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Apocrita Symphyta

Hymenoptera
Hymenoptera
is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera
Hymenoptera
have been described,[2][3] in addition to over 2,000 extinct ones.[4] Females typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. The ovipositor is often modified into a stinger. The young develop through holometabolism (complete metamorphosis)—that is, they have a worm-like larval stage and an inactive pupal stage before they mature.

Contents

1 Etymology 2 Evolution 3 Anatomy 4 Reproduction

4.1 Sex determination 4.2 Thelytoky

5 Diet 6 Classification

6.1 Symphyta 6.2 Apocrita

7 See also 8 References 9 Bibliography 10 External links

Etymology[edit] The name Hymenoptera
Hymenoptera
refers to the wings of the insects, but the original derivation is ambiguous.[5]:42 All references agree that the derivation involves the Ancient Greek πτερόν (pteron) for wing. The Ancient Greek ὑμήν (hymen) for membrane provides a plausible etymology for the term because species in this order have membranous wings. However, a key characteristic of this order is that the hind wings are connected to the fore wings by a series of hooks. Thus, another plausible etymology involves Hymen, the Ancient Greek god of marriage, as these insects have "married wings" in flight. Evolution[edit] The cladogram of external relationships, based on a 2008 DNA
DNA
and protein analysis, shows the order as a clade, most closely related to endopterygote orders including the Diptera
Diptera
(true flies) and Lepidoptera
Lepidoptera
(butterflies and moths).[6][7][8][9]

part of Endopterygota

Antliophora

Diptera
Diptera
(true flies)

Mecoptera
Mecoptera
(scorpionflies)

Boreidae
Boreidae
(snow scorpionflies)

Siphonaptera
Siphonaptera
(fleas)

Trichoptera
Trichoptera
(caddisflies)

Lepidoptera
Lepidoptera
(butterflies and moths)

Hymenoptera
Hymenoptera
(sawflies, wasps, ants, bees)

Hymenoptera
Hymenoptera
originated in the Triassic, with the oldest fossils belonging to the family Xyelidae. Social hymenopterans appeared during the Cretaceous.[10] The evolution of this group has been intensively studied by Alex Rasnitsyn, Michael S. Engel, Gennady Dlussky, and others. This clade has been studied by examining the mitochondrial DNA.[11] Although this study was unable to resolve all the ambiguities in this clade, some relationships could be established. The Aculeata, Ichneumonomorpha, and Proctotrupomorpha
Proctotrupomorpha
were monophyletic. The Megalyroidea
Megalyroidea
and Trigonalyoidea are sister clades as are the Chalcidoidea+Diaprioidea. The Cynipoidea
Cynipoidea
was generally recovered as the sister group to Chalcidoidea
Chalcidoidea
and Diaprioidea
Diaprioidea
which are each other's closest relations. The cladogram is based on Schulmeister 2003.[12][13]

Hymenoptera

Hymenoptera, 250mya

  Xyeloidea
Xyeloidea
(Triassic-present)  

  Tenthredinoidea
Tenthredinoidea

  Pamphilioidea
Pamphilioidea

Unicalcarida

  Cephoidea
Cephoidea
(stem sawflies)

  Siricoidea
Siricoidea
(horntails or wood wasps)

  Xiphydrioidea
Xiphydrioidea
(wood wasps)

parasitism

  Orussoidea
Orussoidea
(parasitic wood wasps)  

"wasp waist" 200mya

 APOCRITA (ants, bees, wasps)

Symphyta

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

Anatomy[edit]

