Parasitoid wasp

Last updated

Megarhyssa macrurus (Ichneumonidae), a parasitoid, ovipositing into its host through the wood of a tree. The body of a female is c. 2 inches (50 mm) long, with an ovipositor c. 4 inches (100 mm) long. Ichneumon wasp (Megarhyssa macrurus lunato) (7686081848).jpg
Megarhyssa macrurus (Ichneumonidae), a parasitoid, ovipositing into its host through the wood of a tree. The body of a female is c. 2 inches (50 mm) long, with an ovipositor c. 4 inches (100 mm) long.
Females of the parasitoid wasp Neoneurus vesculus (Braconidae) ovipositing in workers of the ant Formica cunicularia .
Parasitized white cabbage larvae showing wasp larvae exiting its body, spinning cocoons. Playback at double speed. Adult wasps at normal speed.

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.

Contents

Parasitoid wasp species differ in which host life-stage they attack: eggs, larvae, pupae, or adults. They mainly follow one of two major strategies within parasitism: either they are endoparasitic, developing inside the host, and koinobiont, allowing the host to continue to feed, develop, and moult; or they are ectoparasitic, developing outside the host, and idiobiont, paralysing the host immediately. Some endoparasitic wasps of the superfamily Ichneumonoidea have a mutualistic relationship with polydnaviruses, the viruses suppressing the host's immune defenses. [1]

Parasitoidism evolved only once in the Hymenoptera, during the Permian, leading to a single clade called Euhymenoptera, [2] but the parasitic lifestyle has secondarily been lost several times including among the ants, bees, and vespid wasps. As a result, the order Hymenoptera contains many families of parasitoids, intermixed with non-parasitoid groups. The parasitoid wasps include some very large groups, some estimates giving the Chalcidoidea as many as 500,000 species, the Ichneumonidae 100,000 species, and the Braconidae up to 50,000 species. Host insects have evolved a range of defences against parasitoid wasps, including hiding, wriggling, and camouflage markings.

Many parasitoid wasps are considered beneficial to humans because they naturally control agricultural pests. Some are applied commercially in biological pest control, starting in the 1920s with Encarsia formosa to control whitefly in greenhouses. Historically, parasitoidism in wasps influenced the thinking of Charles Darwin. [3]

Parasitoidism

Two strategies found among parasitoidal wasps: Ectoparasites are usually idiobiont, endoparasites koinobiont. Parasitoid Strategies Koinobiont Compared to Idiobiont.svg
Two strategies found among parasitoidal wasps: Ectoparasites are usually idiobiont, endoparasites koinobiont.

Parasitoid wasps range from some of the smallest species of insects to wasps about an inch long. Most females have a long, sharp ovipositor at the tip of the abdomen, sometimes lacking venom glands, and almost never modified into a sting. [4]

Parasitoids can be classified in a variety of ways. They can live within their host's body as endoparasitoids, or feed on it from outside as ectoparasitoids: both strategies are found among the wasps. Parasitoids can also be divided according to their effect on their hosts. Idiobionts prevent further development of the host after initially immobilizing it, while koinobionts allow the host to continue its development while they are feeding upon it; and again, both types are seen in parasitoidal wasps. Most ectoparasitoid wasps are idiobiont, as the host could damage or dislodge the external parasitoid if allowed to move or moult. Most endoparasitoid wasps are koinobionts, giving them the advantage of a host that continues to grow larger and remains able to avoid predators. [4]

Spider wasp (Pompilidae), an idiobiont, carrying a jumping spider she has just paralysed back to her nest, where she will lay an egg on it. IndianSpiderWasp.JPG
Spider wasp (Pompilidae), an idiobiont, carrying a jumping spider she has just paralysed back to her nest, where she will lay an egg on it.

Hosts

Many parasitoid wasps use larval Lepidoptera as hosts, but some groups parasitize different host life stages (egg, larva or nymph, pupa, adult) of nearly all other orders of insects, especially Coleoptera, Diptera, Hemiptera and other Hymenoptera. Some attack arthropods other than insects: for instance, the Pompilidae specialise in catching spiders: these are quick and dangerous prey, often as large as the wasp itself, but the spider wasp is quicker, swiftly stinging her prey to immobilise it. Adult female wasps of most species oviposit into their hosts' bodies or eggs. More rarely, parasitoid wasps may use plant seeds as hosts, such as Torymus druparum . [5]

Some also inject a mix of secretory products that paralyse the host or protect the egg from the host's immune system; these include polydnaviruses, ovarian proteins, and venom. If a polydnavirus is included, it infects the nuclei of host hemocytes and other cells, causing symptoms that benefit the parasite. [6] [7]

Parasitised moth caterpillar.jpg
An apparently healthy moth caterpillar feeds, grows, and molts...
Parasitised moth caterpillar with wasp larvae (cropped).jpg
...but endoparasitic koinobiont wasp larvae eventually fill its body and kill it.

