Wasp

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Wasp
Temporal range: Jurassic–Present
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Vespula germanica Richard Bartz.jpg
A social wasp, Vespula germanica
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Euarthropoda
Class: Insecta
Superorder: Hymenopterida
Order: Hymenoptera
Groups included
Cladistically included but traditionally excluded taxa

A wasp is any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. The Apocrita have a common evolutionary ancestor and form a clade; wasps as a group do not form a clade, but are paraphyletic with respect to bees and ants.

Insect Class of invertebrates

Insects or Insecta are hexapod invertebrates and the largest group within the arthropod phylum. Definitions and circumscriptions vary; usually, insects comprise a class within the Arthropoda. As used here, the term Insecta is synonymous with Ectognatha. Insects have a chitinous exoskeleton, a three-part body, three pairs of jointed legs, compound eyes and one pair of antennae. Insects are the most diverse group of animals; they include more than a million described species and represent more than half of all known living organisms. The total number of extant species is estimated at between six and ten million; potentially over 90% of the animal life forms on Earth are insects. Insects may be found in nearly all environments, although only a small number of species reside in the oceans, which are dominated by another arthropod group, crustaceans.

Hymenoptera Order of insects

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

Apocrita suborder of insects

The Apocrita are 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 between the first two segments of the actual abdomen; the first abdominal segment is fused to the thorax, and is called the propodeum. Therefore, it is general practice, when discussing the body of an apocritan in a technical sense, to refer to the mesosoma and metasoma rather than the "thorax" and "abdomen", respectively. The evolution of a constricted waist was an important adaption for the parasitoid lifestyle of the ancestral apocritan, allowing more maneuverability of the female's ovipositor. The ovipositor either extends freely or is retracted, and may be developed into a stinger for both defense and paralyzing prey. Larvae are legless and blind, and either feed inside a host or in a nest cell provisioned by their mothers.

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 the ovipositor is often modified instead into a sting 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 wasps, 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. Unlike true parasites, the wasp larvae eventually kill their hosts. Solitary wasps parasitize almost every pest insect, making wasps valuable in horticulture for biological pest control of species such as whitefly in tomatoes and other crops.

Yellowjacket predatory wasps of the genera Vespula and Dolichovespula

Yellowjacket or Yellow jacket is the common name in North America for predatory social wasps of the genera Vespula and Dolichovespula. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of these are black and yellow like the eastern yellowjacket Vespula maculifrons and the aerial yellowjacket Dolichovespula arenaria; some are black and white like the bald-faced hornet, Dolichovespula maculata. Others may have the abdomen background color red instead of black. They can be identified by their distinctive markings, their occurrence only in colonies, and a characteristic, rapid, side-to-side flight pattern prior to landing. All females are capable of stinging. Yellowjackets are important predators of pest insects.

Hornet genus of insects

Hornets are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to 5.5 cm (2.2 in) in length. They are distinguished from other vespine wasps by the relatively large top margin of the head and by the rounded segment of the abdomen just behind the waist. Worldwide, there are 22 recognized species of Vespa, Most species only occur in the tropics of Asia, though the European hornet, is widely distributed throughout Europe, Russia, North America and Northeast Asia. Wasps native to North America in the genus Dolichovespula are commonly referred to as hornets, but are actually yellowjackets.

Vespidae family of insects

The Vespidae are a large, diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps and many solitary wasps. Each social wasp colony includes a queen and a number of female workers with varying degrees of sterility relative to the queen. In temperate social species, colonies usually only last one year, dying at the onset of winter. New queens and males (drones) are produced towards the end of the summer, and after mating, the queens hibernate over winter in cracks or other sheltered locations. The nests of most species are constructed out of mud, but polistines and vespines use plant fibers, chewed to form a sort of paper. Many species are pollen vectors contributing to the pollination of several plants, being potential or even effective pollinators, while others are notable predators of pest insect species.

Wasps first appeared in the fossil record in the Jurassic, and diversified into many surviving superfamilies by the Cretaceous. 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 5 centimetres (2.0 in) 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 ). The smallest wasps are solitary chalcid wasps in the family Mymaridae, including the world's smallest known insect, with a body length of only 0.139 mm (0.0055 in), and the smallest known flying insect, only 0.15 mm (0.0059 in) long.

The Jurassic period was a geologic period and system that spanned 56 million years from the end of the Triassic Period 201.3 million years ago (Mya) to the beginning of the Cretaceous Period 145 Mya. The Jurassic constitutes the middle period of the Mesozoic Era, also known as the Age of Reptiles. The start of the period was marked by the major Triassic–Jurassic extinction event. Two other extinction events occurred during the period: the Pliensbachian-Toarcian extinction in the Early Jurassic, and the Tithonian event at the end; however, neither event ranks among the "Big Five" mass extinctions.

The Cretaceous is a geologic period and system that spans 79 million years from the end of the Jurassic Period 145 million years ago (mya) to the beginning of the Paleogene Period 66 mya. It is the last period of the Mesozoic Era, and the longest period of the Phanerozoic Eon. The Cretaceous Period is usually abbreviated K, for its German translation Kreide.

Asian giant hornet species of insect

The Asian giant hornet, including the subspecies Japanese giant hornet, colloquially known as the yak-killer hornet, is the world's largest hornet, native to temperate and tropical Eastern Asia. They prefer to live in low mountains and forests, while almost completely avoiding plains and high-altitude climates. V. mandarinia creates nests by digging, co-opting pre-existing tunnels dug by rodents, or occupying spaces near rotted pine roots. It feeds primarily on larger insects, colonies of other eusocial insects, tree sap, and honey from honey bee colonies. The hornet has a body length of 45 mm (1.8 in), a wingspan around 75 mm (3.0 in), and a stinger 6 mm (0.24 in) long, which injects a large amount of potent venom.

Wasps have appeared in literature from Classical times, as the eponymous chorus of old men in Aristophanes' 422 BC comedy Σφῆκες (Sphēkes), The Wasps , and in science fiction from H. G. Wells's 1904 novel The Food of the Gods and How It Came to Earth , featuring giant wasps with three-inch-long stings. The name "Wasp" has been used for many warships and other military equipment.

Aristophanes ancient Athenian comic playwright

Aristophanes, son of Philippus, of the deme Kydathenaion, was a comic playwright of ancient Athens. Eleven of his forty plays survive virtually complete. These provide the most valuable examples of a genre of comic drama known as Old Comedy and are used to define it, along with fragments from dozens of lost plays by Aristophanes and his contemporaries.

