ettudom sz án é y m ológia i r n i é e h K c a s T e r T Myrmecophily of Maculinea butterflies in the Carpathian Basin (Lepidoptera: Lycaenidae) A Maculinea boglárkalepkék mirmekofíliája a Kárpát- medencében (Lepidoptera: Lycaenidae) PhD Thesis Tartally András Department of Evolutionary Zoology and Human Biology University of Debrecen Debrecen, 2008. Ezen értekezést a Debreceni Egyetem TTK Biológia Tudományok Doktori Iskola Biodiverzitás programja keretében készítettem a Debreceni Egyetem TTK doktori (PhD) fokozatának elnyerése céljából. Debrecen, 2008.01.07. Tartally András Tanúsítom, hogy Tartally András doktorjelölt 2001-2005 között a fent megnevezett Doktori Iskola Biodiverzitás programjának keretében irányításommal végezte munkáját. Az értekezésben foglalt eredményekhez a jelölt önálló alkotó tevékenységével meghatározóan hozzájárult. Az értekezés elfogadását javaslom. Debrecen, 2008.01.07. Dr. Varga Zoltán egyetemi tanár In memory of my grandparents Table of contents 1. Introduction......................................................................................... 9 1.1. Myrmecophily of Maculinea butterflies........................................................ 9 1.2. Why is it important to know the local host ant species?.............................. 9 1.3. The aim of this study.................................................................................... 10 2. Materials and Methods..................................................................... 11 2.1. Taxonomy and nomenclature...................................................................... 11 2.1.1. Glaucopsyche, Maculinea or Phengaris?.............................................. 11 2.1.2. Separation of Alcon Blues..................................................................... 11 2.1.3. Technical comments on some terms...................................................... 12 2.2. The biology of studied butterflies................................................................ 13 2.2.1. Host plants ............................................................................................ 13 2.2.2. Myrmecophily....................................................................................... 14 2.3. Field surveys and determination................................................................. 15 2.4. Host ant specificity ....................................................................................... 17 2.5. Laboratory experiments .............................................................................. 17 3. The host ant use of predatory Maculinea butterflies ..................... 19 3.1. The host ant use of M. teleius....................................................................... 19 3.1.1. Sites investigated................................................................................... 19 3.1.2. Results................................................................................................... 19 3.1.3. Discussion............................................................................................. 22 3.2. The host ant use of M. nausithous ............................................................... 27 3.2.1. Results on western Hungarian M. nausithous populations .................... 27 3.2.2. Results on isolated Transylvanian populations...................................... 32 3.3. The potential host ants of M. arion ............................................................. 35 4. The host ant use of cuckoo Maculinea butterflies .......................... 39 4.1. Field records ................................................................................................. 39 4.1.1. Sites investigated................................................................................... 39 4.1.2. Results................................................................................................... 40 4.1.3. Discussion............................................................................................. 41 4.2. Laboratory experiments .............................................................................. 49 4.2.1. The accelerated development of M. ‘rebeli’ caterpillars ....................... 49 4.2.2. Is Manica rubida a potential host ant of the Alcon Blues?.................... 53 5. Other recorded taxa connected with the Maculinea-Myrmica relationship ............................................................................................ 59 5.1. Parasitoids .................................................................................................... 59 5.2. Microdon myrmicae ...................................................................................... 61 5.3. Rickia wasmannii .......................................................................................... 62 6. Conclusions and perspectives........................................................... 65 6.1. Evolutionary and ecological aspects ........................................................... 65 6.2. Conservation biology.................................................................................... 69 Summary.................................................................................................. 71 Összefoglalás ............................................................................................ 75 Acknowledgements.................................................................................. 79 References ................................................................................................ 81 List of publications .................................................................................. 91 Fig. 1. ....................................................................................................................... 20 Fig. 2. ....................................................................................................................... 30 Fig. 3. ....................................................................................................................... 36 Fig. 4. ....................................................................................................................... 42 Table 1. .................................................................................................................... 21 Table 2. .................................................................................................................... 31 Table 3. .................................................................................................................... 37 Table 4. .................................................................................................................... 43 Table 5. .................................................................................................................... 51 Table 6. .................................................................................................................... 55 Myrmecophily of Maculinea butterflies in the Carpathian Basin 1. Introduction 1.1. Myrmecophily of Maculinea butterflies Myrmecophily (close association with ants) is a well-known phenomenon in several insect taxa (e.g. HÖLLDOBLER & WILSON 1990). The lycaenid butterflies are particularly interesting in this respect, because the caterpillars of most studied species are associated with ants. Facultative and obligate myrmecophily and the entire range from mutualism to parasitism can be observed among these butterflies (e.g. FIEDLER 1991, 2001, PIERCE & al. 2002). The genus Maculinea is known as the best-known example of obligate parasitism, where after feeding on a host plant for the first three larval instars, the fourth instar caterpillar must complete its development in a host ant nest (e.g. THOMAS & al. 1989; see Chapter 2.2.2, Appendix VI: 1-8). 1.2. Why is it important to know the local host ant species? Maculinea butterflies are often scored as globally endangered (e.g. MUNGUIRA & MARTÍN 1999, VAN SWAAY & WARREN 1999, SETTELE & al. 2002, THOMAS & SETTELE 2004, SETTELE & al. 2005, IUCN 2006; their populations are declining also in the Carpathian Basin: BÁLINT 1991, 1993). Knowledge of the host ant species has been shown to be crucial for the protection of these endangered butterflies (e.g. MUNGUIRA & MARTÍN 1999, SETTELE & al. 2005, IUCN 2006) because the availability of host ants is usually more limiting to Maculinea populations than that of the food plants (e.g. ELMES & THOMAS 1992). Furthermore, the identification of the local host ant species can unravel the evolution of this type of parasitic interaction (e.g. ELMES & al. 1998, ALS & al. 2004). In their seminal work, THOMAS & al. (1989) found that each European Maculinea butterfly depends on a single host ant species, at least in a single habitat. Subsequent studies have refined this by demonstrating that host-ant specificity varies between regions and, in addition, multiple host ant use was observed in some 9 A. Tartally: PhD Thesis regions (e.g. ELMES & al. 1994, ALS & al. 2002, STANKIEWICZ & SIELEZNIEW 2002, STEINER & al. 2003). Therefore data should be collected over the geographical range of Maculinea butterflies. 1.3. The aim of this study The aim of this study is to give a review (according to Papers I-V) on the host ant use of Maculinea populations in the Carpathian Basin. Such knowledge has been missing so far despite a recent book with a detailed chapter on lycaenid butterflies of this region (BÁLINT 1996) and despite the increasing number of local myrmecologists (see e.g. SZABÓ-PATAY 1910, SOMFAI