Studies in phylogeny and biosystematics of bees [Elektronische Ressource] : the bee genus Andrena (Andrenidae) and the tribe Anthophorini (Apidae) (Insecta: Hymenoptera: Apoidea) / vorgelegt von Andreas Dubitzky

Andreas Dubitzky

Informations

• Publié par: ludwig-maximilians-universitat_munchen
• Publié le: 01 janvier 2005
• Nombre de lectures: 32
• Langue: English
• Poids de l'ouvrage: 12 Mo

Extrait

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All nomenclaturically relevant acts in this thesis have to be regarded as unpublished according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 discussion18
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 gr