STUDIES ON THE ECOLOGY AND CONSERVATION OF BUTTERFLIES IN EUROPE VOL.2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model Edited by Josef Settele, Elisabeth Kuhn & Jeremy Thomas .. Conference Proceedings, UFZ LeipzigHalle, December 2005 Studies on the Ecology and Conservation of Butterflies in Europe i STUDIES ON THE ECOLOGY AND CONSERVATION OF BUTTERFLIES IN EUROPE Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model Edited by Josef Settele, Elisabeth Kühn & Jeremy A. Thomas ii J. Settele, E. Kühn & J. A. Thomas This page intentionally left blank Studies on the Ecology and Conservation of Butterflies in Europe iii Studies on the Ecology and Conservation of Butterflies in Europe Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model Edited by Josef Settele, Elisabeth Kühn & Jeremy A. Thomas Sofia-Moscow 2005 iv J. Settele, E. Kühn & J. A. Thomas STUDIES ON THE ECOLOGY AND CONSERVATION OF BUTTERFLIES IN EUROPE Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model Edited by Josef Settele, Elisabeth Kühn & Jeremy A. Thomas Pensoft Series Faunistica No 53 ISSN 1312-0174 First published 2005 ISBN 954-642-256-8 © PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. Pensoft Publishers Geo Milev Str. 13a, 1111 Sofia, Bulgaria Fax: +359-2-967-40-71
[email protected] www.pensoft.net Printed in Bulgaria, November 2005 Studies on the Ecology and Conservation of Butterflies in Europe v Contents Preface ................................................................................................................................................. xiii MacMan flyer .................................................................................................................................... xvii Section 3. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – overview paper and films ........................................................................................ 1 Parasitic versus mutualistic myrmecophiles among the Lepidoptera: the state of the art and where to go Konrad Fiedler .......................................................................................................................................... 3 MACULINEA - The fascinating world of Large Blue butterflies André Künzelmann, Thomas Falkner & Doris Böhme .......................................................................... 4 The Threatened Maculinea - Conservation biology as applied to humid zones Alain Rojo de la Paz ................................................................................................................................ 6 Section 3.1. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Maculinea as indicators .......................................................................................... 7 Spiders (Arachnida, Araneae) of a Maculinea alcon – M. teleius pSCI in NW Italy Marco Isaia, Simona Bonelli, Marco Montani, Guido Badino & Emilio Balletto .................................. 9 Maculinea habitats in Hungary: Orthoptera assemblages Antal Nagy, István A. Rácz & Zoltán Varga .................................................................................... 16 Habitat preferences of Myrmica (Hymenoptera: Formicidae) ant species in Maculinea arion (Lepidoptera: Lycaenidae) sites in South-Western Germany Regina Pauler-Fürste & Manfred Verhaagh ......................................................................................... 22 Myrmica ants as keystone species and Maculinea arion as an indicator of rare niches in UK grasslands Zoe Randle, David J. Simcox, Karsten Schönrogge, Judith C. Wardlaw & Jeremy A. Thomas ........... 26 Maculinea and myrmecophiles as sensitive indicators of grassland butterflies (umbrella species), ants (keystone species) and other invertebrates Jeremy A. Thomas, Ralph T. Clarke, Zoe Randle, David J. Simcox, Karsten Schönrogge, Graham W. Elmes, Judith C. Wardlaw & Josef Settele ................................................................ 28 Assemblages of butterflies and burnets in Maculinea habitats of Hungary Zoltán Varga, László Peregovits & Julianna V. Sipos ......................................................................... 32 ............ Piotr Skórka........ 75 Does the colony structure of Myrmica rubra affect the adoption of Maculinea nausithous? Holger Loritz.... Graham W................................................ Josef Settele & Karsten Schönrogge ............ Piotr Nowicki & Michal Woyciechowski ....................... 61 Oviposition behaviour in the myrmecophilous butterfly Maculinea alcon (Lepidoptera: Lycaenidae) Simona Bonelli... 51 Section 3........P....... rebeli................ Sándor Csősz....... Ewa Sliwinska........... 80 Oviposition in Maculinea alcon butterflies Edina Prondvai....................... Martin Musche & Josef Settele .. 69 The effect of ant communities and spatial pattern for Maculinea nausithous Uta Glinka & Josef Settele ....................... Kühn & J.. András Tartally.................. Mariann Bíró & Zoltán Varga ..............2.... a sympatric social parasite to the Maculinea in moist grassland ecosystems Simona Bonelli........ Emese Szitta.. E................................. Elmes..... Francesca Barbero & Emilio Balletto .. Christian Anton...... 45 The ant communities on meadows in the Kraków region Magdalena Witek..... Worgan....... Thomas Maculinea habitats: diversity of vegetation................................ 73 Egg-laying behaviour of Maculinea rebeli Hirschke........................................ 1904 Ádám Kőrösi ...................... 53 Parasitism of the predatory Maculinea nausithous by the parasitoid Neotypus melanocephalus Christian Anton................................ Sophie Everett.......... 65 Host specificity in Microdon myrmicae................................ Emma Napper........................Stankiewicz............. 78 Do ant cues influence the oviposition preference in the myrmecophilous Maculinea nausithous? Martin Musche... Judit Bereczki.......................... 57 Egg-laying preferences of the xerophilous ecotype of Maculinea alcon (= M............................. Andrea Tóth.................. Wardlaw................ Ervin Árnyas........................................................................................ composition and cenological relegation Julianna Varga-Sipos & Zoltán Varga ............. Andrea Tóth................... S......... Martin Musche.. Ania M....... Andrew D... Cantarino........ Andrea Crocetta.. Emilio Balletto & Karsten Schönrogge ......................... Andras Tartally.................... Settele................... Ferenc Kassai. 74 Behavioural aspects of adoption of Maculinea caterpillars by Myrmica ants Szabolcs Lengyel.. Judith C............................... 72 Contrasting egg laying behaviour of the ecotypes of Maculinea alcon in Hungary Ferenc Kassai & László Peregovits ........................ Lepidoptera: Lycaenidae) in the Aggtelek National Park Ervin Árnyas................. Judit Bereczki.. Christian Anton.... Katalian Pecsenye & Zoltán Varga ........................ 55 Which factors determine the population density of the predatory butterfly Maculinea nausithous? Christian Anton................... Andrew Worgan & Josef Settele .... A.............. Vladimir Hula & Josef Settele .....................vi J.. 82 .......... Marcin Sielezniew. László Peregovits & János Kis ..................... Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Functional and trophic relations in Maculinea systems .... Zoltán Varga............. ......................................... Stankiewicz........ Nicolas Mouquet............................... Ewa Sliwinska & Michal Woyciechowski ... Judit Bereczki.... 99 Host ant specificity and integration rate with Myrmica ants in larvae of Maculinea teleius butterflies Magdalena Witek.................. Hochberg.... Jeremy A......... rebeli in Poland: distribution.. Piotr Nowicki.... 105 Biennialism and host ant specificity in Maculinea teleius larvae Magdalena Witek................... Charles.......... Judith C.... Marcin Sielezniew & Jarosław Buszko .............. Piotr Nowicki.... and the expansion of species’ range Thomas Hovestadt .................................3.. 123 ...... Marcin Sielezniew & Jeremy A............. Thomas................. the Allee-effect... 88 Maculinea alcon and M................Studies on the Ecology and Conservation of Butterflies in Europe vii Acoustical interactions between Maculinea alcon and its host ant Karsten Schönrogge...................... Karsten Schönrogge.......... Michael E..... Graham W..... Clarke................ Thomas...... 120 The control of emigration................................ 109 Results of the mark-release-recapture studies of a Maculinea rebeli population in the Aggtelek karst (N Hungary) between 2002-2004 Ervin Árnyas.................. Elmes.. connections with parasitoids and host plants András Tartally & Zoltán Varga ..... Topham............................................ Graham W...... M......... David Tesar........... nausithous and M......................................... Alexander Singer & Joseph Hale ................ David R. Elmes....................... E..................... Piotr Nowicki..... environmental variability....................... 94 Primary hosts.................................... 111 Modelling the local population dynamics of Maculinea and their spatial interactions with their larval foodplant and Myrmica ant species Ralph T............................ Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Population ecology of Maculinea .............. M................ 121 Within-patch movement limitation in two species of Maculinea butterflies? Analysis of MRR data using randomisation procedures Thomas Hovestadt & Piotr Nowicki .... Piotr Skórka........................................... 122 Emigration and its consequences for the survival of metapopulations Thomas Hovestadt & Hans Joachim Poethke .... habitats.. Piotr Skórka.................................. Thomas ....... Wardlaw.... teleius. 115 A review of the role of dispersal for population persistence in Maculinea Thomas Hovestadt .................. Andrea Tóth & Zoltán Varga ........... alcon Ewa Sliwinska........... 90 Host-ant specificity of Maculinea species in Hungary........... secondary hosts and ‘non-hosts’: common confusions in the interpretation of host specificity in Maculinea butterflies and other social parasites of ants Jeremy A................................................. Nash & Michal Woyciechowski ................... 107 Section 3..... host ant specificity and parasitoids Anna M................... David J................ Ewa Sliwinska & Michal Woyciechowski ........ Ania Stankiewicz............................................... Simcox & Josef Settele ... 84 The key to the caterpillars and pupae of M............................ ......................................... Ágnes Vozár & Lilla Barabás ...... Piotr Skórka & Michal Woyciechowski . 151 Assessing the presence/absence of Maculinea nausithous: a comparison of adult and preimaginal stages Josef Settele...... Christian Anton............ 131 Habitat-use of wetland Maculinea species – a case study Ádám Kőrösi ........................ 134 Population dynamics in the genus Maculinea revisited: comparative study of sympatric M................ 128 Annual and spatial variations in population structure – A case study of Maculinea alcon and Maculinea teleius Ferenc Kassai.. Jeremy A............................... 150 Estimation of butterfly population sizes using pre-imaginal stages exemplified by Maculinea butterflies (Lepidoptera: Lycaenidae) Manfred Alban Pfeifer..... Aleksandra Pępkowska.................. Martin Musche & Anett Richter ........................ Holger Loritz.............. Thomas Analysis of inter-patch dispersal data for Maculinea nausithous and M......... 125 An ESS model for the evolution of growth polymorphism in the social parasite Maculinea rebeli Thomas Hovestadt...... Germany Thomas Hovestadt.......... Sarah Gwillym... Josef Settele ................................... alcon and M............................... E..................... teleius and M. Noémi Örvössy............... southern Poland Piotr Nowicki.............. Uta Glinka.........viii J.................. László Peregovits.................................... Tobias Degen & Hans-Joachim Poethke ........................... Ádám Kőrösi..................... Graham Elmes................. 152 ................ 144 Structure of sympatric populations of M.................................................. 136 Landscape scale research in butterfly population ecology – Maculinea case study Piotr Nowicki... Magdalena Witek & Michał Woyciechowski ....... Kühn & J......................................... the Allee-effect and the recolonisation of empty habitats Thomas Hovestadt....................................... Ágnes Vozár & Ferenc Kassai ............. 124 Control of emigration................... Simona Bonelli............................... Josef Settele........... nausithous in the Kraków region..................................................... teleius in a fragmented landscape in northern Bavaria... A............. Piotr Skórka....... teleius Piotr Nowicki...................... Settele...................... Joanna Kudłek........ 132 Studying the population structure of Maculinea arion ligurica Ádám Kőrösi.. Uta Glinka & Josef Settele ..... Oliver Mitesser.............................. Thomas & Michal Woyciechowski ................... Magdalena Witek................ Noémi Örvössy........... 140 A review of population structure of Maculinea butterflies Piotr Nowicki............................. 130 Analysis of within-habitat patch movement of some Maculinea species Ádám Kőrösi ............................................................................................................. 126 Goodness of sampling in Maculinea butterflies Ferenc Kassai ...... Jeremy Thomas & Michael Hochberg ............... Birgit Binzenhöfer........ 133 Simplified method of estimating butterfly population size with mark-release-recapture Piotr Nowicki & Josef Settele ..................... László Peregovits.......... Francesca Barbero & Emilio Balletto ........... Judith C. Boomsma ........................ Magdalena Witek & Michal Woyciechowski ..... Piotr Nowicki....................................................... Nash.................. 178 The relationship between genetic diversity......................... Josef Settele.. Jeremy A.... Magdalena Witek & Michal Woyciechowski ........... Simona Bonelli.................. Katalin Pecsenye & Zoltán Varga . Elmes & Ralph T..... 157 Geographical versus food plant differentiation in Alcon Blue Populations (Lepidoptera: Lycaenidae) in Northern Hungary Judit Bereczki............ Joanna Kudłek................ Ian R.......... Boomsma ................. Thomas & Karsten Schönrogge ................ Wardlaw. 182 ........................ Karsten Schönrogge..... Worgan.. Andras Tartally..... Laszlo Peregowitz............. 163 Pattern of genetic differentiation in the Maculinea alcon species group (Lepidoptera: Lycaenidae) in Central Europe Judit Bereczki................... Ebsen................. Simcox..................... Nash & Jacobus J......... Marcin Sielezniew. Aleksandra Pępkowska................. Katalin Pecsenye..4...................... David N.. 159 Temporal and spatial structure of genetic variation in the Alcon Blue (Lepidoptera: Lycaenidae) populations in Northern Hungary Judit Bereczki........ Dirk Maes.................................. Irma Wynhoff... Ebsen............................ Zoltan Varga....... David J. Nash... Elmes.......... Sophie Everett & Jacobus J...... 171 Multiple radiation of species and eco-types in the genus Myrmica Graham W....... 167 Cryptic Myrmica species among the hosts of Maculinea butterflies Jon R......... Ania Stankiewicz....................... Wynne................................... alcon in southern Scandinavia in relation to the conservation of these species Andreas E..... Jon R......... David Tesar... Boomsma ................. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Population genetics and physiology of Maculinea and Myrmica ants ... Clarke . Ian Wynne..... David R....... Graham W. Inga Zeisset.............................. David R........................... Ewa Sliwinska................P... László Peregovits & Zoltán Varga ..... 180 Using genetic markers to study the phylogeny............................. Lomborg... Andrew D.......... Michal Woyciechowski........ Sandor Csósz........... Katalin Pecsenye & Zoltán Varga . Christian Anton............ 153 A simple method for estimating worker population size in Myrmica ant nests Piotr Skórka.... Piotr Nowicki..... Nash & Jacobus J.... 174 A population genetic study of Maculinea arion and M..................................................................... 154 Section 3...... distribution and ecology of Maculinea butterflies David R........ 172 Variation in chemical profiles of Maculinea and their Myrmica hosts across Europe Sophie Everett...... nausithous Piotr Skórka..........Studies on the Ecology and Conservation of Butterflies in Europe ix Mobility patterns of Maculinea teleius and M.................................. Walter Durka & Josef Settele . genetic population structure and sexual reproduction in Sanguisorba officinalis (Rosaceae) Martin Musche........ Martin Musche.... Inga Zeisset.... ............ Andrea Tóth. Boomsma & Michal Woyciechowski ........ Settele.. 215 Influence of mowing on the persistence of two endangered Large Blue Butterfly Maculinea species Karin Johst............. 225 ............... Marcin Sielezniew & Anna M............ Josef Settele & Frank Wätzold ........................ Martin Drechsler............................................ Stankiewicz .... Judit Bereczki................ 206 The distribution and ecology of Maculinea teleius and M....... Alexander Blinov & Michal Woyciechowski ..............................x J................................................ 219 Changing a butterfly’s landscape – persistence of the Dusky Large Blue in managed grasslands Holger Loritz & Josef Settele .................................................... Karin Johst........... László Peregovits & Zoltán Varga ........... 218 The other side of the coin: the economic value of butterfly conservation Nele Lienhoop.......... 189 The genetic structure of the Maculinea teleius (Lepidoptera: Lycaenidae) populations in Hungary Katalin Pecsenye................ Thomas & Josef Settele ....... Thomas Non-LTR retrotransposons from Large Blue butterfly Maculinea teleius: the diversity of CR1-like elements Olga Novikova.................... Frank Wätzold................................. 184 Genetic differentiation among the Maculinea species (Lepidoptera: Lycaenidae) in eastern Central Europe Katalin Pecsenye. Judit Bereczki......... 192 Effective population size of Maculinea teleius in southern Poland Ewa Sliwinska............... Jacobus J......... Karin Johst.............................. Giselher Kaule & Josef Settele ........... 210 A model-based approach for designing cost-effective compensation payments for the conservation of endangered species in real landscapes Martin Drechsler................ Piotr Nowicki..... Ewa Sliwinska....... Borbála Tihanyi......... Holger Bergmann & Josef Settele ........................ Andrea Tóth & Zoltán Varga ............... 205 Protection of low yielding Sanguisorba officinalis grasslands as habitat of the Large Blue butterflies Maculinea nausithous and Maculinea teleius – model calculations on the efficiency of agri-environmental schemes Holger Bergmann.............. 199 Section 3............................................... Kühn & J................................................... Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Conservation and management for Maculinea ... 214 Effects of connecting strategies on the Large Blue Butterfly Maculinea nausithous – a case study Sabine Geißler-Strobel.................... Martin Drechsler..................... nausithous in Poland Jarosław Buszko.. Judit Bereczki & Zoltán Varga ...................................................................................... 203 MacMan at schools Anna Amirowicz & Michal Woyciechowski ...................... E.... 221 Effects of human land-use on availability and quality of habitats of the Dusky Large Blue butterfly Holger Loritz & Josef Settele ........ A.............. Susanne Koschel & Josef Settele ................. Jeremy A.......... Katalin Pecsenye............................................................ 196 Patterns of genetic differentiation in the Hungarian Maculinea arion (Lepidoptera: Lycaenidae) populations Andrea Tóth..................................5.............. ........ Kristian Malačič.......................................... Ádám Kőrösi........ Simcox........... Jana Bouberlová......... Ralph T......................... 238 Field research based development of management guidelines for the protection of species and ecosystems of the Habitats Directive – A case study of “wetland Maculinea species” in Bavaria Christian Stettmer & Markus Bräu ..................................................... Jarosław Buszko & Anna M....... Elmes................. Zoe Randle................................................... Graham W................................... Clarke.................... 253 .......................................... Karsten Schönrogge............................................... Péter Batáry........................................................................... Martin Drechsler..Studies on the Ecology and Conservation of Butterflies in Europe xi Patterns of resource allocation and adaptive response to mowing in the plant Sanguisorba officinalis (Rosaceae) Martin Musche & Josef Settele ................ 240 A supporting tool for decision-making in Maculinea management Karin Ulbrich.......................... Dušan Zitňan & Ján Kulfan .............................. Holger Bergmann & Josef Settele ........ László Peregovits ...................................................................... Josef Settele & Michal Woyciechowski ...... 230 Maculinea arion in Poland: distribution........... Piotr Nowicki..................................... ecology and conservation prospects Marcin Sielezniew.......................................... 229 How endangered is Maculinea nausithous? Josef Settele ........... Hana Veselá & Jiří Cibulka .. 228 Habitat use and effect of habitat management on Maculinea teleius Noémi Örvössy...................... Frank Wätzold................................ 245 Conservation of Maculinea Species in Slovakia ˇ Ľubomíra Vavrová.......... 234 Grassland butterflies profit from succession but suffer from invasions – a case study from Southern Poland Piotr Skórka.......... Thomas ....... 249 General overview of the status of Maculinea butterflies in Poland Michal Woyciechowski .................. Stankiewicz .............. 251 Distribution and autecology of Maculinea teleius and M.................................... nausithous (Lepidoptera: Lycaenidae) in Northeast Slovenia Valerija Zakšek............................ Frenk Rebeušek & Rudi Verovnik ................ Josef Settele & Jeremy A........................... 231 Science and socio-economically-based management to restore species and grassland ecosystems of the Habitats Directive to degraded landscapes: the case of Maculinea arion in Britain David J.... Karin Johst................................. Ágnes Vozár......................................................................................................................... 247 Conservation of Maculinea populations affected by a waterway construction in Přelouč (Czech Republic) in the view of Czech University of Agriculture research team Vladimír Vrabec. 239 Contrasting management requirements of Maculinea arion across latitudinal and altitudinal climatic gradients in west Europe Jeremy A Thomas & David J Simcox .............. ......................................................................................... 257 Bibliography on Maculinea ecology and related topics (state: September 2005) Elisabeth Kühn............ E......... A...... Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Maculinea Bibliography .................. 285 Index of latin butterfly names ........ Settele........ 259 Author index .......................................xii J.............. 287 ...........6.. Jeremy Thomas & Josef Settele .............................. Thomas Section 3....................................................... Sarah Gwillym.................................. Kühn & J................................................... Held at UFZ LeipzigHalle on 5th to 9th of December 2005.5 million Euro to restore up to 70 km of degraded grassland ecosystems. in Switzerland. in numbers that dwarf the previous workers in this field. Inevitably. not just as objects worthy of greater conservation effort in their own right but also in recognition of their wider usefulness as sensitive indicators of environmental change (especially of habitat degradation. one product of the two EU programmes has been the training across Europe of a new generation of excellent young scientists. interest and the classy images have also made butterflies increasingly useful tools in education. Europe has seen ever larger projects involving butterflies as tools for science. to study both the ecology of endangered Maculinea species and their usefulness as ‘super-indicators’ in conservation. along the Atlantic coast of Cornwall. the second. Hand-in-hand with the increased use of butterflies in pure and applied biology has been a burgeoning popular interest in them as objects to be noticed and enjoyed.net). far more rapidly than any scientist of the 1980s could have foreseen. watching and photography have largely replaced as leisure activities the more specialised and male-dominated hobby of collecting. Matching this has been a parallel advance in the priority afforded to butterflies by global. led by Josef Settele (UFZ.macman-project. films and videos: some are described in these volumes. this growing knowledge. On the contrary.de). 25 million Euro to build on the outskirts of London the biggest (by far) walk-through exhibition of living butterflies in the world.Studies on the Ecology and Conservation of Butterflies in Europe xiii Preface The use of butterflies as model systems in biological research has increased enormously over the past two decades.europeanbutterflies. eventually containing more than a quarter of a million (exotic) butterflies and expected to attract many more than the quarter of a million visitors that annually visit its sister butterfly house. and . national and voluntary conservation bodies. ensure that current interest in butterflies is not a passing phase. www.fi/science/fragland) to use questions about butterfly metapopulations to train PhDs and exchange post-doctoral researchers across European nations.thanks to the new technologies – the beauty of butterflies has spawned an unprecedented number of high quality images. the National Lottery Fund has approved funding of >2. Across Europe. These. conservation and leisure.helsinki. Papiliorama. Germany. highly skilled in butterfly ecology and conservation. it was composed of 10 sections which were divided into . fragmentation and climate change) and as umbrella species whose targeted conservation benefits wider communities of lesser-known. Finland. Prominent among these are the granting of planning permission to the Butterfly World Trust to invest c. The conference “Ecology and Conservation of Butterflies in Europe” brought together most of the leading and new butterfly biologists and conservationists of Europe. and many other developments. In science. threatened species. www. targeted for native butterflies (especially Maculinea arion) and associated wildlife. the EU recently funded two massive programmes of research – Fragland and MacMan – the first led by Illka Hanksi (University of Helsinki. NGO butterfly conservation societies enjoy unprecedented growth. In the first five years of the 21st century. Today butterfly gardening. culminating in the foundation in 2004 of the continentalscale “Butterfly Conservation Europe” (www. Also in the UK in 2005 (and one of the ‘babies’ of the MacMan programme). Its broad aims are twofold. The acronym MacMan stands for MACulinea Butterflies of the Habitats Directive and European Red List as Indicators and Tools for Habitat Conservation and MANagement. reproduced in this book on the pages after this preface. and will continue – we hope for many years – as a self-sustaining European Network of insect conservation biologists. Full details of partners. The first volume “General Concepts and Case Studies” encompasses the “Ecology of Butterflies” (3 sections) and the “Conservation of Butterflies and Global Change” (two sections). to develop standards for monitoring Maculinea butterflies as indicators and tools for grasslands and their management. over and above the small ‘community modules’ of species with which they directly interact.and intraspecific variation in their functional ecology across the continent’s gradients of phylo-geography. their status as globally threatened species. In practice. Thomas two conference blocks. and because of their fascinating life-styles that involve living as social parasites within red Myrmica ant colonies for 11-23 months during the late larval and pupal stages. and their associated foodplants. was to assess the suitability of Maculinea butterflies as indicators of biodiversity along a European transect. but are also umbrella species whose targeted conservation is likely to enhance a wide range of other desirable but endangered species in their communities. Maculinea butterflies are not only unusually sensitive indicators of biodiversity richness and particularly of change in response to environmental degradation of the wide range of grasslands they inhabit (fen. This volume contains 90 extended abstracts or mini-papers of one to several pages. In addition to the . First. it recognises that Europe’s five accepted species of Maculinea butterfly have achieved iconic status in Europe (and the world) due to their beauty. therefore. xerophytic grassland). The MacMan project formally runs from February 2002 to late January 2006 and involves eight collaborating institutions from six European nations. 13 sub-contractors ranging from the Polish Academy of fine arts to practical conservation organisations and other users. and a total of about 60 European ecologists. Settele. The second aim of MacMan was based on the hypothesis – backed by anecdotal observation and theory – that. One aim of the MacMan scientists was to generate the fundamental knowledge essential to conserve Europe’s five Maculinea systems by increasing our understanding of inter. humid heath. it includes eleven additional collaborating groups. describing new developments in a diversity of aspects of recent Maculinea research. preceded by a brief initial period of feeding on a specific foodplant(s). while the second volume “Species Ecology along a European Gradient: Maculinea Butterflies as a Model” contains 5 sections and encapsulates the final meeting of the four-year EU Framework V MacMan project. One rationale supporting this hypothesis was that the host Myrmica ants of Maculinea are keystone species which have a disproportionate impact on shaping the communities in the biotopes that they inhabit. due to their specialised life-styles and dependency on two larval resources. is important for conservation not only because these are globally endangered systems in their own right but because the Maculinea have been flagship insects ever since their selection in the 1970s by the World Conservation Union (IUCN) as one of three global priorities for butterfly conservation (the other two being Queen Alexandra’s Birdwing of Papua New Guinea and the Mexican overwintering roosts of monarch butterflies). which are reflected in these two volumes of Proceedings.xiv J. E. ants and specialist parasitoids. climate and altitude. Kühn & J. and to promulgate these monitoring standards and to promote the conservation and restoration of viable meta-populations of Maculinea and their associated wildlife across European landscapes. The second aim of MacMan scientists. A. associates and programmes are given in the MacMan ‘flyer’. Thus the ability to conserve Europe’s dwindling Maculinea populations. moist hay meadow. dry calcareous grassland. Finally. and single-site and metapopulation models ranging from phenomenological and spatial models of the population dynamic interactions within Maculinea community modules (Clarke) to ESS models of evolutionary ecology (Hovestadt et al). It includes six papers on assessment of decline or changing status. it is supplemented by a few review papers or those covering work presented in the earlier years of MacMan. (3)‘Population biology of Maculinea’. six more on host specificity in various parts of Europe. this book has five main sections. anonymously. eight papers giving management advice for conserving populations across various parts or gradients of Europe. and Thomas et al explore the wider role of Maculinea. other myrmecophiles. Randle et al document the successful use of M. together with papers on cryptic speciation and species richness in their Myrmica hosts (Ebsen et al. acoustical communication (Schönrogge et al) and biennialism (Witek et al) in larvae. although by no means all the early work is represented. ants and parasitoids. In addition to the closing keynote lecture of Konrad Fiedler (Austria) that places social parasitism in a wider context. including six papers on oviposition preferences. arion as an umbrella species for the restoration of species-rich grassland. Elmes et al) and inter. (4)‘Population genetics and physiology of Maculinea and Myrmica’ includes ten papers describing mainly intra-specific patterns of genetic variation in Maculinea species across Europe. (5)‘Conservation and management for Maculinea’ applies the knowledge from the previous four sections. broadly reflecting the eight workpackages of the MacMan programme of research: (1)‘Maculinea as indicators’ complements earlier analyses of biodiversity on occupied and unoccupied M. butterflies and burnets (Varga et al).and intra-specific variation in the chemical cues used by Maculinea larvae to mimic different ant hosts (Everett et al). focuses on measuring population parameters such as variation in numbers and habitat quality in different parts of Europe (11 papers). and butterflies in general as sensitive or typical indicators of other invertebrates. as its name implies. parasitoids (Anton et al) and other social parasites that coexist with Maculinea (Bonelli et al). dispersal and the Allee effect (9 papers). and others on population ecology (Árnyas et al). many generated as a result of the MacMan programme. Orthoptera (Nagy et al). plants and vertebrates. foodplants. “Parasitic versus mutualistic myrmecophiles among the Lepidoptera: the state of the art and where to go”. While in the UK. The conference was possible only through the support of many friends and colleagues. (2)‘Functional and trophic relations in Maculinea systems’ presents much new information on the basic ecology and behaviour of Maculinea species. alcon sites in Belgium by comparing the species richness of spiders (Isaia et al). by at least two colleagues. as well as five papers (including models) in which socio-economic considerations are a important parameter in conservation recommendations. to practical conservation questions and solutions. ants (Witek et al) and vegetation (Varga-Sipos & Varga) of Maculinea and other grasslands in Hungary or Poland. as well as much other MacMan research. The contributions of both proceedings volumes have been peer refereed. In particular we thank colleagues from .Studies on the Ecology and Conservation of Butterflies in Europe xv papers delivered at the 2005 conference “Ecology and Conservation of Butterflies in Europe”. whose help is greatly acknowledged. we present a bibliography of 447 key papers for Maculinea research and conservation. we thank the EU whose generous funding made this exciting and useful programme possible. E. Susan Walter and Sylvia Ritter. Thomas UFZ: André Künzelmann. Christiane Viehrig. from CONFIRM Ltd: Hildegard Feldmann & Ogarit Uhlmann. Klaus Henle. Annette Schmidt. NERC): Graham Elmes. Georg Teutsch. Monique Franke. Josef Settele. Kühn & J. A. Silke Rattei. Dirk Immisch. Zoe Randle and Nicky Gammans. Sarah Gwillym. Martin Sharman and Sybille van den Hove for all their support on behalf of the European Commission. Doris Böhme. Andreas Staak. Judith Wardlaw. Finally. Martin Musche. Settele. We are also indebted to Frank Nolden. Dorset (UK) October 2005 .xvi J. Christian Anton. Dana Weinhold. Sabine Rott. and Karin Zaunberger. and from the Centre of Ecology & Hydrology (CEH Dorset. Peter Fritz and Stefan Klotz (all UFZ) and to Mark Bailey (CEH) for the scientific and administrative support of biodiversity research in general and of research on butterfly ecology and conservation in particular. Heike Wolke. Elisabeth Kühn & Jeremy Thomas Halle (Germany). Ellen Selent. Karsten Schönrogge. Studies on the Ecology and Conservation of Butterflies in Europe xvii MacMan . Settele. Kühn & J. E. A. Thomas .xviii J. Studies on the Ecology and Conservation of Butterflies in Europe xix . Thomas . Settele.xx J. Kühn & J. A. E. Studies on the Ecology and Conservation of Butterflies in Europe xxi . xxii J. A. Thomas . E. Settele. Kühn & J. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – overview paper and films . nausithous in Poland 1 Section 3.The distribution and ecology of Maculinea teleius and M. Marcin Sielezniew & Anna M.2 Jarosław Buszko. Stankiewicz This page intentionally left blank . © PENSOFT Publishers The distribution Sofia – Moscow J. and the resulting ecological and evolutionary dynamics. Althanstr. 3 Parasitic versus mutualistic myrmecophiles among the Lepidoptera: the state of the art and where to go Konrad Fiedler University of
[email protected] nectar-mediated mutualisms between ants and other organisms. Not unexpectedly. Recent studies on the nectar secretions of facultative myrmecophiles confirmed earlier hypotheses that more intimately ant-associated species secrete more valuable nectar rich in amino acids. rather than honest signalling.ac. Since the quality and quantity of nectars containing melezitose is poor. and ecology of Maculinea teleiusthe EcologySettele. These have become textbook examples for the evolutionary ecology of parasite-host systems. ant-parasitic species account for only a small fraction of extant Lycaenidae diversity. A-1090 Vienna. The proximate mechanisms mediating these associations. These parasite-host systems largely corroborate predictions of the geographic mosaic model of coevolution and may also support speciation. . In view of that context. Division of Population Ecology. This full integration into the social code of ants allows for finely tuned population specific host ant use. ant-parasitic lycaenid butterfly larvae employ the strategy of active chemical mimicry to integrate into host-ant colonies. E.and Conservation of Butterflies Vol. and backing up current themes on honeydew. Kühn &in Polandin Europe nausithous J. p. but not all attendant ants. chemical mimicry cannot explain the establishment and maintenance of stable associations. However. However. the vast majority of myrmecophiles among the lycaenids does not maintain species. Melezitose may serve as an advertisement signal to some.or genus-specific associations with ants.A. just as in other nest inquilines of ants and other social insects. Thomas (Eds) 2005 3 Studies on and M. this does not appear to be an honest signal. There is growing. some lines of further research will be addressed that should be followed to uncover the mechanisms and consequences of ant-attendance in these more ‘typical’ lycaenid species. Institute of Ecology and Conservation Biology. is common among facultatively antattended lycaenid butterflies. are far less well understood than in the more ‘atypical’ obligate myrmecophiles. Cost-benefit estimates and a range of behavioural data also suggest that manipulation.at In the past two decades much research has centred on ant-parasitic butterflies such as the Maculinea-Phengaris clade. albeit largely circumstantial evidence that similar rules apply to lycaenid species which have obligate mutualistic associations with ants. AUSTRIA Contact: konrad. which is known from homopteran honeydew. Less intimately myrmecophilous species also secrete the trisaccharide melezitose. 14. Here. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. J. J.A. Thomas © 4 PENSOFT Publishers André Künzelmann, Thomas Falkner & Doris Böhme the EcologySettele, E. Kühn & of Butterflies(Eds) 2005 Studies on and Conservation in Europe Sofia – Moscow Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, pp. 4-5 MACULINEA - The fascinating world of Large Blue butterflies DVD documentation 20 minutes - A film production of the UFZ Centre for Environmental Research, Leipzig-Halle in the Helmholtz Association André Künzelmann, Thomas Falkner & Doris Böhme UFZ – Centre for Environmental Research Leipzig-Halle, Public Relations Department, Permoserstr. 15, 04318 Leipzig, Germany Contact:
[email protected] Despite its pretty, filigree appearance, the large blue is characterised by what is for butterflies a distinctly brutal lifestyle, often involving the death of other creatures, namely ants. Like most other butterflies, the large blues lay their eggs on selected plants. However, the caterpillars do not complete their life cycle on these plants. Instead they become parasites of red ants. These ants carry the caterpillars voluntarily into their nests, apparently assuming that they belong to their own brood. Once in the nest, the caterpillars consume the ant eggs and larvae or, like cuckoo chicks, are fed by the worker ants. This extraordinary lifestyle puts the large blue among the most remarkable insects in Europe. Unfortunately, their double dependency – on the right plant species and the right species of ant – means that there are only a few places in the wild where the necessary conditions are to be found. Consequently, it is no surprise that all five European Maculinea species appear on many Red Lists of threatened species. Even the smallest changes in land use or climate can lead to a situation where the right host ants are no longer found in combination with the right plants and thus whole butterfly populations are wiped out. This film provides an insight into the world of the blue butterfly and shows its importance in interaction with nature. It highlights the fact that the butterflies cannot easily adapt to the constantly growing land-use demands of humans and shows how their disappearance sounds an alarm for our environment. The film was produced by the UFZ Centre for Environmental Research Leipzig-Halle to illustrate a typical approach adopted in the field of biodiversity research and its communication; based on activities and results of the EU FP5 RTD project MacMan (www.macman-project.de). Additional footage for the film has kindly been provided by David Nash, Marcin Sielezniew and Anna Stankiewicz. The film was supported by many friends and colleagues, to whom we are very thankful: Andrea Grill (ANL Laufen, Germany), Andreas Nunner (Bioplan, Germany), Anna Stankiewicz (Museum and Institute of Zoology, Warsaw, Poland), Burkhard Beinlich (Landschaftss- MACULINEA - The fascinating world of Large Blue butterflies 5 tation Höxter, Germany), Christian Stettmer (ANL Laufen, Germany), David Nash (University of Copenhagen, Denmark), Jeremy Thomas (CEH Dorset, UK), Marcin Sielezniew (Warsaw Agriculture University, Poland), Martin Warren (Butterfly Conservation, UK), Robert Völkl (ANL Laufen, Germany), Tobias Schiefer (ANL Laufen, Germany) and Ulrike Möhring (Berlin, Germany). And from UFZ Leipzig-Halle (Germany): Alexander Singer, Christian Anton, Elisabeth Kühn, Holger Loritz, Josef Settele, Karin Johst, Martin Drechsler, Martin Musche and Uta Glinka. Research within the project MacMan (Maculinea butterflies as Indicators and Tools for Management) is a RTD Project funded by the EU within the 5th Framework Programme (EVK2-CT2001-00126). J. Settele, E. Kühn & J.A. Thomas (Eds) 2005 © 6 PENSOFT Publishers Martin Musche, & Josef Settele Christian Anton, Studies on the Ecology and Conservation of Butterflies in Europe Sofia – Moscow Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, p. 6 The Threatened Maculinea - Conservation biology as applied to humid zones Alain Rojo de la Paz Université du Maine, Service de Biologie Animale, Faculté des Sciences & Techniques, Avenue Olivier Messiaen, 72085 LE MANS Cedex 9 Contact:
[email protected] Scientific advisors of film project: Fabrice Darinot, Alain Rojo de la Paz & Yves Rozier Realisation: Alain Monclin Most ecosystems have been modified by man. Their preservation has become a major concern and is the very aim of conservation biology. This scientific documentary was realized in the natural reserve of Lavours marshes (AIN, France) and in laboratory at the University of Maine (Le Mans, France). It has pedagogic vocation and presents the conservation biology through the case of the humid zones, using the example of Maculinea butterflies (Lepidoptera: Lycaenidae). Indeed, the three species of Maculinea flying in the european humid zones are found together in the Lavours reserve (M. teleius, M. alcon, M. nausithous). The interest of these butterflies in better understanding the importance of conservation biology is due to their peculiar life-cycle that is presented here by the instance of Maculinea alcon: the caterpillar starts to grow in a plant but will only mature after several months stay in the nest of a red ant. Thus, protecting the Maculinea means protecting the whole system: butterfly, but also the plants and ants it depends on. If one partner of the butterfly, host-plant or host-ant, should disappear, so will the butterfly. Besides the life cycle of the Maculinea, the film shows how the scientists supply the land managers with both information and tools to help them make their decisions and also shows the manner the Lavours reserve is managed to preserve the Maculinea butterflies. Maculinea species are among many others, animals and plants, which are threatened with extinction. The application of conservation biology should help to prevent the extinction of some of these species. It is not just a question of conserving humid zones and their incredible biodiversity: all natural habitats are concerned. Land management has become indispensable. Spiders (Arachnida, Araneae) of a Maculinea alcon – M. teleius pSCI in NW Italy 7 Section 3.1. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Maculinea as indicators 8 Marco Isaia et al. This page intentionally left blank © PENSOFT Publishers Spiders (Arachnida, Sofia – Moscow J. J.A. Thomas Araneae) of a MaculineaStudies on– M. teleius pSCI in&NW Italyin Europe alcon the EcologySettele, E. Kühn of Butterflies(Eds) 2005 9 and Conservation Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, pp. 9-15 Spiders (Arachnida, Araneae) of a Maculinea alcon – M. teleius pSCI in NW Italy Marco Isaia, Simona Bonelli, Marco Montani, Guido Badino & Emilio Balletto University of Turin, Department of Human and Animal Biology, Via Accademia Albertina 13; 10123 Turin; Italy Contact:
[email protected] INTRODUCTION The overall importance of an SCI (Site of Communitarian Importance) may be partially consequential of other aspects than those having promoted its protection. In many cases, in fact, in depth studies of various faunistic groups will offer new conservation perspectives and foster fascinating challenges for multidisciplinary work-teams. Maculinea butterflies are typical representatives of the endangered European biodiversity. All Maculinea species are listed by the IUCN as globally threatened and their conservation is requested by the Habitats Directive. In a broader context, however, it may be important to investigate whether ‘single-species’-based measures, implemented for their conservation, will create suitable conditions also for other species, or to assess the suitability of Maculinea butterflies as biodiversity indicators. Since fens and wet meadows are known to be among the European habitats types that have the highest spider species diversity (Villepoux 1993), the aim of this work is to investigate spider biodiversity at a Maculinea teleius-M. alcon site. As Pesarini (1995) has pointed out in the check-list of the Italian species, the Italian spider fauna is far from being well-known. This work, therefore, will also contribute to a better understanding of Italian spiders. STUDY AREA The NATURA-2000 pSCI “Monte Musiné - Laghi di Caselette (IT1110081)” is located in Piedmont (N-W Italy, 360 m), about 20 km W of Turin, at the bottom of a xeric alpine valley. The site was proposed because of the presence of 8 Habitats Directive (Annexes 2 and/or 4) species, which are also mentioned in the Appendix II of the Berne Convention, (Zerynthia polyxena, Lycaena dispar, Coenonympha oedippus, Euphydryas aurinia, Maculinea arion, M. teleius, Euplagia quadripunctaria, Saga pedo.. As a consequence of the presence of two Maculinea species (M. alcon [Denis & Schiffermüller], 1775 and M. teleius) occurring in strict cohabitation, a small proportion (3 ha) of the pSCI has been studied since 1997. Roberts (1985). More exactly. Q2H. Since 1970 all the area has been lightly cattle grazed and meadows have been irregularly mown for fodder. Because of their striking appearance. May. 1772. Q1G. The area is dominated by a wet meadow with Molinia caerulea Moench. and in spite of their faunistic importance. Argiope bruennichi Scopoli. Spider species were identified on the basis of papers by Simon (1914-1937). All spiders observed during a 30 minutes sampling time were collected with a pooter. MATERIAL AND METHODS Spiders were collected from October 2002 to November 2003. were studied with a stereomicroscope (up to 50x). 250 by sight hunting on standard 9 m2 areas (16 families. 2) were also directly identified in the field in each plot during sampling. As Marc et al. 46 genera and 52 species were collected. The species’ distribution in plots was analysed with Systat® for Windows. filled with 20 ml of ethylene glycol. Heimer and Nentwig (1991).10 Marco Isaia et al. spiders were collected at 12 plots by: • Pitfall traps. Q3B. Q2L). Some other unmistakable species (eg. belonging to 20 families. Quercus pubescens and some Pinus sylvestris. (Fig. Fig. Q2G. For other materials. Grimm (1985). Q2I. Argiope bruennichi (Scopoli. several sampling methods were combined. Lugetti & Tongiorgi (1969).. 21 were caught randomly in the study Fig. 31 genera and 34 species). Q2C. 1) were collected. 2. 154 specimens were sampled by pitfall traps (12 families. Dolomedes fimbriatus (Clerck. Specimens. The surrounding areas are covered with sparse woods with Betula pendula. plots were sorted into 2 groups: (i) 5 plots were chosen in meadows mown once a year (Q1A. Q3D) and (ii) 7 plots in meadows that were not managed since 1990 (Q2B. July and October 2003. most male palps and female epigynes were detached and cleared in clove oil for examination of diagnostic features. (1999) have recommended for all works on spider biodiversity. which made field identification easy. The traps were replaced once a month. 1757) Fig. Q3C. 1. from October 2002 to November 2003. Q2F. Nomenclature follows Platnick’s (2003). 1772) . Sight-hunting was performed in April. RESULTS 425 spiders. Tongiorgi (1966). Depending on the type of grass management. only a few specimens of Dolomedes spp. fixed and stored in ethanol 70% immediately after sampling. 24 genera and 26 species). • Sight-collecting on sample areas of 9 m2. Number of species collected at each plot by the two methods. Araneae) of a Maculinea alcon – M. 1838 NESTICIDAE Nesticus cellulanus (Clerck. 1851) Adults (Months) VII II IV V V I VIII . Taxa DYSDERIDAE Dysdera crocata C L . Fig. Figs 4 and 5 show spider assemblages at each plot. Koch. 1841) Ceratinella brevipes (Westring. 1881 LINYPHIIDAE Centromerus sylvaticus (Blackwall. Steatoda phalerata Panzer. 1. 1757) THERIDIIDAE Robertus sp. Generalised distribution types of sampled spider species. 4. Koch. the generalised distribution types of sampled species are shown in Fig. List of the species found in Caselette’s wetland. 3. 3.Spiders (Arachnida. Fig. 14 genera and 15 species). Months of sampling for each species are illustrated. Table 1. teleius pSCI in NW Italy 11 area (10 families. separately represented for the 2 sampling methods. The species list is shown in Tab. 1801 Theridion impressum L. 1875) DICTYNIDAE Altella sp. 1905) Argiope bruennichi (Scopoli. 1876) Trochosa terricola Thorell. 1817) Pardosa lugubris (Walckenaer. 1841) Trochosa robusta (Simon.L. 1802) Pardosa nigra (C. -Cambridge. Koch. ANYPHAENIDAE Anyphaena accentuata (Walckenaer. Koch.L. 1823 LYCOSIDAE Alopecosa mariae Dahl. -Cambridge. Koch. P. 1757) Aulonia albimana (Walckenaer.L. 1802) LIOCRANIDAE Agroeca lusatica (C. 1881) Tallusia experta (O. 1875) Porrhomma microphtalmum (O. 1841 Tegenaria sp. 1763) Araniella opisthographa (Kulczynski. 1834) Pirata latitans (Blackwall.12 Marco Isaia et al. 1805) Hogna radiata (Latreille. 1772) Cercidia prominens (Westring. 1878 AGELENIDAE Agelena gracilens C. Continued. 1856 PISAURIDAE Dolomedes fimbriatus (Clerck. 1831) Hypsosinga sanguinea (C. 1757) Pisaura mirabilis (Clerck. 1845) Mangora acalypha (Walckenaer. Adults (Months) II X XI II VII-IX X V-VII VI V-VII VI VI V-VIII IV-XI III-VII V V-VI V IV-V VI X X VI V V-VII V-VI VII-X IV XI . 1871) Styloctetor stativus (Simon. 1908 Alopecosa pulverulenta (Clerck.-Cambridge. P. Table 1. 1802) ZORIDAE Zora armillata Simon. 1861) Hypsosinga pygmaea (Sundevall.L. 1757) OXYOPIDAE Oxyopes ramosus (Martini e Goeze. Koch. 1871) TETRAGNATHIDAE Pachygnatha clercki Sundevall. P. 1778) ARANEIDAE Agalenatea redii (Scopoli. Taxa Gongylidiellum vivum (O. Dolomedes fimbriatus. 1871) Adults (Months) V-IX VII VIII V X V-VI VI V-VI VI V-VI V VIII VII-VII 13 DISCUSSION The peculiarity of this site (a wet meadow encapsulated in an overall xeric area) is paralleled by an interesting spider assemblage.L. Pirata latitans). 1826) Talavera aequipes (Pickard-Cambridge. Hogna radiata) as well as grassland species (i. 5). 1839) Zelotes gr. Alopecosa mariae.e. The spider community contains some highly hygrophilous species (i. Araneae) of a Maculinea alcon – M. Temporal trends showed clear seasonal changes. 1866) Micaria sp. 1757) Tibellus oblongus (Walckenaer. Pitfall traps and sight-collected samples were markedly different in terms of spider assemblages. 1802) GNAPHOSIDAE Drassodes lapidosus (Walckenaer.e. diversity. subterraneus SPARASSIDAE Micrommata virescens (Clerck. dominance. pitfall traps sampled the highest number of specimens.L. Hypsosinga sanguinea and Pisaura mirabilis). Koch. In spring (April 2003) and summer (August 2003). teleius pSCI in NW Italy Table 1. Thanatus coloradensis was recorded here for the first time in Italy. 1757) Myrmarachne formicaria (Degeer. Koch. 1802) Drassyllus praeficus (C.L. showing high biodiversity levels. Koch. The statistical analysis of spider assemblages (number of species. 1757) PHILODROMIDAE Thanatus formicinus (Clerck. Continued. 1802) THOMISIDAE Xysticus lanio C. Several rare species (i. Taxa MITURGIDAE Cheiracanthium erraticum (Walckenaer. . 1837) Zelotes electus (C.e. euriecious/stenoecious species ratio) did not allow us to highlight any significant difference among plots.Spiders (Arachnida. showing no influence of grass management type on spider communities. Koch. some xeric and Mediterranean species (i. Trachyzelotes pedestris (C. e. To sum up we can assess that this well-preserved Maculinea site also represents a suitable habitat for a highly diverse spider community. Styloctetor stativus) have been found. 1778) Phlegra fasciata (Hahn. The highest number of sight-sampled specimens was recorded in October 2003. because of the increased number of specimens belonging to weaver species (fig. 1835 SALTICIDAE Evarcha arcuata (Clerck. The highest family richness was recorded in spring. Dolomedes plantarius.L. Germany. Spinnen Mitteleuropas .. A. Naturwiss. Nat. F. CANARD. Spider assemblages according to sampling methods and seasons. & NENTWIG W. Paul Parey. 1969. 5. Fig. YSNEL. Die Gnaphosidae Mitteleuropas .Abh.. MARC P. & TONGIORGI P.. Ricerche sul genere Alopecosa Simon (Araneae – Lycosidae). Spiders (Araneae) useful for pest limitation and bioindication.Ed. Ver. Berlin und Hamburg. (B) 76: 1-100. 318 pp.. 1999.14 Marco Isaia et al. 543 pp. Tosc. . 1991. REFERENCES GRIMM U. Sc. Atti Soc.. LUGETTI G. Agricolture. 1985. Ecosystems and Environment 74: 229-273. Hamburg (NF) 26.. HEIMER S. Boll. University of Michigan. Don Cameron.The spiders of Great Britain and Ireland.. 134(8): 275-334. SIMON E. of Arachnology. Available on-line at: http://research. Edited by Peter Merrett. Calderini. 1993. 1:229 pp. 1914-1937. Catania. Comp. 1985 . teleius pSCI in NW Italy 15 PESARINI C. Colchester. Harv.K. Ed. 361-370. Les Arachnides de France. Mulo Paris. Gioenia Sci. Arachnida Araneae. 14th Eur. The World Spider Catalog. Colloq. Check list delle specie della fauna italiana. VI (1-5) . Accad. 1995.Harley Books.. VILLEPOUX O. In Minelli A. 26. Bologna.. Version 4. Zool. Vol. 1966. e La Posta S. British Arachnological Society and H.Bull. Proc. Nat. 23:1-42.Spiders (Arachnida. . Ruffo S. Italian wolf spiders of the genus Pardosa (Araneae.5.org/entomology/spiders/catalog/index. Araneae) of a Maculinea alcon – M.Ed. Lycosidae) ... TONGIORGI P....html ROBERTS M. 2:204 pp. (eds.. PLATNICK N. The American Museum of Natural History. Etude de la rèpartition des araignèes d’une zone humide.). Mus. 1298 pp. J. 2003. 3:256 pp.amnh.. U. Some meso-xerophilous habitats show relatively rich assemblages of Orthoptera (over 20 spp. Generally.). István A. H-4010 Debrecen. Isophya stysi. István A. arion-site on the Kaszonyi-hill in NE Hungary proved to be the richest site in the species of community interests: Isophya stysi.O. Only one species is typical for all xero-mesophilic habitats.B. rebeli sites. 1). Faculty of Sciences. They were sampled in most sites by sweeping.B. the weakly populated M. The highest species numbers (34 and 33) were observed at two different sites with rather diverse vegetation and also very rich assemblies of butterflies: on the higher plateau of Szilice (Hungarian side. Kühn & J. because they prefer a dense tall-forb vegetation (e.g.unideb.). rebeli-sites of the Slovakian karst and Bükk Mts (21-27 spp. Settele. Saga pedo) and 10 species (2 protected spp.). E.© 16 PENSOFT Publishers Antal Nagy. they were also sampled by pitfall traps in six reference sites of the Aggtelek karst area to measure the effectiveness of sweeping. Hungary Contact: zvarga@tigris. Department of Evolutionary Zoology and Human Biology. H-4010 Debrecen. arion habitat of the Aggtelek karst area and also in the dolomitic rupicolous grassland site of M. The species numbers per site show large contrasts. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. low species numbers were recorded in some extremely dry sites such as the short-grass M. 3.: Arcyptera fusca. arion-site of “Szölöhegy” in the same area and at the rather weakly populated M. Gastropoda). alcon-M. Pholidoptera transsylvanica and Odontopodisma rubripes.O. Seven species are widely distributed in the habitats studied (no protected spp.). with counting and releasing of the “in situ” identified individuals. pp. Aggtelek karst area) and on the high central plateau of the Zempléni Mts (Gyertyánkuti meadows). The floristically rather valuable M.hu Orthoptera assemblages were studied in 23 sites in Hungary and in one neighbouring site of the Slovakian karst (Table 1). and with several biogeographically significant species in other groups (Coleoptera: Carabidae. Hungary 2 Evolutionary Genetics and Conservation Biology Research Group of the Hungarian Academy of Sciences. We used the IndVal method to subdivide the Orthoptera assemblages of the xero-mesophilic habitats (Fig. P. Stenobothrus eurasius). as well. while the second site is one of the few known M. However. 3. The first site belongs to the weakly populated M. However. arion-site on the Teresztenye-plateau also belongs to the richest sites concerning its Orthoptera assemblages (30 spp. Rácz & Zoltán Varga Sofia – Moscow J. P. Another M. rebeli-sites like sites 1 and 3 in the Aggtelek karst area as at the unpopulated site 2. rebeli in the Buda hills (Nagyszénás). Pholidoptera transsylvanica) or they are connected to special edaphic conditions (as Paracaloptenus caloptenoides. with nearly the same number of species at the strongly populated M. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. teleius sites of NE Hungary. the occurrence of biogeographically significant Orthoptera species does not show any correlation with strong populations of Maculinea species. 16 further species characterise the xerothermic habitats of the colline level (2 protected species: Poecilimon fussi. 16-21 Maculinea habitats in Hungary: Orthoptera assemblages Antal Nagy2. Rácz1 & Zoltán Varga12 1 University of Debrecen. Stenobothrus eurasius*) of the sub-montane level (both are .A. Omocestus rufipes. Some humid meadows proved to be also rich in species. 6 and 18). They mostly consist of generally distributed ubiquist species. Chrysochraon dispar. such as the Gyertyánkuti meadows in the Zempléni Mts mentioned above. M.g. The Thamnobionta are also subordinated. Further species (e. Paracaloptenus caloptenoides*) are typical for the higher elevations. rebeli and M. discolor. since they need the tall-forb structure of xerothermic vegetation which is unsuitable for M. Metrioptera roeseli.g. The Tettigonoidea-Acridoidea proportion shows a close connection with the general richness of the assemblages. The Geobiont life-form of Acridoidea only occurs in some dry habitats with open structure of swards (sites 3. The more homogenous Molinieta with low floristic diversity usually show a rather low number of species. They can be considered as “typical species” according to the Habitats’ Directive pointing at the “favourable conservation status” of the sites. Omocestus viridulus. the marshy meadow in the Kiskunság and also in the heath-like variation of the pre-Alpine marshy meadows (Junco-Molinietum nardetosum). Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). Chorthippus parallelus.Maculinea habitats in Hungary: Orthoptera assemblages 17 “rebeli” and “rebeli-arion” habitats). Stethophyma grossum. from the Aggtelek karst area to the pre-Alpine area of W Transdanubia. Thus. nausithous-teleius habitats are: Conocephalus fuscus. Tetrix subulata. The generally distributed species of M. some others (e. brachyptera. 5. Tettigonia cantans. Mecostethus alliaceus. . Chrysochraon dispar) for the mesic or humid habitats. C. the most diverse Orthoptera assemblages have been observed predominantly outside of the strongly populated Maculinea sites. Euthystira brachyptera. the biogeographically significant ones: Pholidoptera transsylvanica*. the heath-like site (Calluno-Genistetum) near Aggtelek. arion. rebeli-site+). slope of karstic dolina (500 m. Rácz & Zoltán Varga Fig. arion-site+). Op: Ocsisnya-ridge. DNy: Eastern margin of the “Dénes” karstic dolina. upper plateau (550 m.18 Antal Nagy. M2: Mogyorós-ridge. bottom of karstic dolina (500 m. arion-° and rebeli-site°). southern slope of karstic dolina (500 m. bottom of karstic dolina (500 m. M1: Mogyorós-ridge. Szt: Xeric grassland with Stipa joannis. E: Szőlő-hegy. Dlakv: Eastern slope of “Dénes” karstic dolina (500 m. arion-site+). Results of the IndVal Analysis of the Orthoptera assemblages (Cluster: Bray-Curtis. M3: Mogyorós-ridge. rebeli-site°). western slope of karstic dolina (500 m. not populated). István A. arion-site°). rebeli-site°). + : strong population. upper plateau (500 m. 20ta: Haragistya plateau. arion-site). rebeli-site°). Ol: Ocsisnya-ridge. unpopulated). 20tf: Haragistya plateau. upper edge of karstic dolina (500 m. not populated). Ob: Ocsisnya-ridge. 1. southern slope of karstic dolina (550 m. rebeli-site°). bottom of karstic dolina (unpopulated). Dtdp: Southern margin of the “Dénes” karstic dolina (500 m. abandoned hayfield (400 m. SzhK: Szőlő-hegy middle level (350 m. arion-site°). rebeli-site°). SzhA: Szőlő-hegy lower level (300 m. ° : weak population . MISSQ) The studied sites: SzhF: Szőlő-hegy upper level (400 m. 24. arion sites in the Aggtelek and Slovakian karst area: 1.M. Tohonya-ridge N of Jósvafő. Plateau of Silica.and M. 21. Rakaca-valley near Meszes. Kecskeláb-rét. N of Jósvafő (M. Maculinea rebeli and M. arion ligurica-site in hilly part of the Bereg lowland. teleius sites in the Zempléni-Mts: 12. 11. rebeli. 3. teleius sites in the Cserehát hilly region: 18. nausithous sites in SW-Hungary. 22-26. Gyertyánkuti-meadows. Maculinea alcon. Maculinea alcon. Bükkszentkereszt. N of Jósvafő. HU side. rebeli. 6. 20. 5. HU side. 4. alcon-M. 18-19. Carex-Alopecurus meadow near Szentgyörgyvölgy. 23. Barakonyi-valley. NardoCallunetum. Teresztenye-plateau in the S part of the Aggtelek karst. Succiso-Molinietum near Apátistvánfalva. Deschampsia facies. 12-13. 8. 7. Drahos-meadow. M.Table 1. teleius-site in the Kiskunság lowland area. 13. 25-26. arion-sites near Aggtelek: 16. Nagymező. 17. 4 1 0 1 0 1 0 1 0 0 0 1 0 0 0 1 1 0 0 0 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 0 0 1 1 0 1 1 0 0 1 1 0 0 0 1 1 1 1 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 0 1 0 1 0 0 0 1 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 1 0 0 1 0 1 0 1 0 0 0 1 0 0 1 0 0 0 1 0 0 0 1 1 1 1 1 0 0 1 1 0 0 1 0 0 0 1 0 0 1 0 1 1 1 1 0 0 0 1 1 0 1 0 0 0 0 0 1 1 0 1 1 1 0 0 0 0 1 1 0 1 1 0 0 1 0 0 1 1 1 1 1 1 0 1 0 1 1 0 0 1 0 0 1 0 1 1 1 0 1 0 0 0 0 0 1 1 1 0 1 0 1 1 0 0 1 1 0 1 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 1 1 1 0 0 0 1 1 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 1 1 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 1 1 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 1 1 0 1 1 0 0 0 0 1 1 0 0 0 0 0 1 0 0 1 0 1 1 1 1 0 0 0 1 Species 1 2 3 Maculinea habitats in Hungary: Orthoptera assemblages Phaneroptera falcata 1 Phaneroptera nana 0 Isophya kraussi 0 Isophya stysi +* 0 Poecilimon fussi + 0 Leptophyes discoidalis + 0 Leptophyes albovittata 1 Barbitistes constrictus 0 Conocephalus fuscus 0 Conocephalus discolor 0 Tettigonia viridissima 1 Tettigonia cantans 0 Metrioptera roeseli 0 Metrioptera brachyptera 0 Metrioptera bicolor 1 Platycleis grisea 1 Pholidoptera transsylvanica +* 0 Pholidoptera aptera 0 Pholidoptera griseoaptera 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 1 19 . 10. Maculinea rebeli sites in the Bükk Mts: 9. Örség: 22. Maculinea teleius-M. M. Szőlőhegy E of Jósvafő. teleius site in the Szatmári-lowland. Molinietum. 2. Slovakian Karst. Xeric site NE of the Tohonya-ridge. M. 15. Molinietum typicum and Molinietum Nardetosum near Kétvölgy. arion-site). poorly populated by M. Succiso-Molinietum near Gödörháza. Kuriszlán NNE of Jósvafő. Lófő-tisztás. Plateau of Silica. Maculinea teleius. 9-11. 14. arion site. densely populated by M. Orthoptera in Maculinea habitats of Hungary (binary data of the relevés of 2002) 1-8. 19.M. M. NW of Hacava. Zádielska planina. Maculinea rebeli site in Buda hilly region: Nagyszénás. 16-17. 20 Table 1. Continued. 4 1 1 1 1 0 1 1 0 1 0 0 0 0 1 0 0 0 1 1 0 0 1 0 0 1 0 1 0 1 0 1 1 1 0 0 0 0 0 1 0 1 1 1 0 0 0 1 0 1 1 0 1 0 1 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 1 1 0 1 1 1 0 0 1 1 1 1 1 0 0 1 0 1 1 0 0 0 0 1 1 0 0 0 0 1 0 1 1 1 0 0 1 0 1 1 1 1 1 1 0 1 1 0 0 0 0 1 1 0 0 0 0 1 0 1 1 1 0 0 1 0 0 0 1 1 1 1 0 1 1 0 1 0 0 1 0 1 0 0 0 1 1 0 0 1 0 0 1 0 1 1 1 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 1 1 0 0 0 1 0 0 1 0 1 1 1 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 1 1 1 0 0 0 0 1 1 1 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 1 1 0 1 0 1 0 0 0 0 1 1 1 0 0 1 0 0 1 0 0 0 0 1 0 0 1 1 1 1 0 0 0 1 0 0 0 0 1 0 1 0 0 1 0 0 1 0 0 0 0 1 0 0 0 1 1 1 0 1 0 1 0 1 0 0 1 1 0 0 0 1 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 1 0 1 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 1 1 1 1 0 0 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 1 1 0 1 1 0 0 0 0 0 1 1 1 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 1 1 0 0 0 1 0 0 1 1 0 0 0 1 0 1 0 0 1 0 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Species 1 2 3 Antal Nagy. Rácz & Zoltán Varga Pholidoptera fallax 1 Pachytrachis gracilis + 0 Rhacocleis germanica + 0 Decticus verrucivorus 1 Gampsocleis glabra + 0 Ephippigera ephippigera 0 Gryllus campestris 1 Acheta deserta 1 Oecanthus pellucens 0 Odontopodisma schmidtii + 0 Odosntopodisma rubripes +* 0 Pseudopodisma nagyi 1 Paracaloptenus caloptenoides +* 0 Calliptamus italicus 1 Stethophyma grossum 0 Parapleurus alliaceus 0 Chrysocraon dispar 0 Euthystira brachyptera 1 Oedipoda coerulescens 0 Psophus stridulus 0 Stauroderus scalaris 0 Stenobothrus lineatus 1 Stanobothrus eurasius +* 0 Stanobothrus nigromaculatus 0 Stenobothrus crassipes 1 Omocestus viridulus 0 Omocestus rufipes 1 Omocestus haemorrhoidalis 1 1 0 0 1 1 0 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 1 0 1 1 1 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 1 1 0 0 0 1 0 0 1 0 1 1 . István A. * : species of Community Interest (Annex II-IV) 21 . Continued.Table 1. 4 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 18 14 24 9 17 6 0 1 1 1 0 1 0 0 0 1 0 0 0 1 0 1 0 1 1 1 1 1 0 1 1 0 1 1 0 0 0 0 0 1 1 0 0 0 1 0 0 1 0 1 0 1 0 1 1 0 0 1 0 0 1 1 0 1 0 1 0 1 1 0 0 0 0 1 1 1 0 1 0 1 0 1 1 1 0 1 0 0 0 0 0 1 1 1 0 1 1 0 0 1 0 0 1 0 0 1 1 0 0 1 0 0 0 1 0 0 0 0 1 1 0 1 1 1 1 1 0 1 0 0 0 1 1 1 0 1 0 1 0 0 0 0 0 1 0 0 0 1 0 1 0 1 1 1 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 1 0 1 0 1 0 1 0 0 1 1 0 1 1 1 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 1 0 1 0 1 0 1 0 0 0 0 1 1 14 25 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Species 1 2 3 Omocestus petraeus Chorthippus albomarginatus-oschei Chorthippus paralellus Chorthippus montanus Chorthippus dorsatus Glyptobothrus apricarius Glyptobothrus brunneus Glyptobothrus biguttulus Glyptobothrus mollis Euchorthippus declivus Gomphocerippus rufus Arcyptera fusca + Tetrix subulata Tetratetrix bipunctata Mantis religiosa + 0 0 0 0 1 0 1 0 1 1 1 1 0 0 0 1 1 0 1 0 1 0 1 1 0 1 0 0 0 0 1 0 1 0 1 1 1 0 1 0 1 0 0 1 1 Number of species 24 20 23 23 27 34 19 30 24 27 15 22 33 20 27 12 28 10 Maculinea habitats in Hungary: Orthoptera assemblages + : protected species in Hungary. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. An important key factor for the survival of the Large Blue Butterfly is the occurrence of Myrmica sabuleti (Meinert. METHOD We investigated the above mentioned parameters and studied M. Elmes et al. It is especially important to know this in regard to habitat management measurements. Germany Contact: regina@pauler. insolation.de INTRODUCTION Maculinea arion (Linnaeus. 1846) and Myrmica schencki (Emery. PROBLEM Myrmica sabuleti lives in different niches across Europe (Elmes & Wardlaw 1982. but a certain soil temperature range seems to be crucial for its presence. 72076 Tübingen. Thomas 2002). Settele. Kühn & J. sabuleti and the cooccurring Myrmica species in two M. manfred. Erbprinzenstr. sabuleti has different habitat requirements in terms of temperature. sabuleti in the range of M. pp. 1894). 1758). 22-25 Habitat preferences of Myrmica (Hymenoptera: Formicidae) ant species in Maculinea arion (Lepidoptera: Lycaenidae) sites in South-Western Germany Regina Pauler-Fürste1 & Manfred Verhaagh2 ¹ University Tübingen. 1860) in sufficient numbers. arion distribution. HYPOTHESIS We hypothesized that M.verhaagh@smnk.© 22 PENSOFT Publishers Regina Pauler-Fürste & Manfred Verhaagh Sofia – Moscow J. Seifert 1986. 1998). 1989. 1996. D-76133 Karlsruhe. because it seems to be the only suitable host species for the caterpillars of this butterfly (Thomas et al. 1988. 1758) is an obligate myrmicophilous lycaenid butterfly with one centre of palaearctic distribution in Central Europe (Wynhoff 1998). each .net. E. Germany ² State Museum of Natural History. vegetation structure and vegetation parameters in our latitudes than the co-occurring Myrmica species Myrmica rubra (Linnaeus.A. arion habitats in the Swabian Jura in SW-Germany. Myrmica scabrinodis (Nylander. Institute of Zoology. Department of Entomology. This takes the form of an along-latitude ecological compensation (Thomas 1998). Auf der Morgenstelle 28. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. which does not allow generalizations of research results to be made concerning the ecology and habitat requirements of the host ant M. 13. scabrinodis (scab) and M. F (indicator value for humidity. schencki (sch) in boxplots. Highly significant differences lead to rank positions in a temperature gradient.outlier. 1996 and unpublished data). M. rubra. . M. and 75.extreme value. 1. Differences in preferences for these parameters were investigated through statistical analysis. percentile. Differences in temperature between habitat and nest sites of the M. arion (Linnaeus. We took the “indicator values” (“Zeigerwerte”) for every occurring plant species from Ellenberg et al. (1991) to calculate the vegetation parameters. sabuleti occurred at the investigated sites among a broad range of vegetation types. plant species composition and inclination at the nest-entrances of nearly 500 Myrmica nests. average (median). rubra (rub). highest and lowest value. from very short to relatively tall vegetation on southern-exposed slopes. RESULTS Temperature measurements The temperature values of the test-sites were compared against each other. inclination and exposure. 1860) and co-occurring M. sabuleti (sab). 1991) for the Myrmica species: M. sabuleti preferred warmer microhabitats compared to M. M. We used beneath surface temperature (electronic data records) from various testsites which differed in vegetation type. Boxplots with: O . 1758) were investigated through statistical analysis.Habitat preferences of Myrmica ant species in Maculinea arion sites 23 population with about 320 butterflies estimated per season (Pauler-Fürste et al. 1758) host ant M. with SPECIES Fig. sabuleti (Meinert. * . inclination was measured with an inclination gauge. PD-values (Seifert 1986) describe the density of above surface vegetation. rubra (Linnaeus. 25. We also characterised vegetation structure. M. Ellenberg et al. arion in SW-Germany (Pauler-Fürste et al. arion habitat concerning exposure and inclination of a patch. 1995) with corresponding abundance of T. 8. sabuleti is strongly dependent on abiotic components like differences in exposure and inclination of the site. nitrate. “summer” and “autumn” were also highly significant. humidity. 