# Hymenoptera

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**Hymenoptera** is a large [order](/source/Order_(biology)) of [insects](/source/Insect), comprising the [sawflies](/source/Sawflies), [wasps](/source/Wasp), [bees](/source/Bee), and [ants](/source/Ant). Over 150,000 living species of Hymenoptera have been described,[1][2] in addition to over 2,000 extinct ones.[3] Many of the species are [parasitic](/source/Parasitoid_wasp). Females typically have a special [ovipositor](/source/Ovipositor) for inserting eggs into hosts or places that are otherwise inaccessible. This ovipositor is often modified into a [stinger](/source/Stinger). The young develop through [holometabolism](/source/Holometabolism) (complete [metamorphosis](/source/Metamorphosis))—that is, they have a wormlike larval stage and an inactive pupal stage before they reach adulthood.

## Etymology

The name *Hymenoptera* comes from [Ancient Greek](/source/Ancient_Greek) [ὑμήν](https://en.wiktionary.org/wiki/%E1%BD%91%CE%BC%CE%AE%CE%BD) (*humḗn*) 'membrane' and [πτερόν](https://en.wiktionary.org/wiki/%CF%80%CF%84%CE%B5%CF%81%CF%8C%CE%BD) (*pterón*) 'wing'.[4]

## Evolution

Molecular analysis finds that Hymenoptera is the earliest branching group of [Holometabola](/source/Holometabola).[5]

Hymenoptera originated in the [Triassic](/source/Triassic), with the oldest fossils belonging to the family [Xyelidae](/source/Xyelidae). Social hymenopterans appeared during the [Cretaceous](/source/Cretaceous).[6] The evolution of this group has been intensively studied by [Alex Rasnitsyn](/source/Alex_Rasnitsyn), [Michael S. Engel](/source/Michael_S._Engel), and others.[7]

Phylogenetic relationships within the Hymenoptera, based on both morphology and molecular data, have been intensively studied since 2000.[8] In 2023, a molecular study[8] based on the analysis of [ultra-conserved elements](/source/Ultraconserved_element) confirmed many previous findings and produced a relatively robust phylogeny of the whole Order, although the relationships among the Symphyta lineages are still a source of active research [9][10]. Basal superfamilies are shown in the cladogram below.

## Anatomy

Hymenopterans range in size from very small to large insects. Their [mouthparts](/source/Insect_mouthparts) are adapted for chewing, with well-developed [mandibles](/source/Mandible_(insect_mouthpart)) (ectognathous mouthparts). Many species have further developed the mouthparts into a lengthy [proboscis](/source/Proboscis), with which they can drink liquids, such as [nectar](/source/Nectar).[11] Hymenopterans usually have two pairs of wings, but some solitary wasps and worker ants don't. They typically have large [compound eyes](/source/Compound_eye) with three simple eyes, [ocelli](/source/Ocelli).[12]

The forward margin of the hind wing bears a number of hooked bristles, or "[hamuli](/source/Hamuli)", which lock onto the fore wing, keeping them held together ([hamuli wing coupling](/source/Wing_coupling#Hamuli_wing_coupling)). 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[13] other insects, especially in the smaller species.

In the more ancestral hymenopterans, the [ovipositor](/source/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](/source/Stinger), and the [eggs](/source/Egg_(biology)) are laid from the base of the structure, rather than from the tip, which is used only to inject [venom](/source/Venom). The sting is typically used to immobilize prey, but in some wasps and bees may be used in defense.[6]

Hymenopteran larvae typically have a distinct head region, three thoracic segments, and usually nine or 10 abdominal segments. In the suborder [Symphyta](/source/Symphyta), the [eruciform](/source/Eruciform) larvae resemble [caterpillars](/source/Caterpillar) in appearance, and like them, typically feed on leaves. They have large chewing mandibles, three pairs of thoracic limbs, and, in most cases, six or eight abdominal [prolegs](/source/Proleg). Unlike caterpillars, however, the prolegs have no grasping spines, and the antennae are reduced to mere stubs. Symphytan larvae that are wood borers or stem borers have no abdominal legs and the thoracic legs are smaller than those of non-borers.