Bombus muscorum
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 mouthparts are adapted for chewing, with well-developed mandibles (ectognathous mouthparts). Many species have further developed the mouthparts into a lengthy proboscis, with which they can drink liquids, such as nectar. They have large compound eyes, and typically three simple eyes, 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 ovipositor is blade-like, and has evolved for slicing plant tissues. In the majority, however, it is modified for piercing, and, in some cases, is several times the length of the body. In some species, the ovipositor has 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 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[edit] Sex determination[edit] Main article: Haplodiploid sex-determination system Among most or all hymenopterans, sex is determined by the number of chromosomes an individual possesses.[15] Fertilized eggs get two sets of chromosomes (one from each parent's respective gametes) and develop into diploid females, while unfertilized eggs only contain one set (from the mother) and develop into 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.[15] 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 as a male, instead. This is especially likely to occur in an individual whose parents were siblings or other close relatives. Diploid
Diploid
males are known to be produced by inbreeding in many ant, bee, and wasp species. Diploid
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[edit] Main article: Thelytoky Some hymenopterans take advantage of parthenogenesis, the creation of embryos without fertilization. Thelytoky
Thelytoky
is a particular form of parthenogenesis in which female embryos are created (without fertilisation). The form of thelytoky in hymenopterans is a kind of automixis in which two haploid products (proto-eggs) from the same meiosis fuse to form a diploid zygote. This process tends to maintain heterozygosity in the passage of the genome from mother to daughter. It is found in several ant species including the desert ant Cataglyphis cursor,[19] the clonal raider ant Cerapachys biroi,[20] the predaceous ant Platythyrea punctata,[21] and the electric ant (little fire ant) Wasmannia auropunctata.[22] It also occurs in the Cape honey bee Apis mellifera capensis.[23] Oocytes that undergo automixis with central fusion often have a reduced rate of crossover recombination, which helps to maintain 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. Colonies of this species which have more homozygous queens will age more rapidly, resulting in reduced colony survival.[24] Diet[edit] Different species of Hymenoptera
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
Parasitoid
wasp A huge number of species are parasitoids as larvae. The adults inject the eggs into a host, which they begin to consume after hatching. For example, the eggs of the endangered Papilio homerus
Papilio homerus
are parasitized at a rate of 77%, mainly by Hymenoptera
Hymenoptera
species.[25] Some species are even 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[edit]

Symphyta, without a waist: the sawfly Arge pagana

Apocrita, with narrow waists: the wasp Vespula germanica

The Hymenoptera
Hymenoptera
are divided into two groups; the Symphyta
Symphyta
which have no waist, and the Apocrita
Apocrita
which have a narrow waist.[26] Symphyta[edit] The suborder Symphyta
Symphyta
includes the sawflies, horntails, and parasitic wood wasps. The group may be paraphyletic, as it has been suggested that the family Orussidae
Orussidae
may be the group from which the Apocrita arose. They have an unconstricted junction between the thorax and abdomen. The larvae are herbivorous, free-living eruciforms, with three pairs of true legs, prolegs (on every segment, unlike Lepidoptera) and ocelli. The prolegs do not have crochet hooks at the ends unlike the larvae of the Lepidoptera. Apocrita[edit] The wasps, bees, and ants together make up the suborder (and clade) Apocrita, characterized by a constriction between the first and second abdominal segments called a wasp-waist (petiole), also involving the fusion of the first abdominal segment to the thorax. Also, the larvae of all Apocrita
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[edit]

List of bees, wasps and ants of Great Britain Hymenoptera
Hymenoptera
genome database Insects in literature
Insects in literature
(ant, bee, wasp) Worker policing

References[edit]

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Endopterygota
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Bibliography[edit]

