 European Lycaenidae and 1 Riodinidae species with 2 ant species revealed significant differences between myrmecophilous and myrmecoxenous caterpillars. A functional dorsal nectary organ is decisive for stable ant-associations. Further aspects of the function of the myrmecophilous organs, and the appHcation of the proposed experimental method to comparative surveys are discussed. 2. ) The presence of myrmecophilous organs in lycaenid larvae and the shape of their interactions with ants are intimately correlated with the higher classification of the family. The hypothesis of ancestral myrmecophily is rejected, the decisive dorsal nec- tary organ being an important synapomorphy of the most advanced subfamily Ly- caeninae. The characteristic states of myrmecophily are discussed for all higher lycaenid taxa. 3. ) Only trophobiotic ant taxa are involved in non-aggressive interactions with ly- caenids. Obligatory myrmecophily is mainly confined to ecologically dominant ants that form large, long-lived colonies, and most of such relationships occur in only a Hmited number of lycaenid Hneages, suggesting the occurrence of phyletic preadapta- tions for obligatory myrmecophily in these particular taxa. 4. ) Lycaenidae caterpillars mainly feed on plants of the subclass Rosidae with a distinct predilection of Fabales and Santalales. The hostplant patterns of the higher lycaenid taxa are described. There is little overlap between hostplant patterns of the Lycaenidae and other butterfly famihes. In contradiction to a recent hypothesis, no close correla- tion between myrmecophily and the predilection of Fabales or Santalales was found, with roughly 50 % of the myrmecophiles utilizing different foodplants. Neither obligate nor facultative myrmecophiles consistently have a wider hostplant range than myrmecoxenous species, indicating that myrmecophily has only exceptionally influenc- ed the hostplant relationships of lycaenid caterpillars. 5. ) The geographic distribution of myrmecophily is intimately correlated with the distribution of the higher lycaenid taxa. Gross patterns of lycaenid distribution point towards an important role of plate tectonics in the separation of the major lineages. The lycaenid faunas of 8 regions are systematically described, and the proportions of ant- associated species are estimated. A clear north-south disparity in the proportion or degree of myrmecophily is not discernible. 6. ) Some hypotheses concerning the evolution of myrmecophily in relation to the phylogeny of the Lycaenidae are proposed. Ant-associations have evolved in parallel in 2 lineages (Miletinae and Lycaeninae). In particular, specializations in, and reductions of, myrmecophily are discussed. 7. ) An Appendix summarizes data on hostplants and myrmecophily of more than 1000 Lycaenidae species. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at INTRODUCTION Lycaenid myrmecophily: the general framework The insect order Lepidoptera comprises 150,000—200,000 described species of which only 13,000—15,000