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A wasp is any
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. The wasps do not constitute a clade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade
Aculeata Aculeata is a subclade of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group cannot ...
can
sting Sting may refer to: * Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger * Irritating hairs or prickles of a stinging plant, or the plant itself Fictional characters and entities * Sting (Middle-earth ...
their prey. The most commonly known wasps, such as yellowjackets and hornets, are in the family Vespidae and are eusocial, living together in a nest with an egg-laying queen and non-reproducing workers. Eusociality is favoured by the unusual haplodiploid system of sex determination in Hymenoptera, as it makes sisters exceptionally closely related to each other. However, the majority of wasp species are solitary, with each adult female living and breeding independently. Females typically have an ovipositor for laying eggs in or near a food source for the larvae, though in the
Aculeata Aculeata is a subclade of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group cannot ...
the ovipositor is often modified instead into a
sting Sting may refer to: * Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger * Irritating hairs or prickles of a stinging plant, or the plant itself Fictional characters and entities * Sting (Middle-earth ...
used for defense or prey capture. Wasps play many ecological roles. Some are predators or pollinators, whether to feed themselves or to provision their nests. Many, notably the
cuckoo wasp Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic colors ...
s, are kleptoparasites, laying eggs in the nests of other wasps. Many of the solitary wasps are parasitoidal, meaning they lay eggs on or in other insects (any life stage from egg to adult) and often provision their own nests with such
hosts A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ...
. Unlike true parasites, the wasp larvae eventually kill their hosts. Solitary wasps parasitize almost every
pest insect Economic entomology is a field of entomology, which involves the study of insects that benefit or harm humans, domestic animals, and crops. Insects that cause losses are termed as pests. Some species can cause indirect damage by spreading diseases ...
, making wasps valuable in
horticulture Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
for biological pest control of species such as whitefly in tomatoes and other crops. Wasps first appeared in the fossil record in the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
, and diversified into many surviving superfamilies by the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
. They are a successful and diverse group of insects with tens of thousands of described species; wasps have spread to all parts of the world except for the polar regions. The largest social wasp is the Asian giant hornet, at up to in length; among the largest solitary wasps is a group of species known as tarantula hawks, along with the giant scoliid of Indonesia (''
Megascolia procer ''Megascolia procer'', the giant scoliid wasp, is a solitary wasp in the family Scoliidae found across the Oriental region. It is one of the largest wasps in the world, with a wingspan of 11.6 cm. Measurement scale on Figure 1. Descriptio ...
''). The smallest wasps are solitary parasitoid wasps in the family
Mymaridae The Mymaridae, commonly known as fairyflies or fairy wasps, are a family of chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. Fairyflies are very tiny insects ...
, including the world's smallest known insect, with a body length of only , and the smallest known flying insect, only long. Wasps have appeared in literature from Classical times, as the eponymous chorus of old men in Aristophanes' 422 BC comedy '' The Wasps'', and in
science fiction Science fiction (sometimes shortened to Sci-Fi or SF) is a genre of speculative fiction which typically deals with imaginative and futuristic concepts such as advanced science and technology, space exploration, time travel, parallel uni ...
from H. G. Wells's 1904 novel ''
The Food of the Gods and How It Came to Earth ''The Food of the Gods and How It Came to Earth'' is a science fiction novel by H. G. Wells, first published in 1904. Wells called it "a fantasia on the change of scale in human affairs. . . . I had hit upon he ideawhile working out ...
'', featuring giant wasps with three-inch-long stings. The name 'Wasp' has been used for many warships and other military equipment.


Taxonomy and phylogeny


Paraphyletic grouping

The wasps are a cosmopolitan paraphyletic grouping of hundreds of thousands of species, consisting of the narrow-waisted clade Apocrita without the ants and
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
s. The Hymenoptera also contain the somewhat wasplike but unwaisted Symphyta, the sawflies. The term ''wasp'' is sometimes used more narrowly for members of the Vespidae, which includes several eusocial wasp lineages, such as yellowjackets (the genera ''
Vespula ''Vespula'' is a small genus of social wasps, widely distributed in the Northern Hemisphere. Along with members of their sister genus ''Dolichovespula'', they are collectively known by the common name yellowjackets (or yellow jackets) in North ...
'' and ''
Dolichovespula ''Dolichovespula'' is a small genus of social wasps distributed widely throughout the Northern Hemisphere. The yellow and black members of the genus are known by the common name yellowjackets in North America, such as '' Dolichovespula norwegica' ...
''), hornets (genus ''Vespa''), and members of the subfamily Polistinae.


