File: <hymenopt.htm>                                          [For educational purposes only]        Glossary            <Principal Natural Enemy Groups >             <Citations>             <Home>

 

HYMENOPTERA   [Latest Classification]

 

Please refer to the following links for details on this groups:

 

     Hymenoptera = Photos-1,  Photos-2

 

 

          Description & Behavior    Principal Families    References

 

 

Description & Behavior

 

          Hymenoptera is one of the largest orders of insects that includes the sawflies, wasps, bees, and ants. There are over 130,056 identified species.  The name relates to the heavy wings of the insects, and comes from the Greek for hymen membrane and pteron for wing. The hindwings are connected to the forewings by a series of hooks called hamuli.

 

          Females usually have a special ovipositor for inserting eggs into hosts or otherwise remote places. The ovipositor also is usually modified into a stinger. The immatures develop through complete metamorphosis with a worm-like larval stage and an inactive pupal stage before they mature.

 

          The order dates from the Triassic, the oldest fossils belonging to the family Xyelidae. Social hymenopterans first appeared during the Cretaceous. The evolution of this group has been intensively studied by A. Rasnitsyn, M. S. Engel, G. Dlussky (see References). 

 

          The size ranges from medium to large, usually with two pairs of wings. The  mouthparts are for chewing, with well-developed mandibles. Many species have further developed the mouthparts into a lengthy proboscis, with which they can imbibe liquids, such as nectar. They have large compound eyes, and usually three ocelli.

 

          The forward margin of the hindwing bears a number of hooked bristles, or "hamuli", which lock onto the forewing, holding them together. The smaller species may have only two or three hamuli on each side, but the largest wasps may have a considerable number, keeping the wings gripped together very  tightly. The wings have few veins compared with many other insects, especially in the smaller species.

 

          In the more primitive species, the ovipositor takes the shape of a blade, and has adapted to slicing plant tissues. However most species have it modified for piercing, and, in some cases, is several times the length of the body. The ovipositor may also become modified as a stinger, and the eggs are laid from the base of the structure, rather than from the tip, which is used only for boring and  to inject venom. The stinger is used to immobilise prey, but in some wasps and bees may be used for defence.

 

          The larvae of the primitive species resemble caterpillars, and like them, they feed on leaves. They have large chewing mandibles, three thoracic limbs, and usually a number of abdominal prolegs. But unlike caterpillars, the prolegs have no grasping spines, and the antennae are greatly reduced.

 

          The larvae of other Hymenoptera more closely resemble maggots, and are adapted to life in a protected environment. This may be the body of a host, or a cell in a nest, where the adults will care for the larva (e.g., Bethylidae). Such larvae have soft bodies with no limbs. Defecation occurs when they reach adulthood due to having an incomplete digestive tract, probably to avoid contaminating their surroundings.

 

          Sex is determined by the number of chromosomes that an insect has.  Fertilized eggs receive two sets. chromosomes (one from each parent's respective gametes), and so develop into diploid females, while unfertilized eggs only contain one set (from the mother), and so develop into haploid males; the act of fertilization is under the voluntary control of the egg-laying female, a process that is known as haplodiploidy. 

          The actual genetic mechanisms of haplodiploid sex determination may be more complex than simple chromosome number. In many Hymenoptera, sex is actually determined by a single gene locus with many alleles. In these species, haploids are male and diploids heterozygous at the sex locus are female, but occasionally a diploid will be homozygous at the sex locus and develop into a male. This is common in an individual whose parents were siblings or other close relatives. Diploid males are known to be produced by inbreeding in ants, bees and wasps.

 

          One result of haplodiploidy is that females on average actually have more genes in common with their sisters than they do with their own daughters. Because of this, cooperation among kindred females may be unusually advantageous, and may contribute to the multiple origins of eusociality within this order. Different species show a wide range of feeding habits. The most primitive forms are usually herbivorous, feeding on leaves or pine needles. Stinging wasps are predators, and will provide their larvae with immobilised prey, while bees feed on nectar and pollen

 

          Quite a few species are parasitoids as larvae. The adults inject the eggs into a paralysed host, which the larvae begin to consume after hatching. Some species are even hyperparasitoids, with the host itself being another parasitoid insect. Habits intermediate between those of the herbivorous and parasitoid forms occur in some species that inhabit the galls or nests of other insects, utilizing their food, and even killing and devouring the occupant.

