Rhamphomyia Longicauda
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''Rhamphomyia longicauda'', the long-tailed dance fly, is a species of fly commonly found in eastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
that belongs to the family
Empididae __NOTOC__ Empididae is a family of flies with over 3,000 described species occurring worldwide in all the biogeographic realms but the majority are found in the Holarctic. They are mainly predatory flies like most of their relatives in the Empid ...
and part of the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
of dance flies
Empidoidea The Empidoidea are a large monophyletic superfamily of true flies, the sister taxon to the Muscomorpha (Cyclorrhapha). These two groups are sometimes united in the unranked taxon Eremoneura. There are some 10,000 known species within Empidoid ...
. It is included in the subgenus ''Rhamphomyia''. This species of fly is most known for sex role reversal during courtship, as females put on exaggerated displays and congregate in
leks Lek or LEK may refer to: * Lek mating, of animals * Albanian lek, currency * Lek (magazine), Norway * Lek (pharmaceutical company), now part of Sandoz * Lek (river), Netherlands * De Lek, Netherlands fiefdom * L.E.K. Consulting, firm * Leung ...
to attract males. Females cannot hunt for food, so they receive protein from
nuptial gift Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simply gametes in order to improve the reproductive fitness of the donor. Often, such a gift will improve the fit ...
s brought to them by males. Female dependence on males for nutrition is the principal cause for sex role reversal in this species of fly.


Description


Males

While this fly species is termed the long-tailed dance fly and its
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
nomenclature Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. (The theoretical field studying nomenclature is sometimes referred to as ''onymology'' or ''taxonymy'' ). The principl ...
resembles the same meaning (''Rhamphomyia'' = “crooked fly;” ''longicauda'' = longtailed), the “tail” actually references the male
genitalia A sex organ, also known as a reproductive organ, is a part of an organism that is involved in sexual reproduction. Sex organs constitute the primary sex characteristics of an organism. Sex organs are responsible for producing and transporting ...
, which forms an elbowed erection towards the rear of the
abdomen 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 ...
. Common to many male
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advance ...
, the
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
section of the eyes is larger than the
ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
section. The lengthened dorsal section, in addition to shortened
wings A wing is a type of fin that produces both lift and drag while moving through air. Wings are defined by two shape characteristics, an airfoil section and a planform. Wing efficiency is expressed as lift-to-drag ratio, which compares the bene ...
, aids the male's ability to approach the female from beneath during mating.


Females

Females have black scales that protrude forwards and backwards on their middle and
hind legs A hindlimb or back limb is one of the paired articulated appendages ( limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-H ...
. They also have small hairs that cover the sides of the legs and tan abdomens. In comparison to males, females have larger and inflatable pleural sacs, larger wings,
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of 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 di ...
length, and hind
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
l length, which are the principal
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
body parts. The ventral region of the female
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
is larger than the dorsal region, which helps them locate encroaching males that are pursuing mating. Males prefer females that have longer wings and shorter tibiae.


Territoriality

During the
mating season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and ch ...
(May–July), females congregate in
leks Lek or LEK may refer to: * Lek mating, of animals * Albanian lek, currency * Lek (magazine), Norway * Lek (pharmaceutical company), now part of Sandoz * Lek (river), Netherlands * De Lek, Netherlands fiefdom * L.E.K. Consulting, firm * Leung ...
10 minutes before sunset and remain together for up to 40 minutes just after dawn or before dusk. Females return to the same lek every night, even if already inseminated, unless there is significant inclement weather, like continued rain or winds above 14 km/h. Unique to this species, the lek site is not a grounded surface; they hover above the aggregation site in a horizontal plane .3 m-.6 m above the ground/vegetation under an opening in an otherwise enclosed
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
.


Food resources

Females cannot hunt for food: they only receive protein from
nuptial gift Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simply gametes in order to improve the reproductive fitness of the donor. Often, such a gift will improve the fit ...
s brought to them by males. Female dependence on males for nutrition is the principal cause for sex role reversal in this species of fly. Males hunt an hour before sunset within a range of 30m from the lek site. They rapidly scour the area in a figure-8 flight pattern looking for food. Males will eat any prey they can find, but they normally capture swarming
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s. More specifically, most tend to prey on flies (Diptera'')'', which make up 70% of their combined
prey 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 ki ...
. They also feed on
mayflies Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are aquatic insects belonging to the order ...
(Ephemeroptera) and
caddisflies The caddisflies (order Trichoptera) are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the basis ...
(Trichoptera), and infrequently feed on male
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s,
moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...
s, and
mosquito Mosquitoes, the Culicidae, are a Family (biology), family of small Diptera, flies consisting of 3,600 species. The word ''mosquito'' (formed by ''Musca (fly), mosca'' and diminutive ''-ito'') is Spanish and Portuguese for ''little fly''. Mos ...
es. The size of their captures varies significantly, ranging from 2–4.5mm in one 5-year study.