Fossils

Hymenoptera in the form of Symphyta ( Xyelidae) first appeared in the fossil record in the
Lower Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a ...
. Apocrita, wasps in the broad sense, appeared in the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
, and had diversified into many of the extant superfamilies by the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
; they appear to have evolved from the Symphyta. Fig wasps with modern anatomical features first appeared in the Lower Cretaceous of the Crato Formation in Brazil, some 65 million years before the first fig trees. The Vespidae include the extinct genus ''
Palaeovespa ''Palaeovespa'' is an extinct genus of wasp in the Vespidae subfamily Vespinae. The genus currently contains eight species, five from the Priabonian stage Florissant Formation in Colorado, United States two from the middle Eocene Baltic amber dep ...
'', seven species of which are known from the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
rocks of the Florissant fossil beds of Colorado and from fossilised
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than ...
in Europe. Also found in Baltic amber are crown wasps of the genus ''
Electrostephanus ''Electrostephanus'' is an extinct genus of crown wasp in the hymenopteran family Stephanidae, and is the only genus placed in the subfamily Electrostephaninae. The genus contains four described species, ''E. brevicornis'', ''E. neovenatus'', ...
''.


Diversity

Wasps are a diverse group, estimated at well over a hundred thousand described species around the world, and a great many more as yet undescribed. For example, almost every one of some 1000 species of tropical
fig trees ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species exten ...
has its own specific fig wasp ( Chalcidoidea) that has co-evolved with it and pollinates it. Many wasp species are parasitoids; the females deposit eggs on or in a host
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
on which the larvae then feed. Some larvae start off as parasitoids, but convert at a later stage to consuming the plant tissues that their host is feeding on. In other species, the eggs are laid directly into plant tissues and form galls, which protect the developing larvae from predators, but not necessarily from other parasitic wasps. In some species, the larvae are predatory themselves; the wasp eggs are deposited in clusters of eggs laid by other insects, and these are then consumed by the developing wasp larvae. The largest social wasp is the Asian giant hornet, at up to in length. The various tarantula hawk wasps are of a similar size and can overpower a spider many times its own weight, and move it to its burrow, with a sting that is excruciatingly painful to humans. The solitary giant scoliid, ''
Megascolia procer ''Megascolia procer'', the giant scoliid wasp, is a solitary wasp in the family Scoliidae found across the Oriental region. It is one of the largest wasps in the world, with a wingspan of 11.6 cm. Measurement scale on Figure 1. Descriptio ...
'', with a wingspan of 11.5 cm, has subspecies in Sumatra and
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
; it is a parasitoid of the Atlas beetle ''
Chalcosoma atlas The Atlas beetle (''Chalcosoma atlas'') is a very large species of beetle in the family Scarabaeidae, found in Southeast Asia. Males have three prominent horns. The species is named for Atlas, the giant of Greek mythology who supported the ski ...
''. The female giant ichneumon wasp ''
Megarhyssa macrurus ''Megarhyssa macrurus'', also known as the long-tailed giant ichneumonid wasp or long-tailed giant ichneumon wasp, is a species of large ichneumon wasp. It is a parasitoid, notable for its extremely long ovipositor which it uses to deposit an egg ...
'' is long including its very long but slender ovipositor which is used for boring into wood and inserting eggs. The smallest wasps are solitary parasitoid wasps in the family
Mymaridae The Mymaridae, commonly known as fairyflies or fairy wasps, are a family of chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. Fairyflies are very tiny insects ...
, including the world's smallest known insect, '' Dicopomorpha echmepterygis'' (139 micrometres long) and ''
Kikiki huna ''Kikiki'' is a genus of fairyfly wasps containing a single species, ''Kikiki huna'', known from Hawaii, Costa Rica, Nagarcoil and Trinidad. At (150 μm), it is the smallest flying insect known . It is a close relative of wasps in the genus '' ...
'' with a body length of only 158 micrometres, the smallest known flying insect. There are estimated to be 100,000 species of ichneumonoid wasps in the families
Braconidae The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
and Ichneumonidae. These are almost exclusively parasitoids, mostly using other insects as hosts. Another family, the Pompilidae, is a specialist parasitoid of spiders. Some wasps are even parasitoids of parasitoids; the eggs of '' Euceros'' are laid beside lepidopteran larvae and the wasp larvae feed temporarily on their haemolymph, but if a parasitoid emerges from the host, the hyperparasites continue their life cycle inside the parasitoid. Parasitoids maintain their extreme diversity through narrow specialism. In Peru, 18 wasp species were found living on 14 fly species in only two species of ''
Gurania ''Gurania'' is a genus of flowering plants belonging to the family Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important ...
'' climbing squash. File:Megascolia procer MHNT dos.jpg, ''
Megascolia procer ''Megascolia procer'', the giant scoliid wasp, is a solitary wasp in the family Scoliidae found across the Oriental region. It is one of the largest wasps in the world, with a wingspan of 11.6 cm. Measurement scale on Figure 1. Descriptio ...
'', a giant solitary species from
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
in the Scoliidae. This specimen's length is 77mm and its wingspan is 115mm. Measurement scale on Figure 1. File:Ichneumon_wasp_(Megarhyssa_macrurus_lunato)_(7686081848).jpg, ''
Megarhyssa macrurus ''Megarhyssa macrurus'', also known as the long-tailed giant ichneumonid wasp or long-tailed giant ichneumon wasp, is a species of large ichneumon wasp. It is a parasitoid, notable for its extremely long ovipositor which it uses to deposit an egg ...
'', a parasitoid. The body of a female is 50mm long, with a c. 100mm ovipositor File:Wasp with Orange-kneed tarantula.JPG, Tarantula hawk wasp dragging the tarantula ''
Megaphobema mesomelas ''Megaphobema'' is a genus of tarantulas that was first described by Reginald Innes Pocock in 1901. They look similar to members of '' Pamphobeteus'' except for its legs; the third and fourth pairs of legs are much larger and stronger than the fir ...
'' to her burrow; it has the most painful sting of any wasp.


