Howardula | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Nematoda |
Class: | Secernentea |
Order: | Tylenchida |
Family: | Allantonematidae |
Genus: | Howardula Cobb, 1921 |
Howardula is a genus of nematode that infests the larvae of mushroom-feeding flies, beetles, and other insects. Various Howardula species and strains infest mushroom-feeding Drosophila, including Howardula aoronymphium and Howardula neocosmis . [1] Howardula husseyi can infest the mushroom phorid Megaselia halterata . [2]
The life cycle of Howardula nematodes begins inside the host fly, where infective juvenile nematodes are released by either the fly anus or ovipositor onto mushrooms. There, the juvenile nematodes mature and mate. Mated females will then pierce the cuticle of a fly larva using a specialized stylet, and enter the fly hemolymph (insect blood) where the nematode resides. Over the course of the fly host's metamorphosis, the female nematode matures into an adult stage called the motherworm. After the adult fly emerges from its pupa, the motherworm begins releasing juvenile nematodes directly into the hemocoel, renewing the cycle. [1]
Other insect-infecting nematodes such as Steinernema and Heterorhabiditis are used as biological control agents, as they efficiently kill their insect prey. However Howardula species require their hosts to survive into adulthood. As such, while Howardula infects insects like other nematode parasites, the "goals" of these nematodes differs drastically, leading to distinct host-parasite interactions.
The apple maggot, also known as the railroad worm, is a species of fruit fly, and a pest of several types of fruits, especially apples. This species evolved about 150 years ago through a sympatric shift from the native host hawthorn to the domesticated apple species Malus domestica in the northeastern United States. This fly is believed to have been accidentally spread to the western United States from the endemic eastern United States region through contaminated apples at multiple points throughout the 20th century. The apple maggot uses Batesian mimicry as a method of defense, with coloration resembling that of the forelegs and pedipalps of a jumping spider.
The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking flight. This behaviour is a source of one of their alternate names, scuttle fly. Another vernacular name, coffin fly, refers to Conicera tibialis. About 4,000 species are known in 230 genera. The most well-known species is cosmopolitan Megaselia scalaris. At 0.4 mm in length, the world's smallest fly is the phorid Euryplatea nanaknihali.
Elaeophora is a genus of parasitic nematodes which live attached to the interior surfaces of major arteries, veins and/or heart chambers in various large mammal hosts. Infestation with Elaeophora species is referred to as elaeophorosis. The species of Elaeophora have been found in Africa, Asia, Europe, and North America. Despite the fact that they produce aneurysms in the arteries and heart of their hosts which measure up to 2 cm in diameter, overt clinical symptoms of infestation are seldom reported, with the notable exception of E. schneideri infestation in sheep, elk, and moose.
Entomopathogenic nematodes (EPN) are a group of nematodes, that cause death to insects. The term entomopathogenic has a Greek origin, with entomon, meaning insect, and pathogenic, which means causing disease. They are animals that occupy a bio control middle ground between microbial pathogens and predator/parasitoids. Although many other parasitic thread worms cause diseases in living organisms, entomopathogenic nematodes are specific in only infecting insects. Entomopathogenic nematodes (EPNs) live parasitically inside the infected insect host, and so they are termed as endoparasitic. They infect many different types of insects living in the soil like the larval forms of moths, butterflies, flies and beetles as well as adult forms of beetles, grasshoppers and crickets. EPNs have been found all over the world in a range of ecologically diverse habitats. They are highly diverse, complex and specialized. The most commonly studied entomopathogenic nematodes are those that can be used in the biological control of harmful insects, the members of Steinernematidae and Heterorhabditidae. They are the only insect-parasitic nematodes possessing an optimal balance of biological control attributes.
Rotylenchulus reniformis, the reniform nematode, is a species of parasitic nematode of plants with a worldwide distribution in the tropical and subtropical regions.
Heterodera sacchari, the sugarcane cyst nematode, mitotic parthenogenic sedentary endoparasitic nematode. This plant-parasitic nematode infects the roots of sugarcane, and the female nematode eventually becomes a thick-walled cyst filled with eggs. Aboveground symptoms are species specific and are similar to those caused by other Heterodera species. Symptoms include: stunted and chlorotic plants, and reduced root growth. Seedlings may be killed in heavily infested soils.
The microfilaria is an early stage in the life cycle of certain parasitic nematodes in the family Onchocercidae. In these species, the adults live in a tissue or the circulatory system of vertebrates. They release microfilariae into the bloodstream of the vertebrate host. The microfilariae are taken up by blood-feeding arthropod vectors. In the intermediate host the microfilariae develop into infective larvae that can be transmitted to a new vertebrate host.
The Filarioidea are a superfamily of highly specialised parasitic nematodes. Species within this superfamily are known as filarial worms or filariae. Infections with parasitic filarial worms cause disease conditions generically known as filariasis. Drugs against these worms are known as filaricides.
