Johan Liljeblad, John M. Heraty, Steve Heydon, Jean-Yves Rasplus, Lars Krogmann, Peter Jansta, Hannes Baur, Simon Van Noort, Roger A. Burks, Ana Dal Molin, D. Chris Darling, Andrew Polaszek, Gary A. P. Gibson, James B. Munro, Matthew J. Yoder, John T. Huber, Jeremiah George, Michael W. Gates, Gérard Delvare, Alex Gumovsky, James B. Woolley, Stefan Schmidt, Serguei V. Triapitsyn, John D. Pinto, Astrid Cruaud, Jason L. Mottern, Elizabeth A. Murray, Department of Entomology, School of Medicine-University of California, Department of Evolution, Ecology, and Organismal Biology, Ohio State University [Columbus] (OSU), Centre de Biologie pour la Gestion des Populations (UMR CBGP), Centre de Coopération Internationale en Recherche Agronomique pour le Développement (Cirad)-Institut National de la Recherche Agronomique (INRA)-Centre international d'études supérieures en sciences agronomiques (Montpellier SupAgro)-Université de Montpellier (UM)-Institut de Recherche pour le Développement (IRD [France-Sud])-Institut national d’études supérieures agronomiques de Montpellier (Montpellier SupAgro), Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro)-Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro), Agriculture and Agri-Food [Ottawa] (AAFC), Swedish University of Agricultural Sciences (SLU), Institute for Genome Sciences, University of Maryland School of Medicine, University of Maryland System-University of Maryland System, Department of zoology, Charles University [Prague] (CU), I.I. Schmalhausen Institute of Zoology of NASU, National Academy of Sciences of Ukraine (NASU), Texas A&M University System, University of California [Davis] (UC Davis), University of California, Natural History Museum [Oslo], University of Oslo (UiO)-University of Oslo (UiO), Staatliche Naturwissenschaftliche Sammlungen Bayerns (SNSB), Department of Natural History, Royal Ontario Museum, Department of Ecology and Evolutionary Biology, University of Toronto, United States Department of Agriculture (USDA), Museum Histoire naturelle, Iziko South African Museum, Department of Zoology, University of Cape Town, Illinois Natural History Survey, University of Illinois System, and National Science Foundation (OL EF-0341149, PEET DEB-0730616, SVV-2012-265206) / Alexander von Humboldt Foundation (Germany) / SFFR (Ukraine) / BELSPO (Belgium) / Genoscope, project @Speed-Id
Cladistics Cladistics 29 (2013) 466–542 10.1111/cla.12006 A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera) John M. Heraty a, *, Roger A. Burks a,b , Astrid Cruaud a,c , Gary A. P. Gibson d , Johan Liljeblad a,e , James Munro a,f , Jean-Yves Rasplus c , Gerard Delvare g , Peter Jansˇ ta h , Alex Gumovsky i , John Huber j , James B. Woolley k , Lars Krogmann l , Steve Heydon m , Andrew Polaszek n , Stefan Schmidt o , D. Chris Darling p,q , Michael W. Gates r , Jason Mottern a , Elizabeth Murray a , Ana Dal Molin k , Serguei Triapitsyn a , Hannes Baur s , John D. Pinto a,t , Simon van Noort u,v , Jeremiah George a and Matthew Yoder w a Department of Entomology, University of California, Riverside, CA, 92521, USA; b Department of Evolution, Ecology and Organismal Biology, Ohio State University, Columbus, OH, 43210, USA; c INRA, UMR 1062 CBGP CS30016, F-34988, Montferrier-sur-Lez, France; d Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, ON, K1A 0C6, Canada; e Swedish Species Information Centre, Swedish University of Agricultural Sciences, PO Box 7007, SE-750 07, Uppsala, Sweden; f Institute for Genome Sciences, School of Medicine, University of Maryland, Baltimore, MD, 21201, USA; g Cirad, INRA, UMR 1062 CBGP CS30016, F-34988, Montferrier-sur-Lez, France; h Department of