Sociality


Social wasps

Of the dozens of extant wasp families, only the family Vespidae contains social species, primarily in the subfamilies Vespinae and Polistinae. With their powerful stings and conspicuous
warning coloration Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...
, often in black and yellow, social wasps are frequent models for Batesian mimicry by non-stinging insects, and are themselves involved in mutually beneficial Müllerian mimicry of other distasteful insects including bees and other wasps. All species of social wasps construct their nests using some form of plant fiber (mostly wood pulp) as the primary material, though this can be supplemented with mud, plant secretions (e.g., resin), and secretions from the wasps themselves; multiple fibrous brood cells are constructed, arranged in a honeycombed pattern, and often surrounded by a larger protective envelope. Wood fibres are gathered from weathered wood, softened by chewing and mixing with saliva. The placement of nests varies from group to group; yellow jackets such as ''
Dolichovespula media The median wasp (''Dolichovespula media'') is a species of social wasp of the family Vespidae found throughout Europe and Asia. It builds aerial paper nests often in shrubs or trees, and occasionally under the eaves of buildings. It is most comm ...
'' and '' D. sylvestris'' prefer to nest in trees and shrubs; ''
Protopolybia exigua ''Protopolybia exigua'' is a species of vespid wasp found in South America and Southern Brazil. These neotropical wasps, of the tribe Epiponini, form large colonies with multiple queens per colony. ''P. exigua'' are small wasps that find nourish ...
''attaches its nests on the underside of leaves and branches; '' Polistes erythrocephalus'' chooses sites close to a water source. Other wasps, like ''
Agelaia multipicta ''Agelaia multipicta'' is a swarm-founding, highly eusocial wasp that lives in Mexico, Argentina, Trinidad and southern Brazil. It nests in natural cavities such as hollow trees and aggressively defends the nest from ants, who are brood predators ...
'' and ''
Vespula germanica ''Vespula germanica'', the European wasp, German wasp, or German yellowjacket, is a species of wasp found in much of the Northern Hemisphere, native to Europe, Northern Africa, and temperate Asia. It has spread and become well-established in m ...
,'' like to nest in cavities that include holes in the ground, spaces under homes, wall cavities or in lofts. While most species of wasps have nests with multiple combs, some species, such as '' Apoica flavissima'', only have one comb. The length of the reproductive cycle depends on latitude; ''Polistes erythrocephalus'', for example, has a much longer (up to 3 months longer) cycle in temperate regions.