Host size is important for the development of the parasitoid, as the host is its entire food supply until it emerges as an adult; small hosts often produce smaller parasitoids. [8] Some species preferentially lay female eggs in larger hosts and male eggs in smaller hosts, as the reproductive capabilities of males are limited less severely by smaller adult body size. [9]

Hornworm with parasitic wasp cocoons Hornworm with parasitic wasp coccoons.jpg
Hornworm with parasitic wasp cocoons

Some parasitoid wasps mark the host with chemical signals to show that an egg has been laid there. This may both deter rivals from ovipositing, and signal to itself that no further egg is needed in that host, effectively reducing the chances that offspring will have to compete for food and increasing the offspring's survival. [10] [11]

Life cycle

Potter wasp (Eumeninae), an idiobiont, building mud nest; she will then provision it with paralysed insects, on which she lays her eggs; she then seals the nest and provides no further care for her young. Potter Wasp building mud nest near completion.JPG
Potter wasp (Eumeninae), an idiobiont, building mud nest; she will then provision it with paralysed insects, on which she lays her eggs; she then seals the nest and provides no further care for her young.

On or inside the host the parasitoid egg hatches into a larva or two or more larvae (polyembryony). Endoparasitoid eggs can absorb fluids from the host body and grow several times in size from when they were first laid before hatching. The first instar larvae are often highly mobile and may have strong mandibles or other structures to compete with other parasitoid larvae. The following instars are generally more grub-like. Parasitoid larvae have incomplete digestive systems with no rear opening. This prevents the hosts from being contaminated by their wastes. The larva feeds on the host's tissues until ready to pupate; by then the host is generally either dead or almost so. A meconium, or the accumulated wastes from the larva is cast out as the larva transitions to a prepupa. [12] [13] Depending on its species, the parasitoid then may eat its way out of the host or remain in the more or less empty skin. In either case it then generally spins a cocoon and pupates. As adults, parasitoid wasps feed primarily on nectar from flowers. Females of some species will also drink hemolymph from hosts to gain additional nutrients for egg production. [14]

Polydnavirus-wasp mutualism: the virus protects koinobiont wasp eggs and larvae from immune suppression by the host's hemocytes. Pdv-host.svg
Polydnavirus-wasp mutualism: the virus protects koinobiont wasp eggs and larvae from immune suppression by the host's hemocytes.

Mutualism with polydnavirus

Polydnaviruses are a unique group of insect viruses that have a mutualistic relationship with some parasitic wasps. The polydnavirus replicates in the oviducts of an adult female parasitoid wasp. The wasp benefits from this relationship because the virus provides protection for the parasitic larvae inside the host, (i) by weakening the host's immune system and (ii) by altering the host's cells to be more beneficial to the parasite. The relationship between these viruses and the wasp is obligatory in the sense that all individuals are infected with the viruses; the virus has been incorporated in the wasp's genome and is inherited. [15] [16] [17]

Host defenses

Parasitoid wasp (Ichneumonidae) pointing ovipositor at cinnabar moth larva, just after ovipositing. The larva wriggles vigorously to try to avoid the attack. Parasitoid wasp pointing ovipositor at cinnabar moth larva.jpg
Parasitoid wasp (Ichneumonidae) pointing ovipositor at cinnabar moth larva, just after ovipositing. The larva wriggles vigorously to try to avoid the attack.

The hosts of parasitoids have developed several levels of defence. Many hosts try to hide from the parasitoids in inaccessible habitats. They may also get rid of their frass (body wastes) and avoid plants that they have chewed on as both can signal their presence to parasitoids hunting for hosts. The egg shells and cuticles of the potential hosts are thickened to prevent the parasitoid from penetrating them. Hosts may use behavioral evasion when they encounter an egg laying female parasitoid, like dropping off the plant they are on, twisting and thrashing so as to dislodge or kill the female and even regurgitating onto the wasp to entangle it. The wriggling can sometimes help by causing the wasp to "miss" laying the egg on the host and instead place it nearby. Wriggling of pupae can cause the wasp to lose its grip on the smooth hard pupa or get trapped in the silk strands. Some caterpillars even bite the female wasps that approach them. Some insects secrete poisonous compounds that kill or drive away the parasitoid. Ants that are in a symbiotic relationship with caterpillars, aphids or scale insects may protect them from attack by wasps. [18] [19]

