Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

Molecular analyses of the Apocrita (Insecta : Hymenoptera) suggest that the Chalcidoidea are sister to the diaprioid complex

Lyda R. Castro A B and Mark Dowton A

A Institute of Biomolecular Sciences, School of Biology, University of Wollongong, NSW 2522, Australia.

B Corresponding author. Email:

Invertebrate Systematics 20(5) 603-614
Submitted: 16 January 2006  Accepted: 21 August 2006   Published: 12 October 2006


Despite recent efforts, hypothesised phylogenetic relationships among apocritan wasps remain unresolved. In this study, molecular analyses were employed to analyse a dataset that included the 16S, the 28S and the COI genes of 87 apocritan representatives. Partial sequences of the 18S gene were also generated and added to this dataset. The topological effects of outgroup choice, method of phylogenetic analysis, and inclusion of the 18S data were systematically investigated, with particular focus on the relationship of the Chalcidoidea with other members of the Proctotrupomorpha (Platygastroidea, Proctotrupoidea, Cynipoidea). We report that ingroup topology was sensitive to the choice of outgroup, the method of phylogenetic analysis, and inclusion of 18S data. However, the Proctotrupomorpha were always monophyletic, and the Chalcidoidea were recovered, in every analysis except one, as the sister to the diaprioid complex (Diapriidae + Monomachidae + Maamingidae). The single exception, where the Chalcidoidea + Platygastroidea were recovered, utilised a more distant outgroup (Symphyta : Cephidae : Hartigia), maximum parsimony, and excluded the 18S data. Our results suggest the Chalcidoidea + (Diapriidae + Monomachidae + Maamingidae) relationship is more likely.

Additional keywords: Bayesian analysis, maximum parsimony, partitioned analysis, phylogeny, 18S rDNA.


Belshaw R. Dowton M. Quicke D. L. J. Austin A. D. 2000 Estimating ancestral geographical distributions: a Gandwanan origin for aphid parasitoids? Proceedings of the Royal Society of London. Series B. Biological Sciences 267 491 496 DOI

Bergsten J. 2005 A review of long-branch attraction. Cladistics 21 163 193 DOI

Chen Y. Xiao H. Fu J. Huang D.-W. 2004 A molecular phylogeny of eurytomid wasps inferred from DNA sequence data of 28S, 18S, 16S and COI genes. Molecular Phylogenetics and Evolution 31 300 307 DOI

Cunningham C. W. 1997 Is congruence between data partitions a reliable predictor of phylogenetic accurancy? Empirically testing and iterative procedure for choosing among phylogenetic methods. Systematic Biology 46 464 478 DOI

Dowton M. Austin A. D. 1995 Increased genetic diversity in mitochondrial genes is correlated with the evolution of parasitism in the Hymenoptera. Journal of Molecular Evolution 41 958 965 DOI

Dowton M. Austin A. D. 1997 Evidence for AT-transversion bias in wasp (Hymenoptera: Symphyta) mitochondrial genes and its implications for the origin of parasitism. Journal of Molecular Evolution 44 398 405 DOI

Dowton M. Austin A. D. 1998 Phylogenetic relationships among the microgastroid wasps (Hymenoptera: Braconidae): combined analysis of 16S and 28S rDNA genes, and morphological data. Molecular Phylogenetics and Evolution 10 354 366 DOI

Dowton M. Austin A. D. 2001 Simultaneous analysis of 16S, 28S, COI and morphology in the Hymenoptera: Apocrita – evolutionary transitions among parasitic wasps. Biological Journal of the Linnean Society 74 87 111 DOI

Dowton M. Austin A. D. 2002 Increased congruence does not necessarily indicate increased phylogenetic accuracy – the behavior of the incongruence length difference test in mixed-model analyses. Systematic Biology 51 19 31 DOI

Dowton M. Austin A. D. Dillon N. Bartowsky E. 1997 Molecular phylogeny of the apocritan wasps: the Proctotrupomorpha and Evaniomorpha. Systematic Entomology 22 245 255

Early J. W. Masner L. Naumann I. D. Austin A. D. 2001 Maamingidae, a new family of proctotrupoid wasp (Insecta: Hymenoptera) from New Zealand. Invertebrate Taxonomy 15 341 352

Gaston K. J. (1993). Spatial patterns in the description and richness of the Hymenoptera. In ‘Hymenoptera and Biodiversity’. (Eds J. LaSalle and I. D. Gauld.) pp. 277–293. (CABI: Wallingford, UK.)