<i>The Wasps</i> comedy by Aristophanes

The Wasps is the fourth in chronological order of the eleven surviving plays by Aristophanes, the master of an ancient genre of drama called 'Old Comedy'. It was produced at the Lenaia festival in 422 BC, a time when Athens was enjoying a brief respite from the Peloponnesian War following a one-year truce with Sparta.

Science fiction Genre of speculative fiction

Science fiction is a genre of speculative fiction that has been called the "literature of ideas". It typically deals with imaginative and futuristic concepts such as advanced science and technology, time travel, parallel universes, fictional worlds, space exploration, and extraterrestrial life. Science fiction often explores the potential consequences of scientific innovations.

Taxonomy and phylogeny

Wasps are paraphyletic, consisting of the clade Apocrita without ants and bees, which are not usually considered to be wasps. The Hymenoptera also contain the somewhat wasplike Symphyta, the sawflies. The familiar common wasps and yellowjackets belong to one family, the Vespidae. Wasps are Paraphyletic.svg
Wasps are paraphyletic, consisting of the clade Apocrita without ants and bees, which are not usually considered to be wasps. The Hymenoptera also contain the somewhat wasplike Symphyta, the sawflies. The familiar common wasps and yellowjackets belong to one family, the Vespidae.
Palaeovespa florissantia, a fossil wasp (Vespinae) from the Eocene rocks of the Florissant fossil beds of Colorado, c. 34 mya Palaeovespa florissantia.jpg
Palaeovespa florissantia , a fossil wasp (Vespinae) from the Eocene rocks of the Florissant fossil beds of Colorado, c. 34 mya

Paraphyletic grouping

The wasps are a cosmopolitan paraphyletic grouping of hundreds of thousands of species, [1] [2] consisting of the narrow-waisted Apocrita without the ants and bees. [3] The Hymenoptera also contain the somewhat wasplike but unwaisted Symphyta, the sawflies.

Ant family of insects

Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the Cretaceous period, about 140 million years ago, and diversified after the rise of flowering plants. More than 12,500 of an estimated total of 22,000 species have been classified. They are easily identified by their elbowed antennae and the distinctive node-like structure that forms their slender waists.

Bee clade of insects

Bees are flying insects closely related to wasps and ants, known for their role in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea and are presently considered a clade, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families. They are found on every continent except Antarctica, in every habitat on the planet that contains insect-pollinated flowering plants.

Sawfly suborder of insects

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 their eggs. The name is associated especially with the Tenthredinoidea, by far the largest superfamily, with about 7,000 known species; in the entire suborder, there are 8,000 described species in more than 800 genera. The suborder Symphyta is paraphyletic, consisting of several basal groups within the order Hymenoptera. Symphyta includes all the species within Hymenoptera that are not wasps or the descendants of wasps.

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 and Dolichovespula), hornets (genus Vespa), and members of the subfamily Polistinae.

Yellowjackets band

Yellowjackets is an American jazz fusion band.

<i>Vespula</i> genus of insects

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 in North America. Vespula species have a shorter oculomalar space and a more pronounced tendency to nest underground than Dolichovespula.

<i>Dolichovespula</i> genus of insects

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, along with members of their sister genus Vespula. In a study on the nesting biology of Dolichovespula, a colony of D. maculata with 771 workers was reported as having the largest recorded population count.

Fossils

Male Electrostephanus petiolatus fossil from the Middle Eocene, preserved in Baltic amber Neotype male of Electrostephanus petiolatus Brues in Baltic amber (AMNH B-JWJ-260).png
Male Electrostephanus petiolatus fossil from the Middle Eocene, preserved in Baltic amber

Hymenoptera in the form of Symphyta (Xyelidae) first appeared in the fossil record in the Lower Triassic. Apocrita, wasps in the broad sense, appeared in the Jurassic, and had diversified into many of the extant superfamilies by the Cretaceous; they appear to have evolved from the Symphyta. [4] 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. [5]

The Vespidae include the extinct genus Palaeovespa , seven species of which are known from the Eocene rocks of the Florissant fossil beds of Colorado and from fossilised Baltic amber in Europe. [6] Also found in Baltic amber are crown wasps of the genus Electrostephanus . [7] [8]

Diversity

Megascolia procer, a giant solitary species from Java in the Scoliidae. This specimen's length is 7.7 cm and its wingspan is 11.5 cm. Megascolia procer MHNT dos.jpg
Megascolia procer , a giant solitary species from Java in the Scoliidae. This specimen's length is 7.7 cm and its wingspan is 11.5 cm.

Wasps are a diverse group, estimated at over a hundred thousand described species around the world, and a great many more as yet undescribed. [10] [lower-alpha 2] For example, there are over 800 species of fig trees, mostly in the tropics, and almost all of these has its own specific fig wasp (Chalcidoidea) to effect pollination. [11]

Megarhyssa macrurus, a parasitoid. The body of a female is c. 2 inches (51 mm) long, with an ovipositor c. 4 inches (100 mm) long Ichneumon wasp (Megarhyssa macrurus lunato) (7686081848).jpg
Megarhyssa macrurus , a parasitoid. The body of a female is c. 2 inches (51 mm) long, with an ovipositor c. 4 inches (100 mm) long

Many wasp species are parasitoids; the females deposit eggs on or in a host arthropod 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. [11]

Tarantula hawk wasp dragging an orange-kneed tarantula to her burrow; this species has the most painful sting of any wasp. Wasp with Orange-kneed tarantula.JPG
Tarantula hawk wasp dragging an orange-kneed tarantula to her burrow; this species has the most painful sting of any wasp.

The largest social wasp is the Asian giant hornet, at up to 5 centimetres (2.0 in) in length. [13] The various tarantula hawk wasps are of a similar size [14] 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. [12] The solitary giant scoliid, Megascolia procer , with a wingspan of 11.5 cm, [9] has subspecies in Sumatra and Java; [15] it is a parasitoid of the Atlas beetle Chalcosoma atlas . [16] The female giant ichneumon wasp Megarhyssa macrurus is 12.5 centimetres (5 in) long including its very long but slender ovipositor which is used for boring into wood and inserting eggs. [17] The smallest wasps are solitary chalcid wasps in the family Mymaridae, including the world's smallest known insect, Dicopomorpha echmepterygis (139 micrometres long) and Kikiki huna with a body length of only 158 micrometres, the smallest known flying insect. [18]

There are estimated to be 100,000 species of ichneumonoid wasps in the families Braconidae and Ichneumonidae. These are almost exclusively parasitoids, mostly utilising other insects as hosts. Another family, the Pompilidae, is a specialist parasitoid of spiders. [11] 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. [19] 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 climbing squash. [20] [21]

Social wasps

Social wasps constructing a paper nest Active Wasp Nest.jpg
Social wasps constructing a paper nest

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, 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 fibers 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 and D. sylvestris prefer to nest in trees and shrubs; Protopolybia exigua attaches its nests on the underside of leaves and branches; Polistes erythrocephalus chooses sites close to a water source. [22]

Social wasps on their nest

Other wasps, like Agelaia multipicta and Vespula germanica, 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. [23] 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. [24]

Solitary wasps

Potter wasp building mud nest, France. The latest ring of mud is still wet. Potter Wasp building mud nest near completion.JPG
Potter wasp building mud nest, France. The latest ring of mud is still wet.