Solitary wasps

The vast majority of wasp species are solitary insects. Having mated, the adult female forages alone and if it builds a nest, does so for the benefit of its own offspring. Some solitary wasps nest in small groups alongside others of their species, but each is involved in caring for its own offspring (except for such actions as stealing other wasps’ prey or laying in other wasp's nests). There are some species of solitary wasp that build communal nests, each insect having its own cell and providing food for its own offspring, but these wasps do not adopt the division of labour and the complex behavioural patterns adopted by eusocial species. Adult solitary wasps spend most of their time in preparing their nests and foraging for food for their young, mostly insects or spiders. Their nesting habits are more diverse than those of social wasps. Many species dig burrows in the ground.
Mud dauber Mud dauber (or "mud wasp" or "dirt dauber") is a name commonly applied to a number of wasps from either the family Sphecidae or Crabronidae which build their nests from mud; this excludes members of the family Vespidae (especially the subfamily E ...
s and pollen wasps construct mud cells in sheltered places. Potter wasps similarly build vase-like nests from mud, often with multiple cells, attached to the twigs of trees or against walls. Predatory wasp species normally subdue their prey by stinging it, and then either lay their eggs on it, leaving it in place, or carry it back to their nest where an egg may be laid on the prey item and the nest sealed, or several smaller prey items may be deposited to feed a single developing larva. Apart from providing food for their offspring, no further maternal care is given. Members of the family
Chrysididae Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic color ...
, the cuckoo wasps, are kleptoparasites and lay their eggs in the nests of unrelated host species.


Biology


Anatomy

Like all insects, wasps have a hard exoskeleton which protects their three main body parts, the head, the mesosoma (including the thorax and the first segment of the abdomen) and the metasoma. There is a narrow waist, the petiole, joining the first and second segments of the abdomen. The two pairs of membranous wings are held together by small hooks and the forewings are larger than the hind ones; in some species, the females have no wings. In females there is usually a rigid ovipositor which may be modified for injecting venom, piercing or sawing. It either extends freely or can be retracted, and may be developed into a stinger for both defence and for paralysing prey. In addition to their large compound eyes, wasps have several simple eyes known as ocelli, which are typically arranged in a triangle just forward of the
vertex Vertex, vertices or vertexes may refer to: Science and technology Mathematics and computer science *Vertex (geometry), a point where two or more curves, lines, or edges meet *Vertex (computer graphics), a data structure that describes the position ...
of the head. Wasps possess mandibles adapted for biting and cutting, like those of many other insects, such as grasshoppers, but their other mouthparts are formed into a suctorial proboscis, which enables them to drink nectar. The larvae of wasps resemble
maggot A maggot is the larva of a fly (order Diptera); it is applied in particular to the larvae of Brachycera flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and crane flies. En ...
s, and are adapted for life in a protected environment; this may be the body of a host organism or a cell in a nest, where the larva either eats the provisions left for it or, in social species, is fed by the adults. Such larvae have soft bodies with no limbs, and have a blind gut (presumably so that they do not foul their cell).


Diet

Adult solitary wasps mainly feed on nectar, but the majority of their time is taken up by foraging for food for their carnivorous young, mostly insects or spiders. Apart from providing food for their larval offspring, no maternal care is given. Some wasp species provide food for the young repeatedly during their development (
progressive provisioning Progressive provisioning is a term used in entomology to refer to a form of parental behavior in which an adult (most commonly a hymenopteran such as a bee or wasp) feeds its larvae directly after they have hatched, feeding each larva repeatedly un ...
). Others, such as potter wasps (Eumeninae) and sand wasps ('' Ammophila'', Sphecidae), repeatedly build nests which they stock with a supply of immobilised prey such as one large caterpillar, laying a single egg in or on its body, and then sealing up the entrance ( mass provisioning). Predatory and parasitoidal wasps subdue their prey by stinging it. They hunt a wide variety of prey, mainly other insects (including other Hymenoptera), both larvae and adults. The Pompilidae specialize in catching spiders to provision their nests. Some social wasps are omnivorous, feeding on fallen fruit, nectar, and carrion such as dead insects. Adult male wasps sometimes visit flowers to obtain nectar. Some wasps, such as ''
Polistes fuscatus ''Polistes fuscatus'', whose common name is the dark or northern paper wasp, is widely found in eastern North America, from southern Canada through the southern United States. It often nests around human development. However, it greatly prefers ...
'', commonly return to locations where they previously found prey to forage. In many social species, the larvae exude copious amounts of salivary secretions that are avidly consumed by the adults. These include both
sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or do ...
s and
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
s, and may provide essential protein-building nutrients that are otherwise unavailable to the adults (who cannot digest proteins).


