Villa alternata

Ge­o­graphic Range

Villa al­ter­nata is a bee fly species found on every con­ti­nent ex­cept Antarc­tica. Bom­byli­idae are a rel­a­tively early di­verg­ing lin­eage within the large ra­di­a­tion of flies known as Brachyc­era (short-horned flies). This genus of bee flies evolved be­fore the breakup of Pangea. (Great­head, 1981)

Habi­tat

Since Villa al­ter­nata is found on all con­ti­nents ex­cept Antarc­tica, it is found in a large va­ri­ety of habi­tats with vary­ing rain­fall, tem­per­a­ture and soil types. Its habi­tat must be rich in its' host species (noc­tuid moths and tene­bri­onid bee­tles). These bee flies are found in dunes, sandy scrub re­gions, forests, grass­lands, taiga, marshes, and bogs. (Falck, 2009; Hull, 1973; Kits, et al., 2008; Yeates and Lam­bkin, 1994)

Phys­i­cal De­scrip­tion

This Bom­byli­idae species has a short pro­boscis. These bee flies are cov­ered in hairs and scales that vary from golden yel­low to black and white, often with bands of bright scales on the ab­dom­i­nal ter­gites and with tufts of black and white or yel­low­ish hairs at the end of the ab­domen. The wings of Villa al­ter­nata are al­most en­tirely clear with a slight brown hue at the wing base. One pale pile ex­ists on each pleu­ron. On the sides of seg­ment three of the ab­domen is a black pile. Tarsal claws of forelegs are much smaller than those of other legs. The fore tibia have ei­ther promi­nent spines or the sides of ab­domen are found to have long, flat­tened scales, but the fly never has both char­ac­ters. Villa al­ter­nata typ­i­cally has a rounded head. The males have a bril­liant mat of sil­very pata­gial scales. On the wing of genus Villa, the fork­ing of the ra­dial sec­tor takes place at or near the r-m cross-vein, and never at a greater dis­tance than the length of that vein. A sec­toral cross-vein may be pre­sent, but is usu­ally ab­sent and con­tact of cells 1M-2 and u-1 are not longer than the base of Cu-1. The an­ten­nal style is minute or ab­sent and there are no pul­villi in most of the species. The species of this genus are more or less densely clothed with pile and to­men­tum.

Since their habi­tat can vary so much, the ap­pear­ance of Villa may not be strictly uni­form. Pop­u­la­tions found in areas of high tem­per­a­ture, low rain­fall and sandy soil are pre­dom­i­nantly lighter in color than those found in areas of lower tem­per­a­ture, higher rain­fall and heav­ier soil. (Falck, 2009; Kits, et al., 2008)

  • Range length
    10 to 17 mm
    0.39 to 0.67 in

De­vel­op­ment

Bee flies are holometabolous and de­velop through hy­per­me­ta­mor­pho­sis in which the 1st in­star larva is in­quiline, liv­ing in a sand cham­ber and left to find a host by it­self. The lar­vae com­monly par­a­sitizes the lar­vae of Noc­tuid moths (Noc­tu­idae) or in some areas tene­bri­onid bee­tles (Tene­bri­on­idae). Once the larva finds a host and pen­e­trates the host's nests, where it turns into a seden­tary par­a­sitoid. The pupa is equipped with spines/spikes that en­able it to break free from the host pupa. It drills out of the nest, where it emerges as a fly­ing adult equipped with sex or­gans. (Falck, 2009; Yeates and Lam­bkin, 2013)

Re­pro­duc­tion

Fe­male bee flies fly to el­e­vated land­marks solely for the pur­pose of mat­ing with males that have ag­gre­gated there. This phe­nom­e­non is known as "hill­top­ping." No in­ter­ac­tive courtship be­hav­ior was de­tectable prior to midair cou­pling to en­gage in mat­ing. Fe­males are free to choose mates but males have been ob­served in­ter­cept­ing a fe­male as she flies over him. It is not known whether fe­males can pre­vent mat­ings after being grabbed. Cop­u­la­tions last over 100 min­utes on av­er­age.