The vast majority of wasp species are solitary insects. [11] [25] 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. [25]

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. [25] Mud daubers and pollen wasps construct mud cells in sheltered places. [26] Potter wasps similarly build vase-like nests from mud, often with multiple cells, attached to the twigs of trees or against walls. [27]

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, the cuckoo wasps, are kleptoparasites and lay their eggs in the nests of unrelated host species. [25]

Biology

Anatomy

European hornet, Vespa crabro Hornet-vespa.jpg
European hornet, Vespa crabro

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. [28] It either extends freely or can be retracted, and may be developed into a stinger for both defence and for paralysing prey. [29]

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 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. [30]

The larvae of wasps resemble maggots, 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). [31]

Sand wasp Bembix oculata (Crabronidae) feeding on a fly after paralysing it with its sting Wasp August 2007-23.jpg
Sand wasp Bembix oculata (Crabronidae) feeding on a fly after paralysing it with its sting

Diet

Black wasp Sphex pensylvanicus (Sphecidae) feeding on nectar Sphex pensylvanicus.jpg
Black wasp Sphex pensylvanicus (Sphecidae) feeding on nectar

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. [25] Some wasp species provide food for the young repeatedly during their development (progressive provisioning). [32] Others, such as potter wasps (Eumeninae) [33] and sand wasps ( Ammophila , Sphecidae), [34] 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). [35]

Spider wasp (Pompilidae) dragging a jumping spider (Salticidae) to provision a nest IndianSpiderWasp.JPG
Spider wasp (Pompilidae) dragging a jumping spider (Salticidae) to provision a nest

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. [25] The Pompilidae specialize in catching spiders to provision their nests. [36]

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 , commonly return to locations where they previously found prey to forage. [37] In many social species, the larvae exude copious amounts of salivary secretions that are avidly consumed by the adults. These include both sugars and amino acids, and may provide essential protein-building nutrients that are otherwise unavailable to the adults (who cannot digest proteins). [38]

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 chromosomes and develop from fertilized eggs. Males, called drones, have a haploid (n) number of chromosomes and develop from an unfertilized egg. [29] 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. [25] Experimental infection of Muscidifurax uniraptor with the bacterium Wolbachia induced thelytokous reproduction and an inability to produce fertile, viable male offspring. [39]

Inbreeding avoidance

Females of the solitary wasp parasitoid Venturia canescens can avoid mating with their brothers through kin recognition. [40] 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. [40] Sibling-mating avoidance reduces inbreeding depression that is largely due to the expression of homozygous deleterious recessive mutations. [41]

Ecology

As pollinators

Minute pollinating fig wasps, Pleistodontes: the trees and wasps have coevolved and are mutualistic. CSIRO ScienceImage 11188 Fig wasp.jpg
Minute pollinating fig wasps, Pleistodontes : the trees and wasps have coevolved and are mutualistic.

While the vast majority of wasps play no role in pollination, a few species can effectively transport pollen and pollinate several plant species. [42] Since wasps generally do not have a fur-like covering of soft hairs and a special body part for pollen storage (Pollen basket) as some bees do, pollen does not stick to them well. [43] 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. [44]

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. [45]

The Agaonidae (fig wasps) are the only pollinators of nearly 1000 species of figs, [43] 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. [46]

As parasitoids

The parasitoidal ichneumon wasp Dolichomitus imperator ovipositing through wood, using its immensely long ovipositor to lay its eggs inside hidden larvae, detected by vibration Dolichomitus imperator Eiablage-02-R Bartz.jpg
The parasitoidal ichneumon wasp Dolichomitus imperator ovipositing through wood, using its immensely long ovipositor to lay its eggs inside hidden larvae, detected by vibration
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. 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.

Most solitary wasps are parasitoids. [47] 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. [48]

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 , parasitise horntails, large sawflies whose adult females also have impressively long ovipositors. [49] Some parasitic species have a mutualistic relationship with a polydnavirus that weakens the host's immune system and replicates in the oviduct of the female wasp. [11]

One family of chalcid 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. [50] [51] [52]

The Chrysididae, such as this Hedychrum rutilans, are known as cuckoo or jewel wasps for their parasitic behaviour and metallic iridescence. Goudwesp.jpg
The Chrysididae, such as this Hedychrum rutilans , are known as cuckoo or jewel wasps for their parasitic behaviour and metallic iridescence.

As parasites

Many species of wasp, including especially the cuckoo or jewel wasps (Chrysididae), 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 , which lays its eggs in the nests of other paper wasps (specifically Polistes dominula ), and whose larvae are then fed directly by the host. [53] [54] 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. [55] According to Emery's rule, social parasites, especially among insects, tend to parasitise species or genera to which they are closely related. [56] [57] For example, the social wasp Dolichovespula adulterina parasitises other members of its genus such as D. norwegica and D. arenaria . [58] [59]

As predators

European beewolf Philanthus triangulum provisioning her nest with a honeybee Bee wolf.jpg
European beewolf Philanthus triangulum provisioning her nest with a honeybee

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 is difficult to establish. [60]

The roughly 140 species of beewolf (Philanthinae) hunt bees, including honeybees, to provision their nests; the adults feed on nectar and pollen. [61]

As models for mimics

The wasp beetle Clytus arietis is one of many Batesian mimics of wasps. Clytus arietis (Linne, 1758) (3989861203).jpg
The wasp beetle Clytus arietis is one of many Batesian mimics of wasps.

With their powerful stings and conspicuous warning coloration, 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. [62]

Bee-eaters such as Merops apiaster specialise in feeding on bees and wasps. Pair of Merops apiaster feeding detail.jpg
Bee-eaters such as Merops apiaster specialise in feeding on bees and wasps.