Parasitoid wasps are vulnerable to hyperparasitoid wasps. Some parasitoid wasps change the behavior of the infected host, causing them to build a silk web around the pupae of the wasps after they emerge from its body to protect them from hyperparasitoids. [20]

Hosts can kill endoparasitoids by sticking haemocytes to the egg or larva in a process called encapsulation. [21] In aphids, the presence of a particular species of γ-3 Pseudomonadota makes the aphid relatively immune to their parasitoid wasps by killing many of the eggs. As the parasitoid's survival depends on its ability to evade the host's immune response, some parasitoid wasps have developed the counterstrategy of laying more eggs in aphids that have the endosymbiont, so that at least one of them may hatch and parasitize the aphid. [22] [23]

Certain caterpillars eat plants that are toxic to both themselves and the parasite to cure themselves. [24] Drosophila melanogaster larvae also self-medicate with ethanol to treat parasitism. [25] D. melanogaster females lay their eggs in food containing toxic amounts of alcohol if they detect parasitoid wasps nearby. The alcohol protects them from the wasps, at the cost of retarding their own growth. [26]

Evolution and taxonomy

Evolution

Based on genetic and fossil analysis, parasitoidism has evolved only once in the Hymenoptera, during the Permian, leading to a single clade. All parasitoid wasps are descended from this lineage. The narrow-wasted Apocrita emerged during the Jurassic. [27] [28] [29] [30] The Aculeata, which includes bees, ants, and parasitoid spider wasps, evolved from within the Apocrita; it contains many families of parasitoids, though not the Ichneumonoidea, Cynipoidea, and Chalcidoidea. The Hymenoptera, Apocrita, and Aculeata are all clades, but since each of these contains non-parasitic species, the parasitoid wasps, formerly known as the Parasitica, do not form a clade on their own. [30] [31] The common ancestor in which parasitoidism evolved lived approximately 247 million years ago and was previously believed to be an ectoparasitoid wood wasp that fed on wood-boring beetle larvae. Species similar in lifestyle and morphology to this ancestor still exist in the Ichneumonoidea. [32] [33] However, recent molecular and morphological analysis suggests this ancestor was endophagous, meaning it fed from within its host. [30] A significant radiation of species in the Hymenoptera occurred shortly after the evolution of parasitoidy in the order and is thought to have been a result of it. [31] [33] The evolution of a wasp waist, a constriction in the abdomen of the Apocrita, contributed to rapid diversification as it increased maneuverability of the ovipositor, the organ off the rear segment of the abdomen used to lay eggs. [34]

The phylogenetic tree gives a condensed overview of the positions of parasitoidal groups (boldface), amongst groups (italics) like the Vespidae which have secondarily abandoned the parasitoid habit. The approximate numbers of species estimated to be in these groups, often much larger than the number so far described, is shown in parentheses, with estimates for the most populous also shown in boldface, like "(150,000)". Not all species in these groups are parasitoidal: for example, some Cynipoidea are phytophagous.

Hymenoptera

Sawflies Xyelapusilla.jpg

parasitoidism

Orussoidea (parasitoid wood wasps, 85) Orussus coronatus.jpg

Apocrita

Ichneumonoidea (150,000) Ichneumon wasp (Ichneumonidae sp) female (cropped).jpg

Cynipoidea (3,000) Cynips sp beentree.jpg

other Superfamilies

Aculeata

Chrysididae (jewel wasps, 3000) Chrysididae jewel wasp.jpg

Vespidae (wasps, hornets, 5000) European wasp white bg.jpg

Mutillidae (velvet ants, 3000) Velvet ant (Mutillidae) (25808496580) (cropped).jpg

Pompilidae (spider wasps, 5000) Spider Wasp (cropped).JPG

other families

Scoliidae (560) Black-Flower-Wasp.jpg

Formicidae (ants, 22,000) Meat eater ant feeding on honey02.jpg

Apoidea

Sphecidae (700) Specimen of Podalonia tydei (Le Guillou, 1841).jpg

Bembicinae (1800) Bembix sp.jpg

other families

Pemphredoninae (aphid wasps, 1000) Pemphredon sp.-pjt1 (cropped).jpg

Philanthinae (1100) Dorsal view cerceris.jpg

Anthophila (bees, 22,000) Apis mellifera (in flight) (cropped).jpg

stinging
wasp waist
evolved once

Taxonomy

Trissolcus (family Platygastridae) on Chinavia eggs Trissolcus on Chinavia eggs.jpg
Trissolcus (family Platygastridae) on Chinavia eggs
Housefly pupae killed by parasitoid wasp larvae (probably Pteromalidae). Each pupa has one hole through which a single adult wasp has emerged after feeding on the housefly larva. Housefly pupae killed by wasp larvae.jpg
Housefly pupae killed by parasitoid wasp larvae (probably Pteromalidae). Each pupa has one hole through which a single adult wasp has emerged after feeding on the housefly larva.