 

          The suborder Symphyta includes the sawflies, horntails, and parasitic wood wasps. The group may be paraphyletic, as it has been thought that the family Orussidae may be the ancestral group of the Apocrita. They have an unconstricted junction between the thorax and abdomen, and the larvae of free-living forms are herbivorous, have legs, prolegs (on every segment, unlike Lepidoptera and ocelli.  The wasps, bees, and ants together make up the suborder Apocrita, characterized by a constriction between the first and second abdominal segments called a petiole, that also involves the fusion of the firs abdominal segment to the thorax. Also, the larvae of all Apocrita do not have legs, prolegs, or ocelli

 

Principal Families & Subfamilies

 

Symphyta (Chalastogastra)--sawflies & horntails

    (Sawflies, mostly phytophagous) <General Characteristics>

 

  Orussoidea

    Orussidae <Overview>; <Adults> & <Juveniles>

 

  Tenthredinoidea

    Tenthredinidae <Overview>; <Adults> & <Juveniles>

 

Apocrita (Clistogastra, Petiolata)-- parasitic/predatory

 

 Apocrita Aculeata (stinging Hymenoptera)

  

  Apoidea <Overview>

    Andrenidae <Habits>; <Adults> & <Juveniles>

    Anthophoridae (= Nomadidae) <Habits>;  <Adults> & <Juveniles>

    Apidae <Habits>; <Adults> & <Juveniles>

    Halictidae <Habits>; <Adults> & <Juveniles>

    Colletidae (= Hylaeidae) <Habits>; <Adults> & <Juveniles>

    Ctenoplectridae <Habits>; <Adults> & <Juveniles>

    Fideliidae <Habits>; <Adults> & <Juveniles>

    Hylaeidae <Habits>; <Adults> & <Juveniles>

    Megachilidae <Habits>; <Adults> & <Juveniles>

    Melittidae <Habits>; <Adults> & <Juveniles>

    Nomadidae <Habits>; <Adults> & <Juveniles>

    Oxaeidae <Habits>; <Adults> & <Juveniles>

    Stelidae <Habits>; <Adults> & <Juveniles>

    Stenotritidae <Habits>; <Adults> & <Juveniles>

    Sphecidae (= previously under Sphecoidea) <Habits>; <Adults> & <Juveniles>

      Ampulicinae <Habits>; <Adults> & <Juveniles>

      Astatinae <Habits>; <Adults> & <Juveniles>

      Bembicinae <Habits>; <Adults> & <Juveniles>

      Cercerinae <Habits>; <Adults> & <Juveniles>

      Crabroninae <Habits>; >Adults> & <Juveniles>

      Larrinae <Habits>; <Adults> & <Juveniles>

      Mellininae <Habits><Adults> & <Juveniles>

      Nyssoninae <Habits>; <Adults> & <Juveniles>

      Pemphredoninae <Habits>; <Adults> & <Juveniles>

      Philanthinae <Habits>; <Adults> & <Juveniles>

      Pseninae <Habits>; <Adults> & <Juveniles>

      Sphecinae <Habits>; <Adults> & <Juveniles>

      Stizinae <Habits>; <Adults> & <Juveniles>

      Trypoxylinae <Habits>; <Adults> & <Juveniles>

 

  Chrysidoidea  ( = Bethyloidea)  <Overview>

    Bethylidae <Habits>; <Adults> & <Juveniles>

    Chrysididae <Habits>; <Adults> & <Juveniles>

    (+ Cleptidae <Habits>); <Adults> & <Juveniles>

    Dryinidae <Habits>; <Adults> & <Juveniles>

    Embolemidae <Habits>; <Adults> & <Juveniles>

    Plumariidae <Habits>; <Adults> & <Juveniles>

    Sclerogibbidae <Habits>; <Adults> & <Juveniles>

    Scolebythidae <Habits>; <Adults> & <Juveniles>

 

  Vespoidea <Overview>

    Bradynobaenidae <Habits>; <Adults> & <Juveniles>

    Formicidae <Habits>; <Adults> & <Juveniles> [formerly

         Formicoidea <Overview>]

    Eumenidae <Habits>; <Adults> & <Juveniles>

    Methocidae <Habits>; <Adults> & <Juveniles>

    Sierolomorphidae <Habits>; <Adults> & <Juveniles>

    Vespidae <Habits>; <Adults> & <Juveniles>

    Scoliidae <Habits>; <Adults> & <Juveniles> [formerly as

                Scolioidea <Overview>]

