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Published: 2007-12-21
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Phylogeny and Classification of Hymenoptera*

S-225 Ag. Sci. Building-N, Department of Entomology, University of Kentucky, Lexington, KY 40546-0091, USA
Hymenoptera Diaprioidea taxonomy cladistics life history Insecta


An overview of recent advances in our understanding of the higher phylogeny and classification is presented. Molecular and morphological cladistic and pre-cladistic studies are summarized. A superfamily-level classification of the Hymenoptera is offered to reflect recent advances in our understanding of the phylogenetic relationships of the Hymenoptera. It differs from most recent classifications in the recognition of the Diaprioidea, to include Diapriidae, Monomachidae, and Maamingidae.


  1. Alexander, B.A. (1992) An exploratory analysis of cladistic relationships within the superfamily Apoidea, with special reference to sphecid wasps (Hymenoptera). Journal of Hymenoptera Research, 1, 25–61.

    Ashmead, W.H. (1896) The phylogeny of the Hymenoptera. Proceedings of the Entomological Society of Washington, 3, 323–336.

    Askew, R.R. & Shaw, M.R. (1986) Parasitoid communities: their size, structure and development. In Waage, J. & D. Greathead. Insect Parasitoids. Academic Press. London, pp 225–264.

    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.

    Austin, A.D., Field, S.A. (1997) The ovipositor system of scelionid and platygastrid wasps (Hymenoptera: Platygastroidea): comparative morphology and phylogenetic implications. Invertebrate Taxonomy, 11, 1–87.

    Basibuyuk, H.H. & Quicke, D.L.J. (1995). Morphology of the antenna cleaner in the Hymenoptera with particular reference to non-aculeate families (Insecta). Zoologica Scripta, 24, 157–177.

    Basibuyuk, H.H. & Quicke, D.L.J. (1997) Hamuli in the Hymenoptera. Journal of Natural History 31, 1563–1585.

    Basibuyuk, H.H. & Quicke, D.L.J. (1999a) Grooming behaviours in the Hymenoptera (Insecta): potential phylogenetic significance. Zoological Journal of the Linnean Society, 125, 349–382.

    Basibuyuk, H.H. & Quicke D.L.J. (1999b) Gross morphology of multiporous plate sensilla in the Hymenoptera (Insecta). Zoologica Scripta, 28, 51–67.

    Basibuyuk, H.H., Quicke, D.L.J., Rasnitsyn, A.P. & Fitton, M.G. (2000) Morphology and sensilla of the orbicula, a sclerite between the tarsal claws, in the Hymenoptera. Annals of the Entomological Society of America, 93, 625–636.

    Belshaw, H.H., Herniou, R.E., Gimeno C., Fitton, M.G. &. Quicke, D.L.J. (1998) Molecular phylogeny of the Ichneumonoidea (Hymenoptera) based on D2 expansion region of 28S rDNA.

    Beutel, R.G., Vilhelmsen, L. (2007) Head anatomy of Xyelidae (Hexapoda: Hymenoptera) and phylogenetic implications. Organisms, Diversity and Evolution, 7, 207–230.

    Brothers, D.J. (1975) Phylogeny and classification of the aculeate Hymenoptera, with special reference to Mutillidae. University of Kansas Science Bulletin, 50, 483–648.

    Brothers, D. J. (1999) Phylogeny and evolution of wasps, ants and bees (Hymenoptera, Chrysidoidea, Vespoidea and Apoidea). Zoologica Scripta, 28, 233–249.

    Brothers, D.J. & Carpenter, J.M. (1993) Phylogeny of Aculeata: Chrysidoidea and Vespoidea. Journal of Hymenopteran Research, 2, 227–302.

    Burke, H.E. (1917) Oryssus is parasitic. Proceedings of the entomological Society of Washington, 14, 87–89.

    Campbell, B., Heraty, J.M., Rasplus, J.Y., Chan, K., Steffen-Campbell, J., & Babcock, C. (2000) Molecular Systematics of the Chalcidoidea Using 28S-D2 rDNA. In: Austin, A. & Dowton, M. (Eds.), The Hymenoptera: Evolution, Biodiversity and Biological Control. CSIRO Publishing, Melbourne. pp. 57–71.

    Carmean, D. & Kimsey L. (1998) Phylogenetic revision of the parasitoid wasp family Trigonalidae. Systematic Entomology 23, 35–76.

