Pollenia Rudis
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''Pollenia rudis'', the common cluster fly, is a species of
fly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
in the family
Polleniidae Polleniidae is a family of flies in the order Diptera. There are at least 6 genera and more than 190 described species placed definitively in Polleniidae, and other genera whose placement here is considered uncertain. The largest genus is '' Pol ...
.Capinera, John; Heath, Allen. “Cluster Fly, ''Pollenia rudis'' (Fabricius) and P. pseudorudis Rognes (Diptera: Calliphoridae) ” Encyclopedia of Entomology: Springer Science+Business Media B.V. 2008. pg 932-935 ''Pollenia rudis'' is also known as the attic fly,Ridge, Gale E. "Cluster Flies ''Pollenia rudis'' (Fabricius) The Connecticut Agricultural Experiment Station 2007-04-06 http://www.ct.gov/CAES/cwp/view.asp?a=2815&q=376718 Retrieved on 2009-03-17."Cluster Flies" Cornell Cooperative Extension Nassau County 2003-01 Retrieved on 2009-03-17. the loft fly, ''pollenie du lombric''
rench The Rench is a right-hand tributary of the Rhine in the Ortenau (Baden (Land), Central Baden, Germany). It rises on the southern edge of the Northern Black Forest at Kniebis near Bad Griesbach im Schwarzwald. The source farthest from the mouth is ...
"''Pollenia rudis'' (Fabricius, 1794" ITIS Report 2003 https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=151647 Retrieved on 2009-03-17. and the buckwheat fly.Lyon, William F. "Cluster and Face Flies" Ohio State University Extension Fact Sheet Retrieved on 2009-03-17. During the autumn and winter months, ''Pollenia rudis'' can be found overwintering inside attics or lofts. This sluggish species can be found “clustering” near the interior windows of a warm structure.Howard, Leland O. “An Account of its Dangerous Activities and the Means of Destroying it” The House Fly – Disease Carrier, 1911. pg 236-240. This species is widely distributed throughout the United States, Canada, and Europe and is considered a pest species in structures. ''P. rudis'' can be found wherever their host earthworm, the ''
Allolobophora ''Allolobophora'' is a genus of annelid belonging to the family Lumbricidae The Lumbricidae are a family of earthworms. About 33 lumbricid species have become naturalized around the world, but the bulk of the species are in the Holarctic re ...
''"Cluster Fly" Varment Guard 2007 http://www.varmentguard.com/pestlibrary/CLUSTER%20FLY.pdf Retrieved on 2009-03-17. genera (also known by the genus name: Aporrectodea), occurs. These earthworms are typically located in well-drained, silt-loam soil with grass cover.Dille, James F.; Kirby, Clay A. "Cluster Flies" The University of Maine Pest Management 2008-19-08http://pmo.umext.maine.edu/factsht/cluster.htm Retrieved on 2009-03-17. During the summer, ''P. rudis'' can be found in fields and open areas. It is only when there is a sudden drop in temperature that the cluster fly shifts to the interior of structures, holes in trees, loose bark, or other crevices and cavities.


History

The common name "cluster fly" was derived from the clustering behavior in adults of this species in attics and lofts. The common name, "buckwheat fly", is derived from the odor of buckwheat honey the species gives off when they are crushed.Alm, Steven R. "Cluster Fly" University of Rhode Island GreenShare Factsheets 199

Retrieved on 2009-03-17.
''Pollenia rudis'' was first documented by
Johan Christian Fabricius Johan Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is cons ...
in 1794. At the time, Fabricius listed the
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and species as ''Musca rudis''.Lintner, J. A. “Pollenia rudis (Fabr.). The Cluster Fly.” Ninth Report on the Injurious and Other Insects of the State of New York For the Year 1892, 1893. pg 309-314. This taxonomy was changed in 1830 by André Jean Baptiste Robineau-Desvoidy to ''Pollenia rudis''.Rognes, Knut. “The Taxonomy of the Pollenia rudis species-group in the Holarctic Region (Diptera: Calliphoridae)” Systematic Entomology, 1987. Vol. 12, pg 475-502.Riley, C. V. “The ‘Cluster Fly’” American Naturalist, January 1883. Vol. CVII No. 1, pg 82-83. The change of genus to '' Pollenia'' (suggested by the pollen of flowers) occurred for Muscids having, among other features, the thorax covered with "down-like clothing". ''P. rudis'' has also been previously described under the name of ''Musca familiaris'' in 1869 by Dr. T.W. Harris. The cluster fly is a European species and the date of its introduction into the United States is not known. This species gained particular attention in the United States when Dr. W. H. Dall, of the
Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums and education and research centers, the largest such complex in the world, created by the U.S. government "for the increase and diffusion of knowledge". Founded ...
, published an article in the Proceedings of the U.S. National Museum for 1882. Dr. Dall secured specimens of ''P. rudis'' for identification. Dr. Dall also documented the species appearance in Geneva, NY thirty years prior to his publication. ''P. rudis'' may have been introduced to the United States upon slow sailing vessels in the cooler months of the year that traveled from Europe. This is possible due to the hibernation behaviors of the adult cluster flies to seek shelter for overwintering. The species also could have been transported to North America in the ballast of ships containing soil and the cluster fly host, earthworms.Jacobs, Steven B. "Cluster Flies" Penn State Entomological Notes 2003-03 http://www.ento.psu.edu/Extension/factsheets/clusterflies.htm Retrieved on 2009-03-17.