       [ formerly Pompiloidea <Overview>]

    Rhopalosomidae <Habits>; <Adults> & <Juveniles>

    Pompilidae (= Psammocharidae) <Habits>; <Adults> &  <Juveniles>

       [formerly Tiphioidea <Overview>]

    Tiphiidae <Habits>; <Adults> & <Juveniles>

    Sapygidae <Habits>; <Adults> & <Juveniles>

        [formerly Mutillidae <Habits>; <Adults> & <Juveniles>]

        [formerly Thynnidae <Habits>; <Adults> & <Juveniles>]

 

  Apocrita Parasitica (nonstinging Hymenoptera)

  

  Ceraphronoidea <Overivew>

    Calliceratidae <Habits>; <Adults> & <Juveniles>

    Ceraphronidae <Habits>; <Adults> & <Juveniles>

    Megaspilidae <Habits>; <Adults> & <Juveniles>

 

Chalcidoidea <Overview>; <General References>

    Agaonidae <Habits>; <Adults> & <Juveniles>

    Aphelinidae <Habits>; <Adults> & <Juveniles>

    Chalcididae <Habits>; <Adults> & <Juveniles>

    Elasmidae <Habits>; <Adults> & <Juveniles>

    Encyrtidae <Habits>; <Adults-1> & <Adults-2>; & <Juveniles>

    Eucharitidae (= Eucharidae) <Habits>; <Adults> & <Juveniles>

    Eulophidae <Habits>; <Adults> & <Juveniles>

    Eupelmidae <Habits>; <Adults> & <Juveniles>

    Eurytomidae <Habits>; <Adults> & <Juveniles>

    Leucospidae <Habits>; <Adults> & <Juveniles>

    Mymaridae <Habits>; <Adults> & <Juveniles>

    Ormyridae <Habits>; <Adults> & <Juveniles>

    Perilampidae <Habits>; <Adults> & <Juveniles>

 

    Pteromalidae <Habits>; <Adults> & <Juveniles>

      (Asaphinae) <Habits>; <Adults> & <Juveniles>

      (Austroterobiinae <Habits>; <Adults> & <Juveniles>

      (Austrosystasinae) <Habits>; <Adults> & <Juveniles>

      (Brachyscelidiphaginae) <Habits>; <Adults> & <Juveniles>

      (Ceinae) <Habits>; <Adults> & <Juveniles>

      (Cerocephalinae) <Habits>; <Adults> & <Juveniles>

      (Chromeurytominae) <Habits>; <Adults> & <Juveniles>

      (Cleonyminae) <Habits>; <Adults> & <Juveniles>

      (Coelocybinae) <Habits>; <Adults> & <Juveniles>

      (Colotrechinae) <Habits>; <Adults> & <Juveniles>

      (Cratominae) <Habits>; <Adults> & <Juveniles>

      (Diparinae) <Habits>; <Adults> & <Juveniles>

      (Ditropinotellinae) <Habits>; <Adults> & <Juveniles>

      (Elatoidinae) <Habits>; <Adults> & <Juveniles>

      (Erotolepsiinae) <Habits>; <Adults> & <Juveniles>

      (Eunotinae) <Habits>; <Adults> & <Juveniles>

      (Eutrichosomatinae) <Habits>; <Adults> & <Juveniles>

      (Herbertinae) <Habits>; <Adults> & <Juveniles>

      (Keiraninae) <Habits>; <Adults> & <Juveniles>

      (Leptofoeninae) <Habits>; <Adults> & <Juveniles>

      (Louriciinae) <Habits>; <Adults> & <Juveniles>

      (Macromesinae) <Habits>; <Adults> & <Juveniles>

      (Miscogasterinae) <Habits>; <Adults> & <Juveniles>

      (Neodiparinae) <Habits>; <Adults> & <Juveniles>

      (Nefoeninae) <Habits>; <Adults> & <Juveniles>

      (Ormocerinae) <Habits>; <Adults> & <Juveniles>

      (Panstenoninae) <Habits>; <Adults> & <Juveniles>

      (Parasaphodinae) <Habits>; <Adults> & <Juveniles>

       (Pireninae) <Habits>; <Adults> & <Juveniles>

      (Pteromalinae) <Habits>; <Adults> & <Juveniles>

      (Spalangiinae) <Habits>; <Adults> & <Juveniles>

      (Storeyinae) <Habits>; <Adults> & <Juveniles>

 