    Carpenter, J.M. (1986) Cladistics of the Chrysidoidea (Hymenoptera). Journal of the New York Entomological Society, 94, 303–330.

    Carpenter, J.M. (1999) What do we know about chrysidoid relationships? Zoologica Scripta, 28, 215–231.

    Carpenter, J.M. & Wheeler, W.C. (1999) Towards simultaneous analysis of morphological and molecular data in Hymenoptera. Zoologica Scripta, 28, 251– 260.

    Castro, L.R. & Dowton, M. (2005) The position of the Hymenoptera within the Holometabloa as inferred from the mitochondrial genome of Perga Condei (Hymenoptera: Symphyta: Pergidae). Molecular Phylogenetics and Evolution 34, 469–479.

    Castro, L.R. & Dowton, M. (2006) Molecular analysis of the Apocrita (Insecta: Hymenoptera) suggest that the Chalcidoidea are sister to the diaprioid complex. Invertebrate Systematics, 20, 603–614.

    Ceresa-Gastaldo, L. & Chiappini, E. (1994) Observations on the cocoon of Oligosita krygeri Girault, (Hymenoptera: Trichogrammatidae) oophagous parasitoid of Cicadella viridis (L.) (Homoptera: Cicadellidae). Norwegian Journal of Agricultural Sciences. Supplement, 16, 131–140.

    Clausen, C.P. (1940) Entomophagous Insects. McGraw-Hill, New York, 688pp.

    Clausen, C.P. (1978) Introduced Parasites & Predators of Arthropod Pests and Weeds: A World Review. United States Department of Agriculture, Agriculture Handbook No. 480, Washington D.C.

    Colazza, S. & Bin, F. (1992) Introduction of the oophage Edovum puttleri Griss. (Hymenoptera: Eulophidae) in Italy for the biological control of Colorado potato beetle. Redia, 75, 203–225.

    Danforth, B.N., Sipes, S., Fang, J. & Brady, S.G. (2006) The history of early bee diversification based on five genes plus morphology. Proceedings of the National Academy of Science, 103, 15118–15123.

    DeBach, P. & Rosen, D. (1991) Biological Control by Natural Enemies, 2nd Edition. Cambridge University Press, Cambridge. 456 pp.

    Deans, A.R., Gillespie, J.J. & Yoder, M.J. (2006) An evaluation of ensign wasp classification (Hymenoptera: Evaniidae) based on molecular data and insights from ribosomal RNA secondary structure. Systematic Entomology, 31, 517–528.

    Dowton, M. (1999) Relationships among the cyclostome braconid (Hymenoptera: Braconidae) subfamilies inferred from a mitochondrial tRNA gene rearrangement. Molecular Phylogenetics and Evolution, 11, 283–287.

    Dowton, M. & Austin, A.D. (1994) Molecular phylogeny of the insect order Hymenoptera: Apocritan relationships. Proceedings of the National Academy of Sciences of the United States of America, 91, 9911–9915.

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

    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.

    Dowton, M., Austin A.D. & Antolin, M. F. (1998) Evolutionary relationships among the Braconidae (Hymenoptera: Ichneumonoidea) inferred from partial 16s rDNA gene sequences. Insect Molecular Biology, 7, 129–150.

    Dowton, M., Austin, A.D., Dillon, N., & Bartowsky, E. (1997) Molecular phylogeny of the apocritan wasps: the Proctotrupomorpha and Evaniomorpha. SystematicEntomology, 22, 245–255.

    Dowton, M., Belshaw, R., Austin, A.D. & Quicke, D.L.J. (2002) Simultaneous molecular and morphological analysis of braconid relationships (Insecta: Hymenoptera: Braconidae) indicates independent mt-tRNA gene inversions within a single wasp family. Journal of Molecular Evolution, 54, 210–226.

    Eberhard, W.G. (2000) The natural history and behavior of Hymenoepimecis argyraphaga (Hymenoptera: Ichneumonidae) a parasitoid of Plesiometa argyra (Araneae: Tetragnathidae). Journal of Hymenoptera Research, 9, 220–240.

    Evans, H.E. (1987) Order Hymenoptera. In: Stehr, F.W. (Ed.), Immature Insects. Vol. 1. Kendall/Hunt, Iowa, pp. 597–710.

    Fernández, F. & Sharkey, M.J. (2006) Introducción a los Hymenoptera de la Región Neotropical, Serie Entomología Colombiana, Sociedad Colombiana de Entomología, Bogotá D.C., Colombia, 896 pp.