With rare exceptions, larvae of the suborder [Apocrita](/source/Apocrita) have no legs and are [maggotlike](/source/Maggot) in form, 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. In parasitic forms, the head is often greatly reduced and partially withdrawn into the prothorax (anterior part of the thorax). Sense organs appear to be poorly developed, with no ocelli, very small or absent antennae, and toothlike, sicklelike, or spinelike mandibles. They are also unable to defecate until they reach adulthood due to having an incomplete digestive tract (a blind sac), presumably to avoid contaminating their environment.[6] The larvae of stinging forms ([Aculeata](/source/Aculeata)) generally have 10 pairs of spiracles, or breathing pores, whereas parasitic forms usually have nine pairs present.[14]

## Reproduction

### Sex determination

Main article: [Haplodiploid sex-determination system](/source/Haplodiploid_sex-determination_system)

Among most or all hymenopterans, sex is [determined](/source/Sex-determination_system) by the number of [chromosomes](/source/Chromosome) an individual possesses.[15] Fertilized eggs get two sets of chromosomes (one from each parent's respective [gametes](/source/Gamete)) and develop into [diploid](/source/Diploid) females, while unfertilized eggs only contain one set (from the mother) and develop into [haploid](/source/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.[6] This phenomenon is called [haplodiploidy](/source/Haplodiploidy).

However, the actual genetic mechanisms of haplodiploid sex determination may be more complex than simple chromosome number. In many Hymenoptera, sex is 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](/source/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 have more genes in common with their sisters than they do with their daughters. Because of this, cooperation among kindred females may be unusually advantageous and has been hypothesized to contribute to the multiple origins of [eusociality](/source/Eusociality) within this order.[6][17] 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](/source/Worker_policing).[18]

Another consequence is that hymenopterans may be more resistant to the deleterious effects of [inbreeding](/source/Inbreeding_depression). As males are haploid, any recessive genes will automatically be expressed, exposing them to natural selection. Thus, the [genetic load](/source/Genetic_load) of deleterious genes is purged relatively quickly.[19]

### Thelytoky

Main article: [Thelytoky](/source/Thelytoky)

Some hymenopterans take advantage of [parthenogenesis](/source/Parthenogenesis), the creation of [embryos](/source/Embryo) without [fertilization](/source/Fertilization). [Thelytoky](/source/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](/source/Meiosis) fuse to form a diploid zygote. This process tends to maintain [heterozygosity](/source/Zygosity) in the passage of the genome from mother to daughter. It is found in several ant species including the desert ant *[Cataglyphis cursor](/source/Cataglyphis_cursor)*,[20] the clonal raider ant *[Cerapachys biroi](/source/Cerapachys_biroi)*,[21] the predaceous ant *[Platythyrea punctata](/source/Platythyrea_punctata)*,[22] and the electric ant (little fire ant) *[Wasmannia auropunctata](/source/Wasmannia_auropunctata)*.[23] It also occurs in the Cape honey bee *[Apis mellifera capensis](/source/Apis_mellifera_capensis)*.[24]

[Oocytes](/source/Oocyte) that undergo automixis with central fusion often have a reduced rate of [crossover recombination](/source/Chromosomal_crossover), which helps to maintain [heterozygosity](/source/Heterozygosity) and avoid [inbreeding depression](/source/Inbreeding_depression). Species that display central fusion with reduced recombination include the ants *[Platythyrea punctata](/source/Platythyrea_punctata)*[22] and *[Wasmannia auropunctata](/source/Wasmannia_auropunctata)*[23] and the Cape honey bee *Apis mellifera capensis*.[24] In *A. m. capensis*, the recombination rate during meiosis is reduced more than tenfold.[24] In *W. auropunctata* the reduction is 45 fold.[23]

Single queen colonies of the narrow headed ant *[Formica exsecta](/source/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.[25]

## Diet

Different species of Hymenoptera show a wide range of feeding habits. The most primitive forms are typically phytophagous, feeding on flowers, pollen, foliage, or stems. Stinging wasps are predators, and will provision their larvae with immobilized prey, while bees feed on nectar and [pollen](/source/Pollen).

