Mallophora Bomboides
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''Mallophora bomboides'', also known as the
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
bee killer, is a predaceous species of robber fly of the family
Asilidae The Asilidae are the robber fly family, also called assassin flies. They are powerfully built, bristly flies with a short, stout proboscis enclosing the sharp, sucking Insect mouthparts#Hypopharynx, hypopharynx. The name "robber flies" reflects t ...
that feeds primarily on
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s. ''M. bomboides'' is a noteworthy instance of
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
given its close resemblance to its prey, the bumblebee. These bees are typically found in the Eastern and Southern regions of the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
like
South Carolina South Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders North Carolina to the north and northeast, the Atlantic Ocean to the southeast, and Georgia (U.S. state), Georg ...
and
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
.


Physical description

''M. bomboides'' typically have three basal
abdominal The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
tergites A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'margin'. ...
densely covered with yellow
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and ...
s. The fourth and fifth tergites have black hairs, and the final two segments have pale hairs. The
ventrum The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
of the abdomen is adorned with yellow hairs. The average body length of ''M. bomboides'' is about 25 mm.


Distribution


Geographical distribution

''M. bomboides'' occurs in the
southeastern United States The Southeastern United States, also known as the American Southeast or simply the Southeast, is a geographical List of regions in the United States, region of the United States located in the eastern portion of the Southern United States and t ...
in states including
North Carolina North Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, South Carolina to the south, Georgia (U.S. stat ...
,
South Carolina South Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders North Carolina to the north and northeast, the Atlantic Ocean to the southeast, and Georgia (U.S. state), Georg ...
,
Georgia Georgia most commonly refers to: * Georgia (country), a country in the South Caucasus * Georgia (U.S. state), a state in the southeastern United States Georgia may also refer to: People and fictional characters * Georgia (name), a list of pe ...
,
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
,
Alabama Alabama ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South, Deep Southern regions of the United States. It borders Tennessee to the north, Georgia (U.S. state), Georgia to the east, Florida and the Gu ...
, and
Mississippi Mississippi ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South regions of the United States. It borders Tennessee to the north, Alabama to the east, the Gulf of Mexico to the south, Louisiana to the s ...
.


Phenology

Flight time in Florida is from April through December, though they are most commonly seen in late
summer Summer or summertime is the hottest and brightest of the four temperate seasons, occurring after spring and before autumn. At or centred on the summer solstice, daylight hours are the longest and darkness hours are the shortest, with day ...
.


Habitat

Adults typically live in open habitats often in the vicinity of apiaries, perching on stalks of weeds or tips of shrubs from which they launch their attacks.


Life Cycle


Larva

Specific feeding habits of
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e of ''M. bomboides'' are unknown, but other ''Mallophora'' larvae are ectoparasites on scarabaeid
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
larvae in the soil.


Adult

Adults live in open
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
s near apiaries. They perch vertically angled on plants from which they launch their attacks on their bumblebee prey.


Mimicry


History

The similar appearances of some
flies 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 hindwin ...
and
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s have been noted since the time of Aristotle. In fact, in these ancient times, the confusion between the two insects led to the disproven notion that bees undergo
spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from non-living matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could ...
from decaying carcasses. British entomologists William Kirby and
William Spence William Guthrie Spence (7 August 1846 – 13 December 1926), was an Australian trade union leader and politician, played a leading role in the formation of both Australia's largest union, the Australian Workers' Union, and the Australian Labor ...
first theorized in 1817 that flies gained an advantage by resembling bees, reasoning that the physical similarity of
hoverflies Hoverflies, also called flower flies or syrphids, make up the insect family (biology), family Syrphidae. As their common name suggests, they are often seen Hover (behaviour), hovering or nectaring at flowers; the adults of many species feed main ...
from genus
Volucella ''Volucella'' is a genus of large, broad-bodied, dramatic hover-fly, hover-flies. They have distinctive Wiktionary:plumose, plumose Arista (biology), aristae and the face is extended downward. They are strongly Insect migration, migratory and mal ...
to
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s enabled the invasion of bee nests. Having entered the bee nests without arousing suspicion, the flies then lay eggs and depart without being attacked. The same idea was later put forth by British naturalist
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was an English naturalist, explorer, geographer, anthropologist, biologist and illustrator. He independently conceived the theory of evolution through natural selection; his 1858 pap ...
in 1871.


Initial classification

Classifying categories of mimicry in 1890, Edward Bagnell Poulton placed this into the category of
aggressive mimicry Aggressive mimicry is a form of mimicry in which predation, predators, parasites, or parasitoids share similar signalling theory, signals, using a harmless model, allowing them to avoid being correctly identified by their prey or host (biolog ...
, a deceptive mechanism in which one species resembles another in order to approach it without arousing suspicion to carry out a detrimental end. However,
Gregory Bateson Gregory Bateson (9 May 1904 – 4 July 1980) was an English anthropology, anthropologist, social sciences, social scientist, linguistics, linguist, visual anthropology, visual anthropologist, semiotics, semiotician, and cybernetics, cybernetici ...
criticized this view in 1892 by pointing out that the Volucella example fits much better as an instance of protective mimicry, now commonly known as
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
. By appearing as bees, palatable flies gain protection from
predators Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
that recognize bees as noxious and therefore unappetizing. Bateson argued with the counterexamples that Volucella females entered bumblebee nests belonging to species that they did not mimic and that a European species with similar habits actually benefited the host because the fly
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e, once hatched, acted as
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
s inside the nest.