The parasitoid wasps are paraphyletic since the ants, bees, and non-parasitic wasps such as the Vespidae are not included, and there are many members of mainly parasitoidal families which are not themselves parasitic. Listed are Hymenopteran families where most members have a parasitoid lifestyle. [35]

Symphyta:

Apocrita:

Interactions with humans

Biological pest control

Encarsia formosa, an endoparasitic aphelinid wasp, bred commercially to control whitefly in greenhouses Encarsia formosa, an endoparasitic wasp, is used for whitefly control.jpg
Encarsia formosa , an endoparasitic aphelinid wasp, bred commercially to control whitefly in greenhouses
Trioxys complanatus, (Aphidiinae) ovipositing into a spotted alfalfa aphid, a commercial pest in Australia. CSIRO ScienceImage 2357 Spotted alfalfa aphid being attacked by parasitic wasp.jpg
Trioxys complanatus, (Aphidiinae) ovipositing into a spotted alfalfa aphid, a commercial pest in Australia.

Parasitoid wasps are considered beneficial as they naturally control the population of many pest insects. They are widely used commercially (alongside other parasitoids such as tachinid flies) for biological pest control, for which the most important groups are the ichneumonid wasps, which prey mainly on caterpillars of butterflies and moths; braconid wasps, which attack caterpillars and a wide range of other insects including greenfly; chalcidoid wasps, which parasitise eggs and larvae of greenfly, whitefly, cabbage caterpillars, and scale insects. [37]

One of the first parasitoid wasps to enter commercial use was Encarsia formosa , an endoparasitic aphelinid. It has been used to control whitefly in greenhouses since the 1920s. Use of the insect fell almost to nothing, replaced by chemical pesticides by the 1940s. Since the 1970s, usage has revived, with renewed usage in Europe and Russia. [38] In some countries, such as New Zealand, it is the primary biological control agent used to control greenhouse whiteflies, particularly on crops such as tomato, a particularly difficult plant for predators to establish on. [39]

Commercially, there are two types of rearing systems: short-term seasonal daily output with high production of parasitoids per day, and long-term year-round low daily output with a range in production of 4–1000 million female parasitoids per week, to meet demand for suitable parasitoids for different crops. [40]

In culture

Parasitoid wasps influenced the thinking of Charles Darwin. [lower-alpha 2] In an 1860 letter to the American naturalist Asa Gray, Darwin wrote: "I cannot persuade myself that a beneficent and omnipotent God would have designedly created parasitic wasps with the express intention of their feeding within the living bodies of Caterpillars." [3] The palaeontologist Donald Prothero notes that religiously-minded people of the Victorian era, including Darwin, were horrified by this instance of evident cruelty in nature, particularly noticeable in the Ichneumonidae. [42]

Notes

  1. ↑ Trioxys complanatus has been introduced to Australia to control the spotted alfalfa aphid. [36]
  2. ↑ Darwin mentions "parasitic" wasps in On the Origin of Species , Chapter 7, page 218. [41]

Related Research Articles

<span class="mw-page-title-main">Hymenoptera</span> Order of insects comprising sawflies, wasps, bees, and ants

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. Many of the species are parasitic. Females typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. This ovipositor is often modified into a stinger. The young develop through holometabolism — that is, they have a wormlike larval stage and an inactive pupal stage before they reach adulthood.

<span class="mw-page-title-main">Parasitism</span> Relationship between species where one organism lives on or in another organism, causing it harm

Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria, sleeping sickness, and amoebic dysentery; animals such as hookworms, lice, mosquitoes, and vampire bats; fungi such as honey fungus and the agents of ringworm; and plants such as mistletoe, dodder, and the broomrapes.

<span class="mw-page-title-main">Biological pest control</span> Controlling pests using other organisms

Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also involves an active human management role. It can be an important component of integrated pest management (IPM) programs.

<span class="mw-page-title-main">Parasitoid</span> Organism that lives with its host and kills it

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. 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.

