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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 pairs ...
of the narrow-waisted suborder
Apocrita Apocrita is a suborder of insects in the order Hymenoptera. It includes wasps, bees, and ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from Symphyta by the narrow "waist" ( petiole) formed ...
of the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
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 monophyly, monophyletic lineage within the ...
nor an
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
; this excludes the broad-waisted
sawflies Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
(Symphyta), which look somewhat like wasps, but are in a separate suborder. The wasps do not constitute a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, 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-eart ...
their prey. The most commonly known wasps, such as
yellowjacket Yellowjacket or yellowjacket is the common name in North America for predatory social wasps of the genus, genera ''Vespula'' and ''Dolichovespula''. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of ...
s and
hornet Hornets (insects in the genus ''Vespa'') are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to in length. They are distinguished from other vespine wasps by th ...
s, are in the family
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
and are
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
, living together in a nest with an egg-laying queen and non-reproducing workers. Eusociality is favoured by the unusual
haplodiploid Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...
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 The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
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-eart ...
used for defense or prey capture. Wasps play many ecological roles. Some are
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s 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
kleptoparasite Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when fo ...
s, 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 Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
of species such as
whitefly Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described. Description and taxonomy The A ...
in
tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
es and other crops. Wasps first appeared in the fossil record in the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), 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 J ...
, 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 th ...
. 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 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 ...
, at up to in length; among the largest solitary wasps is a group of species known as
tarantula hawk A tarantula hawk is a spider wasp (Pompilidae) that preys on tarantulas. Tarantula hawks belong to any of the many species in the genera ''Pepsis'' and ''Hemipepsis.'' They are one of the largest parasitoid wasps, using their sting to paralyze the ...
s, 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 In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
wasps in the family
Mymaridae The Mymaridae, commonly known as fairyflies or fairy wasps, are a family (biology), family of chalcid wasp, chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. F ...
, 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 Classical antiquity (also the classical era, classical period or classical age) is the period of cultural history between the 8th century BC and the 5th century AD centred on the Mediterranean Sea, comprising the interlocking civilizations of ...
, as the eponymous chorus of old men in
Aristophanes Aristophanes (; grc, Ἀριστοφάνης, ; c. 446 – c. 386 BC), son of Philippus, of the deme In Ancient Greece, a deme or ( grc, δῆμος, plural: demoi, δημοι) was a suburb or a subdivision of Athens and other city-states ...
' 422 BC comedy ''
The Wasps ''The Wasps'' ( grc-x-classical, Σφῆκες, translit=Sphēkes) is the fourth in chronological order of the eleven surviving plays by Aristophanes. It was produced at the Lenaia festival in 422 BC, during Athens' short-lived respite from the ...
'', 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 unive ...
from
H. G. Wells Herbert George Wells"Wells, H. G."
Revised 18 May 2015. ''
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 In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
grouping of hundreds of thousands of species, consisting of the narrow-waisted
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Apocrita Apocrita is a suborder of insects in the order Hymenoptera. It includes wasps, bees, and ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from Symphyta by the narrow "waist" ( petiole) formed ...
without the
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s 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 monophyly, monophyletic lineage within the ...
s. The Hymenoptera also contain the somewhat wasplike but unwaisted
Symphyta Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
, the sawflies. The term ''wasp'' is sometimes used more narrowly for members of the
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
, which includes several
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
wasp lineages, such as
yellowjacket Yellowjacket or yellowjacket is the common name in North America for predatory social wasps of the genus, genera ''Vespula'' and ''Dolichovespula''. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of ...
s (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'' ...
''),
hornet Hornets (insects in the genus ''Vespa'') are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to in length. They are distinguished from other vespine wasps by th ...
s (genus ''Vespa''), and members of the subfamily
Polistinae The Polistinae is a subfamily of eusocial wasps belonging to the Family (biology), family Vespidae. They are closely related to the more familiar wasps (“yellowjackets” as they are called in North America) and true hornets of the subfamily V ...
.


Fossils

Hymenoptera in the form of Symphyta (
Xyelidae The Xyelidae are a comparatively species-poor family of sawflies comprising about 80 Extant taxon, extant species in five genera worldwide and is the only family in the superfamily Xyeloidea. The fossil record of the family is extensive, comprisi ...
) 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 un ...
. Apocrita, wasps in the broad sense, appeared in the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), 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 J ...
, 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 th ...
; they appear to have evolved from the Symphyta.
Fig wasps Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while the ...
with modern anatomical features first appeared in the
Lower Cretaceous Lower may refer to: *Lower (surname) *Lower Township, New Jersey *Lower Receiver (firearms) *Lower Wick Lower Wick is a small hamlet located in the county of Gloucestershire, England. It is situated about five miles south west of Dursley, eight ...
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 (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
rocks of the
Florissant fossil beds The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones and ...
of
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
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 1 ...
in Europe. Also found in Baltic amber are crown wasps of the genus '' Electrostephanus''.


