Phylogenetic implications of the mesosomal skeleton in Chalcidoidea (Hymenoptera, Apocrita) – tree searches in a jungle of homoplasy

This study arose from the first author’s PhD thesis, which was conducted in the Zoological Museum Hamburg and supported by a fellowship from the University of Hamburg. The first author is greatly indebted to his supervisor Rudolf Abraham for advice and encouragement throughought the project and for numerous discussions on pteromalid and chalcidoid phylogeny. Renate Walter (ZMH) kindly assisted with preparing many of the SEM pictures. The staff from the Entomological Department of the Zoological Museum, Copenhagen is acknowledged for extensive help and for maintaining a great working atmosphere during two visits made by the first author. These visits were financed by the German Academic Exchange Service (April–July 2004) and the SYNTHESYS project of the European Union (May–August 2005). The second author acknowledges the Danish Natural Science Council for general support. Gary Gibson (CNC, Ottawa) thoroughly revised an earlier version of this manuscript and made substantial comments that greatly improved the paper. Special thanks are also due to John Heraty and Johan Liljeblad (University of California, Riverside) for organising the ‘Chalcid workshop’ (September 2005) on morphological character systems which provided essential information on character scoring and coding in Chalcidoidea. The following individuals donated specimens for our study or helped with the identification: Kathleen Campbell (University of California, Riverside, USA), Gérard Delvare (CIRAD, Montpellier, France), John Huber (CNC, Ottawa, Canada), Fiona Impson (ARC-Plant Protection Research Institute, South Australia), Michael Ohl (Museum für Naturkunde, Humboldt University Berlin, Germany), Ralph Peters (University of Hamburg, Germany), Matthias Schöller (Biological Consultancy, Berlin, Germany), Kai Schütte (University of Hamburg, Germany), Johannes Steidle (Institute for Zoology, University of Hohenheim, Germany), and Marc Török (Hamburg, Germany).

Bendel-Janssen M. (1977). Zur Biologie, Ökologie und Ethologie der Chalcidoidea. Mitteilungen aus der Biologischen Bundesanstalt für Land- und Forstwirtschaft 176, 1–163.

Bouček Z. (1963) A taxonomic study in Spalangia Latr. (Hymenoptera, Chalcidoidea). Acta Entomologica Musei Nationalis Pragae 35, 429–512. [verified 8 August 2006.]

Quicke D. L. J., Fitton M. G., Tunstead J. R., Ingram S. N., Gaitens P. V. (1994) Ovipositor structure and relationships within the Hymenoptera, with special reference to the Ichneumonoidea. Journal of Natural History 28, 635–682.
| Crossref | GoogleScholarGoogle Scholar |

Rasnitsyn A. P. (1980) Origin and evolution of Hymenoptera. Trudy Paleontologicheskogo Instituta [In Russian.] 174, 1–192 .

Rasnitsyn A. P., Basibuyuk H. H., Quicke D. L. J. (2004) A basal chalcidoid (Insecta: Hymenoptera) from the earliest Cretaceous or latest Jurassic of Mongolia. Insect Systematics & Evolution 35, 1–13.

Rasplus J.-Y., Kerdelhué C., Le Clainche I., Mondor G. (1998) Molecular phylogeny of fig wasps. Agaonidae are not monophyletic. Comptes Rendus de l'Académie des Sciences. Series III. Sciences de la Vie 321, 517–527.
| Crossref | GoogleScholarGoogle Scholar |

Riek E. F. (1970). Hymenoptera. In ‘The Insects of Australia: A Textbook for Students and Research Workers’. pp. 867–959. (Melbourne University Press: Melbourne, Australia.)

Ronquist F., Nordlander G. (1989) Skeletal morphology of an archaic cynipoid, Ibalia rufipes. Entomologica Scandinavica. Supplementum 33, 1–60.