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. [63] The honey buzzard attacks the nests of social hymenopterans, eating wasp larvae; it is the only known predator of the dangerous [64] Asian giant hornet or "yak-killer" ( Vespa mandarinia ). [65] Likewise, Roadrunners are the only real predators of tarantula hawk wasps. [66]

Relationship with humans

Encarsia formosa, a parasitoid, is sold commercially for biological control of whitefly, an insect pest of tomato and other horticultural crops. 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 of tomato and other horticultural crops.
Tomato leaf covered with nymphs of whitefly parasitised by Encarsia formosa Tomate Blatt Eier Weisse Fliege parasitiert.jpg
Tomato leaf covered with nymphs of whitefly parasitised by Encarsia formosa

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, when wasp colonies stop breeding new workers; the existing workers search for sugary foods and are more likely to come into contact with humans; if people then respond aggressively, the wasps sting. [67] 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. [68] Stings are usually painful rather than dangerous, but in rare cases, people may suffer life-threatening anaphylactic shock. [69]

In horticulture

Some species of parasitic wasp, especially in the Trichogrammatidae, are exploited commercially to provide biological control of insect pests. [2] [70] For example, in Brazil, farmers control sugarcane borers with the parasitic wasp Trichogramma galloi . [71] 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. [72] Several species of parasitic wasp are natural predators of aphids and can help to control them. [73] For instance, Aphidius matricariae is used to control the peach-potato aphid. [74]

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. [75] The club has an amateur side called Wasps FC. [76]

Among the other clubs bearing the name are a basketball club in Wantirna, Australia, [77] and Alloa Athletic F.C., a football club in Scotland. [78]

In fashion

Wasp waist, c. 1900, demonstrated by Polaire, a French actress famous for this silhouette Polaire, French actress 5.jpg
Wasp waist, c. 1900, demonstrated by Polaire, a French actress famous for this silhouette

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. [79] 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. [80] [81]

In literature

The Ancient Greek playwright Aristophanes wrote the comedy play Σφῆκες (Sphēkes), The Wasps , first put on in 422 BC. The "wasps" are the chorus of old jurors. [82]

H. G. Wells made use of giant wasps in his novel The Food of the Gods and How It Came to Earth (1904): [83]

It flew, he is convinced, within a yard of him, struck the ground, rose again, came down again perhaps thirty yards away, and rolled over with its body wriggling and its sting stabbing out and back in its last agony. He emptied both barrels into it before he ventured to go near. When he came to measure the thing, he found it was twenty-seven and a half inches across its open wings, and its sting was three inches long. ... The day after, a cyclist riding, feet up, down the hill between Sevenoaks and Tonbridge, very narrowly missed running over a second of these giants that was crawling across the roadway. [83]

Detail of Botticelli's Venus and Mars, 1485, with a wasp's nest on right, probably a symbol of the Vespucci family who commissioned the painting. Botticelli-Venus and Mars (cropped).jpg
Detail of Botticelli's Venus and Mars, 1485, with a wasp's nest on right, probably a symbol of the Vespucci family who commissioned the painting.

Wasp (1957) is a science fiction book by the English writer Eric Frank Russell; it is generally considered Russell's best novel. [85] In Stieg Larsson's book The Girl Who Played with Fire (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. [86]

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:

I own that I cannot see as plainly as others do, and as I should wish to do, evidence of design and beneficence on all sides of us. There seems to me too much misery in the world. I cannot persuade myself that a beneficent and omnipotent God would have designedly created the Ichneumonidae with the express intention of their feeding within the living bodies of caterpillars, or that a cat should play with mice. [87]

In military names

HMS Wasp (1880), one of nine Royal Navy warships to bear the name HMS Wasp (1880).jpg
HMS Wasp (1880), one of nine Royal Navy warships to bear the name

With its powerful sting and familiar appearance, the wasp has given its name to many ships, aircraft and military vehicles. [88] Nine ships and one shore establishment of the Royal Navy have been named HMS Wasp, the first an 8-gun sloop launched in 1749. [89] Eleven ships of the United States Navy have similarly borne the name USS Wasp, the first a merchant schooner acquired by the Continental Navy in 1775. [90] The eighth of these, an aircraft carrier, gained two Second World War battle stars, prompting Winston Churchill to remark "Who said a Wasp couldn't sting twice?" [88] In the Second World War, a German self-propelled howitzer was named Wespe, [91] while the British developed the Wasp flamethrower from the Bren Gun Carrier. [92] In aerospace, the Westland Wasp was a military helicopter developed in England in 1958 and used by the Royal Navy and other navies. [93] The AeroVironment Wasp III is a miniature UAV developed for United States Air Force special operations. [94]

See also

Notes

  1. Specimen measured from photograph.
  2. Methods to estimate species diversity include extrapolating the rate of species descriptions by subfamily (as in the Braconidae) until zero is reached; and extrapolating geographically from the species distribution of well-studied taxa to the group of interest (say, the Braconidae). Dolphin et al found a correlation between the predicted numbers of undescribed species by these two methods, doubling or tripling the number of species in the group. [10]

Related Research Articles

Ovipositor Anatomical structure for laying eggs

The ovipositor is an organ used by some animals 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 typically its form is adapted to functions such as transmitting the egg, preparing a place for it, and placing it properly. In some insects the organ is used merely to attach the egg to some surface, but in many parasitic species it is a piercing organ as well.

Parasitoid organism that spends a significant portion of its life history attached to or within a single host organism in a relationship where the host is ultimately killed

A parasitoid is an organism that lives in close association with its host and at the host's expense, and which sooner or later kills it. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

Paper wasp

Paper wasps are vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct water-resistant nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrella wasps, due to the distinctive design of their nests.

Ichneumonoidea superfamily of insects

The superfamily Ichneumonoidea contains the two largest families within Hymenoptera: Ichneumonidae and Braconidae. The group is thought to contain as many as 100,000 species, many of which have not yet been described. Like other parasitoid wasps, they were long placed in the "Parasitica", variously considered as an infraorder or an unranked clade, now known to be paraphyletic.

<i>Polistes</i> genus of insects

Wasps of the cosmopolitan genus Polistes are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. It is also the single largest genus within the family Vespidae, with over 300 recognized species and subspecies. Their innate preferences for nest-building sites leads them to commonly build nests on human habitation, where they can be very unwelcome; although generally not aggressive, they can be provoked into defending their nests. All species are predatory, and they may consume large numbers of caterpillars, in which respect they are generally considered beneficial. The European paper wasp, Polistes dominula, was introduced into the US about 1981 and has quickly spread throughout most of the country, in most cases replacing native species within a few years. This species is very commonly mistaken for a yellow jacket, as it is black, strongly marked with yellow, and quite different from the native North American species of Polistes. The cuckoo wasp, Polistes sulcifer, is an obligate social parasite, whose only host is P. dominula. Polistes annularis, whose species name is Latin for "ringed", is also known for its distinctive red body color. Polistes metricus adults malaxate their insect prey by chewing them into a pulp, sucking out and ingesting the body fluids, then feeding the rest of the morsel to their larvae. The most widely distributed South American wasp species, Polistes versicolor, is particularly common in the southeastern Brazilian states. This social wasp is commonly referred to as the yellow paper wasp due to the distinct yellow bands found on its thorax and abdomen. Polistes wasps can be identified by their characteristic flight; their long legs dangle below their bodies, which are also more slender than a yellow jacket.