3. 1996).g. 1991) and nitrogen content of the soil (N = indicator value for nitrate. which could be characterized more concretely (Pauler-Fürste in prep. Ellenberg et al. sabuleti thus seems to tolerate a relatively broad range of vegetation structure. 1996) Thymus pulegioides does not (Pauler-Fürste et al. 1991). Therefore we predict that the occurrence of M. Ellenberg et al. for details see Pauler et al. temperature.weekly grazing. Fig. Types of land use (1 – 8.cutting 2 times per year. Vegetation parameters.24 Regina Pauler-Fürste & Manfred Verhaagh a highly significant difference in temperature over the entire period of investigation. 1995). the main host plant species of M.grazing once a year for 1-2 weeks. light.). CONCLUSIONS These results provide direct evidence for the hypothesis that the four investigated Myrmica species have different habitat preferences in M. Influence of land use on the abundance of Thymus pulegioides (modified from Pauler et al. 2. ph-value and plant density. Differences in the seasonal sections “spring”. Ellenberg et al. summer and autumn in the different types of land use: 1. 1991) were highly significant and lead to rank positions for the four Myrmica species (e. The abundance Fig. 1). 1995.grazing once in 14 days. 7. . except T (indicator value for temperature. pulegioides according to the scale of Londo (1975) together with humidity (F= indicator value for humidity.grazing once a year or once in two years for less than 1 week.daily grazing (temporary fold). While the host ant M. 4. Frequency of grazing in spring. rather more than on slight differences in vegetation structure in our region. Pauler-Fürste et al. vegetation structure and insolation Differences in all parameters mentioned.cutting once a year or once in two years.daily grazing. 6. 2. 5. 2. (1998).R. J. REFERENCES Ellenberg. (1991). Pedobiologia. G. arion habitats in SW-Germany in order to increase the coverage of Thymus to a sufficient degree (Poethke et al. by Myrmica ants. P. Thomas. SUBSIDIARY CONCLUSION M. Effects of latitude. J..W.. D. J. Hochberg. 275-281). Maculinea arion (large blue butterfly).. 1996). Scripta Geobotanica. 2). Wynhoff.Augsburg: Naturbuch Verlag. Host specificity among Maculinea butterflies in Myrmica ant nests.W. 39-46. Clarke. Wirth. Journal of Insect Conservation. Oecologia. 1-124.J.. (1994). (1988).(1st ed. Londo. E. Göttingen: Goltze. G. Zeigerwerte von Pflanzen in Mitteleuropa. Journal of Insect Conservation. Poethke. C. R. Thomas. Elmes. (1975).. H. H. Therefore intensive seasonal grazing is very important for the management of M. and Paulißen. and Settele. 197-206. and Caucasia (Hymenoptera. Vegetatio 33 (1). more light and reduction of nitrate in the soil. M. Abhandlungen und Berichte des Naturkundemuseums Görlitz. Seifert. W. Larval niche selection and evening exposure enhance adoption of a predacious social parasite. Weber.Habitat preferences of Myrmica ant species in Maculinea arion sites 25 of T. D. Pauler-Fürste et al. and Woychiechowski M. pulegioides increases with higher grazing intensity (Fig. Düll. J. Aspects on the Population Vulnerability of the Large Blue Butterfly.). Abhandlungen und Berichte des Naturkundemuseums Görlitz 62. Kaule. 16. A taxonomic revision of the Myrmica species of Europe. G. Elmes. Glaucopsyche (Maculinea) arion. Wardlaw. Settele... J. Variations in populations of Myrmica sabuleti and Myrmica scabrinodis. can be found on regularly and intensively seasonal grazed sites (Pauler et al.A. 1-75. 15-27.. Individuenbasierte Modelle als Entscheidungshilfen im Artenschutz. 147-186. The decimal scale for releves of permanent quadrats.. A. Nachrichten des Entomologischen Vereins Apollo.C. 18. Henle (Eds. R. Pauler-Fürste. B. 59. . and Simcox. Oecologia. (1995). Pauler. I. A. 90-97. Griebeler.. 1995. (2002). Kaule. due to several reasons like competitive advantage.J. C. R. Ameisen: beobachten. J. Species Survival in Fragmented Landscapes (pp. sabuleti may also occur in adequate nest densities on not very intensively grazed sites on south-facing slopes in our latitudes. and Settele. In: J.. (1998). 67-78. 2.. (1998). M. Wardlaw. 531-537. E. J. Seifert. in South-West Germany. Poschlod & K. (1996). Zeitschrift für Ökologie und Naturschutz. Seifert B. T. J. (1986). Thomas. (1996).Werner. 1994). however. Significant higher densities of the Large Blue Butterfly and therefore more stable populations. and Pauler. M. 23. Untersuchungen zur Autökologie des Schwarzgefleckten Ameisenbläulings Maculinea arion LINNAEUS 1758 (Lepidoptera: Lycaenidae) in SüdwestDeutschland. G. Verhaagh. 248pp. & Wardlaw. R. Elmes. 452-457. altitude and climate on the habitat and conservation of the endangered butterfly Maculinea arion and its Myrmica host ant.. E. V. The ecology of Myrmica ants in relation to the conservation of Maculinea butterflies. 3. Formicidae). B. G. Asia Minor.). A. (1989). W. 61-64. R. Journal of Insect Conservation. Thomas. 132. G. 79. bestimmen. (1982). Margules. H. Vergleichende Untersuchungen zur Habitatwahl von Ameisen (Hymenoptera: Formicidae) im mittleren und südlichen Teil der DDR.. The recent distribution of the European Maculinea species. 2. Dordrecht: Kluwer. J. both Viola species were significantly more abundant in the soil near Myrmica nests. In the field. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. A key question was: Does management for M. UK Contact: ZR@ceh. Simcox. Dorchester. Settele. and various butterflies). The specific aim of management has been to achieve a wide distribution of flowering Thymus praecox in co-existence with high densities of the ant Myrmica sabuleti.ac. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. neither Viola was attractive to the other ants (mainly two Formica species) that inhabit these study sites. at least 20 Red Data Book species have increased under this management. although it is unclear whether this is an inherent or an acquired trait. through grazing quite heavily in spring and autumn in order to create the warm soil micro-climate required at these times of year by M. Zoe Publishers Sofia – Moscow J. This resulted from the attractiveness of elaiosomes on the seeds of both Viola species.3 cm height on well drained southern-facing slopes. b). sabuleti under UK climates (Thomas et al 1998 a. this is achieved by maintaining early successional grassland swards of 1 cm . Judith C. pp. a cockroach. where they are believed to receive protection from herbivores (enemy-free space) and perhaps superior soil conditions for establishment. but the relative density of Viola riviniana to V. Karsten Schönrogge. it is also necessary to burn the Ulex scrub on cycles of 5-7 years. where the elaiosomes were eaten and the seeds then scattered around the nest. 26-27 Myrmica ants as keystone species and Maculinea arion as an indicator of rare niches in UK grasslands Zoe Randle. On the acid grassland sites of south-west England. Kühn & J. Thomas NERC . Wardlaw & Jeremy A. as well as more common but characteristic species ranging from grasshoppers and tiger beetles to plants. David J. E. At least two Myrmica species preferred the species of Viola that coincided most with its niche. However. On restoration sites around Dartmoor. Dorset DT2 8ZD. beetles. Winfrith Newburgh. In practice. selene and their foodplants Viola lactea and V.© 26 PENSOFT Randle et al. CEH Dorset.Centre for Ecology & Hydrology. Among the RDB species that have benefited are four coupled species: the butterflies Boloria euphrosyne and B.uk Our studies indicate that alpha and beta biodiversity increase on former and potential sites managed to restore the optimum habitat of Maculinea arion in the UK (Elmes & Thomas 1992. lactea was primarily determined by . arion merely create a scarce niche shared by a guild of other species. Winfrith Technology Centre. ants. or is there a direct impact due to the increase in Myrmica sabuleti or other Myrmica ant colonies? We found that there was a positive association between Myrmica and Viola densities. Each Viola species was highly attractive to all Myrmica species tested in the lab and was carried to their nests. plants and various insects (robber and bee flies. including rare birds.A. bugs. The aim was to investigate the processes explaining their increases and to explore any interactions each might have with various Myrmica species. riviniana. Thomas 1999). Journal of Insect Conservation 2: 39-46. Wardlaw. 1991 Rare species conservation: case studies of European butterflies. & Clarke. euphrosyne is generally in steep decline elsewhere in Britain.Myrmica ants as keystone species and Maculinea arion as an indicator of rare niches 27 microclimate. Spellerberg. lactea. with V. . J. M. lactea replacing V. Eds. G. Thomas.A. J..A. Dempster & I. We conclude that targeted management for Maculinea arion in the UK has profoundly beneficial effects on the wider community of organisms characteristic of its grasslands. 1998. & Morris. R. D.W. coinciding with Myrmica ruginodis colonies. G. Clarke.G.A. & Hochberg. euphrosyne has increased greatly on two experimental sites. As a result. 1999 The large blue butterfly – a decade of progress.. In The scientific management of temperate communities for conservation. Biodiversity and Conservation.F. We found that B. Boloria selene inhabits a cooler niche. Blackwells. Chapman & Hall. McLean. Ed by J. riviniana growing in more shaded vegetation.. M.T.G. Elmes..P. J. B. I.A. Hochberg. Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). REFERENCES Elmes. due to the restoration of rare niches within the biotope in combination with the direct (keystone) effect of high densities of Myrmica ants impacting upon mutualists and other plants and animals. sabuleti. Thomas. with about 80% of known populations going extinct in the past 30 years and the surviving ones declining to an average of < 10% their former densities. Elmes. British Wildlife 11: 22-27 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). 1: 155-169 Thomas. Symposia of the Royal Entomological Society 19: 261-290. and a large new population has established on a third site following its creation (from dense scrub and bracken) in the first year of the MacMan project. In Insect population dynamics: in theory and practice. riviniana in the hottest niches.A. Simcox. J. In contrast. & Thomas..W. R. J. B. euphrosyne oviposition was apparently influenced by microclimate as well as by the growth form of Viola.. Management for M. 1992 The complexity of species conservation: interactions between Maculinea butterflies and their ant hosts. euphrosyne.J.E. straddling the warmest microclimates occupied by V.T. Oxford. the density of B.W. G. J. altitude and climate on the habitat and conservation of the endangered butterfly Maculinea arion and its Myrmica ant hosts. riviniana and the coolest ones occupied by V. Goldsmith. BES Symposium 29: 149-197. Like all the Viola-feeding fritillaries. arion increases both the larval food plants and the specific niche selected by each Boloria species. Boloria euphrosyne is extremely selective during oviposition and deposits eggs on only a small minority of the available violet plants (Thomas 1991). but especially that of B.E. 1998 Effects of latitude. London Thomas. M. ovipositing mainly on V. In contrast. and happens to coincide with the niche of M.C. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Winfrith Newburgh. Dorchester. 28-31 Maculinea and myrmecophiles as sensitive indicators of grassland butterflies (umbrella species). David J. Using new analyses of comparative changes in UK species that take account of recording artefacts (Thomas 2005). E.Centre for Ecology & Hydrology. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Karsten Schönrogge1. Department of Community Ecology. Thomas et al. as May. Graham W. and it is the rare and local species in a taxon .ac. Sofia – Moscow J. comparisons of the proportion of species believed to have become extinct in different taxonomic groups will be biased if the groups being compared experienced different levels of past recording. Elmes1. and are social parasites. about 40% are beetles. This occurs because the early species’ lists for under-sampled groups contain a disproportionately high representation of common widespread species. conservationists have inevitably relied on extrapolations from a few well-recorded. CEH Dorset. Butterflies are the only practical group to use in most parts of the world. Wardlaw1 & Josef Settele2 NERC . of which most are known from a single specimen or site (Anon 2003).that are especially prone to extinction.which tended not to have been recorded in the first place . Of these. even more sensitive in their population responses than the mutualistic myrmecophiles? It was recently suggested that the widespread use of butterflies as indicators of change is inappropriate because butterflies have apparently experienced amplified extinction rates in Britain compared to other insect groups (Hambler & Speight 2004). UK 2 UFZ . Lawton & Stork (1995) and McKinney (1999) point out. Kühn & J. With such poor data on which to base priorities. pp. 06120 Halle. However.© 28 PENSOFT Publishers Jeremy A. Winfrith Technology Centre. Nevertheless. conspicuous insect groups in order to assess change among invertebrates. but are they representative of other taxa? Moreover. among butterflies.92). 4. Simcox1. we showed that butterflies are surprisingly representative of change in most of the other main insect groups (Fig 1). Thomas1. Dorset DT2 8ZD. do the roughly 25% of butterfly species that interact with keystone ant species (myrmecophiles) provide such an early warning system. Ralph T. exemplified by the genus Maculinea. despite the closeness of fit (r2 = 0. Zoe Randle1.A.uk 1 Only one fifth of the estimated 4-5 million species of insects living in the world have been discovered and described to date. are the more specialist species particularly sensitive to environmental change and do they provide an early warning system. ants (keystone species) and other invertebrates Jeremy A. it should be recognised that butterflies do not represent the inhabitants of two types of habitat in which declines are known . alerting us to changes that will eventually impact on more generalist species? In particular. Judith C. Germany Contact: jat@ceh. Clarke1.Centre for Environmental Research Leipzig-Halle. Theodor-Lieser-Str. Settele. Least squares fitted line: Q = 8. Among the different species of UK butterfly experiencing change. (6) 47 ant. respectively.Maculinea and myrmecophiles as sensitive indicators of grassland butterflies 29 Fig. Finally. among myrmecophilous and other butterflies. ( ) (11) Butterflies. we have recently shown that butterflies have experienced greater declines than plants and birds in the UK (Thomas et al 2004). rotting) trees and freshwater systems. of which myrmecophiles comprise 36% of the total (our calculation). (ii) Butterflies are sensitive indicators of future change in vertebrates and plants. Hence butterflies (and by implication other insects) can be considered particularly sensitive bio-indicators of environmental change compared to plants and vertebrates. (4) 266 hoverfly. because their reaction to environmental degradation or perturbations is amplified and rapid.0996U . In contrast. the social parasite Maculinea arion was one of four species to go extinct in the 20th century in the UK.92. Warren et al (2001) demonstrated that specialist species. Furthermore. We tentatively conclude that: (i) Butterflies are useful (average) indicators of similar changes in other less conspicuous insect groups. (9) 32 mosquito. Percentage (Q) of insect (+ spider) groups considered to have become extinct in c. comprising 18% of the extinct butterfly species of that nation. ( ) Other groups: (1) 900 other Macrolepidoptera. the second most rapid type of butterfly to experience population extinctions on these same sites were the mutualistic myrmecophiles. (3) 612 weevil. during the past quarter century. . 1900. had declined to a significantly greater extent than generalists.001.13e-0. arion has a history of being the species of butterfly whose population generally went extinct first on 32 individual UK grassland sites that were monitored over a period of 150 years (Fig 2). p < 0. non-myrmecophilous UK specialists (sensu Pollard & Eversham 1995) persisted for longer (Fig 2). r2 = 0. albeit unrepresentative of saproxylic and freshwater species. Interestingly. 1900-1987 in relation to the percentage (U) of native species in current British lists that were unknown in c. in the UK. M. (10) 26 bumblebee species. (2) 622 spider. of which 10% of species are myrmecophiles. From Thomas & Clarke 2004 to have been high: saproxylic (ancient. For these we recommend. In addition to these comparative changes of insects. whereas in the Netherlands three out of four Maculinea species went extinct. increased monitoring of the whole habitat and of adult Odonata. (5) 154 macro-Brachyra. (7) 43 dragonfly. This represents regional extinctions at a scale that is 2-3 orders of magnitude greater than that of a typical butterfly meta-population. 1. (8) 38 grasshopper-cricket. ÷2 = 26. S. REFERENCES Anon 2003 Monitoring conservation for biodiversity. Fig. Melanargia galathea). H. J. May. Euphydryas aurinia. mutualistic myrmecophile extinct before non-myrmecophilous specialist. 1563 R.. Asher. ÷2 = 13. R.7. 1879-81 . 2df. C. A. Pyrgus malvae. ed A. G. 13. birds and plants and the global extinction crisis. D. Argynnis adippe. Preston. N.. pp 23-36. In Extinction rates.7. Social parasite extinct before mutualistic myrmecophile. B. C. M. C. Lawton. J. In Ecology and conservation of butterflies. 2df. Hamearis lucina. p<0. J. myrmecophiles are more sensitive to environmental change than other specialist (or generalist) species.. Biol. E.. & Clarke. giving this genus a useful role as an early warning system of future change. R.. Lawton. The Royal Society.001. M. 1995 Butterfly monitoring. Lawton. R. hatched. J. 1999 High rates of extinction and threat in poorly studied taxa. & Speight.T. Conserv.. Science 305. OUP. M. 339-357 Thomas. J. 1563-4 Thomas. L. vertical Lysandra coridon. 12731281. 2004 Extinction rates and butterflies. p<0. Roy. McKinney. Science 303. The order in which 14 butterfly species went extinct in 32 UK grasslands that were frequently monitored for periods of up to 150 years (sources: J A Thomas and BRC archival data.D. R. 2004 Extinction rates and butterflies. and the socially parasitic group Maculinea appears to be more sensitive still. non-myrmecophilous specialists (Boloria euphrosyne. Science 305. Fox. Black. Stork 1995 Assessing extinction rates. London Thomas. H. J. white. B. London. Chapman & Hall. dots Thecla betulae). 2005 Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups. Thomas et al. J. left diagonal Lysandra bellargus.. selene. R. Oxford. Phil Trans Roy Soc B 360. Horizontal lines represent different species.. B. Clarke.30 Jeremy A. D. Telfer. pp. M. 2004 Comparative losses of British butterflies. Pullin. interpreting the changes. E & Eversham. A. A. social parasite (Maculinea arion).001 (iii) Among butterflies. G M Spooner pers comm). Pollard. Leptidea sinapis. Greenwood. Hambler. mutualistic myrmecophile (right diagonal Cupido minimus. Hipparchia semele. May. 1-24. 2. Eds J. M. T. H. J. G. M. A. P. Fox. G.. Telfer. D. K. C. S. D... Huntley. Jeffcoate. Hill J. N. Asher..Maculinea and myrmecophiles as sensitive indicators of grassland butterflies 31 Warren. D.. G. B.... Thomas. & Thomas. Willis. S. Nature 414: 65-69 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126) . Moss... Harding. J. M. 2001 Rapid responses of British butterflies to opposing forces of climate and habitat change. J. J. S. Greatorex-Davies... B. Roy.. R. Jeffcoate. Brenthis hecate+. 1088 Budapest. P. Callophrys rubi. Z. skippers and burnets were estimated by linear census along transects in at least 3 seasons (W Hungary: Őrség. Adscita budensis+. H-4010 Debrecen. loti.unideb. pp. Some localised species occur only at a few sites. site 1-2 (2002) are presented in Table 2. Melitaea punica kovacsi*+. Mellicta athalia. 32-44 Assemblages of butterflies and burnets in Maculinea habitats of Hungary Zoltán Varga1. butterflies. coridon.2. Sipos on the EcologySettele. Hungary Contact: zvarga@tigris. Widely distributed species of these habitats are: Thymelicus silvestris.13. Studies and Conservation in Europe Sofia – Moscow Vol. P. Table 1) These assemblages are generally rich in species. Plebeius sephirus*. P. 55-99 species per site have been observed.hu At some intensively studied sites. E. Zempléni Mts: 2 sites). Aggtelek karst: 3 sites. lineola.B. P. These data are summarised in Table 1. arcania. Bükk Mts. NE Hungary. Zygaena brizae+. Z. or the protected species in Hungary: Papilio machaon*. Baross u. bellargus.O. C. J. Protected and European Red Book species* and several biogeographically significant components+ are characteristic for this habitat type. 4 sites. Brenthis ino*. Ochlodes venatus. Nagyszénás-hill. arionsite of the Teresztenye-plateau (99 spp. Aricia artaxerxes issekutzi+. ephialtes pannonica. Polyommatus semiargus. P. 2 sites. Transdanubia: Vértes Mts. Pyrgus carthami.g. Melitaea didyma. Department of Evolutionary Zoology and Human Biology. Coenonympha glycerion. Boloria selene*. trivia+. Sipos2 1 University of Debrecen. . Faculty of Sciences. Lepidoptera assemblages of the xerothermic habitats (sites 1-10 and 18-19. chloros+ . Leptidea morsei*. britomartis. aurelia. decoloratus*. Colias alfacariensis. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Hungary 2 Evolutionary Genetics and Conservation Biology Research Group.: Thymelicus acteon*..A. C. M.!). Iphiclides podalirius*. as Zerynthia polyxena*. Cupido osiris*+. Mesoacidalia aglaja. Zygaena purpuralis. as Lycaena dispar rutila*.O. László Peregovits3 & Julianna V. Z.J. Melanargia galathea. Z. Polyommatus admetus*. M. in mesic habitats also. 3. Plebeius argus. M. Fabriciana adippe. Lycaena virgaureae. Some species of Community Interest have a relatively wide distribution. A.g. on the M. Hungary 3 Hungarian Natural History Museum. T.B. Minois dryas. Kühn & of Butterflies(Eds) 2005 Zoltán Varga. Lopinga achine*. e. In several other sites the diurnal Lepidoptera species that were present were only noted. amandus*. Department of Zoology. P. Z. Adscita globulariae. carniolica. angelicae. L. 3. detailed data of the Aggtelek karst. Conclusions were formulated regarding the management of the sites. Thomas © 32 PENSOFT Publishers László Peregovits & Julianna V. meleager. Arethusana arethusa. filipendulae. Boloria dia. Lycaena alciphron*. tityrus. H-4010 Debrecen. most of them e. Leptidea sinapis/reali. Eriogaster lanestris*. Species associated with forested habitats. 45 and 45. Discussion: generalist vs. Brenthis ino). These sites are generally poor in protected and Red List species. (no protected spp. aethiops. Some few xerothermic species (ponto-mediterranean and southerncontinental) proved to be confined to E (NE) Hungary (Thymelicus acteon. Sideridis lampra (Seseli osseum). Species with over two-thirds presence in humid (“alcon-teleius-nausithous”) habitats were characterised as euryoecous meso-hygrophilic species (two protected species: Boloria selene. The species which occured in both habitat types with over one-third presence were characterised as “meso-xerophilous” spp.g. in the Aggtelek karst area). The mountain meadows of the Zemplén Mts and also the pre-Alpine colline areas of Transdanubia are often rather rich in species (highest numbers are: 73 and 72 species respectively). we will publish these data in special papers. They do not belong to the „standard” set of species of these habitats. Polyommatus amandus.g. Cupido osiris. specialist species As quantitative censuses were only carried out at a few. P. Saturnia pavonia (Rosa sp. Species with over two-thirds presence in dry (“rebeli-arion”) habitats were characterised as euryoecious xerophilic species (only one protected sp. Since they do not refer strictly to these sites. 20-25) These assemblages show several extremes in their faunal composition. We considered the species as “generalists” which occur in both types of habitats and were detected at least in two-thirds of all relevés. we only used a rather simple method for the characterisation of the butterfly assemblages of the habitats. 14. Aricia artaxerxes issekutzi. L. Proserpinus proserpina* (Epilobium sp. since larvae have been found on the food-plants. 15 and 22 sites: 22. The data of light-trapping do not refer strictly to any site. Hemaris tityus* (Knautia arvensis). 25 and 26 species). Assemblages of humid meadows and heath-like habitats (Table 1. ligea) are typical for the pre-Alpine and pre-Carpathian habitats.Assemblages of butterflies and burnets in Maculinea habitats of Hungary 33 Other groups of moths have also been regularly studied by daily observation and light trapping near to sites 1-2. as Lemonia dumi*. Thus. with only some regionally characteristic species belonging to this category: Lycaena hippothoe* and Euphydryas aurinia* in the pre-Alpine area. E.). Euphydryas maturna*. There is only one protected species among them (Papilio machaon). and Argyronome laodice* in NE of the country. more intensively studied sites (e. medusa.g. thus we do not repeat these data here. Malacosoma castrensis (Euphorbia cyparissias). Plebeius idas. there are also some (meso-) hygrophilous species with limited . sites 11-17.).: Lycaena alciphron). Apatura* and Limenitis* spp. The Erebia species (E. Some grass-sedge dominated habitats are rather poor in species (e. botanically more diverse sites from the same area are rich in species (e. Slovakia and Romania also E. the M. etc. Hyles galii (Galium verum). such as Parnassius mnemosyne*. sephirus. E. 13. these are all protected species. 72 species). Alternatively. Melitaea telona kovacsi). in Slovenia. The data on night-active moth species of these sites were collected mostly by light-trapping. most of our data refer on the presence or absence of species at the sites. while neighbouring. 16 and 23-24 sites: 56. rebeli-site on the Tohonya-ridge near Jósvafő. only occur occasionally at Maculinea sites. catax* (on Prunus spinosa scrubs at the edge of the site). in addition they are already published in the volume on the fauna of the Aggtelek National Park. Nymphalis antiopa*. Some protected species* and also some rare food-plant specialists regularly occur at these sites.). Some wide generalists occur in both types of habitats. taraxaci* (polyphagous). ) and Eastern Carpathians of Transylvania usually have almost unicolorous blackish-brown females. J. light brownish-grey underside) are also present among the individuals of the “alconoid” populations (using Gentiana pneumonanthe as the initial food plant). the “rebeloid” populations of the Slovenian karst (Gentiana cruciata. mostly in W Hungary (protected: Lycaena hippothoe. 1758) . and has a taller but patchy vegetation structure with large sprout-colonies of Gentiana cruciata. lutea-feeding on the Nanos Mts. On the contrary. The flying period of the imagos covers the last 10-12 days of June and first 10-15 days of July on both sites. the other also Gentiana pneumonanthe (observation of A. This form was relegated to the nominotypic Maculinea arion arion (Linnaeus. pannonicus. while the populations at the abandoned. rebeli xerophila populations were only observed. Tóth and Z. Nevertheless. due to abandoned mowing of the meadows. The flying period of the “Bükk” populations starts about 5-10 days later than in the Aggtelek karst area.5 km from the basic „reference site” (vegetation relevés. tall-grass sites are weak. the Maculinea species inhabit diverse habitats populated by numerous protected species (Table 3). Argynnis laodice). underside brownish-gray with greenish-blue colouration at the basis of the hindwings. estimation of individual number of the population by MRR. Surprisingly. bluish fore wing discus with black spots of the females. externally different phaenological strains. survey of host ants. Thus. marschallianus. These butterflies are generally medium-sized. etc. arion is now probably the most threatened of all forms of Maculinea in Hungary. Tartally. However. more shiny bluish violet colouration with narrow black margin in the males. rebeli xerophila populations were only found at some similar. We have known for several years that Maculinea arion is represented in Hungary by two. Their habitats are often former. with the exception of some populations of the Aggtelek karst area. An other strong population of M. from occasional grazing. This „ecotype” of M.). László Peregovits & Julianna V. However. and G. in NE Hungary M. they do not occur in the extremely xerothermic. invertebrate sampling.g.or end of June. estimation of egg numbers on Gentiana plants.34 Zoltán Varga. Th. rebeli xerophila was located at a distance of about 1. these characters (e. Phenology and bionomy of some Maculinea spp. some rather weak M. They use early (May-June) flowering Thymus spp. neighbouring sites. A similar situation was observed in the Bükk Mts where two moderately strong and several rather weak M. and are on the wing to mid. Varga) as the initial food plant. tourism). These populations are distributed mainly in central and northern Hungary. praecox) as initial food-plants. abandoned pastures. Th. Euphydryas aurinia*). This site is only occasionally mown. The specimens of these populations have a “rebeloid” appeareance. as a rule. declining.or end of May. in colline-submontane elevations. Relatively stronger populations occur at moderately disturbed sites (i. depending on the weather conditions. rupicolous steppic and swampy-boggy habitats. (Th. they are generally in serious decline. All these data indicate that. that the “typical” population of the Hochschwab area also shows a considerable amount of variation. One of the stronger populations alternatively uses Gentianella austriaca (observation of A. where the host ant species cannot survive either. but also in NE Hungary (protected: Aricia eumedon. The populations of the higher part of the karst area are rather weak. Sipos distribution. we have also observed. Butterflies belonging to the first “ecotype” emerge in short-grass steppic swards at mid. upperside of the wings with dark violet-blue ground-colour and with broad dark margin. rebeli xerophila proved to be the most frequent at a site in the Aggtelek karst region with relatively short-grass structure with only a 30-40% cover value of tall grasses and with very abundant large sprout-colonies of Gentiana cruciata. with predominantly dark. Tartally). blackish-brown females.e. They use mostly Origanum vulgare as the initial foodplant and lay their eggs into the closed flower capitula (Kaszonyi-hill in NE Hungary. but do not occur at the lowland. The black spots of the upperside are often vanishing. Vértes Mts in Transdanubia). This form was relegated to the central Mediterranean subspecies: Maculinea arion ligurica (Wagner. NE Hungary). The underside is silvery grey with pure bluish colouration on the basis of the hindwings. This ecotype seems to be less threatened. However. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). since it can colonise transitional habitats (clearcut sites). with silvery-bluish uppersides and narrow dark margins of the wings. 1904). .Assemblages of butterflies and burnets in Maculinea habitats of Hungary 35 The butterflies of the other „ecotype” emerge in the last days of June or early July and populate two different types of habitats: edges or clear-cuttings of xerothermic oak forests with abundant Origanum vulgare or heath-like dwarf-scrub communities (Calluno-Genistetum germanicae) with cushions of Thymus pulegioides. These populations are distributed from SW Hungary to the NE of the country. also the oviposition was also observed on Thymus pulegioides (Aggtelek. The imagoes are generally large. Bükkszentkereszt. NW Hungary. Grazed grasslands on the hillfoot of the Polovnik Mts. Molinietum typicum near Kétvölgy. S part of the Aggtelek karst. László Peregovits & Julianna V. near to the Soca river. arion ligurica-site in hilly part of the Bereg lowland. Maculinea rebeli-site in the Vértes hills near Vérteskozma. flower-rich montane hayfields. Örség area: 21. 7. 25. Sampling periods: Middle-end of May and early-middle July. 18-20. 10. Some sites in W Hungary (18. Bükk-plateau. Molinietum. 23. Torockó. Zádielska planina near Hacava. 19. Nagybajcs. Ágfalva. 6. Faunal lists of butterflies and burnets in Maculinea habitats of Hungary and neighbouring areas Sites 1-17: NE Hungary. teleius site in the Szatmári-lowland. Maculinea teleius-M. 15. Molinietum Nardetosum near Kétvölgy.M. Central Hungary. Bükkplateau.M. Maculinea arion-site on the Teresztenye-plateau. Junco-Molinietum. 21-25. Carex-Alopecurus meadow near Szentgyörgyvölgy. . M. Kiskunság lowland area. Lófő-tisztás. 28. early August 2002. sites as in the vegetation relevés and Orthoptera samplings. Maculinea rebeli-sites in the Bükk Mts: 4. 4-6. Tohonya-ridge N of Jósvafő. 21-25: SW Hungary. calcareous rupicolous grassland. 13. NW Hungary. 3. 8. Drahos-meadow. 26-27: NW Hungary. calcareous rupicolous grassland. Barakonyi-valley. Kupás-valley near Gyilkos-tó (Lacul Rosu). Transylvania. 5. 9. 1-3. Nagymező. 20.26. M. Maculinea teleius-sites near Aggtelek: 14. Maculinea alcon . Aggtelek karst. NE of Jósvafő. Slovenian karst: Maculinea rebeli-sites. 22. 32-34: Romania. teleussite near Kunpeszér. 25. Kecskeláb-rét. teleius sites in the Cserehát hilly region: 16. Maculinea alcon . 12. Romania. 4 1 1 1 0 1 0 1 1 0 0 0 0 1 0 1 1 0 1 1 0 1 0 1 1 0 0 0 0 0 0 1 1 1 1 0 0 1 0 1 1 1 0 0 0 1 0 1 1 1 1 1 0 1 0 0 0 0 0 1 0 1 0 1 1 1 1 1 1 1 0 1 1 0 0 1 0 1 0 1 1 1 0 0 0 1 0 1 1 0 0 1 0 0 0 0 0 1 1 0 0 1 0 1 1 0 0 0 0 1 1 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 1 1 1 1 1 0 1 1 1 1 0 0 0 0 1 0 1 1 1 0 0 0 0 1 1 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 1 1 1 1 1 0 0 1 0 0 1 0 0 1 0 0 0 0 1 1 1 0 0 1 0 1 1 1 0 0 1 0 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 0 0 0 1 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 1 1 1 0 0 1 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 1 1 1 0 0 1 1 1 0 0 0 0 0 0 1 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 1 0 0 0 1 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 1 0 0 0 0 0 0 Zoltán Varga. Xero-mesophilous tall-grass meadow on the Trnovski Gozd. 17. N of Jósvafő. 26) and the sites outside of Hungary (27-33) have been surveyed only in the summer periods of 2002 and 2003. 28-32. 2. middle of May and end of July 2003. Sipos Species 1 2 3 Erynnis tages Thymelicus silvestris Thymelicus lineola Thymelicus actaeon Ochlodes venatus Hesperia comma Pyrgus malvae Pyrgus carthami Pyrgus serratulae Pyrguy alveus Spialia orbifer/sertorius Carcharodus alceae Carcharodus flocciferus Heteropterus morpheus Carterocephalus palaemon Papilio machaon 1 1 0 0 1 0 1 1 0 0 1 0 1 0 1 1 1 1 0 0 1 0 1 1 0 0 0 0 1 0 1 1 1 1 1 0 1 1 1 0 0 0 0 0 0 0 1 1 . Gyertyánkuti-meadows. Vargyas-gorge.36 Table 1. 14-15. N Hungary. 16-17. 28-31: Slovenia. 33. 31. Junco-Molinietum near Apátistvánfalva. alcon-M. 29. 11-12. 32. Kuriszlán. Maculinea rebeli-sites in Aggtelek and Slovakian karst area: 1. Deschampsia facies. Grazed meadow on the Nanos-plateau. Succiso-Molinietum near Gödörháza. Rakaca-valley near Meszes. 27. Transylvania: Maculinea rebeli-sites. Maculinea arion-site on the plateau of Silica. teleius sites in the Zempléni-Mts: 11. M. Maculinea rebeli-site in Buda hilly region: Nagyszénás. nausithous sites in SW-Hungary. 32-33. 34. Aggtelek karst. 24. 18. Maculinea arion-site near Sóshartyán. Tall-forb karstic meadow on the Nanos-plateau. 30. Table 1. 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Species 1 2 3 Assemblages of butterflies and burnets in Maculinea habitats of Hungary Iphiclides podalirius Zerynthia polyxena Parnassius apollo Parnassius mnemosyne Colias alfacariensis Colias crocea Colias erate Colias hyale Gonepteryx rhamni Leptidea sinapis (reali) Leptidea morsei Pieris brassicae Pieris rapae Pieris napi Pontia edusa Anthocharis cardamines Neozephyrus quercus Thecla betulae Satyrium spini Satyrium w-album Satyrium ilicis Satyrium acaciae Callophrys rubi Lycaena dispar rutila Lycaena virgaureae Lycaena alciphron Lycaena tityrus Lycaena hippothoe Lycaena phlaeas Cupido argiades Cupido alcetas Cupido decoloratus Cupido minimus Cupido osiris 1 0 0 0 1 0 0 0 1 1 0 0 1 1 1 1 0 0 0 0 0 1 1 1 1 1 1 0 1 1 0 1 1 0 1 0 0 0 1 0 0 0 1 1 0 0 1 1 0 1 0 0 1 0 0 0 1 1 1 1 1 1 1 1 0 0 1 0 1 0 0 1 1 0 0 0 1 1 0 1 1 1 0 0 1 1 0 0 0 0 1 0 1 0 1 0 1 1 0 0 1 0 1 0 0 0 1 1 0 0 1 1 0 0 1 1 1 1 1 0 0 1 0 0 1 0 1 0 1 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 1 1 0 0 1 1 0 1 0 0 0 0 0 0 1 0 1 0 1 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 1 1 0 0 1 1 0 1 0 0 0 0 0 0 1 1 1 0 1 1 1 0 0 0 1 0 1 1 0 0 1 0 0 0 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 1 0 1 0 1 1 0 0 1 1 1 0 0 1 1 0 0 0 1 1 1 0 1 1 0 1 1 1 1 0 0 1 1 0 1 0 1 0 1 0 0 0 0 0 1 0 0 0 1 0 1 0 1 1 0 0 1 1 1 1 0 0 0 0 0 1 1 1 0 0 1 0 1 1 0 1 1 0 1 1 0 0 1 0 1 0 1 1 0 1 1 1 1 1 1 1 1 0 1 1 1 1 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 1 0 1 0 0 0 0 1 1 1 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 1 1 0 1 0 0 0 1 0 0 1 1 1 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1 0 1 1 1 1 0 0 1 0 0 0 0 0 1 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 0 1 1 0 1 0 1 0 0 0 0 1 1 1 0 1 0 1 1 0 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 1 0 1 0 0 1 0 0 0 1 1 1 0 1 0 1 0 0 0 1 0 1 0 0 0 1 0 0 0 1 1 0 1 1 1 0 0 1 0 1 0 1 1 1 0 0 0 0 0 1 1 0 1 1 0 1 0 0 0 1 1 1 0 1 1 0 1 1 1 1 1 1 0 1 0 1 0 1 0 0 0 1 0 1 1 1 0 0 0 1 0 0 0 1 1 1 1 1 1 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 1 1 0 1 0 0 0 0 0 0 0 1 0 0 1 0 1 1 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 1 1 0 1 0 0 0 0 0 0 1 1 1 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 1 1 0 1 0 0 0 1 0 0 0 1 1 0 1 1 0 0 0 0 0 0 1 0 0 0 1 0 0 1 1 1 0 0 1 1 0 1 1 0 0 0 0 0 1 1 1 0 1 1 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 0 0 0 0 0 1 0 1 0 1 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 1 1 0 1 0 1 0 0 0 0 0 0 0 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 0 1 0 0 1 0 0 1 0 1 0 1 0 0 0 0 0 1 0 1 0 1 0 1 1 0 0 1 0 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 37 . Continued. Table 1. 4 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 1 1 0 1 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 0 1 1 0 1 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 1 0 1 0 0 0 0 1 1 0 1 1 1 1 0 1 1 1 1 0 0 1 0 0 0 0 1 0 1 0 1 0 0 0 1 0 1 0 0 0 0 1 0 0 1 1 1 1 0 1 1 1 1 0 0 0 0 0 1 0 0 0 0 1 1 1 1 1 1 0 1 0 0 0 0 1 0 0 0 0 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 1 1 0 0 0 0 1 1 0 1 1 1 0 0 0 0 0 0 1 1 0 0 1 0 0 1 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1 1 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 1 0 0 1 0 0 1 1 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 1 0 1 0 1 1 1 1 0 0 0 0 0 1 0 0 1 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 1 1 0 1 0 0 0 0 1 1 0 0 1 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 1 1 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 0 0 1 0 1 0 1 0 1 1 0 0 0 0 0 1 0 1 0 0 0 0 1 1 0 0 0 0 0 1 1 0 1 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 1 0 0 0 1 1 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 38 Species 1 2 3 Zoltán Varga. László Peregovits & Julianna V. Sipos Celastrina argiolus Plebeius argus Plebeius idas Plebeius argyrognomon Plebeius sephirus Pseudophilotes schiffermuelleri Scolitantides orion Glaucopsyche alexis Maculinea rebeli Maculinea alcon Maculinea teleius Maculinea nausithous Maculinea arion arion Maculinea arion ligurica Aricia agestis Aricia artaxerxes issekutzi Polyommatus semiargus Polyommatus icarus Polyommatus thersites Polyommatus amandus Polyommatus dorylas Polyommatus coridon Polyommatus bellargus Polyommatus meleager Apatura iris Apatura ilia Neptis sappho Neptis rivularis Limenitis populi Limenitis camilla Limenitis reducta Nymphalis polychloros Nymphalis antiopa Aglais urticae 0 1 1 1 0 1 0 1 1 0 0 0 0 0 0 1 1 1 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 0 0 0 1 0 0 1 1 1 0 0 1 1 0 1 0 0 0 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 1 0 1 1 1 0 0 1 1 1 1 0 1 0 0 0 1 0 1 1 1 . Continued. Table 1. Continued. 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Species 1 2 3 Assemblages of butterflies and burnets in Maculinea habitats of Hungary Inachis io Polygonia c-album Cynthia cardui Vanessa atalanta Araschnia levana Melitaea phoebe Melitaea telona kovacsi Melitaea trivia Melitaea didyma Melitaea cinxia Melitaea diamina Mellicta athalia Mellicta britomartis Mellicta aurelia Euphydryas maturna Euphydryas aurinia Boloria dia Boloria selene Boloria euphrosyne Brenthis ino Brenthis daphne Brenthis hecate Issoria latonia Mesoacidalia aglaia Fabriciana adippe Fabriciana niobe Argynnis paphia Argynnis pandora Argynnis laodice Melanargia galathea Hipparchia semele Hipparchia fagi Hipparchia ferula Brintesia circe 1 0 1 1 1 0 1 1 1 1 0 1 1 1 0 0 1 0 0 1 0 1 1 1 1 1 1 0 0 1 0 1 0 1 1 0 1 1 1 0 1 1 1 1 0 1 1 1 0 0 1 0 0 1 1 1 1 1 1 1 1 0 0 1 0 1 0 1 1 1 1 1 0 0 0 1 1 1 0 1 1 1 0 0 1 1 1 1 0 0 1 1 1 1 1 0 0 1 0 1 0 1 1 1 1 1 1 0 0 0 1 1 0 1 1 1 0 0 1 0 0 0 0 0 1 1 1 0 1 0 0 1 0 1 0 1 1 1 1 0 0 0 0 0 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 1 0 0 0 0 1 0 1 0 1 1 0 0 1 1 0 1 1 1 0 0 1 1 0 1 0 0 1 1 1 1 1 0 0 1 0 0 0 0 1 1 1 1 1 0 1 1 1 1 0 1 1 1 0 0 1 0 1 1 1 1 1 1 1 0 1 1 1 1 0 1 0 1 0 1 0 1 0 0 1 1 1 1 0 1 1 1 0 0 1 0 0 1 0 1 1 1 1 1 1 0 0 1 1 1 0 1 1 1 1 1 0 1 0 1 1 1 0 1 0 1 0 0 1 0 0 0 0 1 1 0 0 0 1 0 0 1 1 1 0 1 1 1 1 1 1 1 0 0 0 1 1 1 0 0 1 0 1 1 0 1 1 0 1 0 0 0 1 0 0 1 0 0 0 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 0 1 1 0 1 1 0 0 1 1 0 1 0 1 1 0 0 0 0 1 1 0 0 1 0 0 0 1 1 1 1 1 1 0 0 1 1 0 1 1 0 0 1 1 1 1 0 1 1 0 1 0 0 1 1 1 0 1 1 0 0 0 0 0 1 0 0 1 0 1 1 0 0 1 0 1 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 1 1 0 1 1 1 0 0 1 1 0 1 0 1 0 0 0 1 0 0 1 0 1 1 1 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 1 1 1 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 0 0 0 1 1 1 0 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 1 0 1 1 0 1 1 1 0 1 1 1 0 1 1 1 1 1 1 1 0 1 1 0 0 1 0 1 0 0 1 0 1 0 1 0 1 1 1 0 1 0 0 1 0 1 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 1 0 0 1 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 0 1 1 0 0 1 0 0 1 0 0 0 1 1 0 0 1 1 0 0 0 1 1 1 1 0 0 0 1 1 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 0 0 0 1 1 1 0 1 0 0 0 1 0 0 1 0 0 1 0 0 0 0 1 1 1 0 1 1 0 0 0 1 1 1 0 0 0 1 1 1 1 0 1 0 0 1 1 0 1 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 1 0 1 0 1 1 1 0 1 0 1 0 0 0 0 0 1 1 1 0 1 1 1 1 0 0 1 0 1 1 1 1 1 1 0 0 1 0 1 1 1 0 1 1 1 0 0 1 0 0 0 1 1 0 1 1 0 1 0 0 1 0 0 1 1 1 1 1 1 0 1 0 0 1 1 0 1 0 1 0 0 1 0 0 0 1 0 0 1 1 1 1 0 0 1 0 1 0 1 0 1 1 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 1 0 1 1 1 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 0 1 1 0 0 1 0 1 0 0 0 0 0 39 . Sipos Arethusana arethusa Minois dryas Maniola jurtina Hyponephele lycaon Hyponephele lupina Aphantopus hyperanthus Erebia ligea Erebia euryale Erebia medusa Erebia aethiops Coenonympha pamphilus Coenonympha glycerion Coenonympha arcania Pararge aegeria Lasiommata megera Lasiommata maera Lopinga achine Adscita budensis Adscita globulariae Adscita chloros Procris statices Zygaena purpuralis/osterodensis Zygaena brizae Zygaena carniolica Zygaena loti Zygaena viciae Zygaena angelicae/transalpina Zygaena filipendulae Zygaena lonicerae Zygaena ephialtes pannonica 1 1 1 1 0 1 0 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 0 1 0 0 1 1 1 1 1 1 0 1 1 1 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 0 1 1 0 1 1 1 1 1 1 0 1 1 0 1 0 1 1 0 0 1 1 1 1 1 1 Number of species 87 85 86 72 55 69 99 85 68 74 56 73 45 22 56 25 45 66 73 35 42 54 26 45 72 22 25 54 57 62 49 6 10 29 . László Peregovits & Julianna V. Continued.Table 1. 4 0 1 1 0 0 1 0 0 0 0 1 1 1 1 0 1 0 0 1 0 1 1 0 1 1 1 1 1 1 0 0 0 1 0 0 1 0 0 0 0 1 1 1 1 0 1 0 0 1 0 1 1 0 0 1 1 1 1 1 0 0 0 1 1 0 1 0 0 1 0 1 1 1 1 0 1 0 0 1 1 1 1 0 0 1 1 1 1 1 0 1 1 1 0 0 1 0 0 1 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 0 1 1 0 1 1 1 1 0 1 1 0 0 1 0 0 0 0 1 0 1 1 1 1 0 0 1 0 0 1 0 0 1 0 0 1 0 1 0 1 1 0 0 1 0 0 0 0 1 1 1 1 1 1 0 0 1 0 1 1 0 1 1 0 0 1 0 1 0 1 1 1 0 1 0 0 1 1 1 1 1 1 0 1 0 0 0 0 1 1 0 0 1 1 0 1 1 0 0 1 1 1 0 1 0 0 1 1 1 1 1 1 0 1 0 0 1 0 1 1 0 0 1 1 1 1 1 0 0 1 1 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 1 0 1 1 0 1 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 1 0 1 1 0 0 0 0 1 0 1 1 1 1 0 0 0 0 0 1 0 1 1 0 1 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 1 1 1 0 0 1 1 1 1 0 1 1 0 0 1 0 1 1 1 1 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 0 1 0 0 1 0 1 1 0 1 0 1 0 0 0 0 0 1 0 0 1 1 0 1 0 0 0 1 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 0 1 1 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 1 1 0 1 0 0 1 1 1 1 1 1 0 1 0 0 1 0 1 1 0 1 1 0 0 1 1 1 1 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 1 0 0 0 0 1 1 1 0 1 0 0 0 0 1 0 0 1 1 0 1 1 0 0 0 0 1 1 0 1 1 0 0 0 1 1 1 1 0 1 0 0 0 0 1 1 0 1 1 0 1 1 0 0 0 1 1 1 0 1 1 0 0 1 1 1 1 1 0 1 0 0 0 0 1 1 0 1 1 0 1 1 0 X 1 0 1 0 0 0 1 0 0 1 0 0 1 0 1 1 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 40 Species 1 2 3 Zoltán Varga. the same subdivision. 11-13. N of Jósvafő. Tohonya-ridge Species Erynnis tages Thymelicus silvestris Ochlodes venatus Pyrgus malvae Pyrgus carthami Spialia orbifer Carcharodus flocciferus Carterocephalus palaemon Papilio machaon Iphiclides podalirius Pieris rapae Pieris napi Pontia edusa Anthocharis cardamines Colias alfacariensis Satyrium acaciae Callophrys rubi Lycaena dispar rutila Lycaena virgaureae Lycaena alciphron Lycaena phlaeas Everes argiades Everes decoloratus Plebeius argus Plebeius idas Plebeius argyrognomon Glaucopsyche alexis Maculinea alcon xerophila Cyaniris semiargus Polyommatus dorylas Polyommatus icarus Polyommatus coridon Polyommatus bellargus Polyommatus meleager Melitaea trivia Melitaea didyma Melitaea cinxia Melitaea telona kovacsi Mellicta athalia Mellicta britomartis Mellicta aurelia Clossiana dia Brenthis ino Brenthis hecate Issoria latonia Mesoacidalia aglaia Fabriciana adippe Fabriciana niobe Argynnis paphia Cynthia cardui 1/1 1/2 1/3 1/4 1/5 2/1 2/2 2/3 2/4 2/5 2 0 0 9 0 2 0 2 1 1 0 4 0 7 6 0 2 2 0 3 0 0 0 29 0 4 4 0 0 0 9 0 2 0 5 0 3 2 0 0 0 8 0 0 2 0 0 0 0 0 5 0 0 6 0 0 0 1 2 0 0 9 0 4 5 0 4 0 0 0 1 5 0 43 0 3 7 0 0 2 14 0 3 0 9 0 2 3 2 0 0 9 0 0 5 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 7 0 4 2 0 1 0 0 0 6 3 0 37 0 0 3 0 0 0 6 0 0 0 6 0 9 0 0 0 0 5 0 0 3 0 0 0 0 3 7 1 0 0 0 0 0 0 0 2 1 5 2 3 7 0 0 0 0 0 2 0 0 34 2 0 3 0 0 0 8 0 0 0 2 0 11 0 0 0 0 11 0 0 0 0 0 0 0 2 0 2 0 4 0 0 0 0 5 0 0 5 0 8 5 0 3 0 0 1 3 4 0 18 0 0 0 0 2 0 11 0 0 0 5 0 6 0 1 0 0 4 0 0 0 0 0 0 0 2 0 0 1 0 4 0 0 0 2 4 0 1 0 0 9 2 0 0 17 0 0 0 0 9 0 0 0 4 0 0 13 2 0 4 0 5 0 0 23 3 4 0 2 1 0 7 3 0 3 0 0 0 0 0 1 0 0 0 1 3 0 1 0 0 2 5 0 0 8 0 0 0 0 14 3 0 0 9 0 0 18 7 0 3 0 6 0 0 18 5 9 0 4 0 0 11 8 1 2 0 0 1 3 0 0 0 0 0 0 1 0 0 0 0 5 0 0 0 9 0 0 0 2 12 1 0 0 2 0 0 11 0 0 0 0 2 0 0 14 0 11 0 0 0 1 9 5 0 0 1 0 5 2 0 0 0 0 0 3 1 0 0 0 0 11 0 0 0 6 0 0 0 3 16 0 0 0 5 0 0 9 2 0 0 0 3 0 0 17 0 8 0 1 0 0 4 7 0 0 0 0 2 0 2 3 0 1 0 2 2 0 3 1 0 8 1 0 0 11 0 1 0 0 11 0 0 0 4 0 1 11 0 0 3 0 8 0 0 9 0 14 0 2 4 0 5 2 1 5 0 Ó1 14 3 0 5 0 2 0 3 8 3 2 30 2 26 25 0 10 2 0 4 12 12 0 161 2 7 17 0 2 2 48 0 5 0 27 0 31 5 3 0 0 37 0 0 10 0 0 0 0 7 Ó2 0 8 6 2 8 0 1 0 8 11 0 2 1 0 34 8 0 0 51 0 1 0 5 62 4 0 0 24 0 1 62 11 0 10 0 24 0 0 81 8 46 0 9 5 1 36 25 2 10 1 K1 III I 0 IV 0 I 0 II III II II V I V V 0 IV I 0 II IV III 0 V I II IV 0 I I V 0 II 0 V 0 V II II 0 0 V 0 0 III 0 0 0 0 III K2 0 III III I III 0 I 0 IV V 0 III I 0 V III 0 0 V 0 1 0 II V II 0 0 V 0 I V III 0 III 0 V 0 0 V II V 0 IV II I V V II III I . in daily repetitions 5 x 150 m. Butterfly assemblages. May and 1-3. as in the vegetation relevés (A1-15 – E 1-15) Aggtelek karst. Linear transsects.Assemblages of butterflies and burnets in Maculinea habitats of Hungary 41 Table 2. July 2002. Species Vanessa atalanta Inachis io Araschnia levana Coenonympha pamphilus Coenonympha glycerion Coenonympha arcania Lasiommata megera Lasiommata maera Lopinga achine Maniola jurtina Hyponephele lycaon Erebia medusa Aphantopus hyperanthus Melanargia galathea Hipparchia fagi Brintesia circe Arethusana arethusa Adscita budensis Adscita globulariae Adscita chloros Zygaena purpuralis Zygaena brizae Zygaena carniolica Zygaena loti Zygaena angelicae Zygaena filipendulae Zygaena lonicerae Zygaena ephialtes pannonica 1/1 1/2 1/3 1/4 1/5 2/1 2/2 2/3 2/4 2/5 0 2 1 2 0 0 0 0 0 0 0 6 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 27 1 1 2 3 0 0 0 0 0 0 0 11 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 27 0 1 0 1 0 0 0 0 0 0 0 8 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 19 0 0 0 4 0 0 1 0 0 0 0 10 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 20 0 2 5 0 0 0 0 0 0 0 0 16 0 0 0 0 0 2 0 0 0 1 0 0 0 0 0 0 22 0 0 0 0 1 0 0 0 2 17 0 0 2 27 0 7 0 0 0 1 7 0 11 8 2 6 0 2 34 0 1 0 0 0 2 0 0 0 26 0 0 1 32 0 12 0 0 1 0 4 0 18 4 3 11 2 0 32 0 0 0 2 4 0 0 0 0 18 0 0 0 17 0 6 2 0 0 0 0 0 4 3 0 5 0 0 26 0 0 0 3 5 0 0 0 0 11 0 0 0 22 0 9 0 0 0 0 0 0 1 7 2 4 0 0 26 0 1 0 0 2 0 0 1 0 31 2 0 8 19 2 14 0 0 3 2 2 0 12 11 4 8 1 0 43 Ó1 1 6 8 10 0 0 1 0 0 0 0 51 0 0 0 0 0 2 0 0 0 3 0 0 0 0 0 0 Ó2 0 2 0 5 12 2 0 1 2 103 2 0 3 117 2 48 2 0 4 3 13 0 46 33 11 34 3 2 K1 I IV III IV 0 0 I 0 0 0 0 V 0 0 0 0 0 I 0 0 0 II 0 0 0 0 0 0 K2 0 II 0 II IV I 0 I I V I 0 III V I V I 0 II II III 0 V V IV V II I . László Peregovits & Julianna V.42 Zoltán Varga. Sipos Table 2. Continued. NE Hungary. alcon-M. teleius-M. NE Hungary. 12) M. 5) M. 7) M. rebeli and M. sandstone. teleius-habitats of the Cserehát hills. 10) M. N Hungary.habitats of the Aggtelek karst. lower level (350-400 m). Species Aglais urticae Apatura ilia Apatura iris Argynnis pandora Argyronome laodice Aricia artaxerxes issekutzi Boloria euphrosyne Boloria selene Brenthis ino Chersotis fimbriola Cucullia gnaphalii Cucullia lucifuga Cucullia xeranthemi Cupido osiris Dichagyris musiva Euphydryas aurinia Gortyna borelii Hemaris fuciformis Hemaris tityus Inachis io Iphiclides podalirius Lemonia dumi Lemonia taraxaci Leptidea morsei Limenitis populi Lopinga achine Lycaena alciphron Lycaena dispar Lycaena hippothoe Maculinea alcon Maculinea arion arion Maculinea arion ligurica Maculinea nausithous Maculinea rebeli Maculinea teleius Melitaea ogygia kovacsi 1 1 r r 1 0 1 0 0 1 0 1 1 1 0 1 0 0 0 1 1 1 1 1 0 r 1 1 1 0 0 r 0 0 1 0 1 2 1 r r 0 r 1 1 1 1 1 0 1 0 0 1 0 0 0 0 1 1 0 1 1 r 1 1 1 r 0 1 0 0 r 0 0 3 1 0 r r r 1 r r 1 0 0 0 r 1 0 0 0 0 1 1 1 0 0 1 0 1 0 r 0 0 1 1 0 r 0 1 4 1 1 r 0 0 1 1 1 1 0 0 0 0 0 1 0 0 1 0 1 1 0 1 0 0 1 1 r 1 0 1 0 0 r 0 0 5 1 1 0 0 0 1 1 1 1 0 0 1 0 0 0 0 0 1 0 1 1 r 1 0 0 0 r r 1 0 r 0 0 1 0 0 6 1 r r 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 7 1 r 0 0 0 0 r 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 8 1 1 0 0 r r r 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 1 0 1 0 0 0 1 0 0 1 0 9 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 10 1 1 r 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 r 0 0 0 0 r 1 1 r 1 0 r 0 0 1 0 11 1 r 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 r 1 0 0 0 0 0 1 0 1 0 0 0 0 1 0 12 1 r 0 0 0 0 r 1 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 1 r 1 0 0 1 0 1 0 13 1 1 r 0 0 0 1 1 0 0 0 0 0 0 0 1 0 1 1 1 1 1 0 0 r 0 0 1 1 1 0 r 1 1 1 0 14 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 r 0 1 0 r EU 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + + 0 0 0 0 0 0 + 0 + 0 + 0 0 + + + 0 + 0 . Beregi-lowland. 4) M.Assemblages of butterflies and burnets in Maculinea habitats of Hungary Table 3. NE Hungary. arion) habitats of the acidic gravel ridge near Aggtelek. 14). N Hungary. N Hungary. teleius habitats on the Szatmár-lowland. 13) M. rebeli-habitats of the Bükk-plateau. nausithous habitats N from Balaton. N Hungary. EU: Species included into the HD Annex II. nausithous habitats in the Örség area. higher level (800-850 m). arion-habitats of Sóshartyán. S of Aggtelek karst. 11) M. W Hungary. higher level (500-550 m).habitats of the Teresztenye plateau. alcon-M. 2) M. teleius habitats on the mountain meadows of the ZemplénMts. teleius-(M. teleius-M. higher level (750-850 m). N Hungary. 9) M. in low individual number + in column EU: The species is included into the Annex II-IV of the Habitat Directive 43 Explanation of the columns: 1) Maculinea rebeli-habitats of the Aggtelek karst. 8) M. Protected butterfly and moth species at the Maculinea sites of Hungary 1: The species regularly occur at the site (in moth: light trap data) r: The species only occasionally occur. arion. arion. alcon-M. 6) M. Káli-depression. rebeli-habitats of the Slovakian karst. N Hungary. 3) M. Nagyszénás hill in the Buda Mts. arion-habitat on the Kaszonyi-hill. 44 Zoltán Varga. Species Papilio machaon Parnassius mnemosyne Photedes captiuncula Plebeius idas Plebeius sephirus Polyommatus amandus Polyommatus admetus Proserpinus proserpina Satyrium ilicis Satyrium w-album Vanessa atalanta Zerynthia polyxena 1 1 0 0 1 0 0 0 1 1 0 1 0 2 1 1 1 1 0 0 0 0 0 0 1 0 3 1 0 0 1 0 0 1 r 1 0 1 r 4 1 1 1 1 0 0 0 0 0 0 1 0 5 1 1 0 0 0 0 0 0 0 1 1 0 6 1 0 0 0 1 0 0 0 1 0 1 0 7 1 0 0 0 0 1 0 1 0 0 1 1 8 1 0 0 1 0 0 0 0 0 0 1 0 9 1 1 0 0 0 0 0 0 0 0 1 0 10 1 1 0 1 0 0 0 0 0 1 1 0 11 1 0 0 0 0 0 0 1 0 0 1 0 12 1 0 0 0 0 0 0 1 0 0 1 0 13 1 1 0 1 0 0 0 0 0 0 1 0 14 1 1 0 0 0 0 0 0 1 0 1 0 EU 0 0 0 0 0 0 0 + 0 0 0 + . Continued. Sipos Table 3. László Peregovits & Julianna V. H-4010 Debrecen. mowing at the end of the vegetation period. The propagula of these species were obviously present in the seed bank of the soil.. arion habitats in NE Hungary Several Maculinea rebeli sites of the Aggtelek karst belong to the Pannonian semi-dry swards of the alliance Cirsio pannonicae-Brachypodion pinnati. as well. In addition also some „pioneer” species appeared on the bare patches as Adonis aestivalis. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group. However. where the change of the vegetation structure is slow. Faculty of Sciences. Bükk Mts. Kühnrelegation (Eds) 2005 45 Studies on and cenological and Conservation of Butterflies in Europe Vol. rebeli proved to be more xerothermic than the neighbouring.g. E. P.A. abandonment. .B. The sites of the Bükk Mts. Aggteleki-karszt: Kuriszlán. but they are decreasing or fluctuating in some sites of the Bükk Mts. 3.: Lófö-meadow) is unclear as a consequence of secondary succession. rebeli population. Cirsium arvense. Dominant weeds of this site are: Agrimonia eupatoria. The life-form spectra of the vegetation clearly reflect the different types of management vs. The active management of this strongly degraded site started in 2001 (cutting of Rosa scrubs.O. Bifora radians. & J.© PENSOFT Publishers Maculinea habitats: Sofia – Moscow J. Consolida regalis. also in protected species (Appendix: Table 1) in the Aggtelek karst and in the Bükk Mts. Department of Evolutionary Zoology and Human Biology. pp.hu Dry and semi-dry swards: Maculinea rebeli and M.B. one of the latter sites situated on the plateau between the Tohonya and Lófej valleys was heavily overgrazed and –trampled by horses in the mid-90’s. 1). M. partly harrowed) which resulted in the increase of the M. The floristic composition of the M.unideb. Hungary Contact: zvarga@tigris. Berteroa incana. 3. rebeli sites is generally rich in species. where the vegetational succession seems to be faster and the Gentiana cruciata plants are often overgrown by tall grasses. Thomas diversity of vegetation. 45-50 Maculinea habitats: diversity of vegetation. Stenactis annua.2 1 DUFS . composition and cenological relegation Julianna Varga-Sipos2 & Zoltán Varga1.O. The phytocenological relegation of some abandoned hayfields of these mountains (e.University of Debrecen. compositionthe EcologySettele. H-4010 Debrecen. since this area was a traditional agricultural area until the 1970’s. The Ellenberg-Zólyomi ecological characteristics (TWRN) show a typical gradient. Seemingly similar sites with or without the occurrence of Maculinea rebeli populations were compared in the Aggtelek karst area . rebeli populations of some abandoned hayfields are strong (Aggtelek karst: Kuriszlán). mostly represent various transitions between semi-dry swards and mesophilous mountaine meadows (AnthyllidoFestucetum rubrae). where the unsuitable sites for M. Papaver rhoeas. strongly populated sites (Fig. P. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. certain habitats in this area are short-grass swards. The swards of the Aggtelek karst are only studied phytocenologically. as well. Sipos & Zoltán Varga It is difficult to characterise the vegetation of the Maculinea arion-sites. the dark green dots of the humid mountain hayfields (alcon-teleius habitats) of the same region and the dark violet dots are the humid habitats of W Hungary (alcon. teleius.g. arion (early flying population). They are either edaphic grasslands (e. sandstone rupicolous swards in Nógrád county. The M. The dark blue + marks symbolise the Deschampsio-Molinieta (teleius habitats) of NE Hungary. humid-subatlantic trend. . nausithous habitats). arion habitats of the Pannonian area. too. on dolomitic ridges in the Aggtelek karst area. 1. Thus the gradient clearly shows a xerotheric-continental vs. rebeli and more regularly by M. Calculated according to the Ellenberg-Zólyomi ecological characteristica of the plant species occurring in the vegetation relevés.) or abandoned former pastures in hilly part of northern Hungary. the light yellowish-green + marks and the light yellowish-green dots symbolise the semi-dry grasslands populated by M. Nevertheless. rebeli.46 Julianna V. Fig. etc. an ecological gradient of the Maculinea sites was constructed. The general vegetation characteristics of the central and western European sites (short grass nutrient-poor swards with cushions of Thymus pulegioides) are different from most of the M. arion habitats of the calcareous part of the Aggtelek karst area represent the alliance Cirsio pannonicae-Brachypodion pinnati. This trend is also clearly reflected by the butterfly and grasshopper assemblages. The most xerothermic sites (red dots) are short-grass calcareous steppic grasslands only scarcely populated by M. The orange + marks. The floristically richest stands of Molinieta extend in the pre-Alpine colline area (Örség) on one hand.Maculinea habitats: diversity of vegetation. June). forming a metapopulation system due to the high flying capacity of M. Other M. M. These sites are abandoned pastures under extreme edaphic conditions (acidic gravel). but they are often dominated by Alopecurus pratensis. arion habitats are either xerothermic forest „skirts” or even clear-cut areas with abundant Origanum vulgare. arion populations are not abundant. Humid and marshy meadows: Maculinea alcon. The other. nausithous habitats. 1). The flying period of the imagos is also synchronised here with the main flowering period of the initial foodplant (second half of July. teleius and M.. These sites are interspersed with densely populated habitats of M. but relatively stable. which is a widely distributed initial food-plant of M. they are only small fragmented patches within a forested „matrix”. more diverse type of the swards (Hypochoerio-Brachypodietum) occur at lower altitudes (Teresztenye plateau. Their external appeareance is similar to the AdriatoMediterranean subspecies: M. Populations living at such sites are often subjected to extreme fluctuations. Junco-Molinietum juncetosum and Junco-Molinietum nardetosum (Table 4). Betula pendula and Populus tremula. M. arion on Thymus marschallianus and Th. arion. However. These successionally transitional sites are highly unstable and it is hard to relegate them to any „classical” cenotaxa. Table 1). The external appeareance of the imagoes also corresponds here to M. Teucrium montanum and Thymus spp. composition and cenological relegation 47 and they mostly belong to two typical associations. arion also forms a relatively stable metapopulation system here over the more or less connected grassland patches. Thus. The humid or marshy grasslands of the Pannonian region often become dry during the summer season. arion in this region (as opposed to other Maculinea spp. kosteleckianus) at both types of sites. these M. They mostly belong to the alliances Molinion coeruleae and Deschampsion caespitosae. arion in some other landscapes). These are mainly characterised by the co-occurrence of Maculinea teleius and . In W Hungary (and also in the neighbouring part of Slovenia) three types of Molinia meadows are widely distributed: Junco-Molinietum typicum. The flying period of the imagoes seems to be synchronised with the flowering period of these initial food-plants (2 half of May. They are often rich in protected plant species. pannonicus (= Th. and in the northern sub-Carpathian part of the Zemplén Mts on the other. 1904). 1758). The short-grass Poo badensis-Caricetum montanae of the dolomitic ridges at medium altitudes (500-550 m) has a patchy vegetation structure with abundant cusheon-forming chamaephyta as Alyssum montanum. Their appeareance corresponds to the nominotypical Maculinea a. Thus. arion sites have been found recently in the Aggtelek karst region with heath-like vegetation (mosaic of Calluno-Genistetum germanicae and Nardo-Callunetum) with abundant cushions of Thymus pulegioides. Festuca pratensis and Carex spp. Globularia elongata. The flying period of the imagoes also co-incides here with the main flowering period of the initial food-plant (from mid-July to early August). about 400 m). with the beginning of re-forestation by pioneer scrubs and trees such as Juniperus communis. Both types of site are rich in protected plant species (see: Appendix. teleius (see below). especially in the eastern part of the region. thus the management of these sites will be rather important during the next years. arion ligurica (Wagner. Further M. we could observe a gradient in the ecological characteristics of these grasslands from southwest to northeast (Fig. arion. and to M. These sites are stabilised by the grazing of the red deer and have shown hardly any changes during the last 20-30 years. the alternative initial food-plant of M. as well. early August). We observed the oviposition of M. arion (Linnaeus. too. 1904). Thus. arion ligurica (Wagner. Trifolium pratense. Several ovipositions were observed and photographed on this locally important food plant. rebeli populations on the slopes of karstic dolinas with strong stands of G. alcon. often fenced and only periodically used. humid plateau of the Trnovski Gozd M. alcon. where the initial food-plant Gentiana cruciata grows in abundance. rebeli often seems to be restricted. Leontodon hispidus. teleius is still much more widely distributed. Neighbouring fertilised areas with intensified use were floristically extremely poor and unpopulated by M. nausithous and only rarely of M. The picture is rather varied. M. P. in the Hochschwab area (Styrian Alps) M. C jacea. Aster bellidiastrum. Kamnik Alps). Ranunculus acris. as they are either abandoned or mowed in summer by heavy machines. Vegetation of Maculinea habitats in the Alps and Slovenian karst We could also compare the vegetation of the sites populated by M. we could find some strong M. Festuca rubra. to the mowed edges of roads. In some areas weak M. together with its initial food-plant G. are characterised by co-occurrences of M. Cirsio cani-Festucetum pratensis and Caricetum caespitosae). teleius only. Unfortunately. trivialis. on the grazed meadows of the same area M. The wet Molinia grasslands of N-NE Hungary are different in the Aggtelek karst (Junco-Molinietum deschampsietosum and juncetosum). rebeli in Slovenia and at the „classical” site Hochschwab area in Austria. rebeli. cruciata for oviposition. rebeli seems to be restricted to some few. teleius and M. Such areas are floristically diverse with the dominant species: Anthoxanthum odoratum. in the Szatmári-lowland (a poor variation of Junco-Molinietum and Alopecuretum) and in the Zempléni-Mts. (a very diverse mountaine Molinietum with numerous mountaine hayfield species). although M. In Austria. cruciata can be extremely high. Crepis aurea. although the mowing period at these sites (early July) is unsuitable for the survival of freshly emerging larvae. they are often in unfavourable situations.g. in the Cserehát hilly region (Junco-Molinietum. Despite this. However. rebeli populations may occur also in heavily overgrazed sites (e. Cynosurus cristatus. Sipos & Zoltán Varga M. Poa alpina. On the high. rather fragmented „classical” sub-alpine pastures. Trisetum flavescens. rebeli densely populates the nature-like tall-forb meadows of the high karstic plateau of the Nanos Mts where Gentiana lutea grows abundantly. whereas the habitats of the Szatmári-lowland and Zempléni-Mts. Centaurea pseudophrygia. cruciata and floristically highly diverse vegetation showing some similarity to the sites of the Aggtelek karst. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). .48 Julianna V. The habitats in the Aggtelek and Cserehát region are characterised by the frequent occurrence of M. cruciata. rebeli uses G. In such „low input” sites the abundance of butterflies and also of the eggs on G. rebeli-habitats of the Bükk-plateau. N Hungary. NE Hungary. arion) habitats of the acidic gravel ridge near Aggtelek. N Hungary. arion-habitat on the Kaszonyi-hill. teleius-(M. teleius habitats on the Szatmár-lowland. 8) M. sandstone. Beregi-lowland. 11) M. W Hungary.habitats of the Aggtelek karst. alcon-M. teleius-habitats of the Cserehát hills. arion. teleius-M. brymii Anacamptis pyramidalis Anemone sylvestris Aster amellus Astragalus exscapus Calamagrostis varia Carduus collinus Carex appropinquata Carex davalliana Carex hartmannii Carex lasiocarpa Carex paniculata Carex umbrosa Carlina acaulis Centaurea sadleriana Centaurea triumfettii Chamaecytisus albus Dactylorhiza fuchsii Dactylorhiza incarnata Dactylorhiza maculata Dactylorhiza majalis Dactylorhiza sambucina Dianthus deltoides Dianthus superbus Dictamnus albus Dracocephalum austriacum Echium maculatum Eriophorum latifolium Erysimum odoratum Gentiana asclepiadea Gentiana cruciata 1 0 0 1 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 2 0 0 1 0 0 0 1 0 1 1 0 0 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 1 0 1 0 1 3 0 0 1 0 0 0 1 0 1 1 0 0 1 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 1 1 0 1 0 1 4 0 0 1 0 0 0 1 1 1 1 0 0 1 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 1 5 0 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 6 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 7 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 8 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 10 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 11 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 1 0 0 0 1 0 0 0 12 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 13 1 0 0 0 0 1 0 0 0 0 0 1 0 1 1 0 1 1 0 0 0 0 0 0 1 0 1 1 0 1 0 0 0 1 0 0 0 14 1 0 0 0 1 0 0 0 0 0 0 1 0 1 0 1 1 1 1 0 0 0 0 0 1 1 1 1 0 1 0 0 0 1 0 1 0 EU + + + . 7) M. rebeli and M. higher level (500-550 m). lower level (350-400 m). N Hungary. N Hungary. 2) Weakly populated Maculinea habitats on the wineyard hills of the Aggtelek karst (300-400 m). composition and cenological relegation Table 1. 3) M. higher level (800-850 m). N Hungary. NE Hungary. 4) M. Káli-depression. Protected plant species on the Maculinea habitats in Hungary 49 Explanation of the columns: 1) Maculinea rebeli-habitats of the Aggtelek karst. 12) M. arion-habitats of Sóshartyán. N Hungary. teleius habitats on the mountain meadows of the Zemplén-Mts. alcon-M. 5) M. rebeli-habitats of the Slovakian karst. alcon-M. arion-habitat of the Teresztenye-plateau. included into the HD Annex II. 15. Species Achillea ptarmica Adenophora liliifolia Adonis vernalis Alchemilla hybrida Alchemilla xanthochlora Allium suaveolens Alyssum montanum ssp. 13) M. NE Hungary.Maculinea habitats: diversity of vegetation. nausithous habitats N from Balaton. 9) M. column: Species of Community Interests. S part of the Aggtelek karst (350-400 m). 14) M. nausithous habitats in the Örség area. higher level (750-850 m). 6) M. teleius-M. 10) M. 50 Julianna V. Continued. Species Gentiana pneumonanthe Gentianella austriaca Gentianella livonica Gentianopsis ciliata Gladiolus imbricatus Gymnadenia conopsea Hemerocallis lilio-asphodelus Iris pumila Iris sibirica Iris variegata Jurinea mollis Lathyrus pannonicus Linum flavum Linum hirsutum Linum tenuifolium Orchis morio Orchis pallens Orchis purpurea Orchis tridentata Orchis ustulata Ornithogalum pyramidale Parnassia palustris Peucedanum officinale Polygala major Polygonum bistorta Primula elatior Prunella grandiflora Pulsatilla grandis/slavica Pulsatilla patens Sorbus aria Sorbus domestica Stipa joannis Stipa pulcherrima Stipa tirsa Thalictrum aquilegiifolium Trollius europaeus Veratrum album Veronica paniculata Number of protected species 1 0 0 0 0 0 1 0 0 0 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 19 2 0 0 0 1 0 1 0 1 0 0 1 1 1 1 1 1 0 1 1 1 1 0 0 1 0 0 1 1 0 0 1 1 0 1 0 0 0 0 29 3 0 0 0 0 0 1 0 1 0 1 1 1 1 1 1 0 0 0 1 1 1 0 0 1 0 0 1 1 0 1 0 1 0 1 0 0 0 0 26 4 0 0 1 1 0 1 0 0 0 0 1 1 1 0 1 0 0 0 0 1 0 0 0 1 0 0 1 1 0 1 0 0 0 0 0 0 0 0 23 5 0 0 0 0 1 1 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 1 0 1 0 1 1 1 0 0 0 0 0 0 0 0 16 6 1 0 1 1 0 1 0 0 0 0 1 1 0 0 0 1 0 1 0 0 0 0 0 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 20 7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 3 8 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 2 9 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 10 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 7 11 1 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 15 12 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 13 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 14 14 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 23 EU + + . Sipos & Zoltán Varga Table 1. 6% 45. where Maculinea population structure were studied. Ewa Sliwinska. with the proportion of species being as follows: Myrmica scabrinodis 69%.5% 43. Institute of Environmental Sciences.© PENSOFT Publishers The distribution Sofia – Moscow J. and Formica spp. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. rubra M. scabrinodis M. rubra M. Piotr Skórka. are presented in Tables 1 (site1) and 2 (site2). p. Settele. rubra is rare on site 2 may serve as an explanation for low numbers of Maculinea nausithous there.5%. . Piotr Nowicki & Michal Woyciechowski UJAG .Jagiellonian University. We found 704 Myrmica nests. rubra 18. Kühninof Butterflies(Eds) 2005 teleius the Ecology nausithous & J. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). sabuleti 0. M. M. E. The results of the Myrmica species composition on two sites. Species of Myrmica M. Thomas in Europe 51 and Conservation Poland Vol.uk The aims of the study were: 1) to investigate the abundance and distribution of ant species in the Kraków region and 2) to detect which Myrmica species co-occur with Maculinea butterflies in this region. The studies were carried out from May to September 2002. Lasius flavus.6%.5% 79% 9. even though the proportion may vary slightly.75% 12. Ant nests were searched on 7 sites selected for surveying the butterfly community. and ecology of MaculineaStudies on and M. M. 51 The ant communities on meadows in the Kraków region Magdalena Witek. In addition.9%. Species of Myrmica M. on two sites where Maculinea population structure was intensively studied.A.3% Sugar cubes 43. ruginodis 11. scabrinodis M. ruginodis Table 2. 30-387 Kraków. Other ant species found included Lasius niger.7% 7.1% 1. Gronostajowa 7. Furthermore.4% Sugar cubes 11. the fact that M.9% 65.5% The above comparison proves that the order of dominance of a particular Myrmica species remains the same in both methods. Poland Contact:
[email protected]% Nests 26. Table 1. ruginodis Nests 53.co. we also used a suger cube method to determine ant species composition. 52 Jarosław Buszko. Stankiewicz This page intentionally left blank . Marcin Sielezniew & Anna M. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Functional and trophic relations in Maculinea systems .2.Chaetocnema conducta (Motschulsky) and its Kindred Species in the Afrotropical Region 53 Section 3. Martin Musche & Josef Settele This page intentionally left blank .54 Christian Anton. Centre for Environmental Research Leipzig-Halle. M. We analysed the impact of host density. Theodor-Lieser-Str. 55-56 Parasitism of the predatory Maculinea nausithous by the parasitoid Neotypus melanocephalus Christian Anton. P < 0.A. High densities of caterpillars on the food plant may lead to the overexploitation of ant resources. In addition. nausithous caterpillars feed inside the flower heads of Sanguisorba officinalis (Thomas et al. GLM. On the population level.de Parasitoid wasps are major enemies of the early stages of most butterflies.anton@ufz. 06120 Halle. we carried out behavioural studies in the field to analyse the exploitation of flower heads hosting M. c2 = 4. high host densities may not always be advantageous to foraging N. 1998). and habitat-related parameters on parasitism by N. n = 26. In this study we analyse the factors influencing the parasitism of the predacious lycaenid Maculinea nausithous. nausithous caterpillars and the size of the habitat patch. Kühn & J. E. Settele. pp. the density of caterpillars did not influence the parasitism rate.05 . Before being integrated into the nest of Myrmica ants. Department of Community Ecology. While mutualistic lycaenids are predominantly tended by ants. Caterpillars of the family Lycaenidae are often associated with ants which gains them protection against enemies (Pierce et al. nausithous populations. caterpillars are parasitised by the wasp Neotypus melanocephalus. host population size. caterpillars of parasitic blues do not have ant guards. melanocephalus. Germany Contact: christian. 1 Relationship between the proportion of parasitised M. nausithous caterpillars. Fig. 4. however. 2002).1. melanocephalus in 26 M.© PENSOFT Publishers Chaetocnema conducta Sofia – Moscow J. Thomas (Motschulsky) and its Kindred Speciesthe EcologyAfrotropicalof Butterflies(Eds) 2005 55 Studies on in the and Conservation Region in Europe Vol. Despite their concealed feeding. Thus. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Martin Musche & Josef Settele UFZ . N. J. Oecologia.E. Thomas. & Hochberg. . Braby. D. P < 0. This study shows that N. Maculinea nausithous occurs at higher density in small patches (Anton et al. Temporal host refuges of M.3. 154. 261-290. M. melanocephalus is a large solitary parasitoid.P. 23-36.9 99. The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat. officinalis Behaviour of N. Mathew. REFERENCES Cronin.6. Doak.05 Non-exploited 19. Chapman and Hall. Annual Review of Ecology and Systematics. Surprisingly.002 small-sized habitat patches displayed higher parasitism than large patches (Fig. unpublished results).. D. 2000. & Strong.3 127.3 ± 1. Martin Musche & Josef Settele Table 1 Behaviour of N. Insect population dynamics in theory and practice (ed.G. Foraging N. Elmes. Clarke.7 ± 0.5 2. many flower heads were not probed by N.4 ± 2. 2002.. However.3±0. 18.. The ecology and evolution of ant association in the Lycaenidae (Lepidoptera). R. t = 3.F. A.002).8 flower heads W = 43. P. pp. melanocephalus probed occupied flower heads only. This might be due to small caterpillars feeding inside the flower cups that N. R. D. Non-exploited flower heads were left very soon whereas oviposition prolonged foraging. n = 26. Dispersal-dependent oviposition and the aggregation of parasitism.5 Significance P < 0. J..1 ± 5. melanocephalus although they hosted caterpillars. 1999.0 P < 0. melanocephalus allocates foraging time to hosts in an adaptive way. E. American Naturalist.6 s .7 ± 0.0 vs. Lohman. ÷2 = 4.6 ± 30.05). Heath. melanocephalus might not be able to oviposit through. McLean)..2 W = 5. A. J. 47.51 P = 0. 556-56 Pierce. B. 733-771. and spent more time on flower heads that were successfully exploited (127.8 ± 26. Doak (2000) argued that increasing parasitism in small and isolated patches might be correlated with travel time and Cronin and Strong (1999) showed that parasitism was dispersal-dependent. M. but considering that N. melanocehalus. but would need to be tested in future. G.9 ± 2. 1).1. Dempster and I.6 P = 0. 122.001 0.W. This might lead to an increased reproduction of parasitoids.6 s ± 30. M.05 0..A.T. Symposia of the Royal Entomological Society 19. P = 0.. F. however.8 2. London. & Travassos. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Rand. At present. too. melanocephalus ± 1SE [s] Searching Exploited flower heads Probing Oviposition Grooming Resting 13. melanocephalus on exploited and non-exploited flower heads of S.4 ± 2. by J. Maculinea nausithous populations in small-sized habitat patches experienced higher parasitism than those in larger patches (GLM. T.2 26.3 ± 22. (1998) Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). J.0 12 45 Time spent on flower ± 1SE [s] n 75.56 Christian Anton.6 W = 206 t = .3 ± 6. nausithous caterpillars might contribute to heterogeneity in parasitism.7 W = 126 P < 0.5 18. we lack information on the dispersal behaviour of N. we suggest that it is a strong disperser.. 1994) or by immigration behaviour (Hill et al. P = 0. P = 0. Department of Zoology.33 = 12.© PENSOFT Publishers factors Which Sofia – Moscow J. but not with the density of the host plant S. Germany 2 Mendel University of Agriculture and Forestry Brno.7. rubra in edge habitats (Dauber & Wolters. 613-01 Brno. 1993. 1997. Once inside the ant nest. officinalis (F1.52). rubra. Also. P = 0. Department of Community Ecology. Maculinea species employ two kinds of feeding strategies. The adult density of cuckoo-feeding species positively correlates with the density of the foodplant (Elmes et al. This field study indicates that the density of M. Egg densities were positively correlated with both plants (F1. This study aimed to answer the question whether the density and distribution of the food-plant Sanguisorba officinalis or the density of the host-ant Myrmica rubra limits the density of the predatory M. Prior to the adoption by Myrmica worker ants Maculinea caterpillars feed on a specific food-plant. Martin Musche1. In this study. Vladimir Hula2 & Josef Settele1 UFZ .43. We used the density of eggs.33 = 5. The term community module refers to small groups of species that interact either directly or indirectly. Kühn & of Butterflies(Eds) 2005 density of the and Conservation J. Ovaskainen. 57-59 Which factors determine the population density of the predatory butterfly Maculinea nausithous? Christian Anton1. It is unknown whether this effect was due to larger colony sizes of M.001) and host ants (F1. Zemedelska 1. 1998). 2004). the density of caterpillars responded to host plant density (F1.001).7. Thomas et al. but benefits from habitat edges. . Czech Republic Contact: christian. lower caterpillar mortality through parasitoids (Roland. Predatory Maculinea species experience scramble competition inside ant nests. Theodor-Lieser-Str.. P < 0. Thomas in Europe predatory butterfly 57 Vol. Small-sized patches displayed higher densities of butterflies than large ones. 1996). E. smaller habitat patches were shown to have higher densities of adult M. 06120 Halle. pp.Centre for Environmental Research Leipzig-Halle. rubra (F1.de 1 Butterflies of the genus Maculinea and their food resources constitute a community module (Holt. P < 0.A.anton@ufz. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.02). Agronomical Faculty.33 = 6. higher recolonization probability of M. rubra in case of overexploitation.5. 2004). 1996. nausithous.33 = 11. 4. determine the population Studies on the EcologySettele.33 = 0.02). caterpillars and adult butterflies separately as response variables in multiple regression models. nausithous.. 2002. officinalis and the low efficiency of feeding might lead to a close relationship between butterflies and ants in the predatory M. Kruess & Tscharntke. but see Wahlberg et al. In previous studies eggs were counted to calculate the density of adult butterflies. nausithous butterflies is limited by the density of M.0. nausithous.5.33 = 12. whereas cuckoo-feeding Maculinea are contest competitors (Thomas and Elmes 1998). The density of butterflies positively correlated with the density of M. It is suggested that the density of predatory Maculinea responds more strongly to the density of host-ants than to the density of food-plants. P < 0.002) and host ant density (F1. Higher density of the food-plant S. 52). 65.33 = 0.02). f) relationship between adults and ants (F1. J. a parasitic butterfly of red ant colonies. Thomas. Clarke. 17. G. Biodiversity and Conservation. 901-915. C.K.02). & Lewis. Hill.58 Christian Anton et al. Acta Oecologia. P = 0.002). d) relationship between caterpillars and ants (F1. (1996) Effects of habitat patch size and isolation on dispersal by Hesperis comma butterflies: implications for metapopulation structure. 13.33 = 5. R. Fig. M. P = 0.. respectively.001). b) relationship between eggs and ants (F1. nausithous stages and the density of food-plants (first column) and host-ants (second column).33 = 6. Thomas.33 = 11..T. 725-735.5.T. P < 0.D. 61-80. e) relationship between adults and plants (F1. Elmes. P = 0.E. P < 0. & Hochberg.A. Journal of Animal Ecology.33 = 12. V.33 = 12.. & Wolters. (2004) Edge effects on ant community structure and species richness in an agricultural landscaped. (1996) Empirical tests of specific predictions made from a spatial model of the population dynamics of Maculinea rebeli. J. 1. REFERENCES Dauber.0. Relationship between the density of M.5. O.001).7.W.43.7. . a) relationship between eggs and plants (F1. P < 0. c) relationship between caterpillars and plants (F1. J. Science. 93. 25-30.F. 264. Elmes. Clarke. . Roland. M. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). pp. T.P.. Insect population dynamics in theory and practice (ed.. & Tscharntke. & Hochberg. Oxford. by J.E. Blackwell Science. Symposia of the Royal Entomological Society 19. London. pp.W. R. Dempster and I. species loss. Kruess. (1997) Community modules. 333-350. and biological control.D.A. J.G.T. Multitrophic interactions in terrestrial systems (ed. Chapman and Hall. (1994) Habitat fragmentation. Thomas.C. Brown). (1998) Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). Oecologia. McLean). Gange and V. 261-290. J. R.K. by A.Which factors determine the population density of the predatory butterfly 59 Holt. G. 1581-1584. A. (1993) Large-scale forest fragmentation increases the duration of a tent caterpillar outbreak. 60 Christian Anton et al. This page intentionally left blank . 88%) than that of intact sprouts (70.unideb. Kühn & of Butterflies(Eds) 2005 Maculinea Thomas in Europe alcon 61 Studies on the Ecology and Conservation Vol. Settele. Lepidoptera: Lycaenidae) in the Aggtelek National Park Ervin Árnyas1.68) in the years before reconstruction. Judit Bereczki1. pp. P. of which an area of 9500 m2 was designated as our sampling area and divided into squares of 10x10m. 1993 and 1998). Food plants had been mapped and eggs counted in the habitat even before the area was reconstructed. 3.hu We studied the egg-laying preferences of the xerophilous ecotype of Maculinea alcon (= Maculinea rebeli) in the Tohonya valley within the Aggtelek karst region (N Hungary). 1). while the ratio of intact fertile sprouts (48. 1).© PENSOFT Publishers Egg-laying Sofia – Moscow J. P. 50 and 40 randomly selected squares in 2003.O. sterile and bitted sprouts. In the years studied the percentage of grazed sprouts was much lower (7.2 1 DUFS .66) in the number of intact fertile sprouts per square in the years following habitat reconstruction. while eggs were counted only on 20. Because the number of the sample squares was different throughout the study.3.94%) was higher. 61-64 Egg-laying preferences of the xerophilous ecotype of Maculinea alcon (= M.University of Debrecen. As the survey was done with essentially the same methods as in the earlier years (1992. we compared the average counts per square (Fig. which was probably due to the overpopulation of roe deer in the 90s. Gentiana cruciata colonies were mapped in the whole sampling area. Department of Evolutionary Zoology and Human Biology. it was possible to study the effect of the habitat reconstruction on the number of the eggs. there was a significant increase (14. In the years before the habitat reconstruction. Faculty of Sciences.8) was rather low. Hungary Contacts: arnyaser@delfin. The population of Maculinea alcon is distributed over an area of about 3 ha of the above territory. J. H-4010 Debrecen. Before the . the ratio of grazed sprouts (35. the area has been managed by regular mowing and selective cutting of the scrub at the end of the vegetation period. After the treatment (mowing and cutting of scrubs). However. Since the Gentiana sprouts were mapped and also the eggs were counted in every year.O.B. 2). Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group. since the number of roe deer was controlled at the beginning of 2000. rebeli.97%). The number of intact fertile sprouts per square was very low (2.A. and it had been decreasing before the management started.B. Andrea Tóth1. Fig. Since 2002. Before the management the average number of eggs per quadrate (6. the results could be compared. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. 3. The two periods showed significant differences in the percentage of grazed and fertile sprouts. The other change was found in the distribution of eggs on the intact. this number significantly increased (160. H-4010 Debrecen.14%) was lower than in the years after the reconstruction (Fig. 2004 and 2005 respectively. Katalian Pecsenye1 & Zoltán Varga1. preferences of the xerophilous ecotype of E. The average counts of intact fertile sprouts (dotted columns) and eggs (hatched columns) per square in the years of the study. the number of intact fertile sprouts in the squares and the total egg count in each year. The dashed line indicates the beginning of the management of the area. After 2002. We could hardly find any eggs on sterile. grazed and green-fly (aphid) infected sprouts. dotted bar: intact fertile sprouts. The table. as the proportion of intact sprouts was essentially lower. The start of the habitat reconstruction is marked by the dashed line. 2.2%). Striped bar: intact sterile sprouts.9%). thus females laid their eggs almost exclusively on them (98. Fig. We have also examined the relationship between the number of the eggs and the number of sprouts (ramets) belonging to a sprout-colony. The greatest number of the eggs was found on . shows the number of squares.62 Ervin Árnyas et al. however. treatment the females had also laid some eggs on the sterile and bitted sprouts (15. 1. the number of intact and fertile sprouts significantly increased. The distribution of sprouts counted in the years of sampling. blank bar: grazed sprouts. Fig. Dotted bars: the number of sprouts. Adax: adaxial surface of the leaves. verticil 25 3 12 3 . verticil 77 17 62 6 4. As the number of ramets per sprout-colony increased the chance of egg-laying also increased. the fourth one 0. the third one 2. Thus they are easily detected by the females. We suppose that colonies with 7-8 ramets are optimal. Comparing the proportion of eggs on different parts of the host plant (Table 1). often sterile. The ramets of the colonies with over-abundant sprouts are generally lower. since these are sufficiently large and their ramets are also tall enough.Egg-laying preferences of the xerophilous ecotype of Maculinea alcon 63 the sprout-colonies with a medium number (3-4) of ramets (Fig. Table 1. 1. consequently they suit less for egg-laying. The number of eggs on the food plant by verticilia and organs.3%. verticil 130 20 232 28 3. The second verticil carried 7. We could find practically no eggs on the lowest verticilia. We found significantly more eggs on the colonies with 5-6 or more ramets than could be expected from their frequency in the quadrates. The distribution of eggs on the verticilia showed the expected proportions. Abax: abaxial surface of the leaves. flowers and flower-buds belonging the uppermost verticil. we have to consider that these colonies were the most frequent in the quadrates (45-50%). empty bars: the number of eggs. because it is easier for the searching females to find the larger sprout-colonies in the vegetation. The largest fraction (89%) of eggs was found on the leaves. 3).8%. the upper surface of the Fig. Nevertheless. We have found a strong negative correlation between the number of the eggs and the number of verticilia.7% of the eggs. 3. The distribution of the sprouts and eggs among the sprout colonies with different number of ramets in the randomly chosen squares (5000 m2). verticil Flower Stalk Adax Abax 1444 37 2932 571 2. the height of the ramets and the number of verticila with flowers/flower buds essentially influenced the egg laying preferences of females. 4a. Compared to the 90’s. this ratio still indicates that the flowers were the most attractive parts of the plant for the females. This conclusion is also supported by the fact that flowers only appear during the last part of the flight period of imagos. however. the number of ramets of the Gentiana sprout-colonies. In addition. . as well. Fig.64 Ervin Árnyas et al. Considering. In summary. the large surface area of leaves compared to the flowers. 4a). the flowers and flower-buds proved to be the most attractive for the females.01% of the eggs on the stems of the plants. significantly more eggs were found on the adaxial surface of the leaves and on verticilia with flowers/flower buds than on other parts of the plant. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). we found a fourfold increase in the number of eggs per ramet. During our surveys (in 2004) we have found significantly higher number of eggs on taller than on shorter ramets (Fig. The short grass structure of the sward maintained by mowing and grazing proved to be beneficial both for the growth of the sprout colonies and for the Maculinea population. we also searched for the association between the number of the eggs and some properties of the ramets. Furthermore. The proportion of the eggs on the leaves compared to the flowers and flower-buds was 2:1. Significant positive correlation was also found between the number of eggs and the number of verticilia with flowers/flower buds (Fig. Correlation between the height of the sprouts (cm) and the number of eggs laid on them Fig. 4b). 4b. thus the possibility of egg-laying is limited on them. We have only found 0. Correlation between the number of the verticilia with flowers and the number of eggs laid on them leaves. 1758) and Maculinea rebeli (Hirschke. [1779]). arion. rebeli. 10123 Turin. behaviour in the myrmecophilous butterflyE.it INTRODUCTION Four Maculinea species are present in Italy M. 1) and are the most endangered. alcon and M. Francesca Barbero & Emilio Balletto University of Turin. 1904) are wide ranging: M. while M. The 2 hygrophilous species. Maculinea arion (Linné. Settele. are restricted to the extreme north of Italy (Fig. Andrea Crocetta.© PENSOFT Publishers Oviposition Sofia – Moscow J. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.A. The two mainly xerophilous species. M. 1. Department of Human and Animal Biology. rebeli occurs from the Alps to the Central Italian region of Molise. Maculinea alcon ([Denis & Schiffermüller]. Via Accademia Albertina 13. 1775) and Maculinea teleius (Bergsträsser.bonelli@unito. 65-68 Oviposition behaviour in the myrmecophilous butterfly Maculinea alcon (Lepidoptera: Lycaenidae) Simona Bonelli. in contrast. teleius. J. arion reaches the south of the Italian peninsula. Kühn & of Butterflies(Eds) 2005 Maculinea Thomas in Europe alcon 65 Studies on the Ecology and Conservation Vol. Italy Contact: simona. M. pp. . Fig. by Candelo (BI). two contrasting hypotheses exist. other parts are subject to sheep grazing and still other parts have not been managed for the past 15 years. this SCI has been listed among the 36 Italian Prime Butterfly Areas (Balletto et al. (i) van Dyck and his co-workers (2000) hypothesised that M. at the foothills of the south-western Alps near the French border. alcon and M. The site is a small (3. as well as the study area. both butterflies parasitize Myrmica tulinae Elmes. With regard to which plants are chosen by females for egg laying. For this reason. all Molinia meadows used to be cattle grazed. Some of the meadows are mown every year. we have been monitoring M. Do butterflies choose gentians for egg laying? If so. alcon is present on an area of about 2 ha. vi) plants’ stem diameters. v) distance of plant from the nearest Myrmica nest. Aktaç. Our study area lies within the broader Site of Communitarian Interest of “Monte Musiné– Laghi di Caselette. Lycaena dispar and Zeryntia polyxena) are found. teleius population dynamics by the MarkRelease-Recapture (MRR) method.5 ha. 4. The other site is also part of a natural reserve (Riserva Naturale Orientata della Vauda).Brich Zumaglia Mont Préve” Regional Park. alcon to oviposit. Two additional Maculinea alcon sites were investigated because of their different management regimes. teleius occur in cohabitation is located in Piedmont. ii) turf height. leaf and flower). At out study site. We catalogued flowers by their blooming stage (from green buds to open and faded flowers). In summer 2001. iv) number larval exit holes. Most of them. Maculinea arion. Parts of the sward are burnt on average every 2 years. In 2002 there were 971 eggs (176 exit holes). The first is a Molinia coerulea meadow within the “Baragge .5 ha) wet meadow dominated by Molinia coerulea. The area is subject to a mild mowing regime and light sheep grazing. alcon is 3. Since 1997. while some meadows have not been managed for about 10 years. suggested that oviposition occurs on high quality plants. we recorded egg-positions on the basis of the plant’s nodes (counted from the apex) and parts (stalk. where another 3 endangered species (i. Both species had large year-to-year fluctuations. iii) number and position of M. others every two years. Coenonympha oedippus and M. (ii) Thomas and Elmes (2002). arion was assessed by MRR. that we wished to test. 2004). alcon and M. e. We visited each gentian once a week and noted: i) plant height. we addressed the following questions: 1.Bessa . are now mown for fodder. More specifically. The same meadow is inhabited also by two other endangered butterfly species: Euphydryas aurinia and Coenonympha oedippus. we investigated 484 eggs and noted 103 exit holes. On each plant. in contrast. The only Italian site were M. we monitored respectively 55 and 75 gentians used by M. Until the 70’s. In 2005 the population size of M.66 Simona Bonelli et al. 2. Radchenko. 5. alcon females oviposit on those gentian plants that are closest to a host-ant nest. 3. 6. how do butterflies choose gentians for egg laying? On part(s) of the food plant do butterflies choose for egg laying? Do females choose any further among suitable buds? Do females clump eggs on suitable buds? Does butterfly choice of plant parts affect initial larval survival? MATERIALS & METHODS In 2001 and 2002. . 2002. The area with M. alcon eggs. Oviposition behaviour in the myrmecophilous butterfly Maculinea alcon 67 In 20 (5x5 m) plots, chosen in variously mown areas, we evaluated: i) plant density, height, distance from the nearest Myrmica nest and bud numbers of individual G. pneumonanthe; ii) Myrmica. tulinae nest densities; iii) the position of individual Maculinea alcon eggs; iv) the number of Sanguisorba officinalis plants; v) the height of the grass surrounding each gentian plant; vi) temperature (by a Buttons® data logger); vii) Myrmica nest size (described as nest diameter [max> 20 cm min<15 cm mean 15 e 20 cm]) Data were analysed with SYSTAT ® for Windows. We estimated population sizes by FisherFord’s method. RESULTS & DISCUSSION In 2001-2002 the flight period at our site was from the 20 July to 23 August. We marked 244 adults (98 females) in 2001 and 327 (163 females) in 2002. The flight peak was around the middle of August. By CRM, we estimated that 424 adults were present at our site in 2001 and 726 in 2002. In summer 2001, we investigated 484 eggs and noted 103 exit holes. In 2002 there were 971 eggs (176 exit holes). During the orientation phase of oviposition, M. alcon females were attracted by the most luxuriant and robust gentian plants. (Females choose the highest and most robust plants having several buds but without flowers in full bloom) On the chosen plant females select immature buds, growing as close as possible to the plant’s apex. Females avoid plants, with blue flowers and they are not influenced by plant closeness to a prospective host ant nest. In more detail: 1) Egg distribution on gentians is clumped and non-random (C2 33.89, DF 2, P<0.001) The numbers of gentians that were not chosen, or were chosen by at least 2 different females, far exceed those expected on the basis of random-choice hypothesis. Eggs distribution proved non-random. On each plant, eggs were generally laid by two females, rarely by three or more females. 2a) Females did not choose plants on the basis of their closeness to a potential hostant colony. In fact, avoided plants were not growing farther from a potential host nest than eggs-bearing plants (KW: P=0.646 – 1096 cases). Furthermore we were unable to demonstrate any correlation between the number of eggs laid on each plant and the distance from the nearest host ant nest. In this respect, however, we need to keep in mind the very high density of Myrmica tulinae nests at the site (0.62 nests/m2). As a consequence, all gentians probably laid inside the foraging range of at least one ant colony. 2b) Plants chosen for oviposition and eggs numbers on each plant were positively correlated with some characters of the plant’s phenology Eggs-bearing gentian plants were different in aspect from the avoided ones. They are higher and show more blue bud, but have fewers flower in full bloom. The number of eggs laid per plant was positively correlated with the plants’ stem diameters (r=15.80 P<0.001) and sward height (r=0.38 P=0.02). 68 Simona Bonelli et al. A strong correlation exists between egg number and bud s number (Spearman ñ=0.541, p<0.001, 75 cases). 3a) Females avoided leaves and stalks and actively chose buds (86% of eggs in 2001, 90% in 2002). 3b) Buds were in their initial (but not very early) blooming stage: the numbers of eggs laid on the different bud categories was statically different (KW, 2001: test stat 17.218 (201 cases), P=0.004. 2002: test stat 24.742 (376 cases), P<0.001). 3c) Females actively chose green buds. Eggs laid on these buds amounted to 65% in 2001 and 86% in 2002. 4a) Eggs were consistently laid as close as possible to the plant’s apex (Kendall’s coefficient of concordance W=0.350 (2001), P=0.004; W= 0.536 (2002), P<0.001). Preference for the apical part of plant was constant among samples and between the two years of study. Since gentians have more flowers in their apical part than lower down along the stem, and since the plant’s apex is the most apparent part of the plant, with respect to the surrounding vegetation, it is possible that this choice is driven by “visibility” rather than by other factors. 4b) The total number of eggs per node is correlated with the number of flowers per node (Spearman. 2001: ñs=0.985 (10 cases), P<0.001; 2002: ñs=0.957 (10 cases), P<0.001). At the apex of the gentian plant there are more flowers. 5) The observed number of eggs laid per bud was low. On average they were 2.1 in 2001 and 2.3 in 2002. This behaviour will limit larval competition inside the bud. 7a) Immature (at egg-laying time) flowers provided the highest larval success. 7b) Larval success did not vary between eggs laid at the plant’s top or in other locations. Gentians showing the above reported features were more likely to be present in the irregularly mown parts of the site, both at Caselette and Candelo, as well as in burnt or un-mown meadows at Vauda. In fact eggs density is significantly (KW: p=0.048; Mann-Whitney U test statistic: p=0.003) different between meadows subject to different management regimes. Although the highest concentration of gentians was observed in the yearly mown swards, females mainly chose plants growing in irregularly mown areas (once in several years), where gentians had scarce recruitment, host ant density is low but were ant nests are bigger in size. These empirical results therefore support both the hypothesis that Maculinea oviposition is not mediated by the presence of ants, and the theoretical predictions of a recent Macman model describing the responses of interacting M. alcon, G. pneumonanthe and M. scabrinoids populations to management (Mouquet et al. in press). REFERENCES Mouquet, N. Thomas, J. A., Elmes, G. W., Clarke, R. T. & Hochberg, M. E. “in press”. Conserving community modules: a case study of the endangered lycaenid butterfly Maculinea alcon. Ecology. Thomas, J.A. & Elmes, G.W. (2001) Foodplant niche selection rather than the presence of ant nests explains oviposition patterns in the myrmecophilous butterfly genus Maculinea. Proc Roy Soc B: 268, 471-477 van Dyck, H., Oostermeijer, J.G.B., Talloen, W., Wynhof, I., Feenstra, V. & van der Hidde, A. (2000) Does the presence of ant nests matter for oviposition to a specialised myrmecophilous Maculinea butterfly? Proc Roy Soc B 267, 861-866. © PENSOFT Publishers Host specificity Sofia – Moscow J. E. J.A. Thomas in Microdon myrmicae, a sympatric socialEcologySettele,to Kühn & of Butterflies(Eds) 2005 the Maculinea in Europe 69 Studies on the parasiteConservation and Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, pp. 69-71 Host specificity in Microdon myrmicae, a sympatric social parasite to the Maculinea in moist grassland ecosystems Simona Bonelli1, Andrew D.P. Worgan2, Sophie Everett2, Emma Napper2,3, Graham W. Elmes2, Ania M.Stankiewicz 4, Marcin Sielezniew5, Judith C. Wardlaw2, S. Cantarino1, Andras Tartally6, Emilio Balletto1 & Karsten Schönrogge2 University of Turin, Department of Human and Animal Biology, Via Accademia Albertina 13; 10123 Turin; Italy 2 NERC - Centre for Ecology & Hydrology, CEH Dorset, Winfrith Technology Centre, Winfrith Newburgh, Dorchester, Dorset DT2 8ZD, UK 3 Rothamsted Research, Biological Chemistry Division, Harpenden, AL5 2JQ, UK Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland 5 Department of Applied Entomology, SGGW - Warsaw Agriculture University, Nowoursynowska 159, 02-776 Warszawa, Poland 6 DUFS - University of Debrecen, Faculty of Sciences, Department of Evolutionary Zoology and Human Biology, P.O.B. 3, H-4010 Debrecen, Hungary Contact:
[email protected] 1 4 Species once regarded as single entities are increasingly found to be ecologically and genetically diverse, with many displaying local adaptations and several containing cryptic species. Understanding variation within morpho-species is vital both for the preservation of genetic diversity and practical conservation management. Obligate social parasites are extreme specialists equipped to infiltrate insect societies and feed on the resources inside nests (Thomas et al., 2002; Thomas & Settele, 2004). Most are very localised compared to their hosts and may consist of multiple cryptic species adapted to different host species (Als et al., 2004; Thomas & Settele, 2004). Social parasites such as Maculinea butterflies or Microdon hoverflies are abundantly represented in national and international red data lists, yet being highly adapted to infiltrate host ant societies these are also species where the formation of local races or even cryptic species might be expected (Schönrogge et al., 2002). For its simple life-history involving a short lived (<10 days) adult stage outside the host ant nest, while both the larval and pupal development occur inside the ant nests, the obligate myrmecophile hoverfly Microdon mutabilis is arguably more typical for myrmecophiles than Maculinea species. Yet it is exceptional in that within a population on the Isle of Mull (Scotland) egg mortality from ant attack increased sharply in Formica lemani colonies over short distances from the mothers natal nest suggesting that these flies are adapted not at the species level, but at population or even sub-population level (Elmes et al., 1999). While this is often referred to 70 Simona Bonelli et al. as extreme host specificity, there are in fact few appropriate studies from other myrmecophile species (Thomas et al., 2005). Also such extreme host specificity stands in contrast to historical records that list various Formica and Lasius species among the hosts of M. mutabilis (Andries, 1912; Donisthorpe, 1927). Within the British Isles we found a sharp distinction between Microdon using F. lemani in Scotland and Ireland, but Myrmica scabrinodis, a host not previously recorded, in the south-west of England (Schönrogge et al., 2002). In Scotland and Ireland together we opened 1680 nests of F. lemani and found 422 (25%) contained Microdon larvae or pupae, while we found no Microdon in 337 nests of M. scabrinodis. Twenty out of 1955 nests (1%) of other ant species also contained Microdon larvae, but it is thought that the original host colony might have vacated nests, probably under the predation pressure of the Microdon, and that other ant colonies might have moved subsequently into the nest site (Schönrogge et al., 2002). In contrast, 65 M. scabrinodis nests out of 162 (40%) were found to host Microdon larvae, while only 3 out of 113 (3%) nests of other ant species contained Microdon. F. lemani is not found in the habitats where M. scabrinodis hosts Microdon. Using morphometric measurements particularly on the pupae, we recently showed that British populations consist of two sibling species, M. myrmicae using M. scabrinodis while M. mutabilis is associated with F. lemani (Schönrogge et al., 2002). Since M. myrmicae was described from Britain and because under MacMan Myrmica colonies have been studied intensively, its presence has been confirmed for Ireland (Speight, 2002) and Poland (Stankiewicz, 2003), we know of records from France and Germany and for the current study we have sampled this species in Britain, Poland, Hungary and Italy. In seven populations on the European continent sampled M. myrmicae lives sympatrically with other myrmecophiles including Maculinea alcon, M. nausithous and M. teleius (in one location all four species co-occur) and in at least one population M. alcon and M. myrmicae have been found to co-inhabit the same ant colonies. It is reasonable to assume that the distribution of M. myrmicae includes the whole of Europe and possibly, like M. mutabilis, even the western and eastern Palaearctic. In contrast to M. mutabilis, M. myrmicae has now been recorded with 2 possibly 3 Myrmica species: M. scabrinodis, M. gallieni and M. tulinae although the identity of the latter has yet to be confirmed. We will use morphometric measurements on both the adults and pupae similar to Schönrogge et al. (2002). However, Elmes et al. (1999) also argued that M. mutabilis eggs should be protected by chemical mimicry because: 1. Mimetic compounds, cuticular hydrocarbons, have been found on the larvae of nearctic Microdon species and were described to be bio-synthesised by the larvae (Howard et al., 1990a; Howard et al., 1990b). 2. Microdon eggs that were kept in isolation in sterile vials for a few days before being introduced to ant nests were increasingly more likely to be attacked by workers from the maternal colony but less likely to be attacked by workers from a distant colony, the longer the interval between oviposition and introduction to ants. This suggests a chemical compound or compounds that are to some degree volatile (Elmes et al. 1999). 3. Other mimetic mechanisms used by the adult or larval stages of myrmecophiles, such as behaviour or acoustic signals, should not be possible for eggs. Following this argument we will study the surface chemistry of the eggs for both M. mutabilis and M. myrmicae and investigate whether M. myrmicae could contain further cryptic species with reference to their host use, or alternatively the likelihood that this species while Myrmica specific, is more generalist than its sister species M. mutabilis. Host specificity in Microdon myrmicae, a sympatric social parasite to the Maculinea LITERATURE 71 Als, T.D., Vila, R., Kandul, N.P., Nash, D.R., Yen, S.H., Hsu, Y.F., Mignault, A.A., Boomsma, J.J., & Pierce, N.E. (2004) The evolution of alternative parasitic life histories in large blue butterflies. Nature, 432, 386-390. Andries, M. (1912) Zur Systematik, Biologie, und Entwicklung von Microdon Meigen. Zeitschrift für Wissenschaftliche Zoologie, 103, 300-361. Donisthorpe, H.J.K. (1927) The guests of British ants - their habits and life-histories George Routledge and Sons Ltd., London. Elmes, G.W., Barr, B., Thomas, J.A., & Clarke, R.T. (1999) Extreme host specificity by Microdon mutabilis (Diptera : Syrphidae), a social parasite of ants. Proceedings of the Royal Society of London Series BBiological Sciences, 266, 447-453. Howard, R.W., Akre, R.D., & Garnett, W.B. (1990a) Chemical mimicry in an obligate predator of Carpenter Ants (Hymenoptera, Formicidae). Annals of the Entomological Society of America, 83, 607-616. Howard, R.W., Stanley-Samuelson, D.W., & Akre, R.D. (1990b) Biosynthesis and chemical mimicry of cuticular hydrocarbons from the obligate predator, Microdon albicomatus Novak (Diptera, Syrphidae) and its ant prey, Myrmica incompleta Provancher (Hymenoptera, Formicidae). Journal of the Kansas Entomological Society, 63, 437-443. Schönrogge, K., Barr, B., Wardlaw, J.C., Napper, E., Gardner, M.G., Breen, J., Elmes, G.W., & Thomas, J.A. (2002) When rare species become endangered: cryptic speciation in myrmecophilous hoverflies. Biological Journal of the Linnean Society, 75, 291-300. Speight, M.C.D. (2002) Two controversial additions to the Irish insect list: Microdon myrmicae Schönrogge et al. and Pipiza festiva Meigen (Diptera: Syrphidae). Bulletin of the Irish Biogeographical Society, 26, 143-153. Stankiewicz, A. (2003) Hoverfly Microdon myrmicae SCHONROGGE et al., 2002 (Diptera: Syrphidae) in Poland Polish Journal of Entomology, 72, 145. Thomas, J.A., Knapp, J.J., Akino, T., Gerty, S., Wakamura, S., Simcox, D.J., Wardlaw, J.C., & Elmes, G.W. (2002) Parasitoid secretions provoke ant warfare. Nature, 417, 505-506. Thomas, J.A., Schönrogge, K., & Elmes, G.W. (2005). Specializations and host associations of social parasites of ants. In Insect Evolutionary Ecology (eds M.D.E. Fellowes, G.J. Holloway & J. Rolff), pp. 479 518. CABI Publishing, Wallingford, UK. Thomas, J.A. & Settele, J. (2004) Evolutionary biology - Butterfly mimics of ants. Nature, 432, 283-284. Research has partly been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). J.A. Thomas © 72 PENSOFT Publishers Jarosław Buszko, Marcin Sielezniew & Anna M. StankiewiczJ. Settele, E. Kühn & of Butterflies(Eds) 2005 Studies on the Ecology and Conservation in Europe Sofia – Moscow Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, p. 72 The effect of ant communities and spatial pattern for Maculinea nausithous Uta Glinka & Josef Settele UFZ - Centre for Environmental Research Leipzig-Halle, Department of Community Ecology, Theodor-Lieser-Str. 4, 06120 Halle, Germany Contact:
[email protected] The development of Maculinea nausithous is obligately dependent on the occurrence of ants. For ant communities in general, competition is a major organizing principle and probably the hallmark of ant ecology. We therefore tried to determine general nest patterns in ant communities, where Maculinea nausithous is present or where it is not but where the host plant occurs. Furthermore, we investigated how density, size or distance of ant nests affected the occurrence or the population size of M. nausithous. Data on species richness and nest abundance of ants were collected at three plots in each of the 24 managed grasslands. Within those, the position and characteristics of all ant nests where mapped. RESULTS All habitats are characterised by a low diversity of ant species. The most common community consisted of only three species M. rubra, M. scabrinodis and L. niger. In contrast, nest densities were quite different. There was a clear dominance of M. rubra in half of the habitats. The mean nest density was 17 nests per 100m². In contrast, compared with M. scabrinodis dominated in just 6 cases with an mean abundance of 10 nests per 100 m². Ant species richness was not related to vegetation cover. We tested the given ant nest pattern against different null models (e.g. pattern one fixed, pattern 2 random). The spatial distribution of M. rubra nests was mostly randomly. This indicates that there is no territorial behaviour around the nest. Fields with a high nest density of M. rubra exhibit additionally a higher population size of M. nausithous. Also, nest size of M. rubra was positively correlated with population size of butterflies. For M. rubra we found no correlation between nest size and nest density. Furthermore, a method of simplification for the assessment of ants in Maculinea habitats was carried out by means of ant baiting in comparison to ant nest searching. It was shown, that the results of both methods are quite similar with regard to species diversity as well as to the quantity of a particular species. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Females laid most of their eggs on the flowers of G. 73 Contrasting egg laying behaviour of the ecotypes of Maculinea alcon in Hungary Ferenc Kassai & László Peregovits HNHM . We believe. while the upper side of leaves was preferred in the case of G. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). In this study host plant preference and the distribution pattern of eggs laid on the host plant are compared of the marshland and xerophilous ecotypes of Maculinea alcon. Both ecotypes preferred the upper node of the shoots. and ecology of MaculineaStudies on and M. rebeli. We regard the two major groups as xerophil (M. the relationship between the number of flowers and the number of eggs can be described with a gaussian function. since the latter depends on whether there is a host ant nest within foraging range of the selected plant specimen. eggs may have different survival probabilities on different parts of the host plant. Thomas in Europe 73 and Conservation Poland Vol. Egg counting was carried out in 1992 in a dolomite grassland at Nagy-Szénás (N Hungary) and in 1999 on a marshland habitat at Kunpeszér (C Hungary). cruciata. pneumonanthe. In the case of Maculinea species. From the last two parameters we calculated the apparency of shoots.© PENSOFT Publishers The distribution Sofia – Moscow J. In the case of the marshland ecotype. alcon species group is still uncertain. E.nhmus.A. pneumonanthe. namely: number of flowers (number of nodes with flowers in the case of G. The mean number of eggs per shoot was higher in the case of G. alcon). height of the surrounding vegetation. Taxonomy of the M. Furthermore. that the observed trends are similar in other populations. Settele.Department of Zoology. The main hostplant of the xerophilous ecotype is Gentiana cruciata. Parameters describing the structure of the shoot were measured. and some were also found in the case of the other foodplant. pneumonanthe. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. host plant selection and finding the host ant species are coupled.hu In general. M. Baross u. while the marshland ecotype used in most cases is G. tolistus) and as marshland ecotypes (M. Stems were randomly selected and eggs were counted at different parts of the shoots. cruciata). Kühninof Butterflies(Eds) 2005 teleius the Ecology nausithous & J. Hungary Contact: perego@zoo. Hungarian Natural History Museum H-1088 Budapest. xerophila. . The more prominent shoots received more eggs in the case of the xerophilous ecotype. selection of the appropriate host plant specimen is essential for the survival of the larvae of a butterfly. where the optimum number of flowers is six. M.zoo. height. cruciata. The number of shoots without eggs was extremely high in the case of G. although the pattern formed by these behavioural trends depends strongly on the extent to which the host plant resource is limited. 13. p. Fourth instar larvae drop off the plant and wait to be found and adopted by foraging ant workers on the soil surface. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 1904 Ádám Kőrösi HNHM . E. 861-866.nhmus. attempts at mating. Clarke Sofia – Moscow J.hu Larvae of Maculinea butterflies are obligate social parasites of Myrmica ants. Talloen W. REFERENCES Thomas. . We studied females of Maculinea rebeli. Settele. Hidde A van der.A. alcon occupying dry calcareous grasslands and ovipositing mainly on Gentiana cruciata. & Elmes. Hungary Contact: korosi@zoo. (ii) Females prefer plants that are in a phenophase suitable for oviposition (since small larvae need young buds). 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. This phase lasts for 1 week. G. a species which has been recently demonstrated to be an ecological type of M. J. Elmes & Ralph T. the caterpillars spend some weeks on the foodplant consuming the developing seeds. p. All these studies examined the spatial distribution of eggs and ant nests. so significantly more egg-laying will be detected on these plants – at least in the first section of the flight period. For this reason it is crucial for the butterfly to lay the eggs on a plant within the foraging range of a host ant nest. but not the mechanism that generates the observed pattern. There have been many studies published about egg-laying preferences of Maculinea alcon sensu lato. After hatching. Hungarian Natural History Museum H-1088 Budapest. some of them argued that egg-laying depends on presence of host ants (Van Dyck et al. (2001) Foodplant niche selection rather than the presence of ant nests explains oviposition patterns in the myrmecophilous butterfly genus Maculinea. cruciata shoots and number of laid eggs were recorded in order to test some hypothesis on ovipositing preferences: (i) females prefer gentian plants standing in the foraging range of a host ant nest. Feenstra V. Proceedings of the Royal Society B. so different plants are used for egg-laying in the beginning and in the end of the flight period. Baross u. while others contradict this and emphasize the role of suitable phenophase of the foodplant (Thomas & Elmes 2001). 13.Department of Zoology. 267. 268.zoo.© 74 PENSOFT Publishers Graham W. which means that they can develop only in nests of host ant species. Van Dyck H.A. Kühn & J. Wynhoff I (2000) Does the presence of ant nests matter for oviposition to a specialized myrmecophilous Maculinea butterfly? Proceedings of the Royal Society B. Females were tracked for 1-2 hours and all ovipositing events. 471-477. 2000). number of visited G.W. (iii) Due to high spatial variation in caterpillars’ survival ’risk spreading’ is an optimal strategy. Oostermeijer JGB. 74 Egg-laying behaviour of Maculinea rebeli Hirschke. so movement pattern of ovipositing females has some directionality in order to maximize the net displacement. We studied the behaviour of both the ants and the caterpillars (see also: Elmes et al. 3. 2001). whereas.2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group. 75-77 Behavioural aspects of adoption of Maculinea caterpillars by Myrmica ants Szabolcs Lengyel1.B.A. 10% and 0% in the first. We added Maculinea alcon and M. alcon and M.University of Debrecen. pp. Emese Szitta2. Adoption during 10 min was rare (30%) in the first round of trials. Hungary Contact: szabolcs@delfin. The number of ants licking negatively influenced 10-min adoption. The number of physical contacts by ants. H-4010 Debrecen.© PENSOFT Publishers Behavioural Sofia – Moscow J. Adoption patterns were not influenced by the source (locality) of the ant colony or whether the colony contained queens. rebeli caterpillars to colonies of Myrmica scabrinodis ants collected in the field at M. time spent smelling positively affected 24-hr adoption and caterpillar survival. Hungary 2 DUFS . and short-term rejection was more often associated with 24-hr adoption than in 2003. rebeli sites and recorded the behaviour of caterpillars and ants by a video camera for 10 min and for 5 min 24 hours later. Mariann Bíró2 & Zoltán Varga1. András Tartally2. 3.O. both its total time per trial and the number of participating ants. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. when 10-min adoption was associated with 24-hr rejection (caterpillar death). Settele. H-4010 Debrecen.B.hu 1 Adoption of Maculinea (Lepidoptera: Lycaenidae) caterpillars by Myrmica (Hymenoptera: Formicidae) ants is a crucial stage of the butterfly life cycle because mortality can reach 85% at this stage under laboratory conditions (Als et al. Faculty of Sciences. respectively) because caterpillars died in later trials. and became more frequent in the second (45%) and third rounds (90%) in both butterfly species. (1) In the adoption experiment in 2003. Caterpillars did not survive if they were not adopted after 24 hrs. 24-hr adoption changed in the opposite direction (80%. whereas four caterpillars actively crawled into the colony. . In 2004. E. second and third rounds. However. and smelling.unideb. P. were positively related to 24-hr survival of the caterpillar. Department of Evolutionary Zoology and Human Biology. Thirty-six out of 60 caterpillars showed some form of movement during the trials and caterpillars moved towards the ant colony more often than to any other directions. the behaviour of the ants was more repeatable.O. However. P. whereas the frequency of long-term (24-hr) adoption was high in round 1 and became rare by round 3. we were interested in whether the success of adoption is related to the behaviour of either the caterpillar or the ants. we repeated the experiment in M. rebeli by using larger arenas and ant colonies and with colonies containing queens/broods. Kühn & J. We used caterpillars only once and ant colonies once per week for three weeks. similarly to the results of 2003. Short-term (10-min) adoption was equally frequent in the three rounds. Thomas aspects of adoption of Maculinea caterpillars Conservation of Butterflies(Eds) 2005 75 Studies on the Ecology and by Myrmica ants in Europe Vol. 1991) during the process of adoption by conducting two experiments. G. and their Myrmica host ants: wild adoption and behaviour in ant-nests.R. teleius (43%). co-occurring non-host ants (Myrmica ruginodis for Maculinea rebeli. prey on the ant brood. i. rebeli caterpillars did not move from the centre of the arena. 2001: Adoption of parasitic Maculinea alcon caterpillars (Lepidoptera: Lycaenidae) by three Myrmica ant species. However. J. and caterpillars survived better if they were adopted only later (i. n = 76) and M. teleius caterpillars actively crawled to the chambers.W. & Boomsma. after 10 min). D.e..J. Fourth instar naive caterpillars of M. Some M. We predicted that caterpillars will move towards and spend more time in the vicinity of host ants than non-host ants and near the control chamber than near the non-Myrmica chamber. rebeli (“cuckooing” caterpillars. .. Although M. These experiments show that behaviour can be important in understanding how adoption of Maculinea caterpillars by ants occurs. the behaviour of ants can be used to predict the success of adoption. 1991: Larvae of Maculinea rebeli. REFERENCES Als. fed by the ants. Elmes. In summary. i. quick adoption was associated with low caterpillar survival. The difference in the distance crawled was highest in the host vs. teleius caterpillars than to cuckooing M. control and in the non-host-Myrmica vs. Nash.. Fiedler 1990) and if they differentiate between host and non-host ant colonies.. and there also was no clear effect of treatment (i. rather. pairwise combinations). Myrmica rubra for Maculinea teleius).g. Our results showed that the predatory Maculinea teleius caterpillars had more body movements and crawled more than the cuckoo caterpillars of M. J. non-Myrmica ant comparisons. non-host) vs. Each of the three ant species was collected on the site where the caterpillars were from. J.. n = 62) were introduced in the middle of rectangle-formed arenas. whereas 16% of M. teleius (predatory caterpillars. a large-blue butterfly. the difference was not statistically significant. co-occurring non-Myrmica ants (Lasius niger) or no ants (controls). e. rebeli.. (2) In a caterpillar choice experiment we tested if the caterpillars are able to search actively for the ant nests (see e. rebeli chose the predicted half of the arena more often (55%) than M. moving towards host vs. Thomas & Elmes 1998. ACKNOWLEDGEMENTS Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126).. they sometimes moved actively towards colonies. rebeli caterpillars.e.e.76 Szabolcs Lengyel et al. The behaviour of caterpillars appeared to be related to their life history because movements were more characteristic to predatory M. Both species of caterpillars started active movements to neutral directions first and the frequency of choosing the predicted direction (e. Caterpillars were not passive participants in adoption. Thomas. There also were no differences in the time until the first movement.e. T.C. Animal Behaviour 62: 99-106. Thomas & Elmes 1998.A.D.g. The chambers were attached to the arena so that it was possible for the caterpillar to perceive the odours and sounds of the ants but the possibility of direct physical contact between the ants and the caterpillars was excluded.g.. – Journal of Zoology (London) 223: 447-460. & Wardlaw. At both ends of the arenas small chambers contained pairwise combinations of host ant species (Myrmica scabrinodis). the non-expected direction did not differ between the species. e.g. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). G. K. 1990: New information on the biology of Maculinea nausithous and M. . 1998: Higher productivity at the cost of increased host-specificity when Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation. teleius (Lepidoptera: Lycaenidae).A. – Nota lepidopterologica 12: 246-256. & Elmes. J.W.Behavioural aspects of adoption of Maculinea caterpillars by Myrmica ants 77 Fiedler. – Ecological Entomology 23: 457-464. Thomas. p < 0. Theodor-Lieser-Str. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Department of Community Ecology. Dorchester. UK Contact: Holger. CEH Dorset. Winfrith Newburgh. E. followed and aggression behaviour observed for ten minutes. Christian Anton1. We conducted 104 adoptions on 60 nests (94 % adoption rate).054). nausithous-caterpillars. five meadows and five fallow patches. Kühn & J. Pearson’s Correlation of adoption time and caterpillar weight was positive and significant (rp = 0. shows a highly variable colony structure in these different habitats [1].A. To test for influence of aggression levels on adoption behaviour. number of workers per nest. Colony structure at these sites can be considered multi-colonial. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. 60 ant nests were used for adoption experiments with fourth instar M. We hypothesised that ant colonial structure affects Maculinea adoption behaviour by altering colony aggression levels.Centre for Environmental Research Leipzig-Halle. There was no effect of habitat type on aggression levels.and multi-colonial in the UK and Finland [1].de.01). 78-79 Does the colony structure of Myrmica rubra affect the adoption of Maculinea nausithous? Holger Loritz1. We conducted inter-colonial aggression experiments with 90 ant nests collected along a transect of 100-150 m at ten different sites. Josef Settele1 & Karsten Schönrogge2 UFZ . At five sites there was a significant increase of aggression level with increasing inter-nest distance.Centre for Ecology & Hydrology. number of queens per nest and habitat type as independent variables. Single alien worker ants were introduced into experimental
[email protected] and p = 0. unicolonial in the invasive regions of North America. Christian. Settele. and at three further sites significance levels were marginally significant (p-values between p = 0. Excavated nests were split into “home” and “experimental” nests.54. 06120 Halle. . Winfrith Technology Centre. uni. Colony structure is also variable across the ant’s range.© 78 PENSOFT Publishers et al. Holger Loritz Sofia – Moscow J. 4. pp. the red ant Myrmica rubra.de 1 The Dusky Large Blue (Maculinea nausithous) inhabits different habitats ranging from regularly managed meadows and pastures to unmanaged and overgrown fallow land.Anton@ufz. We performed an analysis of covariance with mean adoption time as the dependent variable and nest aggression level. Dorset DT2 8ZD. Caterpillars were introduced into “home-nest” units and observed for 30 minutes. Only the number of workers per ant nest had a significant positive effect on adoption time. Its secondary important habitat resource. Germany 2 NERC . REFERENCES [1] Seppä.Does the colony structure of Myrmica rubra affect the adoption of Maculinea nausithous? ACKNOWLEDGEMENTS 79 We thank Franziska Faber. Walin (1996): Sociogenetic organization of the red ant Myrmica rubra. & L. P. This study has been funded by the EC within the RTD project “MacMan” (EVK2-CT-2001-00126). Ulrike Mühle and Christina Otto for help in the laboratory. Behavioral Ecology and Sociobiology 38 (3):207-217 . After feeding on the flower heads of Sanguisorba officinalis larvae of the Dusky Large Blue (Maculinea nausithous) are adopted by workers of the host ant Myrmica rubra and carried into the ant nest. pp. CEH Dorset. Winfrith Newburgh. Christian Anton1. Andrew Worgan2 & Josef Settele1 UFZ . nausithous is influenced by olfactory cues of the host ant. UK Contact: Martin. Instead. or whether host plant related characters determine the selection of the oviposition site. Germany 2 NERC . Winfrith Technology Centre. Host plant choice of the co-occuring M. Deposition of eggs on plants in close vicinity to the host ant should increase the survival of M. rubra nests from the onset of the flight season. The ant cue did not have any effects on the oviposition preference of M. We counted the number of eggs and measured host plant traits that might be related to the oviposition preference of M. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 80-81 Do ant cues influence the oviposition preference in the myrmecophilous Maculinea nausithous? Martin Musche1. 06120 Halle. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. . Dorset DT2 8ZD. all plants were removed from the field and brought to the lab. Settele. A field experiment was established to investigate whether host plant choice in M. teleius was similar to that of M.Musche@ufz. The survival of the caterpillars after leaving the host plant depends on the probability of being found by the host ant species. nausithous. officinalis were contaminated with soil from M. host plant traits as flowerhead size and phenology significantly influenced the distribution of eggs. flower heads with non-ant soil were used. Kühn & J. E. nausithous.de 1 Butterflies of the genus Maculinea are obligatory associated with specific ants of the genus Myrmica. female behaviour was independent from differences in butterfly density and did not change throughout the flight period. nausithous. Moreover. 4. Our findings seem to corroborate those found in other myrmecophilous lycaenids.Centre for Ecology & Hydrology.A. nausithous larvae. The result confirms the suggestion that host ant dependent oviposition appears to be a disadvantageous strategy in the face of resource limitation within ant colonies and the immobility of caterpillars (Thomas & Elmes 2001). Theodor-Lieser-Str. As a control.© 80 PENSOFT Publishers et al. Martin Musche Sofia – Moscow J. Dorchester. Flower heads of S. Inside the nest.Centre for Environmental Research Leipzig-Halle. caterpillars feed on the ant brood. Department of Community Ecology. After two weeks. 471-477 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). & Elmes. .W. Proc Roy Soc B: 268. (2001) Foodplant niche selection rather than the presence of ant nests explains oviposition patterns in the myrmecophilous butterfly genus Maculinea. G. nausithous? REFERENCES 81 Thomas. J.A.Do ant cues influence the oviposition preference in the myrmecophilous M. Baross u. M. At one of the study sites where Myrmica schencki was found to be an occasional host. we think that avoiding such plants might be beneficial to ants.University of Debrecen. 1088 Budapest. Judit Bereczki2. 2000). M.A. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Department of Evolutionary Zoology and Human Biology. a food resource of larvae in the phase of early development. there were more eggs on food-plants where M. 3. M. Maculinea alcon females lay their eggs on Gentiana food-plants on which the eggs hatch and the larvae start developing by feeding on developing seeds. Furthermore.© 82 PENSOFT Publishers et al. Our results do not support the hypothesis that egglaying Maculinea females are able to choose individual food-plants that are visited by host ants. This care is provided by Myrmica ants. Andrea Tóth2.hu 1 Maculinea are among the few genera in butterflies in which the larvae need care. Zoltán Varga2. László Peregovits1 & János Kis3 HNHM . Although we do not know anything about a possible underlying mechanism in ants to recognize such plants. at one of the study sites. Kühn & J. Dept. schencki was absent than where they were present. Hungary 2 DUFS . H-4010 Debrecen. alcon. Ervin Árnyas2. . and the species of ant hosts may vary among different populations of each Maculinea species. Edina Prondvai Sofia – Moscow J. REFERENCES Thomas. Therefore.B. &
[email protected] Natural History Museum. Sándor Csősz1. Proceedings of the Royal Society B. whereas other studies concluded that ant host presence had not influenced oviposition (Thomas & Elmes 2001). G. 82-83 Oviposition in Maculinea alcon butterflies Edina Prondvai1. schencki might avoided Gentiana loaded with many eggs. alcon females laid more eggs on food-plants with a high amount of flowers. E. 268. Department of Zoology.O. J.13. It has been suggested that females recognize food-plants that are growing within the range of ant hosts (Van Dyck et al. We studied food-plant choice in several Hungarian populations of Maculinea alcon in relation to the presence of host ants and the amount of flowers on food-plants. Settele. P. 471-477. pp. (2001) Foodplant niche selection rather than the presence of ant nests explains oviposition patterns in the myrmecophilous butterfly genus Maculinea.A. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Ecology Contact: kis.W. schencki’s microhabitat preference might not overlap at this site with the oviposition preference of M. Alternatively. There was not such relationship in other Myrmica species. The ants then collect the larvae and take them to their nests. the reproductive success of Maculinea females depends on their success in finding a suitable food-plant within the foraging range of their specific Myrmica species. Hungary 3 Szent István Univ. Faculty of Sciences.szie. Ferenc Kassai1. 861-866. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). Hidde A van der. 267. Oostermeijer JGB. . Feenstra V. Wynhoff I (2000) Does the presence of ant nests matter for oviposition to a specialized myrmecophilous Maculinea butterfly? Proceedings of the Royal Society B.Oviposition in Maculinea alcon butterflies 83 Van Dyck H. Talloen W. UK 2 La Grosterie. Poland 4 Department of Applied Entomology. Polish Academy of Sciences. Similarly. CEH Dorset. Kühn & J. Wilcza 64. pp. Sofia – Moscow J. Winfrith Newburgh. all ant-tended mutualists and social parasites appear to have the ability to stridulate via a file-and-scraper stridulatory organ (Fig 1). the ability to produce sound is widespread among lycaenid caterpillars in both mutualistic species and social parasites. Electron micrograph of the stridulatory organ found between abdominal segments of a Maculinea alcon pupa . Winfrith Technology Centre. However. Fig. 84-87 Acoustical interactions between Maculinea alcon and its host ant Karsten Schönrogge 1. 1. Charles2. 2002 and therein). Marcin Sielezniew4 & Jeremy A. M. Dorchester. Ania Stankiewicz3. Nowoursynowska 159.© 84 PENSOFT Publishers Karsten Schönrogge et al. 02-776 Warszawa.uk 1 It has long been known that ants have the ability to produce sounds by stridulation although the function of such calls and how they are perceived. Wardlaw1.A. Poland Contact: ksc@ceh. Judith C.ac. whether as air waves or substrate vibration. SGGW .Centre for Ecology & Hydrology. Le Mesnil Amand 50450. Settele. E. E. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. is still largely unknown and might well vary between species (Hölldobler and Wilson 1990). Normandy. 00-679 Warszawa.Warsaw Agriculture University. Thomas1 NERC . Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. There is some debate whether stridulations within the family of Lycaenidae is universal or whether some non-ant-attended species might be mute (Pierce et al. Dorset DT2 8ZD. Topham1. France 3 Museum and Institute of Zoology. were highly variable and an analysis of whether the Maculinea calls were more similar to that of their respective host ant species showed no discernible pattern. scabrinodis (Devries 1991. pupae. Here we recorded caterpillars and pupae of Maculinea alcon together with the larvae.Acoustical interactions between Maculinea alcon and its host ant 85 The caterpillars of a wide range of riodinid and lycaenid caterpillars were recorded. Ant brood never emitted any sounds during our recordings and also the small number of queens available appeared to be mute. the methods used in that study involved immobilising adult ants above the microphone. including all five European Maculinea species. and it appears likely that the caterpillar calls may be able to elicit behavioural responses in the ants (DeVries et al. 2. Housed inside its sound dampening cover. DeVries et al. Our digital recording device is a bench top set up with noise reduction electronics (Fig. 2). Recording equipment includes A. B. However. The equipment records digitally into a computer and is powered by a 12V gel-cell to reduce electrical interference. etc. under ambient light. a second microphone with C sound subtraction electronics. M. pulse rates. the recording microphone with nest chamber. they are not immobilised. dominant frequencies. Also. and were highly stressed during the recordings. 1993). . and both the ants and caterpillars were at room temperature. sabuleti and M. ruginodis. alcon were much closer than in the original study. recordings are carried out in the dark. and the calls of the latter were compared to the calls of the worker ants of Myrmica rubra. and while the movement of adult ants is restricted during recordings. 1993). although call frequencies were not matched very closely. DeVries et al. the ant and caterpillar calls encompassed a similar range from a few hundred Hz to a few kHz. Our results were largely consistent with those of DeVries et al. other characteristics such as the fundamental frequencies. M. and therefore closely resembled those of the worker ants. Pupae also emitted stridulations that were very similar to those of caterpillars. Fig. (1993) concluded that while the general “shape” of the Maculinea calls producing sets of pulses was strikingly similar to that of adult Myrmica ants and different from calls from other non-parasitic lycaenid species. adult workers and queens of their host ant Myrmica scabrinodis. (1993) although the frequency ranges of host ant species and caterpillars of M. Of the five behaviours. alcon and those of M. workers approached the speakers that were broadcasting Maculinea alcon or worker ant sounds significantly more often and antennated the speaker more often in comparison to the white noise. To our knowledge these are the first behavioural trials involving Maculinea stridulations. rebeli. which appears from this preliminary study to be closer than DeVries et al (1993) originally thought. pupal-stridulations and d. the variation within each species’ call is substantial and the authors themselves suggest that some of it might be attributable to the immobilisation of the ants during recording (DeVries et al. It still appears reasonable to interpret the stridulations as a “call for attention”. alcon caterpillars by the ants had no significant effect on the results. If this assay supports a role for acoustic cues in the Maculinea – ant interaction. future studies will aim to establish their role in determining host specificity. Also.86 Karsten Schönrogge et al. In addition to the analysis of the recordings we established a behavioural assay where the sounds were played back to worker ants using computer generated white noise as a control. while six colonies were naïve and had no previous contact. antennating the speaker. for instance. traversing the speaker. approaching the speaker. teleius are more than 1kHz higher in frequency (DeVries et al. with four arenas attached to play a. There was no significant difference in repulsion between the different noise sources. caterpillar-. that the dominant frequency of M. workers of 12 colonies were used in direct observations (30 minutes each). Also. white noise as a control. and being repelled (approach and quickly run away). It is intriguing. nausithous and M. is about 130Hz higher than that of M. alcon caterpillars previously and were termed experienced. six of the colonies had reared M. 1993). previous experience of M. Play back device with 6 modified MP3 players. but that chemical cues such as cuticular hydrocarbons might be more important on contact. which allows playing 6 tracks to 12 arenas simultaneously. however. 3. In total. b. c. . While the mean dominant frequencies found in that study for the Myrmica species did not vary that widely. Each trial was replicated four times and analyses were carried out on the average responses. Fig. ant worker-. scoring each of five behaviours ranging from resting on the speaker. F. Heath. E. A. Rand. LITERATURE Devries. Hölldobler. and thus to study the relative importance of sound characters such as pulse repetition frequencies. Wilson.. amplitudes.Acoustical interactions between Maculinea alcon and its host ant 87 1993). functional. Springer Verlag. Call production by myrmecophilous riodinid and lycaenid butterfly caterpillars (Lepidoptera): morphological. . 1993.. 2002. P. B. N. it will be possible to generate sound artificially that varies only in particular variables. acoustical. Travassos. D. 1991. R. Annual Review of Entomology 47:733-771. M. P. Braby. Novitatis 3025:1 . J. Mathew. B. J. 1990. and M. and Thomas. duty cycles and of course their frequencies. Research has partly been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Comparison of Acoustical Signals in Maculinea Butterfly Caterpillars and Their Obligate Host Myrmica Ants. Cocroft. and E. O. Berlin. The ecology and evolution of ant association in the Lycaenidae (Lepidoptera). Pierce. J. Biological Journal of the Linnean Society 49:229-238. Lohman. By further elucidating the acoustic characteristics of the calls of both Maculinea and Myrmica species. The ants. D. and evolutionary patterns.23. J. DeVries. A. J. B..A. Poland UKBH . teleius M. nausithous and M. David R. Gronostajowa 7. DK-2100 Copenhagen Ø. Department of Population Biology. Settele.© 88 PENSOFT Sliwinska et al. teleius. In this stage. 1. nausithous and M. E. M. Institute of Biology. nausithous and M. As for M. 1). there are significant morphological similarities between the species in their larval and pupal stages. Institute of Environmental Sciences. nausithous M. Piotr Nowicki1. its pili are transparent and thicker.A. alcon caterpillars. alcon are similar to those of M. but straight and thicker. which should make assigning them to the species simple and accurate. The eminent interspecific differences include the appearance and distribution of hairs (pili and setulae) covering the body surface of caterpillars. Universitetsparken 15. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. including host ant specificity. teleius.University of Copenhagen.Jagiellonian University. M. M. alcon Ewa Sliwinska1. Our aim was to present characteristic features of the caterpillars of the three butterflies. 30-387 Kraków. and those on the head and above legs are considerably (3 to 5 times) longer. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Centre for Social Evolution. nausithous. teleius. Ewa Publishers Sofia – Moscow J. teleius has short. Most of the above differences are most pronouced in the third instar. The third (above) and fourth (bottom) instars of M. alcon Fig. teleius. dark and slightly bent pili. 88-89 The key to the caterpillars and pupae of M.pl 1 2 Three species of myrmecophilous Maculinea butterflies. Denmark Contact: ewa-sliwinska@wp. The pili of M. often occur sympatrically and therefore the correct identification of the larvae and pupae is crucial for studying various aspects of their ecology. M. genetic research and conservation biology. Nash2 & Michal Woyciechowski1 UJAG . caterpillars can be easily distinguished by the length and colour of the pili (Fig. . M. which are uniform over the entire body surface. alcon. M. Kühn & J. However. pp. alcon 89 M. shiny cuticle. teleius and M. The pupa of M. Considering between-instar differences within species. and remains this way until pupation. 2). alcon In the fourth (final) instar M. M. teleius is immediately recognizable due to the presence of characteristic long and thick setulae covering the top side of its thorax. but large and wide in M. The pupae morphology of M. nausithous. 2.The key to the caterpillars and pupae of M. nausithous and M. teleius. nausithous and M. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). nausithous Fig. teleius. with a transparent. fourth instars can be identified by the fold of cuticula coat (so-called ‘hump’). M. dull cuticle (Fig. The caterpillar in initially cylindrical in general shape. alcon is more slender. The developing pupa of this species appears yellowish brown. Apart from this. with clearly visible and distinct body segments. The other two species lack these structrures. with a thick. Both are stout. teleius. The pupae of M. M. but just before eclosion the wing pattern and colour becomes clearly visible through the transparent cuticle. The fourth instar of M. which completely covers the joint section between head and thorax as well as a large part of hemisphaera. exudatorium is small and narrow in M. teleius and M. . The cuticle of this species is particularly smooth and shiny. alcon is characterised by transparent setae occurring across the surface of the body. and this becomes much more noticeable as the caterpillar develops in the ant nest. nausithous cannot readily be distinguished from each other. alcon. J. isolation and the short life period of adults. The species occurs in three regions. 2003). 2003). pneumonanthe separated by Phragmition. It has been found in 35 10 km grid squares so far and its national status is assessed as ‘vulnerable’ (Buszko & Nowacki. On neutral or alkaline grounds M. is situated in the vicinity of the town and extends over the south and west hill slopes.l. supporting the strongest population. (500 m a. Settele. s.).. probably on the verge of extinction in spite of an abundance of food plant.5-2 ha. 90-93 Maculinea alcon and M. Thomas © 90 PENSOFT Publishers Anna M. Hence some others undoubtedly still await discovery. It is still likely that the species is more widespread in the south-east and some sites are unknown because of the small area. rebeli is the rarest species of Maculinea in Poland. alcon is most often encountered in Molinion meadows or their degenerated forms such as Nardus grassland near Warsaw. and Beskid Niski Mts. In mid 1990 another very large population with at least 1000 butterflies on the wing in one favourable season was found in the vicinity of Przemyśl (270-320 m a. (Buszko. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. 00-679 Warszawa. habitats. estimated as ‘endangered’ (Buszko & Nowacki. Kühn & of Butterflies(Eds) 2005 Studies the Ecology and Conservation in Europe Sofia – Moscow Vol. 1997). Wilcza 64. pp. Gagarina 9. In spite of the species’ rarity. In 2005 only about two thousands eggs were counted. Now it is rare in this locality. Marcin Sielezniew2 & Jarosław Buszko3 SGGW .s. host ant specificity and parasitoids Anna M. Magnocaricion elatae. On acid soils it occurs in Nardo-Callunetea types of vegetation including heathlands and grassy meadows. 2002). Metapopulations inhabiting calcareous fens in the Polesie region occur in patches of Molinion medioeuropaeum with G.pl 1 The sites of Maculinea alcon are scattered across southern and eastern Poland but the species occurs also in isolated localities in Biebrza Basin and Wielkopolska.A.waw. Poland. E. quite a wide range of habitats is used in Poland. Salicetum pentandro-cinereae and Carici elongate-Alnetum. Department of Applied Entomology. Department of Animal Ecology. Most of the sites are very small and cover an area 0. all in southern and south-eastern Poland. Contact: ams@miiz.) support just a few dozen specimens (Sielezniew et al.Warsaw Agriculture University. l. Marcin Sielezniew & JarosławonBuszkoJ. Stankiewicz1. The site near Przemyśl. Poland 3 Nicolas Copernicus University. For the first time it was recorded in the Pieniny Mts. Institute of Ecology and Environmental Protection. rebeli in Poland: distribution.. Three small sites discovered in 2002 in the Bieszczady Mts. Gentiana cruciata covers a somewhat wider . Polish Academy of Sciences. The strongest populations were encountered in the Polesie region and in the Holy Cross Mts. especially when we consider the fact that the main larval food plant Gentiana pneumonanthe is much more widespread. 