Sex determination

In wasps, as in other Hymenoptera, sex is determined by a haplodiploid system, which means that females are unusually closely related to their sisters, enabling kin selection to favour the evolution of eusocial behaviour. Females are diploid, meaning that they have 2n
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
s and develop from fertilized eggs. Males, called drones, have a haploid (n) number of chromosomes and develop from an unfertilized egg. Wasps store sperm inside their body and control its release for each individual egg as it is laid; if a female wishes to produce a male egg, she simply lays the egg without fertilizing it. Therefore, under most conditions in most species, wasps have complete voluntary control over the sex of their offspring. Experimental infection of ''
Muscidifurax uniraptor ''Muscidifurax uniraptor'' is a species of wasp (the taxonomic order Hymenoptera) in the family Pteromalidae. The species does not currently have a common name. ''M. uniraptor'' is a pupal parasitoid of synanthropic filth-breeding Diptera and is ...
'' with the
bacterium Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
'' Wolbachia'' induced thelytokous reproduction and an inability to produce fertile, viable male offspring.


Inbreeding avoidance

Females of the solitary wasp parasitoid ''Venturia canescens'' can avoid mating with their brothers through kin recognition. In experimental comparisons, the probability that a female will mate with an unrelated male was about twice as high as the chance of her mating with brothers. Female wasps appear to recognize siblings on the basis of a chemical signature carried or emitted by males. Sibling-mating avoidance reduces
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
that is largely due to the expression of homozygous deleterious recessive mutations.


Ecology


As pollinators

While the vast majority of wasps play no role in pollination, a few species can effectively transport pollen and pollinate several plant species. Since wasps generally do not have a fur-like covering of soft hairs and a special body part for pollen storage (
pollen basket The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive. Other species of bees have scopae instead. Ety ...
) as some bees do, pollen does not stick to them well. However it has been shown that even without hairs, several wasp species are able to effectively transport pollen, therefore contributing for potential pollination of several plant species. Pollen wasps in the subfamily Masarinae gather nectar and pollen in a crop inside their bodies, rather than on body hairs like bees, and pollinate flowers of '' Penstemon'' and the water leaf family, Hydrophyllaceae. The Agaonidae ( fig wasps) are the only pollinators of nearly 1000 species of figs, and thus are crucial to the survival of their host plants. Since the wasps are equally dependent on their fig trees for survival, the coevolved relationship is fully mutualistic.


As parasitoids

Most solitary wasps are parasitoids. As adults, those that do feed typically only take nectar from flowers. Parasitoid wasps are extremely diverse in habits, many laying their eggs in inert stages of their host ( egg or pupa), sometimes paralysing their prey by injecting it with venom through their ovipositor. They then insert one or more eggs into the host or deposit them upon the outside of the host. The host remains alive until the parasitoid larvae pupate or emerge as adults. The Ichneumonidae are specialized parasitoids, often of Lepidoptera larvae deeply buried in plant tissues, which may be woody. For this purpose, they have exceptionally long ovipositors; they detect their hosts by smell and vibration. Some of the largest species, including '' Rhyssa persuasoria'' and ''
Megarhyssa macrurus ''Megarhyssa macrurus'', also known as the long-tailed giant ichneumonid wasp or long-tailed giant ichneumon wasp, is a species of large ichneumon wasp. It is a parasitoid, notable for its extremely long ovipositor which it uses to deposit an egg ...
'', parasitise horntails, large sawflies whose adult females also have impressively long ovipositors. Some parasitic species have a mutualistic relationship with a polydnavirus that weakens the host's
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
and replicates in the oviduct of the female wasp. One family of chalcidoid wasps, the Eucharitidae, has specialized as parasitoids of ants, most species hosted by one genus of ant. Eucharitids are among the few parasitoids that have been able to overcome ants' effective defences against parasitoids.


As parasites

Many species of wasp, including especially the cuckoo or jewel wasps (
Chrysididae Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic color ...
), are kleptoparasites, laying their eggs in the nests of other wasp species to exploit their parental care. Most such species attack hosts that provide provisions for their immature stages (such as paralyzed prey items), and they either consume the provisions intended for the host larva, or wait for the host to develop and then consume it before it reaches adulthood. An example of a true brood parasite is the paper wasp ''
Polistes sulcifer ''Polistes semenowi'' is a species of paper wasp in the genus ''Polistes ''that is found in southeastern and southern central Europe, as well as central Asia, and was until 2017 erroneously known by the name ''Polistes sulcifer'', while a differe ...
'', which lays its eggs in the nests of other paper wasps (specifically ''
Polistes dominula The European paper wasp (''Polistes dominula'') is one of the most common and well-known species of social wasps in the genus '' Polistes''. Its diet is more diverse than those of most ''Polistes'' species—many genera of insects versus mainly ...
''), and whose larvae are then fed directly by the host. Sand wasps '' Ammophila'' often save time and energy by parasitising the nests of other females of their own species, either kleptoparasitically stealing prey, or as brood parasites, removing the other female's egg from the prey and laying their own in its place. According to Emery's rule, social parasites, especially among insects, tend to parasitise species or genera to which they are closely related. For example, the social wasp ''
Dolichovespula adulterina ''Dolichovespula adulterina'' is a species of parasitic social wasp found in the Palearctic region. ''D. adulterina'' feeds on a variety of foods, including insects, spiders, arthropods, meat, molluscs, fruit, nectar, and larval secretions. ''D ...
'' parasitises other members of its genus such as '' D. norwegica'' and '' D. arenaria''.