Parasitoid wasp evolutionary grade of hymenopteran superfamilies

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 causing the death of these hosts. Different species specialise in hosts from different insect orders, most often Lepidoptera, though some select beetles, flies, or bugs; the spider wasps (Pompilidae) exclusively attack spiders.

<i>Polistes gallicus</i> species of insect

Polistes gallicus is a fairly common species of paper wasp found in various parts of Europe, excluding England, Denmark, and Scandinavia, from warmer climates to cooler regions north of the Alps. The distribution of P. gallicus also extends into northern regions of Africa, Israel, Iran, and even parts of China and Russia. Nests of these social insects are created in these various conditions. The Polistes species uses an oral secretion to construct their nests, which consist of a combination of saliva and chewed plant fibers. This structural mixture physically protects the nest from various harsh elements and from weathering over time.

<i>Polistes chinensis</i> species of insect

Polistes chinensis is a polistine vespid wasp in the cosmopolitan genus Polistes, and is commonly known as the Asian, Chinese or Japanese paper wasp. It is found in East Asia, in particular China and Japan. The subspecies P. chinensis antennalis is an invasive species in New Zealand, having arrived in 1979.

<i>Polistes annularis</i> species of insect

Polistes annularis is a species of paper wasp which lives throughout the Caribbean and in parts of North America. Its species name is Latin for "ringed" and it is known for its distinct red body color. It builds its nest under overhangs near bodies of water that minimize the amount of sunlight penetration. It clusters its nests together in large aggregations, and consumes nectar and other insects. Its principal predator is the ant, although birds are also known to prey on it. It is a primitively eusocial wasp, meaning that all individuals develop the capacity for reproduction, regardless of social caste. This primitive eusociality has been seen in bees as well, including the sweat bee, Lasioglossum zephyrum. As such, P. annularis demonstrates behavior typical of other polistine wasps, and has a dominance hierarchy, relatively small colony size, and a female-biased sex ratio. Unlike other wasps, P. annularis is relatively robust in winter conditions, and has also been observed to store honey in advance of hibernation. It is closely related to P. major, P. buysonni, and others in the subgenus Aphanilopterus, and slightly less related to the more common P. bellicosus, P. carolina, P. metricus, and P. fuscatus.

<i>Polistes metricus</i> species of insect

Polistes metricus is a wasp native to North America. In the United States, it ranges throughout the southern Midwest, the South, and as far northeast as New York, but has recently been spotted in southwest Ontario. A single female specimen has also been reported from Dryden, Maine. Polistes metricus is dark colored, with yellow tarsi and black tibia. Nests of Polistes metricus can be found attached to the sides of buildings, trees, and shrubbery.

Hunting wasp

Hunting wasps are members of various taxa of the insect order Hymenoptera. Their habits and affinities vary in many ways, but all practise parental care of their larvae in that they capture prey, usually insects, to feed their larvae. Whether solitary or social, most species construct some form of protection or nest in which they hide the prey and in which the larvae can feed and pupate in reasonable security.

<i>Brachymeria</i> genus of insects

Brachymeria is a genus of parasitic wasps in the family Chalcididae. Over 300 species are known worldwide, all of them parasites of insect larvae. Most species are black with limited yellow markings, and like most chalcidid wasps, they have enlarged hind femora. The female typically lays eggs inside the larva of a lepidopteran using its ovipositor. Although mostly parasitic on Lepidoptera, a few are hyperparasites, or attack other types of insect larvae. The adult parasites emerge typically from the host pupa. Some species have been used in biological control.

<i>Polistes exclamans</i> species of insect

Polistes exclamans is a social wasp and is part of the family Vespidae of the order Hymenoptera. It is found throughout the United States, Mexico, the Bahamas, Jamaica and parts of Canada. Due to solitary nest founding by queens, P. exclamans has extended its range in the past few decades and now covers the eastern half of the United States, as well as part of the north. This expansion is typically attributed to changing global climate and temperatures. P. exclamans has three specific castes, including males, workers, and queens, but the dominance hierarchy is further distinguished by age. The older the wasp is, the higher it is in ranking within the colony. In most P. exclamans nests, there is one queen who lays all the eggs in the colony. The physiological similarities between the worker and queen castes have led to experiments attempting to distinguish the characteristics of these two castes and how they are determined, though males have easily identifiable physiological characteristics. Since P. exclamans live in relatively small, open combed nests, they are often subject to predators and parasites, such as Chalcoela iphitalis, Elasmus polistis, and birds. P. exclamans have defense and recognition strategies that help protect against these predators and parasites.

Polistes carnifex, also known as the executioner wasp, is a neotropical vespid wasp in the cosmopolitan genus Polistes, widely known for its extremely painful and potent sting. The wasp is native to Central and South America. It is a very large yellow and brown paper wasp that establishes small colonies which build nests under the eaves of buildings or suspended from branches. The colonies are often founded by solitary queens, many times in the form of winter aggregations to wait until environmental conditions are more suitable for building a colony. Not all nests have a female with developed ovaries. Foraging adults bring nectar and caterpillars back to the nest to feed to the developing larvae which are individually housed in separate cells in the nest. Adaptations include mandibles and teeth. Worker-queen conflict, worker policing, and dominance interactions all come into play for the species. Polistes carnifex is not territorial and has not been observed to be parasitized. These species tend to live from 6 months to 18 months, making it the longest-living wasp known to date.

Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed at about this time as it is present in all three groups of Apocrita, the wasps, bees and ants.

<i>Polistes biglumis</i> species of insect

Polistes biglumis is a species of social wasp within Polistes, the most common genus of paper wasp. It is distinguished mainly by its tendency to reside in montane climates in meadows or alpine areas. Selection pressure from the wasp's environment has led to several idiosyncrasies of its behavior and life cycle with respect to its relative species in the genus Polistes. It alone among paper wasps is often polyandrous. In addition, it has a truncated nesting season that gives rise to unique competitive dynamics among females of the species. P. biglumis wasps utilize an odor based recognition system that is the basis for all wasp to wasp interaction of the species. The wasp's life cycle is highly intertwined with that of Polistes atrimandibularis, an obligate social parasite wasp that frequently invades the combs of P. biglumis wasps.