    Rotoitidae <Habits>; <Adults> & <Juveniles>

    Signiphoridae <Habits>; <Adults> & <Juveniles>

    Tanaostigmatidae <Habits>; <Adults> & <Juveniles>

    Tetracampidae <Habits>; <Adults> & <Juveniles>

    Torymidae <Habits>; <Adults> & <Juveniles>

    Trichogrammatidae <Habits>; <Adults> & <Juveniles>

 

Cynipoidea <Overview>

    Cynipidae <Habits>; <Adults> & <Juveniles>

      (Charipinae) <Habits>; <Adults> & <Juveniles>

      (Cynipinae) <Habits>; <Adults> & <Juveniles>

      (Eucoilinae) <Habits>; <Adults> & <Juveniles>

    Figitidae <Habits>; <Adults> & <Juveniles>

    Ibaliidae <Habits>; <Adults> & <Juveniles>

    Liopteridae <Habits>; <Adults> & <Juveniles>

   

  Evanioidea <Overview>

    Aulacidae <Habits>; <Adults> & <Juveniles>

    Evaniidae <Habits>; <Adults> & <Juveniles>

    Gasteruptiidae <Habits>; <Adults> & <Juveniles>

 

  Ichneumonoidea <Overview>

     Apozygidae <Habits>; <Adults> & <Juveniles>

     Braconidae <Habits>; <Adults-1> & <Adults-2> & <Adults-3>  &  <Juveniles>

     Aphidiinae <Habits>; <Adults> & <Juveniles>

     Ichneumonidae <Habits>; <Adults-1> & <Adults-2> &

           <Adults-3>; & <Juveniles>

 

    Agriotypinae <Habits>; <Adults> & <Juveniles>

    Paxylommatidae <Habits>; <Adults> & <Juveniles>

 

  Megalyroidea <Overview>

    Megalyridae <Habits>; <Adults> & <Juveniles>

 

 ymarommatoidea (or Serphitoidea)

      Mymerommatidae (= Callimomidae) <Habits>; <Adults> &  <Juveniles>

 

    Platygastroidea

       Platygastridae (= Platygasteridae) <Habits>; <Adults> & <Juveniles>

        Scelionidae <Habits>; <Adults> & <Juveniles>

 

  Proctotrupoidea <Overview>

    Austroniidae <Habits>; <Adults> & <Juveniles>

    Diapriidae <Habits>; <Adults> & <Juveniles>

    Heloridae <Habits>; <Adults> & <Juveniles>

      [formerly Mesoserphidae <Habits>; <Adults> & <Juveniles>]

    Monomachidae <Habits>; <Adults> & <Juveniles>

      [formerly Monoserphidae <Habits>; <Adults> & <Juveniles>]

    Pelecinidae <Habits>; <Adults> & <Juveniles>

    Peradeniidae <Habits>; <Adults> & <Juveniles>

    Proctotrupidae (= Serphidae) <Habits>; <Adults> & <Juveniles>

    Roproniidae <Habits>; <Adults> & <Juveniles>

    Serphidae (now with Proctotrupidae) <Habits>; <Adults> & <Juveniles>

    Vanhorniidae <Habits>; <Adults> & <Juveniles>

 

  Stephenoidea <Overview>

    Stephanidae <Habits>; <Adults> & <Juveniles>

 

  Trigonalyoidea <Overview>

    Trigonalyidae <Habits>; <Adults> & <Juveniles>

 

 

 

References:   Please refer to  <biology.ref.htm>, [Additional references may be found at:  MELVYL Library]

 

Achterberg, C. van. 1982. Familietabel van de Hymenoptera in Noordwest‑Europa. Wetenschappelijke Mededelingen van de Koningklijke Nederlandse Natuurhistorische Vereniging 152. 50 pp.

 

Arnett, R.H., Jr. 1985. Hymenoptera (wasps, ants, and bees). Pages 402‑471 in Amer. insects: a handbook of the insects of Amer. north of Mexico. Van Nostrand Reinold, New York, USA. 850 pp.

 

Berland, L., and F. Bernard. 1951a. Hymenopteroides (Symphytes et Terebrants). Pages 771‑975 in Grasse, P.‑P. Traite de zoologie, anatomie, systematique, biologie. Tome X. Insectes superieurs et hemipteroides (Premier fascicule). Masson, Paris, France. 975 pp.