    Flanders, S.E. (1938) Cocoon formation in endoparasitic chalcidoids. Annals of the Entomological Society of America, 31, 167–180.

    Gaston, K.J. (1991) The magnitude of global insect species richness. Conservation Biology 5, 283–296.

    Gibson, G.A.P. (1985) Some pro- and mesothoracic characters important for phylogenetic analysis of Hymenoptera, with a review of terms used for structures. Canadian Entomologist, 117, 1395–443.

    Gibson, G.A.P. (1986) Evidence for monophyly and relationships of Chalcidoidea, Mymaridae, and Mymarommatidae (Hymenoptera: Terebrantes). Canadian Entomologist, 118, 205–240.

    Gibson, G.A.P. (1993) Groundplan structure and homology of the pleuron in Hymenoptera based on a comparison of the skeletomusculature of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Memoirs of the Entomological Society of Canada, 165, 165–187.

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

    Gibson, G.A.P., Heraty, J.M. & Woolley, J.B. (1999) Phylogenetics and classification of Chalcidoidea and Mymarommatoidea - a review of current concepts (Hymenoptera: Apocrita). Zoologica Scripta, 28, 87–124.

    Goulet, H. & Huber, J.T. (1993) Hymenoptera of the World: An identification guide to families. Agriculture Canada 1894E, 688 pp.

    Gourlay, E.S. (1951) Notes on the insects associated with Pinus radiata in New Zealand. Bulletin of Entomological Research, 42, 21–22.

    Grissell, E. E. (1999) Hymenopteran diversity: Some alien notions. American Entomologist, 45, 235–244.

    Hanson, P.E. & Gauld, I.D. (2006). Hymenoptera de la regin neotropical. Memoirs of the American Entomological Institute, 77, 1–994.

    Hawkins, B.A. (1993) Refuges, host population dynamics and the genesis of parasitoid diversity., In: LaSalle, J. & Gauld, I.D. (Eds.) Hymenoptera and Biodiversity. CAB International, Wallingford, UK, pp 235–256.

    Hedrick, R.W. & Parker, J. D. (1997) Evolutionary genetics and genetic variation of haplodiploids and x-linked genes. Annual Review of Ecology and Systematics, 28, 55–83.

    Hennig. W. (1969). Die Stammesgeschichte der Insekten. Kramer, Frankfurt am Main, Germany, 436 pp.

    Hennig. W. (1981). Insect Phylogeny. John Wiley, New York, 514 pp.

    Heraty, J.M., Woolley, J.B.& Darling, D.C. (1994) Phylogenetic implications of the mesofurca and mesopostnotum in Hymenoptera. Journal of Hymenopteran Research, 3, 241–77.

    Hölldobler, B. & Wilson, E.O. The Ants. Harvard University Press, Cambridge, 732 pp.

    Jennings, J.T. & Austin, A.D. (2000) Higher-level phylogeny of the Aulacidae and Gasteruptiidae (Hymenoptera: Evanioidea). In: Austin, A.D. & Dowton M. (Eds) The Hymenoptera: Evolution, Biodiversity and Biological Control. CSIRO Publishing, Melbourne, pp. 154–164.

    Jennings, J.T. & Austin, A.D. (2004) Biology and host relationships of aulacid and gasteruptiid wasps (Hymenoptera: Evanioidea): a review. In: Rajmohana, K., Sudheer, K., Girish Kumar, P., & Santhosh, S. (Eds) Perspectives on Biosystematics and Biodiversity. University of Calicut, Kerala, India, pp. 187–215

    Johnson, N.F. (1988) Midcoxal articulations and the phylogeny of the order Hymenoptera. Annals of the Entomological Society of America, 81, 870–81.

    Köristensen, N.P. (1991) Phylogeny of the extant hexapods. In: Naumann, I. (Ed.), The Insects of Australia (2nd edition). Melbourne University Press, pp. 125–140.

    Königsmann, E. (1976). Das phylogenetische System der Hymenoptera. Teil 1: Einfuhrung, Grundplanmerkmale, Schwestergruppe und Fossilfunde. Deutsche Entomologische Zeitschrift, 23, 253–79.

    Königsmann, E. (1977) Das phylogenetische System der Hymenoptera. Teil 2: Symphyta. Deutsche Entomologische Zeitschrift, 24, 1–40.