<i>Chrysis ignita</i> species of insect

Chrysis ignita, also known as the ruby-tailed wasp, is a species of cuckoo wasps. Cuckoo wasps are kleptoparasites – they lay their eggs in the nests of other wasp species and their young consume the eggs or larva of the host wasp for sustenance. These wasps have a number of adaptions which have evolved to equip them for their life cycle. Chrysis ignita parasitize mason bees in particular. Ruby-tailed wasps have metallic, armored bodies, and can roll up into balls to protect themselves from harm when infiltrating the nests of host bees and wasps. Unlike most other Hymenopterans, cuckoo wasps cannot sting. Chrysis ignita is found across the European continent.

Protopolybia chartergoides, also known as Pseudochartergus chartergoides, is a species of wasp within the genus Protopolybia. It is a social wasp found in southern Central America and northern South America.

References

  1. Broad, Gavin (25 June 2014). "What's the point of wasps?". Natural History Museum. Retrieved 18 June 2015.
  2. 1 2 "Wasp". National Geographic.
  3. Johnson, Brian R.; Borowiec, Marek L.; Chiu, Joanna C.; Lee, Ernest K.; Atallah, Joel; Ward, Philip S. (2013). "Phylogenomics Resolves Evolutionary Relationships among Ants, Bees, and Wasps" (PDF). Current Biology. 23 (20): 2058–2062. doi:10.1016/j.cub.2013.08.050. PMID   24094856.
  4. Gillott, Cedric (6 December 2012). Entomology. Springer. pp. 302–318. ISBN   978-1-4615-6915-2.
  5. "World's oldest fig wasp fossil proves that if it works, don't change it". University of Leeds. 15 June 2010. Retrieved 5 August 2015.
  6. Poinar, G. (2005). "Fossil Trigonalidae and Vespidae (Hymenoptera) in Baltic amber". Proceedings of the Entomological Society of Washington. 107 (1): 55–63.
  7. Engel, M.S.; Ortega-Blanco, J. (2008). "The fossil crown wasp Electrostephanus petiolatus Brues in Baltic Amber (Hymenoptera, Stephanidae): designation of a neotype, revised classification, and a key to amber Stephanidae". ZooKeys (4): 55–64. doi:10.3897/zookeys.4.49.
  8. Yunakov, N.N.; Kirejtshuk, A.G. (2011). "New genus and species of broad-nosed weevils from Baltic amber and notes on fossils of the subfamily Entiminae (Coleoptera, Curculionidae)". ZooKeys (160): 73–96. doi:10.3897/zookeys.160.2108. PMC   3253632 . PMID   22303121.
  9. 1 2 Sarrazin, Michael; Vigneron, Jean Pol; Welch, Victoria; Rassart, Marie (5 November 2008). "Nanomorphology of the blue iridescent wings of a giant tropical wasp Megascolia procer javanensis (Hymenoptera)". Phys. Rev. E 78 (5): 051902. arXiv: 0710.2692 . Bibcode:2008PhRvE..78e1902S. doi:10.1103/PhysRevE.78.051902. PMID   19113150. Measurement scale on Figure 1.
  10. 1 2 Dolphin, Konrad; Quicke, Donald L. J. (July 2001). "Estimating the global species richness of an incompletely described taxon: an example using parasitoid wasps (Hymenoptera: Braconidae)". Biological Journal of the Linnean Society. 73 (3): 279–286. doi:10.1111/j.1095-8312.2001.tb01363.x.
  11. 1 2 3 4 5 Godfray, H.C.J. (1994). Parasitoids: Behavioral and Evolutionary Ecology. Princeton University Press. pp. 3–24. ISBN   978-0-691-00047-3.
  12. 1 2 Williams, David B. "Tarantula Hawk". DesertUSA. Retrieved 13 June 2015.
  13. Backshall, Steve (2007). Steve Backshall's venom: poisonous animals in the natural world. New Holland Publishers. p. 147. ISBN   978-1-84537-734-2.
  14. Carwardine, Mark (2008). Animal Records. Sterling Publishing. p. 218. ISBN   978-1-4027-5623-8. Pepsis heros, which has a body length of up to 5.7cm (2 1/4 in) and a maximum wingspan of 11.4 cm (4 1/2 in).
  15. Betrem, J. G.; Bradley, J. Chester (1964). "Annotations on the genera Triscolia, Megascolia and Scolia (Hymenoptera, Scoliidae)". Zoologische Mededelingen. 39 (43): 433–444.
  16. Piek, Tom (2013). Venoms of the Hymenoptera: Biochemical, Pharmacological and Behavioural Aspects. Elsevier. p. 173. ISBN   978-1-4832-6370-0.
  17. Cranshaw, W. (5 August 2014). "Pigeon Tremex Horntail and the Giant Ichneumon Wasp". Colorado State University Extension. Retrieved 15 June 2015.
  18. Huber, John; Noyes, John (2013). "A new genus and species of fairyfly, Tinkerbella nana (Hymenoptera, Mymaridae), with comments on its sister genus Kikiki, and discussion on small size limits in arthropods". Journal of Hymenoptera Research. 32: 17–44. doi:10.3897/jhr.32.4663.
  19. Ward, D.F.; Schnitzler, F.R. (2013). "Ichneumonidae: Eucerotinae: Euceros Gravenhorst 1829". Landcare Research. Retrieved 15 June 2015.
  20. Westlake, Casey (13 March 2014). "More to biological diversity than meets the eye: Specialization by insect species is the key". Iowa Now. University of Iowa. Retrieved 18 June 2015.
  21. Condon, M. A.; Scheffer, S. J.; Lewis, M. L.; Wharton, R.; Adams, D. C.; Forbes, A. A. (2014). "Lethal interactions between parasites and prey increase niche diversity in a tropical community". Science. 343 (6176): 1240–1244. Bibcode:2014Sci...343.1240C. doi:10.1126/science.1245007. PMID   24626926.
  22. Carlos A. Martin P.; Anthony C. Bellotti (1986). "Biologia y comportamiento de Polistes erythrocephalus" [Biology and behavior of Polistes erythrocephalus](PDF). Acta Agron (in Spanish). 36 (1): 63–76. Retrieved 14 October 2014.