Mating


Courting

Before entering a lek, females rest on nearby vegetation and swallow air to inflate expandable pouches on the sides of the abdomen. The size of the abdomen is made to look even bigger by wrapping their large, scaly legs around their abdomen. Males enter female leks and discriminate first for female wing size. Model experiments showed that males give sexual preference to females with larger abdomen size. In females, there is a positive correlation between larger pleural sacs and greater egg length/number. Female abdomen size has evolutionarily correlated with extent of
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
maturation in other species of ''
Rhamphomyia ''Rhamphomyia'' is a genus of dance flies, in the fly family Empididae. It contains more than 600 species in 8 subgenera. Species The species of ''Rhamphomyia'' are arranged into subgenera, as follows: ;'' Aclonempis'' Collin, 1926 :*'' R ...
,'' and females take advantage of this pre-established male preference to gain mates. Overall abdomen size does correlate to more developed egg maturity in ''R. longicauda'', but inflation decreases the accuracy of body size as an indicator of egg maturity by 49%, making this a deceptive signal to entice males.


Leks

Females form leks in groups of 10–100 at sites under openings of an otherwise closed
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
. They return to the lek site every night to continue receiving nuptial gifts from the males; no virgins are found after almost two weeks of lek formation. Females hover over the lek site to provide them with backlighting so that they appear as silhouettes in order to exaggerate their size and deceive males. Stratification within leks was observed, as females with larger tibiae hover at lower positions in the swarm to receive the most copulations. Males never enter a lek without prey but go immediately to lek sites after catching
prey 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 ki ...
. They enter leks from below, or if they arrive before the females congregate, they fly by the swarm area several times and wait on nearby vegetation. Once females congregate, males fly below the females until she drops to him to receive his nuptial gift and mate.


Copulation

Males pass through the female swarm several times before finding a suitable mate. The male then ascends directly beneath the female about 1–2 cm below her, at which point the female drops down to the male. The two then fly away together at a faster speed away from the swarm, during which time the male gives the female the nuptial gift and recedes into copulating position. Flight pace slowed at the beginning of copulation. Nuptial flights last about 40 minutes.


Female-female interaction

Larger females hover at lower positions in the swarm to receive the most copulations. Horizontal positioning was also of importance, as central females also gained from more frequent encounters with males. However, unlike many male leks, females do not engage in physical altercations to receive better positioning within the lek.


Assortive mating

Females with the largest
ornaments An ornament is something used for decoration. Ornament may also refer to: Decoration *Ornament (art), any purely decorative element in architecture and the decorative arts *Ornamental turning *Biological ornament, a characteristic of animals tha ...
typically are more likely to attract a male in ''R. longicauda,'' and larger males typically mate with larger females, as they can bear their weight during the
nuptial flight Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as '' Rhamphomyia longicauda''. During the flight, virgin queens mate with males and then land ...
. During copulation, males must withstand the weight of the female and the
nuptial gift Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simply gametes in order to improve the reproductive fitness of the donor. Often, such a gift will improve the fit ...
during the flight, and because of this, there is no
directional selection In population genetics, directional selection is a type of natural selection in which one extreme phenotype is favored over both the other extreme and moderate phenotypes. This genetic selection causes the allele frequency to shift toward the ...
for increased body size or ornament size (pinnate leg scales and pleural sacs). Rather, female ''R. longicauda'' undergo
stabilizing selection Stabilizing selection (not to be confused with negative or purifying selection) is a type of natural selection in which the population mean stabilizes on a particular non-extreme trait value. This is thought to be the most common mechanism of ...
, as a female being too large lowers
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the capability to produc ...
since many males will not be able to perform copulation with her.


Enemies


Predators

The ''
Tetragnatha ''Tetragnatha'' is a genus of long-jawed orb-weavers found all over the world. It was first described by Pierre André Latreille in 1804, and it contains hundreds of species. Most occur in the tropics and subtropics, and many can run over water. ...
'' spider is a known predator of dance flies. It makes webs on the periphery of female swarms, making flies near the edge of the swarm more vulnerable to attack.


Victim dispersal

Another consequence to the sex role reversal is that females are also preyed upon at a higher frequency than males. Despite not
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
for food, more females are killed by spiders even in locations where both sexes are traversing because of the risky flight patterns females perform in order to get attention from males. This includes flying closer and more erratically towards nearby
vegetation Vegetation is an assemblage of plants and the ground cover they provide. It is a general term, without specific reference to particular Taxon, taxa, life forms, structure, Spatial ecology, spatial extent, or any other specific Botany, botanic ...
where spiders lay their webs. Researchers also note that the enlarged ornaments may also trigger higher predation rates. Neither enlarged ornaments nor risky flight patterns are observed in male dance flies.


External links

*


References

{{Taxonbar, from=Q13987289 Rhamphomyia Articles created by Qbugbot Insects described in 1861