    Königsmann, E. (1978a) Das phylogenetische System der Hymenoptera. Teil 3: Terebrantes (Unterordnung Apocrita). Deutsche Entomologische Zeitschrift, 25, 1–55.

    Königsmann, E. (1978b) Das phylogenetische System der Hymenoptera. Teil 4: Aculeata (Unterordnung Apocrita). Deutsche Entomologische Zeitschrift 25, 365–435.

    Kukalova-Peck, J. & Lawrence, J.F. (2004) Relationships among coleopteran suborders and major endoneopteran lineages: Evidence from hind wing characters. European Journal of Entomology, 101, 95–144.

    LaSalle, J. (2003) Parasitic Hymenoptera, biological control and biodiversity. In: LaSalle J. & Gauld, I. (Eds), Hymenoptera and Biodiversity. CAB International Wallingford, UK, pp. 197–215.

    Laurenne, N.M., Broad, G.R. & Quicke, D.L.J. (2006) Direct optimization and multiple alignment of 28S D2–D3 rDNA sequences: problems with indels on the way to a molecular phylogeny of the cryptine ichneumonid wasps (Insect: Hymenoptera). Cladistics, 22, 442–473.

    Liljeblad, J. & Ronquist, F. (1998) A phylogenetic analysis of higher-level gall wasp relationships. Systematic Entomo-logy, 23, 229–252.

    Liu, Z., Engel, M.S. & Grimaldi, D.A. (2007) Phylogeny and geological history of the cynipoid wasps (Hymenoptera, Cynipoidea). American Museum Novitates, 3583, 1–48.

    Melo, G.A.R. (1999) Phylogenetic relationships and classification of Apoidea (Hymenoptera), with emphasis on the crabronid wasps. Natural History Museum, University of Kansas Scientific Papers, 14, 1–55.

    Michener, C.D. (2000) The bees of the world. John Hopkins University Press, Baltimore and London, 913 pp.

    Murphy, N.P. Carey, D., Castro, L.R., Dowton, M., & Austin, A.D. (2007) Phylogeny of the platygastroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and cytochrome oxidase genes: implications for the evolution of the ovipositor system and host relationships. Biological Journal of the Linnean Society, 91, 653–669.

    Naumann, I.D. & Masner, L. (1985). Parasitic wasps of the proctotrupid complex: a new family from Australia and a key to world families (Hymenoptera: Proctotrupoidea sensu lato). Australian Journal of Zoology, 33, 761–783.

    Nieves-Aldrey, J.L., & Fontal-Cazalla F.M. (1999) Filogenia y evolucin del orden Hymenoptera, In: Meli, A., de Haro, J.J., Mendez, M. & Ribera, I. (eds.) Evolucin y Filogenia de Arthropoda. Monogrphico SEA No. 26. Zaragoza, pp 459–474.

    Nieves-Aldrey, J.L., Fontal-Cazalla, F.M.& Fernández, F. (2006) Filogenia y evolucin en Hymenoptera. In: Fernández, F. & Sharkey, M.J. (Eds). Introducción a los Hymenoptera de la Región Neotropical, Serie Entomología Colombiana, Sociedad Colombiana de Entomología, Bogotá D.C., Colombia.pp. 37–55.

    Nuttall, M.J. (1980). Insect parasites of Sirex. Forest and timber insects in New Zealand, 47 (unpaginated).

    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.

    Oeser, R. (1961) Vergleichend-morphologissche Untersuchungen uber den Ovipositor der Hymenopteren. Mitteilungen aus dem Zoologischen Museum in Berlin, 37, 1–119.

    Ohl, M. & Bleidorn, C. (2006) The phylogenetic position of the enigmatic wasp family Heterogynaidae based on molecular data, with description of a new, nocturnal species (Hymenoptera: Apoidea). Systematic Entomology, 31, 321–337.

    Peterson, A. (1948) Larvae of Insects, an Introduction to Nearctic Species. Part I. Lepidoptera and Plant Infesting Hymenoptera. Privately published. Columbus, Ohio, 315 pp.

    Pitz, K.M., Dowling, A.PG, Sharanowski, B.J., Boring, C.A, Seltmann, K.C. & Sharkey, M.J. (2006) Phylogenetic relationships among the Braconidae (Hymenoptera: Ichneumonoidea) as proposed by Shi et al.: a reassessment. Molecular Phylogenetics and Evolution, 43, 338–343.