Aguiar, Alexandre P.; Deans, Andrew R.; Engel, Michael S.; Forshage, Mattias; Huber, John T.; Jennings, John T.; Johnson, Norman F.; Lelej, Arkady S.; Longino, John T.; Lohrmann, Volker; Mikó, István; Ohl, Michael; Rasmussen, Claus; Taeger, Andreas; Yu, Dicky Sick Ki (30 August 2013). "Order Hymenoptera". Zootaxa. 3703 (1): 51–62. doi:10.11646/zootaxa.3703.1.12. , in Zhang, Z.-Q. (ed.) Animal Biodiversity: An Outline of Higher-level Classification and Survey of Taxonomic Richness (Addenda 2013) Capinera, John L., ed. (2008). Encyclopedia of Entomology (2nd ed.). Dordrecht: Springer. ISBN 978-1-4020-6242-1.  Carus, Julius Victor; Gerstaecker, C.E.A., eds. (1863). Handbuch der zoologie. Zweiter Band. Leipzig: Engelmann.  Dallas, W.S. (1867). Insecta. pp. 195–484. , in Günther (1867) Dowton, M.; Austin, A. D. (11 October 1994). "Molecular phylogeny of the insect order Hymenoptera: apocritan relationships". Proceedings of the National Academy of Sciences. 91 (21): 9911–9915. Bibcode:1994PNAS...91.9911D. doi:10.1073/pnas.91.21.9911. PMC 44927 . PMID 7937916.  Gerstaecker, A (July 1867). "Ueber die Gattung Oxybelus Latr. und die bei Berlin vorkommenden Arten derselben". Zeitschrift für die gesammten Naturwissenschaft. 30 (VII): 1–144.  Goulet, Henri; Huber, John T., eds. (1993). Hymenoptera
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of the world: An identification guide to families (PDF). Ottawa: Agriculture Canada. ISBN 0-660-14933-8. Archived from the original (PDF) on 2016-03-05.  Gullan, P. J.; Cranston, P. S. (2014). The Insects: An Outline of Entomology (Fifth ed.). Wiley Blackwell.  Günther, A.C.L.G., ed. (1867). The Record of Zoological Literature vol. iv. London: van Voorst.  (see The Record of Zoological Literature) Grimaldi, D. and Engel, M.S. (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. CS1 maint: Multiple names: authors list (link) Hennig, Willi (1969). Die Stammesgeschichte der Insekten. Frankfurt: Waldemar Kramer.  Mao, Meng; Gibson, Tracey; Dowton, Mark (March 2015). "Higher-level phylogeny of the Hymenoptera
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inferred from mitochondrial genomes". Molecular Phylogenetics and Evolution. 84: 34–43. doi:10.1016/j.ympev.2014.12.009. PMID 25542648.  Rasnitsyn, A.P. and Quicke, D.L.J. (2002). History of Insects. Kluwer Academic Publishers. pp. 242–254. ISBN 1-4020-0026-X. CS1 maint: Multiple names: authors list (link) Rasnitsyn, A.P. (1988). "An outline of evolution of hymenopterous insects (order Vespida)". Oriental Insects. 22: 115–145.  Seltmann, Katja Chantre ́ (December 2004). Building web-based interactive keys to the hymenopteran families and superfamilies (PDF). College of Agriculture, University of Kentucky. Archived from the original (M Sc Thesis) on 3 December 2011.  Sharkey, Michael J (2007). "Phylogeny and Classification of Hymenoptera" (PDF). Zootaxa. 1668: 521–548.  Sharkey, Michael J.; Carpenter, James M.; Vilhelmsen, Lars; Heraty, John; Liljeblad, Johan; Dowling, Ashley P.G.; Schulmeister, Susanne; Murray, Debra; Deans, Andrew R.; Ronquist, Fredrik; Krogmann, Lars; Wheeler, Ward C. (February 2012). "Phylogenetic relationships among superfamilies of Hymenoptera". Cladistics. 28 (1): 80–112. doi:10.1111/j.1096-0031.2011.00366.x.  Song, Sheng-Nan; Tang, Pu; Wei, Shu-Jun; Chen, Xue-Xin (16 February 2016). "Comparative and phylogenetic analysis of the mitochondrial genomes in basal hymenopterans". Scientific Reports. 6: 20972. Bibcode:2016NatSR...620972S. doi:10.1038/srep20972. PMC 4754708 . PMID 26879745. 

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v t e

Extant Hymenopteran families

Kingdom: Animalia Phylum: Arthropoda Class: Insecta Subclass: Pterygota Infraclass: Neoptera Superorder: Endopterygota

S y m p h y t a

Xyeloidea

Xyelidae

Pamphilioidea

Megalodontesidae Pamphiliidae
Pamphiliidae
(web-spinning sawflies)

Tenthredinoidea

Argidae
Argidae
(argid sawflies) Blasticotomidae
Blasticotomidae
(fern sawflies) Cimbicidae
Cimbicidae
(cimbicid sawflies) Diprionidae
Diprionidae
(conifer sawflies) Pergidae
Pergidae
(pergid sawflies) Tenthredinidae
Tenthredinidae
(common sawflies)

U n i c.