= Reclassification

= In the same paper in which he discussed aggressive mimicry in order to account for ''Volucella'' appearances, Poulton also discussed family Asilidae flies that prey upon aculeate Hymenoptera as adults. ''M. bomboides'' bear a highly specific resemblance to their prey, which Poulton classified as
protective mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
. However, Lincoln Brower et al. demonstrated in 1960 that this phenomenon was, in fact, Batesian mimicry.


Aggressive mimicry

Adult ''M. bomboides'' prey on bumblebees, their mimetic models, to a great extent. Though their attacks are swift, it is possible that the resemblance of the fly to the prey and mode enhances the likelihood of its success. The same argument for aggressive mimicry in Volucella flies could apply in ''M. bomboides''. Consistent with other members of the family Asilidae, larvae are thought to be predaceous, feeding on soft-bodied insects in the soil. In terms of aggressive mimicry, the female flies of ''M. bomboides'' may enter into bumblebee nests to lay eggs. Due to their mimetic resemblance, ''M. bomboides'' females could then escape recognition and attack. Once hatched, the fly larvae could consume bumblebee larvae as a food source. However, all this is merely speculative and further experimentation will help elucidate the validity of this model.


Batesian mimicry

Experiments conducted by Brower et al. demonstrate that ''M. bomboides'' is a Batesian mimic of its bumblebee model and prey, ''B. americanorum'' (now more commonly known as ''
Bombus pensylvanicus ''Bombus pensylvanicus'', the American bumblebee, is a threatened species of bumblebee native to North America. It occurs in eastern Canada, throughout much of the Eastern United States, and much of Mexico.Hatfield, R., et al. 2015''Bombus pensyl ...
''), which is noxious to predators such as the toad '' Bufo terretris'' due to its sting. In a 1921 experiment by
Geoffrey Douglas Hale Carpenter G.D. Hale Carpenter MBE (26 October 1882 in Eton, Berkshire – 30 January 1953 in Oxford) was a British entomologist and medical doctor. He worked first at the London School of Hygiene and Tropical Medicine, and in Uganda, on tse-tse flies a ...
, one monkey ate a wasp-like asilid presented to it with caution and another rejected the asilid as food altogether. However, the second monkey was willing to eat a non-mimetic asilid, suggesting that the first asilid is a true Batesian mimic. In other words, mimicry of an unpalatable species will lend species such as ''M. bomboides'' a form of protection from predators who have learned their lesson from an unpleasant previous attempt. In another experiment by Lloyd Morgan in 1896, it was discovered that a
moorhen Moorhens—sometimes called marsh hens—are medium-sized water birds that are members of the rail family (Rallidae). Most species are placed in the genus ''Gallinula'', Latin for "little hen." They are close relatives of coots. They are ...
chick that had eaten and presumably been stung by a bumblebee rejected future offers of bumblebees, even with removed stings, as well as mimetic drone flies. An inexperienced chick was willing to accept these as food. In addition to supporting the hypothesis for Batesian mimicry, this experiment suggests that the noxious quality of the bumblebees is their sting. This was further supported by a series of feeding experiments conducted using
toad Toad (also known as a hoptoad) is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands. In popular culture (folk taxonomy ...
s and
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s by Hugh B. Cott in 1936.


Requirements

In order that Batesian mimicry should evolve and remain an evolutionarily stable strategy, several prerequisites are deemed necessary. # Prospective models and mimics must occur together. # Models must outnumber mimics. # Models must possess a noxious and advertise this by means of conspicuous coloration. For mimetic asilids like ''M. bomboides'', these organisms attack their aculeate Hymenoptera models and will seek habitats abundant in their prey, thereby ensuring
sympatry In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
. All these conditions hold for the ''M. bomboides'' with their models, ''B. americanorum'' in a 1960 conducted by Brower et al. in south central
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
. The toad ''B. terrestris'' was used as caged predators to demonstrate that, despite their night foraging and lack of color vision, they can learn to reject bumblebees on sight alone and confuse mimetic flies with their apian hosts as well.


Diet

As a member of the family Asilidae, ''M. bomboides'' preys on various
aculeate Aculeata is an infraorder of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group canno ...
Hymenoptera 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 typi ...
species. ''M. bomboides'' in particular prey on
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s such as ''
Bombus pensylvanicus ''Bombus pensylvanicus'', the American bumblebee, is a threatened species of bumblebee native to North America. It occurs in eastern Canada, throughout much of the Eastern United States, and much of Mexico.Hatfield, R., et al. 2015''Bombus pensyl ...
'', to which ''M. bomboides'' bears remarkable resemblances.


Reproduction

''M. bomboides'' lay their eggs in
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
.


Economic importance

There are few instances of financials losses to beekeepers due to depredations of bee killers such as ''M. bomboides'', but Florida is one of a few states where such losses have been reported as noteworthy. Little Lake City experienced an attack of over hundreds of these insects against bee hives in July 2008.


References


External links

* {{Taxonbar, from=Q10790674 Asilidae Articles created by Qbugbot Insects described in 1821