    Pilgrim, E.M., von Dohlen, C.D. & Pitts, J.P. (in prep.) Oh Brother, where art thou Vespoidea? Molecular phylogenetics indicates the superfamily is paraphyletic and contains Apoidea.

    Powell, J.A. & Turner, W.J. (1975) Observations on ovipositional behavior and host selection in Orussus occidentalis (Hymenoptera: Siricoidea). Journal of the Kansas Entomological Society, 48, 299–307.

    Prentice, M.A. (1998) The comparative morphology and phylogeny of the apoid wasps (Hymenoptera: Apoidea). Ph.D. Dissertation, University of California, Berkeley, 1439 pp.

    Quicke, D.L.J. & Belshaw, R. (1999) Incongruence between Morphological data sets: an example from the evolution of endoparasitism among parasitic wasps (Hymenoptera: Braconidae). Systematic Biology, 48, 436–454.

    Quicke, D.L.J., Ingram, S.N. & Fitton, M.G. (1992a) Phylogenetic implications of the structure and distribution of ovipositor valvilli in the Hymenoptera. Journal of Natural History, 26, 587–608.

    Quicke, D.L.J., Ingram, S.N., Baillie, H.S. & Gaitens, P.V. (1992b) Sperm structure and ultrastructure in the Hymenoptera (Insecta). Zoologica Scripta, 21, 381 – 402.

    Quicke, D.L.J., Fitton, M.G., Tunstead, J., 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.

    Quicke, D.L.J., Basibuyuk, H.H., Fitton, M.G. & Rasnitsyn, A.P. (1999a) Morphological, palaeontological and molecular aspects of ichneumonoid phylogeny (Hymenoptera, Insecta). Zoologica Scripta, 28, 175–202.

    Quicke, D.L.J., LeRalec, A., & Vilhelmsen, L. (1999b) Ovipositor structure and function in the parasitic Hymenoptera with an exploration of new hypotheses. Rendiconti, 47, 197–239.

    Quicke, D.L.J., Lopez-Vaamonde, C. & Belshaw, R. (1999c) The basal Ichneumonidae (Insecta: Hymenoptera): 28S rDNA considerations of the Brachycyrtinae, Labeninae, Paxylommatinae and Xoridinae. Zoologica Scripta, 28, 203–210.

    Rahman, H., Fitton, M.G., & Quicke, D.L.J. (1998) Ovipositor internal microsculpture in the Braconidae (Insecta, Hymenoptera). Zoologica Scripta, 27, 319–331.

    Rasnitsyn, A.P. (1968) Evolution of the function of the ovipositor in relation to the origin of parasitism in Hymenoptera. Entomological Review, 47, 35–40.

    Rasnitsyn, A.P. (1969) Origin and evolution of the lower Hymenoptera. Trudy Paleontologicheskii Instituta, 123, 1–196.

    Rasnitsyn, A.P. (1980) Origin and evolution of Hymenoptera. Transactions of the Paleontological Institute of the Academy of Sciences of the USSR, 174, 1–192.

    Rasnitsyn, A.P. (1988) An outline of evolution of hymenopterous insects (order Vespida). Oriental Insects, 22, 115–145.

    Rasnitsyn, A.P. (2000) Testing cladograms by fossil record: the ghost range test. Contributions to Taxonomy 69, 251–258.

    Rasnitsyn, A.P. (2002) Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linn, 1758. In: Rasnitsyn, A.P. & Quicke D.L.J. (Eds), History of Insects. Kluwer Academic Publishers, Dordrecht. pp. 242–254

    Rasnitsyn A.P. (2006) Letopis i cladogramma [Fossil record and cladogram]. In: Rozhnov S.V. (Ed.). Evolutsiya biosphery i bioraznoobraziya [Evolution of biosphere and biodiversity]. KMK Press, Moscow, pp. 39–48.

    Rasnitsyn, A.P., Ansorge, J. & Zhang, H. (2006) Ancestry of the orussid wasps, with description of three new genera and species of Karatavitidae (Hymenoptera = Vespida: Karatavitoidea stat. nov.). Insect Systematics and Evolution, 37, 179–190.

    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 and Evolution, 35, 123–135.

    Rawlings, G.B. (1957) Guiglia schauinslandi (Ashmead) (Hym. Orussidae), a parasite of Sirex noctilio (Fabricius) in New Zealand. The Entomologist, 90, 35–36.