Cephoidea

Cephidae
Cephidae
(stem sawflies)

Siricoidea

Anaxyelidae
Anaxyelidae
(cedar wood wasps) Siricidae (horntails)

Xiphydrioidea

Xiphydriidae
Xiphydriidae
(wood wasps)

Orussoidea

Orussidae
Orussidae
(parasitic wood wasps)

A p o c r i t a

U n i c a l c a r i d a

P a r a s i t i c a

Ichneumonoidea

Braconidae
Braconidae
(braconids) Ichneumonidae
Ichneumonidae
(ichneumon wasps)

Ceraphronoidea

Ceraphronidae Megaspilidae

Proctotrupomorpha

Platygastroidea

Platygastridae Scelionidae

Cynipoidea

Austrocynipidae Cynipidae (gall wasps) Figitidae Ibaliidae Liopteridae

Proctotrupoidea
Proctotrupoidea
(s.str.)

Proctorenyxidae Roproniidae Heloridae Pelecinidae Peradeniidae Proctotrupidae Vanhorniidae

Diaprioidea

Austroniidae Diapriidae Maamingidae Monomachidae

Mymarommatoidea

Mymarommatidae

Chalcidoidea (chalcid wasps)

Agaonidae (fig wasps) Aphelinidae Chalcididae Encyrtidae Eucharitidae Eulophidae Eupelmidae Eurytomidae Leucospidae Mymaridae (fairyflies) Ormyridae Perilampidae Pteromalidae Rotoitidae Signiphoridae Tanaostigmatidae Tetracampidae Torymidae Trichogrammatidae

Evanioidea

Aulacidae Evaniidae
Evaniidae
(ensign wasps) Gasteruptiidae

Stephanoidea

Stephanidae

Megalyroidea

Megalyridae

Trigonaloidea

Trigonalidae

A c u l e a t a

Chrysidoidea

Bethylidae Chrysididae (cuckoo wasps) Dryinidae Embolemidae Plumariidae Sclerogibbidae Scolebythidae

Vespoidea

Rhopalosomatidae
Rhopalosomatidae
(rhopalosomatid wasps) Vespidae
Vespidae
(paper wasps, potter wasps, pollen wasps, yellowjackets, hornets)

Tiphioidea

Bradynobaenidae Tiphiidae (tiphiid wasps)

Thynnoidea

Chyphotidae Thynnidae
Thynnidae
(flower wasps) Sierolomorphidae
Sierolomorphidae
(sierolomorphid wasps)

Pompiloidea

Mutillidae
Mutillidae
(velvet ants) Myrmosidae Pompilidae (spider wasps) Sapygidae
Sapygidae
(sapygid, or club-horned wasps)

Scolioidea

Scoliidae
Scoliidae
(scoliid wasps)

Formicoidea

Formicidae (ants)

Apoidea

Spheciformes (sphecoid wasps)

Ampulicidae
Ampulicidae
(cockroach wasps) Crabronidae
Crabronidae
(sand wasps, bee wolves) Heterogynaidae Sphecidae
Sphecidae
(thread-waisted wasps)

Anthophila (bees)

Andrenidae
Andrenidae
(mason bees) Apidae
Apidae
(honey bees, bumblebees, cuckoo bees, carpenter bees, orchid bees, stingless bees) Colletidae
Colletidae
(plasterer bees) Halictidae
Halictidae
(sweat bees) Megachilidae
Megachilidae
(mason bees, leafcutter bees) Melittidae Stenotritidae

Italic are paraphyletic groups Based on Malm and Nyman (2015) and Peters et al (2017)

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Eusociality

Topics

Evolution of eusociality Presociality Social insects

Gamergate Group selection Haplodiploidy Identity in social insects Kin recognition Kin selection Sexual selection in social insects Thelytoky Worker policing