Zoology, Charles University, Vinicna 7, CZ-128 44, Praha 2, Czech Republic; i Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kiev, 30 01601, Ukraine; j Natural Resources Canada, c/o Canadian National Collection of Insects, 960 Carling Ave, Ottawa, ON, K1A 0C6, Canada; k Department of Entomology, Texas AM l Department of Entomology, State Museum of Natural History Stuttgart, Rosenstein 1, 70191, Stuttgart, Germany; m Bohart Museum of Entomology, University of California, Davis, CA, 95616, USA; n Department of Entomology, Natural History Museum, London, SW7 5BD, UK; o Staatliche Naturwissenschaftliche Sammlungen Bayerns, Zoologische Staatssammlung, M€ unchhausenstr. 21, 81247, Munich, Germany; p Department of Natural History, Royal Ontario Museum, Toronto, ON, M5S 2C6, Canada; q Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, ON, M5S 1A1, Canada; r Systematic Entomology Laboratory, USDA, ARS, PSI, c/o National Museum of Natural History, Washington, DC, 20013, USA; s Abt. Wirbellose Tiere, Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, 3005, Bern, Switzerland; t PO Box 2266, Waldport, OR, 97394, USA; u Natural History Department, Iziko South African Museum, PO Box 61, Cape Town, 8000, South Africa; v Department of Zoology, University of Cape Town, Private Bag, Rondebosch, 7701, South Africa; w Illinois Natural History Survey, University of Illinois, Champaign, IL, 61820, USA Accepted 19 September 2012 Abstract Chalcidoidea (Hymenoptera) is extremely diverse with an estimated 500 000 species. We present the first phylogenetic analysis of the superfamily based on both morphological and molecular data. A web-based, systematics workbench mx was used to score 945 character states illustrated by 648 figures for 233 morphological characters for a total of 66 645 observations for 300 taxa. The matrix covers 22 chalcidoid families recognized herein and includes 268 genera within 78 of 83 subfamilies. Morphological data were analysed alone and in combination with molecular data from ribosomal 18S (2105 bp) and 28S D2–D5 expansion regions (1812 bp). Analyses were analysed alone and in combined datasets using implied-weights parsimony and likelihood. Proposed changes in higher classification resulting from the analyses include: (i) recognition of Eriaporidae, revised status; (ii) recognition of Cynipencyrtidae, revised status; (iii) recognition of Azotidae, revised status; (iv) inclusion of Sycophaginae in Agaonidae, revised sta- tus; (v) reclassification of Aphelinidae to include Aphelininae, Calesinae, Coccophaginae, Eretmocerinae and Eriaphytinae; (vi) inclusion of Cratominae and Panstenoninae within Pteromalinae (Pteromalidae), new synonymy; (vii) inclusion of Epichrysomalli- nae in Pteromalidae, revised status. At a higher level, Chalcidoidea was monophyletic, with Mymaridae the sister group of Rotoiti- dae plus the remaining Chalcidoidea. A eulophid lineage was recovered that included Aphelinidae, Azotidae, Eulophidae, Signiphoridae, Tetracampidae and Trichogrammatidae. Eucharitidae and Perilampidae were monophyletic if Eutrichosomatinae (Pteromalidae) was included, and Eupelmidae was monophyletic if Oodera (Pteromalidae: Cleonyminae) was included. Likelihood recovered a clade of Eupelmidae + (Tanaostigmatidae + (Cynipencyrtus + Encyrtidae). Support for other lineages and their impact on the classification of Chalcidoidea is discussed. Several life-history traits are mapped onto the new phylogeny. © The Willi Hennig Society 2013. © 2012 Her Majesty the Queen in Right of Canada Cladistics © The Willi Hennig Society 2013 Reproduced with the permission of the Minister of Agriculture and Agri-Food Canada.