Ter­ri­to­r­ial fights be­tween males be­fore choos­ing a mate in­volve aer­ial col­li­sions dur­ing which mod­i­fied spines on the wing mar­gins pro­duce scars on the bod­ies of op­po­nents. Ter­ri­tory own­ers and mat­ing males are not dif­fer­ent in size or age from the re­main­der of the male pop­u­la­tion. (Yeates and Dod­son, 1990)

After mat­ing, Villa al­ter­nata adult fe­males find a crack or hole in the ground and po­si­tion them­selves to make a sand cham­ber. The fe­males then fill this cham­ber with sand, dust, saw-dust, or other fine par­ti­cles. This be­hav­ior is un­der­taken to pre­vent the eggs from stick­ing to­gether, as they are flicked off the fe­male and land one by one to the ground where host species are in high den­sity. In ovipo­si­tion, the fe­male gives the ab­domen a down-ward flick, touches hind tarsi to the ground, hov­ers again for a sec­ond or two, re­leases an­other egg. Hun­dreds of eggs are de­posited in the sand cham­ber in a mat­ter of hours. Villa ap­pear com­monly in the months of May to Au­gust in North Amer­i­can tem­per­ate cli­mates, so re­pro­duc­tion likely takes place in the spring and sum­mer. (Falck, 2009; Toft, 1983)

  • Breeding season
    Mating takes place in the spring and summer.

There seems to be no known parental in­vest­ment by male bee flies, but there may be a pos­si­bil­ity for the stor­age of nu­tri­ent rich ejac­u­late into the fe­male dur­ing cop­u­la­tion. Cop­u­la­tion for Bom­byli­idae lasts 2 to 15 min­utes, pro­vid­ing the po­ten­tial for trans­fer of large quan­ti­ties of sperm or other com­po­nents into the fe­male for use to de­velop and sus­tain off­spring. Fe­male in­vest­ment lies in the en­ergy and re­sources they in­vest to­wards the for­ma­tion of the zy­gote and the col­lect­ing and glu­ing of small par­ti­cles of sub­strate to eggs be­fore flick­ing them off their bod­ies. (Yeates and Great­head, 1997)

  • Parental Investment
  • pre-hatching/birth
    • provisioning

Lifes­pan/Longevity

The lifes­pan of Villa al­ter­nata is not known, but in the Amer­i­can south­west, the ma­jor­ity of noted Villa al­ter­nata carry out their adult lives in Au­gust. It is likely that adults live for about a month. (Toft, 1983)

  • Average lifespan
    Status: wild
    1 months

Be­hav­ior

Bee flies are very mo­bile, hov­er­ing or skill­fully fly­ing in their adult life. Bee flies are soli­tary as adults ex­cept for the hours when they mate or when males par­take in ter­ri­to­r­ial fights. Ter­ri­to­r­ial fights for mates be­tween males in­volve aer­ial col­li­sions, usu­ally within range of the hill­top­ping area. Bee fly males have mod­i­fied spines on the wing mar­gins that can cut into the bod­ies of male op­po­nents. Bom­byli­idae are di­ur­nal and most ac­tive in the pres­ence of sun­light. Lar­vae stay in groups in the host in­sect be­fore emerg­ing. (Kits, et al., 2008; Yeates and Dod­son, 1990)

Com­mu­ni­ca­tion and Per­cep­tion

Most Dipteran par­a­sitoids that rely on im­ma­tures for host con­tact pro­duce ac­tively search­ing lar­vae. Such plani­d­i­form lar­vae have evolved in­de­pen­dently in fam­i­lies such as Bom­byli­idae which al­lows lar­vae to find hosts in­ac­ces­si­ble to adult bee flies. Lit­tle is known about the type of per­cep­tion used for lar­vae to do so. There is lit­tle known about so­cial be­hav­iors in Bom­byli­idae, but it is likely that vi­sion and chemore­cep­tion are used. (Feener and Brown, 1997)

Food Habits

Villa al­ter­nata is a pollen eater and nec­tar feeder. It has a rel­a­tively short pro­boscis in re­la­tion to other Bom­byli­idae. This species is known to feed on small flow­ers such as Chrysotham­nus vis­cid­i­florus in the com­mon Aster­aceae fam­ily. Such feed­ing is noted in the Amer­i­can South­west. Fe­male bee flies spend large amounts of time feed­ing, up to 8 times that of males. Food ob­tained by adults has the po­ten­tial to con­sti­tute a sub­stan­tial part of the nu­tri­tional in­vest­ment in re­pro­duc­tion. (Toft, 1983)