2002).) which is the core spot of this butterfly in Poland (Sielezniew et al. M. Nowoursynowska 159. (Świętokrzyskie Mts. 02-776 Warszawa. Poland 2 Museum and Institute of Zoology. 87-100 Toruń. Stankiewicz. everywhere in Poland. rebeli ( ) in Poland . comm. The present distribution of Maculinea alcon (•) and M. In most seasons butterflies are on the wing for about 3 weeks from mid June until early July. Als et al. alcon develops.s.g. The first adults emerge in the warmest patches of the site. forest fringe community Origanetalia and fresh meadow (Arrhenatheretum elatioris).) and one site near Komańcza. Only G. host ant specificity 91 area of calcium rich soils than that of the butterfly population. Tartally & Csősz. Females lay eggs mainly on G. as was also noted in France. cruciata on xerothermic meadows surrounded by calcareous fens where both plant species are sympatric (Sielezniew & Stankiewicz. 2003). G... Gentianella amarella were also recorded from eastern Poland (W. in M. alcon are on the wing usually from early July until late August although considerable variation between sites and years was observed. making a transitional type of habitat between dry grassland Festucetalia valesiaceae.Maculinea alcon and M. One of them consists of the narrow edge of a ground road near a stream (with some signs of grazing). The flight period of M. Adults of M. pneumonanthae but sometimes other gentians are also used as secondary larval food plants e. southern Netherlands and Hungary (Elmes et al. 2004a). Michalczuk pers. rebeli is the shortest of all Maculinea species in Poland. M. Gentianella ciliata. Spain. The second includes a flat. 2002. Warm springs accelerate emergence and the first males may be observed in the first days of June. The vegetation is classified as xerothermic grasslands Festucetalia valesiaceae mixed with Origanetalia and light scrub Prunetalia. scabrinodis nests.l. 2004).e. south-facing slopes with very short turf.. scabrinodis is the most common ant species in patches with larval food Fig. The two remaining localities are very different. i. However in a cold summer the last adults may fly until the end of July. cruciata has been recorded as a larval food plant. M.). Similar topography and vegetation characterize the site in Pieniny Mts. 1994. habitats. On most sites. 1. elevated area adjacent to the road covered by plants of Arrhenatheretum as well as ruderal plant species with traces of cultivation (Sielezniew et al. rebeli in Poland: distribution. (500-600 m a. .. A. scabrinodis. G. M. rebeli with M.. 2005). 2005b). rebeli being a distinct species (Als et al. Mignault.. Manguira. is almost identical chemically. J. Thomas. rugulosa nest in the small population near Komańcza. The studies carried out near Przemyśl showed that the most important host ant of M. . Z.. 2004). 693–696. & Nowacki. alcon seem to be most similar in respect of host-ant specificity to the Hungarian ones (Tartally & Csősz. Only 3 pupae from one among hundreds of examined nests were infested. 55-68. Elmes. Spain (Thomas et al. A. Despite the lack of morphological traits. 3P04G02624). Hence we may assume that there is probably no geographic variation in host ant specificity in Poland. (1994) Differences in host-ant specificity between Spanish. Vila. Boomsma. Sielezniew. 403-414. scabrinodis was completely absent (Stankiewicz et al. 2002). M. Nota lepidopterologica 27.D. & Boomsma. Toruń. N. habitat and caterpillar growth rate are enough to consider both groups of populations as separate conservational units in Poland. (2002) Geographical variation in host-ant specificity of the parasitic butterfly Maculinea alcon in Denmark. Kandul. J...A. J.D.. M. Ecological Entomology 27. European Journal of Entomology 101. D. L. 386-390. butterfly larvae and pupae were also found in M. 2004b). rebeli studies in Poland were supported by the State Committee for Scientific Research (grant no.R. In Red list of threatened animals in Poland (ed. (2002) Lepidoptera. Dutch and Swedish populations of the endangered butterfly Maculinea alcon (Denis et Schiff. 80-87. R.. rebeli is M. while M. recorded in less than 20% of sites in the Alps (Thomas et al. Unidentified wasps of the genus Ichneumon were reared from pupae collected at the same sites. The parasitation rate is very high. It is still unknown if parasitoids reared from ‘cuckoo’ Maculinea in Poland are the same species as Ichneumon eumerus.J. Stankiewicz. It was usual for half of the nests of this species under gentians to be infested. 1989) and Lithuania (Stankiewicz et al. Polish populations of both M. T. However. Memorabilia Zoologica 48. REFERENCES Als.. Sielezniew. vandeli is thought to be a temporary social parasite of M.92 Anna M. pp. M. Hsu. (2004a) Gentiana cruciata as an additional host plant of Maculinea alcon on a site in Eastern Poland.R. and proved to be an equally effective host (Sielezniew & Stankiewicz. Nature 432..W. Nash. Głowaciński). D.. J. One infested pupa was also noticed in the Polesie region. J. exceeding 50 %. Turpress. & Stankiewicz. There is also a single record of the presence of M.) (Lepidoptera). Marcin Sielezniew & Jarosław Buszko plant. the parasitization rate seems to be very low.E. sabuleti. was found in 2004 in the Przemyśl population. scabrinodis colonies were also exploited but the parasitization rate was considerably lower. Buszko. alcon and M. sabuleti was very rare and M. M. the differences in phenology. Als. T. [in Polish]. & Pierce.. schencki. Yen. which is absent near Przemyśl dominated the turf. 2004b). We have never observed M.. morphologically very similar to the one reared from M. 91-93. Instytut Ochrony Przyrody PAN. O. (2004b) Simultaneous exploitation of Myrmica vandeli and M. Kraków. rebeli pupae in a M. Buszko.J. (1997) A distribution atlas of butterflies in Poland 1986-1995. M. The latest genetic studies have challenged the idea of M. That ant species. N. Hammarstedt. 2004). A. rebeli and M. the main host in France. alcon. Martin. vandeli colonies – an ant species that is absent at other sites (Sielezniew & Stankiewicz. J. & Stankiewicz. J.G.. scabrinodis (Hymenoptera: Formicidae) colonies by the endangered myrmecophilous butterfly Maculinea alcon (Lepidoptera: Lycaenidae).. In the Świętokrzyskie region in southern Poland. (2004) The evolution of alternative parasitic life histories in large blue butterflies. Nash. The unidentified Ichneumon parasitoid. & van der Made J. Oecologia 79. 425-457. Wakamura S.W. (1989) Host specificity among Maculinea butterflies in Myrmica ant nests. & Elmes G.J.A. 51-54..Maculinea alcon and M. 1904 (Lepidoptera: Lycaenidae). M. Polish Journal of Entomology 74. rebeli in Poland: distribution. S.. (2005a) The first record of Myrmica rugulosa Nylander. M.C..M. Knapp J. Thomas. Tartally. A. 309-317. J. & Górnicki. Sielezniew. M... Nature 417.W. A. Sielezniew. A. & Woyciechowski. 99-103. Przeglad Zoologiczny 47. Elmes.C. Ł. Simcox D. & Barański. G. Myrmecologische Nachrichten 7. Akino T. Stankiewicz. Wardlaw. (2003) On the distribution and the ecology of Mountain Alcon Blue. Természetvédelmi Közlemények 11. M. (2005b) Myrmica schencki rears Maculinea rebeli in Lithuania: new evidence for geographical variation of host-ant specificity of an endangered butterfly. A... & Švitra. 2002: Parasitoid secretions provoke ant warfare. A. G.. 211-220. . 505 – 506.M.J. habitats. host ant specificity 93 Sielezniew.. & Csősz. [in Polish] Stankiewicz.A. Łuczaj. M. Maculinea rebeli Hirschke (Lepidoptera: Lycaenidae) in Poland. Wardlaw J.. Gerty S. (2004) Data on the ant hosts of the Maculinea butterflies (Lepidoptera: Lycaenidae) of Hungary. 1849 (Hymenoptera: Formicidae) as a host-ant of Maculinea rebeli Hirschke. J. Stankiewicz. [in Hungarian] Thomas J. apart from the potential confusion caused by occasional nest switches by different ant species during the pupal stage. (Lepidoptera: Lycaenidae) are obligate parasites of Myrmica (Hymenoptera: Formicidae) colonies in Europe during most of their development. The latter was important because there were some Myrmica . gallienii. Tartally 2005a. Faculty of Sciences. pers.© 94 PENSOFT Publishers & Zoltán Varga András Tartally Sofia – Moscow J. Maculinea rebeli specimens were mostly found in Myrmica scabrinodis. M. salina nest. S. see also: Tartally & Csősz 2004. specioides nest. Department of Evolutionary Zoology and Human Biology.) situated within 2 m of Maculinea initial host plants (Tab. salina as hosts (see: Als et al. E. salina. These are the first records of Maculinea alcon using Myrmica salina. vandeli nests. Hungary 2 HAS-DU Evolutionary Genetics and Conservation Biology Research Group. Host-ant specificity may vary between regions (Elmes et al. rubra and once in a M. Knowledge of the host ant species has been shown to be crucial for the protection of these endangered butterflies (Thomas et al. Munguira & Martin 1999).B. it reveals which ant colonies reared Maculinea larvae through to nearadulthood rather than which colonies retrieved young IV instar larvae but later killed them (Thomas et al. pp. 2004: Supplementary Tab. Myrmica rubra was the only host ant of Maculinea nausithous. specioides and once in a M. 141 infected nests contained a total of 890 Maculinea specimens (larvae. 1994). This work was conducted to obtain data on the host ants of Maculinea butterflies in Hungary by two methods between 2000 and 2005: (1) Field surveys were made just before the flying periods by examining Myrmica spp. pupae and exuvia). rebeli caterpillars in artificial Myrmica nests. Maculinea teleius specimens were mainly found in Myrmica scabrinodis but often in M. lonae. therefore data should be collected over the geographical range of a butterfly species’ distribution. Kühn & J. arion ligurica specimens in ant nests. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 3. salina and M.A. P. Settele. and of Maculinea teleius using Myrmica specioides and M. connections with parasitoids and host plants András Tartally1 & Zoltán Varga1. nests (determination according to: Seifert 1988. Csősz. comm. 1989.O. We could not find Maculinea arion arion and M. 2005).University of Debrecen. Hungary Contact: tartally@delfin. sabuleti. 10).hu Larvae of Maculinea spp. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.O. In 23 sites. P. (2) Laboratory examinations were done by testing the adoption and survival of Maculinea alcon and M. M. of Maculinea rebeli using Myrmica lonae and M.2 1 DUFS . salina. H-4010 Debrecen. schencki and sometimes in M. M. 94-98 Host-ant specificity of Maculinea species in Hungary. Tartally & Varga 2005). 1. M. H-4010 Debrecen. Our aim was to confirm our field records and find new potential host ant species.B. This is the recommended time of year to investigate host specificity in the field because. Maculinea alcon specimens were found predominantly in Myrmica scabrinodis and sometimes also in (its close relatives) M. M.unideb. 3. A similarly accelerated development of M. specioides M. lobicornis M. up to one and a half months when we introduced them to artificial Manica rubida (Hymenoptera: Formicidae) nests (Tartally 2004). Potential and recorded Myrmica host ants of Maculinea spp. lonae + — + — — + + + — — — + — — — + — — + — The different symbols show that the Myrmica sp. alcon were completed with Myrmica gallienii and M. not present in any of the sites (—). So. nausithous M. Furthermore the potential host ant species of M. During these laboratory observations we found a surprising phenomenon as seven specimens of M. species whose density was low at the examined Maculinea sites and there was little chance of finding nests of these ants near Maculinea host plants. caterpillars was at least 5 in average in the infected nests ( ). it would be desirable to confirm this result also by field data. specioides nests. present and used in less than 50% of the sites. scabrinodis and M. Myrmica rubra M. However. teleius M. we can not say that these Maculinea alcon caterpillars would be able to finish their development at the artificial Myrmica gallienii and M. the number of Maculinea sp.g. M. scabrinodis M. rebeli despite of their overlapping morphological characters (e. present and used in at least 50% of the sites. In the case of Maculinea rebeli we also found a potential host ant species which was not recorded during our fieldwork: six out of ten introduced M. the number of Maculinea sp. However. rebeli caterpillars successfully pupated in a Myrmica lobicornis nest (which is a rare ant in Hungary). ruginodis 95 M. caterpillars was less than 5 in average in the infected nests ( ). caterpillars was at least 5 in average in the infected nests ( ). 1989). However. present and used in less than 50% of the sites. rebeli. Beuret 1954). alcon larvae was never observed (Tartally 2005b). these artificial colonies and the caterpillars died of an infection during the winter. 2004).Host-ants. the number of Maculinea sp. alcon and M. Thomas et al. was: present minimum in one site but was not used (+). arion arion M. parasitoids and host plants of Maculinea spp. schencki Maculinea alcon M. sabuleti and M. in Hungary. The laboratory observations confirmed our field results about the host ants of M. Of course. salina M. rebeli M. the recent literature showed that the genet- . sabuleti M. arion ligurica — — — + + + + + — — — — — + + + — — — — — — — + — — — + — — — M. vandeli M. specioides because the M. present and used in at least 50% of the sites. and we should also note that Myrmica nests kept in captivity are generally more tolerant of any species of Maculinea parasite in thier midst than are wild nests subject to the stress and depravations of competition in the field (Elmes et al. by their different host ant and initial host plant species (e. caterpillars was less than 5 in average in the infected nests ( ). salina nests.g. in Hungary Table 1. the number of Maculinea sp. gallienii M. The knowledge of the host ant specificity is also of interest from evolutionary and taxonomic point of view because the earlier literature separated Maculinea alcon and M. Another interesting observation was that several caterpillars of Maculinea alcon and M. salina colonies. rebeli survived and increased in size for a number of weeks. rebeli pupated after only about a month in artificial Myrmica scabrinodis. this observation also needs to be confirmed by field data. alcon caterpillars grew similarly well in the colonies of these two Myrmica species as they did in M. teleius pupae. pneumonanthe which is a traditional host plant of the M. 2004. 2005). 2004. rebeli larvae were found in the basins at all the three (Gentiana cruciata. alcon (see also: Sielezniew & Stankiewicz 2004). scabrinodis and M.96 András Tartally & Zoltán Varga ic and morphologic differentiation between these two traditionally separated species is rather low (Als & al. specimens from M. 1). Furthermore. There was another interesting observation at another site within the Bükk Mountains where the M. Pech et al. Munguira & Martin (1999) did not mention the latter two as M. For this purpose. G. we also found a M. rebeli population in the Bükk-Plateau (N-Hungary) which used mainly the traditional M. M. During the field observations we recorded several Maculinea pupae infected by ichneumon wasps (Hymenoptera: Ichneumonidae). salina) which were used by both Maculinea rebeli and M. 1. alcon. Bereczki et al. hoverflies (Diptera: Syrphidae) were often found in the investigated Myrmica nests. Fourth instar M. A Neotypus melanocephalus wasp was bred from a Maculinea teleius pupa (Tartally 2005c) and unidentified Ichneumon sp. The success of these three plant species for M. cruciata but additionally used the Gentianella austriaca for host plant (Fig. Moreover. rebeli host plants. Maculinea rebeli eggs on a Gentianella austriaca plant at Bükkszentkereszt (photo by T. rebeli host plant (Gentiana cruciata) but additionally used the G. rebeli mainly used G. pneumonanthe and Gentianella austriaca) plant species. Kapás) . larvae and puparia of unidentified Microdon sp. These Fig. Here we found two Myrmica species (M. rebeli and also from M. rebeli host was tested. plants with eggs of the butterfly were collected and were kept in glasses of water placed in plastic basins. alcon in Hungary. R. N. A. Pech.. alcon and M. Vila.. M. in Hungary 97 social parasite larvae and their puparia were recorded from several Maculinea alcon. Y. G. D. S. Mignault. Pecsenye. Beuret. K. J....A.. Hammarstedt. Elmes.W. 2004: Gentiana cruciata as an additional host plant of Maculinea alcon on a site in eastern Poland (Lycaenidae) – Nota Lepidopterologica 27: 91-93. 27 (2004): 303-308. 6: 23-27. parasitoids and host plants of Maculinea spp. 1988: A taxonomic revision of the Myrmica species of Europe. Kuehn and J. Entomologia Experimentalis et Applicata 110: 53-63.J. 2005a: Myrmica salina (Hymenoptera: Formicidae) as a host of Maculinea alcon (Lepidoptera: Lycaenidae). Strasbourg. A. Tartally. S. Kandul.W. 2004: Is Manica rubida (Hymenoptera: Formicidae) a potential host of the Maculinea alcon (Lepidoptera: Lycaenidae) group? – Myrmecologische Nachrichten...Host-ants. Yen. Sielezniew. Tartally.A. & Zravý.-F.-H. rebeli seems to be lower in Hungary than it is known from Western Europe. (Eds. 204-222. Seifert. the variability of the host ant. A. – Nota Lepidopterologica. sabuleti nest.E. Konvička. Dutch and Swedish populations of the endangered butterfly. Wardlaw. – Sociobiology 46: 39-43. & van der Made. Bereczki. host plant and parasitoid species of the Hungarian Maculinea populations is usually higher than it is known from Western Europe. Z.. & Martin. J. J. Tóth. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT-2001-00126). Z. – Journal of Zoological Systematics and Evolutionary Research. M. 43: 157-169... Nash. ACKNOWLEDGEMENTS To: E. Hsu. A. T.. – Nota Lepidopterologica.. teleius and M. 2004: The evolution of alternative parasitic life histories in Large Blue butterflies. 97. & Stankiewicz A. Munguira. 2004: Phylogeny of Maculinea blues (Lepidoptera: Lycaenidae) based on morphological and ecological characters: evolution of parasitic myrmecophily. Formicidae).. pp. – Memorabilia Zoologica 48: 55-68. 1994: Differences in host-ant specificity between Spanish. Teil II.. Tartally. G. Thomas. These facts reflect the value of these populations and the importance of their protection and examination. Settele. – Cladistics 20: 362-375. As our results show the ecological difference between M. Martin. A. – Nature 432: 386-390. M.M. R. J. L. J. Asia Minor and Caucasica (Hymenoptera. M. M. rebeli. 28: 65-67. B. Munguira. J.P. O. K. REFERENCES Als. N. Schnrögge. J..A.) 1999: Action Plan for the Maculinea butterflies in Europe. – Nature and Environment.L. Boomsma. nausithous sites in Myrmica scabrinodis but in some cases in M. 2005: Pattern of genetic differentiation in the Maculinea alcon species group (Lepidoptera: Lycaenidae) in Central Europe. & Thomas. & Varga. 2005b: Accelerated development of Maculinea rebeli larvae under artificial conditions (Lycaenidae). No. Tartally.D. E. . J. gallienii and in one case in a M. H. & Pierce. Council of Europe Publishing. Peregovits. At the same time.C. Csősz. M. P. J. 2004: Food stress causes differential survival of socially parasitic larvae of Maculinea rebeli (Lepidoptera: Lycaenidae) integrated in colonies of host and non-host Myrmica species (Hymenoptera. 2005c: Neotypus melanocephalus (Hymenoptera: Ichneumonidae): the first record of a parasitoid wasp attacking Maculinea teleius (Lycaenidae). Maculinea alcon (Denis et Schiff.L. 64 pp. – Abhandlungen und Berichte des Naturkundemuseums Görlitz 62: 1-75.) (Lepidoptera). – Basel (Entomologische Gesellschaft).. Fric. 1954: Die Lycaeniden der Schweiz. Formicidae). Elmes. Tartally. M.. K.. Schönrogge. Holloway.98 András Tartally & Zoltán Varga Tartally. Wardlaw. G. J. G.W.A. & Woyciechowski. – Myrmecologische Nachrichten. S.W.. 7: 55-59 Thomas. & Elmes.A. J. (Data on the ant hosts of the Maculinea butterflies (Lepidoptera: Lycaenidae) of Hungary. A.) – Természetvédelmi Közlemények 11: 309-317. J. Elmes. 2005: Specialisations and host associations of social parasites of ants. Fellowes. & Varga. Thomas. Z. & Csõsz. A. pp 479-518. J. 1989: Host specificity among Maculinea butterflies in Myrmica ant nests.. In Evolutionary Ecology Eds Rolff. G. . – Oecologia 79: 452-457. 2005: Myrmica rubra (Hymenoptera: Formicidae): the first data on host-ant specificity of Maculinea nausithous (Lepidoptera: Lycaenidae) in Hungary. 2004: Adatok a Maculinea boglárkalepkék (Lepidoptera: Lycaenidae) kárpát-medencei hangyagazdáiról. Symposia of the Royal Entomological Society XXI.C. M.. it had been believed that Maculinea were generalist parasites of any Myrmica species and perhaps also of Lasius flavus (Ford 1945). Winfrith Technology Centre. at least among the predacious Maculinea (Als et al 2004). Sampling more widely across Europe soon revealed two regional host switches in M. UK. Simcox1 & Josef Settele2 NERC . Thomas1. as demonstrated by several papers in this volume. nausithous mid-way between these two groups (Thomas & Elmes 1998). ruginodis. Hitherto. Considerable advances have been made by the collaborating partners of MacMan. CEH Dorset. The discovery that the currently recognised morpho-species may consist of several cryptic species.Centre for Environmental Research Leipzig-Halle. and that (where known) these different forms are associated with host switches (Thomas & Settele 2004). makes clarification of host specificity across Europe (and beyond) even more urgent. Kühn & of Butterflies(Eds) 2005 ‘non-hosts’: J. but must co-exist with an adequate density of the single host species to which each butterfly is adapted (Thomas et al 2005). These results reveal a much more complex pattern of host specificity across Europe than was once predicted. Karsten Schönrogge1.Centre for Ecology & Hydrology. Theodor-Lieser-Str. Dorset DT2 8ZD. secondary hosts andStudies on the EcologySettele. some in the Pays Bas and Denmark exploiting M. rubra (Elmes et al 1994). In practice.© PENSOFT Publishers Primary Sofia – Moscow J. including the apparent existence of multiple . made in west Europe. pp. alcon. and northern populations using M. Department of Community Ecology. teleius and M. Thomas hosts. 06120 Halle. whereas M. David J. suggested a simple pattern with each of the five recognised species of Maculinea exploiting a single but different host Myrmica species (Thomas et al 1989). scabrinodis. Dorchester.common confusions in Europe 99 and Conservation Vol. and this means that successful conservation requires targeted management for the optimum habitat of the essential ant species (Thomas et al 1998). Germany Contact: jat@ceh. This result indicated that successful conservation required that each species’ early larval foodplant must flower not simply near any of the 3-6 species of Myrmica that typically inhabit its comparatively broad niche within grassland. 2 UFZ .uk 1 An important advance in the conservation of endangered species of Maculinea butterfly in Europe was the discovery that species or populations were host-specific to a single species of Myrmica ant (Thomas 1980).ac. Our first studies of host specificity. Graham W. 99-104 Primary hosts. sites supporting persistent populations of Maculinea generally have higher densities of host ant than the minimum required. secondary hosts and ‘non-hosts’: common confusions in the interpretation of host specificity in Maculinea butterflies and other social parasites of ants Jeremy A. Models and empirical studies suggest that the mimimum density of host ant required by a cuckoo species of Maculinea is about 10% co-occurrence of the Maculinea population on (the preferred growth form of) flowering Gentiana and host ant. Elmes1. with M. 4.A. arion require >50% co-occurrence. E. with southern populations exploiting M. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. if laboratory ants are stressed to simulate more natural conditions (which is hard to achieve in captivity without the whole culture dying). Schönrogge et al 2004). especially where based on small samples or lab studies. ruginoids. Here we build upon earlier assessments (Thomas et al 1989.more than one host on the same site (Thomas et al 1989. Elmes et al 1991). each is predominantly exploiting a different morph or cryptic speices of ‘M. Further research is urgently required into Myrmica taxonomy. various papers in this volume). ii) ‘Adoption’ experiments. then host specificity quickly becomes apparent. Even within Europe. Thus. particularly within the morpho-species M. and M. M. although in both cases survival may be slightly higher with the natural host or kin ant colony (e. MacMan scientists are advised to retain set or frozen ant samples for possible re-assessment in the future. Myrmica rubra. and arose from their inability (later acknowledged) to identify ants of the genus Myrmica from Tetramorium and even Lasius niger (Thomas et al 1989). This is because. Added to this. because well-nourished ants kept under lab conditions typically behave quite differently from colonies in the wild. just as almost any species of Myrmica will successfully rear the larvae of any other Myrmica species (Elmes unpublished). a true measure of retrieval of larvae into ant nests in the wild. we list a few pitfalls that can mislead or can bias the data. certain Myrmica species are notoriously difficult to distinguish (especially as workers) and it is only 30 years since M. Thomas & Wardlaw 1992. because of various situations or artefacts that can cause confusion. Some of these results are detailed and clear-cut. It is well known that almost any species of Maculinea can be reared with any species of Myrmica under benign lab conditions. scabrinodis. Laboratory results should always be interpreted with caution. when considerable time . Modern field reports are much more reliable: even so. Before considering our interpretation of genuine multiple-exploitation. but instead use them as food for their own brood (Elmes et al 2004. MULTIPLE HOST USE ON SINGLE SITES It is undoubtedly true that certain Maculinea species or populations genuinely exploit – and that others appear to exploit . laevinodis were inter-changeable at various times in the first half of the 20th century. the Myrmica workers partly treat their ‘foraging areas’ as extensions of the brood chambers. sabuleti was unequivocally recognised as a good species rather than as a variety of M. hosts in some species or locations. Artefacts from laboratory studies i) Rearing experiments. Simulating ‘adoption’ by placing Maculinea larvae in the foraging chamber of a captive nest is not. because starving or stressed worker ants with hungry brood no longer tolerate imperfect mimics in their nests. Thomas et al. an ever wider range of sibling species is being described. which we suspect encompasses several more good but ‘cryptic’ species (this volume). in our experience.g. Thomas et al 2005) and list some of the more common pitfalls that should be considered when interpreting host specificity. ANT IDENTIFICATION The main mistakes which became rooted in the early literature were made by two principal workers on Maculinea. the names. However. in most (especially small) captive nests. Meanwhile.100 Jeremy A. but others. need careful interpretation before conservation strategies are founded upon them. Elmes et al 1998. scabrinodis. scabrinodis’. It is still perfectly possible that on sites where where Maculinea alcon and Maculinea teleius (and Microdon myrmicae) co-occur. especially with the predacious species. this may well represent the propensity for individual caterpillars to become integrated with host societies after retrieval (or be rejected by non-host ones). alcon). In every study where observations have been made in the wild (although we know of none made yet of M. it is a common feeding strategy to exploit more than one ant colony. This. alcon or M. such as Maculinea. nausithous. we therefore recommended sampling in the late larval or pupal stage (typically May-July). and because Maculinea larvae have relatively simple chemical profiles on leaving their plants (Akino et al 1999 & Macman results). carrying in a fresh supply of grubs (Thomas & Wardlaw 1992).for a few weeks of months in ‘non-host’ nests before eventually dying. Date of sampling in the field Host specificity at the stage of larval retrieval from beneath food-plants is unlikely to be a possibility in the evolution of traits in brood mimics. one further artefact can arise. However. because larvae may persist – and indeed grow . rebeli because cuckoo caterpillars are such effective mimics of ant brood that they act as a focus for their ant colonies even when no ant brood is present. 2002). the first ant species (and generally individual) to encounter the larva is almost always the one that carries it to the nest. secondary hosts and ‘non-hosts’: common confusions 101 differences have been reported in the retrieval of M. but not with M. For MacMan studies. arion has a remarkable ability to fast. M.T. However in their pupal stages. because the workers of any species of Myrmica will generally retrieve the larvae of any other Myrmica species if these are artificially placed into their foraging ranges (Elmes unpublished). we recorded genuine survival from . causing the colony to desert the nest site because it is the ant brood that provides the cohesion and focus to the workers. arion caterpillar will typically draw in two additional ant colonies in this way before pupation. either immediately in the case of M. unpublished) suggest that an individual M. In M. and may live for many days until a neighbouring ant colony buds into the vacated nest. nausithous (Thomas 1984. Our models (R. sampling nests of Myrmica for Maculinea in late summer or autumn often gives misleading results. all Maculinea species are prone to switches occurring in the species of ant occupying their nest site. alcon by different Myrmica species in captivity. rebeli). teleius (Thomas 1984. Clarke et al. clearly. but it does not indicate what proportion of caterpillars are retrieved into different ant nests in the first place. can lead to mistakes in measuring host specificity in the field. and although the majority of these are of the same species. arion and M. 2002). teleius and perhaps M. empirical studies indicate that a change in the species of Myrmica is not unusual (Thomas & Wardlaw 1992). Thus.Primary hosts. rebeli (Elmes et al 1991) and M. As a consequence. we have observed indiscriminate retrieval of Maculinea larvae into the nests of all Myrmica species present on grassland sites (see refs above). Essentially. Distinguishing between host specificity in the retrieval of caterpillars into nests and in their subsequent survival is important because it profoundly affects model predictions of the conditions under which Maculinea populations can persist. or after the classic long behavioural interaction first described by Frohawk in the cases of M. Primary and Secondary hosts: sources or sinks in ‘good’ or ‘bad’ years and sites? In early studies of Maculinea. the caterpillar eats all the grubs in its nest. arion and M. which indicates that a caterpillar has survived at least until pupation in that nest. arion at least. and as such be a sensitive bioassay to foretell future survival. when we followed the survival of several hundred eggs to adults (over several years in the cases of M. The same process is likely to occur with M. and hence were unsuitable for oviposition.sabuleti colonies were attainable. scabrinodis-using populations) and M. in about 20% of emergences of M. it is important first to compare the survival of Maculinea caterpillars in their colonies with that in the colonies of the primary ant host species. secondary hosts are not absolute sinks but may play an important role for population persistence in years with extreme events. . that host specificity is amplified in extreme unfavourable years. it is unclear whether M. sabuleti nests was about five times higher with other Myrmica species. which quite frequently survived with M. this pattern has largely been confirmed by many recent MacMan studies (this volume). Poland) and M. When assessing the value (or harm) of having high densities of ‘secondary’ host Myrmica species present on a site. teleius. lives) can genuinely persist for several years on sites where the majority of a population is not supported by M. arion. and that the vast majority of eggs were laid on Origanum in neighbouring moist-soiled land where M. but there are indications that in central Europe it occasionally may. Myrmica workers are much more tolerant of any species of Maculinea within their nests than when conditions are stressful (Elmes et al 2004. arion is <1. It is also possible that certain sites provide such optimum environments for more than one Myrmica species in most years that a large number of nests of mixed ant species are tolerant. conversely. Regional host switches apart. when really they are partial sinks. for which detailed life-tables of survival exist for six and five years respectively (Thomas et al 1998). a population would rapidly decline to extinction on any site that supported only the secondary host ant species of M. arion. scabrinodis. For example. On present knowledge. alcon (especially the M. and almost never in the case of M. with the important exceptions of apparently multiple-host use in certain populations of M. Thomas et al. rebeli. Despite this. Models and empirical data suggest that a minimum of 55% co-occurrence of primary host (M. rebeli (in Hungary. teleius. For M. sabuleti occurred in Oland Sweden. the most tolerant of all Myrmica hosts. It is easy to imagine that this applies more generally in certain favourable years in the field (and. Elmes et al 1998). This difference is such that the secondary hosts represent patches of misleading pseudo-sources within a site. arion in the UK (Thomas 2002. for example during droughts). arion were reported in 1993). Thomas & Elmes 1998. alcon (north Denmark). in habitats where Myrmica rubra. scabrinodis. Thomas et al 1998. This may seem obvious. this occurred most commonly in M. arion and M. we found that virtually all Thymus larval food-plants failed to flower. in addition to its main host M. arion. Anecdotal observations suggest that there may also be some temporal and spatial variation in survival with ‘secondary’ host species within a population. 2005. sabuleti) and host plant is necessary for ë to exceed one for M. in the areas where M. Schönrogge et al 2004). which are discussed later. larval survival in M. rubra as well as in its more usual host M. the most ‘generalist’ of the Maculinea species (or perhaps that most fortunate to co-exist. rubra dominated. This is consistent with lab experiments that indicate that if food or other conditions are particularly benign. at least enough survived for the population to persist and then recover within a few years once the M. Although the large majority of larvae failed to survive (only two individual adult M. very rarely in the case of M. In other words. where the intrinsic growth rate (ë) of M. nausithous (Thomas et al 1989. 2005). in the drought summer of 1991. rebeli (schenckiusing populations). and in practice most known populations survive only on sites where much higher host ant densities exist (Thomas & Elmes 1998).102 Jeremy A. retrieval to eclosion in a few colonies of ‘non-host’ Myrmica species on some sites (Thomas et al 1998). In general. but on current knowledge is probably possible only for M. scabrinodis. W.. H. Boomsma.W.. T. Simcox. 403-414 Als. Nash. Nash.J. Ecological Entomology 27.. Als et al 2001. scabrinodis. Journal of Zoology 223: 447-460 Elmes. Mignaault. Vila. alcon with M. & Wardlaw.A.A. as elsewhere in Europe do the scabrinodis‘races’ of M. R.. Yen. Clarke. but exploiting different ants. (iv) ‘Mixed-host’ cuckoo populations occur in regions situated near the boundaries where major European host switches exist in both M. (2004) The evolution of alternative parasitic life histories in large blue butterflies. Exactly how these populations function is still unknown. N..E. D. 386-390 Elmes. rubra and M. J. a Large Blue butterfly. rebeli sites in Hungary and Poland (this volume). J.C..C. Thomas et al 2005. N. J. secondary hosts and ‘non-hosts’: common confusions 103 The cuckoo species of Maculinea pose an intriguing question. & Pierce. J. 99-106.T. 2002).. alcon in North Denmark exploit a mixture of M.. J. these species should be more host specific than the predacious species at the scale of a population. D. University of Aarhus Als.. teleius and M. Kandul. but with more generalist males fertilising both types of female. and in many populations this is indeed the observed case (Thomas & Elmes 1998.. R. Alternatively.A. T.C. Animal Behaviour 62.. rubra or M.. D.. Thomas. and Boomsma. J. although the very elegant Macman field studies in Denmark are clarifying the situation (see papers by D R Nash). (2001) Adoption of parasitic Maculinea alcon caterpillars (Lepidoptera: Lycaenidae) by three Myrmica ant species. G. J. (1998) The ecology of Myrmica ants in relation to the conservation of Maculinea butterflies. G. D. J. alcon and M. with each morph adapted to mimic a different host species. They may either consist of hybrids. and Boomsma. not all of which are mutually exclusive: (i) ‘Single-host’ populations inhabit sites where the conditions are so benign for Myrmica that ‘non-host’ ant species will tolerate ill-adapted parasites (as occurs in captive rearing. nausithous do. PhD thesis. apparent multiple host use has been reported from a few M. Hochberg.R. we can hypothesise at least four scenarios. ruginodis) (Als 2001. current theories about infiltration of host societies require major revision. and their Myrmica host ants: wild adoption and behaviour in ant nests. R. A. K & Thomas. Als. R. Journal of Insect Conservation 2: 67-78 Elmes. M.J.T.Primary hosts. Thomas. Possibly. Schönrogge et al 2004).E. exactly in the same way as the closely related (but morphologically distinct) species M. see above) (ii) ‘Mixed-host’ cuckoo populations contain truly generalist individuals pre-adapted to – or which can adapt themselves to – more than one host. rebeli. D. with genetically distinct females functioning like true cryptic species.. Ent Exp et Appl 110: 53-63 . ruginodis colonies (although all seem to be completely incompatible with M. However. D. REFERENCES Als. (1991) Larvae of Maculinea rebeli. In theory. More recently. (2004) Food stress causes differential survival of socially parasitic larvae of Maculinea rebeli (Lepidoptera: Lycaenidae) integrated in colonies of host and non-host Myrmica species (Hymenoptera. D. if they resemble cuckoo birds. Formicidae). Wardlaw. Schõnrogge. this is a maternal trait. (2001) Evolutionary and ecological interactions between Maculinea parasites and their Myrmica host.. it has been recognised for some years that certain populations of M. Nash. If so. (2002) Geographical variation in host-ant specificity of the parasitic butterfly Maculinea alcon in Denmark. Wardlaw. J. For the moment. J. each apparent population might consist of two sympatric populations using the same foodplant. because of the close level of mimicry required for caterpillars to out-compete the ant larvae for worker attention.D. G.W. Nature 432. T. J. perhaps supplemented by immigrants from each region. (iii) ‘Mixed-host’ cuckoo populations contain polymorphic larvae. Clarke.P. (1989) Host specificity among Maculinea butterflies in Myrmica ant nests. Symposia of the Royal Entomological Society 19: 261-290. (2004) Butterfly mimics of ants Nature 432. Journal of Chemical Ecology 30: 91-107 Thomas. Thomas. (1980) Why did the large blue become extinct in Britain? Oryx 15: 243-247.. Everett. E. M.. J. J. Elmes.. S. Peters. (1992) The capacity of a Myrmica ant nest to support a predacious species of Maculinea butterfly. McLean. K. J.C. (2004) Changes in chemical signature and host specificity from larval retrieval to full social integration in the myrmecophilous butterfly Maculinea rebeli. London Thomas. J. Chapman & Hall. & Elmes.A. J.G. & Elmes. J. K..W. G. M. 283-284 Thomas.W. Oecologia 122: 531-537 Thomas.A. Thomas et al.. Ecological Entomology 23: 457-464 Thomas. J. R. CABI. Ed by J. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). Oecologia 91: 101-109. In Insect Evolutionary Ecology pp 475.F.A. Reading Thomas. In Insect population dynamics: in theory and practice. J..B (1945) Buttefrlies Collins.A. Thomas. G.104 Jeremy A. Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). Elmes. Schönrogge. J.C. & Settele. Biological Conservation 28: 325-347. & Hochberg. Wardlaw.C. Oecologia 79: 452-457 Thomas. Dempster & I. (1984) The behaviour and habitat requirements of Maculinea nausithous (the dusky large blue butterfly) and M.A. G. 514.W.J. Woyciechowski.E. (1998) Higher productivity at the cost of increased host-specificity when Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation. (1998).. Wardlaw. & Wardlaw. Maculinea arion (large blue butterfly).W. . Thomas. A.A. J.A.A. (2005) Specialisations and host associations of social parasites of ants. J.A..T. G.A. by Myrmica ants. (2002) Larval niche selection and evening exposure enhance adoption of a predacious social parasite. Ford. J. teleius (the scarce large blue) in France. London Schönrogge..W. G. J. J. & Elmes. Institute of Environmental Sciences. teleius use as the host ant four species of Myrmica: M. Gronostajowa 7.co. Number of nests of particular species of Myrmica used as host ants by larvae of Maculinea teleius in the Kraków region (data from 2003 and 2004). 2. rubra workers after disturbance of ant nests Table 1. showed that: 1) about 11 % of all Myrmica nests are parasited by Maculinea teleius larvae.© PENSOFT Publishers Using butterfly monitoring Sofia – Moscow J. of all nests 1317 323 237 40 % 8. endangered in Europe. Until recently it was believed that each species of Maculinea has evolved to parasite a single. teleius larvae adopted by different Myrmica species (Tab. E. pp. scabrinodis (Tab. 1).7 14 17 15 Table 2. M. J.uk Maculinea is a butterfly genus. 105-106 Host ant specificity and integration rate with Myrmica ants in larvae of Maculinea teleius butterflies Magdalena Witek. Kühn & of Butterflies(Eds)105 Studies on the Ecology butterfly indicator and Conservation in Europe Vol. of experimental nests 36 34 % of nests with adoption 58 76 Mean time of adoption 393 min 180 min . M. Piotr Skórka.A. In the case of Maculinea teleius it was thought that Myrmica scabrinodis is a main host ant for this species. The aim of our studies was to investigate host ant specificity and integration rate with Myrmica ants in larvae of M. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. and different Myrmica species. rugulosa and M. whose larvae parasitise nests of Myrmica ants. rubra No. 2) larvae of M. Results of laboratory experiments showed that: (1) there is no difference in the proportion of M. Species M.Jagiellonian University. scabrinodis. Poland Contact: mawitus@yahoo.) and (2) Myrmica rubra adopt larvae of M. Ewa Sliwinska & Michal Woyciechowski UJAG . Piotr Nowicki. teleius. rubra ants and mean time of adoption (results of laboratory experiment). 2). ruginodis (Tab. rubra M. rubra. rugulosa No. 30-387 Kraków. Species M. Results collected in the Kraków region of southern Poland for two years (2003 and 2004). scabrinodis M. Percent of adopted larvae of Maculinea teleius by Myrmica scabrinodis and M. ruginodis M. teleius more quickly than M. scabrinodis M. teleius were not be carried by Myrmica scabrinodis or M. Results of another experiment showed that larvae of M. Thomas 2005 data to develop a European grassland Settele. teleius larvae with their hosts is very weak.106 Magdalena Witek et al. We also conclude that the integration of M. teleius shows quite plastic behaviour in respect to host ants and can use every Myrmica species that co-occurs with it. . These results suggest that M. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). and always were ignored by ants. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126).and two-year developing larvae coexist. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. teleius. For a long time it was believed that Maculinea larvae live for eleven months inside Myrmica nests and then pupate. southern Poland in 2003 and 2004. Myrmica species. 107 Biennialism and host ant specificity in Maculinea teleius larvae Magdalena Witek. a predatory species of Maculinea. Gronostajowa 7. scabrinodis. The aim of our studies was to investigate host ant specificity and larval development of M. Piotr Nowicki. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)107 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 30-387 Kraków.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. Piotr Skórka. p. It was thought that each species of Maculinea has evolved to parasite a single. and different. We also conclude that M. Kühn J. .Jagiellonian University. Ewa Sliwinska & Michal Woyciechowski UJAG . Results of laboratory experiments showed that: (2) there is no difference in the proportion of M. teleius larvae adopted by different Myrmica species.A.co.uk Maculinea butterflies possess highly specialized life cycles. Poland Contact: mawitus@yahoo. The studies were carried out in Kraków region. (4) there is no difference in larval survival rate in nests of different ant species for at least 30 weeks since adoption. These results suggest that within a single population one. Institute of Environmental Sciences. with larvae which parasitise nests of Myrmica ants. Settele. Our laboratory and field data indicated that (1) body mass distribution of pre-pupation larvae was bimodal. teleius shows quite plastic behaviour in respect to host ants and can use every Myrmica species of co-occurring with them. teleius more quickly than M. (3) Myrmica rubra adopt larvae of M. E. 108 Maurizio Biondi and Paola D’Alessandro This page intentionally left blank . 3. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Population ecology of Maculinea .Results of the mark-release-recapture studies of a Maculinea rebeli population 109 Section 3. 110 Ervin Árnyas et al. This page intentionally left blank . H-4010 Debrecen. 233 butterflies were recaptured only once.O. 3. see: Als et al. Their overall decline is connected with their vulnerabilty due to their obligatory myrmecophilous life cycle and with the restriction and fragmentation of their habitats. Varga 2004).44%) were recaptured.B. We chose a sample area of 150 x 50m for our studies in June 2002.O. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. a total of 806 butterflies were marked. E. so each day can be regarded as one sampling occasion. Andrea Tóth1 & Zoltán Varga1. most populations use only one initial food-plant for oviposition. In addition. sex ratio. of which 566 were male and 240 female. The habitat of the population is an area of approximately 3 hectares on a gently sloping ridge between the valleys Tohonya and Lófej. Thomas 2005 mark-release-recapture studies of on Maculinea rebeliKühn & of Butterflies(Eds)111 population in Europe Studies a the Ecology and Conservation Vol.2 1 DUFS . 493 males and 195 females. 780 individuals (507 males and . In 2003. and 81 at least three times. In 2002 a total of 688 individuals was marked. During the 20 sampling days of 2004. In 2003. 291 males and 123 females were recaptured. 1999. during the 10 days of sampling. The recapture rate was 51. Faculty of Sciences. P.hu The Large Blues of the genus Maculinea are among the most threatened butterflies throughout Europe. The calculations were done with the help of MARK and Popan programs. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group.36%. and the basic parameters of population dynamics can be estimated on the basis of the ratio of the marked vs. The captured specimens are individually marked and then released. Of the 688 marked butterflies. 34 at least three times. 2005) shows the most specialised form of myrmecophyly with „cuckooing larvae”. During the sampling period. 2003 and 2004. The Maculinea alcon-rebeli species group (according to some recent results only one species. 1004.University of Debrecen. Department of Evolutionary Zoology and Human Biology. H-4010 Debrecen. Varga et al. Pech et al. recapture probability. Bereczki et al. extensively surveyed already in the 90’es (V.unideb. number of individuals). pp. Judit Bereczki1. unmarked specimens captured (survival rate. Hungary Contacts:
[email protected]. 111-114 Results of the mark-release-recapture studies of a Maculinea rebeli population in the Aggtelek karst (N Hungary) between 2002-2004 Ervin Árnyas1.© PENSOFT Publishers the Results of Sofia – Moscow J. 173 butterflies were recaptured only once. 2004. P. Sipos et Varga 1997. J. The method of repeated marking-releaserecapture (MRR) was used in order to estimate the parameters characterizing the population. After a certain time sampling is repeated. We selected for our population ecological studies a local deme of the butterfly in a wellstudied site of the Aggtelek National Park. Settele. N of the village Jósvafõ. 2000. 3. The essence of the method is to capture as many individuals during the sampling period as possible. Sampling was carried out daily.A. 292 (42. 273 females) were marked.946 0. while recapture probability varies with time and is identical for both males and females.769 0. 66 at least three times. Estimate recapture rate (p) of the models with the best fit to the data set of 2002. i.e. 333 individuals were recaptured. Estimates of survival rate (Phi) of the models with the best fit to the data set of years 2002. The GOF test indicated that both baseline models fit the data sets of their respective years properly. 2003 and 2004.017 0.744 Standard Error 0.768 Fig. 2003 and 2004 Year 2002 2003 2004 Modell Phi(.) p(t) as the model with most support in the data.)p(t) Sex Males Females Estimate (Phi) 0. while recapture probability varies with time and sex.012 95% CI Lower 0. The margins of error show 95% confidence interval of the estimate. As a result of the model selection procedure on the 2003 data set. 188 butterflies were recaptured only once. i. the model in which the survival rate is time constant and identical for both sexes.806 0. no lack of fit could be statistically demonstrated. 1. The model selection procedure on the data sets of 2002 and 2004 resulted in model Phi(.718 95% CI Upper 0. Phi(g) p(g*t) proved to be the model with the most support in the data.)p(t) Phi(g)p(g*t) Phi(. which corresponds to a 42.857 0.025 0.832 0.7% recapture rate. the model in which survival rate is constant and varies with sex.e.868 0.979 0. Survival rate (phi) shows that less than 20 % of the butterflies disappeared (died or migrated) from one day to the next in 2002 and Table 1.804 0.013 0.112 Ervin Árnyas et al. .838 0. The models of best fit are shown in Table 1 and in Figures 1. On the basis of the estimated values. we found different survival and recapture rates for the two sexes. Females. 2. With increasing wind speed and cloud cover the number of captures tends to drop significantly. tend to fly lower because they are searching for initial food plants in the undergrowth. The diagrams demonstrate well that the ratio of males was much higher in the Fig. The number of recaptured individuals decreased significantly. 2003and 2004 with confidence intervals of 95% . the population can be regarded as closed in terms of migration. Estimated sex ratio for each sampling day in 2002. on the other hand. estimates could only be made for ten days because there were only recaptures on the last sampling day. According to the estimated value of phi. Recapture rate (p) shows a great variation from one day to the next. In 2003.5 kms away) although the flight periods of the two populations overlapped. 25% of the population disappeared overnight in 2004. In 2004. searching for less mobile females.Results of the mark-release-recapture studies of a Maculinea rebeli population 113 2003. because butterflies hiding in the undergrowth are hard to observe. The higher recapture rates of males may be attributed to the fact that they are easier to capture as they fly more often and higher. In 2002. Sex ratio was estimated for each sampling day. the estimated values of the recapture rate were high until day 14 of the sampling period. which can be due to the difference in the way males and females behave. but then there was a long period of drought in the area. This is also supported by the fact that we have been unable to find a single marked individual during genetic sampling of the nearest Alcon Blue population (located about 1. Field experience suggests that the value of (p) may be strongly affected by wind speed and cloud cover. The estimated value was calculated as the ratio of the number of males captured and the total number of the individuals captured on a given day. 114 Ervin Árnyas et al. Fig. 3. Estimates of the number of individuals in 2002-2004 by the Jolly-Seber model. The margins of error reflect the standard error of estimation population in the first two thirds of the sampling period in the years studied. Following the flight period, the ratio of females increased gradually, and the sex ratio approximated 1:1 in the last third of the sampling periods. The results of population estimation agree with the estimates of the sex ratio. The bell shaped curve of 2002 indicates that the sampling period covered the flight period. Although sampling stopped a few days before the end of the flight period in 2003, it still provided a good estimate of the size and composition of the population. In 2004, the flight period was longer than in previous years due to rainy weather in the first half of the period. That year, the adults emerged nearly a week later than in the two previous years. At the peak of flight, the estimated number of the sexes was about 100 less in 2004 than in earlier years. However, when we take into consideration the facts that the flight period in 2004 lasted about one and a half weeks longer than in 2002 or 2003, and that about the same number of individuals were marked in 2004 as in the earlier years, it can be stated that the Tohonya valley has had a stable population of about over a thousand individuals for the past three years. In summary, we conclude that the Tohonya valley population is stable, and not threatened by extinction. However, it is essential that the favourable structure of the vegetation should be maintained by continuing the management of the area started in 2002. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). © PENSOFT Publishers Modelling the local Sofia – Moscow Settele, E. J.A. Thomas 2005 population dynamics of Maculinea and theirJ. spatial Kühn & of Butterflies(Eds)115 interactions in Europe Studies on the Ecology and Conservation Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, pp. 115-119 Modelling the local population dynamics of Maculinea and their spatial interactions with their larval foodplant and Myrmica ant species Ralph T. Clarke1, Nicolas Mouquet2, Jeremy A. Thomas1, Michael E. Hochberg2, Graham W. Elmes1, David Tesar2, Alexander Singer3 & Joseph Hale1 NERC - Centre for Ecology & Hydrology, CEH Dorset, Winfrith Technology Centre, Winfrith Newburgh, Dorchester, Dorset DT2 8ZD, UK 2 ISEM - University of Montpellier II, Bat.22, Equipe Hochberg, Place Eugene Battallion, 34095 Montpellier, France 3 UFZ-Centre for Environmental Research Leipzig-Halle, Department of Ecological Modelling, Permoserstr. 15, 04318 Leipzig, Germany Contact:
[email protected] 1 Maculinea blue butterflies have an intricate relationship with their early larval foodplant and the Myrmica ant species on which they depend for survival to adulthood. At the large spatial scale it is important to understand the factors controlling the dynamics of the butterfly within a landscape composed of several populations on various sites. However, most recorded movements and dispersal of Maculinea through mark-release-recapture studies are less than 500m (Nowicki et al. “this volume”) and many remaining fragmented populations are isolated. It is therefore valid and useful to try to develop models which synthesise our knowledge of the internal local dynamics of individual populations on single sites assuming no emigration or immigration. Estimates of population size inter-annual variability, local persistence rates and growth rates of re-colonisations can also contribute to landscape scale metapopulation studies. Within the MacMan project (http://www.macman-project.de), our single-site modelling of Maculinea extended two existing approaches, one involving deterministic density-dependent interaction models (Hochberg et al. 1992) and the other based on spatially-explicit individual-based stochastic processes (HCET model of Hochberg et al. 1994; Clarke et al. 1997,1998). Firstly, a deterministic community module has been developed for Maculinea alcon (Mouquet et al. “in press”). This model was novel and made realistic by incorporating the life stage (seedling, juvenile, vegetative, reproductive and dormant) dynamics of its larval foodplant Gentiana pneumonanthe, including the density-dependent impact of the butterfly damage to seedpods and the effect of increasing heathland successional age on reducing both gentian productivity and the abundance of nests of the dominant host ant, Myrmica scabrinodis. Model parameters for gentian and butterfly dynamics were estimated from a mixture of long-term (>20 years) field studies and laboratory experiments. Without management of the heathland or natural fires to 116 Ralph T. Clarke et al. reset or prevent succession, the host ant and the gentians and thus M. alcon will tend to disappear. Model simulations were used to assess the management effect of burning, grazing, mowing or sod cutting the heathland at various frequencies (2-15 years) and intensities on the three species’ persistence and population sizes (Table 1). Maculinea rebeli and alcon act as “cuckoos” within the ant nests, being fed by the ants, which provides a form of “contest” competition. In contrast, Maculinea arion predates upon the ant brood, which leads to less efficient “scramble” competition for food such that if too many caterpillars are adopted into a nest, few, if any, survive to adulthood (Thomas & Wardlaw, 1992). We have developed a deterministic model for M. arion (Mouquet et al. 2005). In addition to the density-dependent contest competition of larvae on the flower buds of Thymus (or Origanum), this model includes a form of unimodal density-dependent overall survival of caterpillars within the Myrmica sabuleti host ant nests, where the intensity of within-nest competition can be related to the extent of spatial clumping of egg-laying and thus adoption of caterpillars. Estimates of model parameters used field data from sites in the UK, France and Sweden, but especially from one location in south-west England during 1972-2002, leading to two model test scenarios under (i) average and (ii) maximum/optimum observed conditions. Under otherwise average conditions, adult butterfly numbers track closely the simulated long-term (20 year) cyclical variation in the number of ant nests (Figure 1(a)), whereas reductions from high to very low plant densities initially lead to increases then rapid decreases in butterfly numbers followed by rapid recovery when plant density increases again (Figure 1(b)). In contrast, adult population responses to major cyclical variation in (weather-related) adult fecundity or survival on the plant are delayed by 4-5 years (Figure 1(c)). Survival dynamics operate at different spatial scales over the Maculinea life cycle, from egg laying across a site, to larval competition on individual plants, to adoption and development within ant territories and colonies (plus un-modelled occasional migration between sites). We have tried to incorporate this reality by extending and generalising the spatial-explicit HCET model developed for M. rebeli to other Maculinea and, in particular, for the contrasting predatory species M. arion (Table 2). Model parameters for the case of M. arion were derived from published and unpublished field studies and other unpublished studies of Myrmica ants. Table 1. Estimates of optimum frequency (years) of heathland burning, grazing, mowing or sod cutting on average population densities (per hectare in brackets) of Maculinea alcon, Myrmica scabrinodis and Gentiana pneumonanthe using model parameters and results from Mouquet et al. “in press” ). Management Burning Grazing (strong) Grazing (intermediate) Grazing (weak) Mowing (5cm height) Mowing (10cm height) Sod cutting (5% area) Sod cutting (15% area) Sod cutting (25% area) Maculinea alcon Myrmica scabrinodis Gentiana pneumonanthe 4-6 (36000) 4 (19800) 2-3 (17400) 2 (3000) 2-4 (35300) 4 (19700) 2 (11500) 2 (18100) 4 (8400) 3 (952) 2 (1100) 3 (664) 2 (982) 2 (602) 2 (214) ant and butterfly do not survive 2 (252) 2 (114) ant and butterfly do not survive ant and butterfly do not survive 2 (260) 2 (102) 3 (274) 2 (616) Modelling the local population dynamics of Maculinea and their spatial interactions 117 Fig. 1. Effect of temporal variation (dotted sinusoidal line) in (a) the number of host ant nests, (b) the number of larval foodplants and (c) adult fecundity on adult numbers of M. arion butterflies (circles with solid line); dashed vertical line indicates start of temporal variation once equilibrium has been reached using constant average site conditions (based on model scenario #1 from Mouquet et al. 2005). M. arion caterpillars can cope with starvation periods and so can exploit multiple ant nests by consuming all of the over-wintering ant brood in a nest, causing the workers to disband, and allowing neighbouring healthy nests to bud into the empty site bringing new brood. Model simulations showed that being less host specific and able to exploit multiple (optimally 3) ant nests increases M. arion survival in poor conditions (low plant and host ant density), but can result in over-exploitation of the ants at very high plant densities (Figure 2). As risk of local extinction is 118 Ralph T. Clarke et al. Table 2. Extensions and generalisations of the spatial-explicit individual-based HCET model for Maculinea population dynamics within sites. Stochastic or deterministic temporal variation in habitat quality gradient – representing effects of inter-annual weather or site management Extending from ‘host’/’non-host’ to up to eight competing Myrmica ant species Ant colony distribution extended to 1 colony (territory) per m2 (i.e. 1 m2 cells in model) Territories overlap: ants forage and can adopt caterpillars in neighbouring territories Some (16%) caterpillar tolerance (and potentially survival) in ‘non-host’ Myrmica (i.e. degree of non-host specificity) Scramble competition within individual nests as function of current ant worker and caterpillar numbers Increased within-year inter-colony dynamics (nest abandonment + budding) – M. arion caterpillars can consume brood from 1-4 immigrant colonies to develop User friendly interface to change model parameters Model can be run to simulate effects of: – different physical properties or management – different values of host specificity – annual or geographical variation in climate – edge-of-range versus core populations – functional evolutionary questions Fig. 2. Example of effect of varying Thymus plant density (m-2) on Maculinea arion adult density (ha-1) assuming caterpillar scramble competition within nests with either (a) complete host specificity to Myrmica sabuleti and only exploiting one colony (solid line, closed circles), or (b) 16% probability of tolerance within non- M. sabuleti nests and ability to exploit up to three nests (dotted line, open circles). Modelling the local population dynamics of Maculinea and their spatial interactions 119 greatest in poor conditions, this may explain the evolutionary selection pressure for incomplete host specificity and the ability for caterpillars to starve during temporary periods of colony disbandonment (Thomas et al. 1989). Overall we conclude that conservation strategies for optimal management of Maculinea need to be conducted on a case by case basis. REFERENCES Clarke, R. T., Thomas, J. A., Elmes, G. W. & Hochberg, M. E. 1997.The effects of spatial patterns in habitat quality on community dynamics within a site. Proceedings of the Royal Society, London, Series B., 264, 347-354. Clarke, R. T., Thomas, J. A., Elmes, G. W., Wardlaw, J. C., Munguira, M. L. & Hochberg, M. E. 1998. Population modelling of the spatial interactions between Maculinea, their initial foodplant and Myrmica ants within a site. Journal of Insect Conservation, 2, 29-37. Elmes, G. W., Clarke, R. T., Thomas, J. A. & Hochberg, M. E. 1996. Empirical tests of a spatial model of the population dynamics of Maculinea rebeli, a parasitic butterfly of red ant colonies. Acta OEcologica, 17, 61-80. Hochberg, M. E., Thomas, J. A. & Elmes, G. W. 1992. A modelling study of the population dynamics of a large blue butterfly, Maculinea rebeli, a parasite of red ant nests. Journal of Animal Ecology, 61, 397–409. Hochberg, M. E., Clarke, R. T., Elmes, G. W. & Thomas, J. A. 1994. Population dynamic consequences of direct and indirect interactions involving a large blue butterfly and its plant and red ant hosts. Journal of Animal Ecology, 63, 375-391. Mouquet, N. Thomas, J. A., Elmes, G. W., Clarke, R. T. & Hochberg, M. E. “in press (a)”. Conserving community modules: a case study of the endangered lycaenid butterfly Maculinea alcon. Ecology. Mouquet, N. Thomas, J. A., Elmes, G. W., Clarke, R. T. & Hochberg, M. E. (2005). Population dynamics and conservation of a highly specialized predator: A case study of Maculinea arion. Ecological Monographs, 75, 525-542. Nowicki, P., Woyciechowski, M. & Settele, J. “this volume” A review of population structure of Maculinea butterflies. Thomas, J. A., Elmes, G. W., Wardlaw, J. C. & Woyciechowski., M. 1989. Host specificity among Maculinea butterflies in Myrmica ant nests. Oecologia 79:452-457. Thomas, J. A. & Wardlaw, J. C. 1992. The capacity of a Myrmica ant nest to support a predacious species of Maculinea butterfly. Oecologia 91:101-109. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). de Dispersal is a key process affecting the fitness of individuals as well as the persistence of populations.A. I first give an overview over the factors potentially promoting the evolution of dispersal in butterflies and what kind of dispersal strategies we might expect to find. Habitat succession and environmental variability are probably the most relevant factors driving dispersal in butterflies. I use different approaches to analyze displacement data collected in MRR studies of two different Maculinea species in Poland and Germany. In practice. Germany Contact: hovestadt@biozentrum. I will demonstrate that this has important implications for the predicted persistence of populations. Field Station Fabrikschleichach. Glashüttenstrasse 5.University of Würzburg. 120 A review of the role of dispersal for population persistence in Maculinea Thomas Hovestadt UWUERZ .uni-wuerzburg. Many ecologists consider dispersal to be a rare and special event in the life-time of an organism. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Settele. Sofia – Moscow J. .© PENSOFT Publishers 120 Thomas Hovestadt et al. I will discuss reasons why dispersal may be comparatively rare in Maculinea species. animals move around most of the time and it is hard to separate “every-day” movement from dispersal-motivated displacement. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. 96181 Rauhenebrach. These data do not only indicate that Maculinea species presumably establish home-ranges but also that at best a small fraction of the displacement observed is related to dispersal. Kühn & J. E. In addition dispersal is likely to be context-dependent. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). p. In this talk. DIE will lead to a more rapid colonization of empty patches than DDE. DDE is characterized by exceptional migration episodes in good years and no to little emigration in poor to average years. Colonization success under DIE is much more sensitive to an Allee-effect than under DDE. this pattern is reversed as distance becomes progressively larger.uni-wuerzburg. p. however.A. i.e.University of Würzburg. Germany Contact: hovestadt@biozentrum. 121 The control of emigration. In a structured landscape colonization of unoccupied patches may only be successful if a critical number of immigrants simultaneously arrive in that patch. in situations where species strongly respond to environmental variability. Using a numerical and individual-based simulation approach I demonstrate that with density-dependent emigration (DDE) chances for rapid range expansion are much higher than with density-independent dispersal (DIE). the Allee-effect. The difference between density-independent and -dependent dispersal is especially large if an Allee effect prevents establishment from a small number of immigrants.de Most species show rather static (or slowly shrinking) distributions over the landscape. Field Station Fabrikschleichach. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). environmental variability. The model is consistent with observations of long static species distributions with occasional episodes of massive range expansion. The discrepancy between the two scenarios are strongest when populations are very dynamic. and the expansion of species’ range Thomas Hovestadt UWUERZ . in recent years in several cases attributed to the effect of global warming.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. Settele. Glashüttenstrasse 5. This can lead to periods of static or even retreating species distribution interfered by moments of explosive “Big-Bang” range expansion in exceptionally good years. with very small numbers of migrants in “ordinary” or poor years but exceptionally large numbers in good years. At close distance. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)121 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Kühn J. occasionally rapid range expansion have been reported. E. 96181 Rauhenebrach. In those years even remote patches may be reached by sufficient numbers of immigrants to establish a new population. This is a consequence of the skewed distribution in the number of emigrants. . However. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. J. For both species net-displacement did hardly change with the time-lag between mark and – recapture. Germany 2 UJAG . Thomas © PENSOFT Publishers 122 Christian Anton. & Josef Settele on the EcologySettele. Institute of Environmental Sciences. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). 96181 Rauhenebrach. Subsequently. 5. . Kühn & of Butterflies(Eds) 2005 Studies and Conservation in Europe Sofia – Moscow Vol. p. Field Station Fabrikschleichach. If boundaries of patches become barriers for movement we cannot easily compare observed net-displacement distances with those predicted by theoretical dispersal kernels. Martin Musche. 30-387 Kraków. After randomisation the null-model always fitted better to the data than the negative exponential model. Glashüttenstrasse 5. which may have serious implications for their conservation. Poland.uni-wuerzburg. 4. during a MRR study on the two butterfly species: Maculinea nausithous and M.A. we used the procedure to generate “optimal” estimates for displacement parameters for three different dispersal kernels. We applied the procedure to a data-set collected in the vicinity of Kraków. Gronostajowa 7.de 1. Statistical models for the analysis of net-displacement data extracted from mark-releaserecapture (MRR) studies are frequently based on the assumption of movement in unlimited space.University of Würzburg. E. Poland Contact: hovestadt@biozentrum. a simple negative exponential model usually gave the best fit. but a kernel with P(r)~r*exp(-r) always allowed for even closer fit between original and simulated data. Contrary to expectation.Jagiellonian University.and species-specific. The estimates for mean displacement distance d was both site.J. teleius. 2. We developed a permutation procedure to compare observed net-displacement distances with those expected under the null-model of a uniform distance probability distribution within patch-boundaries. 122 Within-patch movement limitation in two species of Maculinea butterflies? Analysis of MRR data using randomisation procedures Thomas Hovestadt1& Piotr Nowicki2 1 UWUERZ . This in turn suggests that these butterflies establish home ranges within habitat patches. When directly fitted to the empirical distance probability distribution. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. the data analyses indicate the existence of within-patch displacement limitation in both species. 3. both modes of emigration increase survival of the metapopulation by ca.University of Würzburg. 96181 Rauhenebrach. (3) With high dispersal success. For isolated patches. Kühn J. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Settele. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)123 Studies on the Ecology and Conservation of Butterflies in Europe Vol. p. . Field Station Fabrikschleichach.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J.2. one order of magnitude. (4) With low dispersal success.uni-wuerzburg. We use individual-based simulations to compare the consequences of density-independent (DIE) and density-dependent emigration (DDE) for the extinction risk of local populations and a two-patch metapopulation. (1) Emigration always reduces the survival chances of source populations as long as it is not compensated by immigration. extinction risk is nearly insensitive to emigration as long as emigration probability remains below 0. With DDE. How these two opposing processes interact to define overall persistence depends on the way emigration is controlled. E.de Dispersal is a key process for the persistence of metapopulations. any individual emigrating from a local population also increases the risk of local extinction. only DDE can improve the global survival chances of the metapopulation. Germany Contact: hovestadt@biozentrum. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). However. extinction risk continuously increases with emigration rates in the DIE scenario. 123 Emigration and its consequences for the survival of metapopulations Thomas Hovestadt & Hans Joachim Poethke UWUERZ . Glashüttenstrasse 5.A. (2) With DDE the risk of extinction is always smaller than with DIE. Josef Settele3 UWUERZ . i. the mean dispersal distance was larger) when fitted only to the data with dispersal between patches that did not share a border: for M. Germany Contact: hovestadt@biozentrum. The observations suggest that individuals of both species eventually establish home ranges spanning across different local sites. Data were analysed to quantify the dispersal range of the species and to check for evidence of an area effect in immigration. & Josef Settele on the EcologySettele. p. Thomas © PENSOFT Publishers 124 Christian Anton. 4.e. Germany 2 Friedhofstrasse 1.Centre for Environmental Research Leipzig-Halle. suggest that for both species the system should not be considered to be a metapopulation. nausithous (n=218).A. teleius mean dispersal distance 2/α ≈ 500m (n=18). For both species more than 50% of individuals were recaptured in another patch than the one where they were originally marked. in the much larger complete data set for M.de 1 We have investigated mark-recapture data collected in the year 1994 in a fragmented landscape in northern Bavaria. as well as estimates for the dispersal parameter α. 124 Analysis of inter-patch dispersal data for Maculinea nausithous and M. teleius in a fragmented landscape in northern Bavaria. 96181 Rauhenebrach.uni-wuerzburg. However. We performed analyses for the whole data-set (for each species) and for a reduced data-set that included only the transfer of individuals between sites that did not share a border. For both species the fitted dispersal parameter α was smaller (i. Glashüttenstrasse 5. Germany for the two Maculinea species M. 06120 Halle. 97475 Zeil. For the data-set including only transfers between patches without a common border. immigration was roughly proportional to patch circumference. Germany Thomas Hovestadt1. The number of recapture events from sites other than the one where the animals were originally marked. Field Station Fabrikschleichach. Birgit Binzenhöfer2. for M. Kühn & of Butterflies(Eds) 2005 Studies and Conservation in Europe Sofia – Moscow Vol. In both species we found no correlation between the time lag between marking and recapture and the distance between the patch of marking and the patch of recapture. a mixed kernel model led to a highly significantly improved fit of the model with most of the dispersal (94%) being over short distances only (mean distance of 126m) and 6% over much larger distances (mean distance of 1340m). we found significant scaling of immigration to area I~Ac with c ≈ 0.5. J. This indicates that the movement of individuals across the landscape can probably not be approximated by a random walk. nausithous and M.e. teleius. E.University of Würzburg. nausithous 2/α ≈ 420m (n=157). Martin Musche. Theodor-Lieser-Str. The study sites were distributed over a distance of > 6km. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.J. . Germany 3 UFZ . Department of Community Ecology. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). uni-wuerzburg. . can prevent the (re-)colonization of empty habitat patches. We use individual-based simulations to model the colonization of an empty habitat patch by an annual organism with density-dependent population growth underlying environmental as well as demographic stochasticity. 125 Control of emigration. Glashüttenstrasse 5. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)125 Studies on the Ecology and Conservation of Butterflies in Europe Vol. p. 96181 Rauhenebrach. (3) As long as either the Allee-effect is weak or dispersal mortality is low DIE is more likely to lead to rapid (re-)colonization than DDE.de The Allee-effect.e. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Tobias Degen & Hans-Joachim Poethke UWUERZ . (1) Under DIE. the Allee-effect and the recolonisation of empty habitats Thomas Hovestadt. E. the distribution of emigrants is close to normal. we compare the colonization success either assuming density-independent (DIE) or density-dependent emigration (DDE). low population growth at low population densities. i.A. Germany Contact: hovestadt@biozentrum. Specifically. Settele. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.University of Würzburg. Kühn J. they emigrate in large numbers. Field Station Fabrikschleichach.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. under DDE it is highly leptokurtic. (4) When the Allee-effect or dispersal mortality become large DDE is more likely to lead to the successful establishment of a new population.e. in many years no individuals emigrate but sometimes. i. The likelihood of establishing a new population will depend on the number of immigrants arriving within a generation. (2) Even a small to moderate Allee-effect prevents a successful colonization from a small number of immigrants. The ESSmodel accounts for (i) the added mortality risk experienced by SDL. (ii) the effect of competition among larvae for the limited resources the ants can provide. We present an analytic model to investigate and explain under which conditions the evolution of such a dimorphism can be adaptive in spite of an added mortality risk for the SDL in their second winter. Jeremy Thomas2 & Michael Hochberg3 1 UWUERZ . Settele. Dorchester. (iv) for an existing competitive advantage of SDL over newly entering FDL “priority effect”. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. .University of Würzburg. Dorset DT2 8ZD. 126-127 An ESS model for the evolution of growth polymorphism in the social parasite Maculinea rebeli Thomas Hovestadt1. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Kühn & J. (ii) Whatever the kin-competition.University of Montpellier II. Field Station Fabrikschleichach. the remaining three quarters delay development into the second summer and emerge only after two years (“slow developing larvae”. the fraction of larvae delaying development should never surpass a value of 0. Sofia – Moscow J.22. While approximately one quarter of the larvae entering a host ant colony grow and develop within one year (“fast developing larvae” FDL). (iii) Even with a minor priority effect the evolution of even higher frequencies of delayed development than those observed empirically becomes likely.Centre for Ecology & Hydrology.uni-wuerzburg. Winfrith Technology Centre. E. Place Eugene Battallion. pp. . SDL). Glashüttenstrasse 5. UK 3 ISEM . (iii) for the potential competition among sibling larvae. There is compelling evidence that this polymorphism is not just the result of temporal or energetic constraints but is genetically controlled. Bat. France Contact: hovestadt@biozentrum. the larvae of M.© PENSOFT Publishers 126 Thomas Hovestadt et al.A. parasitic butterflies of the genus Maculinea are one of the few known examples which exhibit a polymorphism in their developmental strategy. 34095 Montpellier. CEH Dorset. Equipe Hochberg. Germany 2 NERC . Graham Elmes2.5 in the absence of a priority effect.5.de Among insects. After a short growth period on the host plant. The model allows us to draw the following conclusions: (i) Without a priority effect a minimum level of kin-competition must be involved to explain the evolution of delayed development. rebeli are carried into the nests of their host ants to complete their development there. Oliver Mitesser1. (iv) This could only be prevented by rather unrealistically high degrees of relatedness among larvae (kin-competition). Winfrith Newburgh. (v) the tendecy of infected ant-colonies to move away from the vicinity of host plants (segregation). 96181 Rauhenebrach. (v) Segregation is a further important factor stabilizing the fraction of SDL closer to 0. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). rebeli 127 M.An ESS model for the evolution of growth polymorphism in M. The model presented here may be of general applicability for systems where maturing individuals compete in smaller subgroups of related individuals. rebeli provides an example of reduction of competition and especially sibling competition by dispersal in time. . There were some failures in the model selection procedure. The minimum number alive was acceptable above 60% SI compared to the 100% DSI values. 6th and 9th occasion. Standard error decreased rapidly below 25 percent in the case of apparent survival and below 55% in the case of the recapture probability. 128-129 Goodness of sampling in Maculinea butterflies Ferenc Kassai HNHM . At 20% sampling intensity. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. It can be concluded. The number of captured individuals and the number of captures strongly depended on ESI. Changing of the daily sampling intensity (DSI) was modelled by randomly reducing the number of daily captures from 100% to 5%. By eliminating every 2nd. Hungary Contact: fe. the time dependent models were better supported than the constant ones. while the number of captured individuals increased assimptoticly after 70% DSI. LS had no impact on model selection procedure. the most supported CJS model was the Phi(. Several aspects of GOS were studied using data obtained from Maculinea ligurica at Vérteskozma (Hungary). Kühn & J. The effect of sampling intensity within the entire study period (ESI) was also examined. 1088