<span class="mw-page-title-main">Tachinidae</span> Family of insects

The Tachinidae are a large and variable family of true flies within the insect order Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in this family commonly are called tachinid flies or simply tachinids. As far as is known, they all are protelean parasitoids, or occasionally parasites, of arthropods, usually other insects. The family is known from many habitats in all zoogeographical regions and is especially diverse in South America.

<span class="mw-page-title-main">Sawfly</span> Suborder of insects

Sawflies are wasp-like insects that are in 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 in the suborder, with about 7,000 known species; in the entire suborder, there are 8,000 described species in more than 800 genera. Symphyta is paraphyletic, consisting of several basal groups within the order Hymenoptera, each one rooted inside the previous group, ending with the Apocrita which are not sawflies.

<span class="mw-page-title-main">Apocrita</span> Suborder of insects containing wasps, bees, and ants

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 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.

<span class="mw-page-title-main">Ichneumonoidea</span> Superfamily of wasps

The superfamily Ichneumonoidea contains one extinct and three extant families, including 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.

<span class="mw-page-title-main">Ichneumonidae</span> Family of wasps

The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25,000 species described as of 2016. However, this likely represents less than a quarter of their true richness as reliable estimates are lacking, along with much of the most basic knowledge about their ecology, distribution, and evolution. It is estimated that there are more species in this family than there are species of birds and mammals combined. Ichneumonid wasps, with very few exceptions, attack the immature stages of holometabolous insects and spiders, eventually killing their hosts. They thus fulfill an important role as regulators of insect populations, both in natural and semi-natural systems, making them promising agents for biological control.

<span class="mw-page-title-main">Hyperparasite</span> Parasite of another parasite

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 other insect orders, the Diptera and Coleoptera (beetles). Seventeen families in Hymenoptera and a few species of Diptera and Coleoptera are hyperparasitic. Hyperparasitism developed from primary parasitism, which evolved in the Jurassic period in the Hymenoptera. Hyperparasitism intrigues entomologists because of its multidisciplinary relationship to evolution, ecology, behavior, biological control, taxonomy, and mathematical models.

<i>Polydnaviriformidae</i> Family of viruses

Polydnaviriformidae ( PDV) is a family of insect viriforms; members are known as polydnaviruses. There are two genera in the family: Bracoform and Ichnoviriform. Polydnaviruses form a symbiotic relationship with parasitoid wasps. Ichnoviriforms (IV) occur in Ichneumonid wasps and Bracoviriforms (BV) in Braconid wasps. The larvae of wasps in both of those groups are themselves parasitic on Lepidoptera, and the polydnaviruses are important in circumventing the immune response of their parasitized hosts. Little or no sequence homology exists between BV and IV, suggesting that the two genera have been evolving independently for a long time.

<span class="mw-page-title-main">Braconidae</span> Family of wasps

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 estimated a total between 30,000 and 50,000, and another provided a narrower estimate between 42,000 and 43,000 species.

<span class="mw-page-title-main">Trigonalidae</span> Family of wasps

Trigonalidae is a family of parasitic wasps in the suborder Apocrita. They are the only living members of the superfamily Trigonaloidea. Trigonalidae are divided into 2 subfamilies; Orthogonalinae and Trigonalinae. These wasps are extremely rare, but surprisingly diverse, with over 90 species in 16 genera, and are known from all parts of the world. It is possibly the sister group to all Aculeata.

<i>Cotesia congregata</i> Species of wasp

Cotesia congregata is a parasitoid wasp of the genus Cotesia. The genus is particularly noted for its use of polydnaviruses. Parasitoids are distinct from true parasites in that a parasitoid will ultimately kill its host or otherwise sterilize it.

<i>Glyptapanteles</i> Genus of wasps

Glyptapanteles is a genus of endoparasitoid wasps found in all continents, except Antarctica. The larvae of Glyptapanteles species are able to manipulate their hosts into serving as bodyguards.

<span class="mw-page-title-main">Wasp</span> Group of insects

A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. The wasps do not constitute a clade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade Aculeata can sting their prey.

<span class="mw-page-title-main">Microgastrinae</span> Subfamily of wasps

Microgastrinae is a subfamily of braconid wasps, encompassing almost 3,000 described species, with an estimated 30,000–50,000 total species. This makes it one of the richest subfamilies with the most species of parasitoid wasps.

