Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Molecular phylogenetics and the evolution of wing reduction in the Baeini (Hymenoptera : Scelionidae): parasitoids of spider eggs

D. Carey A , N. P. Murphy A and A. D. Austin A B
+ Author Affiliations
- Author Affiliations

A Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences, The University of Adelaide, SA 5005, Australia.

B Corresponding author. Email: andy.austin@adelaide.edu.au

Invertebrate Systematics 20(4) 489-501 https://doi.org/10.1071/IS06011
Submitted: 11 April 2006  Accepted: 18 July 2006   Published: 29 August 2006

Abstract

Members of the tribe Baeini are unique among the Scelonidae as endoparasitoids of the eggs of spiders, whereas all other scelionids parasitise insect eggs. The monophyly of the Baeini has previously been assumed based on this host association and several morphological characteristics, none of which are unequivocal. Here we present a phylogeny for the group using sequence data from two genes and 32 ingroup taxa, mostly from Australia. Combined analysis of the nuclear 28S rRNA gene and the mitochondrial CO1 gene showed that the tribe is not monophyletic, because two genera (Neobaeus Austin and Mirobaeoides Dodd) are resolved in a clade separate from the rest of the Baeini. The two most speciose genera, Idris Foerster and Ceratobaeus Ashmead, are not resolved as monophyletic, indicating that a metasomal horn, which acts as a recess for the internally retracted ovipositor, has evolved multiple times within the tribe. Mapping of wing reduction states that occur in the Baeini revealed multiple wing reduction events, possibly associated with adaptations for searching for host egg masses in crytobiotic habitats, and/or penetrating the silk egg sacs of spiders. The molecular phylogeny generated in this study differs significantly from previous morphological phylogenies, and has resolved relationships that have otherwise been intractable in morphological studies because of homoplasy associated with reduction in body size.

Additional keywords: Araneae, Baeus, Ceratobaeus, homoplasy, Idris, ovipositor.


Acknowledgments

We wish to thank Lubomir Masner, Norm Johnson, Mark Dowton and John Jennings for help with specimens and discussions about scelionid systematics, and the Australian Research Council and The University of Adelaide for funding the project.


References


Ashmead W. H. (1893) A monograph of the North American Proctotrypidae. Bulletin of the United States National Museum 45, 1–472. open url image1

Austin A. D. (1981) The types of Australian species in the tribes Idrini, Baeini and Embidobiini (Hymenoptera: Scelionidae: Scelioninae). General and Applied Entomology 13, 81–92. open url image1

Austin A. D. (1983) Morphology and mechanics of the ovipositor system of Ceratobaeus Ashmead (Hymenoptera: Scelionidae) and related genera. International Journal of Insect Morphology & Embryology 12, 139–155.
Crossref | GoogleScholarGoogle Scholar | open url image1

Austin A. D. (1985) The function of spider egg sacs in relation to parasitoids and predators, with special reference to the Australian fauna. Journal of Natural History 19, 359–376. open url image1

Austin A. D. (1986) A taxonomic revision of the genus Mirobaeoides Dodd (Hymenoptera, Scelionidae). Australian Journal of Zoology 34, 315–337.
Crossref | GoogleScholarGoogle Scholar | open url image1

Austin A. D. (1988a). Guarding behaviour, eggmass shape and eggsac in Clubiona robusta L. Koch (Araneae: Clubionidae). In ‘Australian Arachnology’. (Eds A. D. Austin and N. W. Heather.) pp. 87–95. (The Australian Entomological Society: Brisbane, Australia.)

Austin A. D. (1988b) A new genus of baeine wasp (Hymenoptera, Scelionidae) from New-Zealand associated with moss. New Zealand Journal of Zoology 15, 173–183. open url image1

Austin A. D., Field S. A. (1997) The ovipositor system of scelionid and platygastrid wasps (Hymenoptera: Platygastoidea): comparative morphology and phylogenetic implications. Invertebrate Taxonomy 11, 1–87.
Crossref | GoogleScholarGoogle Scholar | open url image1

Austin A. D., Iqbal M. (2005) A new species of Cyphacolus Priesner (Hymenoptera: Scelionidae) from Australia with a discussion of generic relationships within the Baeini. Acta Societatis Zoologicae Bohemicae 69, 17–23. open url image1

Austin A. D., Johnson N. F., Dowton M. (2005) Systematics, evolution, and biology of scelionid and platygastrid wasps. Annual Review of Entomology 50, 553–582.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Brandley M. C., Schmitz A., Reeder T. W. (2005) Partitioned bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards. Systematic Biology 54, 373–390.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Dodd A. P. (1939) Hymenopterous parasites of Embioptera. Proceedings of the Linnean Society of New South Wales 64, 338–344. open url image1

Galloway I. D., Austin A. D. (1984) Revision of the Scelioninae (Hymenoptera, Scelionidae) of Australia. Australian Journal of Zoology 99, 1–138. open url image1

Gillespie J. J., Munro J. B., Heraty J. M., Yoder M. J., Owen A. K., Carmichael A. E. (2005) A secondary structural model of the 28S rRNA expansion segments D2 and D3 for chalcidoid wasps (Hymenoptera: Chalcidoidea). Molecular Biology and Evolution 22, 1593–1608.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Hickman V. V. (1967) New Scelionidae (Hymenoptera) which lay their eggs in those of spiders. Journal of the Entomological Society of Australia (New South Wales) 7, 8–33. open url image1

