Zatypota percontatoria | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Ichneumonidae |
Genus: | Zatypota |
Species: | Z. percontatoria |
Binomial name | |
Zatypota percontatoria Mueller, 1776 [1] | |
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. [2] [3]
The parasitoid that has been recently studied for its mechanism of parasitism. It reproduces by laying eggs within the abdomen of a spider by oviposition, slowly developing as an ectoparasitic koinobiont that will eventually kill its host. The wasp larva protects itself during the parasitism process through neuroparasitism affecting and controlling the behavior of the spider to the extent that they are unable to attack the larva. [3] The mechanism is still fairly unknown, but it is believed that it involves hormones and/or polydnaviruses. [4] The larva will influence the spider to the extent that it will build a web/cocoon for the pupal stage of the parasitoid wasp. Z. percontatoria wasps are distributed worldwide within several different countries, mostly within terrestrial locations that are rich with tree species preferring woodlands. [5]
Members of the order Hymenoptera are both parasitic and non-parasitic as some families continued to evolve as parasites while others lost parasitism as a trait. The family Ichneumonidae is one of the largest in the animal world, containing over 20,000 species that have been described or identified in some way. Although it is difficult to pin down exactly how many species there are within the family, there have been estimates that range from 60,000 to over 100,000. [6]
Z. percontatoria is symmetrical with the females being generally larger. The flagellum, which are the antenna on the wasp's head excluding some of the base section, has around 18-20 segments in the female. The head, as well as most parts of the body, is covered with small hairs known as pubescence. This is especially long around the mouth of the wasp. The area covering the head of the wasp has a grainy exterior. The eyes are bare to the environment and the head appears to be convex upon further inspection. However, the head converges downward when viewed from the lateral side. In the jaw or mandible, the upper tooth is longer and wider than the lower tooth. The head is black with yellowish white sections with the flagellum being light brown. [7]
The mesosoma is rather rigid in terms of flexibility. There can be wrinkles seen increasing closer to the different ends of wasp's mesosoma. The pubescence are shorter on the mesosoma compared to the head. The hairs are short so that the mesosoma appears to be smooth and bare. The mesosoma is a darkened and reddish brown, with yellowish white sections. The legs are a pale brown. [7]
The wasp has two pairs of membranous wings, which are held to the body by small hooks. Veins run through the wings. The fore wings are larger than the hind wings. The average female fore wing length is 3.0-3.8 mm. The legs are slender and held together by segments. The metasoma goes back breaking into five different parts. The wasp will get skinnier the closer to the caudal end. The ovipositor is black and brown. The ovipositor has a strong, slender base that thickens to a tapered point medially used for quick insertion. The wings are brown and clear in the sections that are mainly membranous. The metasoma like the mesosoma is a darkened and reddish brown. [7]
Male Z. percontatoria wasps are similar to females except that they are smaller and darker in color. The flagellum for the males contain 18 segments and they have an average fore wing length of 2.9 mm. [7]
Larvae are usually a pale yellowish white. There are small differences between the parasitic wasps as they are distributed around the world, but this description mainly refers to the Z. percontatoria native to Japan. [7]
The lifecycle begins when a female wasp is able to successfully oviposit an egg within the abdomen of a spider within the family Theridiidae. Z. percontatoria is a unique species in that although the wasps are koinobionts, they are one of the rare cases of being ectoparasitic as well. Although the egg begins inside of the spider, eventually it will hatch into a larva. It does not use an immobilization or paralyzing agent, so the spider develops as the egg/larva develops. As time progresses, the larva bursts out of the spider and continues as an ectoparasitoid. The larva will develop using the nutrients from the spider's hemolymph. When the larva moves out of the abdomen, it remains connected to the hemolymph for nutrients as an ectoparasitoid. How large the larva gets ultimately depends on the size and nutrient availability of the spider. [8]
The spider may continue to forage and take part in activities that ultimately increase the fitness of the larva. However, Z. percontatoria has to be careful because it have to protect itself and the spider from their predators. The spider when it is parasitized is not able to defend itself as it could before it was parasitized. The mechanism is unknown as to how the larva is able to alter the behavior of the spider, but there are multiple hypotheses. One is that it may be controlling the central nervous system directly through the influence of neurons, although it is unlikely as Z. percontatoria connects to the hemolymph not directly associated with the nervous system.