As predators

Many wasp lineages, including those in the families Vespidae, Crabronidae, Sphecidae, and Pompilidae, attack and sting prey items that they use as food for their larvae; while Vespidae usually macerate their prey and feed the resulting bits directly to their brood, most predatory wasps paralyze their prey and lay eggs directly upon the bodies, and the wasp larvae consume them. Apart from collecting prey items to provision their young, many wasps are also opportunistic feeders, and will suck the body fluids of their prey. Although vespid mandibles are adapted for chewing and they appear to be feeding on the organism, they are often merely macerating it into submission. The impact of the predation of wasps on economic
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
is difficult to establish. The roughly 140 species of
beewolf Beewolves (genus ''Philanthus''), also known as bee-hunters or bee-killer wasps, are solitary, predatory wasps, most of which prey on bees, hence their common name. The adult females dig tunnels in the ground for nesting, while the territorial m ...
( Philanthinae) hunt bees, including honeybees, to provision their nests; the adults feed on nectar and pollen.


As models for mimics

With their powerful stings and conspicuous
warning coloration Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...
, social wasps are the models for many species of mimic. Two common cases are Batesian mimicry, where the mimic is harmless and is essentially bluffing, and Müllerian mimicry, where the mimic is also distasteful, and the mimicry can be considered mutual. Batesian mimics of wasps include many species of hoverfly and the wasp beetle. Many species of wasp are involved in Müllerian mimicry, as are many species of
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
.


As prey

While wasp stings deter many potential predators, bee-eaters (in the bird family Meropidae) specialise in eating stinging insects, making aerial sallies from a perch to catch them, and removing the venom from the stinger by repeatedly brushing the prey firmly against a hard object, such as a twig. The honey buzzard attacks the nests of social hymenopterans, eating wasp larvae; it is the only known predator of the dangerous Asian giant hornet or "yak-killer" (''
Vespa mandarinia The Asian giant hornet (''Vespa mandarinia'') or northern giant hornet, including the color form referred to as the Japanese giant hornet, is the world's largest hornet. It is native to temperate and tropical East Asia, South Asia, Mainland Sout ...
''). Likewise, roadrunners are the only real predators of tarantula hawk wasps. File:CSIRO ScienceImage 11188 Fig wasp.jpg, Minute pollinating fig wasps, '' Pleistodontes'': the trees and wasps have coevolved and are mutualistic. File:Latina rugosa planidia.png, '' Latina rugosa'' planidia (arrows, magnified) attached to an ant larva; the Eucharitidae are among the few parasitoids able to overcome the strong defences of ants. File:Goudwesp.jpg, The
Chrysididae Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic color ...
, such as this ''
Hedychrum rutilans ''Hedychrum rutilans'' is a species of cuckoo wasps (insects in the family Chrysididae). The species occurs primarily in Austria, Italy, Bulgaria, Greece, France, Poland, Portugal, Spain, Switzerland and in North Africa.Agnoli, G. L.; Rosa, P ...
'', are known as cuckoo or jewel wasps for their parasitic behaviour and metallic iridescence. File:Bee wolf.jpg,
European beewolf The European beewolf (''Philanthus triangulum''), also known as the bee-killer wasp or the bee-eating philanthus (from the now obsolete synonym ''Philanthus apivorus''), is a solitary wasp that lives in the Western Palearctic and Afrotropics. Alt ...
''Philanthus triangulum'' provisioning her nest with a honeybee File:Clytus arietis (Linné, 1758) (3989861203).jpg, Wasp beetle ''Clytus arietis'' is a Batesian mimic of wasps. File:Pair of Merops apiaster feeding detail.jpg, Bee-eaters such as '' Merops apiaster'' specialise in feeding on bees and wasps.