 

Berland, L., and F. Bernard. 1951b. Hymenopteroides (Aculeates). Pages 976‑1276, in Grasse, P.‑P. Traite de zoologie, anatomie, systematique, biologie. Tome X. Insectes superieurs et hemipteroides (Fascicule II). Masson, Paris, France. 973 pp.

 

Berland, L. 1958a. Atlas des Hymenopteres de France, Belgique, Suisse. I. Tenthredes, Parasites, Porte‑aiguillon (Bethylides). Nouvel Atlas d'Entomologie. Editions N. Boubee, Paris, France. 155 pp.

 

Berland, L. 1958b. Atlas des Hymenopteres de France, Belgique, Suisse. II. Porte‑aiguillons: Bethyloides (fin), Scolioides, Formicoides, Pompiloides, Vespoides, Sphecoides, Apoides. Nouvel Atlas d'Entomologie. Editions N. Boubee, Paris, France. 184 pp.

 

Betts, C. 1986. The hymenopterist's handbook. The amateur entomologist. Vol. VII. Second edition. The Amateur Entomologist's Soc., Middlesex, England. 208 pp.

 

Bischoff, H. 1929. Biologie der Hymenopteren: eine Naturgeschischte der Hautflugler. Julius Springer, Berlin, Germany. 606 pp.

 

Bohart, R.M., and A.S. Menke. 1976. Sphecid wasps of the world: a generic revision. University of California Press, Berkeley, California, USA. ix + 695 pp.

 

Borror, D.J., C.A. Triplehorn, and N.F. Johnson. 1989. Order Hymenoptera, sawflies, parasitic wasps, ants, wasps, bees. Pages 665‑‑744 in An introduction to the study of insects. Sixth edition. Saunders College Publishing, Philadelphia, Pennsylvania, USA. 875 pp.

 

Boucek, Z. 1957. BlanokrVidli‑‑Hymenoptera Pages 35‑406 in Kratochvil, J. ed. KhAc Zvlreny CSR. Nakladetelstvi Ceskoslovenske Akademie Ved Prague, Czechoslovakia. 746 pp.

 

Boucek, Z. 1988. Australasian Chalcidoidea (Hymenoptera): a biosystematic revision of genera of fourteen families, with a reclassification of species. C.A.B. International, Wallingford, England. 832 pp.

 

Brown, W.L., Jr. 1982. Hymenoptera. Pages 652‑680 in Parker, S.P., ed. Synopsis and classification of living organisms. Vol. 2. McGraw‑Hill, New York, New York, USA. 1232 pp.

 

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

 

Brues, C.T., A.L. Melander, and F.M. Carpenter. 1954. Order Hymenoptera. Pages 621‑‑684 in Classification of insects: keys to the living and extinct families of insects, and to the living families of other terrestrial arthropods. Bulletin of the Museum of Comparative Zoology 108. 917 pp.

 

Ceballos, G. 1941‑1943. Las tribus de los himenopteros de Espaha. Consejo Superior de Investigaciones Cientificas. Trabajos del Instituto Espahol de Entomologia, Madrid, Spain. 420 pp.

 

Ceballos, G. 1956. Catalogo de los himenopteros de Espaha. Consejo Superior de Investigaciones Cientificas. Trabajos del Instituto Espahol de Entomolog~a, Madrid, Spain. 554 pp.

 

Clausen, C. P.  1940.  Entomophagous Insects.  McGraw-Hill Book Co., NY & London.  688 p.

 

Comstock, J.H. 1918. The wings of insects. Comstock, Ithaca, New York, USA. 430 pp.

 

Comstock, J.H., and J.G. Needham. 1898‑1899. The wings of insects. The Amer. Naturalist 32‑33.

 

Cooper, B.A. ed. 1945. Hymenopertist's handbook. The Amateur Entomologist 7 (40).160 pp. Reprinted 1969, Department of Agriculture, The University, Leeds, England.

 

Costa Lima, A. da. 1960. Insetos do Brasil. 11E Tomo. Capitulo 30. Hymenopteros 1.a Parte. Escola Nacional de Agronomia, Rio de Janeiro, Brazil. Serie Didactica No. 13. 368 pp.