[email protected] Goodness of sampling (GOS) could provide a tool for optimizing sampling investment in time and space in capture-recapture studies. At certain sampling intensities (60%-70%-85%-90%). We examined the model selection procedure and estimates of CJS models on all dataset variants of the different levels. Settele.13. pp. Recapture probability increased linearly. 3rd. The number of sampling occasions fluctuated. ESI was modelled by eliminating sampling occasions. four ESI levels were obtained. In all cases. and SEs were high at low ESI levels.© PENSOFT Publishers 128 Ferenc Kassai Sofia – Moscow J. Population size was estimated with Pollock’s robust design method at different DSIs. but increased in a similar manner.Hungarian Natural History Museum. Values of p and Phi were fluctuating in a wide range.). the data failed on the GOF test. Baross u. that the CJS models provide acceptable estimates above 70% DSI in the case of the dataset was examined.)p(. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. the number of captures linearly depended on the DSI. Department of Zoology. In this example.A. Both the num- . the best supported model was different than when the entire dataset was used. Estimated population sizes strongly depended on DSI. Reduced DSI caused failures in the model selection procedure. The effect of the length of sampling (LS) on model selection procedure and estimates of CJS models was examined. the value of estimated apparent survival was biased below 25% DSI and it was relatively stable above 25%. As expected. E. thus this model is applicable when sampling effort is extremely intensive. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). The optimal timing of daily sampling investment was examined.Goodness of sampling in Maculinea butterflies 129 ber of captures and the number of captured individuals depended linearly on LS. and 12 a. sampling was most effective between 10a. Estimated values of p and Phi were stable after the 8th day. while the value of SEs decreased rapidly within the first 10 days. Based on the number of specimens captured/time invested index.m.m. . 130 Annual and spatial variations in population structure – A case study of Maculinea alcon and Maculinea teleius Ferenc Kassai. The number of butterflies decreased moving away from the edge of the bogs. In 2002 we selected for sampling only that bog where most of the butterflies were seen. We carried out mark-release-recapture studies for 2 years in the same habitat on two different spatial scales: some preliminary results are presented here. Next year.Hungarian Natural History Museum. sampling was carried out around all bogs. reed and some water cover. Gentiana pneumonanthe and Sanguisorba officinalis occur in huge quantities in a 30-40 m wide belt around the bogs and the density of butterflies (Maculinea alcon and Maculinea teleius) is highest here. Ágnes Vozár & Lilla Barabás HNHM . E.13. The habitat was a network of bogs separated by different types of marshland meadows. Bogs are shallow basins with trees. p. 10×10 m squares were pegged out around the bog. Department of Zoology. bushes. Sofia – Moscow J.zoo. The ratio of the two Maculinea species was similar at different bogs.A.nhmus. Noémi Örvössy. Baross u. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. .© PENSOFT PublishersElmes & Ralph T. Ádám Kőrösi. but the density of butterflies differed. The abundance of foodplants does not seem to influence the number of butterflies around the bogs. Kühn & J. László Peregovits.hu The spatial scale of an investigation determines the population structure that can be observed. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Settele. Hungary Contact: korosi@zoo. 1088 Budapest. Clarke 130 Graham W. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). and to infer the spatial structure of populations.zoo. When a high proportion of individuals in a population is recaptured 5-10 times. alcon and of M. and a separate population of M. 1088 Budapest. p. J. arion ligurica. respectively). Individual tracking is recommended in order to collect much more data about the dispersal and behavior of individuals. The diffusion rate combines both the mean and variance of the movement distances in one single measure. teleius and M. An analysis of net squared displacement was also carried out.13. 7. 131 Analysis of within-habitat patch movement of some Maculinea species Ádám Kőrösi HNHM . Mark-release-recapture studies on three populations of different Maculinea species were carried out in order to describe the pattern of movement of individuals. In both studies the sampling area was divided into unit squares (10×10 m. arion ligurica population (73. and this supported the fit of uncorrelated random walk model. teleius co-occuring in the same habitat.5×7. we could not test for this model because of insufficient data (lack of long recapture histories).03 m2/hour) compared to the others (M.hu Dispersal has a basic effect on local population dynamics and the long-term persistence of a species at the metapopulation scale. In the case of the M.A. alcon population. it provides overall information about the dispersal of a population. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126).nhmus. so the uncorrelated random walk model was accepted. Kühn &Poland (Eds)131 Studies and M. There was no autocorrelation between subsequent moves in the case of the M. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. and incorporates the time factor. Dispersal within habitat patches determines the spatial structure of local populations. Hungary Contact:
[email protected] Natural History Museum. Thomas 2005 and ecology of Maculinea teleiuson the EcologySettele. Department of Zoology. alcon: 183. arion ligurica populations. the duration and length of the moves were calculated. then the data can be tested for random walk. this is likely to be due to the smaller size of the sampling area. and the movement of individuals was characterized by a set of subsequent moves. so the mark–release–recapture of individual adults is applicable for analysis of movement. M. . Baross u. Its value was much lower in the M. Each recapture was considered as a move. We studied one population of M. Although it involves a big simplification. An overall test for independence of subsequent displacements proposed by Swihart & Slade (1985) was adapted.39 m2/hour).85 m2/hour. teleius: 207.© PENSOFTThe distribution Publishers Sofia – Moscow J. The diffusion rate of each population was calculated according to the uncorrelated random walk model. nausithous in of Butterflies in Europe and Conservation Vol.5 m. E. Department of Zoology. The emigration and immigration rates were calculated for all patches and both negatively correlated with patch size.13. p. Kühn & J. even though its population size in 2004 was much lower. In some patches the proportion of residents was zero. Baross u. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Hungary Contact: korosi@zoo. Sofia – Moscow J. This study showed that all species prefer old abandoned patches instead of annually mown ones. nausithous seemed to move through the whole habitat along edges and used the habitat most evenly.nhmus. To study such a case we carried out a MRR study in 2003 and 2004 on a wet meadow. teleius adults increased with the size of patches. M. the mobility of species between patches was analysed and compared. 1088 Budapest. M teleius was the most abundant species. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). nausithous) occur and where their flight periods overlap. we must have some information about the habitat-use of these butterflies. Clarke 132 Graham W. where all of the three wetland Maculinea species (M. nausithous occurs mainly in shaded areas near to forest edges. M. although the quantity of blooming Sanguisorba officinalis was not lower. E. Secondly. even when three Maculinea species occur in the same habitat. teleius. M. 132 Habitat-use of wetland Maculinea species – a case study Ádám Kőrösi HNHM . but in patches that are mown twice a year.hu In order to achieve succesful conservation management on Maculinea sites. Settele. The number of M. . M. In order to describe habitat-use.zoo. alcon is restricted to some smaller parts of the habitat. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. we first examined whether the density pattern of populations is characteristic of the species.© PENSOFT PublishersElmes & Ralph T. while M. irrespective of foodplant abundance.Hungarian Natural History Museum.A. much fewer butterflies were detected. alcon. Previous works suggested that the population size of this obligately fully predacious Maculinea species (such as M. arion) may fluctuate considerably. . in spite of the fact that the habitat and abundance of food plant appears stable. In 2002 only one population was sampled and the estimated number of butterflies was ca. In Hungary it is on the wing in July and lays eggs on Origanum vulgare. Next year two populations. 1088 Budapest. Baross u. 133 Studying the population structure of Maculinea arion ligurica Ádám Kőrösi. were studied and similar densities were estimated.A. Department of Zoology. It occurs mainly at forest edges. László Peregovits. vulgare and serve as corridors for the adult butterflies. Ágnes Vozár & Ferenc Kassai HNHM . Thomas 2005 and ecology of Maculinea teleiuson the EcologySettele. E. patches of foodplant are situated mainly in woodland clearings and are often isolated from each other by several kilometers.hu Maculinea arion ligurica is a phenotype of M. J. Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). p. A 4-year study of a population living in the Vértes Mountains was carried out to investigate the structure and dynamics of the population. On our study site. Verges of roads that connect these patches support high densities of O. 300 specimens in a 2 hectare habitat patch. about 4 km far from each other. arion. In 2004 and 2005 the number of butterflies was so low that sampling was pointless.zoo.© PENSOFTThe distribution Publishers Sofia – Moscow J. Hungary Contact: korosi@zoo. Conservational management of Maculinea habitats must be planned with a full knowledge of the butterflies’ population biology. although the quantity of food plant and the quality of habitat patches did not change considerably.13. This case study tries to draw attention to difficulties of studying this species’ population structure. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. We observed this pattern when Mark-Release-Recapture (MRR) studies were carried out in local populations that form a metapopulation. on woodland clearings and in deforested fields in mountain areas.Hungarian Natural History Museum.nhmus. Noémi Örvössy. Kühn &Poland (Eds)133 Studies and M. nausithous in of Butterflies in Europe and Conservation Vol. Germany Contact:
[email protected]). This is because the sex ratio dynamics within season most often . Gronostajowa 7. e (iv) flight period length (FPL) defined as the time between the first and the last day with >2 individuals.9046׈ ×FPL–1 +0. yet less reliable.Centre for Environmental Research Leipzig-Halle.14) after different life span and season length were accounted for.39). We analysed 23 capture-recapture data sets from closed populations of six Lepidoptera species (Maculinea nausithous. E. 4. but mostly below one third. but not necessarily with seasonal population size. The implication is that population size can be reliably estimated if the three other population parameters are known with the regresˆ ˆ sion equation Npeak = Ntotal /(1. ˆ ˆ The results obtained indicated that the proportion of individuals flying at peak (Npeak /Ntotal ) was highly variably (CV = 0. Moreover. corrected for individuals emerging and dying between capture periods ˆ (ii) peak population size (Npeak ) defined as the maximum daily population size. are cheap and time-efficient.5. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. in turn. pp.edu. ˆ ˆ ˆ (iii) average daily survival rate (φ ) and life span (ˆ ) calculated as e = (1–φ )–1 – 0. M. or 2 individuals of different sexes. 134-135 Simplified method of estimating butterfly population size with mark-release-recapture Piotr Nowicki1 & Josef Settele2 UJAG . but this relationship cannot be applied for detecting peak time. Our aim was to develop a sampling scheme that would retain the accuracy of standard MRR. ˆ in the case of MRR we get estimates of not only ‘snapshot’ (= daily) N. corrected for different capture probabilities for males and females.uj. 06120 Halle. Theodor-Lieser-Str.Jagiellonian University. Poland 2 UFZ . MRR produces more accurate estimates.A. M. Over 90% of the ˆ ˆ ˆ variance in Npeak /Ntotal was explained by the e to FPL ratio.pl 1 Mark-release-recapture (MRR) and transect counts are two standard methods of estimating ˆ butterfly population size (N). Settele. Estimated parameters included: ˆ (i) seasonal population size (Ntotal ). Polyommatus coridon and Zygaena carniolica) according to the Cormack-Jolly-Seber type constrained models. but at much reduced cost. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Erebia sudetica. Kühn & J. which correlate well with daily. The variation decreased considerably (CV = 0. Institute of Environmental Sciences. whereas transect counts yield only relative indices of abundance. (v) daily and seasonal sex ratio. teleius. based on the proportions of individuals captured. being captured.© PENSOFT Publishers & Josef Settele 134 Piotr Nowicki Sofia – Moscow J. but at the same time it is more laborious. Department of Community Ecology. transect counts. but also those of the total (=seasonal) population size. rebeli. e Almost always the peak sex ratio reflected the seasonal one. 30-387 Kraków. Richter A. Settele J (2005) Less input same output – simplified approach for population size assessment in Lepidoptera. (iii) intervals between capture days should correspond with the average life span of investigated butterflies (2–3 days in most cases). Woyciechowski M. Toelke U. Glinka U. Further details of this study are provided by Nowicki et al. Simulations of hypothetical simplified schemes proved that only sampling throughout the 2nd ˆ and 3rd quarter of flight period ensured 100% peak detection. REFERENCES Nowicki P. The accuracy of Ntotal estimated with the regression equation was comparable for intensive schemes (daily sampling) and non-intensive ones (sampling once in two or three days). Henle K. Popul Ecol [in press] Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). and the accuracy of e was highly ˆ sensitive to the number of capture days. (2005). and furthermore the seasonal sex ratio varied greatly so it cannot be assumed a priori. . As a conclusion the following ‘rules of thumb’ for simplified MRR sampling can be proposed: (i) sites should be checked for the presence of flying adults at the beginning and end of the flight period in order to assess its length.Simplified method of estimating butterfly population size with mark-release-recapture 135 followed a tangensoid curve rather than a sigmoid one (as a consequence the period with the sex ratio resembling the seasonal one is very long). (ii) MRR sampling should cover approximately the 2nd and 3rd quarter of the flight period and contain at least 5 sampling days. Holzschuh A. The site is approximately 3-ha wet meadow dominated by Molinia coerulea. but . Francesca Barbero2 & Emilio Balletto2 1 UJAG . 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. 30-387 Kraków. makes this system ideal for a comparative study of population dynamics of both species. teleius and M. 2005c). Populations of both species have been investigated since 1997 with intensive mark-releaserecapture (MRR). Via Accademia Albertina 13. Italy Contact:
[email protected] (White & Burnham 1999). according to mathematical models Maculinea butterflies should rather be regulated by density dependent trends (Hochberg et at. Both capture probabilities and survival were slightly.) (the model with survival and capture probability equal for both sexes and constant over time) as the best fitting the data. M.)p(. Cormack-Jolly-Seber type constrained models (Schwarz & Arnason 1996) were applied according to the procedure described by Nowicki et al. 136-139 Population dynamics in the genus Maculinea revisited: comparative study of sympatric M.Jagiellonian University. further increasing the reliability of the results. northern Italy (45°07' N. Institute of Environmental Sciences. We have decided to test the above predictions using the data provided by long-term Maculinea monitoring in Caselette near Turin. teleius). 1998).A. 1). Survival estimates obtained were within a typical range for Maculinea butterflies (Pfeifer et al. Settele. which makes population dynamics in Caselette independent of any possible effect of immigration. in particular in resources and weather patterns (Dempster 1983). However. 136 Piotr Nowicki Sofia – Moscow J. Simona Bonelli2. Moreover.uj. The data collected were analysed with the with the program MARK 2. E. nausithous.edu. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Each year sampling was conducted regularly on every second day (with very few exceptions) throughout the entire flight period. with the nearest other Maculinea sites being 2 km away. Nowicki et al. but capture probabilities proved to be relatively low (compare Meyer-Hozak 2000. pp. M. It is located in the Susa Valley. rebeli) should be more stable than those of ‘predatory’ ones (M.© PENSOFT Publishers et al. alcon. teleius Piotr Nowicki1. 1994. 2005b). arion. Model selection routine invariable suggested the model ϕ(. (2005b). Gronostajowa 7. 2000. Department of Human and Animal Biology. 07°29' E). alcon. alcon and M. Poland 2 University of Turin. which at this site fly simultaneously from mid July to late August. Sympatric occurrence of M. 10123 Turin. Kühn & J. at an altitude of ca. These models also predicted that populations of ‘cuckoo’ species (M. Nowicki et al. and are thus influenced by the same weather conditions. Thomas et al. 360 m above sea-level. the site is well isolated.pl Population dynamics of most butterfly species are believed to be predominantly affected by environmental variation. M. For both species model parameters were fairly uniform across the years (Fig. alcon (Fig. 150–1900 adults in the case of M. however the difference was surprisingly small (compare Thomas et al.Population dynamics in the genus Maculinea revisited: comparative study 137 Fig. 1998). teleius (Wilcoxon test: P = 0. consequently higher for M. 1. and ca. This may indicate either (i) their interspecific interactions through using the same . teleius.) models as revealed by the analysis of MRR data for M. As theoretically expected the population of ‘cuckoo’ M. Dynamics of both species was highly correlated (r = 0.68 respectively). reflecting difference in adult life spans (2.)p(. alcon was more stable than that of predatory M. The seasonal population size fluctuated within a range of ca.0 days for M. teleius (dark bars) on the Caselette site. teleius vs. teleius (coefficient of variation CV = 0.0–2. CV = 0. Parameter estimates (± SE) of Cormack-Jolly-Seber ϕ(.5–3. whereas difference in survival is most likely a genuine biological phenomenon.002).51 vs.91. 200–2800 adults in the case of M. 2). 2.012 in both cases). P = 0.5 days for M. Higher catchability seems to be an artefact originating from the larger sampling area for this species. alcon (light bars) and M. alcon). expected to reflect basic reproductive rate R0. The maximum year-to-year increase. teleius and 3. J Anim Ecol 63:375–391 Meyer-Hozak C (2000) Population biology of Maculinea rebeli (Lepidoptera: Lycaenidae) on chalk grasslands of Eastern Westphalia (Germany) and implications for conservation. host ant species. rebeli from a set of reproductive parameters (Hochberg et al. 2005a). Richter A. Nowicki et al. Woyciechowski M. Clarke RT. 140-143.A. pp. Glinka U. Witek M. teleius adults and 800 M.2 for M. alcon. Thin lines mark respective carrying capacity levels estimated for both species. or (ii) synchronised effect of environmental variation. e. Thomas JA (1994) Population dynamic consequences of direct and indirect interactions involving a large blue butterfly and its plant and red ant hosts. teleius (thick solid line) on the Caselette site. Woyciechowski M (2005a) Landscape scale research in butterfly population ecology – Maculinea case study. Holzschuh A. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol.g. alcon adults. Settele J (2005b) Less input same output – simplified approach for population size assessment in Lepidoptera. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Skórka P. but at the same time considerably lower than the basic reproductive rate of 6. on both species. 2. In addition. Popul Ecol [in press] . Pepkowska A. Skórka P. Settele J. Thomas JA. Kudlek J. Seasonal population size dynamics of M. Woyciechowski M (2005c) Population ecology of the endangered butterflies Maculinea teleius and M.138 Piotr Nowicki et al. Elmes GW (1992) A modelling study of the population dynamics of a large blue butterfly. Carrying capacity of the Caselette site was estimated at about 1100 M.0 respectively.3 for M. J Anim Ecol 61:397–409 Hochberg ME. These values are in excellent agreement with those recorded for the same species in the Kraków region (4. Settele. 1992). Maculinea rebeli. Fig. alcon (thick broken line) and M. nausithous. was 4. and its implications for conservation. Toelke U. In: J. the analysis of population trends indicated that in both species they were density dependent. Kühn & J.3 and 3. a parasite of red ant nests. Popul Ecol [in press] Nowicki P. REFERENCES Dempster JP (1983) The natural control of populations of butterflies and moths. Biol Rev 58:461–481 Hochberg ME. E. Elmes GW. J Insect Conserv 4:63–72 Nowicki P.2 derived for M. weather patterns. Witek M. Henle K. Nowicki P. Andrick UR. Arnason AN (1996) A general methodology for the analysis of capture-recapture experiments in open populations. London. pp 261–290 White GC.Population dynamics in the genus Maculinea revisited: comparative study 139 Pfeifer MA. Nota Lepid 23:147–172 Schwarz CJ. Chapman and Hall. Biometrics 52:860–873 Thomas JA. Clarke RT. Settele J (2000) On the ecology of a small and isolated population of the Dusky Large Blue Butterfly Glaucopsyche (Maculinea) nausithous (Lycaenidae). Symposia of the Royal Entomological Society 19. Frey W. Bird Study 46:120–138 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). . Burnham KP (1999) Program MARK: Survival estimation from populations of marked animals. McLean IFG (eds) Insect Population Dynamics in Theory and Practice. Hochberg ME (1998) Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). In: Dempster JP. Elmes GW. Joanna Kudłek. One example here may be butterflies of the genus Maculinea.005–33 ha). Foodplant patches were mapped in 2001–2002 with ca. total area of ca. alcon. E. it appears that not all species fit equally well into this theoretical framework. However. These properties are likely to lead to low population turnover in the metapopulations of Maculinea butterflies. nausithous. Poland Contact: nowic@eko. In its classic form it assumes frequent colonisation and extinction processes. 1-metre precision using GPS Magellan ProMark X. as exemplified by Hanski’s (1994) incidence function model. their dynamics and factors affecting both for the functioning of such metapopulations. M. Aleksandra Pępkowska. 1998). The study area comprised a wet meadow complex in the Vistula river valley (50°01' N.001–3 ha. here we only mention them briefly. and M. which in turn increases the importance of local population sizes.uj. Gronostajowa 7. pp. It included 61 patches of Sanguisorba officinalis – the foodplant of M. Subsequently we conducted a GIS analysis of their spatial and environmental parameters in order identify factors affecting presence/absence as well as local densities of the three Maculinea species investigated. occurring in the Kraków region. S. alcon. which are considerably smaller (range: 0. The results of this analysis are to be published in details elsewhere (e. 140 Piotr Nowicki Sofia – Moscow J. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Nowicki et al. and consequently its main interest is focused on spatial presence/absence patterns rather than on abundance patterns.Jagiellonian University. Magdalena Witek & Michał Woyciechowski UJAG . Institute of Environmental Sciences.© PENSOFT Publishers et al. We investigated spatial abundance patterns of three Maculinea species: M.A. usually reaching over 1metre height and >50% of the vegetation cover. teleius. at an altitude of 200–240 m above sea-level.25 ha) or total count on smaller ones in the case of G. 19°54' E). southern Poland.pl Metapopulation theory has become a paradigm in butterfly studies. officinalis patches there are 18 patches of Gentiana pneumonanthe – the foodplant of M. teleius and M. pneumonanthe. nausithous – with the total area exceeding 200 ha (range: 0. which – as compared to other butterflies – are characterised by relatively low mobility (Nowicki et al. Settele. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 140-143 Landscape scale research in butterfly population ecology – Maculinea case study Piotr Nowicki. Kühn & J. In the two following years they were surveyed with Braun-Blanquet method in the case of S.edu. 30-387 Kraków. officinalis. The distances between neighbouring patches are mostly within the range of 100–300 m. officinalis is the dominant plant on these patches. and sampling along 2-metre wide transects on larger patches (>0. Piotr Skórka. 8 ha) and more isolated (distances between neighbouring patches reaching 300–700 m). . Within the S. 2005b) and demographically stable populations (Thomas et al. 2005a).g. nausithous was present on 51 patches in 2003. 50 thousand individuals and slightly over 500 individuals respectively. M. pneumonanthe patches in both years of the study. nausithous and M. alcon in 2003–2004 was assessed with the standard egg count method. decreasing by about one third in 2004 (Fig. . disappearing from 4 of them by 2004. Patches lacking Maculinea populations were significantly smaller and more isolated than the remaining ones.Landscape scale research in butterfly population ecology – Maculinea case study 141 Abundance of M. All S. Nowicki et al 2005b). M. tested and parameterised on three patches intensively investigated with MRR (cf. M. Metapopulation sizes (N ± SE) of three Maculinea species in the Kraków region in 2003 (light bars) and 2004 (dark bars). The overall metapopulation size of M. whereas those of M.78–0. Population densities of M. teleius and M. pneumonanthe surveys. nausithous was estimated with the catch-per-time-unit method. roughly at peak time of both species. officinalis patches in the study area were surveyed for 1–2 hours in fine weather between 9:00 and 17:00 on 17–18 July 2003 and 26 July 2004. 1. which were available for the patches with high enough numbers of recaptures. These daily estimates were converted into seasonal population sizes on the basis of the proportion of individuals flying on survey days. 1992). In the case of M. Abundance of M. The numbers of adults per season were calculated assuming 150 eggs per female and 1:1 sex ratio (as in Hochberg et al. officinalis patches in 2003. Butterflies captured were individually marked and immediately released. teleius and M. where capture frequencies proved to be strongly correlated with butterfly densities. and it apparently managed to colonise 7 new patches in the following year. alcon was found on the same 14 G. 1). alcon there was in turn little year-to-year difference in metapopulation size estimates. which remained at the level of ca. teleius was estimated at almost 150 thousand individuals in 2003. nausithous were significantly affected by patch area (negatively) and its fragmentation (positively). teleius occurred on all 61 S.89). alcon depended predominantly on ˆ Fig. It should be noted that the estimates of daily population sizes yielded by our method were highly concordant with MRR ones (r = 0. conducted simultaneously with G. 10% individuals changing patch per season is high enough to lead to a rapid colonisation of vacant patches. and Dit+1 is its density in 2004.38). 9% and 12% per season (the difference stem from longer individual life span of M.e. This finding was consistant with low inter-patch mobility recorded. In the case of M. 2). nausithous. This allowed to estimate carrying capacity of a 1 ha S.22). nausithous. nausithous. and 3. . officinalis patch at about 1200 M. assumed to be a good approximation of basic reproductive rate R0. at least in the case of the two former species (Fig. alcon. and to derive the ‘rule of thumb’ that these carrying capacities should increase/decrease 5–6 times with ten-fold increase/decrease in patch area. for M. x = 2212 (R2 = 0. which corresponds to respectively ca. Inter-patch movements were observed only between the neighbouring patches. x = 2271 (R2 = 0. Incorporating patch area (i. Our findings confirm relative stability of Maculinea populations as well as low mobility of these butterflies. as proved by Mantel test Fig. teleius: r = 4. teleius as well as M.6% for M. with largest distances covered reaching ca. but at the same time not sufficient to even out population densities and synchronise population trends. 2. The proportion of individuals that changed patch in within 24hour period in 2003 was 4. nausithous. The curve fitted was Dit+1 = rxDit / (x + rDit).142 Piotr Nowicki et al.10). was 4. see Nowicki et al 2005b for details of the estimation). alcon the procedure was not attempted due to inadequate sample size and the fact that year-to-year changes in population density of this species were very small. The maximum year-to-year increase in population density. nausithous adults. For M.35. Plotting observed relative year-to-year changes against initial densities suggested the existence of density dependence of population trends. 350 m for M. Observed dispersal levels of ca. foodplant densities. where Dit is population density on patch i in 2003. teleius adults and 1000 M. For any of the species investigated there was no indication of spatial autocorrelation of either population densities or their relative year-to-year change (Mantel test r always well below 0.2% for M. the main factor influencing densities of both species) into the model considerably improved its fit.30.0 for M. teleius and 3. Relative changes in Maculinea population densities between 2003 and 2004 in relation to densities recorded in 2003.3 for M. often within the confidence limits of yearly estimates. teleius and 200 m for M. nausithous: r = 4. Clarke RT. Kudlek J. Brasilia. Maculinea rebeli. Skórka P. Elmes GW (1992) A modelling study of the population dynamics of a large blue butterfly. a parasite of red ant nests. Settele J. pp.Landscape scale research in butterfly population ecology – Maculinea case study 143 results. but not for M. teleius and M. Popul Ecol [in press] Thomas JA. pp 261–290 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). and its implications for conservation. McLean IFG (eds) Insect Population Dynamics in Theory and Practice. regulated by density dependence trends. nausithous. Woyciechowski M (2005a) Abundance patterns of Maculinea butterflies at landscape scale – implications for metapopulation conservation. 151 Nowicki P. In: Constantino R. Witek M. Goedert D (eds) Conservation biology capacity building and practice in a globalized world. Woyciechowski M (2005b) Population ecology of the endangered butterflies Maculinea teleius and M. nausithous. Abstracts of XIX annual meeting of the Society of Conservation Biology. J Anim Ecol 61:397–409 Nowicki P. alcon. REFERENCES Hanski IA (1994) A practical model of metapopulation dynamics. Skórka P. Witek M. and less likely to become (re)colonised. Pepkowska A. In: Dempster JP. Thomas JA. This may lead to a conservation dilemma of whether to opt for stable systems or higher butterfly numbers. for which patch size and shape matter the most. Elmes GW. Local populations appear thus to act as independent demographic units. J Anim Ecol 63:151–162 Hochberg ME. Foodplant availability is the main factor limiting population densities for M. University of Brasilia. . Hochberg ME (1998) Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). small and isolated patches are more likely to experience population extinctions. On the other hand. Small and highly fragmented patches may support higher densities of the two latter species. Chapman and Hall. Symposia of the Royal Entomological Society 19. London. Poland 2 UFZ .5 times per year) are considerably lower than the basic reproductive rate values used in HTE and HCET models developed for M. 144-149 A review of population structure of Maculinea butterflies Piotr Nowicki1. M. which are more efficient in using their Myrmica ant hosts. arion. should perhaps be regarded as conspecific according to their genetic analysis (Als et al. M.edu.A. however proterandry reduces the effective population size a little (Nowicki et al. 2005c). As expected from evolutionary theory. pp. are more stable than those of predatory species (Table 2). E. 144 Piotr Nowicki Sofia – Moscow J. Dorchester DT2 8ZD. 2005c. only a fraction of individuals flies and can mate on any day of the season. Even though two of them. Their sex ratio is either balanced or slightly female biased (Nowicki et al. which have longer life spans and often shorter flight periods (Table 1). 2005d). 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. particularly within the MacMan project. i. 06120 Halle. alcon. Josef Settele2. Kühn & J. Nowicki et al. However. Maculinea butterflies typically occur in medium-size populations of several hundred individuals. 1998). rebeli).uj. M. The proportion of individuals flying on peak day ranges between 11 and 46% (Nowicki et al. 30-387 Kraków. 2005c). the populations of the ‘cuckoo’ Maculinea.© PENSOFT Publishers et al. Dorset Laboratory. Theodor-Lieser-Str.Centre for Environmental Research Leipzig-Halle. Our aim is thus to present the current state of knowledge of the population ecology of all five European Maculinea species. Due to the fact that adult life span is very short compared to flight period length.e. Centre of Ecology and Hydrology. Germany 3 NERC – Natural Environment Research Council. 2005d).Jagiellonian University. M. The latter is further negatively affected by the temporal fragmentation of populations. rebeli. Nevertheless. but larger populations are not uncommon (Munguira and Martin 1999. Gronostajowa 7. Interesting- . rebeli (Table 2). Thomas3 & Michal Woyciechowski1 UJAG . Jeremy A. The temporal fragmentation is likely to be stronger in wet meadow species (M. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 2004) we have decided to consider them separately for the sake of consistency with the references cited as well as for practical purposes as the two differ considerably in many aspects of their ecology. Institute of Environmental Sciences.pl 1 Munguira and Martin (1999) provided the first comprehensive review of the population structure of Maculinea butterflies. nausithous. 4. over recent years there have been a lot of studies in this field. alcon and M. even the latter group experience smaller demographic fluctuations than most other butterfly species (compare Thomas et al. Nowicki et al. Settele. UK Contact: nowic@eko. teleius) than in dry meadow ones (M. Department of Community Ecology. The maximum growth rates observed in all Maculinea species (2–4. Winfrith Technology Centre. The considerable difference in adult life expectancy between such closely related species could possibly be explained by generally more favourable summer weather on dry habitats. 80–0.1 2.5–4. (2005a) Kassai et al.86 M. Characteristics of adult stage of Maculinea butterflies as revealed by MRR studies.59–0.4 3.2–5.7 days). (2005c) Settele et al. The authors state 2.7 3.0–2. (2005c) Nowicki et al.3 a 2.5 3.74 a M.64–0.0 4. (unpubl.69 0. (2005c) Nowicki et al.) Kassai et al. (2000) Nowicki et al. (2005 a) M. teleius 0.) Pfeifer et al.5–6.84 day-1) and longer adult life span (5. All the life span values are based on MRR model estimates.75–0. 145 .75 M.74 0.59–0. c The value is slightly underestimated since the studies missed the very beginning of the flight period.) Meyer-Hozak (2000) Nowicki et al.2–3.66 M.0–3.72 0. arion 0.3 days. (unpubl. which must be either misprint or miscalculation as they report proper formula used for the calculation.5 3.Table 1.5–3. alcon 0. nausithous 0.77 0. rebeli 0.58–0. 2000) are not included as they are obviously underestimated and dependant on sampling effort.77 a A review of population structure of Maculinea butterflies b Single population had much higher survival rate (0.9–2.63–0. the values based on observed residence time (see review in Pfeifer et al.0 40 23–40 37–48 28–42 25 c 20–31 21–29 24–31 18–36 15–30 c Survival rate [day-1] Individual life span [days] Flight period length [days] Source Nowicki et al.82–0.5 1.3 b 2. Species 2.5–3. (unpubl. Derived from a set of reproductive parameters.3 ca. teleius 0. 1994). alcon 0. (2005a) Nowicki et al.55 a 0. arion 0. Population dynamics parameters of Maculinea butterflies. 1000 ca.2–21 b 2. 400 ca.) Thomas (1995). c The level of 450 adults/ha was reached after intensive management. 450–650 ca. unless stated otherwise. (1998) Nowicki (unpubl. Thomas et al. 20–450 ca.0 6. nausithous 0. Carrying capacity has been scaled to 1-hectare habitat patch.3 4. while basic reproductive rate has been approximated by the maximum recorded year-to-year growth rate. unpubl. Table 2. 1200 ca.2 3. Thomas et al. (1992.51 M. 300 Population size variability Basic reproductive rate (R0) Estimated carrying capacity (K) [adults/ha] Source Nowicki et al.57 Hochberg et al. (2005a) Nowicki et al.68 a b Based on simulation results rather than empirical data. 50–60 c ca.2 4.92 M. (2005b) M. (1998) Griebeler and Seitz (2002) Nowicki et al.51 M.146 Piotr Nowicki et al.3 4. . (2005b.) Nowicki et al. rebeli <0. Species 3. Population size variability is expressed as coefficient of variation (CV) of population size time series.3 3. (2005b) M. nausithous 8 5–24 12 M. (2005b) Binzenhöfer and Settele (2000) Nowicki et al. 50 a M. (2002) Kockelke et al. (1996) Pajari (1992) Binzenhöfer and Settele (2000) Nowicki et al. alcon ca.) Pauler-Fürste et al. (2005d) Nowicki et al.1 ca. 2 100–300 200–400 5. (2005b) M. (2005d) Nowicki et al. (1994. (1996) Benes et al. rebeli 2–5 M. Species 50–300 ca.Table 3. Mobility parameters of Maculinea butterflies. teleius 14 6–24 9 A review of population structure of Maculinea butterflies a The value is seriously overestimated since many movements recorded by the authors should in fact be treated as intra-site one 147 . unpubl.7 100–400 80–300 100–200 200–400 80–300 100–350 2. 3 % individuals changing site Typical distance of inter-site movements [m] Maximum recorded distance of inter-site movements [km] Source Wynhoff et al. arion <1 M.4 5. although maximum recorded distances of movements are in the range of 2–6 km. Wantuch 2005). UFZBericht 2:1–98 Elmes GW. Mignault AA. 1779) und Maculinea teleius (Bergstr. It is also likely that demographically stable ‘cuckoo’ species hardly ever reach very low densities. especially from small sites containing optimum habitat (Elmes and Thomas 1987. Clarke RT. Weidenhoffer Z (2002) Butterflies of the Czech Republic: Distribution and conservation I.g. and thus growth rates observed in their populations in the field will be far below the achievable (intrinsic growth) levels. 2005c).. nausithous and M. teleius. The exchange of individuals between local populations obviously depends on their spatial setting. Settele J (2000) Vergleichende autökologische Untersuchungen an Maculinea nausithous (Bergstr. alcon that combined egg count and MRR of adults (e. Maculinea may live in small isolated populations. Pavlicko A. rebeli. Fric Z.: Lycaenidae) im nördlichen Steigerwald. In addition. The highest carrying capacities have been estimated for M. Thomas JA. Vila R. Pierce NE (2004) The evolution of alternative parasitic life histories in large blue butterflies. such long-distance movements are in fact extremely rare. Nowicki et al. Basel.. 1994) was originally parameterised from a single site. This high fecundity. However. Kandul NP. REFERENCES Als TD. 1998). For all Maculinea species a great majority of inter-site dispersal is limited to less than 500 m (Table 3). Hsu Y-F. due to the high density-dependent mortalities experienced by larvae both on the foodplant and in ant nests (Hochberg et al. Schweizerishes Bund für Naturschutz. Acta Oecol 17:61–80 Elmes GW. whereas the lowest for M. Seitz A (2002) An individual based model for the conservation of the endangered Large Blue Butterfly. mainland-islands systems or typical metapopulations. II. (1992. (2005b) that site area negatively affects its carrying capacity. we found no indication of higher growth rates for ‘cuckoo’ species. and life table counts of larval survival starting with >1000 eggs. and their high values probably reflect the optimum conditions of habitat quality and climate experienced in these centre-of-range populations. Elmes at al. but it only sporadically exceeds 20% (Table 3). which is agreement with the finding of Nowicki et al. compared with those nearer edges of range (where the most acute conservation problems exist). These parameters were confirmed over 5 years in several other Myrmica schencki using populations of M. Meyer-Hozak 2000. Konvicka M. 1998). SOM. 1779) (Lep. It is important to note that densities exceeding our estimated carrying capacities have been frequently reported elsewhere. and the higher intrinsic growth rate reported by Hochberg et al. from a combination of mark-release-recapture (MRR) adult estimates and absolute egg counts.) Tagfalter und ihr Lebensraum. Havelda Z. as modelled by Thomas et al (1998). 1992. arion (Table 2). Ecol Model 156:43–60 . Dvorak J. even in metapopulations they are characterised by low mobility. pp 354–368 Griebeler EM. In: Geiger W (ed.148 Piotr Nowicki et al. 1996. Thomas JA (1987) Die Gattung Maculinea. Nature 432:386–390 Benes J. Boomsma JJ. Nash DR. Thus this high fecundity was not confirmed in later studies on either this species or M. Praha Binzenhöfer B. ly. Hochberg ME (1996) Empirical tests of specific predictions made from a spatial model of the population dynamics of Maculinea rebeli. a parasitic butterfly of red ant colonies. Vrabec V. despite their reportedly twice-higher female fecundity (Thomas et al. Maculinea arion (Lepidoptera: Lycaenidae). which is possibly connected with different ant host abundance. Yen S-H. Thomas et al. a parasite of red ant nests. Popul Ecol [in press] Pajari M (1992) Muurahaissinisiiven (Maculinea arion (L. Kudlek J. J Anim Ecol 63:375–391 Kockelke K. pp 261–290 Wantuch M (2005) Ekologia populacji motyli z rodzaju Maculinea w Poleskim Parku Narodowym. Elmes GW (1992) A modelling study of the population dynamics of a large blue butterfly. Kaule G. pp 275–281 Pfeifer MA. MSc. Nota Lepid 23:147–172 Thomas JA (1995) The ecology and conservation of Maculinea arion and other European species of Large Blue Butterfly. van der Made JG (1996) Effects of habitat fragmentation on the butterfly Maculinea alcon in the Netherlands. Settele J (2000) On the ecology of a small and isolated population of the Dusky Large Blue Butterfly Glaucopsyche (Maculinea) nausithous (Lycaenidae). Henle K (eds. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Pepkowska A. Thesis. Poschlod P. . 1758) (Lepidoptera: Lycaenidae) in southwest Germany.A. Chapman and Hall. MSc. 140-143. London. In: Settele J. Symposia of the Royal Entomological Society 19. Thesis. In: J. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Elmes GW. Kluwer. E. Kaule G. Settele J (2005c) Less input same output – simplified approach for population size assessment in Lepidoptera. Holzschuh A. Pauler-Fürste R.)) populaatiokoon arviointi ja habitaatiivaatimusten tutkiminen kesällä 1990 Pohjois-Karjalan Liperissä. 97. Wynhoff I. and its implications for conservation. Woyciechowski M (2005d) Population ecology of the endangered butterflies Maculinea teleius and M. Oostermeijer JGB. Kühn & J. Balletto E (2005a) Population dynamics in the genus Maculinea revisited: comparative study of sympatric M. Skórka P. Settele. London. Dordrecht. Kluwer. Nowicki P. Thomas JA. Verhaagh M. nausithous. Richter A. Nature and Environment No. Henle K. alcon and M. Elmes GW. pp. pp. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Andrick UR. Toelke U. Strasbourg Nowicki P. Bonelli S. Clarke RT. Jagiellonian University. Settele J (1996) Aspects of the population vulnerability of Glaucopsyche (Maculinea) arion (Linnaeus. Clarke RT. Settele J (1994) Zur Autökologie und Verbreitung des Kreuzenzian-Ameisenbläulings. Scheper M. Kraków. Witek M. Woyciechowski M (2005b) Landscape scale research in butterfly population ecology – Maculinea case study. Glinka U. McLean IFG (eds) Insect Population Dynamics in Theory and Practice. Council of Europe Publishing. 136-139.) Species survival in fragmented landscapes. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. teleius. In: Settele J. Settele J. 1904).A. J Anim Ecol 61:397–409 Hochberg ME. Maculinea rebeli. Thomas JA (1994) Population dynamic consequences of direct and indirect interactions involving a large blue butterfly and its plant and red ant hosts. Martin J (eds) (1999) Action Plan for Maculinea Butterflies in Europe. Skórka P. In: J. Popul Ecol [in press] Nowicki P. Kühn & J. Poschlod P. Frey W. Dordrecht. Hermann G. Settele. Margules C. Woyciechowski M. Nowicki P. In: Pullin AS (ed) Ecology and Conservation of Butterflies. Henle K (eds. Hochberg ME (1998) Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). pp 180–197 Thomas JA. Chapman and Hall. J Insect Conserv 4:63–72 Munguira ML.) Species survival in fragmented landscapes.A review of population structure of Maculinea butterflies 149 Hochberg ME. Witek M. E. Margules C. University of Joensuu. In: Dempster JP. pp 15–23 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Carolinea 52:93–109 Meyer-Hozak C (2000) Population biology of Maculinea rebeli (Lepidoptera: Lycaenidae) on chalk grasslands of Eastern Westphalia (Germany) and implications for conservation. Maculinea rebeli (Hirschke. Barbero F. nausithous in the Kraków region (50o01' N. Skórka P. p. Gronostajowa 7. Sofia – Moscow J. Institute of Environmental Sciences. Kühn & J. . female-biased for both species. 30-387 Kraków.edu. The results indicate that populations of both species are typically stable within their sites. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol.© PENSOFT Publishers 150 Magdalena Witek et al. Poland Contact: nowic@eko. Magdalena Witek. rebeli has been thoroughly investigated through both empirical and modelling studies. E. nausithous. We present the findings of a two-year research on sympatric populations of M. (2005). despite rather short distances (ca. but consistently. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. southern Poland. There was little year-to-year change in their seasonal population sizes. but in contrast less has been known so far about population ecology of predatory Maculinea. The study comprised intensive mark-release-recapture sampling as well as laboratory raising of the former species. nausithous in the Kraków region. The sex ratio was slightly. Woyciechowski M (2005) Population ecology of the endangered butterflies Maculinea teleius and M. Average life span of laboratory raised butterflies kept in ideal conditions was more than 6 days.Jagiellonian University. Piotr Skórka & Michal Woyciechowski UJAG . whereas daily numbers showed higher variation between the two years of the study apparently due to the differences in daily survival rate. southern Poland Piotr Nowicki. Further details of this study are provided by Nowicki et al. Settele. Settele J. Witek M. 150 Structure of sympatric populations of M.uj. The recruitment of both males and females consistently followed a bimodal pattern.pl Among Maculinea butterflies the cuckoo species M. 100 m) separating them. 19o54' E). 25%) changed sites. but it was only 2–3 days in the field. REFERENCES Nowicki P. teleius and M. but their effective population sizes are reduced by ‘temporal fragmentation’ cause by their very short life span in relation to flight period length. and its implications for conservation. teleius and M. more precisely. high site fidelity) and thus landscape corridors seem necessary to enhance metapopulation viability.A. Popul Ecol [in press] Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Relatively small fractions of individuals (max. A further problem for the conservation of Maculinea butterflies is their low inter-site mobility (or. possibly indicating the existence of cohorts of one-year and two-year developers among larvae. The survival was negatively affected by rainy weather. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Theodor-Lieser-Str. E. Department of Community Ecology. Germany 2 UFZ . (2005). Uta Glinka2 & Josef Settele2 UFZ-Centre for Environmental Research Leipzig-Halle. the total population size of adults is only the reproductive output of the former generation(s). For a population of the Dusky Large Blue butterfly (Maculinea nausithous) the size of the egg population was estimated. This was compared with results of a MRR (mark-release-recapture) study – a frequently used approach for the study of adult butterfly populations. Permoserstr. Kühn &Poland (Eds)151 Studies and M. preimaginal stages may also be used in many cases. Glinka U & Settele J (2004): Die Schätzung von Populationsgrößen bei Tagfaltern anhand von Präimaginalstadien am Beispiel von Ameisenbläulingen (Lepidoptera: Lycaenidae: Maculinea). 15. The efficiency of the two methods is question dependant. because it is the reproductive output of the population which will found the next generation and therefore might be more relevant for some ecological studies. Thomas 2005 and ecology of Maculinea teleiuson the EcologySettele. .© PENSOFTThe distribution Publishers Sofia – Moscow J.de 1 Sizes of butterfly populations are mainly estimated in the adult stages. 151 Estimation of butterfly population sizes using pre-imaginal stages exemplified by Maculinea butterflies (Lepidoptera: Lycaenidae) Manfred Alban Pfeifer1. Nevertheless. 04318 Leipzig. Mainzer naturwiss. Archiv 42: 225-244. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Department of Conservation Biology. nausithous in of Butterflies in Europe and Conservation Vol. The knowledge of the egg population size is advantageous. 06120 Halle.A. Detailed results have been published in the German language by Pfeifer et al. Germany Contact: Josef.Centre for Environmental Research Leipzig-Halle. J. 4. In contrast. but first results indicate that eggassessment might be a good alternative to MRR studies of adult populations.Settele @ufz. p. REFERENCE Pfeifer M A. yielded better estimates of the presence or absence of the species than the transect counts. Marcin Sielezniew & Anna M. 2005). 06120 Halle. comparisons of two different assessment methods were conducted between 1989 and 2005. Permoserstr. 15. Kühn & of Butterflies(Eds) 2005 Studies on the Ecology and Conservation in Europe Sofia – Moscow Vol. The heads have been collected and the M. Department of Conservation Biology.A. all flowerheads were dissceted. 4. After a certain period. StankiewiczSettele. Department of Community Ecology. Steiner R. Martin Musche1 & Anett Richter2 UFZ-Centre for Environmental Research Leipzig-Halle.Centre for Environmental Research Leipzig-Halle. Transect counts during the peak flight period have been compared to random samples of Sanguisorba flower heads. the sampling of flower heads. 152 Assessing the presence/absence of Maculinea nausithous: a comparison of adult and pre-imaginal stages Josef Settele1. Sarah Gwillym1. Thomas © PENSOFT Publishers 152 Jarosław Buszko. nausithous larvae which left the heads within a few days after sampling were counted (and then used for genetic analyses). Therefore this method also has been recommended for standard assessments (Settele et al. 256pp Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). even if it was only a very small fraction of the total present on a patch. Furthermore its big advantage is the weather independence and the short stay needed in the field. Uta Glinka1. Stuttgart: Verlag Eugen Ulmer. Holger Loritz1. Reinhardt R. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Christian Anton1. Theodor-Lieser-Str.de 1 During two long term studies on Maculinea nausithous in Germany (one in the Upper Rhine Valley and one in the area around Leipzig). p. Feldmann R (2005): Schmetterlinge – Die Tagfalter Deutschlands.settele@ufz. Thus collecting flowerheads allows many more sites to be sampled per unit time and thus gives a better picture of the distribution in a particular season.J. Germany 2 UFZ . J. E. 04318 Leipzig. Germany contact: Josef. We found that in the vast majority of cases. . REFERENCE Settele J. Kühn &Poland (Eds)153 Studies and M. These sites were called A. A similar pattern was apparent in M. We did not found significant differences between species. although there was tendency in the case of females of M.teleius on site B to cover larger distances than males. For recaptured individuals we measured body weight. In the case of males of M.pl We studied within.patch migration events. Maximum distance covered by M.07) in M. two of which were statistically significant). Maximum distance covered within the site was always lower then maximal dimension of the habitat. We found that the number of migration events was statistically larger in the second half of the season (after 22. 8 ha.onet. and the lowest fraction on site B – the largest and most isolated one. teleius on two sites (A and B). J. 30-387 Kraków. E. During the flying period of butterflies we made markrelease-recapture studies. We found that average distances between captures depended on the patch area. Institute of Environmental Sciences. teleius.Jagiellonian University. wing length and thorax width. B and K and were about 1ha. We detected only 37 between. teleius and M. p. We did not find significant differences between sexes of either species. however M. Poland Contact: skorasp@poczta.© PENSOFTThe distribution Publishers Sofia – Moscow J. Aleksandra Pępkowska. Gronostajowa 7. We found a negative correlation between body weight and distances covered (from six correlation tests. Thomas 2005 and ecology of Maculinea teleiuson the EcologySettele. nausithous in of Butterflies in Europe and Conservation Vol. respectively. We found that the largest fraction of emigrants and immigrants was on the smallest site K for both species. southern Poland. Moreover site B was isolated from others by forest and built areas. teleius seemed to cover larger distances than M. teleius between sites was 706 m and 602 m in the case of M. Joanna Kudłek. Magdalena Witek & Michal Woyciechowski UJAG . As far as within-patch movements are concerned. Piotr Nowicki. nausithous on three sites in Kraków region. and 0. Most of butterflies covered small distances up to 50 m. .A. Butterflies covered larger distances within larger sites. We found the tendency for butterflies with longer wings to cover larger distances (a trend found in four correlations out of 6. nausithous Piotr Skórka. nausithous.6 ha in area. five showed this tendency was detected. marked them and recorded their position with GPS.and between patch mobility of M. We captured individual butterflies. We did not find differences in migration tendency between sexes of the two species. Ewa Sliwinska. We did not found any significant correlation between distances covered and butterfly thorax width. we found that the distribution of distances between captures was strongly positively skewed. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). we found a significant positive correlation between the progression of the season and distances covered. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Data presented here come from the 2003 season. with two being statistically significant). nausithous. nausithous. 153 Mobility patterns of Maculinea teleius and M. 5 cm). P < 0. Myrmica ruginodis: F1. we slightly opened its uppermost part. After ten seconds the stick was put into a plastic bag to prevent ants from escaping and immediately another stick was put into the nest.668. Ant workers have a tendency to climb up on wooden sticks put into their nests. Institute of Environmental Sciences. temperature at the nest and site (a categorical variable). 2006.. therefore.833. 30-387 Kraków. nest was found. From 25 nests tested.5x3. Myrmica scabrinodis: F1. M. Gronostajowa 7. Studies on the Ecology and Conservation in Europe Sofia – Moscow Vol. pp. We used a general regression model to perform a backward stepwise elimination of explanatory variables. We found 27 nests of Myrmica rubra. holding grasses in hands. Interestingly. “A simple and nondestructive method for estimation of worker population size in Myrmica ant nests” which will be published in Insectes Sociaux. P < 0. the backward stepwise procedure of GRM led to final models with only the number of removed workers as a significant explanatory variable (Myrmica rubra: F1. 154-155 A simple method for estimating worker population size in Myrmica ant nests Piotr Skórka.076.onet.Jagiellonian University. Then a stick was placed vertically into the nest. 3 disappeared as well (Fisher exact test P > 0.pl This abstract is a short summary of the paper: Skórka P. Figure 1). R2 = 0.001. assuming that the number of workers removed on sticks is related to the total number of workers within the nests. Once a Myrmica spp. Witek M. R2 = 0. Settele. The method can be a very useful tool in population studies of .663. Immediately after the series of removals the entire nest was excavated and put into a plastic bowl. Woyciechowski M. Magdalena Witek & Michal Woyciechowski UJAG . NS). The method is apparently nondestructive as we did not observe decreased survival of nests surveyed as compared to control nests. A method to estimate the number of workers in Myrmica ant nests on abandoned meadows was developed based on removal of workers. The number of workers removed on sticks.001. regression models for Myrmica rubra. These were the number of workers removed on sticks.J.19 = 101.25 = 52. even when the data was pooled for all species. ruginodis and M. Kühn & of Butterflies(Eds) 2005 154 Piotr Skorka. Thomas © PENSOFT Publishers Magdalena Witek & Michal Woyciechowski J. width 3. scabrinodis were built. Ant workers have a tendency to climb up on the stick. 22 nests of Myrmica ruginodis and 76 nests of Myrmica scabrinodis. For all species.A. To estimate the number of workers in the ant nest we carried out series of six removals of workers from the nest using wooden sticks (length: 20 cm. This procedure was repeated six times in every nest. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Poland Contact: skorasp@poczta. until the first chambers with larvae could be reached. and the number of ants in the excavated nest were counted directly in the field.05.74 = 171. R2 = 0.001.014. the variance explained by the simple regression was still high R2 = 0.. E.394. P < 0.694. only 3 disappeared and from 28 control nests. where ants are used as bioindicators. Y = 1. ants as well as in biodiversity projects. The relationship between number of workers removed from the nest and total number of workers present in the nest (all data ln-transformed) in three Myrmica species. rubra (dashed regression line. Y = 3.431 + 1.712X). Y = 2. ruginodis (dotted regression line. 1. .370X).A simple method for estimating worker population size in Myrmica ant nests 155 Fig. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). The method could give good estimates of mean nest size at the population level and may be useful in conservation of Maculinea butterflies.870 + 0. a) M. c) M.489X). b) M.432 + 1. scabrinodis (solid regression line. Magdalena Witek & Michal Woyciechowski This page intentionally left blank .156 Piotr Skorka. 4. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Population genetics and physiology of Maculinea and Myrmica ants .Geographical versus Food Plant Differentiation in Alcon Blue Populations 157 Section 3. 158 Judit Bereczki. Katalin Pecsenye & Zoltan Varga This page intentionally left blank . In the analysis of the structure of genetic differentiation. Faculty of Sciences. P. Imagos were collected from 11 localities in four subregions of Northern Hungary between 2000 and 2003.A. pp. At the same time. H-4010 Debrecen.O. G. In the analysis of the data. The average FIS value suggested significant heterozygote deficiency within the samples. F-statistics was computed and the total genetic variation was partitioned into within and between population components. The within population variation was also sizeable. Nei’s genetic distances were calculated and UPGMA dendrogram was constructed on the basis of the distance matrix. Enzyme polymorphism was analysed at 16 enzyme loci using polyacrylamide gel electrophoresis. however.B. First we computed F-statistics on restricted data sets. pneumonanthe). the aim of the present study was to analyse the pattern of genetic differentiation among the Alcon Blue populations at a smaller geographic scale. When the samples were grouped according to the food plant they used. A substantial portion of this variation. The results indicated a high level of genetic differentiation among the samples.g. 3. H-4010 Debrecen. That is.O. Katalin Pecsenye1 & Zoltán Varga1. This implied that the geographic scale of the survey was too large to detect the genetic consequences of migration that is the geographic pattern of genetic variation. which did not exhibit an evident species or geographic pattern. 3. 2 1 DUFS . Hierarchical F-statistics and AMOVA was computed to study the pattern of genetic differentiation among the samples. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group.hu In our previous study we have analysed the genetic structure of several Alcon Blue populations in Central Europe. The samples were significantly differentiated at all but two of the 16 loci. Thus. was observed among the samples. PCA analysis was also carried out using the allele frequencies of the samples. Hungary Contact: pecskati@tigris. the level of differentiation was as high among the pneumonanthe type populations as it was in the entire data set (Table 1: Total vs. we first contrasted the “food plant” and the geographic pattern of the between sample variation in Northern Hungary. 159-162 Geographical versus food plant differentiation in Alcon Blue Populations (Lepidoptera: Lycaenidae) in Northern Hungary Judit Bereczki1.University of Debrecen. To avoid “a priori” taxonomical preconclusions we distinguished pneumonanthe and cruciata type populations and had samples from both types. the FST . Settele. Department of Evolutionary Zoology and Human Biology. within one region of Central Europe.klte. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. E. Thomas 2005 versus Food Plant Differentiation in Alcon Blue Kühn & of Butterflies(Eds)159 Populations in Europe Studies on the Ecology and Conservation Vol. e.© PENSOFT Publishers Geographical Sofia – Moscow J. P. the level of genetic differentiation was fairly high. The results of F-statistics indicated a relatively high level of polymorphism.B. J. 024** 0. FST values computed on restricted data sets according to the two types of hierarchy of the samples: food plant vs. Northern Hungary – WSR: within subregion component. A series of hierarchical analyses of genetic variance was computed by Arlequin. Most of the total genetic variation was attributable to the within sample component both in Central Europe and in Northern Hungary (Figure 1).057** FST % 84.082** 0. The results were compared with those obtained for Central Europe.159** 0.011** 0. BS: between samples component. subregions. The distribution of the total genetic variation at different levels of the hierarchy. . Katalin Pecsenye & Zoltan Varga Table 1. Groups of populations Value Total (21) Food plant Region G. The distribution of the between sample variation was. pneumonanthe (12) G. cruciata (9) Karst (5) Bükk (4) Mátra (5) Zemplén (5) 0. %: portion of loci exhibiting significant differentiation among the samples. BR: between region component.160 Judit Bereczki. Central Europe – WR: within region component. These results suggest that the genetic structure of the samples in Northern Hungary exhibited some geographic pattern. ** significant at 0.041** 0. Karst or Bükk or Mátra or Zemplén). BSR: between subregion component.7 90 30 20 10 50 25 values together with the portion of the differentiating loci estimated for the subregions separately were always substantially lower compared to the total (Table 3: Total vs. however different in the two Figure 1. Numbers in brackets: samples in the given group. WS: within samples component.208** 0.01 level. The 95% ellipses show 5 subregions. . gene flow was not effective enough in counterbalancing genetic differentiation. The results of the PCA analyses fully confirmed that the results of the two investigations with different geographic scale were significantly different. AMOVA and hierarchical F-statistics fulfilled our expectations. among populations within a subregions). The results of PCA computed for the M. The differentiation among the subregions in Northern Hungary was as high as that among the populations within the subregions.Geographical versus Food Plant Differentiation in Alcon Blue Populations 161 analyses. The 95% ellipses indicate four large regions. Thus. B: Samples originated from Northern Hungary. therefore. Hence we concluded that the geographic distances among these regions were too large relative to the dispersal ability of the butterflies and. The results of PCA. In Central Europe. In contrast. in Northern Hungary the differences between the subregions explained a substantial portion of it (Figure 1). the geographic pattern of differentiation was not evident. In the present study. While in Central Europe. the variation among the regions only accounted for a low proportion of this variation. the pattern has become much clearer in Northern Hungary. alcon samples at different geographic scale. the 95% ellipses drawn according to the geographic regions of the samples were practically overlapping (Figure 2A).g. The 95% ellipses indicated a certain level of differentiation among the subregions (Figure 2B) Although the sampled populations could be sorted into four large regions in Central Europe. A: Samples originated from Central Europe. we expected certain geographic pattern of genetic variation within the Northern Hungarian region. our working hypothesis was that migration must be more intensive at a smaller geographic scale (e. This indicates that the geographic distances among the populations within these Figure 2. Katalin Pecsenye & Zoltan Varga subregions are comparable with the long distance migration of the Alcon Blues.162 Judit Bereczki. . Namely. are not differentiated clearly from the cruciata type ones of the region. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). the population is pneumonanthe type but the individuals have a “rebeloid” appearance. it is worth noting that the pneumonanthe type populations of the Zemplén Mts. Nevertheless. which is in agreement with the phenotype of the individuals in these populations. The results of F-statistics also indicated a higher level of polymorphism in the cruciata type populations compared to the pneumonanthe type ones (Table 2: FIT). they are exposed to strong bottle neck in every generation. Nei’s genetic distances were calculated and UPGMA dendrogram was constructed on the basis of the distance matrix. Enzyme polymorphism was analysed at 16 enzyme loci using polyacrylamide gel electrophoresis. Kühn & J. which was coupled with a sizeable amount of heterozygote deficiency (Table 1). pp.A. 163-166 Temporal and spatial structure of genetic variation in the Alcon Blue (Lepidoptera: Lycaenidae) populations in Northern Hungary Judit Bereczki1. 3. our goal was to contrast the level of genetic differentiation between nearby populations (spatial pattern) and among consecutive generations within a population (temporal pattern). alcon. and 5 samples from two cruciata type populations in the Bükk Mts.O. To avoid “a priori” taxonomical preconclusions we distinguished pneumonanthe and cruciata type populations. PCA analysis was also carried out using the allele frequencies of the samples. their larvae develop in the nest of various Myrmica species. and two cruciata type populations in the Bükk Mts. H-4010 Debrecen.University of Debrecen. Thomas 2005 structure of genetic variation in on the Alconand Conservation of Butterflies(Eds)163 Studies the Ecology Blue populations in Europe Vol. Hierarchical Fstatistics and AMOVA was computed to study the pattern of genetic differentiation among the samples.B. As a consequence of this special life cycle. The level of genetic differentiation was slightly higher among the samples of the pneumonanthe type populations of the Zemplén Mts. had a slightly lower level of heterozygote frequency. The samples of the two subregions exhibited a similar level of polymorphism. Hungary Contact: pecskati@tigris. H-4010 Debrecen. Owing to the repeated bottle necks Maculinea populations are exposed to the effect of genetic drift.hu Alcon Blues are obligate myrmecophylous species. and Bükk Mts.B. We had 7 samples from two pneumonanthe type populations in the Zemplén Mts. we were able to analyse the structure of genetic variation in both ecotypes of M. Nevertheless. E. P.) of Northern Hungary between 1999 and 2003.klte. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. In this way. which has a temporal and spatial dynamics as well. We have chosen two pneumonanthe type populations from the Zemplén Mts. The average FIS values suggested significant heterozygote deficiency within the samples in both subregions. Katalin Pecsenye1 & Zoltán Varga1.O. Accordingly. In the analysis of the data. than among the cruciata type ones of the Bükk Mts. 3. 2 1 DUFS . the cruciata type populations in the Bükk Mts. F-statistics was computed and the total genetic variation was partitioned into within and between population components. Department of Evolutionary Zoology and Human Biology.© PENSOFT Publishers Temporal and spatial Sofia – Moscow J. Settele. P. Imagos were collected from 4 localities in two subregions (Zemplén Mts. . Faculty of Sciences. nA: mean number of alleles per locus.7 39.091* 0. A: distribution of the total genetic variation. FIS: within sample component of variation. Katalin Pecsenye & Zoltan Varga Table 1.122 0. H: mean frequency of heterozygotes. 1.45 1.044** Fig. Results of F-statistics computed for the samples of the two subregions separately.1 H 0.174** In order to analyse the structure of genetic differentiation.091* 0. B: distribution of the between sample variation.210** FIS 0.01. Parameters of enzyme polymorphism in the Alcon Blue populations averaged for the two regions. Subregion Zemplén Bükk FIT 0. The results suggested that most of the total Table 2. FIS: index representing the within sample variation Subregion Zemplén Bükk nA 1. .174** FST 0. FST: between sample component of variation. **: significant at 0. P: percentage of polymorphic loci. we contrasted the spatial and the temporal components of the total between sample variation in the two subregions separately. Results of AMOVA in the two subregions.164 Judit Bereczki. In the analysis of molecular variation (AMOVA).091 FIS 0. FIT: total genetic variation.158** 0. the total genetic variation can be analysed at the following levels of the hierarchy: within years/generations.5 P 34.05 level. *: significant at 0. between years/generations within a population and between populations in a subregion.073** 0. Each point stands for an individual in the reduced space of variables. some groups of individuals could be detected in all populations (Figure 2). Different symbols represent individuals of consecutive generations. the points representing the individuals comprised a diffuse cloud of points in the reduced space of variables. . Ellipses indicate groups of individuals with similar multilocus genotypes. In the last part of the study. Acon. The individuals of the different generations were not clearly differentiated (Figure 2). Aox. Nevertheless. In all four populations. Results of the PCA analyses. carried out a series of PCA analyses where the genotypic composition of the individuals was involved. we analysed the fine structure of the populations. Idh and Mdh contributed most to the first two axes. 2. We therefore. The results were similar in the four populations. Est. Considering a fairly strong effect of genetic drift in the Large Blue populations we expect a moderate level of variation within the populations coupled with strong genetic differentiation Fig. The distribution of the between sample variation indicated that the populations were not differentiated within any subregion and practically all variation was observed among the years/ generations within a population (Figure 1: B).Temporal and spatial structure of genetic variation in the Alcon Blue populations 165 genetic variation was attributable to the within sample (year/generation) component (Figure 1: A). the results of all analyses demonstrated that the temporal component of differentiation was higher than the spatial one. we also supposed that the temporal component of the genetic variation will be substantial. . Moreover. Allowing for the reproductive strategy of the Alcon Blues we predicted that their populations consist of some unrelated individuals in addition to several groups of relatives. The stochastic processes the Alcon Blue populations experience led to fluctuations in allele frequency. Katalin Pecsenye & Zoltan Varga among them. we detected groups of individuals in the reduced space of variables. the level of differentiation among the samples was high. The results fulfilled our expectations. Even though the sample sizes were relatively small the results seem to support our hypothesis. which exhibit high genetic similarities in their multilocus genotypes. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). which resulted in differentiation both between nearby populations (spatial component of variation) and between consecutive generations within a population (temporal component of variation). At the same time. In order to test our prediction we carried out the PCA analysis using the genotypic composition of the individuals for the four populations separately.166 Judit Bereczki. Although the portion of polymorphic loci was comparatively high. At the same time. We detected a relatively low level of polymorphism within the populations. In all four populations. the average frequency of heterozygotes and especially the mean number of alleles were low. while others consider them as subspecies or simply ecological forms.182** .Hungarian Natural History Museum. 2 1 DUFS . Katalin Pecsenye1.B. nA: mean number of alleles per locus. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group.177** 0. 167-170 Pattern of genetic differentiation in the Maculinea alcon species group (Lepidoptera: Lycaenidae) in Central Europe Judit Bereczki1. Department of Zoology.5 32.5 P 32. Baross u. László Peregovits3 & Zoltán Varga1.J. Nei’s Table 1.O. Some authors distinguish them as separate species of the M. Groups of populations Food plant Region G. Imagos were collected from 27 localities in Central Europe between 1999 and 2003.5 1. Faculty of Sciences.105 0.152** 0. In the analysis of the data.O. H-4010 Debrecen.097 0. rebeli) is quite confused. cruciata North-Hungary Centre of Hungary Transylvania Slovenia Total nA 1.9 29.2 33.5 1. alcon species group.01 level. geographic regions.0 H 0. 1088 Budapest. pp. Romania/ Transylvania and Slovenia.097 0.8 31.0 34.7 30.A. pneumonanthe G.5 1. Parameters of enzyme polymorphism in the Central European Alcon Blue populations averaged according to the two types of hierarchy of the samples: food plant vs. ** significant at 0.094 0. Thomas 2005 © PENSOFT Publishers Pattern of genetic differentiation in the Maculinea alcon Studies on the EcologySettele. E. Central Hungary. Samples originated from four regions: Northern Hungary. Hungary 3 HNHM .193** 0. FIS : index representing within sample variation. Our aim was to study the pattern of genetic differentiation among several populations of these taxa with reference to their taxonomic position.4 1.228** 0.hu The taxonomic status of Alcon Blues living in Central and Western Europe (conventionally: Maculinea alcon and M. P.B.172** 0.klte.093 FIS 0. Department of Evolutionary Zoology and Human Biology. P: percentage of loci polymorphic. Hungary Contact: pecskati@tigris. Enzyme polymorphism was analysed using polyacrylamide gel electrophoresis. F-statistics was computed and the total genetic variation was partitioned into within and between population components. H-4010 Debrecen. H: mean frequency of heterozygotes.079 0. 3. 3. P. 16 enzyme loci were studied.084 0. In all samples. J.172** 0.University of Debrecen.4 1. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.13. Kühn & of Butterflies(Eds)167 species group in Central Europe in Europe and Conservation Sofia – Moscow Vol.5 1. Hierarchical F-statistics and AMOVA were computed to study the pattern of genetic differentiation among the samples. The differences between the two species (Figure 2: Food plant – Among groups) accounted for a slightly lower portion of the total genetic variation than those among the geographic regions (Figure 2: Geographic regions – Among groups). The distribution of the between sample variation indicated a relative high level of differentiation among the years/generations (Figure 2: Between sample component). Nei’s genetic distances were calculated and UPGMA dendrogram was constructed on the basis of the distance matrix.168 Judit Bereczki et al. Fig. The samples belonging to different geographic regions were also randomly distributed among the branches. genetic distances were calculated and a UPGMA dendrogram was constructed on the basis of the distance matrix. remarkable that the samples originating from the same population but collected in different years (generations) were not clustered in the same branch either. This possibly reflects the fact that the majority of individuals in these species have an obligate biennial larval development (Thomas et al 1998). The results suggested that most of the total genetic variation was observed within the samples (Figure 2). First. The next step was to divide the samples according to the geographic regions from which they were collected. 1. The samples exhibited an average level of polymorphism. alcon in central Europe. They were grouped in two ways. A series of hierarchical analyses of genetic variance (AMOVA) was computed by Arlequin. PCA analysis was also carried out using the allele frequencies of the samples. . The parameters of genetic polymorphism were similar in these two groups of populations. The distribution of the total genetic variance at different level of the hierarchy in the 40 samples of M. we contrasted the initial food plant and the geographic pattern of the between sample variation. The samples using different food plants were scattered in all branches. The dendrogram did not show a clear pattern. they were split according to the food plant they use. It is. however. The parameters averaged over the samples of the different regions indicated a higher level of polymorphism in the Northern Hungarian region compared to the others. In the analysis of the structure of genetic differentiation. e. TRY: Transylvania. . rebeli in Central Europe. The two axes explained more than 60% of the total variation. Although some per cent of the total between sample variation was attributable to the differences between the two “species” using different food plants.Pattern of genetic differentiation in the Maculinea alcon species group in Central Europe 169 Fig. The results of the PCA analysis. KMO: Central Hungary. the samples using the same initial food plant (i. The results of the PCA analyses fully confirmed the lack of species pattern in the genetic differentiation among the samples. B: The geographic pattern of variation. this amount of variation was actually less than that explained by the differences among the geographic regions where the samples were collected. When the 95% ellipses were drawn according to the food plants the samples used they were practically overlapping (Figure 2A). The pattern was not much clearer either when the 95% ellipses were drawn corresponding to the geographic origin of the samples. A: The food plant pattern of variation. The results of all statistical analyses suggested a high level of differentiation among the samples without a clear species pattern. The samples exhibited one large cloud of points in the reduced space of variables. SLO: Slovenia. That is. belonging to the same “species”) were as highly differentiated from each other as those using different food plants (i. It thus appears that the pattern of genetic differentiation does not support the species status of M. 2. alcon and M. EMO: Northern Hungary. H: Hochschwab. Although the Transylvanian samples almost completely overlapped with the Northern Hungarian ones they both were slightly differentiated from those originating from Central Hungary (Figure 3B). coming from different “species”).e. C. a social parasite of Myrmica ant colonies. Proc Roy Soc B 265: 1895-1901 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126).W. J. REFERENCE Thomas. .. G.. & Wardlaw. (1998) Polymorphic growth in larvae of the butterfly Maculinea rebeli.A. J. Elmes.170 Judit Bereczki et al. A. scabrinodis complex has been the source of much debate over the years. but also suggests several synonymies within the presently accepted species within the M. E.ku. Settele. scabrinodis sensu strictu. Our phylogenetic investigation additionally supports the status of two recently described species. We present a molecular phylogeny of 17 European species from the M. scabrinodis complex. Boomsma UKBH . DK-2100 Copenhagen Ø. Department of Population Biology. and overlapping species morphologies have lead to several European species being overlooked until recently. David R.University of Copenhagen. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. scabrinodis sensu strictu in Central Europe. scabrinodis sensu strictu lineages in Europe and a microsatellite investigation of reproductive isolation between the mitochondrial lineages in sympatric populations of the two forms. Universitetsparken 15. scabrinodis has been suggested to consist of several ecotypes or even cryptic species. scabrinodis is part of the M. 171 Cryptic Myrmica species among the hosts of Maculinea butterflies Jon R.dk Myrmica scabrinodis is one of the most important host ants for Maculinea butterflies in Europe. Nash & Jacobus J. several of which are used as host by Maculinea butterflies. . scabrinodis complex in general and to evaluate the presence of cryptic species within M. scabrinodis complex based on mitochondrial sequences (COI and Cytochrome B) showing high genetic divergence within Central European M. there are inconsistencies in both its behaviour and interaction with Maculinea butterflies across Europe and M.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. a subdivision of the genus Myrmica represented by roughly 20 described species in Europe. However. scabrinodis complex. With the present investigation we hope to supplement earlier morphological investigations with modern molecular methods to get a better resolution of the M. supporting the division of the species into two well defined subspecies or cryptic species. Kühn J. Ebsen. p. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)171 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Denmark Contact: DRNash@zi. M. Institute of Biology. Centre for Social Evolution. The taxonomy of the M. We will additionally present the initial results of a phylogeographic investigation of the two M. The existence of genetic ecotype lineages might hold crucial information for the understanding of the evolution and dynamics of Maculinea / Myrmica interactions and is of vital importance in Maculinea conservation. uk Recent work has shown that many species of myrmecophiles are so integrated with their host’s social life-style that they have been able to become parasites of ant colonies – social parasites in the broader sense. In the case of the Maculinea alcon/rebeli complex some level of mimicry of the cuticular hydrocarbons of the Myrmica host ants has been demonstrated (Akino et al 2001.nerc. more uniformly distributed”. Even within the species recognised as Maculinea rebeli. E. Clear differences in the “hydrocarbon signatures” have been demonstrated between the main European Myrmica species (potential hosts of Maculinea see this volume) (Elmes et al 2002). yet on others are there more species. using several species on the same site or in the same region. Winfrith Technology Centre. Schönrogge et al 2004). (See other papers in this volume) In any temperate natural or semi-natural habitat in Europe. often with one very abundant species. Kühn & J. developed for alpine M. Dorchester. Settele. and how models developed as part of the MacMan pro- . sabuleti. 172-173 Multiple radiation of species and eco-types in the genus Myrmica Graham W. This shows that in many circumstances only a reduced subset of the original colonising species can persist. The HCET model clearly quantified the dramatic impact that a Maculinea population can have upon a single host population and it is hypothesised that extreme host specificity only pays in certain circumstances where the other ecological and climatic conditions enable a sufficiently robust host population to develop. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. On sites suitable for Maculinea rebeli. one to eight of the common Myrmica species can coexist. defined by the local distribution of the Gentiana foodplant. there appears to be regional differences in host specificity. UK Contact: GWE@wpo. and the community of closely related species coexisting on any site might be determined probabilistically as a consequence of environmental factors combined with disease history. rebeli populations (Hochberg et al 1994). Sofia – Moscow J.A. two to six Myrmica species may be found.ac. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Elmes & Ralph T. We have investigated this by adapting the HCET model. while others appear much more catholic. Some populations more or less exclusively use Myrmica schencki colonies as hosts. CEH Dorset. Dorset DT2 8ZD. This begs the question: “why on some sites are there very few potential host species. This modelling approach is a good example of how a study originally aimed at understanding a rare butterfly that happened to be myrmecophilous has demanded a much greater understanding of the ecology of the host ants. other populations in different geographic regions use M. Clarke 172 Graham W.Centre for Ecology & Hydrology.© PENSOFT PublishersElmes & Ralph T. Clarke NERC . In this respect parasitisation by Maculinea butterflies might be a metaphor for any disease that impacts upon an ant colony’s fitness. pp. to accommodate multiple potential competing host ants living under different conditions of temporal and spatial heterogeneity. E. Peters. Thomas.A. (2002) Interspecific differences in cuticular hydrocarbon profiles of Myrmica ant species are sufficiently consistent to explain host specificity in Maculinea (Large blue) butterflies.W.W. in turn. Thomas. K. Everett. T.A. Knapp. There is good anecdotal evidence that these different “ecotypes” are not equally good hosts for Maculinea and other myrmecophiles.A.. Oecologia 130: 525-535 Hochberg. Akino. J. J. to understand the butterfly dynamics.T. GW (1999) Chemical mimicry and host specificity in the butterfly Maculinea rebeli.T. JJ. G. Proceedings of the Royal Society of London B: 266: 1419-1426 Elmes. that is more than simply a question of nomenclature: “How do we recognise these forms and communicate our results to each other and to practical nature conservationists?” Can cuticular hydrocarbon analysis be a solution? REFERENCES Akino.C. M. Wardlaw.J. Clarke. a social parasite of Myrmica ant colonies.. A. R. (2004) Changes in chemical signature and host specificity from larval retrieval to full social integration in the myrmecophilous butterfly Maculinea rebeli.. Finally. J.J. Yet. S. Journal of Chemical Ecology 30: 91-107 Research has been funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). .. unlike some other insect groups.W. G. studies of host Myrmica ants made as part of the MacMan project have demonstrated that many “species” encompass intra-specific forms that can be ecologically and behaviourally very different from each other. help explain some of the more puzzling apparent contradictions in results obtained by different groups studying different geographical populations of Maculinea. Genetic analyses have suggested that some may even be real “cryptic species”. J. Clarke. JA & Elmes. Current studies are now in hand to test whether our revised understanding of the host ants’ ecologies can.. the rules of nomenclature as applied in ant taxonomy prevent many of these forms being recognised. 1994 Population dynamic consequences of direct and indirect interactions involving a large blue butterfly and its plant and red ant hosts..Multiple radiation of species and eco-types in the genus Myrmica 173 gramme. & Thomas. & Knapp. G. Thomas. J. T. have thrown new light on the population dynamics of the host ants. & Elmes. Journal of Animal Ecology 63: 375-391 Schönrogge. Potentially this poses a problem for ecologists investigating endangered populations of myrmecophiles... R. Elmes. 174-177 Variation in chemical profiles of Maculinea and their Myrmica hosts across Europe Sophie Everett1. 06120 Halle. Baross u. Department of Evolutionary Zoology and Human Biology. Ian Wynne4.A. Laszlo Peregowitz7. First evidence was produced by Akino et al (1999) who both compared the cuticular hydrocarbon (CHC) profiles of French Maculinea rebeli caterpillars and Myrmica workers. Department of Human and Animal Biology. Winfrith Technology Centre. Place Eugene Battallion. Piotr Nowicki5. Department of Community Ecology. (1991). Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 12 University of Turin. Hungary 7 HNHM . Martin Musche11. 4. Irma Wynhof10. Universitetsparken 15. Centre for Social Evolution.B.University of Debrecen. Thomas1 & Karsten Schönrogge1 NERC . P. Polish Academy of Sciences. David N. Elmes1. Bat.uk 1 That caterpillars of Maculinea butterfly species use chemical mimicry to be adopted and infiltrate colonies of their Myrmica host ants was first proposed by Elmes et al. H-4010 Debrecen.P. rebeli CHCs and observed Myrmica worker behaviour. Marcin Sielezniew2. Via Accademia Albertina 13. David Tesar8. Institute of Biology.Jagiellonian University. Poland 3 Museum and Institute of Zoology. . Poland 6 DUFS . Christian Anton11.Hungarian Natural History Museum. Nowoursynowska 159.Centre for Ecology & Hydrology. Michal Woyciechowski5. Kühn & J. Simcox1. Department of Population Biology. Josef Settele11. 30-387 Kraków. Wardlaw1. Denmark 5 UJAG . Foraging ant workers of their host ant M. 6708 Wageningen. Institute of Environmental Sciences.University of Copenhagen. Department of Zoology. Gronostajowa 7. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Andrew D. pp. 1088 Budapest.ac. France 9 Institute of Nature Conservation Kliniekstraat 25 B-1070 Brussels. 10 Mennonietenweg 10. Poland 4 UKBH . Winfrith Newburgh. 10123 Turin. Italy Contact: ssce@ceh. Sofia – Moscow J. Sandor Csósz7. Simona Bonelli12. Nash4. Andras Tartally6. 11 UFZ . David J. DK-2100 Copenhagen Ø.Centre for Environmental Research Leipzig-Halle. and also painted glass dummies with M. Netherlands. Department of Applied Entomology. 00-679 Warszawa. Hungary 8 ISEM . 3. Theodor-Lieser-Str.Warsaw Agriculture University. Belgium. Graham W. Judith C. Wilcza 64. 34095 Montpellier. Dirk Maes9.13. SGGW . CEH Dorset.© PENSOFT Publishers 174 Sophie Everett et al.University of Montpellier II. 02-776 Warszawa. Faculty of Sciences. schencki carried the dummies in the same way that they carried dummies treated with . Settele. Jeremy A. E. Zoltan Varga6. Dorset DT2 8ZD. Worgan1.22. UK 2. Germany. Dorchester.O. Equipe Hochberg. Ania Stankiewicz3. 2004). but might work in more subtle ways than was previ- . the behavioural observations during adoption and the close matches in CHC profiles by Akino et al. (32% similarity) than M. Later studies demonstrated that particularly M. schencki) and non-host (M. it was also generally known that many caterpillars are adopted by the apparently “wrong” Myrmica species in the field. but also for the host specificity observed in the field. (2004) undertaken as part of the MacMan programme suggested that pre-adoption CHC profiles are not sufficient to explain host specificity and postulated that post-adoption synthesis of additional mimetic compounds determines survival especially during periods of stress within ant colonies. alcon described to use M. rebeli caterpillars taken from host (M. (2004) and Elmes et al. M. 2002). M. While there is a strong case that post-adoption “enhanced” chemical mimicry is a strong determinant of caterpillar survival.. schencki (61%) leading to the supposition that the additional hydrocarbons were acquired by contact with the host ants (chemical camouflage) (Akino et al. schencki. which was clearly contradicted by field data. the presence of these chemicals in the CHC profile of M. (2004) studied the post-adoption CHC profiles of M. scabrinodis in southern Europe (Elmes et al. 1989). ruginodis by Thomas et al. However. once adopted even by non-host Myrmica species. rebeli became more similar to that of M. However. 1999). 2002). but which are found in the CHC profile of M. Also. host nests were unchanged. This. (2002) still suggest that preadoption CHC profiles should be adaptive. and under laboratory conditions it is possible to forceadopt Maculinea caterpillars with the “wrong” hosts and have them reared successfully. schencki. further support for chemical mimicry as the basis of host specificity was presented by Nash et al. They found that for caterpillars taken and isolated from the nests. schenckis’ worker profile was to those of other Myrmica ants (<25%) (Akino et al. sabuleti and M. schencki. while ants of other Myrmica species would carry the dummies but drop them eventually. rubra) ant nests. the Maculinea caterpillars should survive equally well with all Myrmica species. After one week in the ants nest the CHC profile of M. 1994. For instance in Poland the main host of M.Variation in chemical profiles of Maculinea and their Myrmica hosts across Europe 175 extracts from their own brood and also as they would adopt the caterpillars in the field. M. Studies by Schönrogge et al. rebeli has been shown to be M. schencki (Steiner et al 2003). ruginodis. (2002) who showed that the CHC profile of M. the caterpillars are able to mimic their natural host. led to the expectation that CHCs at the pre-adoption stage could provide the mechanism not only for the adoption and survival of the caterpillars. rebeli utilise different host ants in different parts of their geographical ranges. a reflex which is suppressed when adopted by the ‘wrong’ host (Schönrogge et al. (1999) and Nash et al. together with the observation at the time that each of the five Maculinea species was adapted to a different host Myrmica species (Thomas et al. North America and Russia were analysed showed clear chemical differentiation and species specific CHC profiles (Elmes et al. rubra in Scandinavia and M. Further studies in which the CHC profiles of five Myrmica species from 49 colonies across the British Isles. whereas those taken from non-host nests were shown to secrete chemicals not found in pre-adoption studies nor in the CHC profiles of their non-hosts. alcon and M. Schönrogge et al. As these chemicals could not have been acquired from any external source. Also the caterpillar CHC profile was more similar to that its host ant. (1989) in the Netherlands was found to use M. Europe. CHC profiles for caterpillars from M. Als et al. here M. sabuleti rather than its western European host. the natural. Also. rebeli suggests that rather than chemical camouflage. local. if all compounds on post-adoption caterpillars were acquired. 1999). alcon closely correlates with those from its host in Denmark. A. a social parasite of Myrmica ant colonies. Poland.. R. Elmes.. 1419-1426. Akino. T.32mm i. 2002 Interspecific differences in cuticular hydrocarbon profiles of Myrmica ants are sufficiently consistent to explain host specificity by Maculinea (large blue) butterflys. The hexane was decanted into clean glass vials and sent back to the CEH laboratories in Dorset for analysis by Gas Chromatography-Mass Spectroscopy (GC/MS). Aliquots of hexane and standard extraction methodologies were sent to collaborators in France. R. a large blue butterfly. Tentschert. Elmes. The GC/MS traces were then analysed visually and using databases of known mass spectra to remove extraneous peaks (Phthalates etc. cleaned mass spectra integrated manually to determine the relative abundance of each chemical and compared with one another using multivariate and nonparametric multidimensional scaling (MDS) on the ranks of the BrayCurtis similarities (Carr 1996). & Clarke. T. J. In the largest. Proceedings of the royal Society B 266. & Boomsma. Als.0b. Hungary and Italy. Denmark. T. Ecological Entomology.R. Japan. J. C.. Using these data we discuss the differences in the pre-adoption CHC profiles between M. Belgium. Als. J.J. Carr.J. A non-polar bonded HP-1 capillary column (50m x 0. alcon and M. W. Knapp. version 4.. Germany. Nash.... Thomas. T. T. & Wardlaw. A. Plymouth.d. J. J. D. Thomas. M. most widespread study of its kind. Nash. & Boomsma. ously thought.176 Sophie Everett et al. UK: Plymouth Marine Laboratories. and their Myrmica host ants: wild adoption and behaviour in ant nests. 1996 Primer user manual (Plymouth routines in multivariate ecological research). 1991 Larvae of Maculinea rebeli. 447-460. Journal of Zoology 223.. 138 . the initial picture of host specificity has become more complex and in order to assess the relevance of the original chemical mimicry hypothesis more chemical analysis of Maculinea and their local Myrmica hosts was required. G. W. Maile. 1999 Chemical mimicry and host specificity in the butterfly Maculinea rebeli. A. J. In: Proceedings of the XIV International Meeting of the IUSSI. chemical extracts from Myrmica and preadoption Maculinea species have been collected using MacMan collaborators from across Europe to determine whether the pre-adoption CHC signatures of Maculinea caterpillars reflect the differences in host ant associations throughout Europe. W.. rebeli and the variation in chemical profiles amongst all five Maculinea species across their European distribution. alcon and M. LITERATURE Akino. Sapporo. a social parasite of Myrmica ants.) and the resultant. Since the MacMan programme has enabled research over a much wider geographical range than any previous study.. D. 2ul of each sample was injected in the splitless mode onto a Hewlett Packard HP5890II GC coupled to an HP5971A MSD operated through HP-ChemStation software.R. p. Thomas. 403-414. 0. G. rebeli. (2002) Geographical variation in host-ant specificity of the parasitic butterfly Maculinea alcon in Denmark.. (2002) Local adaptation and coevolution of chemical mimicry in the butterfly Maculinea alcon.D. J. Jungnickel.52um phase coating thickness) was used to separate the cuticular hydrocarbons and Electron impact Ionisation (EI) was conducted at 70eV with mass scanning from 40-600 m/z. Samples were concentrated under a gentle stream of oxygen-free nitrogen to a final volume of 20ul (caterpillars) or 50ul (ants). G... Oecologica 130. & Elmes. The cuticular hydrocarbons from all 5 Maculinea species and their local host and non-host Myrmica ants were extracted into hexane. J. 27.D. 525-535. J. H. R. the geographical variation with reference to host ant use by M. .. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Thomas. J. F. & Woyciechowski. H. A. M. A. M. B.. Elmes. 91-107. Oecologia 79. W.. Sielezniew. Wardlaw. 2004 Changes in chemical signature and host specificity from larval retrieval to full social integration in the myrmecophilos butterfly Maculinea rebeli. C. Steiner. A. G.Variation in chemical profiles of Maculinea and their Myrmica hosts across Europe 177 Schönrogge.. W. . Everett. Stankiewicz. S. Journal of Insect Conservation 7. 1989 Host specificity among Maculinea butterflys in Myrmica ant nests. J. & Gorniki. 2003 Host specificity revisited: New data on Myrmica host ants of the vulnerable lycaenid butterfly Maculinea rebeli. J. K. M.. & Elmes.. A. 1-6. C. Peters.. Thomas. A. C. Höttinger.. J. Journal of Chemical Ecology 30. G. 452-457.. Wardlaw. J. Schlick-Steiner. pp. arion. Inga Zeisset. E. M. more research is needed for a proper foundation of long-term conservation measures for Maculinea butterflies. Sofia – Moscow J. Here we present the initial results of a study to clarify the detailed population genetic structure and genetic diversity of Maculinea alcon and M. Denmark Contact: DRNash@bi. and another study documented higher sequence divergence among conspecific populations of the predatory species relative to the “cuckoo” species (Als et al. Settele. Centre for Social Evolution. 178-179 A population genetic study of Maculinea arion and M. and from a population in southern Sweden. Boomsma UKBH . Department of Population Biology. the only Danish population. arion the primers had only been tested on one or two individuals. For M. optimization was conducted for more efficient use. and may in part explain the current difference in conservation status of the two species in Denmark and North-Western Europe. These findings have important implications for conservation and reintroduction efforts. alcon in southern Scandinavia in relation to the conservation of these species Andreas E. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. This work will hopefully allow a comparative study with . Universitetsparken 15. and tests were therefore carried out on a greater number of individuals and the microsatellites were found to be polymorphic. A pilot study has also been conducted on a number of museum specimens and resulted in amplification of microsatellites from individuals from the 1960’s and 1970’s using one leg of an adult individual for DNA extraction. arion in Denmark and Sweden using microsatellite markers constructed by Zeisset et al. Nash & Jacobus J. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Kühn & J.dk The two species of Maculinea butterflies that are found in Denmark each represent one of the two known parasitic strategies in the genus: Maculinea alcon whose larvae are fed mouth to mouth by worker ants of the genus Myrmica in addition to feeding on ant brood (the “cuckoo” lifestyle) and M. Work is currently being completed on this material in the laboratory. Additionally.University of Copenhagen.A. Institute of Biology. David R. For the Danish populations.ku. However. whose 4th instar larvae are obligate predators of Myrmica ant brood..© PENSOFT Publishers 178 Andreas E. alcon samples from four population in Sweden and four additional populations in Denmark (all from Jutland) were collected and are also currently being analysed. DK-2100 Copenhagen Ø. Research by Elmes and Thomas (1992) has shown that the predator species are less resistant than the “cuckoos” to habitat disturbances. For M. 2005). their genetic diversity will be compared with individuals collected in 1995 from the same sites. (2005). Wing fragments were sampled from the island of Møn. alcon the primers have already been optimized and their polymorphism has been assessed for populations on the Danish island of Læsø. Lomborg. Lomborg et al. and Thomas. Hsu. A.. D. Mignault. (2005) Microsatellite markers for the large blue butterflies Maculinea nausithous and Maculinea alcon (Lepidoptera: Lycaenidae) and their amplification in other Maculinea species. G. N. I.. REFERENCES Als. alcon in southern Scandinavia 179 recent populations of M..F. J.E. Nature 432. and Pierce. and Boomsma. alcon and so allow an examination of the pattern of genetic change during the decline and/or extinction of several Maculinea populations in the 20th century. Vila.D. 165-168 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). T.. Kandul. Yen. Y.P.. J..R. (1992) Complexity of species conservation in managed habitats: interaction between Maculinea butterflies and their ant hosts. S. N. 155-169 Zeisset.J.A population genetic study of Maculinea arion and M.W. arion and M. Molecular Ecology Notes 5.J.. Nash. . J.. (2004) The evolution of alternative parasitic life histories in large blue butterflies..D. Settele. Als. 386-390 Elmes.A. T. R. Boomsma.A. Biodiversity and Conservation 1..H.. J. Population genetic parameters were estimated using AFLP markers. Both processes may decrease the level of genetic variation and may result in a reduction of reproductive fitness. Department of Community Ecology. 4. J. Theodor-Lieser-Str. genetic population structure and sexual reproduction in Sanguisorba officinalis (Rosaceae) Martin Musche. Kühn & of Butterflies(Eds) 2005 and Conservation in Europe Sofia – Moscow Vol. Germany Contact: Martin. The highest amount of genetic variation was due to differences between individual plants within populations. Population size was not related to the amount of within population genetic diversity but there was a positive correlation between plant density and genetic diversity. percentage germination decreased in populations characterized by a low level of genetic diversity suggesting inbreeding depression. . 06120 Halle.J. Random samples were taken to measure traits related to sexual reproduction.Centre for Environmental Research Leipzig-Halle. Genetic theory predicts that plant populations are susceptible to random genetic drift and inbreeding if gene flow is interrupted or population sizes are small. These findings indicate a large influence of environmental factors on these traits. population size or plant density but largely differed between habitats and populations. The number of seeds per inflorescence and seed weight did not show a relationship with genetic diversity. Fitness traits responded differently to the level of within population genetic diversity. indicating remarkable gene flow counter-balancing the effects of genetic drift.de The perennial meadow plant Sanguisorba officinalis is the single food plant of the endangered butterfly Maculinea nausithous whose early instar caterpillars feed on the flowers and developing seeds. Thomas © PENSOFT Publishers 180 Martin Musche. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. For our study. In this way the genetic population structure of the plant may have a potential impact on the persistence of M.musche@ufz. However. officinalis within an area of 15km x 50km in the Upper Rhine Valley (Germany). nausithous populations as caterpillar survival directly depends on the number and quality of inflorescences.A. pp. Weak differentiation was found between populations but not between both types of habitat. we investigated 24 populations of S. Walter Durka & Josef SetteleStudies on the EcologySettele. We chose populations from different habitats and estimated their population size and density. Pairwise genetic distances between populations were not correlated with geographic distances. This relationship may be the result of fertilisation among a restricted number of genotypes in sparse populations. 180-181 The relationship between genetic diversity. In contrast. Walter Durka & Josef Settele UFZ . In this region the plant occurs frequently and its populations show a low degree of isolation. most variability in germination was explained by maternal seed weight underlining the importance of local environmental conditions. E. However. Therefore. Low gene flow might accelerate the process of genetic erosion if populations become more isolated. non-genetic factors are expected to have a greater impact on the survival of M.The relationship between genetic diversity. genetic population structure 181 Altogether. this conclusion applies only to the study region. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). . Attention should be directed to sparse populations at the range margins of the distribution. officinalis was mainly determined by non-genetic factors. reproductive fitness of S. nausithous caterpillars than the amount of genetic variation within plant populations. Slow-changing sequences can be used to reconstruct phylogeny and patterns of distribution over long time periods. Settele. Boomsma1 1 UKBH . Cytochrome B. Dorset DT2 8ZD. COII. allowed us to reconstruct the phylogeny of the genus. Denmark 2 NERC . The sequencing of several slowly-evolving genes in the Maculinea mitochondrial (COI. Ian R. Winfrith Technology Centre. and to compare extinct with extant populations. to examine their phylogeny and population structure in the light of their interaction with Maculinea butterflies. Wynne1. but we will illustrate some of the possibilities available now using ecological and population genetic . in collaboration with Prof. 182-183 Using genetic markers to study the phylogeny. Winfrith Newburgh. DK-2100 Copenhagen Ø. distribution and ecology of Maculinea butterflies David R.Centre for Ecology & Hydrology.and fast-changing regions of the Maculinea genome. while differences in fast-changing sequences can be used to reconstruct the recent history of populations. It has also been possible to make some tentative inferences about the long-term distribution patterns of the European species that have major implications for their ecology and conservation. Institute of Biology. Sophie Everett2 & Jacobus J. movements and adaptations. CEH Dorset. Universitetsparken 15. During the course of the MacMan project we have identified and sequenced both slow. and so to examine the patterns of life history evolution of Maculinea butterflies. Nash1. as it allows the ecological history of populations to be inferred and their future to be more accurately predicted. Centre for Social Evolution. Ebsen1. pp. Inga Zeisset1. D-Loop) and nuclear (EF1-á.© PENSOFT Publishers 182 David R. Studies of Maculinea butterflies using microsatellite markers are still in their infancy. Dorchester. allowing us to use such regions as genetic markers to examine both evolutionary changes within the genus and population-level changes within the different European Maculinea species. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. The ability to use these markers on museum specimens of Maculinea butterflies will also ultimately allow us to examine changes in the genetic diversity of populations during the last century. Triose Phosphate Isomerase) genome has. Nash et al. Department of Population Biology. their size.dk Examining the spatial and temporal differences in DNA sequences of organisms gives a powerful toolkit to reconstruct their evolutionary history. immigration and emigration.University of Copenhagen.A. The development of highly variable microsatellite markers for Maculinea butterflies has opened up a vast range of possibilities for investigating local population fluctuations. Kühn & J. Sofia – Moscow J. Naomi Pierce’s group in Harvard. E. Jon R. UK Contact: DRNash@bi. Karsten Schönrogge2.ku. We have also used similar techniques to investigate the host ants in the genus Myrmica. Combing population genetic studies with traditional ecological techniques further increases the usefulness of both. This level of differentiation is also reflected in the surface hydrocarbons of the caterpillars. . providing no evidence for local adaptation based on host ant use on Læsø. We have shown that there is no detectable genetic subdivision of populations based on their host ant use. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Although both ant species are used as hosts on the island.Using genetic markers to study the phylogeny. Previous studies have shown that both Myrmica rubra and Myrmica ruginodis are used as host ants on the island. however. at a local scale only one ant species acts as a host. and have examined the genetic and chemical differentiation of different populations based on their geographical distribution and pattern of host use. This island supports some of the largest populations of Maculinea alcon in Western Europe. We will also present the results of initial analyses comparing estimates of population size and movement collected using traditional ecological techniques (Mark-Release-Recapture and following of individual flight paths) with estimates based on population genetic data. During the course of the MacMan project. There are several places on the island. representing the ant species that is numerically dominant in the area. where areas where different host ants are used are separated by only a few tens of meters. we have carried out detailed studies on the distribution and abundance of M. distribution and ecology 183 studies of Maculinea alcon on the Danish island of Læsø. alcon and its host ants on the island. but there is geographical differentiation of populations on the order of 1 km or so. Non-LTR retrotransposons have been found in many eukaryotes investigated up to date.nsc. In this study. Lovsin et al. 630090. Several enzymatic activities can be distinguished in proteins encoded by non-LTR retrotransposons (Malik et al. CR1. RTE. Finally.000 copies (~20 % of the genome) for L1 elements in human (International Human Genome Sequencing Consortium 2001). L2 and I. 2000. allowed to suppose the presence at least nine clades of non-LTR retrotransposons in Maculinea: R2. LOA. The first open reading frame (ORF). The elements of this class have no long terminal repeats and utilize a simpler target-primed reverse transcription (TPRT) mechanism for their retrotransposition (Luan et al.. Novosibirsk. Alexander Blinov1 & Michal Woyciechowski2 1 Siberian Branch of the Russian Academy of Sciences. 10. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Lavrentjev aven. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. it appears that diversity of non-LTR retrotransposons is considerable higher in arthropods than it was supposed previously (Biedler and Tu 2003). Poland Contact: novikova@bionet. 1999.A. The insects’ mobile elements. Russia 2 UJAG . Ewa Sliwinska2. Institute of Cytology and Genetics. R4. . 30-387 Kraków. Volff et al. Institute of Environmental Sciences. R1. The second component is endonuclease.Jagiellonian University. 1993). The known distribution and phylogenetic analysis of non-LTR retrotransposons. Arkhipova and Morrison 2001). based on the reverse transcriptase domains. 1999). Jockey. which is provided by restriction-enzyme-like endonuclease (REL-endo) domains in some elements and by apurinic/apyrimidinic (AP) endonuclease in other elements. Malik and Eickbush 2000. Phylogenetic analysis of non-LTR retrotransposons based on the reverse transcriptase domains allowed 15 phylogenetic clades to be distinguished (Malik et al. pp. Kühn & J. 184-188 Non-LTR retrotransposons from Large Blue butterfly Maculinea teleius: the diversity of CR1-like elements Olga Novikova1. The key component is reverse transcriptase that is present in all non-LTR elements. if present. were intensively investigated using model organisms such as Drosophila melanogaster (Diptera) and Bombyx mori (Lepidoptera). Lovsin et al. encodes a gag-like protein with a function of nucleic acid chaperone (Martin and Bushman 2001). large blue butterflies of the genus Maculinea are examined for the presence of the non-LTR retrotransposons.ru Non-LTR retrotransposons are mobile genetic elements that propagate themselves by reverse transcription of an RNA intermediate. Settele. some elements also contain ribonuclease H (RNase H) domains. Gronostajowa 7. E. 2001. 1999.© PENSOFT Publishers 184 Olga Novikova et al. However. including non-LTR retrotransposons. Sofia – Moscow J. All these clades contain elements known from arthropods (Malik et al. The copy number of these elements may vary from just several copies per genome to over 800. 2001). The PCR products for LOA and CR1 clades from all four species were cloned into pBlueScript vector using standard procedures. The amino acid sequences of newly identified RT and RT from GenBank database were aligned using ClustalW software (Thompson et al. or they can be highly divergent from the known elements. 1. Statistical support for the trees was evaluated by bootstrapping (100 replications) (Felsenstein 1985). 2001). non-LTR retrotransposons clade investigated and results of PCR screening. mori. teleius. RTE and CR1 (Table 1). It is possible that these groups are absent from Maculinea genomes. R4. Jockey. even in distantly related host species. R1. than between the different clades from the same host. and was appropriate for the R4 family from all Maculinea species investigated. nausothous M. and four species of the genus Maculinea were screened by PCR amplification (Table 1). . It was intriguing that all clones for LOA and CR1 clade. except two clones for CR1 clade. Only the new primers the R4 clade amplified a product of ca. 153 clones were isolated. So we decided to design new non-degenerate primers based on these elements. mori elements sequences. alcon M. we obtained no PCR products with Maculinea DNA.result PCR with degenerate primers/ result PCR with non-degenerate primers. The BLAST search against protein database (blastp) showed that most of RT-carried clones for LOA and CR1 clades share a significant similarity with non-LTR retroelement from B. whereas others showed clear similarity with reverse transcriptase of known non-LTR retroelements. other lepidopaterian species. which have appropriate lengths. LOA. there are known elements from B. group A). Among 20 sequenced clones only two from LOA clade and one from CR1 clade did not contain any sequences of reverse transcriptase. DNA fragments of the expected size were obtained for the R4. species Maculinea teleius M. “-/-” . melanogaster. 300 bp and 1000 bp correspondingly) from D. 400 bp. Whereas new primers for R1 and R2 clades were capable of amplifying products of appropriate lengths (ca. This similarity allows degenerate primers to be designed that are capable of detecting retrotransposons belonging to a given superfamily in the diverse taxa.two pairs of primers were used. 1). mori. It seems that primers Table 1. In total. The phylogenetic analysis was implemented to find the respective positions of the obtained clones in the non-LTR retrotransposons evolutionary tree (Fig. formed a common branch on the phylogenetic tree (Fig. Consistent with the spacing of the amino acid sequence domains. “-” – negative result of PCR. “+” – positive result of PCR. None of the investigated species showed positive results with primer combinations for R2. Maculinea species. 1994) and improved manually. Degenerate oligonucleotide primers were designed for eight clades. Phylogenetic trees were generated by Neighbor Joining method using MEGA2 software package (Kumar et al.Non-LTR retrotransposons from Large Blue butterfly Maculinea teleius 185 Reverse transcriptase sequences from known retroelements show more similarity within the same clade. arion R2* -/-/-/-/R1* -/-/-/-/R4* -/+ -/+ -/+ -/+ LOA + + + + Jockey + + + + CR1 + + + + RTE + + + + I - * . However. in each clade. We sequenced ten clones for each LOA and CR1 clades from M. the primers amplified a PCR product of 500 bp in length. and I groups of elements. 96 of them were obtained for LOA clade and 57 for CR1. degenerate primers and non-degenerate primers designed based on the B. The names of source species are given near the names of elements. the real position and distribution of the LOA clade is unknown. The BLAST search of LOA nucleotide sequence against Fig. and obtained in present investigation – bold. In addition. as a consequence of degeneracy. It is possible. and. It seems that arthropods have a high diversity of elements inside the CR1 clade. primers for LOA clade became to be involved in amplification of other elements. designed for the detection of LOA-like elements failed to amplify of the LOA-like sequences but can be used for procuring of elements from CR1 clade. Phylogenetic tree of non-LTR retrotransposons: retrieved from database – bold. italic and underlined. the resent research failed to identify LOA-like elements in the Anopheles gambiae genome (Biedler and Tu 2003). The LOA element was originally described from genome Drosophila silvestris (Felger and Hunt 1992) and subsequently in genomes of D. that elements from LOA clade are absent or present by low copy numbers in genomes of Maculinea butterflies. However. . On the other hand these results contrasted with our results obtained in nonLTR retrotransposons from scorpions’ investigations (unpublished results) and results of Anopheles gambiae whole genome screening (Biedler and Tu 2003). 1998). The bootstraps with number less than 30 were removed. 1. These elements can be subdivided into several groups and construct new families or even clades (Biedler and Tu 2003). subobscura (Blesa and Martinez-Sebastian 1997) and Aedes aegypti (Tu et al.186 Olga Novikova et al. and Q. one dead. Thus. Evolution 39:783-791 International Human Genome Sequencing Consortium (2001) Initial sequencing and analysis of the human genome. insects (Zampicinini et al. 2004). REFERENCES Arkhipova IR. 14:1145-1153 Casa AM. The most popular transposon-based marker method is the Sequence-Specific Amplification Polymorphism approach (S-SAP). Hunt JA (1992) A non-LTR retrotransposon from the Hawaiian Drosophila: the LOA element. Anopheles gambiae: unprecedented diversity and evidence of recent activity. 1. and fungi (Taylor et al.like sequences have been revealed in genomes of many arthropods investigated. Zhang Q. Mobile elements. 2004). 2003). Proc Natl Acad Sci USA 97:10083-10089 Felger I. horizontal transfer of non-LTR retrotransposons. Moreover. On the other hand. therefore the distribution of non-LTR retrotransposons among species generally reflects their phylogenetic relationships. Proc Natl Acad Sci USA 98:14497-14502 Besansky NJ (1990) Evolution of the T1 retroposon family in the Anopheles gambiae complex. are a powerful tool for phylogenetic analysis. Mukabayire O (1994) Q: a new retrotransposon from the mosquito Anopheles gambiae. Brouwer C. Nature 409:860-921 . Non-LTR retrotransposons. It will permit us to show how many clades of non-LTR retrotransposons are present in lepidopterians and to investigate in details a phylogenetic structure of the CR1 clade. group B). also called “transposon display” (Casa et al. Besansky et al. Wessler SR (2000) Inaugural article: the MITE family heartbreaker (Hbr): molecular markers in maize. appear to be fixed. 85:119-130 Felsenstein J (1985) Confidence limits on phylogenies: an approach using the bootstrap. The S-SAP markers were developed for wide range of taxa in particular in plants (Casa et al. group A). is believed to be very rare. 1. Bedell JA. On the other hand two clones share a very high similarity with T1 and Q elements from Anopheles gambiae (Besansky 1990. Morrison HG (2001) Three retrotransposon families in the genome of Giardia lamblia: two telomeric. mori and formed one group with LOA clones (Fig. Mol Biol Evol 7:229-246 Besansky NJ. 2000. non-LTR retroelements are planned to be used in phylogenetic and population polymorphism analysis. Insect Mol Biol 3:49-56 Biedler J. On the one hand the main group of elements like the LOA clones showed high similarity with elements from B. Further investigations of non-LTR retrotransposons will be made for the remaining clades and other Maculinea species. Genetica. insertions are triflingly insignifican in the same independent locus in unrelated lineages. Martinez-Sebastian MJ (1997) bilbo. One more phenomenon. particularly non-LTR retrotransposons. 2000). 1994) (Fig. The integration of a non-LTR retrotransposon to a new place is an irreversible event. Vershinin et al. Kresovich S. Tu Z (2003) Non-LTR retrotransposons in the African malaria mosquito. once inserted in chromosomal DNA. 20:1811-1825 Blesa D.Non-LTR retrotransposons from Large Blue butterfly Maculinea teleius 187 database of arthropods sequences gave the high similarity with oneself and several sequences from other Drosophila species (unpublished data). Another intriguing observation is the clear division of CR1 clones into two groups. Wang L. Mol Biol Evol. The presence of this group in the genome of Maculinea teleius is expected since the T1. Nagel A. a non-LTR retrotransposon of Drosophila subobscura: a clue to the evolution of LINE-like elements in Drosophila. it seems to be that at least two separate groups from the CR1 clade are present in the genome of Maculinea teleius. Mol Biol Evol. Ellis TH (2003) Transposable elements reveal the impact of introgression. Eickbush TH (1993) Reverse transcription of R2Bm RNA is primed by a nick at the chromosomal target site: a mechanism for non-LTR retrotransposition. Blinov A. Nucleic Acids Res 22:4673-4680 Tu Z. Chironomidae) as detected by transposon insertion display. Mol Biol Evol 16:793-805 Malik HS. Cell 82:545-554 . and domestication. Konstantinova P. Genome 47:1154-1163 Zimmerly S. Schartl M (2000) Multiple lineages of the non-LTR retrotransposon Rex1 with varying success in invading fish genomes. position-specific gap penalties and weight matrix choice. Genome 47:519-525 Thompson JD. Mol Cell Biol 21:467-475 Taylor EJ. Eickbush TH (1999) The age and evolution of non-LTR retrotransposable elements. Korting C. Leigh F. Higgins DG. Mol Biol Evol. Bates JA. Cell 72:595-605 Malik HS. Eickbush TH (2000) NeSL-1. Tamura K. Guryev V.188 Olga Novikova et al. Cervella P. Kumar S. Lambowitz AM (1995) Group II intron mobility occurs by target DNAprimed reverse transcription. Perlman PS. Bushman FD (2001) Nucleic acid chaperone activity of the ORF1 protein from the mouse LINE-1 retrotransposon. Genetics 154:193-203 Martin SL. genomic. Burke WD. Gubensek F. and phylogenetic analysis of Lian. Ambrose MJ. Gibson TJ (1994) CLUSTALW: improving the sensitivity of progressive multiple sequence alignment through sequence weighting. Bioinformatics 17:1244-1255 Lovsin N. an ancient lineage of site-specific non-LTR retrotransposons from Caenorhabditis elegans. Sella G (2004) Insertional polymorphism of a non-LTR mobile element (NLRCth1) in European populations of Chironomus riparius (Diptera. Mol Biol Evol 17:1673-1684 Zampicinini G. Aedes aegypti. Lee D (2004) Gypsy-like retrotransposons in Pyrenophora: an abundant and informative class of molecular markers. Mol Biol Evol 18:2213-2224 Luan DD. Isoe J. Kordis D (2001) Evolutionary dynamics in a novel L2 clade of non-LTR retrotransposons in Deuterostomia. in Pisum diversity. Nei M (2001) MEGA2: molecular evolutionary genetics analysis software. Korman MH. Guzova JA (1998) Structural. Knox MR. Jakubczak JL. a novel family of non-LTR retrotransposons in the yellow fever mosquito. Allnutt TR. evolution. 15:837-853 Vershinin AV. Guo H. Mol Biol Evol 20:2067-2075 Volff J-N. rather than transposition. Jakobsen IB. 4 18. E.73 H 0. Faculty of Sciences.05 level. pp. László Peregovits3 & Zoltán Varga1. P. Baross u.8 37. 189-191 Genetic differentiation among the Maculinea species (Lepidoptera: Lycaenidae) in eastern Central Europe Katalin Pecsenye1.O. FIS: index representing the within sample component of variation.47 1.136** 0.101 0.01 level. Hungary. H-4010 Debrecen. Kühn & of Butterflies(Eds)189 Studies on in eastern Central Europe and Conservation in Europe Vol.3 31. N: sample size. 3.klte.hu The aim of the present work was to study the level of genetic variation in the eastern Central European (Slovenia. Hungary 3 HNH -M Hungarian Natural History Museum.129** 0. * significant at 0. teleius (13) M. 3.126** . alcon and M. AMOVA was computed to study the pattern of Table 1. 2 1 DUFS . Species M. arion (3) Total (32) N 27. H-4010 Debrecen.6 14. rebeli (7) M. Judit Bereczki1. P.080 0. nausithous (3) M.A. 1088 Budapest. Borbála Tihanyi1. P: proportion of polymorphic loci.71 1. Hungary Contatcs:
[email protected] 0.097 FIS 0. and Romania/Transylvania) populations of the Large Blues in order to analyse the pattern of differentiation both between and within the species.50 1.6 32. Imagos were collected from 23 localities in eastern Central Europe in 2002. In the analysis of the data. It was also an objective of our investigation to compare the level of differentiation between the two disputed species (M. rebeli) to those among the others.3 24. 14 enzyme loci were studied in all samples. Department of Zoology.089 0.7 42.O.3 22.© PENSOFT Publishers Genetic differentiation Sofia – Moscow J. J.11 nA 1.194 0.B. ** significant at 0. Nei’s genetic distances were calculated and a UPGMA dendrogram was constructed on the basis of the distance matrix.70 1.575 P 28.4 23.38 1.0 54. Andrea Tóth1.135* 0. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.167** 0.218** 0. alcon (6) M.University of Debrecen.13.B. Parameters of polymorphism in the samples of the five Maculinea species. Numbers in brackets indicate the number of samples analysed. F-statistics was computed and the total genetic variation was partitioned into within and between population or between species components. nA: average number of alleles. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group. Enzyme polymorphism was analysed using polyacrylamide gel electrophoresis. H: average observed frequency of heterozygotes. Thomas 2005 among the Maculinea species the EcologySettele. Department of Evolutionary Zoology and Human Biology. 05 . M. UPGMA dendrogram constructed for the five Maculinea species on the basis of Nei’s genetic distances. We were also looking for diagnostic loci among the investigated Maculinea species.190 Katalin Pecsenye et al. alcon pneumonanthe type.837** Nm 1. Both parameters showed that the level of differentiation was much lower within the species than among them (Table 2). refer to M. a. M. rebeli as M. arion (3 p) Total (4 sp) FST 0.: M. Gdh.038** 0. alcon cruciata type. pneu.33 0. alcon (13 p) M. Mdh. cruci. **: significant at 0. a.56 1. rebeli populations using different food plants were not differentiated at the species level (Figure 1). The UPGMA dendrogram built on this distance matrix showed that the M.110** 0. pneu. therefore.12 2. Species M. alcon as M.138** 0. The samples showed an average level of polymorphism. Four loci (Aox. (7 p) M. teleius (13 p) M. FST and Nm values computed on the basis of FST calculated for the Maculinea species separately. alcon and M. Me) had specific allele composition in at least one species. (6 p) M. alcon cruciata type. Table 2.02 6. PCA analysis was also carried out using the allele frequency data of the samples.067** 0. 1. cruci. nausithous (3 p) M.097** 0. a. FST values were computed and the effective number of migrants was estimated using restricted data sets where the samples of the species were analysed in separate series. Numbers in brackets: number of populations (p) or species (sp) in the given group.01 level. genetic differentiation among the samples. M.33 3.48 2. Comparing the parameters estimated for the pneumonanthe and the cruci- Fig.182** 0. arion populations were the most polymorphic among the five species. Parameters of genetic polymorphism were significantly different in the four species. Nei’s genetic distances were calculated using a pooled data set with 5 “composite populations” where the samples belonging to one species were combined.: M. We. a. alcon pneumonanthe type and to M. Our results did not show genetic differentiation among the pneumonanthe and cruciata type M. pneumonanthe type M. Results of the PCA analysis for the 32 samples of the five Maculinea species. alcon (i. the M. alcon populations were not differentiated according to the food plant they use. cruciata type M. The results of the PCA analysis were very obvious. but sometimes multiple use of food plants and host ants) did not result in genetic differentiation among these two types of M. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). alcon populations. alcon samples we could not find a clear difference. alcon populations. concluded that the local ecological adaptation (various habitats. different.Genetic differentiation among the Maculinea species in eastern Central Europe 191 Fig. 2. We therefore. rebeli (i.e. Low level of genetic variation within the populations coupled with strong differentiation among them implies the effect of genetic drift. alcon) and M. alcon) samples were not separated from each other. which is not surprising considering the highly specialised life cycle of the Large Blues. .e. alcon separately to those calculated for all M. ata type populations of M. The samples exhibited four compact clouds of points in the reduced space of variables (Figure 2). Our results indicated that the Large Blues are generally less polymorphic than other European lycaenid butterflies studied. At the same time. This also indicated that M. The two axes explained slightly more than 88% of the total variation. the outcome of all analyses suggested that the level of genetic differentiation was high even among local populations within the species. That is. E. 3. some populations were sampled in two or more years.7 1.226** 0.e. P. Sofia – Moscow J. Altogether 40 samples were collected between 2000 and 2004.6 37.8 1.080 0.5 1.8 15. M. i.109 0. FIS: index of the within population component of genetic variation.116 0.158** 0.7 24. In the analysis of the data.189** 0.7 1.7 26. N: sample size. teleius populations had a moderate level of enzyme polymorphism (Table 1).172** . H-4010 Debrecen. the parameters of polymorphism were first estimated and F-statistics were computed where the total genetic variation was partitioned into within and between population components.1 22.146** 0.7 1.2 29.0 20. Numbers in brackets indicate the number of samples / number of populations.B. Hierarchical F-statistics and AMOVA was computed to study the pattern of genetic differentiation among the samples.hu Maculinea teleius samples originated from 21 populations of 8 geographic regions in Hungary.8 31. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. (3/3) Õrség (9/6) Szigetköz (2/1) Total (40/21) N 65.O.klte.5 22. 3.079 0. PCA analysis was carried out using the allele frequencies of the samples to show the size of overlap in the genetic variation of the populations in a reduced space of variables. 192-195 The genetic structure of the Maculinea teleius (Lepidoptera: Lycaenidae) populations in Hungary Katalin Pecsenye. Judit Bereczki.5 50.3 34.5 1.Evolutionary Genetics and Conservation Biology Research Group. Genotype and allele frequencies were estimated.O. Department of Evolutionary Zoology and Human Biology.01 level.246** 0.087 FIS 0.073 0.237** 0.69 P 29.7 1. P: proportion of polymorphic loci.059 0.© PENSOFT Publishers 192 Katalin Pecsenye et al.2 39. ** significant at 0.076 0. nA: average number of alleles per locus.0 44.83 H 0. Faculty of Sciences.4 16. H: average frequency of heterozygotes. which did not have any alternative allele in any of the investigated samTable 1. Parameters of polymorphism in the Maculinea teleius populations averaged for the 8 regions. Andrea Tóth & Zoltán Varga 1 DUFS .B.6 23. Settele.178** 0. Hungary Contact:
[email protected]** 0. Kühn & J. H-4010 Debrecen.A.35 nA 1. Regions Karst (4/1) Cserehát (8/3) Zemplén (8/4) Bereg (3/1) Kiskunság (3/2) Dunántúli-Mts. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Hungary 2 HAS-UD . Enzyme polymorphism was investigated at 16 loci in all samples using polyacrylamide gel electrophoresis. pp. Nei’s genetic distances were calculated and a UPGMA dendrogram was constructed on the basis of the distance matrix.University of Debrecen. There was only one locus (Got-2).9 1. P.100 0.7 46. The pattern of genetic differentiation was analysed using different methods. The parameters of polymorphism averaged for the 8 regions did not differ significantly from each other.The genetic structure of the Maculinea teleius populations in Hungary 193 ples.056** 0. The outcome of these analyses showed that the level of differentiation within the regions was similar to that estimated for all samples (Table 2). also indicated an average level of genetic variation.085** 0.102** 0. the populations of any region were scattered in different branches of the dendrogram. (3) Õrség (9) Total (40) FST 0. A UPGMA dendrogram was constructed on the basis of the distance matrix.074** Exact test *** *** *** *** * *** Fig. ** significant at 0. Nevertheless.01 level.05 level. . The 21 populations were clustered in the dendrogram independently to their geographic origin. population and sample). The results of F-statistics. teleius samples. This implies that populations originating from different regions were not more differentiated than those of the same one. FST values were computed for the 8 regions separately using restricted data sets. where all samples were involved.028** 0. FST values indicated a significant differentiation among the samples. That is. The results of AMOVA revealed that the highest portion of variation was attributable to the within Table 2. Distribution of variation at different levels of the hierarchy estimated by computing AMOVA. Most of this variation was observed within the samples. All samples and populations exhibited a high level of heterozygote deficiency (Table 1: FIS). * significant at 0. This finding also indicated the lack of geographic pattern in the differentiation of the M.001 level. Numbers in brackets indicate the number of samples. Both parameters were significant in all regions. AMOVA was also computed to analyse the distribution of variation at different levels of the hierarchy (region. Region Cserehát (8) Zemplén (8) Kiskunság (3) Dunántúli-Mts.012* 0. The next step was to calculate Nei’s genetic distances. FST values and the results of exact test estimated for the 8 regions separately. 1. *** significant at 0. This suggests that the genetic composition of the populations exhibits a considerable fluctuation generation by generation. . The points representing the samples composed Fig. it is remarkable that a sizeable portion of the between sample variation was explained by the differences among the regions (Figure 1). The results of the PCA supported those of AMOVA. 2. Fig. teleius populations. Nevertheless. The distribution of the between sample variation of M. the temporal component of variation (within population variation) proved to be higher than the spatial one (Figure 2). a principal component analysis was carried out using the allele frequency data of the samples to study the differentiation among the geographic regions. The entire between sample variation amounted to less than 10% of the total variation. Three regions were chosen for this analysis where we had several samples from three or more populations. Each point indicate a sample in the reduced space of variables. 3. As some of the samples originated from the same population but were collected in different years (generations) it was possible to estimate the temporal component of genetic variation in the M. The ellipses represent the samples of one geographic region. In the last part of the study. sample component (Figure 1).194 Katalin Pecsenye et al. teleius in three regions of Hungary. In all regions. Results of the PCA analysis. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). The high portion of the temporal component in the between sample variation suggested a strong effect of genetic drift affecting the populations and a limited gene flow among them even within a region.The genetic structure of the Maculinea teleius populations in Hungary 195 one diffuse cloud. teleius populations had a moderate level of variation coupled with a low level of genetic differentiation and an overall heterozygote deficiency. The 95% ellipses indicating the samples collected in one region were completely overlapping. The genetic structure of the populations did not have a geographic pattern at the regional level. we concluded that M. In summary. . A. When a formerly continuous population becomes fragmented into several isolated subunits this is expected to substantially reduce the effective population size and to increase the risk of extinction (Frankham et al. teleius. teleius habitat in the Vistula valley near Krakow has had an impact on the effective population sizes of this species. no selection and random mating (Wright 1931) and will thus be a measure of the rate at which genetic variation is lost through genetic drift (Hedrick 2000. Centre for Social Evolution. The effective population size is a key variable when considering the potential of a population to persist. 196 Ewa Sliwinska Sofia – Moscow J. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. E.© PENSOFT Publishers et al. Frankham et al. Poland 2 UKBH . teleius caterpillars were frozen in an . Piotr Nowicki1. 2002). 1990). Nowicki et al. In genetic models. The site has a few dozen patches of the Sanguisorba officinalis host plant for M. pp. MATERIAL AND METHODS Sampling and genetic analysis The study was conducted on a wet meadow complex located in southern Poland. Department of Population Biology. Settele. 2002. Jacobus J. Here we use genetic microsatellite markers to investigate whether recent fragmentation of continuous M. Universitetsparken 15. i. Institute of Environmental Sciences.Jagiellonian University. The M. Boomsma2 & Michal Woyciechowski1 UJAG . The multitrophic relationships of the larvae of this butterfly with the foodplant Sanguisorba officinalis and one or two species of Myrmica host ant make it likely that the population dynamics of this species will be affected by a complex of ecological factors.University of Copenhagen.e. 4 km southwest of the centre of the city of Krakow. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. DK-2100 Copenhagen Ø. 2002). 30-387 Kraków. Denmark Contact: ewa-sliwinska@wp. 1998. 196-198 Effective population size of Maculinea teleius in southern Poland Ewa Sliwinska1.pl 1 INTRODUCTION Maculinea teleius is one of four species of Maculinea butterflies included in the European Red List (IUCN. Stankiewicz & Sielezniew. Its threatened status has made this species a subject of investigations in many European countries including Poland (Figurny & Woyciechowski. Gronostajowa 7. Kühn & J. the effective population size (Ne) is estimated under the assumption of Hardy-Weinberg equilibrium. Institute of Biology. From seven of these subpopulations we collected 30 larvae from Sanguisorba flower buds in August of 2004. 2005b). Beerli 2004.7.048 0.260 0.245 0.001). Population ID K-3 K-10 K-17 K-18 K-71 K-14 K-25 Fis 0. We used two long and five short Markov chains and the number of discarded genealogies per chain was 1000.066 Ln(L) -2096. teleius subpopulations might indicate that there is some inbreeding depression connected to habitat fragmentation. RESULTS We found significant deviations from Hardy-Weinberg equilibrium both within subpopulations and when all samples were pooled.153 0.280 0. To test the relationship between effective population sizes and censused population sizes we used censused population sizes estimated with capture frequencies in 2004 (see Nowicki et al.033 0. “ì” is mutation rate).044 0. Ln(L) means highest likelihood estimation for è. which is most appropriate for microsatellite markers and a migration matrix model with variable è.6.167 0. This hypothesis is reinforced by the positive correlation of the population census data and è.178 0. 2005).Effective population size of Maculinea teleius in southern Poland 197 ultrafreezer (-80oC) shortly after collection and screened for six polymorphic microsatellite loci (Zeisset et al. Results for seven investigated populations: Fis statistics.05). Statistics To test for Hardy-Weinberg equilibrium within and among populations we used G-tests (FSTAT. Statistical significances are based on Pearson correlation coefficients (FSTAT).1. Beerli & Felsenstein 1999).112) (Table 1). DISCUSSION The significant heterozygote deficiencies in all seven M. 2005a). The null hypothesis of Hardy-Weinberg equilibrium within and among (sub)populations could be rejected (P = 0. Given that the Table 1. Goudet 1992). All subpopulations had positive inbreeding coefficients but the correlation between population census estimates and Fis values was not statistically different from zero (give r value and exact P value)(P > 0.524 . We applied a stepwise mutation model and a gamma-distributed mutation rate (ì ) among loci. The correlation between the è values and the population size estimates per (sub) populations was also positive (r = 0.131 Seasonal population size 229 6012 780 196 12241 7055 7188 è (= 4Ne ì) 0. We used the Fst values to estimate the effective population size è = 4Neì using the coalescence approach in the Migrate likelihood estimator (version 1.294 0. and è calculation (in formula shown in the table “Ne“ is effective population size.055 0.086 0. seasonal population size (estimated on the basis of capture frequencies). Inbreeding coefficients (Fis) and coefficients of population subdivision (Fst) were calculated using Weir and Cockerham’s F-statistics (1984) in FSTAT.643) but not significant (P = 0. . Academic Press.W. Estimating F-statistics for the analysis of population structure.A. NY. Evolution 38: 1358-1370. Genetic effective size of a metapopulation. The reintroduction of two Maculinea butterfly species. Stankiewicz A.. nausithous Bgstr. Doctoral Thesis. (2005a) Landscape scale research in butterfly population ecology – Maculinea case study.. Briscoc D. . 140-143. 1999. Evolution in Mendelian populations. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126)..H. The effective size of a subdivided population. IUCN Red list of threatened animals. 165-181. At home on foreign meadows. and M. 1998. A. teleius are more resistant to temporal demographic fluctuations that affect Ne (Hedrick & Gilpin 1997).E. Whitlock M. Wageningen University. Pępkowska A. Felsenstein J.D. Cambridge University Press. Kühn & J. Tropical Nature Conservation and Vertebrate Ecology Group... Genetics of Populations. 1.198 Ewa Sliwinska et al. In Metapopulation Biology: Ecology.. Wright S. 2002. Maximum likelihood estimation of migration rates and population numbers of two populations using a coalescent approach. Genetics 152: 763-773. Kudłek J.).C. Genetics 146: 427-441.W. Settele J.. Host specificity of Maculinea teleius Bgstr. Department of Environmental Sciences. nausithous (Lepidoptera: Lycaenidae). Witek M. The Netherlands. Genetics 16: 96-159. Skórka P. Woyciechowski M.S. 1997. Jones and Bartlett Publishers. 1984.. E.. pp. eds. nausithous. MA.. Population ecology of the endangered butterflies Maculinea teleius and M. 2000. expectation was in fact one-tailed (only a positive correlation could be expected). Annales Zoologici 52: 403-408. Flowerhead selection for oviposition of the sympatric butterfly species Maculinea teleius and M. 1995. Nowicki P. 2001. 1931.E. and M. This suggests that such large unfragmented populations of M. Cockerham C. IUCN. Cambridge. Skórka P. REFERENCES Beerli P.2): a computer program to calculate F-statistics. IUCN 1990. Hedrick P. Sudbury. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.. (Lepidoptera: Lycaenidae): the new insight. Sielezniew M. Pp.. Frankham R..). Population Ecology [in press]. 2002. A qualitative inspection of the data showed that estimates of è were almost equal for subpopulations with ~ 200 to ~ 7000 butterflies. 1997. Gilpin. Hedrick P. Hanski. Entomologia Generalis 23: 215-222. Ist edn. Cambridge Nowicki P. Introduction to conservation genetics. Wynhoff I. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. In: J. Witek M.C. Figurny E.. Journal of Heredity 86: 485-486.. Barton N. this relationship is border-line significant. Weir B.. Goudet J. Woyciechowski M. Ballou J. but that the largest population (K-17) occurring on a non fragmented area with S. Woyciechowski M. Settele. 2005b. and its implications for conservation. FSTAT (vers. Gilpin M. Genetics and Evolution (I. officinalis had a much higher effective population size è (Table 1. Nevertheless. H-4010 Debrecen. Enzyme polymorphism was investigated at 16 enzyme loci in all samples using polyacryamide gel electrophoresis. P. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. arion: M. Hierarchical F-statistics and AMOVA were computed to study the pattern of genetic differentiation among the samples. a. some authors simply consider them as ecological forms.B.8 1.064 0. PCA analysis Table 1.9 15.074** .O. pp.A. summer type) as food plant. Kühn & J. M. spring HU (3/1) M.B. N: sample size. summer HU (8/3) M.hu The taxonomic status of Large Blues (Maculinea arion) living in Central and Western Europe is confused. Settele. a. Faculty of Sciences. Subspecies M. 199-202 Patterns of genetic differentiation in the Hungarian Maculinea arion (Lepidoptera: Lycaenidae) populations Andrea Tóth1.1 44. spring and summer type) or Origanum vulgare (M. the spring type and M. nA: average number of allels per locus. FIS: index of the within population component of genetic variation.8 1. Altogether 15 samples were collected between 2000 and 2004.6 21.University of Debrecen. H: average frequency of heterozygotes.144 0. Nei’s genetic distances were calculated and UPGMA dendrogram was constructed on the basis of the distance matrix. arion. E. 3. The aim of this study was to analyse the genetic differentiation in a few populations of two putative subspecies of M. Katalin Pecsenye1. In the analysis of the data. P: proportion of polymorphic loci. a.7 44. a.8 50. The sampled populations used either Thymus spp. ** significant at 0.70 H 0.139 0. summer TR (3/3) Total (14/7) N 21. H-4010 Debrecen. Thomas 2005 differentiation in the HungarianonMaculineaand Conservation of Butterflies(Eds)199 Studies the Ecology arion populations in Europe Vol.159** 0.klte.79 P 41. arion samples originated from seven populations in Hungary and in Transylvania. a.O. Department of Evolutionary Zoology and Human Biology. Parameters of polymorphism in the Maculinea arion populations averaged for the spring and summer types in Hungary and for the summer type in Transylvania. 3.8 1. Genotype and allele frequencies were estimated on the basis of the enzyme pattern. especially from mountainous areas.© PENSOFT Publishers Patterns of genetic Sofia – Moscow J. the summer type.169 0. Judit Bereczki1 & Zoltán Varga1. P. a. Hungary Contact:
[email protected] 0.2 17.2 1 DUFS . (M.01 level.69 nA 1. Several subspecies have been described. Numbers in brackets indicate the number of samples and the number of populations sampled. ligurica. the parameters of polymorphism were first estimated and Fstatistics were computed where the total genetic variation was partitioned into within and between population components. a. Hungary 2 HAS-UD Evolutionary Genetics and Conservation Biology Research Group.148 FIS 0. summer type HU.001 level. all samples were included. indicating strong differentiation among them. The level of variation was relatively high in the M. FST values and the results of exact test estimated for summer and spring types of M. was carried out using the allele frequencies of the samples to show the size of overlap in the genetic variation of the populations in a reduced space of variables. Both the proportion of polymorphic loci and the average heterozygote frequency were the highest among the Maculinea species. while in the second computation.131** Exact test *** *** *** *** . In the first analysis. however. Our results indicated a high level of genetic variation in the Large Blue populations. summer type TR) was quite sizeable (5%). Hierarchical F-statistics were carried out in two parts. summer HU (8) M a. ** significant at 0. The summer and spring type populations did not differ significantly in any of these parameters. which was the highest among the Maculinea species. arion in the two regions Hungary and Transylvania. The dendrogram constructed on the basis of Nei’s genetic distances did not show any specific pattern in the genetic structure of the populations. arion populations. arion populations practically exist as large. M.200 Andrea Tóth et al. This indicated a fairly high temporal variation within these populations. The results of both analyses indicated that a sizeable portion of the total between sample variation was explained by the differences among the samples collected in different years from the same population. the differentiation between the two types (summer and spring type) only explained 1. open metapopulation systems with effective gene flow between neighbouring habitat patches. arion in Hungary. It was also clear that the differentiation of the Transylvanian samples was due to a sample from one population. In the second analysis. the samples originating from the same population but collected in different years were clustered in separate branches. that is the temporal component of the between sample variation was quite high (Figure 1). This suggests that the two putative subspecies were not differentiated genetically.7% of the total variance. the Hungarian samples were only considered. summer TR (3) Total (15) FST 0. arion collected in Hungary were not differentiated from each other. The results of F-statistics showed that most of the total genetic variation (FIT) occurred between the populations (FST). The level of differentiation within the putative subspecies using different initial food plant was similar to that among all populations. A possible explanation of this phenomenon might be the population structure of the species. a. The high level of variation within the populations was coupled with strong differentiation among Table 2. It thus appears that the Hungarian and Transylvanian samples were more differentiated from each other than the two types of M. Subspecies M. The Hungarian samples of the spring and summer type M. *** significant at 0. Numbers in brackets indicate the number of samples. When all samples were involved the differentiation among the groups (spring type HU. Moreover. The samples of the two types of M. spring HU (3) M.01 level. The results of the two analyses were different.110** 0. a. The result of the PCA analysis fully supported those of the hierarchical F-statistics.147** 0.128** 0. arion were scattered in different branches of the dendrogram. the variation among samples originating from different generations of the same population was also sizeable suggesting a fairly strong effect of genetic drift.Patterns of genetic differentiation in the Hungarian Maculinea arion populations 201 Fig. could not detect genetic differentiation between the spring and the summer type populations collected in Hungary. We. Moreover. 2. . however. which is a general feature of the Maculinea species. Each point represents a sample in the reduced space of variables. Fig. Moreover. two of the three Transylvanian populations seemed to be fairly similar to the Hungarian ones. Distribution of variation at different levels of the hierarchy estimated by computing AMOVA. 1. Nevertheless. Results of the PCA analyses. them. arion populations resemble those in Sweden and the UK in possessing annual and biennial-developing larvae (Schonrogge et al 2000). This effect might also be amplified if Hungarian M. one Transylvanian population was clearly differentiated from all the others. J. (2000) Polymorphic growth rates in myrmecophilous insects.A. 771-777 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). J.C. & Elmes. REFERENCE Schönrogge. Thomas.. Proc Roy Soc B 267. K. Wardlaw. G.W.202 Andrea Tóth et al. . Chaetocnema conducta (Motschulsky) and its Kindred Species in the Afrotropical Region 203 Section 3. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Conservation and management for Maculinea .5. Martin Musche.204 Christian Anton. & Josef Settele This page intentionally left blank . The package consists of posters and badges produced at the Academy of Fine Arts in Kraków and a floppy disc/CD with the information on the MacMan project goals and organization of work. The packages were distributed to 31 primary schools. The teachers who already carried out the lessons submitted the records of activities in a form of reports and photos.uj. p. 30-387 Kraków. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126).edu. 205 MacMan at schools Anna Amirowicz & Michal Woyciechowski UJAG . accordingly to the project schedule. Maculinea butterflies biology. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)205 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 17 secondary schools. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. . The guidance for activities was prepared in a form of an educational package.pl The idea of the MacMan project was disseminated at schools in 2003. Kühn J. 18 middle schools. The promotion will continue in 2004. Institute of Environmental Sciences. Settele.A. Poland Contact: amir@eko.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. E. 7 joint schools and 7 kindergartens in 53 localities of Poland. Gronostajowa 7.Jagiellonian University. the importance and ways of biodiversity conservation as well as exercises for a warm-up and to test students’ knowledge of the subject. 04318 Leipzig. Department of Community Ecology. Kühn & J. All Farms were assumed to have milk quota of 300. Using Linear Programming Methods. D-37073 Göttingen. Department of Economics. 206-209 Protection of low yielding Sanguisorba officinalis grasslands as habitat of the Large Blue butterflies Maculinea nausithous and Maculinea teleius – model calculations on the efficiency of agri-environmental schemes Holger Bergmann1. the objective of this study is to analyze whether existing agri-environmental schemes are the cheapest way to protect extensive grasslands (and thus their butterflies). The land use of the farms was as follows: . 4.A. The conservation of rare species like Maculinea nausithous and M.000 kg/year and use mother cows as a possible alternative use of grassland. 04318 Leipzig. Germany Contact: hbergma1@gwdg. Germany 3 UFZ – Centre for Environmental Research Leipzig-Halle. E. 15.© PENSOFT Publishers 206 Holger Bergmann et al. Josef Settele3 & Frank Wätzold4 Institute of Agricultural Economics. under existing market conditions and production needs. Department of Ecological Modelling. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Germany 4 UFZ – Centre for Environmental Research Leipzig-Halle. Sofia – Moscow J. For this reason the European Union is paying specific support grants to make the use of extensive grasslands economically more attractive. Permoserstr. However. in times of budgetary scarcity the ecological effectiveness of measures must be supplemented by economic efficiency. University of Göttingen. Germany ² UFZ – Centre for Environmental Research Leipzig-Halle. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Karin Johst². Theodor-Lieser-Str. MATERIALS AND METHODS Four different dairy farm organisations were used in this study (Bergmann 2004). However.de 1 INTRODUCTION The farming of grassland in agriculturally unproductive areas makes an important contribution to the protection of butterflies all over Europe. Settele. Permoserstr. the sum of standard gross margins per production unit were maximised. Martin Drechsler². pp. 06120 Halle. teleius on extensively used grasslands plays a key role for the implementation of the Habitats’ Directive and thus contributes especially to the Natura 2000 network. 15. the use of low yielding grasslands is not economically efficient. Using Southwest-German Rhineland-Palatinate as an example. Platz der Göttinger Sieben 5. To determine the compensation payments for the profit lost in adopting alternative mowing regimes for farmers. Based on a literature analysis. Due to its low quality. Effect of different mowing dates on MJ NEL/kg. fodder with content between 4 and 6 MJ NEL per kg DM. we calculated that fodder with an energy content of 6 or more MJ NEL per kg DM (=dry matter) was usable in intensive cattle production. The calculated yields per ha have been qualified by having a closer look at the requirements of cattle nutrition. Using specific literature describing cattle and horse nutrition. the impacts on quality and quantity of extensification measures had been described (Bergmann 2004). and we calculated the compensation payments accordingly. that most farmers in the region used meadows for silage. 1. silage harvested with a first cut in August cannot be used in cattle nutrition. the necessary calculations were based on silage production with a base energy yield per ha of 52 GJ NEL/ha and year. Fig. Due to the fact. the method of standard gross margin calculations has been used. The calculations had to be made by taking into account the fact that the silage as well as the hay produced from grassland under nature protection constraints has limited use in dairy farms. It was assumed that the farmers purchase concentrates as an additional fodder to compensate for the loss of energy yields. was mostly usable in horse and heffer nutrition. DM/ha and their usability Source: Own Calculations based on OPITZ VON BOBERFELD 1994. However. as figure 1 shows.Protection of low yielding Sanguisorba officinalis grasslands as habitat of the Large Blue Farm I Farm II Farm III Farm IV 45 ha arable farm land & 5 ha grassland 25 ha arable farm land & 25 ha grassland 10 ha arable farm land & 40 ha grassland 5 ha arable farm land & 45 ha grassland 207 The level of compensation needed was calculated to test whether the existing agri-environmental scheme is capable of compensating farmers for the losses through the extensification. Based on a function by Opitz von Boberfeld (1994) the effects of different cutting regimes on yield quality and quantity have been calculated.262 . 112 € 16.00 7.00 10.05 4.00 7.332 € 14.00 0.208 Holger Bergmann et al.00 0.11 4.52 29. In addition the use of the products .52 30.00 7.00 7. barley. RESULTS The existing agri-environmental scheme in Rhineland-Palatinate (Germany) pays 204 €/ha for one cut per year after June 14th.69 0.00 8.00 35.50 0. Table 1 shows some of the results summarized for all four farms. Therefore. for mowing regimes with a first cut later than the beginning of August.05 115.00 18.00 24. Production effects of extensification premiums Crop 0 Wheat Rye Barley Triticale Rape Maize (Silage) Fallow Intensive Grassland Extensive Grassland Mother cows Specific farm cost in Sum Costs paid with premium 42.52 0.00 2.75 78.967 € 22.52 0.was not recommended for either cattle production or for horse nutrition.13 68.29 4.) is paid for extensive grassland to protect biodiversity.00 17.25 0.29 0.52 30.00 0. wheat and rape is decreasing and the needed area for maize used in dairy nutrition is increasing.00 25.215 € 225 35.38 54.663 € 11.80 0. Table 1 should be read in the following way: In the second column is the reference level of the calculations. There is paid no premium on nature conservation and the dairy farms produce the most profitable crops.00 0.50 0. 204 €. when the total loss of usability was presumed and a compensation of 1156 €/ha was calculated.etc. The effect is that with increasing level of payments the proportion of barley.00 5. and a third stage after 1 August.542 € Tendency .05 46.91 16.215 € 17.50 0.00 0.422 € 235 34. The results indicated that the existing payment was high enough to compensate dairy farms with a high proportion of grasslands.91 0. The curve of calculated compensations (as figure 1 shows) therefore had three different stages: a first stage until 15 June with compensation amounts about a 200 €/ha. While mixed farm organisations like Farm I and II needed a fixed compensation amount (~232 €/ha).50 110. but not dairy farms with a high proportion of arable land.16 0 0 100 40. farms III and IV required varying amounts depending on their acceptance of the program (from a 0 to a 354 €/ha).00 7.88 0.91 4. wheat.apart from the nutrition of heffers between the 12th and 18th living months .52 0. a second stage from 15 June until 1 August with compensation costs increasing from about a 200 €/ha to 1000 €/ha.96 4.00 10.62 4.91 4.63 0.00 10.00 7.05 36. farmers had to be compensated for the complete loss of usability of the meadow. In the third and the following columns is shown what is happening with the farm production structures if a premium of 100 €/ha (150 €.05 60.051 € 204 35.69 41.05 34.00 8.50 80.144 € Premium supposed 150 36. rape and maize for cattle nutrition. Table 1. The reference level in the calculations has been a first cutting date after 14 June each year.00 2. : Integrating Efficient Grassland Farming and Biodiversity.25 ha of 200 ha are needed for nature conservation purposes. observ. H. R.. The comparison shows that if less than 78. UFZ Berichte 2/2004. Bertke. 62pp. J. that auctioning a system “…makes auctions a valuable tool for governments to use in coping. C. which are known to have negative external effects (Scheringer 2002) and which in the study region require further land area formerly used as grasslands (pers. Furthermore the extensification of grassland will result in an increase of highly intensive conventional maize production or other intensive production systems. p. an example how this can be done is developed by Bertke et al. Scheringer. 336pp.. R. (2004): Berechnung von Kosten für Maßnahmen zum Schutz von gefährdeten MaculineaArten.88. Volume 10. 2005). Tute..). A. if only a part of the area is needed to fulfil specific ecological goals (e. Van der Hamsvoort.) ha of grassland.. it is cheaper to use a farm specific scheme than the actual scheme which pays 204 €/ha.. it is cheaper to implement farm specific schemes by using auctioning systems rather than general mean compensation schemes. R. 153pp. (2005): An outcome-based payment scheme for the promotion of biodiversity in the cultural landscape.. American Journal of Agricultural Economics 79. Grassland Science in Europe. The row “specific farm costs in Sum” shows how much money is needed to compensate the farmers loses by integrating the necessary farm production adjustments.” ACKNOWLEDGEMENTS The study was co-financed by UFZ .. (1997): Auctioning Conservation Contracts: A Theoretical Analysis and an Application.. Ulmer. Hence.63 (68. (2002): Nitrogen on dairy farms: balances and efficiency. Marggraf.25. 407-418 Opitz von Boberfeld (1994): Grünlandlehre. REFERENCES Bergmann. Consequently. . Gerowitt. 78.Protection of low yielding Sanguisorba officinalis grasslands as habitat of the Large Blue 209 The last two rows give an impression of the financial costs.K. Viiralt. 36-39 Latacz-Lohmann. if the target is to extensify e. Leipzig. & Geherman. Hespelt S. it is known from Latacz-Lohmann & Van der Hamsvoort (1997). UTB 1770. Linke.g. V. U. B.g. At least in reaching specific grassland protection targets.. Tallin.Centre of Environmental Research Halle-Leipzig and the EU FP5 RTD project MacMan (EVK2-CT-2001-00126).. Stuttgart.deficiencies in allocating contracts for the provision of non-market goods. J. C. Hohengandern. Isselstein. only dairy farms with a high proportion of grassland (in this case 62%) will participate. E. Although these calculations ignore transaction costs. Excelsior. 54. The row “costs paid with premium” supposes that the existing premium of 204 €/ ha is paid for each extensified grassland hectare. etc. In: Lillak. CONCLUSIONS The results of the present study demonstrate that the existing compensations are not high enough for dairy farms with high proportions of arable land. an increase in scheme participation is directly connected to an increase in maize production and a decrease in mother cow production. E. Marcin Sielezniew2 & Anna M. nausithous are widely distributed in the southern part of Poland and have stable populations. Poland Contact: sielezniew@alpha. About one hundred grid squares are shared with M.J. Gagarina 9. 02-776 Warszawa. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Often the habitats consist of a mosaic vegetation with significant coverage of trees and bushes (Stankiewicz & Sielezniew. nausithous in Poland Jarosław Buszko1. nausithous in Poland is related generally to the range of the food-plant Sanguisorba officinalis. . officinalis in the south of the country provides the discovery of one or both species. Stankiewicz3 1 Nicolas Copernicus University. The northernmost localities reach the vicinity of Warsaw. and on transition zones between very wet areas dominated by Carex or Phragmites and xerothermic meadows or pastures. Current practic shows that practically each new finding of a meadow with S. Thomas © PENSOFT Publishers 210 Jarosław Buszko.sggw. 00-679 Warszawa. StankiewiczSettele. Filipendulo-geranietum or Arrhenaterion. teleius. Salicetum pentandro-cinerae association (Stankiewicz & Sielezniew. M. nausithous is much rarer than M. In fen communities. In the hilly areas in south-eastern Poland they also thrive on slopes with bushes but this situation is rather exceptional. Its northern range limit is distinct and easy detectable. nausithous was recorded in over 160 10 km grid UTM squares across southern Poland. Marcin Sielezniew & Anna M. 210-213 The distribution and ecology of Maculinea teleius and M. where the species almost always occur in the same site. 87-100 Toruń. Compared to M. Poland 3 Museum and Institute of Zoology. In open Maculinea habitats M. where both species are declining more or less severely (van Swaay & Warren. teleius or absent.Warsaw Agriculture University. Institute of Ecology and Environmental Protection. nausithous prefers taller vegetation with trees and bushes of e. 2002).g. Poland 2 SGGW . Nowoursynowska 159. Wilcza 64. teleius. Their national status is estimated as ‘least concern’ (Buszko. Polish Academy of Sciences. which is very local and scarce in northern Poland. Two isolated colonies were also discovered in Puszcza Knyszyńska Forest in north-eastern Poland. The butterflies occur in damp grassy habitats covered by vegetation which can be classified as: Molinion. 1999). 2002).waw.A. Department of Applied Entomology. An isolated site near Chełmno is the northernmost locality of this species in western and central Europe. Kühn & of Butterflies(Eds) 2005 Studies on the Ecology and Conservation in Europe Sofia – Moscow Vol. Department of Animal Ecology. Thus the resources of the species are certainly still under-estimated in Poland. teleius has been recorded in over 150 10 km grid squares so far.pl Maculinea teleius and M. The distribution of M. J. M. they occur on patches of raised land with Molinietum. M. teleius and M. including those with calciumrich soils. 2002) in contrast to many other European countries. pp. The distribution and ecology of Maculinea teleius and M. 2. The present distribution of Maculinea nausithous in Poland . especially in the northern part of its range. nausithous in Poland 211 The adults fly in July and August. The present distribution of Maculinea teleius in Poland Fig. 1. teleius is sympatric with M. In sites where M. In warm seasons. some specimens are already on the wing in mid-June while during cold summers the species can be observed as late as September. teleius adults feed on various flow- Fig. nausithous the former usually appears a little earlier. Although M. in southern Poland. The most important potential threat is intensification of agriculture. 2004. rubra nests in southern Poland. 2002). nausithous is concerned S. As far as M. rubra prefers taller vegetation than M. teleius. gallienii (Stankiewicz & Sielezniew. Open. gallienii – two alternative hosts of M. nausithous is also rare or absent.. M. the undergoing of succession leads finally to the deterioration of habitat quality. Lythrum salicaria.) emerged from pupae collected in M. or higher host specificity. 2002) and is the most specific of all Maculinea species. parasitizing larvae of M. Viccia spp. Many wet meadows undergo inappropriate management (e. rubra. However. ruginodis.g. Eupatorium cannabinum. teleius and M.g. nausithous. officinalis range may be explained by lower dispersal abilities. Urbanization affects some Maculinea populations. but it can also be scattered throughout vegetation. Succisa pratensis. nausithous (Wynhoff. which may contribute to the deterioration of their biotopes. officinalis often forms fields with ten or more individuals per square meter. Apparently the most important factor determining the population size of the butterfly is the number of host ant nests and their distribution. Arrenatherion. Ichneumonidae). which is not used as a host ant either by M. scabrinodis is used (Thomas et al. officinalis is undoubtedly the main plant of adult butterflies.. Unfortunately the lack of traits enabling us to distinguish M. Neotypus pusillus (Hymenoptera. and E. Only males were rarely observed feeding also on S.g. tinctoria. rubra is more abundant.. teleius develops successfully in Poland in colonies of three ant species M. rubra is rare. Stankiewicz & Sielezniew. The sites differ significantly as regards the density of larval food plant..212 Jarosław Buszko. rubra. M. especially in southern Poland. nausithous larvae inside were observed. Trifolium spp. e. rubra (Figurny & Tomaszewicz. scabrinodis and M. this species is rather rare on those sites. teleius or M. Perhaps the butterfly-ant relationship resembles that in Western Europe where mainly M. 1997. On S. . this species seems to be a more effective host (Figurny & Tomaszewicz. it seems to prefer Vicia spp. early mowing) after recent implementations of EU subsidies. e. Poland is still the European refuge of both species and some economic changes contribute temporarily and locally to an improvement in habitat conditions for Maculinea species and the expansion of the space of its sites. 1997. Females ovipositing into flowerheads with M. Marcin Sielezniew & Anna M. have been found recently in two other regions: Polesie and Upper Silesia. Stankiewicz ers available in their habitats. Less frequently the butterflies were recorded on Betonica officinalis. scabrinodis. especially in the vicinity of cities. 2004). is initially a favourable process both for the larval food plant and for the host ants. scabrinodis and drier niches than M. an increase in the proportion of M. officinalis sites with open and rather short vegetation where M. M. teleius in western Poland near the northern S. 1989). Cirsium arvense. cannabiunum. and males were reared from pupae (Stankiewicz et al. At sites with diverse and generally taller vegetation where M. Serratula tinctoria and Sanguisorba officinalis. not especially wet grasslands are dominated by M. as compared with its relative M. Many low productivity grasslands have been abandoned during two last decades. Tartally & Varga. in calcareous fens. S. Allium angulosum and other pink flowers. However. M. 2002). Just a few unidentified wasps (Ichneumon sp. 2001). nausithous in Poland are very similar to data from Hungary (Tartally & Csősz. nausithous develops in Poland exclusively in nests of M. teleius from M. Observations concerning host ant specificity both of M. Relaxation of mowing and grazing pressure or even cessation of any forms of management. nausithous recorded a few dozen years ago from Lower Silesia. 2005). Generally. nausithous at this stage makes any statement impossible. Stankiewicz & Sielezniew. or Calthion communities are replaced by tall herbs and grasses of the Filipendulion type of vegetation. especially M. and thus to their decline.. Absence of M. The results obtained so far show that M. & Sielezniew. Z. A. Ichneumonidae). 309-317.C. In Współczesne kierunki w ekologii – Ekologia behawioralna (eds T. (2002) Lepidoptera. Fragmenta faunistica 47. nausithous Bgstr. J. Doctoral thesis ISBN 90-5808-461-2. Lublin. G. M. (2004) Neotypus pusillus (Hymenoptera. In Red list of threatened animals in Poland (ed. C. WUMCS. M.S. A. (2001). J. Toruń. Puszkar). 80-87. Elmes. (Lepidoptera: Lycaenidae) the new insight. J. 425-457. (2004) Data on the ant hosts of the Maculinea butterflies (Lepidoptera: Lycaenidae) of Hungary. Kraków. Puszkar & L. A. 7. Strasbourg. (1999) Red Data Book of European butterflies (Rhopalocera). 115-120. S. (1997) A distribution atlas of butterflies in Poland 1986-1995. Wageningen Agricultural University. Z. pp. Természetvédelmi Közlemények 11. (1997) Parasitism of Maculinea teleius (Lepidoptera: Lycaenidae) and M. [in Polish]..A. The Netherlands. (2002) Host specificity of Maculinea teleius Bgstr. nausithous in Poland REFERENCES 213 Buszko. I. (1989) Host specificity among Maculinea butterflies in Myrmica ant nests. & Varga. Głowaciński).M. & Nowacki. Stankiewicz. A. Oecologia 79. Sielezniew. Tartally. [in Hungarian] Tartally. No 99. J. Annales Zoologici 53. Nature and Environment. E.M. Council of Europe Publishing.The distribution and ecology of Maculinea teleius and M. 55-59.A.179-184. (2005) Myrmica rubra (Hymenoptera: Formicidae): the first data on host-ant specificity of Maculinea nausithous (Lepidoptera: Lycaenidae) in Hungary Myrmecologische Nachrichten. 403-409. and M. M. Poland [in Polish] Stankiewicz. Thomas. J. & Warren. Figurny-Puchalska. Wynhoff. At Home on foreign meadows: the reintroduction of two Maculinea butterfly species. endoparasite of Maculinea nausithous (Lepidoptera: Lycaenidae) in Poland: new data on distribution with remarks on its biology. Wardlaw. S. rubra nests. Turpress. pp. van Swaay. Buszko. J. nausithous in Myrmica scabrinodis and M. & Sawoniewicz. M.. & Tomaszewicz. & Woyciechowski.W. Instytut Ochrony Przyrody PAN. . & Csősz. Germany Contact: martin. Frank Wätzold1. 06120 Halle. Platz der Göttinger Sieben 5. the methodology is general and applicable elsewhere. and to evaluate an existing conservation scheme. Germany. p. Karin Johst1.A. Department of Ecological Modelling. Theodor-Lieser-Str. The approach is used to develop a cost-effective payment scheme for conservation of two endangered butterfly species (Maculinea nausithous and M. Holger Bergmann2 & Josef Settele3 1 3 UFZ-Centre for Environmental Research Leipzig-Halle. Although the results from our case study are specific to the area and species studied. 37073 Göttingen UFZ . 04318 Leipzig.drechsler@ufz. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. teleius) protected by the EU Habitats Directive in the region of Landau. Permoserstr. Institute of Agricultural Economics.Centre for Environmental Research Leipzig-Halle. 4. Kühn & J. to compare spatially homogeneous and heterogeneous payments. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Germany 2 University of Göttingen.© PENSOFT Publishers and Thomas Fartmann 214 Petra Dieker Sofia – Moscow J. 15. E. Department of Community Ecology. 214 A model-based approach for designing cost-effective compensation payments for the conservation of endangered species in real landscapes Martin Drechsler1. The results of the case study are used to analyse the effect of metapopulation dynamics on the cost-effectiveness of payment schemes. Settele. . 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.de An approach is presented which integrates an economic and an ecological model for designing cost-effective compensation payments for the conservation of endangered species in real landscapes. University of Stuttgart. Although this plant is present in all of the grasslands. . 06120 Halle. Keplerstr. the majority of the population in the study area lives in a few fallow meadows and along the banks of small brooks and open drainage channels. all patches with flowering Sanguisorba officinalis were mapped in Juli/August 1987 (Settele & Geißler 1988). This management not only results in low densities of the host ant. but also by cutting the plants in the period of larval development in the plant. the butterfly’s caterpillars cannot survive. where two different strategies are used to create a connection of habitats along brook banks and open drains. From 1987 to 1995 the number of individuals was recorded by transect counts in every patch once a year during the flight period. The main subpopulations in the ‘Filder’ are within the vicinity of two villages. In one case the mowing regime was adapted to the requirements of the larvae in the Sanguisorba-plants.Centre for Environmental Research Leipzig-Halle.settele @ufz.de 1 INTRODUCTION In the ‘Filder’ (near the city of Stuttgart) in Southwestern-Germany a large metapopulation of a few thousand individuals of Maculinea nausithous existed in 1987. Kühn & J. This is mainly due to mowing. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.Maculinea nausithous (Eds)215 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 4. METHODS In an area of about 60km². Germany Contact: Josef.A. Giselher Kaule1 & Josef Settele2 Institute of Landscape Planning and Ecology. The detailed results of this research work have been published in German language by Geißler-Strobel et al. 215-217 Effects of connecting strategies on the Large Blue Butterfly Maculinea nausithous – a case study Sabine Geißler-Strobel1.© PENSOFT Publishers Effects of connecting Sofia – Moscow Settele. 138 potential patches were registered. Theodor-Lieser-Str. Thomas 2005 strategies on the Large Blue ButterflyJ. Germany 2 UFZ . 70174 Stuttgart. where the hostplant Sanguisorba officinalis grows. (2000). E. in the other bushes and trees have been planted along the brooks in 1989 and since then mowing of grassy vegetation has been abandoned. Department of Community Ecology. Myrmica rubra (Johst et al in press). pp. which in these meadows normally occurs twice or three times a year. Generally the species’ habitat consists of wet or moist grasslands and meadows. 10. the proportion of inhabited patches increased. 31 by 142 individuals in 1995. The aim of connecting strategies need not to be the connectedness (spatial neighbourhood) of habitats with similar physical conditions and structural elements. as are presently often implemented in Germany. Development of transect-counted Maculinea nausithous in the two villages with different connecting strategies . In the village with adapted mowing regimes the number of individuals has doubled. In the village connecting brook banks with trees and bushes the number of observed individuals has been reduced by 50% (Figure 1) and the percentage of inhabited patches decreased drastically between 1987 and 1995. In 30 of 62 patches Sanguisorba officinalis disappeared. but a sufficient connectivity as a spatial arrangement of habitats for particular species (KLEYER et al. Other factors have been an extention of housing areas (n = 6) and drastic land use changes in moist meadows (n = 2). however there were only 10 patches with 56 individuals in 1995. In this case study we found a very high incidence of the butterfly where groups of 7 or more suitable patches are not further than 800m apart from the next occupied group of patches (GEISSLER 1998). 2. Unspecific connecting systems. CONCLUSIONS 1. Thirty one patches were inhabited by 122 Maculinea nausithous individuals in 1987. 1. The main reasons for the decline of Sanguisorba officinalis were changes in the microclimate caused by the shading due to young trees and bushes (n = 9) and abandoned mowing between the planted trees (n = 10). and only two of 78 patches were lost. may even destroy core populations of endangered species! Fig. 1996). Giselher Kaule & Josef Settele RESULTS The two different strategies of connecting brook banks and open drains caused different developments in the metapopulation of Maculinea nausithous (Figure 1). 24 patches were inhabited by 65 individuals in 1987.216 Sabine Geißler-Strobel. Settele. M..).Natur und Landschaft 63 (11): 467-470. Geißler-Strobel. P. G. Species Survival in Fragmented Landscapes. Margules..D.Ph. S.In: J. & S.Effects of connecting strategies on the Large Blue Butterfly Maculinea nausithous REFERENCES 217 Settele. S. Grabenpflege und Biotopverbund im Filderraum. Kluwer. Journal of Applied Ecology. Poschlod & K. C. (1998): Landschaftsplanungsorientierte Studien zu Ökologie. University of Hohenheim. M. Kaule & J. & Thomas. Kleyer. . Settele (2000): Gefährdet Biotopverbund Tierarten? – Langzeitstudie zu einer Metapopulation des Dunklen Wiesenknopf-Ameisenbläulings und Diskussion genereller Aspekte. A.. J. J. (in press) Influence of mowing on the persistence of two endangered Large Blue Butterfly (Maculinea) species. . Henle (eds. Johst. Kaule & J. . Verbreitung. Faculty of Plant Production and Landscape Ecology. J. G. 293-299. Dordrecht: 138-151. K. Geißler (1988): Schutz des vom Aussterben bedrohten Blauschwarzen Moorbläulings durch Brachenerhalt. Settele. Germany. Settele (1996): Landscape fragmentation and landscape planning. — Naturschutz und Landschaftsplanung 32. Geißler. Drechsler. Gefährdung und Schutz der Wiesenknopf-Ameisen-Bläulinge Glaucopsyche (Maculinea) nausithous und Glaucopsyche (Maculinea) teleius. thesis. .. johst@ufz. pp.de. 04318 Leipzig. CEH Dorset. mowing once a year. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Germany 2 NERC . E. As it is not obvious how the timing and frequency of mowing impact on population dynamics. Dorchester. or every second or third year before or after the flight period was appropriate for both species in a variety of landscapes. Winfrith Newburgh. fax: 0341-2353500 1 Conservation biologists have argued that mowing is detrimental for population persistence of two endangered butterfly species Maculinea nausithous and Maculinea teleius. Settele. .© PENSOFT Publishers 218 Thomas Hovestadt et al. 15. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. UK 3 UFZ . We found that optimal mowing regimes for butterfly conservation were sensitive to landscape attributes like the influence of climate on resource availability and the level of parasitism. Winfrith Technology Centre. Department of Community Ecology. Jeremy A. Dorset DT2 8ZD. Theodor-Lieser-Str. Despite this sensitivity.A. Permoserstr. we were able to identify robust mowing regimes appropriate for a wide range of landscape attributes and to derive general management recommendations. 4.Centre for Ecology & Hydrology. Kühn & J. Sofia – Moscow J. However. Our results show that the traditional mowing regime (twice per year with the second cut during the flight period) was always detrimental for the two species at both a local (single population) and regional (metapopulation) scale. 218 Influence of mowing on the persistence of two endangered Large Blue Butterfly Maculinea species Karin Johst1. Thomas2 & Josef Settele3 UFZ-Centre for Environmental Research Leipzig-Halle. Department of Ecological Modelling. Germany Contact: karin. a simulation model was used to investigate whether and how current traditional mowing regimes could be altered to reconcile butterfly conservation with agriculture.Centre for Environmental Research Leipzig-Halle. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Martin Drechsler1. 06120 Halle. Kühn J. The market gives inappropriate signals and hence wrong land-use decisions of butterfly habitats are made. their role as pollinators and important contributors to biodiversity. Susanne Koschel1 & Josef Settele2 UFZ . Germany Contact: nele. the question of how much society is willing to pay for a certain abundance of butterflies) will be the topic of this presentation. Maculinea teleius. Maculinea nausithous and Lycaena dispar will serve as example species. Thomas 2005 Butterfly monitoring in 10 National NatureJ. The valuation of non-market benefits of these butterflies is compli- . The benefit side (i. As a consequence. Although the CV methodology has been greatly improved in the past 30 years. 219-220 The other side of the coin: the economic value of butterfly conservation Nele Lienhoop1. 15. Theodor-Lieser-Str. 04318 Leipzig. costs and benefits of such measures need to be determined and balanced. 06120 Halle. increase in the population of the butterfly species mentioned). some species are declining and face local extinctions due to inappropriate habitat conditions. pp. In essence it uses surveys to elicit people’s preferences in terms of their willingness to pay (WTP) to obtain a positive environmental change (i. 4. their value has so far hardly been reflected in the management of their habitats.e.e.© PENSOFT Publishers Sofia – Moscow Settele. Benefits of the agricultural use of meadows are perceived to be larger or more important than benefits of conservation management for the survival of rare species.Centre for Environmental Research Leipzig-Halle. An economic explanation for this decline is market failure with regards to benefits of rare butterflies. These estimates can then be averaged and aggregated to the population affected by the environmental change to obtain the economic value of the butterfly. Reserves in&France (Eds)219 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Permoserstr. The overall aim of this study is to apply the Contingent Valuation method to obtain preference estimates for at least one of the three species mentioned above. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. In order to determine the adequate (“socially optimal”) level of butterfly conservation.e. people’s preferences expressed in terms of WTP tell us the demand for a certain species. (1995). Department of Community Ecology.de 1 Although butterflies provide society with multiple benefits including “butterfly watching”.Centre for Environmental Research Leipzig-Halle. how much it costs the authorities to enforce conservation for the maintenance of butterfly populations) has been presented by Drechsler et al. In other words. Economics Department. The cost side of butterfly conservation (i. there are still concerns regarding the validity of WTP. even though butterfly conservation may be socially more desirable. E. especially when the environmental change to be valued is complex and unfamiliar.lienhoop@ufz. The Contingent Valuation method (CV) is an increasingly recommended method for valuing non-market benefits of environmental goods. Germany 2 UFZ .A. . Johst. N.. D. 2005: A model-based approach for designing cost-effective compensation payments for conservation of endangered species in real landscapes. D. 24 S. S. N. C..Diskussionspapier. conventional survey methods may not be suitable.. 2004). J. Philip. Land Use Policy (in press)... guided by an experienced moderator. MacMillan. ISSN 1436-140X. Information and Consumer Behaviour (eds: Krarup. the issue can be discussed and people have sufficient time to think about their preferences and state these in terms of their WTP at the end of the meeting.. . Lienhoop. Previous applications of Market Stall (Lienhoop and MacMillan. In order to give participants the opportunity to state well-informed and thoughtful WTP responses. Bergmann. 2002. Edward Elgar. A week-long interval (between the Market Stall meeting and a second elicitation via the telephone) allows people to give the issue further thought. and Philip. Lienhoop.. 249-262 Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). M. In: Environment.). 2005: Valuing Wilderness: Estimation of WTA/WTP Using the Market Stall Approach to Contingent Valuation. discuss with friends or family and. N. 2002: Valuing the Non-market Benefits of Wild Goose Conservation: A comparison of Interview and Group-based Approaches.. L. J. REFERENCES Drechsler. A more elaborate way of obtaining values is the application of a deliberative group-based approach. 49-59. MacMillan. B. Hanlez. Susanne Koschel & Josef Settele cated due to the lack of detailed knowledge among the general public about these particular species. D. 43 (1). F. L. H. and Alvarez-Farizo. if wanted. detailed information about the butterflies can be conveyed to participants. Ecological Economics.. Settele. called Market Stall (MacMillan et al.. 2005) have shown that people are very certain about their WTP and that the validity of estimates is very high.. 2004: New approaches to valuing environmental benefits using contingent valuation.. and MacMillan.. gather more information. Cheltenham.220 Nele Lienhoop. During the group-meetings. and Russell. Potts. K. Wätzold. UFZ. Persistence is higher in small habitat patches. e. we surveyed land-use of habitats of a grassland butterfly species of the Annexes of the Habitats Directive (Maculinea nausithous) and of surrounding grasslands (about 1800 ha) in the Upper Rhine Valley (Germany. These spatio-temporal patterns and dynamics directly translate into patterns of resource availability for organisms living in the landscape. Butterflies are frequently used as surrogates for biodiversity [1]. Rhineland-Palatinate) for four consecutive years (2002-2005). too. Thomas 2005 a butterfly’s landscape – persistencethe EcologySettele. 3). leaving host-plant flowering duration higher than in multiply managed patches. Germany Contact: Holger. Therefore. 4.g. e. g. grazing or tilling) on the patch level and thus create highly dynamic and variable spatio-temporal patterns on a larger scale. Landscape . On the other hand. Furthermore. Kühn & of Blue (Eds)221 Studies on of the Dusky Large Butterflies in Europe and Conservation Vol. Therefore. patterns and dynamics at these two scales may have different effects on species persistence. We use two different time scales for this study: a) long-term landscape composition and structure adjacent to habitat-patches (Fig.Centre for Environmental Research Leipzig-Halle. host-plants in smaller patches are disturbed less frequently. Department of Community Ecology.g. 06120 Halle. 2. With increasing host-plant flowering duration increases patch occupancy probability of M. nausithous through increased resource availability and quality [2]. Theodor-Lieser-Str. On the one hand we compare dynamics of habitat occupancy of the butterfly species at the level of single patches. 1).Loritz@ufz. patterns created by these short-term dynamics should influence species persistence differently compared to long-term landscape patterns. Smaller grassland patches are managed less frequently (Fig. amount and configuration of forests or grasslands. We compare dynamics of habitat occupancy and distribution of the butterfly in relation to spatio-temporal patterns and dynamics of resource availability occurring during the summer season. J. Therefore. e. the landscape scale. E.© PENSOFT Publishers Changing Sofia – Moscow J. Results at the single patch level show that M. 4). 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. and b) short-term spatio-temporal patterns and dynamics of resource availability occurring during each flight-period of the butterfly (Fig. we investigate landscape composition and structure comparing occupied and unoccupied host-plant patches. nausithous occupies smaller host-plant patches more frequently than larger patches (Fig. pp.de Human land-use is the main driving force in cultural landscapes.A. mowing. but also on a higher level of organization. Humans apply different land-use regimes (disturbance regimes. We hypothesise that patterns and processes of human management/farming practice influence species distribution and population abundance not only on the scale of patches. 221-224 Changing a butterfly’s landscape – persistence of the Dusky Large Blue in managed grasslands Holger Loritz & Josef Settele UFZ . [2]). 2. Map of part of the study area showing long-term composition and configuration of land-cover in 2002. 1. nausithous in 2002 (maximum: 40 days. Fig.222 Holger Loritz & Josef Settele Fig. Map of part of the study area showing short-term variation of host-plant flowering duration during flight-period of M. . number of classes of host-plant flowering duration halfed for graphical reasons). Boxplot of logarithmic area-perimeter-ratio of host-plant patches against M. 5). t-Test: p < 0. These results suggest a high importance of refuges for the host-ant.01). composition and structure adjacent to occupied patches of M. Fig.001). quartiles as box and whiskers and outliers as circles. p < 0. tTest: t = 4. 4. .622. 3. In such non-habitat of the butterfly ant populations may recover and build strongholds. small patches with lower area-perimeter-ratio can be invaded easier and quicker by ants.Changing a butterfly’s landscape – persistence of the Dusky Large Blue 223 Fig. From here ants can (re-) colonise habitat patches with host-plants of their parasite M. Differently managed grasslands. Boxplot of logarithmic area of host-plant patches against land-use frequency during summer of 2003 (different letters above whiskers indicate significant differences between classes. nausithous occupancy in 2003 (boxplot shows median as dark thick line. nausithous is on average more divers than adjacent to patches where the species is absent (Fig. forests or small tree patches and extensively managed orchard meadows occur more frequently adjacent to Maculinea-habitat and may act as ant refuges. nausithous as suggested by recent models [3]. Similarly. C. & J. J. J. H. Elisabeth Kühn and Angela Lausch for helpful comments and discussions. Sarah Gwillym. Fox. J. nausithous in 2003 (bars show averages and 95 % confidence intervalls.. M.224 Holger Loritz & Josef Settele Fig. M. pp. Greenwood. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. In: J.. & Thomas.01. (2004) Comparative Losses of British Butterflies. severity and frequency of land-use are crucial factors for the butterfly species. A.. Kühn & J.. B. n = 200) and unoccupied (absence. Lawton. Telfer. R.W. Asher. Birds and Plants and the Global Extinction Crisis.T. Elmes.D. H. This study has been funded by the EC within the RTD project “MacMan” (EVK2-CT-2001-00126). fragmentation effects and questions of scale in cultural landscapes.. . J. G. J. 5. Our data show how intrinsic (patch management practice) and extrinsic factors (landscape context) determine presence or absence of this grassland butterfly.A.. Clarke. (1994): Population dynamic consequences of direct and indirect interactions involving a large blue butterfly and its plant and red ant hosts.. D. U-Test: ** = p < 0.A. Settele. Journal of Animal Ecology 63: 375-391. We conclude that studies which consider short-term spatio-temporal dynamics of resource availability and long-term landscape context may be a promising way to enhance our understanding of (meta-)population dynamics. Science 303: 1879-1881 [2] Loritz.T. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol.. Martin Musche. Clarke. Roy. n = 716) host-plant patches by M. R. Timing. G. D. E. REFERENCES [1] Thomas.E. R. Comparison of landscape composition adjacent to occupied (presence. 225-227 [3] Hochberg.001).. Settele (2005): Effects of human land-use on availability and quality of habitats of the Dusky Large Blue butterfly.. J. ACKNOWLEDGEMENTS We thank Christian Anton. *** = p < 0. Preston. & J. g. Maculinea nausithous depends much on the flowering stages of its larval host-plant.A. 1779) in a grassland landscape (~ 1450 ha grasslands) in the Upper Rhine Valley (Rhineland-Palatinate. human land-use.© PENSOFT Publishers Sofia – Moscow J. In both years maximum . E. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. most grassland plants regenerate quickly. Most habitat with flowering host-plants is available at the end of the flight-period. the Great Burnet Sanguisorba officinalis (L.de Habitat availability and quality are strongly influenced by man in cultural landscapes. grasslands. Thomas 2005 Effects of human land-use on availabilityEcologySettele. The peak of the butterfly’s flight-period and the maximum of available patches with flowering host-plants were not synchronised and differed for three 5-day-cycles (Fig. 2]. continuously changes the landscape within one vegetation-period. data on land-use of all S. During the butterfly’s flight-period (July-August). Species living in such highly dynamic grassland landscapes face very narrow niches for survival. grassland communities are capable of fast regeneration. In grassland landscapes. officinalis-patches did not flower during the butterfly’s flight-period. After a land-use event.Loritz@ufz. Fig. Figure 3 shows increasing occupancy of hostplant patches by M. more than 40 % of S. Given the species’ biology. Kühn habitats (Eds)225 Studies on the and quality of of Butterflies in Europe and Conservation Vol. SW-Germany) during two subsequent years. Maculinea nausithous should therefore respond to different periods of flower availability. Theodor-Lieser-Str. 225-227 Effects of human land-use on availability and quality of habitats of the Dusky Large Blue butterfly Holger Loritz & Josef Settele UFZ . Human land-use alters availability of the resource in time. nausithous. on average only 56 % of the area of potential habitat was available for Maculinea butterflies as habitat in the landscape (ranging from 27 % to 74 %. altering habitats probably within a few days. Thus patches can again be considered as habitat 31-35 days after land-use occured.) where feeding. About 580 ha of land-use plots of the study area were regarded as potential habitat for M. Sanguisorba officinalis on average regenerates after 31-35 days to produce flowers (Fig. So. 06120 Halle. After disturbances. Besides presenceabsence data and quantity of butterflies and its host-plant.Centre for Environmental Research Leipzig-Halle. which serve as a primary resource for M. officinalis-patches was collected for more than 3. 1). nausithous where the larval hostplant was present. Habitat quality for the butterfly was measured as the period of available flowers of the hostplant. e. Habitat is temporally destroyed but regenerates soon after. 2). We investigated how human land-use dynamics influenced habitat availability and quality of the Dusky Large Blue butterfly Maculinea nausithous (Bergstr. mainly by mowing or grazing livestock. Human land-use on grasslands usually removes biomass completely. Germany Contact: Holger. pp. 4. often called a “mini succession” [1.5 months during summer 2002 and 2003 on a basis of 5-day-cycles. nausithous with increasing period of hostplant flower availability during flight-periods of 2002 and 2003. Department of Community Ecology. 2). mating and egg-laying takes place. where they leave the hostplant (~ 8 days for egg and instar [3]). 2. Phenology (first day of 5-day-cycle) of proportion of area with flowering host-plant S. left vertical axis. 4). nausithous butterflies (line. p < 0. 1. n=580 ha) and proportion of observed M. The influence of land-use on hostplant flowering duration turned out to be a major factor of habitat quality. n=455).226 Holger Loritz & Josef Settele Fig.01) occurs after 31-35 days after a land-use event. Land-use and regeneration of the host-plant create a mosaic-landscape of habitat . Fig. right vertical axis. More than 90 % of occupied patches in 2003 showed at least 30 days of flower availability in the previous year. Butterfly occupancy levels of hostplant patches in 2003 increase with increasing duration of flower availability during the previous year. occupancy level is around 50 % and decreases quickly below 20 % when hostplant flowers are available less than half of the main flight-period of the species. A significant change of the amount of flowering plants (spikes) relative to the previous time-step (U-Test. officinalis (bars. A similar pattern arises between following years when extra time is added to the timeframe to consider the development time of the caterpillar (Fig. Regeneration of Sanguisorba officinalis after a land-use event in 2002. This fits well with the known minimum time required for caterpillars to develop into the fourth instar. 2002. Effects of human land-use on availability and quality of habitats 227 Fig. Schuyt. REFERENCES [1] Tscharntke. Occupancy of hostplant patches by Maculinea nausithous in 2003 against periods of hostplant flower availability during flight-period of 2002 (6th July – 3rd September. T. 60 days). (1979): Plant Strategies & Vegetation Processes. Occupancy of hostplant patches by Maculinea nausithous against periods of hostplant flower availability during flight-periods of 2002 and 2003 (6th July – 14th August.A.-J. max. . max. J. 2002: >95 % and 2003: >99 % of observed specimen). (1995): Insect communities. H. [2] Grime. [3] Bink. These habitats show a very high variability of resource quality and availability. patches of different quality for the butterfly species. Haarlem. & Greiler. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126).P. More than 90 % of occupied patches showed at least 30 days of flower availability during 2002. F. 4. and grasslands. 40 days. (1992): Ecologische Atlas van de Dagvlinders van Noordwest-Europa. 3. Wiley & Sons. Annual Review of Entomology 40: 535-558. grasses. Numbers above (2002) and inside bars (2003) indicate sample size of the class. London. Fig. Except for plant height. Thomas © PENSOFT Publishers 228 Jarosław Buszko. Department of Community Ecology. In a second step we examined whether plants from both habitats show a different adaptive response to land use. occurs in a range of habitats which are characterized by different land use intensity. Similar results were obtained for the number of leaves per plant. they may generate selection pressures leading to the adaptation of populations to particular management regimes. The lack of differentiation might be due to the perennial life form of S. Our results show that lower seed weight in pastures does not reduce seedling performance. The fact that offspring from both habitats were similarly able to compensate for the loss of biomass indicates the absence of special adaptations to mowing in meadow plants. In contrast seed weight was significantly higher in plants from meadows compared to plants from pastures what resulted in a larger percentage of germination. number of leaflets per leave. the food plant of the dusky large blue butterfly Maculinea nausithous. 4. artificial mowing and control. Theodor-Lieser-Str. J. Agricultural practices may influence the allocation of resources to growth and reproduction. half of them growing in repeatedly mown meadows and pastures. respectively and measured several plant traits related to growth and reproduction.A. Germany Contact: Martin. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Sanguisorba officinalis. Further. plants from both habitats responded in a similar way to artificial cutting which took place 8 weeks after establishment.Musche@ufz. 06120 Halle. Moreover. and the size of leaves and leaflets. Plants growing in meadows produced relatively fewer shoots. officinalis and gene flow between habitats. p. The result shows that a reduced seed set in meadows is counterbalanced by a higher seed quality. The ratio of ground based leaves to the number of shoots was larger compared to plants from pastures.J. To investigate the impact of land use on resource allocation we selected 24 populations.Centre for Environmental Research Leipzig-Halle. For that purpose we grew seedlings from meadows and pastures in a common environment and attributed them to different treatments. StankiewiczSettele. Kühn & of Butterflies(Eds) 2005 Studies on the Ecology and Conservation in Europe Sofia – Moscow Vol. E. Marcin Sielezniew & Anna M. Otherwise differences might become visible in later developmental stages rather than during first year growth. Offspring originated from meadows and pastures gained a similar amount of biomass during one growing season. Meadow plants also developed fewer inflorescences per shoot and less seeds per inflorescence than plants from pastures. 228 Patterns of resource allocation and adaptive response to mowing in the plant Sanguisorba officinalis (Rosaceae) Martin Musche & Josef Settele UFZ . . the type of habitat influenced all plant traits under study. The plant is able to reproduce clonally as well as by the means of seeds. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model.de Land use has been shown to affect natural plant populations in many ways. nhmus. Depressious marshlands are prevailing. Thomas 2005 A review of population Studies on the EcologySettele. not grazed in 134 quadrats. Kiskunság National Park). so there is a little difference among groups. but also many. Four types of vegetation were identified. J.2005 and 25. grazed in 196 quadrats. Five transects (50×5 m) were pegged out at each shallow area. 118 specimens in strongly grazed quadrats and 110 specimens in not grazed quadrats were observed. Our hypothesis was that the above parameters could affect the distribution of imagoes.© PENSOFT Publishers Sofia – Moscow J. Pitfall traps were used to detect Myrmica species in every second quadrat. The number of imagoes was counted twice a day during the flight period. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. This large area is a mosaic of marshland patches and meadows. Department of Zoology. Eight of these nearby marshy patches were choosen for sampling based on their size.Hungarian Natural History Museum. title: The origin and genezis of the fauna of the Carpathian Basin: diversity. grazing intensity and soil humidity were noted and vegetation height was measured.2005. Ádám Kőrösi. E. directing from the edge of the patch perpendiculary to the meadows.A. p. contract no: 3B023-04. biogeographical hotspots and nature conservation significance. Vegetation type. Altogether 578 butterflies were counted. The study area is situated at Kunpeszér on the Hungarian Great Plain (Central Hungary.zoo. The aim of this study was to improve our knowledge on habitat use and effects of habitat management on Maculinea teleius population. Ágnes Vozár.07. The size of marshy patches and distance of a given quadrat from the edge of the marshy patch can be an important factor as well. László Peregovits HNHM . The transects were divided into 5×5 m quadrats. The number of imagoes was counted on 16 days between 31.08.13. Hungary Contact:
[email protected] Maculinea teleius is very sensitive to habitat management due to its myrmecophily and use of an early and larval host plant (Sanguisorba officinalis). Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT-200100126) and partly by the National R&D Programme. . The vegetation was strongly grazed in 70 quadrats. 229 Habitat use and effect of habitat management on Maculinea teleius Noémi Örvössy. Most specimens. which are characterized by Salix bushes reeds and reedmace. In the vegetation survey the number of Sanguisorba specimens and the number of Sanguisorba flower heads were counted in every quadrats. Péter Batáry. It seems that grazing has no strong effect on adult Maculinea teleius distribution in these conditions. 1088 Budapest. Kühn & of Butterflies(Eds)229 structure of Maculinea butterflies in Europe and Conservation Vol. Baross u. 325 were in grazed quadrats. 88-96. (with corrections of mistakes in the figures in Landschaft + Stadt 22 (4). that the species might suffer from a significant decline in population density which was not apparent from presence absence data alone. Kühn & of Butterflies(Eds) 2005 Studies on the Ecology and Conservation in Europe Sofia – Moscow Vol. p. J. StankiewiczSettele. it is likely that the intensive search for Maculinea nausithous has revealed many new inhabited sites. In conclusion. these new discoveries may well have taken place during a period of general decline which might thus have been overlooked.Settele@ufz. . Theodor-Lieser-Str. Marcin Sielezniew & Anna M. Thomas © PENSOFT Publishers 230 Jarosław Buszko. 230 How endangered is Maculinea nausithous? Josef Settele UFZ . 06120 Halle. J. E. conducted with assistance by Chris Van Swaay) however has shown. — Landschaft + Stadt 22 (3). (1990): Zur Hypothese des Bestandsrückganges von Insekten in der Bundesrepublik Deutschland: Untersuchungen zu Tagfaltern in der Pfalz und die Darstellung der Ergebnisse auf Verbreitungskarten. 4. Department of Community Ecology. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126).J. REFERENCE Settele. Germany Contact: Josef. Therefore it was also regarded as less endangered than originally thought.A. A comparison of all available data of about one century until 1987 with the ones of only one year of intensive assessment (1989) has shown twice as many occupied grids for the one year inventory (Settele 1990).de In recent years intensive research on the presence of Maculinea nausithous has shown that the species (at least in Germany) is rather widespread. A quantitative comparison with transect data (on a subset of patches. however.Centre for Environmental Research Leipzig-Halle. A comparison of presence / absence data in the Upper Rhine Valley between 1989 and 2005 has shown that Maculinea nausithous has annually been found on around 50% of potentially suitable patches. 162-163). 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. For nectar sources they visit mainly the flowers of their larval food plants and Viccia spp. pulegioides and maybe also on Origanum vulgare. Stankiewicz3 SGGW . Polish Academy of Sciences. For the first time in Poland a M.) M. which are typical for this species.A.. Nowoursynowska 159. Poland Contact: sielezniew@alpha. Jarosław Buszko2 & Anna M. 2002). Bieszczady Mts. J. In lowlands (e. Gorce Mts.pl 1 Maculinea arion is widely distributed in the south-eastern part of Poland. Wilcza 64. Gagarina 9. railway tracks and fallow lands. Poland. the rarest Myrmica species on the site in Biebrza National Park (north-eastern Poland) where the survey was carried out (Sielezniew et al. Pieniny Mts.Warsaw Agriculture University. Adults are on the wing from mid June until the beginning of August. where Thymus serpyllum is used as a food-plant.g. arion pupa was found in a M. 231-233 Maculinea arion in Poland: distribution. 2003). E. arion is the most rapid (van Swaay & Warren. It has disappeared from western regions within the last few decades (Buszko. serpyllum is not present there and eggs are deposited on T. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. In the mountains and their foothills (e.sggw. ecology on the EcologySettele. arion are distinguished in Poland. Most of the sites are concentrated in the Podlasie region (northeastern Poland) and in the Kraków-Częstochowa Upland (southern Poland). However. The soils are sandy and covered usually by Festuco-Sedetalia. lobicornis nest. T. Nevertheless. It is also observed on roadsides. arion inhabits warm grasslands and pastures. 00-679 Warszawa. Kühn & of Butterflies(Eds)231 Studies and conservation prospects in Europe and Conservation Vol. Very unusual sites are known from Biebrza National Park where M. ant species composition with more than half of the M. recorded from about a hundred 10 km grid squares. Preliminary studies has just been started. (especially Diantho-Armerietum). sabuleti colonies suggest- . arion is estimated as ‘Endangered’ (Buszko & Nowacki. 02-776 Warszawa. Department of Animal Ecology.g.waw. 2 Nicolas Copernicus University. dry land surrounded by fens. Fluctuations in abundance. Poland. Department of Applied Entomology. 1999). Even in areas where it still persists a severe decline is observed. arion in Poland remain the poorest examined compared to the four other Maculinea species. 1997). Generally two types of habitat of M.© PENSOFT Publishers Maculinea Sofia – Moscow J. population densities are usually low and only in some seasons in particular localities it is possible to see more than a dozen or so specimens per day. Thomas 2005 arion in Poland: distribution. The national status of M. Poland is one of the few countries where the decline of M. ecology and conservation prospects Marcin Sielezniew1. pp. Podlasie) and some southern uplands it inhabits dry meadows and clearings in pine forests. Institute of Ecology and Environmental Protection. make any estimation of the status of individual populations difficult.. 87-100 Toruń. 3 Museum and Institute of Zoology. arion is observed on patches of raised. Butterfly-ant relationships of M. g.. sabuleti-dependent M. but we can not exclude the possibility that expected differences in host-ant specificity reflect only geographical variation. lobicornis nests. 2004). Acquisition of increased knowledge is vital for conservation of the butterfly in Poland and the whole of central and eastern Europe.. the M. Nevertheless. the existence of M. arion is more probable in the grassy habitats of highland regions than on the sandy sites of Podlasie or Polesie regions in the eastern part of the country. type and structure of vegetation) in various climates (Thomas et al. arion may be genetically differentiated. arion in Palearctic (Als et al.. We suspect that at least some Polish populations living in the north-eastern part of the country are characterized by different host-ant specificity in comparison to M. Local adaptations and coevolutions have already been recorded in other representatives of the genus Maculinea.. 1. On a site in the Kraków-Częstochowa Upland we found two pupae and one prepupa in a M.232 Marcin Sielezniew. Considering these results we also suspect that Polish populations of M. The latest genetic studies suggest the existence of a few cryptic species of M. may occur in southern Poland. arion race. Analyzing the vegetation structure. 1994) and M. 2004). arion dependent on the same host M. rebeli (Steiner et al. Jarosław Buszko & Anna M. 2003). 1998). However. This conclusion follows the model of viable population described by Thomas (1995). alcon (Elmes et al. like in Western Europe. regarding the fact that in some very sandy localities in the region M. especially M. sabuleti colony. arion populations studied so far in western Europe. dependent on its ‘classic’ host. sabuleti is rare or completely absent. The present distribution of Maculinea arion in Poland . further records of M. sabuleti had different ecological demands (e. Stankiewicz ed rather that the last species is accepted as the main host. and in other predatory Maculinea species these correlate with ant host switches (Thomas & Settele. arion early stages in M. Studies proved that even various populations of M. Fig. as well as in colonies of other non-sabuleti hosts suggest a more complicated situation in the region. This is connected with the microhabitat preferences of different Myrmica species. Sielezniew.J. Sielezniew. & Clarke.T.. Council of Europe Publishing. N. 386-390. (1994) Differences in host-ant specificity between Spanish.C. in comparison to other Maculinea butterflies. Schlick-Steiner. A..M. J. D. Elmes. G. 7. & Settele. Urgent action should be undertaken to counteract the alarming situation. Głowaciński). [in Polish]. ecology and conservation prospects 233 The main threat for M. are probably required to support viable populations of M. Thomas.Maculinea arion in Poland: distribution. 55-68. N. Hence isolation and fragmentation of habitats are also important negative factors.E. J. M. A.G.. & Warren. Journal of Insect Conservation 2. J. Buszko. M. altitude and climate on the habitat and conservation of the endangered butterfly Maculinea arion and its Myrmica ant hosts. J.. B.E. but they should be rooted in a deep knowledge of species ecology in Poland and a clear determination of negative trends. Manguira. G.. Yen. Although larval food plants are common.A. A. J.. Thomas.. Vila. C. arion indicates a rather low quality of habitat on most of the sites. Stankiewicz. (2003) First observation of one Maculinea arion pupa in a Myrmica lobicornis nest in Poland.. A. Nature and Environment. M. pp. A. Nature 432. A low density of M.W.R. 39-46.. No 99. Relatively large areas. J. as well as increasing general eutrophication of the environment.. arion.. 80-87. Nash. which is a common practice on sandy wastelands. REFERENCES Als. (2004) Butterfly mimics of ants. (2003) Host specificity revisited: New data on Myrmica host ants of the lycaenid butterfly Maculinea rebeli. Buszko. considering that even obtaining knowledge comparable to that which we have about other Large Blues in Poland is extremely time-consuming and laborious.A. Z. & van der Made J. 1-6. Journal of Insect Conservation. (2004) The evolution of alternative parasitic life histories in large blue butterflies. Hammarstedt. Höttinger.. R. & Pierce. & Bystrowski. host ants are less numerous and are restricted only to local patches within sites. 249-250. Kandul. J. Hsu. J. (2002) Lepidoptera.M. Toruñ. J. Thomas. 289-284.) (Lepidoptera). Simcox. Nature 432. Chapman and Hall. L. Kraków. Stankiewicz.. Dutch and Swedish populations of the endangered butterfly Maculinea alcon (Denis et Schiff. (1997) A distribution atlas of butterflies in Poland 1986-1995. Martin. Mignault. Pullin.. van Swaay. Instytut Ochrony Przyrody PAN.J. Turpress. Steiner. and Górnicki. Nota lepidopterologica 25.. (1995) The ecology and conservation of Maculinea arion and other European species of large blue butterfly. (1998): Effects of latitude. & Nowacki. T...W. Elmes. . Wardlaw. M. F.A. (1999) Red Data Book of European butterflies (Rhopalocera). M. arion in Poland is habitat loss resulting from the abandonment of grazing and excessive afforestration with conifers.S. In Red list of threatened animals in Poland (ed.D. In Ecology and Conservation of Butterflies (A. ed). Thomas. R. Boomsma. Hochberg.A. It will be a difficult challenge.C. London. J. Strasbourg. D. J. C.S. Memorabilia Zoologica 48. H. O. the genus Maculinea is the most specialised.© PENSOFT Publishers 234 David J. (ii) manipulations had occurred for long enough (20-30 years) for the results to impact on the community. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. pages 26-27) Although the concept of targeting conservation on a single umbrella species is theoretically attractive (Simberlof 1998). and that project gives hope that the concept of umbrella species is . Insect populations have also proved hard to conserve. Department of Community Ecology. and have often disappeared from landscapes in which their ecosystems and resources remain intact (Thomas 1984.Centre for Environmental Research Leipzig-Halle. Thomas et al 2004). Elmes1. especially myrmecophiles or myrmecophobes (Randle et al. Josef Settele2 & Jeremy A.A. Simcox1. Stewart in press. pp. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. apart from the case of Maculinea arion in the UK. or because the Myrmica ants impact directly on other species. 1991. Settele. Graham W. Thomas1 NERC . and (iii) the impacts on other species had been scientifically monitored for the duration of the project. for example at the Royal Entomological Society’s 2005 symposium “Insect Conservation Biology” (e. to date. 4. Thomas (in press) argued that. involving butterflies. 06120 Halle. including other endangered species. McGeoch in press. Among European butterflies. Against this. Ralph T. Dorset DT2 8ZD. which involved not only an insect but a conservation project in which the following criteria were satisfied: (i) the ecological cause of decline of the target species was understood and had been rectified by manipulating the management of whole sites or landscapes. Winfrith Technology Centre. It is also considered to have the potential to be an umbrella genus for other threatened organisms within grassland communities. UK 2 UFZ . CEH Dorset. with most successful targeted projects. Karsten Schönrogge1. due to the dependency of its five recognised species on the coexistence of specific foodplants and ants of the genus Myrmica (Thomas & Settele 2004).g. 234-237 Science and socio-economically-based management to restore species and grassland ecosystems of the Habitats Directive to degraded landscapes: the case of Maculinea arion in Britain David J. Kühn & J. endangered and challenging to conserve. The only example known to us that fulfils these criteria involves the restoration to Britain of the extinct butterfly Maculinea arion. Dorchester. E. Theodor-Lieser-Str. either because they inhabit similar (endangered) niches or habitats to each Maculinea and host Myrmica species. Sofia – Moscow J. Clarke1.Centre for Ecology & Hydrology. Simcox et al. Sheppard in press). there was no practical example of a successful conservation project on which to make this judgement. Zoe Randle1. Winfrith
[email protected] 1 European insects have generally experienced faster declines than vertebrates or plants. seeking successful examples in practice that involve insects is so elusive that the validity of the concept has been widely questioned. Germany Contact: David. 1995. this information came too late to save the last UK population of M. M. many other rare species have increased here as a consequence. ecological research was started in order to determine whether a key factor in the ecology of the species had been overlooked that might explain the failure of previous conservation measures. Maculinea arion is a globally threatened butterfly and a species of the Habitat Directive. arion from extinction. arion in two regions of S. as soon as fewer than 50% of Thymus. and the populations of its parasite Maculinea arion typically went extinct 2-5 years after grazing was abandoned. using a physiologically similar race from Sweden. Targeted management began on Dartmoor in 1975 both to improve the habitat quality of its last site and to extend the breeding area to two adjoining sites. It became extinct in Britain in 1979. In 1972. to carefully managed former sites. sabuleti was rapidly being displaced by M. These were further extended to form a total of six new breeding areas from 2002 onwards by the owner. A brief review of the rationale driving the British project is given. but a programme was soon proposed to re-establish the species. at least when a strongly myrmecophilous species is involved. and hence its caterpillars. capable of supporting at least three populations. sabuleti) and that this ant had greatly declined on UK sites because it was no longer economic for farmers to graze the thin-soiled ‘unimproved’ swards where the initial foodplant. scabrinodis and other cool-tolerant congeners. grew. Unnoticed. Atlantic Coast and Valleys project: about 3% of the grassland strip and 8% of valleys proposed for restoration are shown. Thymus praecox. 1. when the UK population had sunk to around 250 adults. Unfortunately. 1999). . Dartmoor: the original small breeding area has been improved in quality and extended to new areas. The scale of former restorations and current proposals for M. The National Trust. following 150 years of near continuous decline and after about 50 years of unsuccessful targeted conservation effort. arion was made on Dart- Fig.Science and socio-economically-based management to restore species 235 a valid one in conservation. The first re-introduction of M. coincided with the host ant (Thomas 1980. arion was host specific to a single species of Myrmica (M.W England. under sub-contract to the Macman programme (Fig 1). This revealed that M. and were the principle reason why they were originally listed as UK priority areas for conservation. arion. initial funding for a trial 15 km restoration has been submitted to the UK Heritage Lottery Fund by a . owned predominantly by conservation organisations. Again. as well as occupying both new patches created in 1975. sabuleti to benefit. climate and topography are best suited. The aim is restore the missing species. although it held a diversity of other valued species. Success in Somerset for both M. M. so that general visitors can enjoy the ambience of flower and species-rich grasslands. the flatlands were ploughed and/or fertilised for arable or ‘improved’ grass leys. sabuleti and M. change in other species was monitored on the restoration sites. were grazed to the recommended intensity (for UK climates) at the essential times of year (spring. sabuleti). Simcox et al. others as large as any known in Europe. the expert naturalist can watch rarer species.236 David J. arion. At the time of writing. intensive agriculture along this coastal strip was sustained only by EU subsidy. although much of the land is classified as a European Special Area for Conservation (SAC). which by 2005 contained many patches with high biodiversity levels. By the late 1980s. Shifting economics. arion habitat. Britain aimed for self-sufficiency in food. with Myrmica sabuleti quickly recovering to former levels and the butterfly population tracking model predictions of increase and change for the next twenty-two years. and for farmers to grow the crop for which its soils. but with the wider aim of restoring endangered communities. The Atlantic coastline grasslands had formerly been a stronghold of this butterfly. but the valleys where breeding occurred were soon too overgrown for M. also encompassing around 25 abandoned valleys (Fig 1). To date. sabuleti quickly returned and M. Gradually. due to war and other economic constraints. where many sites had been planted with conifers about 40 years previously. up to 70 km in length. In the 1990s. whilst the valleys were abandoned (Fig 1). During this programme. the region is sadly depauperate. but also in the restoration of high-quality ecosystems that included the increase of many rare species – ranging from UK Red Data Book birds and plants to butterflies. of ‘improved’ coastal grassland. autumn) for M. arion as the flagship or umbrella species. The observation that targeted management for M. National Trust. yet produced a poor return in food. For five decades from the 1930s onwards. The flat ground had not formerly been M. now provide this region with an opportunity to restore its lost ecosystems. and the reform of CAP subsidies. but which had proved uneconomic to maintain. a dependency that was highlighted when foot-and-mouth disease resulted in the coastal footpath being closed throughout 2001. namely natural wildlife. as agricultural techniques developed and. the economy of this region was driven by tourism. The Atlantic Coast and Valleys Project proposes to restore a 250m wide strip. and a large community of endangered Biodiversity Action Plan and Habitats Directive species can increase. Somerset. moor in 1983. and proved a success. smells and sounds of the countryside to its still beautiful topography. flies and cockroaches (Randle et al in press) – encouraged conservation bodies to consider more ambitious restorations that used M. Today. arion resulted not just in increased densities of all Myrmica species (especially M. By the 1990s. this success enabled plans to be implemented to restore the butterfly to three other former regions where grasslands. a consortium including the Somerset Wildlife Trust. This led to a successful bid to use UK Heritage Lottery Funds to restore characteristic calcareous grassland to about 30 sites in the Poldens Hills. Millfield school and the Clarke Trust had restored a series of grassland patches along 8 km of escarpment. arion was successfully restored to three further sites (Thomas 1999). some very small and perhaps temporary. arion and its wider community provided the rationale for a larger proposal for restorations in Devon and Cornwall. these have been colonised by 15 ‘populations’ of M. Wallingford Stewart.Science and socio-economically-based management to restore species 237 consortium containing the local council. I.. CABI. Lewis. O. Simcox. and (ii) that when this occurred. CABI. Roy.. McGeoch. O. R. by R. R. R. & Thomas. arion habitat successfully to degraded and intensively used land. H. Wallingford Simberloff. Blackwells. arion..C. Preston. J.. socio-economists. T. Wallingford Thomas. as it was with its raison de être.J. Thomas. J. G. umbrellas and keystone species: is single-species conservation passé in the landscape era? Biol Conserv 83. Stewart. The application was therefore as much concerned with addressing issues of access.. C. progress and prospects. including the chough (which went extinct despite being the emblem of Cornwall in the same year as M. at the scale of a landscape. Lewis. 1999 The large blue butterfly – a decade of progress. birds and plants and the global extinction crisis. Stewart. the wildlife restorations. O. New. T.D. D. Wallingford Thomas. 1991 Rare species conservation: case studies of European butterflies.... Stewart. In Insect Conservation Biology. Spellerberg. New. local business men. Although the restorations are driven by the requirements of Maculinea arion. R. Science 303. Symposia of the Royal Entomological Society 11: 333-353.. Eds. and thereby by those of many other species.A. D. M. J.. (in press) Myrmica ants as keystone species and Maculinea arion as an indicator of rare niches in UK grasslands. New. In Ecology and conservation of butterflies Ed. A. J. 1995 The ecology and conservation of Maculinea arion and other European species of large blue. CABI. Z. Chapman & Hall.A. Thomas. Clarke. London: Chapter 13: 180-196 Thomas. (i) that it was possible to restore M. Oxford. Ed. this proposal – which will be evaluated during winter 2005 – would never have been plausible without the previous demonstration in Somerset. J. On the other hand.A. Why did the large blue become extinct in Britain? Oryx 15: 243-247. K. farmers. The conservation of butterflies in temperate countries: past efforts and lessons for the future. B. Thomas. D. A. R. economics. 1980. BES Symposium 29: 149-197. ecologists.A. eds A. B. 1984. Fox. J. In Insect Conservation Biology. Lewis. In The scientific management of temperate communities for conservation.. J. J. British Wildlife 11: 22-27 Thomas. Ackery. Wardlaw.I.. eds A. 1998 Flagships.G. In Insect Conservation Biology.A. for the same general reasons). O. M.A. J A (in press) Future perspectives. In: Biology of Butterflies. eds A. J. Schönrogge. T. other characteristic and attractive plants and animals also returned to form the new ecosytems. T. Vane-Wright & P. D Site management or biotope management – and what are insect habitats anyway? In Insect Conservation Biology. CABI. New. REFERENCES Randle. 1879-81 . such large-scale operations cannot be the preserve of a minority group (ecologists) but must necessarily encompass the aspirations and economy of the indigenous human communities. Stewart. & Morris.T. Lawton. training and other local community benefits. D. Greenwood. Goldsmith. eds A. archaeologists. J. M (in press) Insects and bioindication: theory and progress. Telfer. J. 2004 Comparative losses of British butterflies. Academic Press. A (in press) Insect conservation in temperate biomes: issues. R. Asher. Lewis. educationists and many other interested parties. 247-257 Sheppard. J. Pullin.. Settele. 238 Grassland butterflies profit from succession but suffer from invasions – a case study from Southern Poland Piotr Skórka1. Statistical models built for individual species showed that both presence in a habitat patch and abundance could be explained mostly by the two aforementioned variables.© PENSOFT Publishers 238 Maurizio Biondi and Paola D’Alessandro Sofia – Moscow J. 06120 Halle. whereas the lowest were in fallow lands invaded by reed and goldenrod and in mature forests. L. Institute of Environmental Sciences. Germany Contact: skorasp@poczta. Theodor-Lieser-Str. thus the area should be protected. Fallow lands invaded by reed and goldenrod were also included in the analysis. E. In each stage we established three transects in which butterflies were regularly inventoried between the beginning of May and the beginning of September 2004. fallow lands.Jagiellonian University. (2) indicate that meadow restoration is still possible even several decades after abandonment and (3) show that invasion of reed and non-native goldenrods should be prevented. mean number of species per survey and a species diversity index were the highest in fallow lands. 30-387 Kraków. young forests and mature forests. old fallow lands and young forests. A few very rare species included in the Habitat Directive (Lycaena dispar.A. p. We distinguished the following stages of natural succession: extensively mown meadows. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. helle. These results (1) reveal the importance of fallow land for butterflies.Centre for Environmental Research Leipzig-Halle. we showed that the presence and relative abundance of Maculinea butterflies are good indicators of general butterfly species richness and abundance. Poland UFZ . Gronostajowa 7. nausithous) were common on the wet grasslands in Krakow. Josef Settele2 & Michal Woyciechowski1 UJAG . The differences in species number and abundance between the habitat types could be adequately explained by vegetation characteristics such as plant species number and vegetation height. . Moreover. 4. old fallow lands. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). The total number of species and individuals.pl 1 2 We investigated the effects of natural succession and invasions of reed and goldenrod on butterfly communities on wet grassland in Kraków (southern Poland). Department of Community Ecology. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Kühn & J. Maculinea teleius and M.onet. but in some species also by abundance/presence of larval food plant and flower abundance. nausithous. Consequently they indicate that the specific habitat conditions determine. teleius. Seethalerstrasse 6.Maculinea species”: M. their host plants.de.Stettmer@anl. Germany Contact: Christian. E. A main focus of our surveys was the impact of different land use and management patterns on the habitat systems the three species live in. As an associated partner within the MacMan-Project the ANL (Bavarian Academy for Nature Conservation and Landscape Management) was in charge of developing guidelines for the applied conservation and practical management of the three “wetland.de 1 Among European butterflies the fascinating and complex ecology of the genus Maculinea provides a challenging puzzle of questions.A. In addition we established management experiments. which in turn have a strong impact on wetland Maculinea populations and related host communities. investigated habitats of M. nausithous and M.Maculinea arion Butterflies(Eds)239 Studies on the Ecology and Conservation of sites in Europe Vol. M. alcon were situated in the pre-alpine region.© PENSOFT Publishers Sofia – Moscow Settele. alcon. alcon and at three sites for M. 80797 Munich. In summary we can evaluate the principal factors influencing the populations and conclude.braeu@ifuplan. p.bayern. markus. According to their main distribution in Bavaria. Our data show how different mowing regimes affect important habitat parameters. 239 Field research based development of management guidelines for the protection of species and ecosystems of the Habitats Directive – A case study of “wetland Maculinea species” in Bavaria Christian Stettmer1 & Markus Bräu2 ANL . teleius and M. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). alcon and 69 habitats for M. whereas those of both M. which kind of management can enhance long term viability of wetland Maculinea species. teleius and M.and North Bavaria. which kind of mowing regime is well suited and can be applied to a site. Landscape and Nature Conservation. which potentially could influence Maculinea populations. . These experiments were run at four sites for M. we assessed 73 habitats for M. in which different mowing regimes and their impact on Maculinea ecosystems were tested. dealing with the management and conservation of these highly endangered species. 156. Schleißheimer Str. teleius and M. nausithous were situated in South. To assess a maximum range of the various habitat types of each species. 83410 Laufen/Salzach 2 Institute for Environmental Planning. Kühn & J. and host ants as well as the vegetation structure and other factors. nausithous.Bayerische Akademie für Naturschutz und Landschaftspflege. In our studies we investigated the ecology of the three Maculinea species. Thomas 2005 Habitat preferences of Myrmica ant species in J. © PENSOFT Publishers 240 Jeremy A Thomas & David J Simcox Sofia – Moscow J. Settele, E. Kühn & J.A. Thomas (Eds) 2005 Studies on the Ecology and Conservation of Butterflies in Europe Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, pp. 240-244 Contrasting management requirements of Maculinea arion across latitudinal and altitudinal climatic gradients in west Europe Jeremy A Thomas & David J Simcox NERC Centre for Ecology & Hydrology, CEH Dorset, Winfrith Technology Centre, Dorchester, Dorset DT2 8ZD, UK Contact:
[email protected] The myrmecophilous social parasite Maculinea arion is listed in Appendix 4 of the European Habitat Directive, as “LR/nt” in the IUCN 2004 Red List for all species, and as Vulnerable in the IUCN Red List for Invertebrates (Wells & Pyle 1984). Although its populations occur more widely across Europe than those of any other species of Maculinea, large populations of M. arion are extremely rare and in several regions it appears to be experiencing an equal or greater rate of decline and local extinction than its congeners. Moreover, recent studies indicate that certain regional populations of M. arion show such major genetic differentiation that some may eventually be reclassified as distinct cryptic species, even within different parts of Europe (Als et al 2004; Thomas & Settele 2004). So far as is known, the putative sibling species within recognised morpho-species of (especially predacious) Maculinea species are characterised by ant (but not plant) host switches, with populations being adapted to exploit a different species of Myrmica ant in different parts of their ranges. If confirmed, this will increase the priority for conserving regional populations of ‘M. arion’ (Thomas & Settele 2004). Within Europe, one host-switch has been described to date for M. arion: western populations and those in south Poland exploit Myrmica sabuleti (Thomas 1977, Thomas et al 1989, 2005; Sielezniew et al 2003) whereas those in n-e Poland exploit Myrmica lobicornis (Sielezniew et al 2003). Although, like other predacious Maculinea species, M. arion larvae occasionally survive with other Myrmica species on sites, mortality is so much higher in secondary hosts that these represent patches of misleading pseudo-sources within a site, where the intrinsic growth rate (λ) of M. arion is <1: in other words, a population would rapidly decline to extinction on any site that supported only the secondary host ant species of M. arion (Thomas 2002; Thomas et al 1998a, 2005). Models and empirical data suggest that a minimum of 55% co-occurrence of primary host (M. sabuleti) and host plant is necessary for ë to exceed one for M. arion, and in practice most known populations survive only on sites where much higher host ant densities exist (Thomas & Elmes 1998). Despite this, secondary hosts are not total sinks but may play an important role for population persistence in years of extreme weather. For example, in the drought summer of 1992, we found that virtually all larval foodplants failed to flower, and hence were unsuitable for oviposition, in the areas where M. sabuleti occurred in Oland, and that the vast majority of eggs Contrasting management requirements of Maculinea arion across latitudinal 241 were laid on Origanum in neighbouring moist-soiled land where M. rubra dominated. Although the large majority of larvae failed to survive (only two individual adult M. arion were reported in 1992), at least enough survived for the population to persist and then recover within a few years once the M.sabuleti colonies were again attainable. The two regional ant hosts of M. arion in Europe inhabit different niches (Elmes et al 1998) and consequently this morpho-species will require rather different management regimes in north Poland compared to the rest of Europe. To date little is known of the precise ecological requirements of M. locibornis-exploiting populations of M. arion. Here we explore how the niche of M. sabuleti, and hence that of M. arion, varies under different climates across western Europe, and how this, in turn, means that different management regimes are required to maintain optimum habitat. On 68 occupied and unoccupied M. arion sites in west Europe, where M. sabuleti was the known host, we measured Myrmica density (% baits attracting this ant), turf height (using methodology of Stewart et al 2001) and recent management. In the UK (5 sites) we also measured soil (top 1 cm) and air temperature at each sampling point. Study sites were in four regions of UK (Cotswolds, Polden Hills, Dartmoor, Atlantic Coast of Devon and Cornwall), in Sweden (Oland) and in three regions of Central-southern France which, although of similar latitude, had very different altitudes and hence local climates: Dordogne (<200 m altitude), Cevennes (500-1000 m altitude) and Hautes Alps (1500-1800 m altitude). Within individual sites in Britain, we found the familiar gradient of ant niches corresponding to variation in grazing intensity, soil moisture and soil temperature (Elmes et al 1998), with M. schencki confined to the hottest, most sparse and shortest turf, M. sabuleti abundant in turf 1-3 cm tall, and M. cabrinoids and M. rubra or M. ruginodis dominating in progressively taller succesional stages (Fig 1). The schematic illustration shown in figure 1 is calibrated for an unusually warm UK site in the Polden Hills: more typically in the UK, and certainly during the cooler summers of the 1970s, the whole ant gradient is shifted to the left, with M. schencki usually absent (because sites are too cool) and even M. sabuleti often present only in the right-hand half of its niche curve Fig. 1. Schematic diagram of the niches occupied by Thymus and different species of Myrmica ants, calibrated for the warmest sites currently available in the UK. Data from Thomas 1977, Thomas et al (1998a, b), Elmes et al (1998) and Thomas & Simcox unpublished. Niches of the cuckoo-feeder M. rebeli (grey) and the predator M. arion (black) are shown within those of their respective hosts, M. schencki and M. sabuleti. 242 Jeremy A Thomas & David J Simcox (see Fig 2, UK). Fig 1 also shows how the initial Thymus foodplant has a much broader niche in UK grassland swards than does any single species of Myrmica, and how the niche of M. arion is narrower still, because the minimum requirement for a population to persist is for at least 55% of Fig. 2. Distribution of Myrmica sabuleti (black dots, solid black lines) in swards of different heights in three regions of Europe with different local climates. Red lines indicate the minimum density predicted for a M. arion population to persist; blue areas represent the realised niche and peaks the optimum habitat of this butterfly. From Thomas et al (1998), Bourn & Thomas (2002), and Thomas & Simcox unpublished. Contrasting management requirements of Maculinea arion across latitudinal 243 Thymus to co-exist with M. sabuleti. By contrast, the potential niche of M. rebeli (which does not occur in the UK) is broader than that of M. arion, reflecting its more efficient cuckoo-feeding exploitation of its ant host (Thomas et al 2005). Under different climates across Europe, we found major variation in the vegetation structure, and hence management, where optimum densities of M. sabuleti and M. arion occurred (Fig 2 for latitudinal variation). In the UK, as described above, optimum habitat for the host ant and butterfly was restricted to warm, south-facing slopes on certain well-drained soils in southern England, where – in addition to these abiotic attributes - the sward was grazed very short in both early spring (March-early May) and autumn (Sept-late Nov). This ensures that M. sabuleti nurse ants, when transporting their grubs into the upper cells of the nest, can attain the optimum temperatures (20-22oC) for larval growth over a long growing season encompassing 9 months of the year. Initially, such heavy grazing was not welcomed by some nature reserve managers, but in the last decade we have found that large populations of both host ant and butterfly can be supported by grasslands that are not grazed between mid-May and late August, allowing plants to flower and seed and other insects to flourish, including the early larval instars of M. arion when they feed in July on Thymus flowerheads. We also found M. sabuleti supporting M. arion under apparently identical vegetation and management conditions at much lower latitudes, but at high altitude, in the alps north of Grenoble, France. In regions of Europe where the late spring and summer (but not winter) climate is 1-2oC warmer than in the UK, we found that M. arion and the essential high densities of M. sabuleti were not restricted to south-facing slopes but occurred also on flat land. Importantly, optimum habitat was now found in lightly (e.g. extensively) grazed swards (and again in those left ungrazed after early spring), where the sward where Thymus grew was on average 3-10 cm (optimum 5 cm) tall (Fig 2). On the warmest sites in these regions a few eggs are laid on Origanum, a later-flowering foodplant of taller swards. This situation existed not only on Oland (Sweden) (Fig 2) but also at mid altitude in the Cevennes, central France. Finally, under the warm spring-summer climate of Dordogne, central-southern France, we recorded high densities of M. sabuleti and M. arion mainly on flat land where the sward was much taller and indeed shaded the ground, with optimal habitat existing where the vegetation was about 30 cm tall (Fig 2). Here, although Thymus was abundant, we found neither the ant nor butterfly on south-facing slopes, which are presumably too hot since these were dominated by xerophytic ant species. Suitable conditions in Dordogne were found along unmanaged road sides and, especially, on land that had been abandoned for probably 5-10 years, for example on former vineyards and rough patches preserved for La Chasse. Here the sward is so tall that only Origanum is employed as a foodplant. Eventually here, the foodplant, ant and butterfly are displaced in the later stages of grassland and scrub succession, and the optimum management appears to be a major perturbation to a part of the ground within each site every 5-10 years. REFERENCES Als,T. D., Vila, R., Kandul, N., Nash, D.R., Yen, H., Mignaault, A., Boomsma, J.J. & Pierce, N.E. (2004) The evolution of alternative parasitic life histories in large blue butterflies. Nature 432, 386-390 Bourn, N.A.D & Thomas, J.A. (2002) The challenge of conserving butterflies at range margins in Europe. Biol. Conserv. 104: 285-292 Elmes, G.W., Thomas, J.A., Wardlaw, J.C., Hochberg, M.E., Clarke, R.T., Simcox, D.J. 1998 The ecology of Myrmica ants in relation to the conservation of Maculinea butterflies. Journal of Insect Conservation 2: 67-78 244 Jeremy A Thomas & David J Simcox Sielezniew, M., Stankiewicz, A. & Bystrowski, C. (2003) First observation of one Maculinea arion pupa in a Myrmica lobicornis nest in Poland. Nota Lepidopterologica 25: 249-250 Stewart, K. E. J., Bourn, N. A. D. & Thomas, J. A. (2001) An evaluation of three quick methods commonly used to assess sward height in ecology. Journal of Applied Ecology 38: 1148-1154 Thomas, J.A. (1977) Ecology and conservation of the large blue butterfly 2nd Rep. 23 pp, ITE, Huntingdon. Thomas, J.A. (2002) Larval niche selection and evening exposure enhance adoption of a predacious social parasite, Maculinea arion (large blue butterfly), by Myrmica ants. Oecologia 122: 531-537 Thomas, J.A. & Elmes, G.W. 1998 Higher productivity at the cost of increased host-specificity when Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation. Ecological Entomology 23: 457-464 Thomas, J.A., Elmes, G.W., Wardlaw, J.C., Woyciechowski, M. 1989 Host specificity among Maculinea butterflies in Myrmica ant nests. Oecologia 79: 452-457 Thomas, J.A., Clarke, R.T., Elmes, G.W. & Hochberg, M.E. 1998a. Population dynamics in the genus Maculinea (Lepidoptera: Lycaenidae). In Insect population dynamics: in theory and practice. Ed by J.P. Dempster & I.F.G. McLean. Symposia of the Royal Entomological Society 19: 261-290. Chapman & Hall, London Thomas, J.A., Simcox, D.J, Wardlaw, J.C., Elmes, G.W., Hochberg, M.E. & Clarke, R.T. 1998b Effects of latitude, altitude and climate on the habitat and conservation of the endangered butterfly Maculinea arion and its Myrmica ant hosts. Journal of Insect Conservation 2: 39-46. Thomas, J.A., Schönrogge, K. & Elmes, G.W. (2005) Specialisations and host associations of social parasites of ants. In Insect Evolutionary Ecology pp 475, 514, CABI, Reading Thomas, J.A. & Settele, J. (2004) Butterfly mimics of ants Nature 432, 283-284 Wells, S. & Pyle, R.M. (1984) IUCN Red Data Book for invertebrates. IUCN, Gland Research has been part funded by the EC within the RTD project “MacMan” (EV K2-CT2001-00126). © PENSOFTThe distribution Publishers Sofia – Moscow J. J.A. Thomas 2005 and ecology of Maculinea teleiuson the EcologySettele, E. Kühn &Poland (Eds)245 Studies and M. nausithous in of Butterflies in Europe and Conservation Vol. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model, pp. 245-246 A supporting tool for decision-making in Maculinea management Karin Ulbrich1, Martin Drechsler2, Karin Johst2, Frank Wätzold3, Holger Bergmann4 & Josef Settele1 UFZ - Centre for Environmental Research Leipzig-Halle, Department of Community Ecology, Theodor-Lieser-Str. 4, 06120 Halle, Germany 2 UFZ - Centre for Environmental Research Leipzig-Halle, Department of Ecological Modelling, Permoserstr. 15, 04318 Leipzig, Germany 3 UFZ - Centre for Environmental Research Leipzig-Halle, Department of Economics, Permoserstr. 15, 04318 Leipzig, Germany 4 University of Göttingen, Institute of Agricultural Economics, Platz der Göttinger Sieben 5, 37073 Göttingen Contact:
[email protected] 1 While the FFH-Management is in full activity, numerous management plans - more than 250 plans alone in the Saxon Agency for Agriculture – are initiating discussions on nature and species conservation for the next years. Among these are intensive discussions on management measures concerning Maculinea species, such as the prohibition of rolling and schlepping, late mowing and strip farming. The currently practised conservative management in Saxony and in other European regions is far from being economic and sustainable. Up to now, most progressive management regimes have been implemented with the help of promotion payments. However, the feedback is scarce, and little is known about the efficiency of those measures. The immediate threat to Maculinea species requires active management that should be based on an understanding of population dynamics and of economic efficiency as well. Management decisions should be founded on a prognosis based on limited current knowledge rather than arbitrary decisions. To meet those requirements, an internet-based decision supporting tool was developed. The tool is based on the economic-ecological model of Drechsler et al. (this volume) which determines cost-effectiveness of compensation payments for the conservation of endangered Maculinea species. Applying the model of Drechsler et al. to the Landau region (Germany), the quantitative data generated by the ecological and economic modules are combined to calculate the compensation payments necessary for the implementation of a particular mowing regime, to determine which meadows apply the promoted mowing regime, and to assess the effects of this regime on the survival probability of the butterflies. The results are used to evaluate the costeffectiveness of various agro-environmental schemes. 1).246 Karin Ulbrich et al. The decision supporting tool includes chapters with background information on Maculinea ecology as well as on compensation payments and model development. nausithous is assigned to these meadows. They have a great potential to meet the requirements for management actions in nature conservation and will gain better understanding of corporate social responsibility. should be developed in cooperation with perspective users for their pragmatic needs. . and the mean area. occupied by the butterflies. 1. The incidence of M. the user has to select the mowing regime for the given study area and to determine the overall budget for compensation payments. Simulation section of the decision supporting software tool Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). is calculated over the given time period (Fig. based on case-studies and tested in training sessions. In this way. Fig. Contents are visualized by Macromedia Flash and allow individual and interactive reading. the Aarhus Convention is followed which demands effective communication of new knowledge to favour public participation in environmental decision-making. completed by numerous weblinks. teleius and M. A corpus of training tools. The simulation runs result in the calculation of promoted meadows which are mapped on the screen. The simulation section contains a concise user interface which is described in detail. Before starting the simulation. The internet-based software tools are appropriate to make research known to a broader public.
[email protected]. The main goal of the workshop was to present information describing the ecology of Maculinea butterflies (life cycle. Slovakia Contact: vavrova@sopsr. arion. Rescue programmes provide extensive information regarding ecology. 974 01 Banská Bystrica. For invertebrates there is only one rescue programme for Parnassius apollo. Currently.) and MACMAN project. Four of them (M. M.savzv. Dúbravská cesta 9. Štúrova 2. alcon. Lazovná 10.A. there are 11 rescue programmes (mainly for birds and mammals) accepted by the government. egg count.sk 1 All five European species of Maculinea occur in Slovakia: M. alcon. The three latter species are listed in Annex II of Habitats Directive containing specified criteria for evaluation of favourable conservation status of their habitats within Slovakia. arion. host plants. SNC SR prepared and distributed information brochure describing Maculinea species. in Varin. One of the official documents for the protection of species and their habitats is “Rescue programme for endangered species”. The workshop was attended by 21 participants at the department of National Park Mala Fatra Mts. dusan. They also include proposed management activities necessary for conservation of concerned species and habitat. Dušan Zitňan2 & Ján Kulfan3 State Nature Conservancy of Slovak Republic. SNC SR and colleagues from Krakow (Poland) organised a workshop “Conservation and management of Maculinea butterflies in Slovakia” at the end of April 2005. M. teleius and M. Slovakia 3 Slovak Academy of Sciences. teleius and M. M. a table of expected financial expenses to accomplish these activities and maps of species distributions within Slovakia. 247-248 Conservation of Maculinea Species in Slovakia ˇ Ľubomíra Vavrová1. nausithous. Settele. A very important issue of nature conservation is the environmental education of the public. pp. Kühn J. rebeli. Institute of Forest Ecology. foodplants and habitats). The participants had an opportunity to use some of the presented techniques in the field (e. as well as basic information (species description. monitoring and methods for estimation of population size (mark-release-recapture. The brochure contains photos of males and females of each Maculinea species occuring in Slovakia. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. These studies will result in development of effective rescue programme for all Maculinea species within next 3 years. Centre of Nature and Landscape Protection. M. The principal aim of this cooperation is to collect old and recent data of Maculinea distribution in Slovakia and to initiate more intensive research of these butterflies in the field and lab. 960 53 Zvolen. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)247 Studies on the Ecology and Conservation of Butterflies in Europe Vol.sk. 845 06 Bratislava. Close cooperation between State Nature Conservancy of Slovak Republic (SNC SR) and MACMAN countries was initiated in December 2004. their relationship with ants. habitat preferences and the threats of each species.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. sampling of Myrmica ant hosts).g. M. planned for 5 years (2005-2010). Slovakia 2 Slovak Academy of Sciences. Institute of Zoology. . E.zitnan@savba. nausithous) are protected by law. behaviour and protection within European and national legislation). etc. habitat types. Gentiana cruciata all over the studied region. arion use different Thymus and/or the Origanum host plants and are distributed throughout lowlands and mountain meadows in Slovakia. alcon is extremely local and only two stable populations using Gentiana pneumonanthe have been recently discovered. Several populations disappeared or showed considerable declines. The distribution of M. an extensive mapping of Maculinea species’ distributions in Slovakia was initiated in collaboration with specialists from Slovak Academy of Sciences. rebeli is locally distributed along with its host plant. Even more dramatic and irreversible environmental changes are caused by recent development and industrialization of many lowland areas and mountain resorts. M. It is often sympatric with M. M. M. Dušan Zitňan & Ján Kulfan Recently. teleius is local. These data showed that various isolated populations of M. but this species is still common in some regions. insecticides and fertilizers. . The decline of several Maculinea populations is apparently caused by abandoned grazing and intensification of land management including tillage of meadows or overuse of herbicides. teleius. but usually very common in lowlands and mountain valleys of entire Slovak region.248 ˇ Ľubomíra Vavrová. nausithous is restricted to several localities in western and central Slovakia. in 2003: 486 individuals. 2004: 22 individuals.A. In the years 2002 – 2005 19 habitat patches of the butterflies were found in the 5 km radius of the planned construction. M. Piotr Nowicki2. 2003: 16 individuals.©ChaetocnemaPublishers PENSOFT conducta Sofia – Moscow J. There are only a few older data available for the evaluation of the development trend of the blue butterflies. Settele. Poland Contact:
[email protected] 1 In connection with the intended construction of an artificial navigation drop on the Elbe river by Přelouč (Czech Republic). teleius. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. whether the planned construction is realized or not. in 2005: 160 individuals. in 2004: 128 individuals. pp. which is rarer in the Czech Republic (Beneš et al. Before the start of the planned construction the conditions set by the Deparment (Ministry) of Environment of the Czech Republic have to be met. . The main problems being tackled now are: 1) How will the possible destruction of a patch with a large population manifest itself in the whole system. On 11 of them both species are present. E. These conditions demand very strictly that viable populations of both butterflies species are preserved. and the data-set gathered from the beginning of the monitoring is not altogether complete. and also to manage the places occupied by the butterflies close to the threatened meadows so that populations of both species have a capability of long-term existence. Hana Veselá1 & Jiří Cibulka1 Czech University of Agriculture. research is in progress into their population parameters. on the remaining 8 only M. Thomas 2005 (Motschulsky) and its Kindred Species in the Afrotropical&Region (Eds)249 Studies on the Ecology and Conservation of Butterflies in Europe Vol. Gronostajowa 7. nausithous occurs (see Zámečník et Vrabec in prep. the total number of butterflies marked there during individual years shows a considerable fluctuation of the numbers of M. The goal of the Czech University of Agriculture research team is to design and implement an appropriate maintenance regime for the endangered meadows untill the time of their possible destruction.czu. teleius: in 2002: 476 individuals. Kamýcká street 129. 2005: 24 individuals.). Institute of Environmental Sciences. On the two patches which are the sole patches directly in danger of destruction and on several places in their near surroundings. 2002). 30-387 Kraków. CZ – 165 21 2 UJAG . nausithous shows a gradual growth of the population numbers: in 2002: less than 10 individuals. however. Prague 6 – Suchdol. an order was placed for basic research of the butterflies Maculinea teleius and Maculinea nausithous that occur on the route of the planned construction.Jagiellonian University. Czech republic. Kühn J. Jana Bouberlová1. The maintenance of the more important of the endangered patches started in the year 2004 and is targetted especially for the species M. 249-250 Conservation of Maculinea populations affected by a waterway construction in Přelouč (Czech Republic) in the view of Czech University of Agriculture research team Vladimír Vrabec1. if suitable conditions. Zámečník J. (in prep): The occurrence of Maculinea telejus and Maculinea nausithous (Lepidoptera: Lycaenidae) in broader environs of Přelouč town. Havelda Z. Fric Z. that is occurence of a breeding plant and host ants.) (2002): Butterflies of the Czech Republic: Distribution and conservation I.. SOM.. Dvořák J. V. (eds. II... Konvička M. Vrabec 2) Is it possible for the destroyed patches to be replaced by a habitat patch in the near surroundings.. Orig. & Weidenhoffer Z. Vrabec V. & Vrabec V. General situation plan of Přelouč surroundings with habitat patches of Maculinea butterflies marked.250 Vladimír Vrabec et al. are provided there. Praha. 857 pp. Pavlíčko A. 1. REFERENCES Beneš J. .. Fig. alcon) is also found. In the Eastern part (Polesie region) and in the NorthEastern part (near Białowieża NP) it seems that extensive agriculture is conductive to the existence of this butterfly. In the South of Poland there are more mountains and therefore more Southern slopes with suitable conditions for both M. rebeli in Poland. 2001. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. M.A. Institute of Environmental Sciences. A declining trend in the number of plants of this species in many drained meadows is still observed. All European Maculinea are known from Poland and all of them are protected by law. teleius is distributed mainly in the South part of Poland and in this area its food-plant Sanguisorba officinalis is also present. E. There are lots of data on the distribution of Maculinea in Poland. M. There are at least two population of M. J. M.Jagiellonian University. arion is almost exclusively noted in Southern and Eastern Poland.© PENSOFTThe distribution Publishers Sofia – Moscow J. while its host-ant M. They use different ant hosts to M. M. Zając 2001) and my own data is presented. its food-plant Gentiana pneumonanthe is restricted to isolated populations. Czechowski et al.pl Distribution of Maculinea butterflies. nausithous in of Butterflies in Europe and Conservation Vol. associated with wet meadows.uj. teleius because they feed on the same food-plant. their food-plants and potential Myrmica host ants in Poland based on the literature (Buszko 1997. scabrinodis is wildly distributed. Kühn &Poland (Eds)251 Studies and M. Thomas 2005 and ecology of Maculinea teleiuson the EcologySettele. cruciata exists. is represented only by isolated populations in the South-East of Poland and is really endangered by extinction. often with a very low density of plants. alcon and hence require different conservation management. however. 30-387 Kraków. Its main ant-host M. however. since S. In some warm and dry sites in Southern Poland where another food-plant G. and the other one is near the town Rzeszow. alcon. sabuleti and food plant Thymus pulegioides are found all over Poland. although more rare is distributed similarly as M. nausithous. M. their exact occurrence is still not well explained. rubra are present all over Poland. Zając. These two sympatric species are relatively numerous in comparison to other Maculinea.edu. M. pp. rebeli (often treated as a subspecies of M. . officinalis is a common plant on wet and extensively used meadows in Poland. scabrinodis and M. Poland Contact: woycie@eko. and it is well known that all these species are restricted to extensively used meadows or/and grasslands. Gronostajowa 7. 251-252 General overview of the status of Maculinea butterflies in Poland Michal Woyciechowski UJAG . Its potential host-ants. One is on the border with Slovakia in Pieniny Mts. sabuleti and Thymus sp. Zajac M. Research has been funded by the EC within the RTD project “MacMan” (EVK2-CT2001-00126). Distribution atlas of vascular plants in Poland. Radchenko A. 2001. Distribution atlas of butterflies in Poland (Lepidoptera: Papilionidea) 1986-1995. Laboratory of Computer Chorology. Zajac A. Oficyna Wydawnicza Turpress. Museum and Institute of Zoology PAS. Formicidae).252 Michal Woyciechowski REFERENCES Buszko J. Czechowska W.. Institute of Botany. Torun.). 2002. The ants (Hymenoptera.. Warszawa.. Krakow. 1997. . Czechowski W. Jagiellonian University and Fundation of Jagiellonian University. (eds. with the vast majority of sites located north of the Drava river.uni-lj. nausithous is restricted to the northeastern part of the country. Slovenia 2 Andrejci 25a. Antoličičeva 1. Maculinea teleius and M. teleius in the parts of the country where the ranges of both species overlap. abandonment .Studies on the EcologySettele. with a continuous distribution from the Savinja valley eastwards. Slovenia Contact: valerija. Biotechnical Faculty. nausithous have the centre of their distribution in Northeast Slovenia. Although counts of adults were made at each site. teleius and 36 % with M.A. the statistics were limited to the presence or absence of adults. Večna pot 111. The following parameters were selected: patch size. Great fluctuations were observed between the presence of both species in two consecutive seasons. Maculinea teleius is widely distributed in NE Slovenia with isolated colonies also in central and western part of Slovenia. Alltogether 124 plots were selected for the analysis of ecological parameters that could influence the presence of both Maculinea species. habitat type. sites with the food plant Sanguisorba officinalis were mapped in the central region of Slovenske gorice (240 km2).Northeast & of Butterflies in Europe and Conservation Vol. In 2003 Maculinea teleius was present on 69 % and M. 253-256 Distribution and autecology of Maculinea teleius and M.© PENSOFT Publishers Distribution and autecology Sofia – Moscow J. Slovenia 3 Centre for Cartography of Fauna and Flora. presence and time of mowing. . nausithous on 51 % of all patches. To get further insight about the distribution and requirements of both species in Slovenia two preliminary autecological studies were completed. pp. predominantly in the rural hilly countryside of Slovenske gorice and Goričko. nausithous occurs more sporadically than M. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. This could be the reason why M. In 2002 and 2003. Populations in Slovenia are on the southern border of its distribution in central Europe. Kristian Malačič 2. nausithous. SI-9221 Martjanci. Dept. SI-2204 Miklavž na Dravskem polju. nausithous is known from more than 210 localities compared with 25 known sites before 1999.si Up until end of the nineties very little was known about distribution of Maculinea species in Slovenia. abundance of food plant S. Frenk Rebeušek3 & Rudi Verovnik1 1 University of Ljubljana. Thomas 2005 of Maculinea teleius and M. nausithous (Lepidoptera: Lycaenidae) in Northeast Slovenia Valerija Zakšek 1. Currently the number of sites occupied by the species expanded to 300. The distribution of M. KühnSlovenia (Eds)253 nausithous in J. Before the year 1999 the species was known only from 35 localities. With deliberate searches for Maculinea species we were able to fill the gaps in the knowledge about their distribution.overgrowing.zaksek@bf. Currently M. of Biology. SI-1000 Ljubljana. In season 2004 there were just 54 % patches with M. E. The latter has recently obtained the status of a landscape park. officinalis. the presence of host ant species. 2000) and more conservative migration distances (1 km) for each species were used to determine a possible metapopulation structure in Northeast Slovenia. The host ants where widely distributed in the region (Myrmica scabrinodis on 89 % of patches and Myrmica rubra on 43 % of patches). teleius about 10 metapopulations occur in Northeast Slovenia Fig. Fig 3).. 2001) and 5. teleius (Stettmer et al. The difference could possibly be attributed to the rainy season of 2004 which delayed the emergence of adults. Presence of M. b). 2.1 km for M. . and habitat type. Hypothetical metapopulation structure of M. When conservative migration distances were used. nausithous (b) and time of second mowing.254 Valerija Zakšek et al. the populations in Slovenske gorice and Goričko are separated and many isolated populations of both species are present (Fig 2. nausithous (Binzenhöfer and Settele. When maximum known migration distances are used for M. 1. teleius (a) and M. teleius in northeast Slovenia.5 km for M. How many separate metapopulations exist in northeast Slovenia? The maximum dispersal distances of 2. Fig. According to nonparametric ÷2 tests the presence of both species was significantly correlated with the time of the first and second mowing (Fig 1a. The average life-span of adult individuals in the study area was 3. teleius population was 2600 (confidence interval: 1677-7707) and 750 (confidence interval: 324-2940) adults for M. nausithous in northeast Slovenia. nausithous probably occurs in two metapopulations in Northeast Slovenia and there is possibly a connection between populations in Slovenske gorice and Goričko. nausithous species in the eastern part of Goričko on a small area (3. There was a high positive correlation (0. nausithous in Northeast Slovenia 255 Fig. . officinalis. teleius males on a plot in Southeast Goričko.40 days for M. nausithous.25 days for M. 3. We have studied size. Hypothetical metapopulation structure of M. teleius and M.release . The estimated size of the M.5 ha) with 10 separate plots of larval food plant S. The mark . density and migration potential of M. Migration distances of M.92) between Fig.recapture method (MRR) was used and the surveys were made in all suitable days in the year 2003 from 8 July to 3 August. 4.Distribution and autecology of Maculinea teleius and M. and metapopulations between Slovenske gorice and Goričko are probably separated. M. nausithous. teleius and 3. Teil 1: Populationsdynamik.. teleius and M. The M. M.. teleius. nausithous were used to designate Natura 2000 sites in Slovenia (Čelik et al. Nr. 76(6): 278-287. 2005). Settele J. Our current knowledge about the distribution and habitat requirements of M. Düsseldorf: 187-193. the density of larval food plant and the density of M.. Settele J. teleius and 26 % for M. Binzenhöfer B. Habitatmanagement und Schutzmaânahmen für die Ameisenbläulinge Glaucopsyche teleius und Glaucopsyche nausithous.: Lycaenidae) im nördlicen Steigerwald. (in slovene) Geissler-Strobel S. and migrations between most distant plots (500 . Leipzig. was not present on regularly mowed patches with no overgrowing. Bergsträsser 1779 (Lepidoptera: Lycaenidae). NATURA 2000 in Slovenia: butterflies (Lepidoptera). Ljubljana 280 pp. Vergleichende autökologische Untersuchungen an Maculinea nausithous (Bergstr. Verovnik R.600 m) were observed for males of both species. No migrating females of M. 2005. Lasan. teleius was 279 m.. 2001. Čelik T... 2: 1-99. Zur Ökologie und zum Ausbreitungsverhalten von Maculinea nausithous. REFERENCES Binzenhöfer B. In: Populationsökologische Studien an Tagfaltern 2. nausithous.81) between the level of overgrowing of habitats and the density of adult individuals of M. 1779) und Maculinea teleius (Bergstr. Kleinewietfeld S. 2000. 1990. Ausbreitungsverhalten und Biotopverbund.. Gomboc S. ZRC SAZU. nausithous species however.. UFZ-Bericht. Migration levels were high with 40 % for M. Verhandlungen Westdeutscher Entomologen Tag 1989. Hartmann P. teleius adult individuals and a negative correlation (-0. 1779) (Lep.256 Valerija Zakšek et al. Natur und Landschaft. nausithous were observed. .. The maximum distance of migrating female of M. Založba ZRC.). (eds.. Further studies will need to focus on the conservation and preparation of detailed management plans for both species.. Settele J. Only with species specific conservation we will be able to sustain the currently still favourable habitat conditions in Northeast Slovenia for the long term survival of both species. Stettmer C. 6.Bibliography on Maculinea ecology and related topics (state: September 2005) 257 Section 3. Species Ecology along a European Gradient: Maculinea Butterflies as a Model – Maculinea Bibliography . 258 Elisabeth Kühn et al. This page intentionally left blank . T. a social parasite of Myrmica ant colonies.J. The MacMan literature database will be further developed after the funding stage of MacMan has finished. 248-252 (Ulmer.Centre for Ecology & Hydrology. The evolution of alternative parasitic life histories in large blue butterflies (including supplementary information. im Vordertaunus (Lepidoptera: Lycaenidae). 06120 Halle. Ent. 2004.D.© PENSOFT Publishers Bibliography Sofia – Moscow J. 8. Als. 6. Amler. T.can be made available upon request. 3. Akino. T. 1. 403-414 (2002). Sarah Gwillym1. Department of Community Ecology. & Settele. 9. Aagaard. 4. Nunner. as it is planned to continue cooperation under the name MacMan also in the future. Alonso. CEH Dorset. Smithsonian Institution Press. Chemical mimicry and host specificity in the butterfly Maculinea rebeli. 7. Nature 432. UK Contact: Elisabeth. Proceedings of the Royal Society of London Series B-Biological Sciences 266. J. et al.D. K. Adoption of parasitic Maculinea alcon caterpillars (Lepidoptera: Lycaenidae) by three Myrmica ant species. Theodor-Lieser-Str. 99-106 (2001).. (eds. T. (2000). 5. Phylogenetic relationships in brown argus butterflies (Lepidoptera : Lycaenidae : Aricia) from north-western Europe. Agosti.Centre for Environmental Research Leipzig-Halle.. E. J. Winfrith Newburgh.D. M. L. Thomas.D. 1999). 4. D. Dorset DT2 8ZD. 90 (1988). & Boomsma. Winfrith Technology Centre. K. All publications of the data base are also stored in printed versions at UFZ and – particularly if access is otherwise difficult . & Schultz. 259-283 Bibliography on Maculinea ecology and related topics (state: September 2005) Elisabeth Kühn1. Ver.). Über Maculinea nausithous BERGSTR. “The evolution of alternative parasitic life histories in Large Blue butterflies”. A.de The following bibliography lists all known publications with direct relevance to Maculinea ecology and conservation. Nash.. T. D. 27-37 (2002). pp. Amler. & Elmes.J. 2. It is an excerpt of the MacMan literature database (as of 14th of October 2005).Isolation. Kühn & J. Als. Nachr. & Boomsma. Geographical variation in host-ant specificity of the parasitic butterfly Maculinea alcon in Denmark. 9. Nash. Settele. et al. Als.W.. Majer.A. J. 386-390 (2004). Supplementary Information: Nature manuscript 2004-05-19209. Thomas 2005 on Maculinea ecology and related topics (state:and Conservation of2005) (Eds)259 Studies on the Ecology September Butterflies in Europe Vol. T.E. et al.R. Jeremy Thomas2 & Josef Settele1 1 UFZ . Germany 2 NERC . J. T.. 1419-1426 (1999). Biological Journal of the Linnean Society 75. Ecological Entomology 27. et al. D. Stuttgart. Apollo N. und Maculinea teleius BERGSTR. J.D.A.R.Kuehn@ufz. ANTS . . Knapp. Als. K. Kramer. F. J.J.Standard methods for measuring and monitoring biodiversity. Ahrheilger.. G.. Flächenbedarf und Biotopansprüche von Pflanzen und Tieren.R. Populationsbiologie in der Naturschutzpraxis .. pp. 2: Species Ecology along a European Gradient: Maculinea Butterflies as a Model. Dorchester. Animal Behaviour 62. Barnett. Gruttke. 1995. 14. & Konvicka. Die Donau-Brennen zwischen Ingolstadt und Marxheim. Martin. The challenge of conserving grassland insects at the margins of their range in Europe. M.. 22. 1779) (Lep. 8-9-2003.. J..6. Rote Liste gefährdeter Tiere Deutschlands. Species action plan: Large Blue Maculinea arion. J. 62-74 (1985). Wareham. Kepka.D. Anton. 26. Barascud. 1-98 (2000). 17. 1-85. 23. Studie im Auftrag des UFZ. Universität des Saarlandes.A.L. D. Musche. Fachrichtung 6. & Settele. J. Baguette. & Manne. JZS. L. M. 21.. A. 11.Empfehlungen zur Erfassung der Arten des Anhangs II und Charakterisierung der Lebensraumtypen das Anhangs I der . Journal of Evolutionary Biology 12. P.. 294. Bundesamt für Naturschutz (BfN). 611-625 (2000). B. 28.K. Lycaenidae) in central Europe. L. 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J. 140.. 65. 69 Nash.. 210. 174... 234. 28. 234 Settele. 234 Rebeušek. A. J. 174.. M.. L. 196 . 154. Á. 125. M. 152 Gwillym. A.. 45 Skórka. 238 Sliwinska. D. H.. H. 26.. 152. 199 Bergmann. 65. 65. 225. 152... 167. 69 Charles..... 174 Malačič. E. J. 88. J. P.. 132. 174 Boomsma.. 219 Kudłek. 140. 189. 123. 225 Maes. 174 Árnyas. 174. 82.. M. 72. 150. V. J.. 122.... 136. 182. 69. 153. M. 174. 144.. S. 32. 51. 167. 115 Musche.. E... 140. E. 16 Randle. 153.. 57 Isaia. 130. 115. E.T. 90. 9 Balletto. 214. L. A. G. D. 131.. J. 61. 153. 105. 178 Loritz. F. 61...174. O.. 82. 4 Bonelli. 121. 229 Pauler-Fürste. 78. P.. A. 189. T. 107. 238. 115.. 133 Kaule. 180. I. 172.. 178. 73.. R. 134. K. 82. U. 240 Singer. 221. 72. Th. L. 180 Ebsen. 234 Everett. 249 Bräu. A. Th. 189. 229 Pfeifer. 80. 80. B. E. 78. 124... 234 Crocetta. M.Studies on the Ecology and Conservation of Butterflies in Europe 285 Author index Amirowicz. 130. E... 196. 231 Cantarino. A. F. 134.. J. P. 219 Lomborg. 259 Sielezniew. S. 57. 182 Elmes. 122. 206.. 182 Nowicki. 69. J. 159. 69. M. 210. 115. V. 153 Peregovits. H. Z. 133. 151. J. 26. 174.. 28. 133... 74.. 130 Barbero. G. 69. 126 Hovestadt. 51.. 159. 245 Durka. 151.. A. 57. 82. 228. 229 Peregovits. 130. 57. 245. W. 152. 174.. A. R. 171. 206. 219.. S. J. R.. 215. 28.. 82 Rácz.. 3 Geißler-Strobel. 150. 229 Bereczki. 206. 174. 184 Böhme. J. 9 Mouquet. 99. F.. 124 Bíró. 174 Degen. 26.. 82. J. Ch. 215 Kis.. S. 123. 52 Rojo de la Paz. 253 Mitesser... 55. K. 128. 153 Kühn. 215 Glinka. E. K.. 126 Hula. 152. 82 Kőrösi.. 218. 69.. 130. 192. 230.. 180. 115 Hochberg. 239 Buszko. 171. S. 84. 259 Kulfan.. 221.. 4 Fiedler. N. 174. S. 152. 111. 182 Falkner. 28.. A. 245 Binzenhöfer. 231 Simcox. 126 Montani. 75 Blinov. 182. M. 125 Drechsler. 120. 192.. 167. 249 Clarke. 174. 28. 253 Richter. 55.. 218. 163. M. 107. 84.. S. 99. 126. 51. 115 Sipos. 111 Badino. 4 Lengyel.. 245 Kassai. 234.. 78.. 9 Johst. A. 214. 90. 55. 259 Hale. 105. 99.. J. 205 Anton. 136 Barabás. 136 Batáry. 199 Pępkowska... 28.. 184. 84 Cibulka. 61.. 228 Nagy. W. 80. K. E. 99. 144.. 78. 88. 151 Poethke. A. 9. 107. S. 73. 133. N. M. 229 Koschel. 125 Prondvai. 214. 88.... 178. D. J... J. 73. 69. 16 Napper. M. A. 196 Bouberlová. G. K. 136. 206.. D. 9. 189. 214. 124.. 249 Örvössy. 163. .. 115. 88. A. 154 Witek. A. 107. 196. 133. 245 Varga. 140.286 Josef Settele. C. 174 Woyciechowski. 153. 105. 150. 174.. 126. 150. 189 Topham. I. 210. 199 . E.. 154 Worgan. A.. 210. V. F. 189.. 192. 75. 84. 130. I. 82... 94. 199 Vavrová. D. 84. 240. L. 174. R. 245 Witek. 82. 159. 84. 247 Verhaagh. 234.. J. 111. M. 192. 51. 107. 239 Szitta. K... H. 99. 16. 214. 205.. 253 Veselá. 178. 45. 231 Stettmer. 111. M. 51. J. 140. 94. 174. 189. Z. 32. D.. 69. 144. 206. 28. 61. 69. 174 Tesar... 229 Vrabec. 218. R. D.. 75 Tartally. Á. 259 Tihanyi. 251 Wynhoff. 154. P. 80. 249 Wardlaw.. 238. 182 Zakšek.. 174. V. 174 Wynne. 61. 75. 253 Zeisset... 90. 105. 22 Verovnik. 69. 26.. 69... 174.. 28. Thomas Ulbrich. B. 247 Stankiewicz.. 150. 167.. 174 Wätzold.. 84 Tóth. M. 184. E. M. 140. I. A. A. 26. 163. 115 Thomas. 107. 51. 153. Ch. 182 Žitňan. 249 Vozár. Elisabeth Kühn & Jeremy A. M.. 144. 153. 167. 75. 38. 169. 146. 43. 162. 131. 37. 19. 42 arethusa 32. 38. 43 alcon 6. 136. 42. 84. 32. 94 arion ligurica 35. 37. 90. 277. 38. 144. 41 admetus 32. 41 aurelia 32. 42 budensis 32. 43. 161. 43 atalanta 39.Studies on the Ecology and Conservation of Butterflies in Europe 287 Index of latin butterfly names acaciae 37. 42 aegeria 40 aethiops 33. 25. 116. 38 Apatura 33. 92. 41. 95. 278. 40. 117. 85. 41 cardamines 37. 29. 40. 9. 132. 43. 46. 265. 33. 40. 33. 263.19. 30. 267. 248. 281. 65. 118. 270. 170. 284 alcon xerophila 41 alexis 38. 260. 232. 133. 241. 96. 41 camilla 38 captiuncula 44 Carcharodus 36. 260. 251. 189. 43 actaeon 36 adippe 30 adippe 32. 36 carthami 32. 87. 40. 273. 42. 88. 41 Celastrina 38 Chersotis 43 chloros 40. 36. 143. 146. 111. 104. 41 alfacariensis 32. 39. 94. 43 Aphantopus 40. 40. 160. 115. 30. 42 arcania 32. 63. 39. 279. 66. 233. 163. 262. 45. 267 . 41 betulae 30. 275. 33. 33. 41 antiopa 33. 76. 185. 267. 102. 40. 148. 38. 15. 41 brizae 32. 278. 251. 43 argyrognomon 38. 43. 30. 41 carniolica 40. 27. 83. 136. 103. 282. 39. 44 angelicae 40. 33. 42 c-album 39 Callophrys 32. 42 cinxia 41. 179. 17. 178. 23. 42. 26. 141. 137. 239. 38. 93. 24. 190. 240. 179. 36. 38. 236. 40 agestis 38. 101. 113. 145. 33. 67. 134 Carterocephalus 41. 43 Brintesia 39. 42 argiades 37 argiades 41 argiolus 38 argus 32 Argynnis 30. 234. 185. 42 angelicae/transalpina 40 Anthocharis 37. 274. 42 britomartis 32. 242. 168. 172. 42 Arethusana 32. 272. 159. 41. 272. 47. 39. 48. 116. 183. 49. 38. 145. 200. 274. 275. 41 aurinia 9. 42 apollo 37. 264 arion 9. 247. 39. 39. 237. 138. 97. 16. 100. 33. 89. 41 alveus 36 amandus 32. 13. 38. 27. 39. 244. 18. 261. 247. 46. 66. 41. 34. 66 bellargus 30. 231. 149. 243. 82. 41 cardui 39. 189. 247 Araschnia 39. 266. 131. 34. 39. 191. 94. 37 Boloria 26. 140. 36. 130. 232. 43 aglaja 32 alceae 36 alcetas 37 alciphron 32. 199. 133 artaxerxes issekutzi 32. 233. 149. 41. 147. 68. 276. 261. 47. 271. 101. 279 bellargus 32. 164. 73. 269. 166. 43. 190. 22. 86. 148. 178. 65. 32. 147. 280. 34. 91. 61. 268. 43. 41 Argyronome 33. 43 borelii 43 brassicae 37 Brenthis 32. 95. 259. 41 achine 32. 34. 277. 144. 99. 201. 35. 248. 74. 11. 268. 271. 34. 33. 281 arion arion 34. 38. 70. 264 Aglais 38. 119. 33. 44 Adscita 32. 235. 43 Aricia 32. 269. 40. 176. 102. 49. 16. 142. 44 athalia 32. 37. 175. 41. 275. 44 Inachis 39. Thomas galathea 30. 284 circe 39. 41. 216. 145. 198. 283. 42 Glaucopsyche 38. 39. 33. 32. 62. 152. 42 gnaphalii 43 Gonepteryx 37 Gortyna 43 Hamearis 30 hecate 32. 42 malvae 30. 42 hippothoe 33. 41 nausithous 6. 42. 98. 141. 225. 39. 33. 80. 272. 219. 32. 33. 277. 37 decoloratus 41 dia 32. 246. 39. 43 levana 39. 39. 190. 43 ino 32. 89. 48. 138. 267 Mellicta 32. 43 Cyaniris 41 Cynthia 39. 218. 40. 37. 43 Issoria 39. 42 lucifuga 43 lucina 30 lupine 40 Lycaena 9. 42 ligea 40 Limenitis 33. 66. 42. 39. 57. 32. 41. 41 dispar 9. 40. 253. 66 Colias 32. 42 Clossiana 41 Coenonympha 9. 43. 39. 212. 32. 266. 261 idas 33. 36. 40. 269. 41 ephialtes pannonica 40. 134. 122. 33. 265. 41. 43 edusa 37. 41. 37 Minois 32. 189. 37. 41 fagi 39. 248. 250. 238. 223. 148. 43. 88. 38. 38. 70. 39. 46. 270 lycaon 40. 34. 43. 44 Maculinea (not indexed. 42 glycerion 32. 36. 103. 151. 95. 42 Erynnis 36. 102. 222. 149. 41 Maniola 40. 215. 280. 134 crocea 37 Cucullia 43 Cupido 30. 37. 42 maturna 33. 181. 249. 43. 44 morpheus 36 morsei 32. 36. 44 ilia 38. 279 machaon 32. 77. 99. 221. 40. 42 ferula 39 filipendulae 40. 33. 153. 271. 42 laodice 34. 264. 267. 39. 142. 43 Euphydryas 9. 214. 254. 278. 81. 32. 273. 43 musiva 43 napi 37. 41 Melitaea 32. 228. 30. 144. 43 io 39. 265. 255. 275 Euplagia 9 euryale 40 Everes 41 Fabriciana 32. 43 Hesperia 36 Heteropterus 36 Hipparchia 30. 124. 39. 30. 49. 265 hyale 37 hyperanthus 40. 136. 41 jurtina 40. 270 dispar rutila 32. 37. 38. 39 dia 41 diamina 39 Dichagyris 43 didyma 32. 268. 41 comma 36 coridon 30 coridon 32. 33. 41 eumedon 34 euphrosyne 26. 238. 104. 66. 40 mnemosyne 33. 43. 143. 32. 42. 37. 206. 270. 58. 66. 33. 42. 260. 43 loti 40. 97. 41 Lemonia 33. 276. 33.288 Josef Settele. 210. 39. 179. 34. 219. 42 fimbriola 43 flocciferus 36. 150. 41 globulariae 32. 40. 42 megera 40. Elisabeth Kühn & Jeremy A. 41 Mesoacidalia 32 minimus 30. 39. 262. 180. 41 dorylas 38. 275 medusa 30. 41 dryas 32. 37. 226. 43 lineola 32 lonicerae 40. 259. 47. 42 erate 37 Erebia 30. 34. 43 iris 38. 217. 224. 239. 43 Leptidea 30. 42 icarus 38. 40. 41 Hemaris 33. 27. 41. 39 Lasiommata 40. 32. 40 dumi 33. 78. 42 Lopinga 32. 37. 146. 55. 38. 86. 227. 39. 247. 213. 56. 41. mentioned on most pages) maera 40. 140. 42 Hyponephele 40. 72. 17. 282. 41. 41 daphne 39 decoloratus 32. 42 Lysandra 30. 219. 230. 132. 19. 274. 79. 43 Iphiclides 32. 147. 41 fuciformis 43 . 42 latonia 39. 43 ilicis 37. 37. 279. 43. 94. 42 meleager 32. 238. 42 Melanargia 30. 256. 211. 101. 144. 44 podalirius 32. 32 sinapis/reali 32. 282. 149. 41. 43 semele 30. 32. 41 289 spini 37 statices 40 tages 36. 219. 132 . 103. 36. 148. 77. 38. 271. 37. 138. 100. 176. 104. 42. 261. 16. 141. 32. 197. 41 reali 32 rebeli 16. 36. 94. 43 trivia 32. 41 populi 38. 149. 206. 233. 253. 38. 147. 119. 138. 41 taraxaci 43 teleius 6. 66. 248. 18. 239. 269. 259. 173. 44 schiffermuelleri 38 Scolitantides 38 selene 26. 278. 260. 39. 134. 272. 74. 195. 259. 33. 61. 41. 247. 43 polychloros 38 Polygonia 39 Polyommatus 32. 76. 210. 39 semiargus 32. 36. 107. 265. 278. 169. 189. 246. 46. 106. 251. 111. 213. 136. 70. 283 telona kovacsi 33. 41 Plebeius 32. 280. 41 viciae 40 virgaureae 32. 75. 44. 168. 148. 263. 94.Studies on the Ecology and Conservation of Butterflies in Europe Neozephyrus 37 Neptis 38 niobe 39. 279. 41 sinapis 30. 190. 41 Thecla 30. 97. 264. 191. 46. 86. 33. 95.134. 44 xeranthemi 43 Zerynthia 9. 37. 282 reducta 38 rhamni 37 rivularis 38 rubi 32. 150. 38. 42. 255. 89. 243. 43 paphia 39. 153. 104. 32. 41. 39. 134 . 93. 42 pandora 39. thersites 38 Thymelicus 32. 33. 134. 97. 34. 113. 229. 131. 37. 265 tityus 33. 48. 44 serratulae 36 silvestris 32. 66 ogygia kovacsi 43 orbifer 36. 275. 41 orbifer/sertorius 36 orion 38 osiris 32. 36. 266. 38 Ochlodes 32. 184. 174. 116. 124. 214. 40. 65. 37 Spialia 36. 238. 101. 274. 99. 218. 43 Procris 40 proserpina 33. 92. 269. 17. 33. 96. 270. 102. 268. 256. 248. 190. 101. 37. 274. 33. 172. 37. 9. 36. 41 Papilio 32. 33. 44 venatus 32. 86. 217. 33. 41 w-album 37. 105. 281. 137. 40. 189. 277. 261 polyxena 9. 196. 43 Vanessa 39. 19. 91. 251. 41 tityrus 32. 37. 276. 37. 130. 41 sappho 38 Satyrium 37. 44 Zygaena 32. 44 Pontia 37. 41 phoebe 39 Photedes 44 Pieris 37. 267. 143. 41 urticae 38. 260. 38. 126. 177. 143. 37. 102. 80. 49. 192. 232. 187. 198. 254. 136. 145. 17. 147. 41. 65. 44. 211. 175. 247 Phengaris 3. 146. 265. 41 quadripunctaria 9 quercus 37 rapae 37. 62. 96. 41 Nymphalis 33. 142. 194. 73. 37. 32. 280. 36. 38 semiargus 41 sephirus 32. 262. 36. 144. 76. phlaeas 37. 33. 127. 47. 247. 38. 43. 185. 145. 44 Proserpinus 33. 43. 241. 43 palaemon 41. 167. 48. 95. 275. 279. 268. 39. 170. 42 purpuralis/osterodensis 40 Pyrgus 30. 36. 44 Pararge 40 Parnassius 33. 103. 44 Pseudophilotes 38 punica kovacsi 32 purpuralis 32. 45. 193. 36 pamphilus 40. 41 oedippus 9. 212. 88. 58. 19. 90. 122. 150. 27.