The more likely of the hypotheses is that Z. percontatoria is not controlling the spider directly but influencing it through hormones in the endocrine system by activating certain pathways of the spider. The spider does not exhibit any behaviors that are outside of its normal scope, however the usual timing of the behaviors is altered. This is because the larva is using the spider's behaviors to increase its fitness and chance of survival. [9] By increasing the levels of hormones that promote certain actions, the larva alters the normal behavior of the spider to its advantage. The larva also has a mutualistic relationship with polydnaviruses that suppresses the immune system preventing it from making any counter-action. [4] However, if the larva is ever removed the spider returns to normal control and behaviors. [10] The longer that a larva is able to remain on a spider, the longer it will take that spider to recover if the larva is taken off. [11] This action also supports the hypothesis that the change in behavior is most likely the result of an accumulation of hormones stimulated or released by the larva. [12]
Before the spider is fully able to reach adulthood, the larva enters the pupal stage. This stage is the most vulnerable state of the parasitoid wasp's lifecycle. This is the part of the lifecycle that explains why Z. percontatoria prefers the web-building spiders of the family Theridiidae. The larva will begin influencing the spider to build a web. Scientists hypothesize that the larva is somehow able to increase levels of the hormone that encourages behaviors that the spider would conduct before molting or ecdysis. This hormone is elevated levels of 20-OH-ecysone. [13] The web is not the typical type used for capturing prey, but resembles a web similar to one that it would build before overwintering. The web can be seen to be shorter for a stronger build, thickened webs that won't break, and a reinforced frame. [10] The web is stronger, defense-based, and depending on the spider will create a cocoon that will house the larva. This will aid the fitness of the larva when it enters the pupal stage by creating a camouflage, and protecting it from predators and the outside conditions of the environment. [13] [14]
When the web is done, the larva kills the spider host and consumes its body. The larva is then ready to enter the cocoon where a metamorphosis takes place. [15] The pupa will emerge as an adult wasp, usually female as they are able to reproduce starting the cycle again. As an adult, the wasps mainly feed on nectar. Some larva will overwinter during winter. The active months of Z. percontatoria are between March and November varying depending on the climate of the geographical location. [2] [3]
Z. percontatoria, like many parasitoids, is host specific with a narrow range. Z. percontatoria parasitizes members of the arthropod family Theridiidae. The specific host species for Z. percontatoria in a given area varies depending the species of the family Theridiidae that are abundant within that area. Spiders as hosts are able hunters making them difficult prey to parasitize. An additional factor adding to the level of difficulty in ovipositing is that the spider is usually the same size if not larger than the wasp. The wasp has to be big enough to be able to handle itself in a battle with the spider and precise enough to oviposit an egg in the abdomen of the spider. If the wasp is not careful, it will get injured and/or be unsuccessful in an attempt to parasitize the spider. Z. percontatoria prefers female spiders because they forage more than males leading to increased nutrients that the host can provide. The wasps increase their chances by usually parasitizing juvenile spiders that are less able to defend themselves against the wasp. [16] Juvenile spiders are also preferred due to the koinobiont lifecycle of the wasp. On average, they will spend around 34 days parasitizing a host. [17] So adult spiders are better able to defend themselves and may die before parasitism is complete. On the other hand, infant spiders cannot handle the stress of the parasitism. However, Z. percontatoria wasps are adaptable and will change its habits based on the climate, size of the spider population, and the abundance of spiders. As parasitoids, the wasps will keep the spider population stable as to promote a better fitness for itself. There is a correlation between the number of parasitized spider and the abundance of the population. The more spiders there are, the higher the number of spiders that are parasitized. Warmer climates promote faster lifecycles and colder climates will take longer for the lifecycle to be complete. Parasitic wasps are unable to reproduce and survive at any temperatures at or below 5 degrees Celsius. [17]
The rate of parasitism within a given spider population is low, usually around 1%. [18] Spiders will defend themselves and without an immobilization agent are difficult to parasitize. Mortality rates are highest in the egg and larval stages of the parasitoid wasp as well so they may die before they get the chance to finish their lifecycle. Also unless they are able successfully camouflage or protect themselves, predators in the environment can take advantage and kill them as well. With its parasitoid lifecycle taking a great deal of time, there is much that could go wrong resulting in mortality. These are fairly recent studies that need further research to back up and confirm. [3]
Z. percontatoria is Holarctic in distribution. [19] The parasitoid wasps prefer terrestrial woodland locations with a high population of trees. The parasitoid wasp is found in equal distribution in the understory or in the leaves of the canopy, as web-building spiders can be found in both levels. An abundance of plant species and numbers promote species diversity including those of parasitoids and hosts. Parasitoid wasps, although located in many places in the world, are rarely the dominating population in terms of influence or numbers. [20] [5]
Evolution is a process where traits that increase the fitness of an organism are favored and therefore natural selection keeps them in the lineage creating speciation. So for parasites, it is most likely that the way that they harm and feed off another organism was beneficial to them and caused an evolutionary change. The environment most likely could not provide completely for the organism, forcing it to prey on another organism for nutrients forming what would be the basis for parasitism. The complexity of the strategies make it difficult to pin down exactly how they were created. [21] In the order Hymenoptera, parasitoidism only evolved once. This created the bee, ant, and parasitoid wasp species that followed. Parasitism was lost in species that it did not benefit or add to their fitness. For the parasitoid wasps, evolution added a wasp waist and the ability to sting which formed the basis of the mechanisms that Z. percontatoria employs. [22] [4]
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.