Main article: [Parasitoid wasp](/source/Parasitoid_wasp)

A huge number of species are [parasitoids](/source/Parasitoid) 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](/source/Papilio_homerus)* are parasitized at a rate of 77%, mainly by Hymenoptera species.[26] Some species are even [hyperparasitoid](/source/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.[6]

## Classification

The Hymenoptera are divided into two groups; the [Symphyta](/source/Symphyta) which have no waist, and the [Apocrita](/source/Apocrita) which have a narrow waist.[3]

### Symphyta

The suborder [Symphyta](/source/Symphyta) includes the [sawflies](/source/Sawflies), [horntails](/source/Horntail), and [parasitic wood wasps](/source/Orussidae). The group is [paraphyletic](/source/Paraphyletic), as it has been suggested that the family [Orussidae](/source/Orussidae) may be the group from which the [Apocrita](/source/Apocrita) arose. They have an unconstricted junction between the thorax and abdomen. The larvae are herbivorous, free-living, and [eruciform](/source/Eruciform), usually with three pairs of true legs, [prolegs](/source/Proleg) (on every segment, unlike [Lepidoptera](/source/Lepidoptera)) and [ocelli](/source/Ocelli). The prolegs do not have [crochet hooks](/source/Crochet_(insect_anatomy)) at the ends unlike the larvae of the Lepidoptera.[3] The legs and prolegs tend to be reduced or absent in larvae that mine or bore plant tissue, as well as in larvae of [Pamphiliidae](/source/Pamphiliidae).[27]

### Apocrita

The [wasps](/source/Wasp), [bees](/source/Bee), and [ants](/source/Ant) together make up the suborder (and clade) [Apocrita](/source/Apocrita), characterized by a constriction between the first and second abdominal segments called a wasp-waist ([petiole](/source/Petiole_(insect))), also involving the fusion of the first abdominal segment to the [thorax](/source/Thorax_(insect_anatomy)). 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.[clarification needed] In general, the anus only opens at the completion of larval growth.[3]

## Threats

Hymenoptera as a group are highly susceptible to habitat loss, which can lead to substantial decreases in species richness and have major ecological implications due to their pivotal role as plant pollinators.[28]

## See also

- [Hymenoptera Genome Database](/source/Hymenoptera_Genome_Database)
- [Insects in literature](/source/Insects_in_literature) (ant, bee, wasp)
- [Worker policing](/source/Worker_policing)

## References

1. Mayhew, Peter J. (2007). "Why are there so many insect species? Perspectives from fossils and phylogenies". *Biological Reviews*. **82** (3): 425–454. [doi:10.1111/j.1469-185X.2007.00018.x](https://doi.org/10.1111/j.1469-185X.2007.00018.x). [PMID 17624962](https://pubmed.ncbi.nlm.nih.gov/17624962). [S2CID 9356614](https://api.semanticscholar.org/CorpusID:9356614)

1. Janke, Axel; Klopfstein, Seraina; Vilhelmsen, Lars; Heraty, John M.; Sharkey, Michael; Ronquist, Fredrik (2013). "The Hymenopteran Tree of Life: Evidence from Protein-Coding Genes and Objectively Aligned Ribosomal Data". *PLOS ONE*. **8** (8). [Bibcode:2013PLoSO...869344K](https://ui.adsabs.harvard.edu/abs/2013PLoSO...869344K). [doi:10.1371/journal.pone.0069344](https://doi.org/10.1371/journal.pone.0069344). [PMC 3732274](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3732274). [PMID 23936325](https://pubmed.ncbi.nlm.nih.gov/23936325)

