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The Bee Genus Andrena (Andrenidae) and the Tribe Anthophorini (Apidae) (Insecta: Hymenoptera: Apoidea)

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The Bee Genus Andrena (Andrenidae) and the Tribe Anthophorini (Apidae) (Insecta: Hymenoptera: Apoidea) Studies in phylogeny and biosystematics of bees: The bee genus Andrena (Andrenidae) and the tribe Anthophorini (Apidae) (Insecta: Hymenoptera: Apoidea) Dissertation zur Erlangung des Doktorgrades der Fakultät für Biologie der Ludwig-Maximilians-Universität München vorgelegt von Andreas Dubitzky Hebertshausen, 16. Dezember 2005 Erstgutachter: Prof. Dr. Klaus Schönitzer Zweitgutachter: PD Dr. Roland Melzer Tag der Abgabe: 16.12.05 Tag der mündlichen Prüfung: 23.5.06 Disclaimer All nomenclaturically relevant acts in this thesis have to be regarded as unpublished according to Article 8 of the International Code of Zoological Nomenclature, and will become available by separate publications. This dissertation is dedicated to my parents Heinz and Christine Dubitzky, who gave me the opportunity to carry out these studies and continuously supported me with their love and patience. Contents 1. Introduction............................................................................................................1 2. Material and methods............................................................................................4 2.1 Material examined ......................................................................................4 2.1.1 Morphological studies.......................................................................4 2.1.2 Molecular analysis ............................................................................5 2.2 Preparation of male genitalia and female head capsule including mouthparts...................................................................................5 2.3 Light microscopy.........................................................................................7 2.4 Scanning electron microscopy (SEM).........................................................7 2.5 Line drawings..............................................................................................8 2.6 Morphological terminology, abbreviations and measurements ...................8 2.7 Molecular techniques................................................................................10 2.7.1 Collection and preservation of voucher specimens ........................10 2.7.2 DNA-extraction, amplification, sequencing and alignment..............10 2.8 Phylogenetic analysis ...............................................................................11 2.8.1 Selection of taxa .............................................................................11 2.8.2 Character selection.........................................................................12 2.8.3 Cladistic analysis ............................................................................12 3. An evolutionary hypothesis for the genus Andrena Fabricius........................14 3.1 Introduction...............................................................................................14 3.2 Results and discussion .............................................................................18 3.2.1 Phylogenetic analysis of Andrena based on morphological data....18 3.2.2 Molecular evolution of Central European Andrena .........................62 3.2.3 New taxa of the genus Andrena .....................................................67 4. Phylogeny of the world Anthophorini, in particular the genus Habropoda..118 4.1 Introduction.............................................................................................118 4.2 Results and discussion ...........................................................................121 4.2.1 Cladistic analysis of World Anthophorini.......................................121 4.2.2 Cladistic analysis of Old World Habropoda...................................135 4.2.3 Host-parasitoid coevolution: Revision of the species of Habropoda (Anthophorini) and Tetralonioidella (Melectini) of Taiwan.............153 5. Conclusions and outlook..................................................................................192 6. Summary – Zusammenfassung........................................................................195 7. Acknowledgements...........................................................................................201 8. References .........................................................................................................202 Curriculum vitae Appendix 1–3 1. Introduction Natura non facit saltus. (Carl von Linné, 1707-1778) Bees have long exercised a strong and fascinating attraction on humankind. At least since the Mesolithic bees have played an important role in human culture, as evidenced by the 8,000 to 12,000 year old rock drawings at Bicorp (Spain), which are the oldest known representations of bees in Europe. Others found in India and South Africa might be even older (Droege, 1993). The first written reports on bees are hieroglyphic inscriptions on 2,000 to 3,000 year old Egyptian papyrus scrolls (Rudnay & Beliczay, 1987). Aristotle (384-322 B.C.) was the first to compile the available knowledge on beekeeping and the biology of bees, for example, with observations on the continuity of flower visiting behaviour and an account of the sex ratio in a colony of bees (Droege, 1993). However, all early records and studies of bees concentrate exclusively on the honey bee, Apis mellifera L. Other groups of bees are never mentioned due to the overwhelming importance of A. mellifera as honey maker and its fascinating organization of social life. With the introduction of binominal nomenclature by Linnaeus (1758) and the onset of modern taxonomy, the focus of interest increasingly broadened to include the many different groups of bees. Thus based on the work of systematists, who differentiate and categorize bees into various groups, more and more studies were carried out examining the morphology, behaviour, developmental biology or ecology of each group in great detail. These studies showed that non-domesticated bees were of high economic interest, mainly because they play an important part as pollinators for natural vegetation as well as for many crops. The leafcutter bee, Megachile rotundata, for example, is employed worldwide as a pollinator for lucerne (Medicago varia) (Dorn & Weber, 1988). Bumblebees (Bombus) are relied upon for the pollination of tomatoes in greenhouses (Michener, 2000). Nevertheless the great majority of studies carried out today on bees still concern the honey bee, which, because of its useful products such as honey and wax, is the bee of greatest economic importance. Furthermore in scientific terms of physiology and behaviour, the common honey bee can be called the best- known of all insects (Michener, 2000). Today about 17,000 described species of bees are known, and they are separated into 422 genera (Michener, 2000). Bees belong to the insect order Hymenoptera. Within the Hymenoptera they are part of the Aculeata, a group, in which the ovipositor of females has been modified to a sting apparatus. Within the aculeate Hymenoptera, crabronid wasps and bees were found in a cladistic study to be sister groups, and together with the remaining sphecoid wasps they constituted a monophyletic group (Melo, 1999). But what are the characteristic features that separate bees from sphecoid wasps? According to Michener (2000) bees are clearly identifiable by two major aspects. The first concerns the dependence of bees on pollen, which is the only protein-source of the larvae. While nearly all bees provision their larvae with 1. Introduction 2 pollen (except bees of the genus Trigona, who provide their larvae with carrion), all sphecoid wasps are strictly carnivore. The second aspect concerns two different morphological features of bees, the presence of branched or plumose hairs, which are exclusively found in bees and a hind basitarsus which is distinctly broader than the succeeding tarsal segments. From Kirby (1802) to the present, two informal groups of bees have been distinguished relating to the length of their mouthparts (Michener, 2000): the short tongued (S-T) bees, which are characterized by short and not particularly flattened labial palpi, a short galea and a short truncate or acute glossa. In contrast, the first two segments of labial palpi in the long tongued (L-T) bees are elongated and flattened sheathlike. The elongated galea of L-T bees builds a channel in which the elongated glossa can move back and forth. Seven bee families are currently recognized (Michener, 2000). Of them the Stenotritidae, Colletidae, Andrenidae, Halictidae and Melittidae are designated the S-T bees, and the remaining two families, the Megachilidae and the Apidae, comprise the L-T bees. The systematic position of the Melittidae remains dubious, since recent studies (Alexander & Michener, 1995) regard them either as the sister group to all L-T bees or as a paraphyletic group from which the L-T bees evolved. Although members of the Melittidae show more characters typical for S-T bees than L-T bees, they nonetheless exhibit some characteristic features of latter group (Michener & Greenberg, 1980, Michener, 2000). Thanks to the strong interest in bees, numerous studies of taxonomy, general biology and phylogeny have been undertaken and many groups have been thoroughly examined. However, due to the great number of existing genera and species, many taxa still require comprehensive study. This especially applies to evolutionary studies on species-rich taxa that are not as “interesting” as the honey bee and its allies or other groups of bees which attract attention by their remarkable behaviour or biology, e.g.
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