Characteristics


Distinctive markings

All flies can be identified from other species by certain characteristics. They can differ in
thoracic The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
coloring, basicosta coloring, and
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
coloring. Size and shape are aids in identification. ''Pollenia rudis'' eggs are oblong-shaped. They are very small and white. The ''P. rudis''
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e are white with posterior spiracles.Richards P. G. Morrison F. O. “A Summary of Published information on the cluster fly Pollenia rudis (Fabricus) (Diptera: Calliphoridae).” Phytoprotection. 53(2-3). 1972. 103-111. Pg 105. The adult ''Pollenia rudis'' looks like most of the other ''Pollenia'' species such as ''pallida'', and ''dasylpoda''. They are dark gray with checkered black and silvery-black abdomens. A newly emerged fly has many golden hairs on its thorax which may be lost throughout the life of the fly. The stripes on the thorax are not as prominent as on the house fly and the tips of the wings overlap when at rest. The cluster fly is slightly larger than a house fly at 9.525-12.7mm (3/8-1/2 inch) long. The similarities between ''pallida'' and ''rudis'' are seen in the female specimens. ''P. pallida'' has a broad, flattened facial keel. ''P.dasyloda'' has
black head
with yellow tint on the
frons Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insec ...
. Th
basicosta
can be found in many colors ranging from yellow to light brown. Some specimens have black basicosta. The posterior spiracle ranges from yellow in color to light brown. The number of bristles and setae found on this species are characteristic of this species only. There are 2-3 rows of
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
located on the thoracic section and 6-8 strong frontal bristles (bristles are thick setae). They also hav
aristate
antennae.Richards P G. Morrison F O. “A Summary of Published information on the cluster fly ''Pollenia rudis''(Fabricus) (Diptera: Calliphoridae).” Phytoprotection. 53(2-3). 1972. 103-111. Pg 104


Behavior

The behavior of the ''P. rudis'' fly varies with the annual seasons and conditions of the day. During the summer, on a sunny day these flies can be found without much trouble. When it is cold these flies tend to find somewhere warm and dry. They are mainly found in forest or wooded areas during the cold season. The flies tend to frequent dry areas because of their aristae antennae. The aristae are sensitive to minute temperature and pressure changes. During the winter, adult ''P. rudis have a habit of
overwintering Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal activi ...
. This ritual begins when the weather starts to get cold. The flies will inhabit the old tunnels created by past insects. They can also be found in old bird nests, under the bark of trees, or in homes. ''P. rudis'' will overwinter until spring, living off of its own fat.


Food sources

Earthworms are a major source of food for ''Pollenia rudis''. The main species of earthworm that these cluster flies infect ar
Aporrectoda caliginosa
Aporrectoda chlorotica
Eisenia lucens
''Lumbricus rubellus'', and ''Lumbricus terrestris''. Immediately after the larvae hatch, they begin looking for worms. The first
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
larvae eat their way through the integument section of the earthworm’s
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
. While feeding, the ''P. rudis'' larvae leave the spiracles outside of the earthworm. Inside the earthworm, the larvae feed until they are ready to pupate. The adult ''P. rudis'' are, in most cases,
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s. They feed on many types of
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
. Plant sap, fruit, flowers and
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
are common energy pathways for ''P. rudis''. ''P. rudis'' is also attracted to malt extract, acetyl acetate and the
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s in animal meat.


