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* {{cite journal|last1=Dowton|first1=M.|last2=Austin|first2=A. D.|title=Molecular phylogeny of the insect order Hymenoptera: apocritan relationships|journal=[[Proceedings of the National Academy of Sciences]]|date=11 October 1994|volume=91|issue=21|pages=9911–9915|doi=10.1073/pnas.91.21.9911|pmc=44927|ref=harv|pmid=7937916|bibcode=1994PNAS...91.9911D}}
* {{cite journal|last1=Dowton|first1=M.|last2=Austin|first2=A. D.|title=Molecular phylogeny of the insect order Hymenoptera: apocritan relationships|journal=[[Proceedings of the National Academy of Sciences]]|date=11 October 1994|volume=91|issue=21|pages=9911–9915|doi=10.1073/pnas.91.21.9911|pmc=44927|ref=harv|pmid=7937916|bibcode=1994PNAS...91.9911D}}
* {{cite journal|last1=Gerstaecker|first1=A|authorlink=Carl Eduard Adolph Gerstaecker|title=Ueber die Gattung Oxybelus Latr. und die bei Berlin vorkommenden Arten derselben|journal=Zeitschrift für die gesammten Naturwissenschaft|date=July 1867|volume=30|issue=VII|pages=1–144|url=http://www.biodiversitylibrary.org/item/102864#page/17/mode/1up|ref=harv}}
* {{cite journal|last1=Gerstaecker|first1=A|authorlink=Carl Eduard Adolph Gerstaecker|title=Ueber die Gattung Oxybelus Latr. und die bei Berlin vorkommenden Arten derselben|journal=Zeitschrift für die gesammten Naturwissenschaft|date=July 1867|volume=30|issue=VII|pages=1–144|url=http://www.biodiversitylibrary.org/item/102864#page/17/mode/1up|ref=harv}}
* {{cite book|editor1-last=Goulet|editor1-first=Henri|editor2-last=Huber|editor2-first=John T.|title=Hymenoptera of the world: An identification guide to families|date=1993|publisher=Agriculture Canada|location=Ottawa|isbn=0-660-14933-8|url=http://www.esc-sec.ca/aafcmonographs/hymenoptera_of_the_world.pdf}}
* {{cite book|editor1-last=Goulet|editor1-first=Henri|editor2-last=Huber|editor2-first=John T.|title=Hymenoptera of the world: An identification guide to families|date=1993|publisher=Agriculture Canada|location=Ottawa|isbn=0-660-14933-8|url=http://www.esc-sec.ca/aafcmonographs/hymenoptera_of_the_world.pdf|deadurl=yes|archiveurl=https://web.archive.org/web/20160305012733/http://www.esc-sec.ca/aafcmonographs/hymenoptera_of_the_world.pdf|archivedate=2016-03-05|df=}}
* {{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. | title=The Insects: An Outline of Entomology | publisher=Wiley Blackwell | edition=Fifth | year=2014}}
* {{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. | title=The Insects: An Outline of Entomology | publisher=Wiley Blackwell | edition=Fifth | year=2014}}
* {{cite book|editor1-last=Günther|editor1-first=A.C.L.G.|title=The Record of Zoological Literature vol. iv|date=1867|publisher=van Voorst|location=London|url=https://books.google.ca/books?id=i8pDAQAAMAAJ|ref=harv}} (see [[The Record of Zoological Literature]])
* {{cite book|editor1-last=Günther|editor1-first=A.C.L.G.|title=The Record of Zoological Literature vol. iv|date=1867|publisher=van Voorst|location=London|url=https://books.google.ca/books?id=i8pDAQAAMAAJ|ref=harv}} (see [[The Record of Zoological Literature]])

Revision as of 16:00, 29 December 2017

Hymenoptera
Temporal range:
Ma[1]
A digger wasp, Sphex pensylvanicus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Holometabola
Superorder: Hymenopterida
Order: Hymenoptera
Linnaeus, 1758
Suborders

Apocrita
Symphyta

Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described,[2][3] in addition to over 2,000 extinct ones.[4]

Females typically have a special

metamorphosis
)—that is, they have a worm-like larval stage and an inactive pupal stage before they mature.