Apocephalus borealis is a species of North American parasitoid phorid fly that attacks bumblebees, honey bees, and paper wasps. This parasitoid's genus Apocephalus is best known for the "decapitating flies" that attack a variety of ant species, though A. borealis attacks and alters the behavior of bees and wasps. These flies are colloquially known as zombie flies and the bees they infect are colloquially known as zombees. Association with honey bees has so far only been documented from California, South Dakota, Oregon, Washington, British Columbia, and Vermont.
Drosophila falleni is a species of fly found in northeastern North America. A mycophagous insect, it is known to feed on the fruit bodies (mushrooms) of several genera of fungi, including Agaricus, Amanita, Agrocybe, Boletus, Cortinarius, PluteusGrifola, Polyporus, Hypholoma, Russula, Mycena, Stropharia, and Xerula. The fly can be infested by the parasitic nematode Howardula aoronymphium, which enters the fly larvae when it is feeding on mushrooms.
Steinernema scapterisci, the mole cricket nematode, is a species of nematode in the order Rhabditida. It is a parasite of insects in the order Orthoptera, the grasshoppers, crickets and their allies. Native to southern South America, it was introduced into Florida in the United States in an effort to provide a biological control of pest (Neoscapteriscus) mole crickets.
Many species of flies of the two-winged type, Order Diptera, such as mosquitoes, horse-flies, blow-flies and warble-flies, cause direct parasitic disease to domestic animals, and transmit organisms that cause diseases. These infestations and infections cause distress to companion animals, and in livestock industry the financial costs of these diseases are high. These problems occur wherever domestic animals are reared. This article provides an overview of parasitic flies from a veterinary perspective, with emphasis on the disease-causing relationships between these flies and their host animals. The article is organized following the taxonomic hierarchy of these flies in the phylum Arthropoda, order Insecta. Families and genera of dipteran flies are emphasized rather than many individual species. Disease caused by the feeding activity of the flies is described here under parasitic disease. Disease caused by small pathogenic organisms that pass from the flies to domestic animals is described here under transmitted organisms; prominent examples are provided from the many species.
Dendroctonus adjunctus, the roundheaded pine beetle, is a species of bark beetle in the family Curculionidae found in North America. A parasite, the roundheaded pine beetle feeds on and eventually kills pine trees of several species in Guatemala, Mexico, and the Southern United States.
Howardula aoronymphium is a species of nematode that infects specialist mushroom-feeding fruit flies such as Drosophila falleni and Drosophila neotestacea. Mated female nematodes pierce the fly larva cuticle and take up residence in the hemolymph where they mature alongside the fly. When the adult fly ecloses, the nematode motherworm has reached full size and sheds juvenile nematodes into the hemolymph which are eventually excreted by either the fly anus or ovipositor. Howardula nematodes can severely impact fly egg development, as infection can effectively sterilize some species.
Drosophila neotestacea is a member of the testacea species group of Drosophila. Testacea species are specialist fruit flies that breed on the fruiting bodies of mushrooms. These flies will choose to breed on psychoactive mushrooms such as the Fly Agaric Amanita muscaria. Drosophila neotestacea can be found in temperate regions of North America, ranging from the north eastern United States to western Canada.
The Drosophila testacea species group belongs to the Immigrans-tripunctata radiation of the subgenus Drosophila, and contains 4 species: Drosophila putrida, Drosophila neotestacea, Drosophila testacea, and Drosophila orientacea. Testacea species are specialist mushroom-feeding flies, and can metabolize toxic compounds in Amanita mushrooms. The Testacea species group is studied for its specialist ecology, population genetics, and bacterial endosymbionts. The North American species Drosophila neotestacea is perhaps the best-studied of the group for its interactions with parasitic wasps and nematodes, bacterial endosymbionts, and trypanosomatid parasites. Of note, selfish X chromosomes have been discovered in three of the four Testacea group species.
Spiroplasma poulsonii are bacteria of the genus Spiroplasma that are commonly endosymbionts of flies. These bacteria live in the hemolymph of the flies, where they can act as reproductive manipulators or defensive symbionts.
The mushroom phorid fly(Megaselia halterata) is a species of scuttle fly or hump-backed flies in the family Phoridae. "The mushroom phorid" is also used to refer to M. halterata. Megaselia halterata is a common pest of mushroom cultivation, attracted by the aroma of developing fungal mycelium. The larvae damage both the mushroom mycelium and gill tissues. Megaslia halterata can be found worldwide.
The Drosophila quinaria species group is a speciose lineage of mushroom-feeding flies studied for their specialist ecology, their parasites, population genetics, and the evolution of immune systems. Quinaria species are part of the Drosophila subgenus.
Allantonematidae is a family of insect-parasitic nematodes from the order Tylenchida. Allantonematid nematodes infect a variety of insects including beetles, butterflies, flies, thrips, ants, and more. For instance, the nematode Howardula aoronymphium parasitizes mushroom-feeding fruit flies, Formicitylenchus oregonensis parasitizes carpenter ants, and Metaparasitylenchus hypothenemi parasitizes a pest of coffee beans, the coffee berry borer.