<span class="mw-page-title-main">Ibaliidae</span> Family of wasps

The Ibaliidae are a small family of the hymenopteran superfamily Cynipoidea. Ibaliidae differ from most of the cynipoids by the larvae being parasitoids on other wasp larvae in the group Siricidae. The Ibaliidae comprise three extant genera of fairly large wasps, with a total of 20 species, and is a sister group to the rest of the cynipoids except the small subfamily Austrocynipidae.

<i>Zatypota percontatoria</i> Species of wasp

Zatypota percontatoria is a species of parasitoid wasps that is part of the order Hymenoptera and the family Ichneumonidae responsible for parasitizing arachnids, specifically those of the family Theridiidae.

<i>Netelia producta</i> Species of wasp

Netelia producta is a species of ichneumonid wasp in the subfamily Tryphoninae found in Australia.

References

  1. ↑ Herniou, Elisabeth A.; Huguet, Elisabeth; Thézé, Julien; Bézier, Annie; Periquet, Georges; Drezen, Jean-Michel (2013-09-19). "When parasitic wasps hijacked viruses: genomic and functional evolution of polydnaviruses". Phil. Trans. R. Soc. B. 368 (1626): 20130051. doi:10.1098/rstb.2013.0051. PMC   3758193 . PMID   23938758.
  2. ↑ Zhang, Qi; Kopylov, Dmitry S.; Rasnitsyn, Alexandr P.; Zheng, Yan; Zhang, Haichun (November 2020). Smith, Andrew (ed.). "Burmorussidae, a new family of parasitic wasps (Insecta, Hymenoptera) from mid-Cretaceous Burmese amber". Papers in Palaeontology. 6 (4): 593–603. doi:10.1002/spp2.1312. ISSN   2056-2802. S2CID   219039881.
  3. 1 2 "Letter 2814 — Darwin, C. R. to Gray, Asa, 22 May [1860]" . Retrieved 2011-04-05.
  4. 1 2 Gullan, P. J.; Cranston, P. S. (2010). The Insects: An Outline of Entomology (5th ed.). Wiley. pp. 362–370. ISBN   978-1-118-84615-5.
  5. ↑ Cushman R. A. (1916). "Syntomaspis druparum, the apple-seed chalcid". Journal of Agricultural Research. 7: 487–502.
  6. ↑ Miller, Lois K.; Ball, Laurence Andrew (1998). The insect viruses. Springer. ISBN   978-0-306-45881-1.
  7. ↑ Strand, Michael R.; Burke, Gaelen R. (May 2015). "Polydnaviruses: From discovery to current insights". Virology. 479–480: 393–402. doi:10.1016/j.virol.2015.01.018. PMC   4424053 . PMID   25670535.
  8. ↑ Cohen, J. E.; Jonsson, T.; Muller, C. B.; Godfray, H. C. J.; Savage, V. M. (12 January 2005). "Body sizes of hosts and parasitoids in individual feeding relationships". Proceedings of the National Academy of Sciences. 102 (3): 684–689. Bibcode:2005PNAS..102..684C. doi: 10.1073/pnas.0408780102 . PMC   545575 . PMID   15647346. Open Access logo PLoS transparent.svg
  9. ↑ Jones, W. Thomas (1982). "Sex Ratio and Host Size in a Parasitoid Wasp". Behavioral Ecology and Sociobiology. 10 (3): 207–210. doi:10.1007/bf00299686. JSTOR   4599484. S2CID   28101144.
  10. ↑ Couchoux, Christelle; Seppä, Perttu; van Nouhuys, Saskya (2015). "Behavioural and genetic approaches to evaluate the effectiveness of deterrent marking by a parasitoid wasp". Behaviour. 152 (9): 1257–1276. doi:10.1163/1568539X-00003277.
  11. ↑ Nufio, César R.; Papaj, Daniel R. (2001). "Host marking behavior in phytophagous insects and parasitoids". Entomologia Experimentalis et Applicata. 99 (3): 273–293. doi: 10.1046/j.1570-7458.2001.00827.x . Open Access logo PLoS transparent.svg
  12. ↑ Paladino, Leonela Zusel Carabajal; Papeschi, Alba Graciela; Cladera, Jorge Luis (January 2010). "Immature stages of development in the parasitoid wasp, Diachasmimorpha longicaudata". Journal of Insect Science. 10 (1): 56. doi:10.1673/031.010.5601. PMC   3014816 . PMID   20569133. Open Access logo PLoS transparent.svg