 

Costa Lima, A. da. 1962. Insetos do Brasil. 12E Tomo. Capitulo 30. Hymenopteros 2.a Parte. Escola Nacional de Agronomia, Rio de Janeiro, Brazil. Serie Didactica No. 13. 393 pp.

 

Dalla Torre, C.G. de. 1892‑1902. Catalogus hymenopterorum hucusque descriptorum systematicus et synonymicus. Guilelmi Englemann, Leipsig, Germany.

 

Daly, H.V., J.T. Doyen, and P.R. Ehrlich. 1978. Order Hymenoptera (bees, wasps, ants, etc.). Pages 478‑502 in Introduction to insect biology and diversity. McGraw‑Hill, New York, New York, USA. 564 pp.

 

Day, M.C. 1988. Spider wasps: Hymenoptera: Pompilidae. Handbooks for the identification of British insects. Vol. 6, Part 4. Royal Entomol. Soc. of London, London, England. 60 pp.

 

Delvare, G., and H.‑P. Aberlenc. 1989. Ordre Hymenoptera. Pages 163‑‑200 in Les insectes d'Afrique et d'Amerique tropicale. Cles pour la reconnaissance des familles. PRIFAS, CIRAD‑GERDAT, Montpellier, France. 302 pp.

 

Eady, R.D. 1974. The present state of nomenclature of wing venation in the Braconidae (Hymenoptera): its origins and comparison with related groups. Journal of Entomology, Series B, Taxonomy 43:63‑‑72.

 

Fitton, M.G., M.W.R. de V. Graham, Z.R.J. Boucek, N.D.M. Fergusson, T. Huddleston, J. Quinlan, and O.W. Richards. 1978. A check list of British insects. Second edition (completely revised). Part 4: Hymenoptera. Handbooks for the identification of British insects. Vol. XI, Part 4. Royal Entomol. Soc. of London, London, England. 159 pp.

 

Gauld, I. & B. Bolton.  1988.  The Hymenoptera.  British Mus. (Nat. Hist.), Oxford Univ. Press.  332 p.

 

Gauld, I.D., N. Mark Collins, and M.G. Fitton. 1990. The biological significance and conservation of Hymenoptera in Europe. Nature and Environment Series, No. 44. Council of Europe, Strasbourg, France. 47 pp.

 

Gibson, G.A.P. 1985. Some pro‑ and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms for the structures. Canad. Ent. 117:1395‑1443.

 

Grandi, G. 1966. Ordine Hymenoptera. Pages 611‑638 in Institzioni di entomologia generale. Edizioni Calderini, Bologna, Italy. 654 pp.

 

Grimaldi, D. & M. S. Engle. 2005. Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 

 

Handlirsch, A. 1925. Uberordnung und Ordnung: Hymenoptera L. (Hautflugler). Pages 712‑‑825 in Schroder, C., ed. Handbuch der Entomologie. Band III. Gustav Fischer, Jena, Germany. 1201 pp.

 

Hirashima, Y. 1989. Hymenoptera. Pages 541‑‑692 in A checklist of Japanese insects. Entomol. Laboratory, Faculty of Agriculture, Kyushu, University, Kyushu, Japan. 1767 pp.

 

Hoell, H.V., Doyen, J.T. & Purcell, A.H. 1998. Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. p. 320.

 

Hoell, H.V., Doyen, J.T. & Purcell, A.H. 1998. Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. pp. 570–579.

 

Holldobler, B., and E.O. Wilson. 1990. The ants. The Belknap Press of Harvard University Press Cambridge, Massachusetts, USA. 732 pp.

 

Konigsmann, E. 1976. Das phylogenetisches System der Hymenoptera. Teil 1: Einfuhrung, Grundplanmerkmale, Schwestergruppe und Fossilfunde. Deutsche Entomologische Zeitschrift. N.F. Band 23. Heft IV‑‑V:253‑279.

 

Konigsmann, E. 1977. Das phylogenetisches System der Hymenoptera. Teil 2: "Symphyta". Deutsche Entomologische Zeitschrift, N.F. Band 24, Heft I/III:1‑40.

 

Konigsmann, E. 1978a. Das phylogenetisches System der Hymenoptera. Teil 3: "Terebrantes" (Unterordnung Apocrita). Deutsche Entomologische Zeitschrift, N.F. Band 25, Heft I/III: 1 ‑55.