Gibson G. A. P. 1999 Sister-group relationships of the Platygastroidea and Chalcidoidea (Hymenoptera) – an alternative hypothesis to Rasnitsyn (1988). Zoologica Scripta 28 125 138 DOI

Giribet G. Carranza S. Baguna J. Riutort M. Ribera C. 1996 First molecular evidence for the existence of a Tardigrada + Arthropoda clade. Molecular Biology and Evolution 13 76 84

Hall T. A. 1999 BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41 95 98

Hillis D. M. Huelsenbeck J. P. Cunningham C. W. 1994 Application and accuracy of molecular phylogenies. Science 264 671 677

Holland B. R. Penny D. Hendy M. D. 2003 Outgroup misplacement and phylogenetic inaccuracy under a molecular clock-a simulation study. Systematic Biology 52 229 238

Huelsenbeck J. P. Ronquist F. 2001 MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17 754 755

Huelsenbeck J. P., and Ronquist F. (2005). MRBAYES v3.1 (Bayesian analysis of phylogeny. Program and manua. Available at [verified September 2006].

Maniatis T., Sambrook J., and Fritsh E. E. (1989). ‘Molecular Cloning: Laboratory manual.’ (Cold Spring Harbor Laboratory Press: Cold Spring Harbor, New York, USA.)

Nylander J. A. A. Ronquist F. Huelsenbeck J. P. Nieves-Aldrey J. L. 2004 Bayesian phylogenetic analysis of combined data. Systematic Biology 53 47 67

Posada D. Crandall K. A. 1998 Modeltest: testing the model of DNA substitution. Bioinformatics (Oxford, England) 14 817 818 DOI

Prendini L. 2001 Species or supraspecific taxa as terminals in cladistic analysis? Groundplans versus exemplars revisited. Systematic Biology 50 290 300

Rasnitsyn A. P. 1980 The origin and evolution of the Hymenoptera. Trudy Paleontologicheskogo Instituta 174 1 192

Rasnitsyn A. P. 1988 An outline of evolution of the hymenopterous insects (Order Vespida). Oriental Insects 22 115 145

Ronquist F. Huelsenbeck J. P. 2003 MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 1572 1574

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 DOI

Sharkey M. J. Roy A 2002 Phylogeny of the Hymenoptera: a reanalysis of the Ronquist et al. (1999) reanalysis, emphasizing wing venation and apocritan relationships. Zoologica Scripta 31 57 66 DOI

Sunnucks P. Hales D. F. 1996 Numerous transposed sequences of mitochondrial cytochrome oxidase I–II in aphids of the genus Sitobion (Hemiptera: Aphididae). Molecular Biology and Evolution 13 510 524

Thompson J. D. Higgins D. G. Gibson T. J. 1994 CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, positions-specific gap penalties and weight matrix choice. Nucleic Acids Research 22 4673 4680

Vilhelmsen L. 2000 The ovipositor apparatus of basal Hymenoptera (Insecta): phylogenetic implications and functional morphology. Zoologica Scripta 29 319 345

Wheeler W. C. Hayashi C. Y. 1998 The phylogeny of the extant chelicerate orders. Cladistics 14 173 192

Whitfield J. B. 1992 Phylogeny of the non-aculeate Apocrita and the evolution of parasitism in the Hymenoptera. Journal of Hymenoptera Research 1 3 14

Whitfield J. B. Cameron S. A. 1998 Hierarchical analysis of variation in the mitochondrial 16S rRNA gene among hymenoptera. Molecular Biology and Evolution 15 1728 1743

Whiting M. F. Carpenter J. C. Wheeler Q. Wheeler W. C. 1997 The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology. Systematic Biology 46 1 68

Yang Z. 1994 Maximum Likelihood phylogenetic estimation from DNA sequences with variable rates over sites: approximate methods. Journal of Molecular Evolution 39 306 314 DOI

Yeates D. K. 1995 Groundplans and exemplars: paths to the tree of life. Cladistics 11 343 357 DOI

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