Groups

Hymenoptera

Ant Apidae Crabronidae Halictidae Honey bee Vespidae

Mammalia

Blesmol Dwarf mongoose Meerkat

Crustacea

Synalpheus

Thysanoptera

Kladothrips

Hemiptera

Aphididae

Coleoptera

Austroplatypus incompertus

Isoptera

In culture

Bee
Bee
(mythology)

Pioneers, works

Karl von Frisch

The Dancing Bees 1927

Charles Duncan Michener

The Bees of the World 2000

E. O. Wilson

The Ants 1990 Sociobiology: The New Synthesis 1975

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Insect
Insect
orders

Kingdom: Animalia Phylum: Arthropoda (unranked): Pancrustacea Subphylum: Hexapoda

Extant

Monocondylia

Archaeognatha
Archaeognatha
(jumping bristletails)

D i c o n d y l i a

Apterygota *

Thysanura
Thysanura
(Zygentoma) (silverfish, firebrats)

P t e r y g o t a

Palaeoptera

Ephemeropteroidea

Ephemeroptera (mayflies)

Odonatoptera

Odonata
Odonata
(dragonflies, damselflies)

N e o p t e r a

Polyneoptera

Plecoptera
Plecoptera
(stoneflies) Dermaptera (earwigs) Embioptera
Embioptera
(webspinners) Phasmatodea
Phasmatodea
(stick and leaf insects) Notoptera
Notoptera
(ice-crawlers, gladiators) Orthoptera
Orthoptera
(crickets, wetas, grasshoppers, locusts) Zoraptera
Zoraptera
(angel insects)

Dictyoptera

Blattodea
Blattodea
(cockroaches, termites) Mantodea (mantises)

E u m e t a b o l a

Paraneoptera *

Psocodea
Psocodea
(barklice, lice) Thysanoptera
Thysanoptera
(thrips) Hemiptera
Hemiptera
(cicadas, aphids, true bugs)

E n d o p t e r y g o t a

basal

Hymenoptera
Hymenoptera
(sawflies, wasps, ants, bees)

Neuropteroidea

Coleopterida

Strepsiptera
Strepsiptera
(twisted-winged parasites) Coleoptera
Coleoptera
(beetles)

Neuropterida

Raphidioptera (snakeflies) Megaloptera
Megaloptera
(alderflies, dobsonflies, fishflies) Neuroptera
Neuroptera
(net-winged insects: lacewings, mantidflies, antlions)

Panorpida (Mecopterida)

Antliophora

Mecoptera
Mecoptera
(scorpionflies) + Siphonaptera
Siphonaptera
(fleas) Diptera
Diptera
(gnats, mosquitoes, flies)

Amphiesmenoptera

Trichoptera
Trichoptera
(caddisflies) Lepidoptera
Lepidoptera
(moths, butterflies)

Four most speciose orders are marked in bold Italic are paraphyletic groups Based on Sasaki et al. (2013)

Extinct

Aethiocarenodea Alienoptera Archodonata Blattoptera Caloneurodea Campylopteridae Carbotriplurida Coxoplectoptera Diaphanopterodea Eoblattodea Eudiaphanoptera Geroptera Glosselytrodea Heraridea Hypoperlida Lapeyriidae Meganisoptera Megasecoptera Miomoptera Monura Palaeodictyoptera Paoliida Permoplecoptera Protanisoptera Protelytroptera Protephemerida Protodiptera Protorthoptera Protozygoptera Syntonoptera Titanoptera Triadophlebioptera

Extinct incertae sedis families and genera are marked in italic

Wikispecies

Taxon identifiers

Wd: Q22651 ADW: Hymenoptera BugGuide: 59 EoL: 648 EPPO: 1HYMEO Fauna Europaea: 11273 Fossilworks: 70707 GBIF: 1457 ITIS: 152741 NCBI: 7399 WoRMS: 151155

Authority control

LCCN: sh85063592 GND: 41740

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