1. Aguiar, A.P.; Deans, A.R.; Engel, M.S.; Forshage, M.; Huber, J.T.; Jennings, J.T.; Johnson, N.F.; Lelej, A.S.; Longino, J.T.; Lohrmann, V.; Mikó, I.; Ohl, M.; Rasmussen, C.; Taeger, A.; Yu, D.S.K. (2013). "Order Hymenoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal Biodiversity: An Outline of Higher-level Classification and Survey of Taxonomic Richness (Addenda 2013)". *Zootaxa*. **3703**: 1–82. [doi:10.11646/zootaxa.3703.1.12](https://doi.org/10.11646/zootaxa.3703.1.12). [PMID 26146682](https://pubmed.ncbi.nlm.nih.gov/26146682)

1. Carpenter, George Herbert. Vol. 14. p. 177.

1. Kjer, Karl M.; Simon, Chris; Yavorskaya, Margarita; & Beutel, Rolf G. (2016). "Progress, pitfalls and parallel universes: a history of insect phylogenetics". *Journal of the Royal Society Interface*. **13** (121): 121. [doi:10.1098/rsif.2016.0363](https://doi.org/10.1098/rsif.2016.0363). [PMC 5014063](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5014063). [PMID 27558853](https://pubmed.ncbi.nlm.nih.gov/27558853)

1. Howell, H.V.; Doyen, J.T.; Purcell, A.H. (1998). *Introduction to Insect Biology and Diversity*. 2nd ed. Oxford University Press. p. 320. ISBN 978-0-19-510033-4.

1. Peters, Ralph S.; Krogmann, Lars; Mayer, Christoph; Donath, Alexander; Gunkel, Simon; Meusemann, Karen; Kozlov, Alexey; Podsiadlowski, Lars; Petersen, Malte (April 2017). "Evolutionary History of the Hymenoptera". *Current Biology*. **27** (7): 1013–1018. [Bibcode:2017CBio...27.1013P](https://ui.adsabs.harvard.edu/abs/2017CBio...27.1013P). [doi:10.1016/j.cub.2017.01.027](https://doi.org/10.1016/j.cub.2017.01.027). [hdl:2434/801122](https://hdl.handle.net/2434/801122). [PMID 28343967](https://pubmed.ncbi.nlm.nih.gov/28343967)

1. Wikidata item [Q117865968](https://www.wikidata.org/wiki/Q117865968)

1. Wutke, Saskia; Blank, Stephan M.; Boevé, Jean-Luc; Faircloth, Brant C.; Koch, Frank; Linnen, Catherine R.; Malm, Tobias; Niu, Gengyun; Prous, Marko; Schiff, Nathan M.; Schmidt, Stefan; Taeger, Andreas; Vilhelmsen, Lars; Wahlberg, Niklas; Wei, Meicai (2024-10-01). ["Phylogenomics and biogeography of sawflies and woodwasps (Hymenoptera, Symphyta)"](https://www.sciencedirect.com/science/article/pii/S1055790324001362). *Molecular Phylogenetics and Evolution*. **199**. [doi:10.1016/j.ympev.2024.108144](https://doi.org/10.1016/j.ympev.2024.108144). [ISSN 1055-7903](https://www.worldcat.org/issn/1055-7903)

1. Zhang, Y. Miles; Bossert, Silas; Spasojevic, Tamara (January 2025). ["Evolving perspectives in Hymenoptera systematics: Bridging fossils and genomes across time"](https://resjournals.onlinelibrary.wiley.com/doi/10.1111/syen.12645). *Systematic Entomology*. **50** (1): 1–31. [doi:10.1111/syen.12645](https://doi.org/10.1111/syen.12645). [ISSN 0307-6970](https://www.worldcat.org/issn/0307-6970)

1. Snodgrass, R E (1942). ["The Skeleto-muscular Mechanisms of the Honey Bee"](https://repository.si.edu/bitstream/handle/10088/22771/SMC_103_Snodgrass_1942_2_1-120.pdf). *Smithsonian Miscellaneous Collections*. **103** (2). [Archived](https://web.archive.org/web/20220124231429/https://repository.si.edu/bitstream/handle/10088/22771/SMC_103_Snodgrass_1942_2_1-120.pdf) 2022-01-24 at the Wayback Machine. Retrieved 2026-01-30.