Etymology

The name Hymenoptera refers to the wings of the insects, but the original derivation is ambiguous.

hooks. Thus, another plausible etymology involves Hymen
, the Ancient Greek god of marriage, as these insects have "married wings" in flight.

Evolution

The

Diptera (true flies) and Lepidoptera (butterflies and moths).[6][7][8][9]

part of 
Endopterygota
Antliophora

Diptera (true flies)

Mecoptera (scorpionflies)

Boreidae (snow scorpionflies)

Siphonaptera (fleas)

Trichoptera (caddisflies)

Lepidoptera (butterflies and moths)

Hymenoptera (sawflies, wasps, ants, bees)

Hymenoptera originated in the

Alex Rasnitsyn, Michael S. Engel, Gennady Dlussky
, and others.

This clade has been studied by examining the mitochondrial DNA.

Chalcidoidea and Diaprioidea
which are each other's closest relations.

The cladogram is based on Schulmeister 2003.[12][13]

Hymenoptera
Hymenoptera, 250mya

Xyeloidea (Triassic-present)

Tenthredinoidea

Pamphilioidea

Cephoidea (stem sawflies)

Siricoidea (horntails or wood wasps)

Xiphydrioidea (wood wasps)

parasitism

parasitic wood wasps)

"wasp waist" 200mya

APOCRITA (ants, bees, wasps)

Symphyta
Symphyta (red bar) are paraphyletic as Apocrita are excluded.

Anatomy

Bombus muscorum drinking nectar with its long proboscis

Hymenopterans range in size from very small to large insects, and usually have two pairs of wings. Their

ocelli
.

The forward margin of the hind wing bears a number of hooked bristles, or "

hamuli
", which lock onto the fore wing, keeping them held 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 especially tightly. Hymenopteran wings have relatively few veins compared with many other insects, especially in the smaller species.

In the more ancestral hymenopterans, the

eggs are laid from the base of the structure, rather than from the tip, which is used only to inject venom. The sting is typically used to immobilise prey, but in some wasps and bees may be used in defense.[14]

The larvae of the more ancestral hymenopterans resemble caterpillars in appearance, and like them, typically feed on leaves. They have large chewing mandibles, three pairs of thoracic limbs, and, in most cases, a number of abdominal prolegs. Unlike caterpillars, however, the prolegs have no grasping spines, and the antennae are reduced to mere stubs.

The larvae of other hymenopterans, however, more closely resemble maggots, and are adapted to life in a protected environment. This may be the body of a host organism, or a cell in a nest, where the adults will care for the larva. Such larvae have soft bodies with no limbs. They are also unable to defecate until they reach adulthood due to having an incomplete digestive tract, presumably to avoid contaminating their environment.[14]

Reproduction

Sex determination

Main article:
Haplodiploid sex-determination system

Among most or all hymenopterans, sex is

haploid males. The act of fertilization is under the voluntary control of the egg-laying female, giving her control of the sex of her offspring.[14] This phenomenon is called haplodiploidy
.

However, 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.

siblings or other close relatives. Diploid males are known to be produced by inbreeding in many ant, bee, and wasp species. Diploid biparental males are usually sterile but a few species that have fertile diploid males are known.[16]

One consequence 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 has been hypothesized to contribute to the multiple origins of eusociality within this order.[14] In many colonies of bees, ants, and wasps, worker females will remove eggs laid by other workers due to increased relatedness to direct siblings, a phenomenon known as worker policing.[17]

Another consequence is that hymenopterans may be more resistant to the deleterious effects of inbreeding. As males are haploid, any recessive genes will automatically be expressed, exposing them to natural selection. Thus, the genetic load of deleterious genes is purged relatively quickly.[18]

Thelytoky

Main article: Thelytoky

Some hymenopterans take advantage of

Apis mellifera capensis.[23]

heterozygosity and avoid inbreeding depression. Species that display central fusion with reduced recombination include the ants Platythyrea punctata[21] and Wasmannia auropunctata[22] and the honey bee Apis mellifera capensis.[23] In A. m. capensis, the recombination rate during meiosis is reduced more than 10-fold.[23] In W. auropunctata the reduction is 45-fold.[22]

Single queen colonies of the narrow headed ant Formica exsecta illustrate the possible deleterious effects of increased homozygosity. In this ant, colonies with more homozygous queens age more rapidly. The result is reduced colony survival.[24]

Diet

Different species of Hymenoptera show a wide range of feeding habits. The most primitive forms are typically herbivorous, feeding on leaves or pine needles. Stinging wasps are predators, and will provision their larvae with immobilised prey, while bees feed on nectar and pollen.