Huggert L. (1979) Revision of the west Palaearctic species of the genus Idris Foerster, s.l. (Hymenoptera: Proctotrupoidea: Scelionidae). Entomologica Scandinavica Supplement 12, 1–60. open url image1

Huelsenbeck J. P., Ronquist F. (2001) MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics (Oxford, England) 17, 754–755.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Iqbal M., Austin A. D. (1997) Species richness and endemism of baeine wasps (Hymenoptera: Scelionidae) in Australia. Memoirs of Museum Victoria 56, 455–459. open url image1

Iqbal M., and Austin A. D. (1999). Systematics, diversity and host relationships of baeine wasps (Hymenoptera: Scelionidae): parasitoids of spider eggs. In ‘The Other 99% — The Conservation and Biodiversity of Invertebrates’. (Eds W. Ponder and D. Lunney.) pp. 228–231. (Transactions of the Royal Zoological Society of NSW: Mosman, Australia.)

Iqbal M., Austin A. D. (2000a) Systematics of Ceratobaeus Ashmead (Hymenoptera: Scelionidae) from Australasia. Records of the South Australian Museum Monographic Series 6, 1–164. open url image1

Iqbal M., and Austin A. D. (2000b). A preliminary phylogeny for the Baeini (Hymenoptera: Scelionidae): endoparasitoids of spider eggs. In ‘Hymenoptera: Evolution, Biodiversity and Biological Control’. (Eds A. D. Austin and M. Dowton.) pp. 178–187. (CSIRO Publishing: Melbourne, Australia.)

Kieffer J. J. (1926) Hymenoptera, Proctotrupoidea, Scelionidae. Das Tierreich 48, 1–885. open url image1

Kozlov M. A. (1970) Supergeneric groupings of the Proctotrupoidea (Hymenoptera). Entomological Review, Washington 49, 115–127. open url image1

Lockhart P. J., Steel M. A., Hendy M. D., Penny D. (1994) Recovering evolutionary trees under a more realistic model of sequence evolution. Molecular Biology and Evolution 11, 605–612. open url image1

Maddison W. P., and Maddison D. R. (2003). Mesquite: a modular system for evolutionary analysis. Available online at: http://mesquiteproject.org [verified July 2006].

Masner L. (1968) A new scelionid wasp from intertidal zone of South Africa (Hymenoptera: Scelionidae). Annals of the Natal Museum 20, 195–198. open url image1

Masner L. (1972) The classification and interrelationships of Thoronini (Hymenoptera: Proctotrupoidea, Scelionidae). Canadian Entomologist 104, 833–849. open url image1

Masner L. (1976) Revisionary notes and keys to world genera of Scelionidae (Hymenoptera: Proctotrupoidea). Memoirs of the Entomological Society of Canada 97, 1–87. open url image1

Masner L. (1980) Key to genera of Scelionidae of the Holarctic region, with descriptions of new genera and species (Hymenoptera: Proctotrupoidea). Memoirs of the Entomological Society of Canada 113, 1–54. open url image1

Masner L., Huggert L. (1989) World review and keys to genera of the subfamily Inostemmatine with reassignment of the taxa to the Platygastrinae and Sceliotrachelinae (Hymenoptera: Platygastridae). Memoirs of the Entomological Society of Canada 147, 1–214. open url image1

Mathews D. H., Sabina J., Zuker M., Turner D. H. (1999) Expanded sequence dependence of thermodynamic parameters improves prediction of RNA secondary structure. Journal of Molecular Biology 288, 911–940.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Murphy N.P., Carey D., Castro L., Dowton M., Austin A.D. (in press) Phylogeny of the platygastroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and CO1 genes: implications for classification and the evolution of host relationships. Biological Journal of the Linnean Society XX, 000–000. open url image1

Posada D., Crandall K. A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics (Oxford, England) 14, 817–818.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Reeder T. W. (2003) A phylogeny of the Australian Sphenomorphus group (Scincidae: Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships. Molecular Phylogenetics and Evolution 27, 384–397.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Schwarz M. P., Tierney S. M., Cooper S. J. B., Bull N. J. (2004) Molecular phylogenetics of the allodapine bee genus Braunsapis: A–T bias and heterogeneous substitution parameters. Molecular Phylogenetics and Evolution 32, 110–122.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Shimodaira H., Hasegawa M. (1999) Multiple comparisons of log-likelihoods with applications to phylogenetic inference. Molecular Biology and Evolution 16, 1114–1116. open url image1

Simon C., Frati F., Beckenbach A., Crespi B., Liu H., Flook P. (1994) Evolution, weighting and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87, 651–701. open url image1

Swofford D. L. (2003). ‘PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4.’ (Sinauer Associates: Sunderland, MA, USA.)

Wiens J. J. (1998) Combining data with different phylogenetic histories. Systematic Biology 47, 568–581.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Zera A. J., Denno R. F. (1997) Physiology and ecology of dispersal polymorphism in insects. Annual Review of Entomology 42, 207–230.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Zuker M. (2003) Mfold web server for nucleic acid folding and hybridization prediction. Nucleic Acids Research 31, 3406–3415.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1