Predators

''Entomophthora muscae''
or ''Entomophthora schizophorae'' is a fungus that commonly infects adult flies. This fungus causes disease within the fly resulting in a swollen abdomen. This swollen abdomen makes the wings and legs spread apart, ultimately causing the fly to have trouble flying. After some time with this disease, a ''P. rudis'' adult will lose the ability to fly. Without flight, this fly has no way of protecting itself from predators. The main predator of ''P. rudis'' larvae is th
sphecid wasps.
These wasps will sting the fly and inject some poison. Once the fly has died, the wasp will feed the fly to their young. There are also other generalist predators of this fly, such as ants, birds, and certain plants.


Life cycle


Europe

There are very specific differences between the life cycle of ''Pollenia rudis'' in Europe and
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
.Thomson AJ (1973) The biology of Pollenia rudis, the cluster fly (Diptera: Calliphoridae). I. Host location of first-instar larvae. Can Entomol 105:335–341. In Europe, it takes 10–12 months for an egg to fully develop into an adult, resulting in one generation per year. ''P. rudis'' adults of European origin
copulate Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetrat ...
in the autumn , leaving their first instar larvae in a dormant state in the bodies of earthworms over the winter. The larvae will then molt twice over approximately 20 days and then pupate outside of the host. Usually, the pupal stage lasts from 32–45 days, but at high temperatures (e.g. 27°C) the pupal stage can be as short as 7 days.


North America

In North America, ''P. rudis'' eggs generally require 27–39 days to fully develop into an adult. In Canada, 25–30 days are required when the temperature is 23°C, and 11-14 of these days are spent in the pupal stage. Cluster flies in North America overwinter in their adult stage, and copulation takes place in the spring. There are three species in the rudis species complex of North America, and the life cycle of each species may differ. The variety of species in North America may account for the discrepancies between European and North American cluster flies life cycles. Female cluster flies preferentially oviposit eggs in humid areas with dense surface vegetation and high soil moisture.Thomson AJ, Davies DM (1973) The biology of ''Pollenia rudis'', the cluster fly (Diptera: Calliphoridae). II. Larval feeding behaviour and host specificity. Can Entomol 105:985–990 Each egg is either deposited by itself or in a small cluster of about seven eggs. In total, a female cluster fly will lay an average of 100-130 eggs by ovipositing a small group, then crawling or flying some distance before ovipositing each subsequent group of eggs. Once the larvae hatch, they burrow into the soil by following natural pore spaces, such as holes near plant stems or paths that earthworms have already created. By randomly moving through these pores, ''P. rudis'' larvae find their host worms. A larva is prompted to penetrate an earthworm when it senses “penetration inducing factor,” a substance that is present in the slime and coelomic fluid of an earthworm. The larvae then use their
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s to penetrate the dorsal side of a worm. It is important that they find their host quickly, because larvae must penetrate a worm within three days in order to survive. More than one larva can penetrate a single earthworm, and two or more larvae can share a penetration site. Once the host worm begins to decompose and is no longer useful to the larvae, the larvae can either leave to find another host or move to a less decomposed section further down on its host worm. If a first instar larva decides to move to another host, it must penetrate its new host quickly in order to survive. Towards the end of the first instar stage and into the second and third instar stages, the larvae are able to survive longer separations from their hosts. Usually, the first and second instars act as internal
parasite 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 has ...
s while the third instar can parasitize the host and feed on the surface of the host. Currently there are no reports of ''P. rudis'' reducing earthworm populations or causing horticultural problems.


Overwintering

Adult cluster flies in North America are slow-flying insects that are active during warmer months. Once the weather becomes much cooler, ''P. rudis'' will seek shelter, usually in homes and buildings. The flies have also been documented staying in tunnels made by
beetles Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
in timber and in animal burrows. Before overwintering, cluster fly’s abdomens are full of
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers spec ...
globules that may be left over from its larval fat bodies. When spring begins and the flies emerge, they appear shrunken because their fat was used up during the winter.