Main article: Parasitoid wasp

A huge number of species are

hyperparasitoid, with the host itself being another parasitoid insect. Habits intermediate between those of the herbivorous and parasitoid forms are shown in some hymenopterans, which inhabit the galls or nests of other insects, stealing their food, and eventually killing and eating the occupant.[14]

Classification

Symphyta, without a waist: the sawfly Arge pagana
Apocrita, with narrow waists: the wasp Vespula germanica

The Hymenoptera are divided into two groups; the Symphyta which have no waist, and the Apocrita which have a narrow waist.[26]

Symphyta

The suborder

ocelli
. The prolegs do not have crochet hooks at the ends unlike the larvae of the Lepidoptera.

Apocrita

The wasps, bees, and ants together make up the suborder (and clade)

thorax. Also, the larvae of all Apocrita lack legs, prolegs, or ocelli. The hindgut of the larvae also remains closed during development, with feces being stored inside the body, with the exception of some bee larvae where the larval anus has reappeared through developmental reversion. In general, the anus only opens at the completion of larval growth.[27]

See also

References

  1. PMID 22723471
    .
  2. PMID 17624962
    .
  3. PMID 23936325.{{cite journal}}: CS1 maint: unflagged free DOI (link
    )
  4. doi:10.11646/zootaxa.3703.1.12. {{cite journal}}: Explicit use of et al. in: |first5= (help
    )
  5. ^ Grissell, Eric (2010). Bees, Wasps, and Ants: The Indispensable Role of Hymenoptera in Gardens. Timber Press.
  6. doi:10.1111/j.1096-0031.2008.00211.x
    .
  7. ^ Yeates, David K.; Wiegmann, Brian. "Endopterygota Insects with complete metamorphosis". Tree of Life. Retrieved 24 May 2016.
  8. doi:10.1046/j.0300-3256.2001.00095.x
    .
  9. ISBN 978-0-231-50170-5
    .
  10. ISBN 0-19-510033-6.{{cite book}}: CS1 maint: multiple names: authors list (link
    )
  11. ^ Mao M, Gibson T, Dowton M (2014) Higher-level phylogeny of the Hymenoptera inferred from mitochondrial genomes. Mol Phylogenet Evol
  12. doi:10.1046/j.1095-8312.2003.00233.x
    .
  13. ^ Schulmeister, S. "'Symphyta'". Retrieved 28 November 2016.
  14. ^
    ISBN 0-19-510033-6.{{cite book}}: CS1 maint: multiple names: authors list (link
    )
  15. ^
    PMID 15232002
    .
  16. PMID 19551142.{{cite journal}}: CS1 maint: unflagged free DOI (link
    )
  17. ^ Davies, N.R., Krebs, J.R., and West, S.A. An Introduction to Behavioral Ecology. 4th ed. West Sussex: Wiley-Blackwell, 2012. Print. pp. 387–388
  18. OCLC 28576921.{{cite book}}: CS1 maint: multiple names: authors list (link
    )
  19. PMID 15576621
    .
  20. PMID 24508170
    .
  21. ^
    doi:10.1007/s10682-010-9382-5
  22. ^
    PMID 21459760
    .
  23. ^
    PMID 15166151
    .
  24. PMID 19127611
    .
  25. doi:10.3390/insects8030068.{{cite journal}}: CS1 maint: unflagged free DOI (link
    )
  26. PMID 26146682
    .
  27. ISBN 978-0-19-804207-5
    .

Bibliography

External links

Wikimedia Commons has media related to Hymenoptera.
Wikispecies has information related to Hymenoptera.
Look up Hymenoptera in Wiktionary, the free dictionary.
General
Systematics
Regional Lists
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