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 (biology), family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few spe ...
has its own specific
fig wasp Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while th ...
(
Chalcidoidea Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, me ...
) 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 Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
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
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
s, 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 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 ...
, 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 Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
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 List ...
; it is a
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
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 skies ...
''. 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 e ...
'' is long including its very long but slender
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
which is used for boring into wood and inserting eggs. The smallest wasps are solitary
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
wasps in the family
Mymaridae The Mymaridae, commonly known as fairyflies or fairy wasps, are a family (biology), family of chalcid wasp, chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. F ...
, including the world's smallest known insect, ''
Dicopomorpha echmepterygis ''Dicopomorpha echmepterygis'' is the smallest known insect and a species of parasitoid wasp in the family Mymaridae, which exhibits strong sexual dimorphism. The males are blind, apterous, and their body length is only 40% that of females. With ...
'' (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 ''T ...
'' 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 The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family (biology), family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 2 ...
. These are almost exclusively parasitoids, mostly using other insects as hosts. Another family, the
Pompilidae Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group-ne ...
, is a specialist parasitoid of spiders. Some wasps are even parasitoids of parasitoids; the eggs of '' Euceros'' are laid beside
lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
n larvae and the wasp larvae feed temporarily on their
haemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
, but if a parasitoid emerges from the host, the
hyperparasite A hyperparasite, also known as a metaparasite, is a parasite whose host, often an insect, is also a parasite, often specifically a parasitoid. Hyperparasites are found mainly among the wasp-waisted Apocrita within the Hymenoptera, and in two othe ...
s 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 t ...
'' 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 List ...
in the
Scoliidae The Scoliidae, the scoliid wasps, are a family of about 560 species found worldwide. They tend to be black, often marked with yellow or orange, and their wing tips are distinctively corrugated. Males are more slender and elongated than females, wi ...
. 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 e ...
'', a parasitoid. The body of a female is 50mm long, with a c. 100mm ovipositor File:Wasp with Orange-kneed tarantula.JPG,
Tarantula hawk A tarantula hawk is a spider wasp (Pompilidae) that preys on tarantulas. Tarantula hawks belong to any of the many species in the genera ''Pepsis'' and ''Hemipepsis.'' They are one of the largest parasitoid wasps, using their sting to paralyze the ...
wasp dragging the
tarantula Tarantulas comprise a group of large and often hairy spiders of the family Theraphosidae. , 1,040 species have been identified, with 156 genera. The term "tarantula" is usually used to describe members of the family Theraphosidae, although ...
''
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 The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
contains social species, primarily in the subfamilies
Vespinae The subfamily Vespinae contains the largest and best-known eusocial wasps, including true hornets (the genus ''Vespa''), and the "yellowjackets" (genera ''Dolichovespula'' and ''Vespula''). The remaining genus, ''Provespa'', is a small, poorly ...
and
Polistinae The Polistinae is a subfamily of eusocial wasps belonging to the Family (biology), family Vespidae. They are closely related to the more familiar wasps (“yellowjackets” as they are called in North America) and true hornets of the subfamily V ...
. 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 or ...
, often in black and yellow, social wasps are frequent models for
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on bu ...
by non-stinging insects, and are themselves involved in mutually beneficial
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian ...
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 In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
), 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 Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can be ...
. 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 nourishm ...
''attaches its nests on the underside of leaves and branches; ''
Polistes erythrocephalus ''Polistes erythrocephalus'' is a species of paper wasp in the subfamily Polistinae of family Vespidae found in Central and South America. ''P. erythrocephalus'' is a eusocial wasp, meaning that it possesses both reproductive and non-reproductive ...
'' 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 ma ...
,'' 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 ''Apoica flavissima'' is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens ...
'', only have one comb. The length of the reproductive cycle depends on
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pol ...
; ''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 Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
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 Eu ...
s and
pollen wasp Pollen wasps, the Masarinae, are unusual wasps that are typically treated as a subfamily of Vespidae, but have in the past sometimes been recognized as a separate family, "Masaridae", which also included the subfamilies Euparagiinae and Gayellina ...
s construct mud cells in sheltered places.
Potter wasp Potter wasps (or mason wasps), the Eumeninae, are a cosmopolitan wasp group presently treated as a subfamily of Vespidae, but sometimes recognized in the past as a separate family, Eumenidae. Recognition Most eumenine species are black or brown, ...
s similarly build vase-like nests from mud, often with multiple cells, attached to the twigs of trees or against walls.
Predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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 colors ...
, 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 An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
which protects their three main body parts, the
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may ...
, 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 A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
, wasps have several simple eyes known as
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
, 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 In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
adapted for biting and cutting, like those of many other insects, such as
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
s, but their other mouthparts are formed into a suctorial
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
, 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. ...
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 u ...
). Others, such as potter wasps (Eumeninae) and sand wasps ('' Ammophila'',
Sphecidae The Sphecidae are a cosmopolitan family of wasps of the suborder Apocrita that includes sand wasps, mud daubers, and other thread-waisted wasps. The name Sphecidae was formerly given to a much larger grouping of wasps. This was found to be p ...
), 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 Mass provisioning is a form of parental investment in which an adult insect, most commonly a hymenopteran such as a bee or wasp, stocks all the food for each of her offspring in a small chamber (a "cell") before she lays the egg. This behavior i ...
).
Predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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 Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group-ne ...
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 Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
. 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 a ...
'', commonly return to locations where they previously found prey to forage. In many social species, the larvae exude copious amounts of
saliva Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can be ...
ry 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 double ...
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 am ...
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 Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones (ova, oft ...
is determined by a
haplodiploid Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...
system, which means that females are unusually closely related to their sisters, enabling
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like Altruism in animals, altruistic behavio ...
to favour the evolution of eusocial behaviour. Females are
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, 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 are ...
s and develop from fertilized eggs. Males, called drones, have a
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
(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 ...
'' 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 among ...
''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reproduct ...
'' 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 Kin recognition, also called kin detection, is an organism's ability to distinguish between close genetic kin and non-kin. In evolutionary biology and psychology, such an ability is presumed to have evolved for inbreeding avoidance, though animals d ...
. 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. In ...
that is largely due to the expression of
homozygous Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
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 Pollen wasps, the Masarinae, are unusual wasps that are typically treated as a subfamily of Vespidae, but have in the past sometimes been recognized as a separate family, "Masaridae", which also included the subfamilies Euparagiinae and Gayellina ...
gather nectar and pollen in a crop inside their bodies, rather than on body hairs like bees, and pollinate flowers of ''
Penstemon ''Penstemon'' , the beardtongues, is a large genus of roughly 250 species of flowering plants native mostly to the Nearctic, but with a few species also found in the North American portion of the Neotropics. It is the largest genus of flowering ...
'' and the water leaf family, Hydrophyllaceae. The
Agaonidae The family Agaonidae is a group of pollinating and nonpollinating fig wasps. They spend their larval stage inside the fruits of figs. The pollinating wasps (Agaoninae, Kradibiinae, and Tetrapusiinae) are the mutualistic partners of the fig trees ...
(
fig wasp Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while th ...
s) are the only pollinators of nearly 1000 species of
figs The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...
, 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 In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
relationship is fully mutualistic.