  • Animal Foods
  • insects
  • Plant Foods
  • nectar
  • pollen

Pre­da­tion

Birds are preda­tors of Villa al­ter­nata. Bee flies demon­strate Bate­sian mim­icry of Hy­menoptera. The bee flies share sim­i­lar col­ors, pat­terns, and a sim­i­lar stout and woolly body as many wasps and bees. Mim­ic­k­ing wasps and bees, which are known to sting preda­tors, is ben­e­fi­cial to Villa al­ter­nata, since preda­tors may as­sume it is a sting­ing hy­menopteran and avoid try­ing to eat it. (Yeates and Lam­bkin, 1994; Yeates and Lam­bkin, 2013)

  • Anti-predator Adaptations
  • mimic
  • Known Predators

Ecosys­tem Roles

Villa al­ter­nata lar­vae par­a­sitize Noc­tuid moths (Noc­tu­idae), which feed on nec­tar rich flow­ers and woody plants. Noc­tuid moths in turn par­a­sitize agri­cul­tural pests with its cut­worm lar­vae. The bee fly lar­val par­a­sitoids on Noc­tuid moths may im­pact moth pop­u­la­tions in a habi­tat patch, in­di­rectly ef­fect­ing the growth or de­cline of nec­tar rich flow­ers and agri­cul­tural pests in the area.

This genus of bee fly also par­a­sitizes tene­bri­onid bee­tles (Tene­bri­on­idae), which feed on a va­ri­ety grasses such as blue­grass and sage­brush. This also sug­gests that Villa al­ter­nata may have an in­di­rect af­fect on the local suc­cess of such grasses if they are lim­ited in abun­dance. The eco­log­i­cal im­pact of Villa may be broad in re­gards to the species af­fected due to the wide range of habi­tats and cli­mate this genus can live. (Hull, 1973; Kits, et al., 2008; Shel­don and Beed­low, 1988; Yela and Her­rera, 1993)

Species Used as Host

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Villa al­ter­nata is known to be a par­a­sitoid of Noc­tu­idae moths. Noc­tuid moth lar­vae are known to be pests of agri­cul­tural crops. Villa al­ter­nata likely can con­trol these noc­tuid pest pop­u­la­tions, de­creas­ing the po­ten­tial eco­nomic losses re­sult­ing from crop dam­age. Bom­byli­idae are also pol­li­na­tors on many types of flow­er­ing plants, some of which may be ben­e­fi­cial to hu­mans. (Wieg­mann, 2011)

  • Positive Impacts
  • controls pest population

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no known ad­verse ef­fects of Villa al­ter­nata on hu­mans.

Con­ser­va­tion Sta­tus

Villa al­ter­nata has no spe­cial con­ser­va­tion sta­tus. (McGin­ley, 1993)

Other Com­ments

There is lit­tle lit­er­a­ture avail­able on Villa al­ter­nata. More re­search needs to be done on preda­tory threats to the species, be­hav­ioral ten­den­cies, longevity, and gen­eral char­ac­ter­is­tics that V. al­ter­nata has in spe­cific habi­tats.

Con­trib­u­tors

Jayna Sames (au­thor), Uni­ver­sity of Michi­gan Bi­o­log­i­cal Sta­tion, Brian Scholtens (ed­i­tor), Uni­ver­sity of Michi­gan Bi­o­log­i­cal Sta­tion, An­gela Miner (ed­i­tor), An­i­mal Di­ver­sity Web Staff.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

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Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

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Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

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Neotropical

living in the southern part of the New World. In other words, Central and South America.

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Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

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bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

bog

a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

cosmopolitan

having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

diurnal
  1. active during the day, 2. lasting for one day.
ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

internal fertilization

fertilization takes place within the female's body

marsh

marshes are wetland areas often dominated by grasses and reeds.