  23. Sôichi Yamane; Sidnei Mateus; Satoshi Hozumi; Kazuyuki Kudô; Ronaldo Zucchi (2009). "How does a colony of Apoica flavissima (Hymenoptera: Vespidae, Epiponini) maintain a constant temperature?". Entomological Science. 12 (3): 341–345. doi:10.1111/j.1479-8298.2009.00328.x.
  24. Ross, Kenneth G. (1991). The Social Biology of Wasps. Cornell University Press. p. 104. ISBN   978-0-8014-9906-7.
  25. 1 2 3 4 5 6 7 O'Neill, Kevin M. (2001). Solitary Wasps: Behavior and Natural History. Cornell University Press. pp. 1–4, 69. ISBN   978-0-8014-3721-2.
  26. Houston, Terry (October 2013). "Slender mud-dauber wasps: genus Sceliphron". Western Australian Museum. Retrieved 12 June 2015.
  27. Grissell, E. E. (April 2007). "Potter wasps of Florida". University of Florida. Retrieved 12 June 2015.
  28. "Hymenoptera: ants, bees and wasps". Insects and their allies. CSIRO. Retrieved 16 June 2015.
  29. 1 2 Grimaldi, David; Engel, Michael S. (2005). Evolution of the Insects. Cambridge University Press. p. 408. ISBN   978-0-521-82149-0.
  30. Krenn, H. W.; Mauss, V.; Plant, J. (2002). "Evolution of the suctorial proboscis in pollen wasps (Masarinae, Vespidae)". Arthropod Structure & Development. 31 (2): 103–120. doi:10.1016/s1467-8039(02)00025-7. PMID   18088974.
  31. Hoell, H.V.; Doyen, J.T.; Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed. Oxford University Press. pp. 570–579. ISBN   978-0-19-510033-4.
  32. Field, Jeremy (2005). "The evolution of progressive provisioning". Behavioral Ecology. 16 (4): 770–778. doi:10.1093/beheco/ari054.
  33. Grissell, E. E. "Potter Wasps of Florida". University of Florida / IFAS. Retrieved 15 June 2015.
  34. Field, Jeremy (1989). "Intraspecific parasitism and nesting success in the solitary wasp Ammophila sabulosa". Behaviour . 110 (1–4): 23–45. doi:10.1163/156853989X00367. JSTOR   4534782.
  35. Elgar, Mark A.; Jebb, Matthew (1999). "Nest Provisioning In The Mud-Dauber Wasp Sceliphron laetum (F. Smith): Body Mass And Taxa Specific Prey Selection". Behaviour. 136 (2): 147–159. doi:10.1163/156853999501252.
  36. "Spider Wasps". Australian Museum. Retrieved 15 June 2015.
  37. Richter, M. Raveret (2000). "Social Wasp (Hymenoptera: Vespidae) Foraging Behavior". Annual Review of Entomology. 45: 121–150. doi:10.1146/annurev.ento.45.1.121. PMID   10761573.
  38. Wilson, Edward O. (2000). Sociobiology: The New Synthesis. Harvard University Press. p. 344. ISBN   978-0-674-00089-6.
  39. Gottlieb, Yuval; Zchori-Fein, Einat (2001). "Irreversible thelytokous reproduction in Muscidifurax uniraptor" (PDF). Entomologia Experimentalis et Applicata. 100 (3): 271–278. doi:10.1046/j.1570-7458.2001.00874.x.
  40. 1 2 Metzger M, Bernstein C, Hoffmeister TS, Desouhant E (2010). "Does kin recognition and sib-mating avoidance limit the risk of genetic incompatibility in a parasitic wasp?". PLoS ONE. 5 (10): e13505. Bibcode:2010PLoSO...513505M. doi:10.1371/journal.pone.0013505. PMC   2957437 . PMID   20976063.
  41. Charlesworth D, Willis JH (2009). "The genetics of inbreeding depression". Nat. Rev. Genet. 10 (11): 783–96. doi:10.1038/nrg2664. PMID   19834483.
  42. Sühs, R. B.; Somavilla, A.; Köhler, A; Putzke, J. (2009). "Vespídeos (Hymenoptera, Vespidae) vetores de pólen de Schinus terebinthifolius Raddi (Anacardiaceae), Santa Cruz do Sul, RS, Brasil" [Pollen vector wasps (Hymenoptera, Vespidae) of Schinus terebinthifolius Raddi (Anacardiaceae), Santa Cruz do Sul, RS, Brazil]. Brazilian Journal of Biosciences (in Portuguese). 7 (2): 138–143.
  43. 1 2 "Wasp Pollination". USDA Forest Service. Retrieved 5 August 2015.
  44. Sühs; Somavilla; Putzke; Köhler (2009). "Pollen vector wasps (Hymenoptera, Vespidae) of Schinus terebinthifolius Raddi (Anacardiaceae), Santa Cruz do Sul, RS, Brazil". Brazilian Journal of Biosciences. 7 (2): 138–143 via ABEC.
  45. Tepidino, Vince. "Pollen Wasps". USDA Forest Service. Retrieved 5 August 2015.
  46. Machado, Carlos A.; Robbins, Nancy; Gilbert, M. Thomas; Herre, Edward Allen (April 2005). "Critical Review of Host Specificity and Its Coevolutionary Implications in the Fig-fig-wasp Mutualism". PNAS. 102: 6558–65. Bibcode:2005PNAS..102.6558M. doi:10.1073/pnas.0501840102. PMC   1131861 . PMID   15851680.
  47. Sekar, Sandhya (22 May 2015). "Parasitoid wasps may be the most diverse animal group". BBC. Retrieved 14 February 2018.
  48. Quicke, D. L. J. (1997). Parasitic Wasps. pp. Chapter 8 and passim. ISBN   978-0-412-58350-6.
  49. Sezen, Uzay (24 March 2015). "Giant Ichneumon Wasp (Megarhyssa macrurus) Ovipositing". Nature Documentaries.org. Retrieved 15 June 2015.
  50. Lachaud, Jean-Paul; Pérez-Lachaud, Gabriela (2009). "Impact of natural parasitism by two eucharitid wasps on a potential biocontrol agent ant in southeastern Mexico". Biological Control. 48 (1): 92–99. doi:10.1016/j.biocontrol.2008.09.006.
  51. Williams, David F. (1994). "Biology and importance of two eucharitid parasites of Wasmannia and Solenopsis". Exotic ants : biology, impact, and control of introduced species. Boulder, CO: Westview Press. pp. 104–120. ISBN   978-0-8133-8615-7.
  52. Brues, C. T. (1919). "A New Chalcid-Fly Parasitic on the Australian Bull-Dog Ant". Annals of the Entomological Society of America. 12 (1): 13–21. doi:10.1093/aesa/12.1.13.
  53. Ortolani, I.; Cervo, R. (2009). "Coevolution of daily activity timing in a host-parasite system". Biological Journal of the Linnean Society. 96 (2): 399–405. doi:10.1111/j.1095-8312.2008.01139.x.