Relationship with humans


As pests

Social wasps are considered pests when they become excessively common, or nest close to buildings. People are most often stung in late summer and early autumn, when wasp colonies stop breeding new workers; the existing workers search for sugary foods and are more likely to come into contact with humans. Wasp nests made in or near houses, such as in roof spaces, can present a danger as the wasps may sting if people come close to them. Stings are usually painful rather than dangerous, but in rare cases, people may suffer life-threatening anaphylactic shock.


In horticulture

Some species of parasitic wasp, especially in groups such as
Aphelinidae The Aphelinidae are a moderate-sized family of tiny parasitic wasps, with about 1100 described species in some 28 genera. These minute insects are challenging to study, as they deteriorate rapidly after death unless extreme care is taken (e.g., ...
,
Braconidae The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
,
Mymaridae The Mymaridae, commonly known as fairyflies or fairy wasps, are a family of chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. Fairyflies are very tiny insects ...
, and
Trichogrammatidae The Trichogrammatidae are a family of tiny wasps in the Chalcidoidea that include some of the smallest of all insects, with most species having adults less than 1 mm in length, with species of '' Megaphragma'' having an adult body length l ...
, are exploited commercially to provide biological control of
insect pest Economic entomology is a field of entomology, which involves the study of insects that benefit or harm humans, domestic animals, and crops. Insects that cause losses are termed as pests. Some species can cause indirect damage by spreading diseases ...
s. One of the first species to be used was '' Encarsia formosa'', a parasitoid of a range of species of whitefly. It entered commercial use in the 1920s in Europe, was overtaken by chemical pesticides in the 1940s, and again received interest from the 1970s. ''Encarsia'' is used especially in greenhouses to control whitefly pests of tomato and cucumber, and to a lesser extent of aubergine (eggplant), flowers such as marigold, and strawberry. Several species of parasitic wasp are natural predators of aphids and can help to control them. For instance, ''Aphidius matricariae'' is used to control the peach-potato aphid. File:Encarsia formosa, an endoparasitic wasp, is used for whitefly control.jpg, '' Encarsia formosa'', a parasitoid, is sold commercially for biological control of whitefly, an
insect pest Economic entomology is a field of entomology, which involves the study of insects that benefit or harm humans, domestic animals, and crops. Insects that cause losses are termed as pests. Some species can cause indirect damage by spreading diseases ...
of tomato and other horticultural crops. File:Tomate Blatt Eier Weiße Fliege parasitiert.jpg, Tomato leaf covered with nymphs of whitefly parasitised by ''Encarsia formosa''


In sport

Wasps RFC is an English professional
rugby union Rugby union, commonly known simply as rugby, is a close-contact team sport that originated at Rugby School in the first half of the 19th century. One of the two codes of rugby football, it is based on running with the ball in hand. In it ...
team originally based in London but now playing in Coventry; the name dates from 1867 at a time when names of insects were fashionable for clubs. The club's first kit is black with yellow stripes. The club has an amateur side called Wasps FC. Among the other clubs bearing the name are a basketball club in Wantirna, Australia, and
Alloa Athletic F.C. Alloa Athletic Football Club is a Scottish association football semi-professional club based in the town of Alloa, Clackmannanshire. Formed as Alloa in 1880, the football club shortly changed its name to Alloa Association, and then to Alloa At ...
, a football club in
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to ...
.


In fashion

Wasps have been modelled in
jewellery Jewellery ( UK) or jewelry ( U.S.) consists of decorative items worn for personal adornment, such as brooches, rings, necklaces, earrings, pendants, bracelets, and cufflinks. Jewellery may be attached to the body or the clothes. From a w ...
since at least the nineteenth century, when
diamond Diamond is a solid form of the element carbon with its atoms arranged in a crystal structure called diamond cubic. Another solid form of carbon known as graphite is the chemically stable form of carbon at room temperature and pressure, b ...
and emerald wasp brooches were made in
gold Gold is a chemical element with the symbol Au (from la, aurum) and atomic number 79. This makes it one of the higher atomic number elements that occur naturally. It is a bright, slightly orange-yellow, dense, soft, malleable, and ductile ...
and
silver Silver is a chemical element with the symbol Ag (from the Latin ', derived from the Proto-Indo-European ''h₂erǵ'': "shiny" or "white") and atomic number 47. A soft, white, lustrous transition metal, it exhibits the highest electrical ...
settings. A fashion for wasp waisted female silhouettes with sharply cinched waistlines emphasizing the wearer's hips and bust arose repeatedly in the nineteenth and twentieth centuries.