As parasitoids

Most solitary wasps are parasitoids. As adults, those that do feed typically only take nectar from flowers.
Parasitoid wasp Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causin ...
s are extremely diverse in habits, many laying their eggs in inert stages of their host (
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
or
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
), 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 The Ichneumonidae, also known as the ichneumon wasps, Darwin wasps, or ichneumonids, are a family (biology), family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 2 ...
are specialized parasitoids, often of Lepidoptera larvae deeply buried in plant tissues, which may be
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin th ...
y. For this purpose, they have exceptionally long ovipositors; they detect their hosts by smell and vibration. Some of the largest species, including ''
Rhyssa persuasoria ''Rhyssa persuasoria'', also known as the sabre wasp, is a species belonging to the family Ichneumonidae subfamily Rhyssinae. Members of this subfamily, including those of ''Rhyssa'' and the allied ''Megarhyssa'', are also known collectively as g ...
'' 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 e ...
'', parasitise
horntail Horntail or wood wasp are any of the 150 non-social species of the hymenopteran family Siricidae, a type of wood-eating sawfly. The common name "horntail" derives from the stout, spine-like structure at the end of the adult's abdomen, which is u ...
s, large sawflies whose adult females also have impressively long ovipositors. Some parasitic species have a mutualistic relationship with a
polydnavirus A polydnavirus (PDV) is a member of the family (biology), family ''Polydnaviridae'' of insect viruses. There are two genera in the family: ''Bracovirus'' and ''Ichnovirus''. Polydnaviruses form a symbiotic relationship with parasitoid wasps; (i ...
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 The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
of the female wasp. One family of chalcidoid wasps, the
Eucharitidae The Eucharitidae are a family of parasitic wasps.Ayre, G.L. Pseudometagea schwarzii (Ashm.) (Eucharitidae: Hymenoptera), a parasite of Lasius neoniger Emery (Formicidae: Hymenoptera). Canadian Journal of Zoology 40 (1962) : 157-164. Eucharitid ...
, 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 colors ...
), 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 Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own ...
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 ca ...
''), 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 In 1909, the entomologist Carlo Emery noted that social parasites among insects (e.g., kleptoparasites) tend to be parasites of species or genera to which they are closely related.Emery, C. "Über den Ursprung der dulotischen, parasitischen und my ...
, social parasitism (biology), social parasites, especially among insects, tend to parasitise species or genera to which they are closely related. For example, the social wasp ''Dolichovespula adulterina'' parasitises other members of its genus such as ''Dolichovespula norwegica, D. norwegica'' and ''Dolichovespula arenaria, D. arenaria''.