Human importance


Pest status

''Pollenia rudis'' is most commonly known for being a household nuisance. The first reports of cluster flies as pests in homes occurred as early as the nineteenth century, and these flies continue to cause problems today. Cluster flies tend to enter homes and buildings in large masses in late summer or early autumn to seek shelter for the winter months.Howard, Russell "Insects and Arthropods" Diagnostic Services at Michigan State 2006 http://www.pestid.msu.edu/InsectsArthropods/ClusterFlyPolleniarudis/tabid/254/Default.aspx They possess the ability to squeeze their bodies through any exterior crevices of a home, such as cracks around windows and doors, air conditioning vents, screening vents, and loosely hung siding. Once cluster flies enter a home, they usually
hibernate Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
in inaccessible areas between walls and in ceilings until spring when they emerge and seek access to the outdoors. ''P. rudis'' is extremely troublesome to home and business owners, but does not cause any true damage to home structures, textiles, foods, or humans. Piles of dead flies left in the walls can sometimes lead to secondary infestations of carpet or larder beetles and rodents.


Control methods

Attempting to control the cluster fly by controlling their earthworm host is not recommended or effective since flies may originate from more than a mile away from the infestation site. Once ''Pollenia rudis'' enters a home, it is almost impossible to kill enough of the flies to fully eradicate them. In order to prevent cluster flies from entering a house, all exterior cracks and openings should be caulked or sealed. This includes (sealing) light fixtures, electrical outlets, windows, and baseboards. Persistent use of insecticides has also been shown to reduce fly numbers if sprayed on the exterior of a home. However, these methods are not guaranteed for preventing infestations. When cluster flies do invade the home, they are extremely difficult to exterminate. Trying to kill the flies with a fly swatter leads to greasy spots on walls and upholstery as well as a lingering smell of buckwheat honey. Once spring arrives the flies will try to leave on their own in order to lay eggs, but many times they enter other living areas of the home instead. Vacuum cleaners and aerosol insecticides may be used to control visible, sluggish flies, but professional extermination is the most reliable method for eliminating an infestation.


Importance in forensic entomology

Unlike the majority of blow flies in the family Calliphoridae, ''Pollenia rudis'' does not play a large role in the medico-criminal subfield of
forensic entomology Forensic entomology is the scientific study of the colonization of a dead body by arthropods. This includes the study of insect types commonly associated with cadavers, their respective life cycles, their ecological presences in a given environme ...
. While most blow flies are attracted to rotting or decomposing matter, ''P. rudis'' is solely parasitic on earthworms and is unable to complete larval development on
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
.Greenberg, Bernard, and Kunich, John C. Entomology and the Law. 1st ed. Cambridge UP, 2002 This unique characteristic makes it uncommon to see this particular blowfly near forensic investigations. However, the urban subfield of forensic entomology has been closely monitoring ''P. rudis'' because of its tendency to infest buildings and its status as a potential
disease vector In epidemiology, a disease vector is any living agent that carries and transmits an infectious pathogen to another living organism; agents regarded as vectors are organisms, such as parasites or microbes. The first major discovery of a disease vec ...
. Urban entomology, which deals with the insects that affect humans and their immediate environment,Byrd, Dr. J.H. "What is Forensic Entomology" Forensics Entomology 2009 http://www.forensicentomology.com/definition.htm is responsible for investigating economic issues and civil proceedings involving
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. There have been several economically hampering cases of ''P. rudis'' infestation around the world. For example, in New Zealand an entire city’s water reservoir tank was drained due to high levels of fecal coliform bacteria produced by mass amounts of cluster flies residing in the tank. Due to cases like these, scientists have investigated the association of ''Pollenia rudis'' and its disease vector capability. In 1973, a massive infestation of cluster flies in a German hospital triggered an investigation of the relationship between ''P. rudis'' and bacteria pathogen transmission.Faulde, M; Sobe, D; Burghardt, H; Wermter, R. “Hospital infestation by the cluster fly, Pollenia rudis sensu stricto Fabricius 1794 (Diptera: Calliphoridae), and its possible role in transmission of bacterial pathogens in Germany.” International Journal of Hygiene & Environmental Health. 203(3):201-4, 2001 Mar. Careful examination of the flies revealed ''P. rudis'' is only capable of transmitting bacteria that causes
opportunistic infection An opportunistic infection is an infection caused by pathogens (bacteria, fungi, parasites or viruses) that take advantage of an opportunity not normally available. These opportunities can stem from a variety of sources, such as a weakened immune ...
s. These results indicate that “mass infestations of cluster flies occurring in sensitive areas, especially in hospitals, may cause a low, but not neglectable health threat due to mechanical transmission of bacterial pathogens.”


References

{{DEFAULTSORT:Pollenia Rudis Polleniidae Insects described in 1794 Diptera of Europe Taxa named by Johan Christian Fabricius