Ronquist F., Rasnitsyn A. P., Roy A., Eriksson K., Lindgren M. (1999) Phylogeny of the Hymenoptera: a cladistic reanalysis of Rasnitsyn’s (1988) data. Zoologica Scripta 28, 13–50.
| Crossref | GoogleScholarGoogle Scholar |

Sharkey M. J., Roy A. (2002) Phylogeny of the Hymenoptera: a reanalysis of the Ronquist et al. (1999) reanalysis, emphazising wing venation and apocritan relationships. Zoologica Scripta 31, 57–66.
| Crossref | GoogleScholarGoogle Scholar |

Shcherbakov D. E. (1981) Morphology of the pterothoracic pleura in Hymenoptera. 2. Modifications of the ground plan. Zooligicheskii Zhurnal [In Russian, translation by multilingual services division, Secretary of State, Canada.] 60, 205–213..

Snodgrass R. E. (1910). The thorax of the Hymenoptera. Proceedings of the U. S. National Museum 39, 37–92.

Snodgrass R. E. (1942) The skeleto-muscular mechanisms of the honey bee. Smithsonian Miscellaneous Collections 103, 1–120.

Swofford D. L. (1997). ‘PAUP 4, Phylogenetic Analysis Using Parsimony (Version 4.0d.56)’. (Laboratory of Molecular Systematics, Smithsonian Institute: Washington, DC, USA.)

Török M., and Abraham R. (2002). Sampling ground or truly monophyletic? Cladistic analyses applied to the phylogeny of Pteromalidae (Hymenoptera: Chalcidoidea). In ‘Parasitic Wasps: Evolution, Systematics, Biodiversity and Biological Control’. (Eds G. Melika and C. Thuróczy.) pp. 54–72. (Agroinform: Budapest, Hungary.)

Van den Assem J. (1974) Male courtship patterns and female receptivity signal of Pteromalinae (Hym., Pteromalidae), with a consideration of some evolutionary trends and a comment on the taxonomic position of Pachycrepoideus vindemiae. Netherlands Journal of Zoology 24, 253–278.

Viggiani G., Battaglia D. (1984) Male genitalia in the Aphelinidae (Hymenoptera: Chalcidoidea). Researches on Hymenoptera Chalcidoidea. LXXXV. Bolletino del Laboratorio di Entomologia Agraria 'Filippo Silvestri' di Portici 41, 149–171.

Vilhelmsen L. (2000a) Cervical and prothoracic skeleto-musculature in the basal Hymenoptera (Insecta): Comparative anatomy and phylogenetic implications. Zoologischer Anzeiger 239, 105–138.

Vilhelmsen L. (2000b) Before the wasp-waist: comparative anatomy and phylogenetic implications of the skeleto-musculature of the thoraco-abdominal boundary region in basal Hymenoptera (Insecta). Zoomorphology 119, 185–221.
| Crossref | GoogleScholarGoogle Scholar |

Vilhelmsen L., Krogmann L. (2006) Skeletal anatomy of the mesosoma of Palaeomymar anomalum (Blood & Kryger, 1922) (Hymenoptera: Mymarommatidae). Journal of Hymenoptera Research 15, 290-306.

Walker F. (1833) Monographia Chalciditum. Entomological Magazine 1, 367–384.

Weber H. (1926) Der Thorax der Hornisse. Ein Beitrag zur vergleichenden Morphologie des Insektenthorax. Zoologische Jahrbücher. Abteilung für Anatomie und Ontogenie der Tiere 47, 1–100.

Wharton R. A., Marsh P. M., and Sharkey M. J. (1997). ‘Manual of the New World Genera of the Family Braconidae (Hymenoptera). Special Publication 1.’ (The International Society of Hymenopterists: Washington, DC, USA.)

Whitfield J. B., Johnson N. F., Hamerski M. R. (1989) Identity and phylogenetic significance of the mesopostnotum on nonaculeate Hymenoptera. Annals of the Entomological Society of America 82, 663–673.

Wijesekara G. A. W. (1997) Phylogeny of Chalcididae (Insecta: Hymenoptera) and its congruence with contemporary hierarchical classification. Contributions of the American Entomological Institute 29, 1–61.

Woolley J. B. (1988) Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalcidoidea). Systematic Entomology 13, 465–501.

Appendix 1.  Taxa examined
ZMH = Zoological Museum Hamburg; ZMUC = Zoological Museum, University of Copenhagen

Appendix 2.  Character list

Appendix 2a. (continued)

Appendix 2b. (continued)

Appendix 2c. (continued)

Appendix 3.  Data matrix

Appendix 3a. (continued)