1. ["Hymenoptera - Larvae, Metamorphosis, Sociality | Britannica"](https://www.britannica.com/animal/hymenopteran/Features-of-immature-stages). *www.britannica.com*. [Archived](https://web.archive.org/web/20250911234222/https://www.britannica.com/animal/hymenopteran/Features-of-immature-stages) 2025-09-11 at the Wayback Machine. Retrieved 2025-10-31.

1. ["Hymenoptera - Field Guide to Common Texas Insects"](https://texasinsects.tamu.edu/hymenoptera/). *Texas A&M AgriLife Extension*. [Archived](https://web.archive.org/web/20251018193847/https://texasinsects.tamu.edu/hymenoptera/) 18 October 2025 at the Wayback Machine. Retrieved 2 March 2026.

1. Hunt, James H. (2007). [*The Evolution of Social Wasps*](https://books.google.com/books?id=bFMoSxCVSVIC&pg=PA12). Oxford University Press, USA. p. 12. ISBN 978-0-19-804207-5.

1. Cowan, David P. Cowan & Stahlhut, Julie K. (13 July 2004). "Functionally reproductive diploid and haploid males in an inbreeding hymenopteran with complementary sex determination". *PNAS*. **101** (28): 10374–10379. [Bibcode:2004PNAS..10110374C](https://ui.adsabs.harvard.edu/abs/2004PNAS..10110374C). [doi:10.1073/pnas.0402481101](https://doi.org/10.1073/pnas.0402481101). [PMC 478579](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC478579). [PMID 15232002](https://pubmed.ncbi.nlm.nih.gov/15232002)

1. Elias, J.; Mazzi, D.; Dorn, S. (2009). "No Need to Discriminate? Reproductive Diploid Males in a Parasitoid with Complementary Sex Determination". *PLOS ONE*. **4** (6). [Bibcode:2009PLoSO...4.6024E](https://ui.adsabs.harvard.edu/abs/2009PLoSO...4.6024E). [doi:10.1371/journal.pone.0006024](https://doi.org/10.1371/journal.pone.0006024). [PMC 2696080](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2696080). [PMID 19551142](https://pubmed.ncbi.nlm.nih.gov/19551142)

1. Quiñones, Andrés E. & Pen, Ido (June 2017). "A unified model of Hymenopteran preadaptations that trigger the evolutionary transition to eusociality". *Nature Communications*. **8**. [Bibcode:2017NatCo...815920Q](https://ui.adsabs.harvard.edu/abs/2017NatCo...815920Q). [doi:10.1038/ncomms15920](https://doi.org/10.1038/ncomms15920). [PMC 5490048](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5490048). [PMID 28643786](https://pubmed.ncbi.nlm.nih.gov/28643786)

1. Davies, N.R.; Krebs, J.R.; and West, S.A. An Introduction to Behavioral Ecology. 4th ed. West Sussex: Wiley-Blackwell, 2012. pp. 387–388

1. LaSalle, John & David, Gauld, Ian (1993). *Hymenoptera and biodiversity*. C.A.B. International. ISBN 978-0-85198-830-6. [OCLC 28576921](https://www.worldcat.org/oclc/28576921)

1. Pearcy, M.; Aron, S.; Doums, C.; Kelle, L. (2004). ["Conditional use of sex and parthenogenesis for worker and queen production in ants"](https://dipot.ulb.ac.be/dspace/bitstream/2013/18807/1/061PearcyScience04.pdf). *Science*. **306** (5702): 1780–3. [Bibcode:2004Sci...306.1780P](https://ui.adsabs.harvard.edu/abs/2004Sci...306.1780P). [doi:10.1126/science.1105453](https://doi.org/10.1126/science.1105453). [PMID 15576621](https://pubmed.ncbi.nlm.nih.gov/15576621). [S2CID 37558595](https://api.semanticscholar.org/CorpusID:37558595). [Archived](https://web.archive.org/web/20160304043322/https://dipot.ulb.ac.be/dspace/bitstream/2013/18807/1/061PearcyScience04.pdf) 2016-03-04 at the Wayback Machine. Retrieved 2018-04-20.