In literature

The Ancient Greek playwright Aristophanes wrote the
comedy Comedy is a genre of fiction that consists of discourses or works intended to be humorous or amusing by inducing laughter, especially in theatre, film, stand-up comedy, television, radio, books, or any other entertainment medium. The term o ...
play Σφῆκες (''Sphēkes''), '' The Wasps'', first put on in 422 BC. The "wasps" are the chorus of old jurors. H. G. Wells made use of giant wasps in his novel ''
The Food of the Gods and How It Came to Earth ''The Food of the Gods and How It Came to Earth'' is a science fiction novel by H. G. Wells, first published in 1904. Wells called it "a fantasia on the change of scale in human affairs. . . . I had hit upon he ideawhile working out ...
'' (1904): '' Wasp'' (1957) is a
science fiction Science fiction (sometimes shortened to Sci-Fi or SF) is a genre of speculative fiction which typically deals with imaginative and futuristic concepts such as advanced science and technology, space exploration, time travel, parallel uni ...
book by the English writer Eric Frank Russell; it is generally considered Russell's best novel. In Stieg Larsson's book ''
The Girl Who Played with Fire ''The Girl Who Played with Fire'' ( sv, Flickan som lekte med elden) is the second novel in the best-selling ''Millennium'' series by Swedish writer Stieg Larsson. It was published posthumously in Swedish in 2006 and in English in January 2009. ...
'' (2006) and its film adaptation, Lisbeth Salander has adopted her kickboxing ringname, "The Wasp", as her hacker handle and has a wasp tattoo on her neck, indicating her high status among hackers, unlike her real world situation, and that like a small but painfully stinging wasp, she could be dangerous. Parasitoidal wasps played an indirect role in the nineteenth-century evolution debate. The Ichneumonidae contributed to
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
's doubts about the nature and existence of a well-meaning and all-powerful Creator. In an 1860 letter to the American naturalist Asa Gray, Darwin wrote:


In military names

With its powerful sting and familiar appearance, the wasp has given its name to many ships, aircraft and military vehicles. Nine ships and one shore establishment of the
Royal Navy The Royal Navy (RN) is the United Kingdom's naval warfare force. Although warships were used by English and Scottish kings from the early medieval period, the first major maritime engagements were fought in the Hundred Years' War against Fr ...
have been named , the first an 8-gun sloop launched in 1749. Eleven ships of the
United States Navy The United States Navy (USN) is the maritime service branch of the United States Armed Forces and one of the eight uniformed services of the United States. It is the largest and most powerful navy in the world, with the estimated tonnage ...
have similarly borne the name , the first a merchant schooner acquired by the Continental Navy in 1775. The eighth of these, an aircraft carrier, gained two Second World War battle stars, prompting
Winston Churchill Sir Winston Leonard Spencer Churchill (30 November 187424 January 1965) was a British statesman, soldier, and writer who served as Prime Minister of the United Kingdom twice, from 1940 to 1945 during the Second World War, and again from ...
to remark "Who said a Wasp couldn't sting twice?" In the Second World War, a German self-propelled howitzer was named Wespe, while the British developed the Wasp flamethrower from the
Bren Gun Carrier The Universal Carrier, also known as the Bren Gun Carrier and sometimes simply the Bren Carrier from the light machine gun armament, is a common name describing a family of light armoured tracked vehicles built by Vickers-Armstrongs and other ...
. In aerospace, the Westland Wasp was a military helicopter developed in England in 1958 and used by the Royal Navy and other navies. The AeroVironment Wasp III is a
miniature UAV A miniature UAV, small UAV (SUAV), or drone is an unmanned aerial vehicle small enough to be man-portable. Smallest UAVs are called micro air vehicle. Miniature UAVs range from micro air vehicles (MAVs) that can be carried by an infantryman, t ...
developed for
United States Air Force The United States Air Force (USAF) is the air service branch of the United States Armed Forces, and is one of the eight uniformed services of the United States. Originally created on 1 August 1907, as a part of the United States Army Si ...
special operations.


See also

* Bee and wasp stings *
Characteristics of common wasps and bees While observers can easily confuse common wasps and bees at a distance or without close observation, there are many different characteristics of large bee Bees are winged insects closely related to wasps and ants, known for their roles in p ...
* Schmidt sting pain index


Notes


References


Sources

*


External links


Differences between Bees and Wasps

Natural History Museum – Wasps: If you can't love them, at least admire them


– Insect bites and stings

* {{Authority control Biological pest control wasps Extant Jurassic first appearances Insects in culture Paraphyletic groups Pest insects