 

Konigsmann, E. 1978b. Das phylogenetisches System der Hymenoptera. Teil 4: Aculeata (Unterordnung Apocrita). Deutsche Entomologische Zeitschrift, N.F. Band 25, Heft IV ‑ V:365‑435.

 

Krombein, K.V., P.D. Hurd Jr., D.R. Smith, and B.D. Burks. 1979. Catalog of Hymenoptera in Amer. North of Mexico. Vols. 1‑3. Smithsonian Institution Press, Washington, D.C, USA. 2735 pp.

 

Kryzhanovskii, O.L., and S.I. Malyshev. 1963. The Hymenoptera, their origin and evolution. Sovetskaya nauka, Moscow, USSR. 291 pp. [In Russian.]

 

Kukalova‑Peck, J. 1991. Fossil history and the evolution of hexapod structures. Pages 141‑179 in CSIRO, ed. The insects of Australia. A textbook for students and research workers. Second edition. Vol. I. Melbourne University Press, Carlton, Australia. xviii + 542 pp.

 

Landin, B.O. 1971. Hypemoptera. Pages 510‑‑1019 in Faltfauna, Insekter 2:2, pp. 381 ‑1053. Natur och kultur, Storkholm, Sweden.

 

Mackerras, I.M. 1970. Skeletal anatomy. Pages 3 ‑ 28 in The insects of Australia: a textbook for students and research workers. CSIRo ed.Melbourne University Press,  Carlton, Australia. xiii + 1029 pp.

 

Malyshev, S.I. 1959. The Hymenoptera, their origin and evolution. Sovetskaya nauka, Moscow, USSR. 297 pp. [In Russian.]

 

Malyshev, S.I. 1968. Genesis of the Hymenoptera and the phases of their evolution. Methuen, London, England. 319 pp.

 

Masner, L., J.R. Barron, H.V. Danks, A.T. Finnamore, A. Francoeur, G.A.P. Gibson, W.R.M. Mason, and C.M. Yoshimoto. 1978. Hymenoptera. Pages 485‑508, in Danks, H.V., ed. Canada and its insect fauna. Memoirs of the Entomol. Soc. of Canada 108. 573 pp.

 

Mason, W.R.M. 1986. Standard drawing conventions and definitions for venational and other features of wings of Hymenoptera. Proceedings of the Entomol. Soc. of Washington 88:1‑7.

 

Mason, W.R.M. 1990. Cubitus posterior in Hymenoptera. Proceedings of the Entomol. Soc. of Washington 92:93‑97.

 

Michener, C.D. 1944. Comparative external morphology, phylogeny, and a classification of the bees (Hymenoptera). Bulletin of the Amer. Museum of Natural History 82:151‑326.

 

Naumann, I.D. 1992. Hymenoptera (wasps, bees, ants, sawflies). Pages 916‑1000, in CSIRO, ed. The insects of Australia: a textbook for students and research workers. Second edition. Vol. 2, pp. 543‑1137. Melbourne University Press, Carlton, Australia.

 

Nichols, S.W., comp. 1989. The Torre‑Bueno glossary of entomology. The New York Entomol. Soc., New York, New York, USA. 840 pp.

 

Oehlke, J. 1969. Beitrage zur Insektenfauna der DDR: Hymenoptera‑Bestimmungstabellen bis zu den Unterfamilien. Beitrage Entomologie 19:753‑801.

 

Oehlke, J. 1989. Hymenoptera ‑ Hautflugler. Pages 398‑463 in Stresemann, E., H.‑‑J. Hannemann, B. Klausnitzer, and K. Senglaub, eds. Exkursionsfauna fur die Gebiete der DDR und der BDR. Band 2/1 Wirbellose Insekten ­Erster Teil. 8. Auflage. Volk und Wissen, Berlin, Germanv.

 

Pagliano, G., and P. Scaramozzino. 1990. Elenco dei generi di Hymenoptera del mondo. Bollettino della Societa Entomologica Italiana (Supplemento). Vol. 122. 210 pp.

 

Prinsloo, G.L., and G.L. Eardley. 1985. Order Hymenoptera (sawflies, wasps, bees, ants). Pages 393‑451 in Scholtz, C.H., and E. Holm, eds. Insects of Southern Africa. Butterworths, Durban, South Africa. 502 pp.

 

Rasnitsyn, A.P. 1980. Origin and evolution of hymenopterous insects. Trudy Paleontologicheskogo instituta. Akademiya nauk SSSR 174:1‑‑192. [In Russian.]