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

nectarivore

an animal that mainly eats nectar from flowers

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

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oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

parasite

an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death

polymorphic

"many forms." A species is polymorphic if its individuals can be divided into two or more easily recognized groups, based on structure, color, or other similar characteristics. The term only applies when the distinct groups can be found in the same area; graded or clinal variation throughout the range of a species (e.g. a north-to-south decrease in size) is not polymorphism. Polymorphic characteristics may be inherited because the differences have a genetic basis, or they may be the result of environmental influences. We do not consider sexual differences (i.e. sexual dimorphism), seasonal changes (e.g. change in fur color), or age-related changes to be polymorphic. Polymorphism in a local population can be an adaptation to prevent density-dependent predation, where predators preferentially prey on the most common morph.

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

scrub forest

scrub forests develop in areas that experience dry seasons.

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

taiga

Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

Ref­er­ences

Falck, M. 2009. The Nor­we­gian species of Villa Lioy, 1864 (Diptera, Bom­byli­idae). Nor­we­gian Jour­nal of En­to­mol­ogy, 56: 120–130.

Feener, D., B. Brown. 1997. Diptera as Par­a­sitoids. An­nual Re­view of En­to­mol­ogy, 2: 73-97.

Great­head, D. 1981. The Villa group of gen­era in Africa and Eura­sia with a re­view of the gen­era com­pris­ing Thyri­dan­thrax sensu Bezzi 1924 (Diptera: Bom­byli­idae). Jour­nal of Nat­ural His­tory, 15:2: 309-326.

Hull, F. 1973. Bee flies of the world: the gen­era of the fam­ily Bom­byli­idae (1973). Wash­ing­ton D.C.: Smith­son­ian In­sti­tu­tion Press.

Kits, J., S. Mar­shall, N. Even­huis. 2008. The bee flies (Diptera: Bom­byli­idae) of On­tario, with a key to the species of east­ern Canada. Cana­dian Jour­nal of Arthro­pod Iden­ti­fi­ca­tion, 6: 1-52.

McGin­ley, R. 1993. Where's the man­age­ment in col­lec­tion's man­age­ment: plan­ning for im­proved care, greater use, and growth of col­lec­tions.. Na­tional Mu­seum of Nat­ural His­tory, 1: 309-338.

Shel­don, J., P. Beed­low. 1988. Diets of Dark­ling Bee­tles (Coleoptera: Tene­bri­on­idae) Within a Shrub-Steppe Ecosys­tem. En­to­mo­log­i­cal So­ci­ety of Amer­ica, 81: 782-791.

Toft, K. 1983. Com­mu­nity pat­terns of nee­tiv­o­rous adult par­a­sitoids (Diptera, Bom­byli­idae) on their re­sources. Oe­colo­gia, 57: 200-215.

Wieg­mann, B. 2011. Over­com­ing the ef­fects of rogue taxa: Evo­lu­tion­ary re­la­tion­ships of the bee flies. PloS Cur­rents, 5: 1-13.

Yeates, D., D. Great­head. 1997. The evo­lu­tion­ary pat­tern of host use in the Bom­byli­idae (Diptera): a di­verse fam­ily of par­a­sitoid flies.. Bi­o­log­i­cal Jour­nal of the Lin­nean So­ci­ety, 50: 149-185.

Yeates, D., G. Dod­son. 1990. The Mat­ing Sys­tem of a Bee Fly (Diptera: Bom­byli­idae). I. Non-Re­source-Based Hill­top Ter­ri­to­ri­al­ity and a Re­source-Based Al­ter­na­tive. Jour­nal of In­sect Be­hav­ior, 3: 1-15.

Yeates, D., C. Lam­bkin. 1994. "Bom­byli­idae" (On-line). Tree of Life Web Pro­ject. Ac­cessed July 24, 2013 at http://​tolweb.​org/​Bombyliidae.

Yeates, D., C. Lam­bkin. 2013. "Fam­ily Bom­byli­idae- Bee Flies" (On-line). Bug Guide. Ac­cessed Au­gust 09, 2013 at http://​bugguide.​net/​node/​view/​185.

Yela, J., C. Her­rera. 1993. Sea­son­al­ity and life­cy­cles of woody plant feed­ing noc­tuid moths (Lep­i­doptera, Noc­tu­idae) in Mediter­ran­ian Habi­tats. Eco­log­i­cal En­to­mol­ogy, 18: 259-269.