  54. Dapporto, L.; Cervo, R.; Sledge, M. F.; Turillazzi, S. (2004). "Rank integration in dominance hierarchies of host colonies by the paper wasp social parasite Polistes sulcifer (Hymenoptera, Vespidae)". Journal of Insect Physiology. 50 (2–3): 217–223. doi:10.1016/j.jinsphys.2003.11.012. PMID   15019524.
  55. O'Neill, Kevin M. (2001). Solitary Wasps: Behavior and Natural History. Cornell University Press. p. 129.
  56. Deslippe, Richard (2010). "Social Parasitism in Ants". Nature Education Knowledge.
  57. Emery, C. (1909). "Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen". Biologisches Centralblatt. 29: 352–362.
  58. Carpenter, James M.; Perera, Estelle P. (16 March 2006). "Phylogenetic Relationships among Yellowjackets and the Evolution of Social Parasitism (Hymenoptera: Vespidae, Vespinae)". American Museum Novitates. 3507 (1): 1–19. doi:10.1206/0003-0082(2006)3507[1:PRAYAT]2.0.CO;2. hdl:2246/5782.
  59. Dvorak, L. (2007). "Parasitism of Dolichovespula norwegica by D. adulterina (Hymenoptera: Vespidae)" (PDF). Silva Gabreta. 13 (1): 65–67.
  60. Fisher, T.W.; Bellows, Thomas S.; Caltagirone, L.E.; Dahlsten, D.L.; Huffaker, Carl B.; Gordh, G. (1999). Handbook of Biological Control: Principles and Applications of Biological Control. Academic Press. p. 455. ISBN   978-0-08-053301-8.
  61. "Biology of the European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae)". Evolutionary Ecology. University of Regensburg. 5 June 2007. Retrieved 20 June 2015.
  62. Edmunds, M. (1974). Defence in Animals: A Survey of Anti-Predator Defences. Longman. pp. 74, 82–83. ISBN   978-0-582-44132-3.
  63. Forshaw, J. & Kemp, A. (1991). Forshaw, Joseph (ed.). Encyclopaedia of Animals: Birds. Merehurst Press. pp. 144–145. ISBN   978-1-85391-186-6.
  64. "Hornet attacks kill dozens in China". The Guardian. 26 September 2013. Retrieved 18 June 2015.
  65. Cocker, Mark; Mabey, Richard (2005). Birds Britannica. London: Chatto & Windus. pp. 113–114. ISBN   978-0-7011-6907-7.
  66. "Roadrunners". Avian Web. Retrieved 3 May 2012.
  67. "Notes & queries: Why do wasps sting people?". The Guardian. 19 August 2009. Retrieved 10 June 2015.
  68. "Wasps". British Pest Control Association. Retrieved 5 August 2015.
  69. "Allergy to Wasp and Bee Stings". Allergy UK. Retrieved 5 August 2015.
  70. "New wasp parasite being studied". The Royal Society of New Zealand. 20 April 2000. Retrieved 15 July 2013.
  71. Moskvitch, Katia (22 January 2013). "Wasps v moths: Biocontrol uses nature against crop pests". BBC. Retrieved 10 June 2015.
  72. Hoddle, Mark. "Encarsia formosa". Cornell University. Retrieved 12 June 2015.
  73. "Aphid Predators". Royal Horticultural Society. Retrieved 5 August 2015.
  74. C.R. Adams; K.M. Bamford; M.P. Early (22 October 2013). Principles of Horticulture. Elsevier. pp. 71–72. ISBN   978-1-4831-4184-8.
  75. "History 1867–1930 London Wasps". Wasps.co.uk. Archived from the original on 22 July 2014.
  76. "Wasps Football Club". Pitcher. Retrieved 5 August 2015.
  77. "Wantirna Wasps Basketball Club" . Retrieved 5 August 2015.
  78. "Alloa Athletic Football Club". Scottish Professional Football League. Retrieved 10 September 2015.
  79. "Diamond and emerald wasp brooch by Fontanna". A La Vieille Russie. Archived from the original on 10 June 2015. Retrieved 10 June 2015. Crown rose diamond and emerald wasp brooch set in silver and gold. By Fontana French, ca. 1875. Width: 3 inches. $46,500
  80. Kunzle, David. "Fashion and Fetishism". Archived from the original on 21 November 2014. Retrieved 17 June 2015.
  81. Klingerman, Katherine Marie (May 2006). Binding Femininity: The Effects of Tightlacing on the Female Pelvis (PDF). University of Vermont (MA Thesis).
  82. "Ancient Greece – Aristophanes – The Wasps" . Retrieved 10 June 2015.
  83. 1 2 Wells, H. G. (1904). Food of the Gods. Macmillan.
  84. Lightbown, Ronald (1989). Sandro Botticelli: Life and Work. Thames and Hudson. pp. 165–168. ISBN   978-0896599314.
  85. Russell, Eric Frank (1957). Wasp. Gollancz Science Fiction. ISBN   978-0-575-07095-0.
  86. Rosenberg, Robin S.; O'Neill, Shannon; McDonald-Smith, Lynne (2013). The Psychology of the Girl with the Dragon Tattoo: Understanding Lisbeth Salander and Stieg Larsson's Millennium Trilogy. BenBella Books. p. 34. ISBN   978-1-936661-35-0.
  87. "Letter 2814 — Darwin, C. R. to Gray, Asa". 22 May 1860. Retrieved 4 May 2011.
  88. 1 2 "USS Wasp. History". United States Navy . Retrieved 1 July 2016.
  89. Colledge, J. J.; Warlow, Ben (2006) [1969]. Ships of the Royal Navy: The Complete Record of all Fighting Ships of the Royal Navy (Rev. ed.). London: Chatham Publishing. ISBN   978-1-86176-281-8. OCLC   67375475.
  90. USS Wasp Veterans (15 June 1999). USS Wasp. Turner Publishing. pp. 8–11. ISBN   978-1-56311-404-5.
  91. Jentz, Thomas L.; Doyle, Hilary Louis (2011). Panzer Tracts No. 23: Panzer Production From 1933 to 1945. Panzer Tracts.
  92. Bishop, Chris (2002). The Encyclopedia of Weapons of World War II. Sterling Publishing. p. 272. ISBN   9781586637620.
  93. James, Derek N. (1991). Westland Aircraft since 1915. Putnam. p. 365. ISBN   978-0-85177-847-1.
  94. "US Air Force Wasp III Fact Sheet". Archived from the original on 23 September 2015. Retrieved 12 June 2015.

Sources