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.
Kleptoparasitism is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, such as when food is scarce or when victims are abundant. Many kleptoparasites are arthropods, especially bees and wasps, but including some true flies, dung beetles, bugs, and spiders. Cuckoo bees are specialized kleptoparasites which lay their eggs either on the pollen masses made by other bees, or on the insect hosts of parasitoid wasps. They are an instance of Emery's rule, which states that insect social parasites tend to be closely related to their hosts. The behavior occurs, too, in vertebrates including birds such as skuas, which persistently chase other seabirds until they disgorge their food, and carnivorous mammals such as spotted hyenas and lions. Other species opportunistically indulge in kleptoparasitism.
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.
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.
Hymenoepimecis argyraphaga is a Costa Rican parasitoid wasp whose host is the spider Plesiometa argyra. The wasp is unusual in modifying the spider's web building behavior to make a web made of very strong lines designed to support the wasp's cocoon without breaking in the rain.
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.
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.
The Eucharitidae are a family of parasitic wasps. Eucharitid wasps are members of the superfamily Chalcidoidea and consist of three subfamilies: Oraseminae, Eucharitinae, and Gollumiellinae. Most of the 55 genera and 417 species of Eucharitidae are members of the subfamilies Oraseminae and Eucharitinae, and are found in tropical regions of the world.
The Leucospidae are a specialized group of wasps within the superfamily Chalcidoidea, that are ectoparasitoids of aculeate wasps or bees. They are typically mimics of bees or stinging wasps, often black with yellow, red, or white markings, sometimes metallic, with a robust mesosoma and very strong sculpturing. The hind femora are often greatly enlarged, with a row of teeth or serrations along the lower margin as in Chalcididae. The wing has a longitudinal fold. The female ovipositor is sometimes short, but if not, it is recurved and lies along the dorsal side of the metasoma, a unique feature. The males are also unusual, in the fusion of many of the metasomal segments to form a capsule-like "carapace".
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.
Megarhyssa, also known as giant ichneumonid wasps, giant ichneumons, or stump stabbers, is a genus of large ichneumon wasps, with some species known for having the longest ovipositors of any insects. They are idiobiont endoparasitoids of the larvae of wood-boring horntail wasps. The ovipositor can be mistaken for a large stinger. This is a genus of holometabolous insects within subfamily Rhyssinae that includes 37 species and belongs to Ichneumonidae, the family of wasps with the highest biodiversity in the world.
Pimplinae are a worldwide subfamily of the parasitic wasp family Ichneumonidae.
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.
Polysphincta boops is a species belonging to the family Ichneumonidae subfamily Pimplinae.
Behavior-altering parasites are parasites with two or more hosts, capable of causing changes in the behavior of one of their hosts to enhance their transmission, sometimes directly affecting the hosts' decision-making and behavior control mechanisms. They do this by making the intermediate host, where they may reproduce asexually, more likely to be eaten by a predator at a higher trophic level which becomes the definitive host where the parasite reproduces sexually; the mechanism is therefore sometimes called parasite increased trophic facilitation or parasite increased trophic transmission. Examples can be found in bacteria, protozoa, viruses, and animals. Parasites may also alter the host behavior to increase protection of the parasites or their offspring; the term bodyguard manipulation is used for such mechanisms.
Reclinervellus nielseni is one of the spider-ectoparasitoids belonging to the Polysphincta genus-group and utilizes exclusively Cyclosa spiders as hosts. The species is distributed from Britain to Japan but is rather sparse. Host spider species is different in accordance with the region, that is Cyclosa conica in Europe whereas Cyclosa argenteoalba in Japan.
Tetragnatha montana, commonly known as the silver stretch spider, is a species of long-jawed orb weaver from the family Tetragnathidae that has a Palearctic distribution. It preys mostly on flies and mosquitoes. The name silver stretch spider refers to its shiny metallic colour and its habit of extending its legs into a stick like shape.
Zatypota maculata, which is a member of ichneumonid ectoparasitoids of spiders, called the Polysphincta-group, is endemic to Japan and utilizes exclusively Nihonhimea japonica as host.
Acrodactyla quadrisculpta is a species of parasitoid wasp belonging to the family Ichneumonidae.