    Rohdendorf, B.B., & Rasnitsyn, A.P. (1980) Historical development of the class Insecta. Transactions of the Paleontological Institute of the Academy of Sciences of the USSR, 175, 1–270.

    Ronquist, F. (1995) Phylogeny and early evolution of the Cynipoidea (Hymenoptera). Systematic Entomology 20, 309–335.

    Ronquist, F. (1999) Phylogeny, classification and evolution of the Cynipoidea. Zoologica Scripta, 28, 139–164.

    Ronquist, F. & Liljeblad, J. (2001) Evolution of the gall wasp-host plant association. Evolution, 55, 2503–2522.

    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.

    Ross, H.H. (1965) A textbook of Entomology. John Wiley and Sons New York, 519 pp.

    Savard, J., Diethard, T., Richards, S., Weinstock, G.M., Gibbs, R.A., Werren, J.H., Tettelin, H. & Lercher, M.J. (2006) Phylogenetic analysis reveals bees and wasps (Hymenoptera) at the base of the radiation of Holometabolous insects. Genome Research 16, 1334–1338.

    Schulmeister, S. (2003a). Genitalia and terminal abdominal segments of male basal Hymenoptera (Insecta): morphology and evolution. Organisms, Diversity and Evolution, 3, 253–279.

    Schulmeister, S. (2003b) Simultaneous analysis of basal Hymenoptera (Insecta), introducing robust-choice sensitivity analysis. Biological Journal of the Linnean Society 79, 245–275. [For additions to and corrections of errors, see Errata and Additions at]

    Schulmeister, S. (2003c) Review of morphological evidence on the phylogeny of basal Hymenoptera (Insecta), with a discussion of the ordering of characters. Biological Journal of the Linnean Society, 79, 209–243.

    Schulmeister, S., (2003d) Morphology and evolution of the tarsal plantulae in Hymenoptera (Insecta), focusing on the basal lineages. Zoologica Scripta, 32, 153–172.

    Schulmeister, S., Wheeler, W.C., & Carpenter, J.M. (2002) Simultaneous analysis of the basal lineages of Hymenoptera (Insecta) using sensitivity analysis. Cladistics, 18, 455–484.

    Sharkey, M.J. (1994). “Am I mad?”: a response to Scott Shaw. Ichnews 14, 2.

    Sharkey, M.J. & Wahl D.B. (1992) Cladistics of the Ichneumonoidea (Hymenoptera). Journal of Hymenopteran Research, 1, 15–61.

    Sharkey, M.J. & Roy, A. (2002) Phylogeny of the Hymenoptera: a reanalysis of the Ronquist et al. (1999) reanalysis, with an emphasis on wing venation and apocritan relationships. Zoologica Scripta, 31, 57–66.

    Shi, M., Chen, X. & van Achterberg, C. (2005) Phylogenetic relationships among the Braconidae (Hymenoptera: Ichneumonoidea) inferred from partial 16S rDNA, 28S rDNA D2,18S rDNA gene sequences and morphological characters. Molecular Systematics and Evolution, 37, 104–116.

    Stork, N.E. (1991) The composition of the arthropod fauna of Bornean lowland rainforest trees. Journal of Tropical Ecology, 7, 161–180.

    Telenga, N.A. (1952) Proiskhozhdenie i Eevolyutsiya Parazitizma u Nasekomykh-naezdnikov i Formirovanie ikh Fauny v SSSR [Origin and Evolution of Parasitism in Parasitic Wasps and Development of Their Fauna in the USSR], Izdatel'stvo Akademii Nauk Ukrainskoi SSR. Kiev, 137 pp.

    Telenga, N.A. (1969) Origin and Evolution of Parasitism in Parasitic Wasps and Development of Their Fauna in the USSR. Israel Program for Scientific Translations, U.S. Department of Commerce, Springfield, Virginia, 112 pp.

    Tillyard, R.J. (1927) On the ancestry of the order Hymenoptera. Transactions of the Entomological Society of London 75, 307–318.

    Trautwein, M., Wiegmann, B., Kim, J., Bertone, M., Winterton, S. (2006) FlyTree: Pursuing a nuclear gene phylogeny of the Holometabola. In: Symposium: Assembling the Tree of Life For Insects and Their Relatives: Emerging Results in the Pursuit of Large-scale Phylogenies. Annual Meeting of the Entomological Society of America, Indianapolis, Indiana.

    Vilhelmsen, L. (1996) The preoral cavity of lower Hymenoptera (Insecta): comparative morphology and phylogenetic significance. Zoologica Scripta, 25, 143–170.