  13. ↑ "Cotesia congregata - a parasitoid wasp". entnemdept.ufl.edu. Retrieved 2017-10-02.
  14. ↑ Jervis, M. A.; Kidd, N. A. C (November 1986). "Host-Feeding Strategies in Hymenopteran Parasitoids". Biological Reviews. 61 (4): 395–434. doi:10.1111/j.1469-185x.1986.tb00660.x. S2CID   84430254.
  15. ↑ Fleming, J. G.; Summers, M. D. (1991). "Polydnavirus DNA is integrated in the DNA of its parasitoid wasp host". PNAS. 88 (21): 9770–9774. Bibcode:1991PNAS...88.9770F. doi: 10.1073/pnas.88.21.9770 . PMC   52802 . PMID   1946402.
  16. ↑ Webb, Bruce A. (1998). "Polydnavirus Biology, Genome Structure, and Evolution". In Miller, L.K.; Ball, L.A. (eds.). The Insect Viruses. Springer, Boston, MA. pp. 105–139. doi:10.1007/978-1-4615-5341-0_5. ISBN   9781461374374.
  17. ↑ Roossinck, M. J. (2011). "Changes in population dynamics in mutualistic versus pathogenic viruses". Viruses. 3 (12): 12–19. doi: 10.3390/v3010012 . PMC   3187592 . PMID   21994724. Open Access logo PLoS transparent.svg
  18. ↑ Strand, M. R.; Pech, L. L. (1995). "Immunological Basis for Compatibility in Parasitoid-Host Relationships". Annual Review of Entomology. 40: 31–56. doi:10.1146/annurev.en.40.010195.000335. PMID   7810989.
  19. ↑ Gross, P. (1993). "Insect Behavioral and Morphological defences Against Parasitoids". Annual Review of Entomology. 38: 251–273. doi:10.1146/annurev.en.38.010193.001343.
  20. ↑ Tanaka, S.; Ohsaki, N. (2006). "Behavioral manipulation of host caterpillars by the primary parasitoid wasp Cotesia glomerata (L.) to construct defensive webs against hyperparasitism". Ecological Research. 21 (4): 570. doi:10.1007/s11284-006-0153-2. S2CID   23457678.
  21. ↑ Pennacchio, Francesco; Caccia, Silvia; Digilio, Maria Cristina (December 2014). "Host regulation and nutritional exploitation by parasitic wasps". Current Opinion in Insect Science. 6: 74–79. doi:10.1016/j.cois.2014.09.018. ISSN   2214-5745. PMID   32846685.
  22. ↑ Oliver, K. M.; Russell, J. A.; Moran, N. A.; Hunter, M. S. (2003). "Facultative bacterial symbionts in aphids confer resistance to parasitic wasps". Proceedings of the National Academy of Sciences. 100 (4): 1803–7. Bibcode:2003PNAS..100.1803O. doi: 10.1073/pnas.0335320100 . PMC   149914 . PMID   12563031.
  23. ↑ Oliver, K. M.; Noge, K.; Huang, E. M.; Campos, J. M.; Becerra, J. X.; Hunter, M. S. (2012). "Parasitic wasp responses to symbiont-based defence in aphids". BMC Biology. 10: 11. doi: 10.1186/1741-7007-10-11 . PMC   3312838 . PMID   22364271.
  24. ↑ Singer, M. S.; Mace, K. C.; Bernays, E. A. (2009). May, Robin Charles (ed.). "Self-Medication as Adaptive Plasticity: Increased Ingestion of Plant Toxins by Parasitized Caterpillars". PLOS ONE. 4 (3): e4796. Bibcode:2009PLoSO...4.4796S. doi: 10.1371/journal.pone.0004796 . PMC   2652102 . PMID   19274098.
  25. ↑ Milan, N. F.; Kacsoh, B. Z.; Schlenke, T. A. (2012). "Alcohol Consumption as Self-Medication against Blood-Borne Parasites in the Fruit Fly". Current Biology. 22 (6): 488–493. doi:10.1016/j.cub.2012.01.045. PMC   3311762 . PMID   22342747.
  26. ↑ Kacsoh, B. Z.; Lynch, Z. R.; Mortimer, N. T.; Schlenke, T. A. (2013). "Fruit Flies Medicate Offspring After Seeing Parasites". Science. 339 (6122): 947–50. Bibcode:2013Sci...339..947K. doi:10.1126/science.1229625. PMC   3760715 . PMID   23430653.
  27. ↑ Branstetter, Michael G.; Danforth, Bryan N.; Pitts, James P.; Faircloth, Brant C.; Ward, Philip S.; Buffington, Matthew L.; Gates, Michael W.; Kula, Robert R.; Brady, Seán G. (2017). "Phylogenomic Insights into the Evolution of Stinging Wasps and the Origins of Ants and Bees". Current Biology. 27 (7): 1019–1025. doi: 10.1016/j.cub.2017.03.027 . PMID   28376325. Open Access logo PLoS transparent.svg