 

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

 

Rasnitsyn, A.P. 1990. Hymenoptera. Vespida. Pages 177‑205 in Rasnitsyn, A.P., ed. Late Mesozoic insects of eastern Transbaihalia. Nauka Press, Moscow, USSR. 223 pp.

 

Rasnitsyn, A.P. and Quicke, D.L.J.  2002. History of Insects. Kluwer Academic Publishers. pp. 242–254.

 

Rasnitsyn, A.P., and R. Kulicka. 1990. Hymenopteran insects in Baltic amber with respect to the overall history of the order. Prace Museum Ziemi 41:53‑64.

 

Richards, O.W. 1977. Hymenoptera. Introduction and key to families. 2nd edition. Handbooks for the identification of British insects. Vol. VI, Part 1. Royal Entomol. Soc. of London, London, England. 100 pp.

 

Richards, O.W., and R.G. Davies. 1977. Hymenoptera (ants, bees, wasps, ichneumon flies, sawflies, etc.). Pages 1175‑1279 in Imm's general textbook of entomology. 10th edition. Vol. 2. Classification and biology. Chapman and Hall, London, England. 1354 pp.

 

Riek, E.F., 1970. Hymenoptera (wasps, bees, ants). Pages 867‑959 in The insects of Australia. CSIRO, sponsor. Melbourne University Press, Carlton, Australia. xiii + 1029 pp.

 

Riek, E.F., and J.C. Cardale. 1974. Hymenoptera (wasps, bees, ants). Pages 107‑111 in The insects of Australia: a textbook for students and research workers. Supplement 1974. CSIRO, sponsor. Melbourne University Press, Carlton, Australia. 146 pp.

 

Rohwer, S.A., and A.B. Gahan. 1916. Horismology of the hymenopterous wing. Proceedings of the Entomol. Soc. of Washington 19:89‑98.

 

Ronquist, F., and G. Nordlander. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufpes (Hymenoptera: Ibaliidae). Entomologica scandinavica, Supplement No. 33. 60 pp.

 

Ross, H.H. 1936. The ancestry and wing venation of the Hymenoptera. Ann. of the Entomol. Soc. of Amer. 29:99‑111.

 

Ross, H.H. 1937. A generic classification of the Nearctic sawflies (Hymenoptera, Symphyta). Illinois biological monographs 15(2).173 pp.

 

Ross, H.H., C.A. Ross, and J.R.P. Ross. 1982. Hymenoptera. Pages 408--434 in A textbook of entomology. 4th edition. John Wiley and Sons, New York, New York, USA. 666 pp.

 

Schmiedeknecht, 0.1930. Dic Hymenopteren nord‑ und Mitteleuropas. Gustav Fischer, Jena, Germany. 1062 pp.

 

Scholtz, C.H., and E. Holm. 1985. Insects of Southern Africa. Butterworths, Durban, South Africa. 502 pp.

 

Sedivy, J. 1989. Enumeratio Insectarum Bohemoslovakiae. Checklist of Czechoslovak Insects III (Hymenoptera). Acta Faunistica Entomologica Musei Nationalis Pragae. XIX. 194 pp.

 

Snodgrass, R.E. 1935. Principles of insect morphology. McGraw‑Hill, New York, New York, USA. ix + 667 pp.

 

Townes, H.K. 1969. The genera of Ichneumonidae, Part I. Memoirs of the Amer. Entomol. Institute. Number 11. Amer. Entomol. Institute, Gainesville, Florida, USA. 300 pp.

 

Valentine, E.W., and A.H. Walher. 1991. Annotated catalogue of New Zealand. Hymenoptera. DSIR Plant Protection Report No. 4. 84 pp.

 

Whitfield, J.B., N.F. Johnson, and M.R. Hamerski. 1989. Identity and phylogenetic significance of the metapostnotum in nonaculeate Hymenoptera. Ann. of the Entomol. Soc. of Amer. 82:663‑673.

 

Wu, C.F. 1941. Catalogus insectorum sinensium (catalogue of Chinese insects). Vol. 6. Yenching University, Beijing, People's Republic of China.

 

Zahradnik, J. 1985. Bienen, Wespen, Ameisen. Die Hautflugler Mitteleuropas. W. Keller, Stuttgart, Germany. 191 pp.