As predators

Many wasp lineages, including those in the families
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
, Crabronidae,
Sphecidae The Sphecidae are a cosmopolitan family of wasps of the suborder Apocrita that includes sand wasps, mud daubers, and other thread-waisted wasps. The name Sphecidae was formerly given to a much larger grouping of wasps. This was found to be p ...
, and
Pompilidae Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group-ne ...
, 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 Pest (organism), pests is difficult to establish. The roughly 140 species of beewolf (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 aposematism, warning coloration, social wasps are the models for many species of mimic. Two common cases are
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on bu ...
, where the mimic is harmless and is essentially bluffing, and
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian ...
, 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 monophyly, monophyletic lineage within the ...
.


As prey

While wasp stings deter many potential predators, bee-eaters (in the bird family Meropidae) specialise in eating stinging insects, making aerial sallying, 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 Pernis (bird), 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''). Likewise, roadrunners are the only real predators of tarantula hawk wasps. File:CSIRO ScienceImage 11188 Fig wasp.jpg, Minute pollinating
fig wasp Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while th ...
s, ''Pleistodontes'': the trees and wasps have
coevolved In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
and are mutualistic. File:Latina rugosa planidia.png, ''Latina (genus), Latina rugosa'' planidia (arrows, magnified) attached to an ant larva; the
Eucharitidae The Eucharitidae are a family of parasitic wasps.Ayre, G.L. Pseudometagea schwarzii (Ashm.) (Eucharitidae: Hymenoptera), a parasite of Lasius neoniger Emery (Formicidae: Hymenoptera). Canadian Journal of Zoology 40 (1962) : 157-164. Eucharitid ...
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 colors ...
, such as this ''Hedychrum rutilans'', are known as cuckoo or jewel wasps for their parasitic behaviour and metallic iridescence. File:Bee wolf.jpg, European beewolf ''Philanthus triangulum'' mass provisioning, provisioning her nest with a honeybee File:Clytus arietis (Linné, 1758) (3989861203).jpg, Wasp beetle ''Clytus arietis'' is a Batesian mimicry, 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,
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 (biology), family of chalcid wasp, chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. F ...
, and Trichogrammatidae, are exploited commercially to provide biological pest control, biological control of pest insect, insect pests. One of the first species to be used was ''Encarsia formosa'', a parasitoid of a range of species of
whitefly Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described. Description and taxonomy The A ...
. 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 The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
and cucumber, and to a lesser extent of aubergine (eggplant), flowers such as Calendula officinalis, 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 Whiteflies are Hemipterans that typically feed on the undersides of plant leaves. They comprise the family Aleyrodidae, the only family in the superfamily Aleyrodoidea. More than 1550 species have been described. Description and taxonomy The A ...
, an pest insect, insect pest 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 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., a football club in Scotland.


In fashion

Wasps have been modelled in jewellery since at least the nineteenth century, when diamond and emerald wasp brooches were made in gold and silver 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 Aristophanes (; grc, Ἀριστοφάνης, ; c. 446 – c. 386 BC), son of Philippus, of the deme In Ancient Greece, a deme or ( grc, δῆμος, plural: demoi, δημοι) was a suburb or a subdivision of Athens and other city-states ...
wrote the comedy play Σφῆκες (''Sphēkes''), ''
The Wasps ''The Wasps'' ( grc-x-classical, Σφῆκες, translit=Sphēkes) is the fourth in chronological order of the eleven surviving plays by Aristophanes. It was produced at the Lenaia festival in 422 BC, during Athens' short-lived respite from the ...
'', first put on in 422 BC. The "wasps" are the chorus of old jurors.
H. G. Wells Herbert George Wells"Wells, H. G."
Revised 18 May 2015. ''
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 (novel), 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 unive ...
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'' (2006) and its The Girl Who Played with Fire (film), 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'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 have been named , the first an 8-gun sloop launched in 1749. Eleven ships of the United States Navy have similarly borne the name , the first a merchant schooner acquired by the Continental Navy in 1775. The eighth of these, USS Wasp (CV-7), an aircraft carrier, gained two Second World War battle stars, prompting Winston Churchill 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 Universal Carrier, Bren Gun Carrier. 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 developed for United States Air Force special operations.


See also

* Bee and wasp stings * Characteristics of common wasps and bees * 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 Wasps,