  28. ↑ Schulmeister, S. (2003). "Simultaneous analysis of basal Hymenoptera (Insecta), introducing robust-choice sensitivity analysis". Biological Journal of the Linnean Society. 79 (2): 245–275. doi: 10.1046/j.1095-8312.2003.00233.x . Open Access logo PLoS transparent.svg
  29. ↑ Schulmeister, S. "Symphyta" . Retrieved 28 November 2016.
  30. 1 2 3 Peters, Ralph S.; Krogmann, Lars; Mayer, Christoph; Donath, Alexander; Gunkel, Simon; Meusemann, Karen; Kozlov, Alexey; Podsiadlowski, Lars; Petersen, Malte (2017). "Evolutionary History of the Hymenoptera". Current Biology. 27 (7): 1013–1018. doi: 10.1016/j.cub.2017.01.027 . hdl: 2434/801122 . PMID   28343967.
  31. 1 2 Heraty, John; Ronquist, Fredrik; Carpenter, James M.; Hawks, David; Schulmeister, Susanne; Dowling, Ashley P.; Murray, Debra; Munro, James; Wheeler, Ward C. (2011). "Evolution of the hymenopteran megaradiation" (PDF). Molecular Phylogenetics and Evolution. 60 (1): 73–88. doi:10.1016/j.ympev.2011.04.003. PMID   21540117.
  32. ↑ Pennacchio, Francesco; Strand, Michael R. (January 2006). "Evolution of developmental strategies in parasitic hymenoptera". Annual Review of Entomology. 51 (1): 233–258. doi:10.1146/annurev.ento.51.110104.151029. PMID   16332211.
  33. 1 2 Whitfield, James B. (2003). "Phylogenetic Insights into the Evolution of Parasitism in Hymenoptera". Advances in Parasitology. 54: 69–100. doi:10.1016/S0065-308X(03)54002-7. ISBN   978-0-12-031754-7. PMID   14711084.
  34. ↑ Peters, Ralph S.; Krogmann, Lars; Mayer, Christoph; Donath, Alexander; Gunkel, Simon; Meusemann, Karen; Kozlov, Alexey; Podsiadlowski, Lars; Petersen, Malte (April 2017). "Evolutionary History of the Hymenoptera". Current Biology. 27 (7): 1013–1018. doi: 10.1016/j.cub.2017.01.027 . hdl: 2434/801122 . ISSN   0960-9822. PMID   28343967.
  35. ↑ Henri, Goulet; Theodore), Huber, John T. (John; Branch, Canada. Agriculture Canada. Research (1993). Hymenoptera of the world : an identification guide to families. Centre for Land and Biological Resources Research. ISBN   978-0660149332. OCLC   28024976.{{cite book}}: CS1 maint: multiple names: authors list (link)
  36. ↑ Wilson, C. G.; Swincer, D. E.; Walden, K. J. (1982). "The Introduction of Trioxys Complanatus Quilis (Hymenoptera: Aphidiidae), an Internal Parasite of the Spotted Alfalfa Aphid, into South Australia". Australian Journal of Entomology. 21 (1): 13–27. doi:10.1111/j.1440-6055.1982.tb01758.x. S2CID   84996305.
  37. ↑ "Parasitoid Wasps (Hymenoptera)". University of Maryland. Retrieved 6 June 2016.
  38. ↑ Hoddle, M. S.; Van Driesche, R. G.; Sanderson, J. P. (1998). "Biology and Use of the Whitefly Parasitoid Encarsia Formosa". Annual Review of Entomology. 43 (1): 645–669. doi:10.1146/annurev.ento.43.1.645. PMID   15012401.
  39. ↑ "Enforce for Greenhouse Whitefly Control". New Zealand: Bioforce Limited. Retrieved 26 January 2024.
  40. ↑ Smith, S. M. (1996). "Biological control with Trichogramma: advances, successes, and potential of their use". Annual Review of Entomology. 41: 375–406. doi:10.1146/annurev.en.41.010196.002111. PMID   15012334.
  41. ↑ On the Origin of Species , Chapter 7, page 218.
  42. ↑ Prothero, Donald R. (2017). Evolution: What the Fossils Say and Why It Matters. Columbia University Press. pp. 84–86. ISBN   978-0-231-54316-3.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.