1. Oxley, P. R.; Ji, L.; Fetter-Pruneda, I.; McKenzie, S. K.; Li, C.; Hu, H.; Zhang, G.; Kronauer, D. J. (2014). "The genome of the clonal raider ant Cerapachys biroi". *Curr. Biol.*. **24** (4): 451–8. [Bibcode:2014CBio...24..451O](https://ui.adsabs.harvard.edu/abs/2014CBio...24..451O). [doi:10.1016/j.cub.2014.01.018](https://doi.org/10.1016/j.cub.2014.01.018). [PMC 3961065](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3961065). [PMID 24508170](https://pubmed.ncbi.nlm.nih.gov/24508170)

1. Kellner, Katrin & Heinze, Jürgen (2011). "Mechanism of facultative parthenogenesis in the ant Platythyrea punctata". *Evolutionary Ecology*. **25** (1): 77–89. [Bibcode:2011EvEco..25...77K](https://ui.adsabs.harvard.edu/abs/2011EvEco..25...77K). [doi:10.1007/s10682-010-9382-5](https://doi.org/10.1007/s10682-010-9382-5). [S2CID 24645055](https://api.semanticscholar.org/CorpusID:24645055)

1. Rey, O.; Loiseau, A.; Facon, B.; Foucaud, J.; Orivel, J.; Cornuet, J. M.; Robert, S.; Dobigny, G.; Delabie, J. H.; Mariano Cdos, S.; Estoup, A. (2011). "Meiotic recombination dramatically decreased in thelytokous queens of the little fire ant and their sexually produced workers". *Mol. Biol. Evol.*. **28** (9): 2591–601. [doi:10.1093/molbev/msr082](https://doi.org/10.1093/molbev/msr082). [PMID 21459760](https://pubmed.ncbi.nlm.nih.gov/21459760)

1. Baudry, E.; Kryger, P.; Allsopp, M.; Koeniger, N.; Vautrin D.; Mougel F.; Cornuet JM.; Solignac M. (2004). "Whole-genome scan in the lytokous-laying workers of the Cape honeybee (Apis mellifera capensis): central fusion, reduced recombination rates and centromere mapping using half-tetrad analysis". *Genetics*. **167** (1): 243–252. [doi:10.1534/genetics.167.1.243](https://doi.org/10.1534/genetics.167.1.243). [PMC 1470879](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1470879). [PMID 15166151](https://pubmed.ncbi.nlm.nih.gov/15166151)

1. Haag-Liautard C, Vitikainen E, Keller L, Sundström L (2009). ["Fitness and the level of homozygosity in a social insect"](https://web.archive.org/web/20210227132228/http://doc.rero.ch/record/11248/files/haag_flh.pdf). *J. Evol. Biol.*. **22** (1): 134–142. [doi:10.1111/j.1420-9101.2008.01635.x](https://doi.org/10.1111/j.1420-9101.2008.01635.x). [PMID 19127611](https://pubmed.ncbi.nlm.nih.gov/19127611). [S2CID 19566175](https://api.semanticscholar.org/CorpusID:19566175). Archived from [the original](http://doc.rero.ch/record/11248/files/haag_flh.pdf) on 2021-02-27. Retrieved 2019-09-24.