    Vilhelmsen, L. (1997a) Head capsule concavities accommodating the antennal bases in Hymenoptera pupating in wood: possible emergence-facilitating adaptations. International Journal of Insect Morphology and Embryology, 26, 129–138.

    Vilhelmsen, L. (1997b) The phylogeny of lower Hymenoptera (Insecta), with a summary of the early evolutionary history of the order. Journal of Zoological Systematics and Evolutionary Research, 35, 49–70.

    Vilhelmsen, L. (1999) The occipital region in the basal Hymenoptera (Insecta): a reappraisal. Zoologica Scripta, 28, 75–85.

    Vilhelmsen, L. (2000a) 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.

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

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

    Vilhelmsen, L. (2001) Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta). Zoological Journal of the Linnean Society, 131, 393–442.

    Vilhelmsen, L. (2003) Larval anatomy of Orussidae (Hymenoptera). Journal of Hymenoptera Research, 12, 346–354.

    Vilhelmsen, L. (2006) Developments in "symphytan" phylogenetics in the 20th century: towards a consensus. In: Blank, S.M., Schmidt, S. & Taeger, A. (Eds), Recent Sawfly Research: Synthesis and Prospects. Goecke & Evers, Keltern, pp. 31–38.

    Vilhelmsen, L., Isidoro, N., Romani, R., Basibuyuk, H.H. & Quicke, D.L.J. (2001) Host location and oviposition in a basal group of parasitic wasps: the subgenual organ, ovipositor apparatus, and associated structures in the Orussidae (Hymenoptera, Insecta). Zoomorphology, 121, 63–84.

    Wahl, D.B. & Gauld, I.D. (1998) The cladistics and higher classification of the pimpliformes (Hymenoptera: Ichneumonidae). Systematic Entomology, 23, 265–298.

    Wharton, R.A., Shaw, S.R., Sharkey, M.J., Wahl, D.B., Woolley, J.B., Whitfield, J.B., Marsh, P.M. & Johnson, J.W. (1992) Phylogeny of the subfamilies of the family Braconidae: a reassessment. Cladistics, 8, 199–235.

    Wheeler, W.C. (1996) Optimization alignment: the end of multiple sequence alignment in phylogenetics? Cladistics, 12, 1–9.

    Wheeler, W.C., Whiting, M., Wheeler, Q.D. & Carpenter, J.M. (2001) The phylogeny of the extant hexapod orders. Cladistics 17, 113–169.

    Whitfield, J.B. (1992a). Phylogeny of the non-aculeate Aapocrita and the evolution of parasitism in the Hymenoptera. Journal of Hymenopteran Research, 1, 3–14.

    Whitfield, J. B. (1992b) The polyphyletic origin of endoparasitism in the cyclostome lineages of Braconidae (Hymenoptera). Systematic Biology, 17, 273–286.

    Whitfield, J.B. (1997) Molecular and morphological data suggest a common origin for the polydnaviruses among braconid wasps. Naturwissenschaften, 84, 502–507.

    Whitfield, J.B. (1998) Phylogeny and evolution of host-parasitoid interactions in Hymenoptera. Annual Review of Entomology, 43, 129–151.

    Whitfield J.B., Johnson, N.F. & Hamerski, M.R. (1989) Identity and phylogenetic significance of the metapostnotum in nonaculeate Hymenoptera. Annals of the Entomological Society of America, 82, 663–73.

    Whiting, M.F. (2001) Phylogeny of the holometabolous insect orders: molecular evidence. Zoologica Scripta, 31, 3–15.

    Whiting, M.F. (2003) Phylogeny of holometabolous insects: the most successful group of terrestrial organisms. In: J. Cracraft & M. Donoghue eds. Assembling the Tree of Life. Oxford University Press, pp. 345–364.

    Whiting, M.F., Carpenter, J.M., Wheeler, Q.D. & 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.

    Wiegmann, B.M., Kim, J., Trautwein, M.D., Cassel, B.K., Bertone, M., Winterton, S.L., & Yeates, D.K. (submitted) Single-copy nuclear genes resolve the phylogeny of the holometabolous insect orders. Public Library of Science One.

    Zayed, A. & Packer, L. (2005) Complementary sex determination substantially increases extinction proneness of haplodiploid populations. Proceedings of the National Academy of Sciences, 102, 10742–10746.