1. Lehnert, Matthew S.; Kramer, Valerie R.; Rawlins, John E.; Verdecia, Vanessa; Daniels, Jaret C. (2017-07-10). "Jamaica's Critically Endangered Butterfly: A Review of the Biology and Conservation Status of the Homerus Swallowtail (Papilio (Pterourus) homerus Fabricius)". *Insects*. **8** (3): 68. [doi:10.3390/insects8030068](https://doi.org/10.3390/insects8030068). [PMC 5620688](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5620688). [PMID 28698508](https://pubmed.ncbi.nlm.nih.gov/28698508)

1. Baine, Q.; Looney, C.; Monckton, S. K.; Smith, D. R.; Schiff, N. M.; Goulet, H.; Redford, A. J. (April 2022). ["Biology and behavior"](https://idtools.org/sawfly/index.cfm?pageID=1844). *idtools.org*. [Archived](https://web.archive.org/web/20240215053439/https://idtools.org/sawfly/index.cfm?pageID=1844) February 15, 2024 at the Wayback Machine. Retrieved February 15, 2024.

1. Spiesman, Brian J. & Inouye, Brian D. (December 2013). "Habitat loss alters the architecture of plant–pollinator interaction networks". *Ecology*. **94** (12): 2688–2696. [Bibcode:2013Ecol...94.2688S](https://ui.adsabs.harvard.edu/abs/2013Ecol...94.2688S). [doi:10.1890/13-0977.1](https://doi.org/10.1890/13-0977.1). [ISSN 0012-9658](https://www.worldcat.org/issn/0012-9658). [PMID 24597216](https://pubmed.ncbi.nlm.nih.gov/24597216)

## External links

**General**

- [Hymenoptera Anatomy Ontology project](http://hymao.org/)
- [Hymenoptera Anatomy Glossary](http://glossary.hymao.org/)
- [Hymenoptera Forum](https://web.archive.org/web/20080609224812/http://www.forum.hymis.de/) German and International
- [Hymenoptera of North America – large format reference photographs, descriptions, taxonomy](http://www.cirrusimage.com/hymenoptera.htm)
- [International Society of Hymenopterists](http://www.hymenopterists.org/)
- [Bees, Wasps and Ants Recording Society](http://www.bwars.com/) (UK)
- [Ants Photo Gallery](http://www.lasius.narod.ru/) (RU)
- [Sphecos Forum for Aculeate Hymenoptera](https://web.archive.org/web/20060924070018/http://www.sel.barc.usda.gov/selhome/sphecos/sph30ttl.htm)
- [Hymenoptera images](http://morphbank.net/Browse/ByImage/index.php?keywords=&tsnKeywords=hymenoptera&spKeywords=&viewKeywords=whole+body&localityKeywords=&listField1=imageId&orderAsc1=DESC&listField2=&orderAsc2=ASC&listField3=&orderAsc3=ASC&numPerPage=20&goTo=&resetOffset=&activeSubmit=2) on MorphBank (a biological image database)
- [Order Hymenoptera](http://www.insectlifeforms.com/Orders564/ANTS_-_BEES_PLUS_WASPS_HYMENOPTERA_60502050_564.aspx) Insect Life Forms

**Systematics**

- [Hymenopteran Systematics](http://hymenoptera.ucr.edu)
- [Hymenoptera Online](https://web.archive.org/web/20101021045815/http://hol.osu.edu/) 1000+ images

**Regional Lists**

- [Insetos do Brasil](http://www.ufrrj.br/institutos/ib/ento/tomo11.pdf)
- [New Zealand Hymenoptera](http://www.landcareresearch.co.nz/research/biosystematics/invertebrates/hymenoptera/) [Archived](https://web.archive.org/web/20071113190650/http://www.landcareresearch.co.nz/research/biosystematics/invertebrates/hymenoptera/) 2007-11-13 at the Wayback Machine
- [Waspweb](http://www.waspweb.org/Afrotropical_wasps/index.htm) [Afrotropical](/source/Afrotropical) Hymenoptera Excellent images
- [checklist of Australian Hymenoptera](https://web.archive.org/web/20131109211836/http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/HYMENOPTERA)

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Adapted from the Wikipedia article [Hymenoptera](https://en.wikipedia.org/wiki/Hymenoptera) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Hymenoptera?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.