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''Drosophila montana'', colloquially referred to as a fruit fly, is a species of
fly 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 advanced ...
belonging to the family
Drosophilidae The Drosophilidae are a diverse, cosmopolitan family of flies, which includes species called fruit flies, although they are more accurately referred to as vinegar or pomace flies. Another distantly related family of flies, Tephritidae, are true ...
and the genus
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
. It belongs to the montana phylad, which diverged from the D. virilis species group in South Asia before its migration into North America. It is typically found in the western United States, but has been seen in Europe and Asia. There are two color phases of the species, having either a yellowish or a blackish brown thorax. It is the species of Drosophila best adapted to cold environments.


Description

''D. montana'' were first described by Patterson and Wheeler in 1942. ''D. montana'' populations do not display homogeneous coloration. They have two forms of color, either light and dark. These possess a yellowish form or a blackish brown form, respectively. The light form has similar coloration to the species ''D. novamericana''. The primary physical characteristic of the dark form is a blackish brown
thorax 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 c ...
. The dark form of ''D. montana'' also possesses a slightly brownish black stripe along its back, spanning the length of its acrostichal region. They also have legs with brown coloration, and the joints at the tips of their extremities are black. Notable physical similarities exist between ''D. montana'' and ''D. flavomontana''. Early studies mistakenly collected ''D. flavomontana'' when searching for ''D. montana'', and species identification was only possible by observing fertility. While black
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
ria are typically attributed to ''D. montana'' and red puparia to ''D. flavomontana'', genetic analysis has shown ''D. montana'' may produce red puparia. The two species sometimes produce hybrids, suggesting their coexistence.


Distribution and habitat

Populations of ''D. montana'' have been historically found throughout North America, Europe, and Asia⁠—most notably in Vancouver, Canada, Finland, and throughout the United States. Within the US, they are primarily located at high elevations along the Rocky Mountains from northern Idaho to Arizona. They have also been found in the
Sierra Nevada The Sierra Nevada () is a mountain range in the Western United States, between the Central Valley of California and the Great Basin. The vast majority of the range lies in the state of California, although the Carson Range spur lies primar ...
mountain range and at lower altitudes within the
Cascade Range The Cascade Range or Cascades is a major mountain range of western North America, extending from southern British Columbia through Washington and Oregon to Northern California. It includes both non-volcanic mountains, such as the North Cascades, ...
and Coastal Range. ''Drosophila montana'' typically reside near bodies of water within boreal forest regions, primarily in latitudes above 40 degrees North. Populations of ''D. montana'' further south will favor environments of higher elevation. Among all species in the Drosophila genus, ''D. montana'' is the most cold-tolerant species, with necessary qualities to survive the conditions of high altitude environments. Their exceptional capacity among the genus Drosophila for tolerating the cold enables the species to survive in high altitude and high latitude environments, which is reflected in their residences of choice.


Phylogeny

The species belongs to the order of flies
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 advanced ...
and the genus
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
. The genus Drosophila includes the virilis group, which diverged in the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...
period, between 7 and 11 million years ago. This event divided the ''D.'' ''virilis'' species group into two clusters: the virilis and montana phylads''.'' The latter encompasses several lineages including ''D. montana.'' This divergence occurred in South Asia, prior to their dispersion into North America. The populations of ''D. montana'' in North America, Asia, and Europe diverged between 450,000 and 900,000 years ago.


Cold adaptation

The species ''D. montana'' has several distinctive adaptations contributing to their high tolerance of cold. These include a generally elevated resistance and an acclimation response to cold, which can be induced, as well as a photoperiodic female reproductive diapause.


Mating

Adult ''D. montana'' congregate in breeding areas, with multiple males pursuing a single female simultaneously. These breeding sites are generally on decomposing tree
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is c ...
and areas of sap flux. Mating occurs during the spring and summer months, with active reproduction prevalent up through July. By late August, most adults are in diapause and reproduction ceases. Like many species of Drosophila in temperate regions, there may be only one generation produced per year. This is likely facilitated by the ability to diapause.


Sexual conflict

Sexual conflict regarding the length of copulation is common for ''D. montana''. Females prefer much shorter mating durations, and engage in physical struggles to attempt to throw off a male partner. However, males are often able to maintain sexual contact much longer than females dictate. This allows males to induce females' resistance to copulations, thus preventing the occurrence of females remating. This activity functions as a means of mate guarding.


Courtship songs

Similar to many species of Drosophila, ''D. montana'' emit "songs" from wing vibrations. These songs are performed while courting females and are a requirement to initiate mating. Females will convey attraction to males by spreading their wings, to which the male will respond by initiating copulation. Males' songs are subject to significant sexual selection in the species. Songs with fast qualities, such as high
carrier frequency In telecommunications, a carrier wave, carrier signal, or just carrier, is a waveform (usually sinusoidal) that is modulated (modified) with an information-bearing signal for the purpose of conveying information. This carrier wave usually has a ...
, are associated with males with high reproductive success. Overlapping frequencies with environmental background noise can interfere with females' ability to detect the songs, which may compromise mate selection. With respect to the frequency of males' courtship songs, average
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
is approximately 14%. This differs significantly from other courtship song characteristics, which differ only by several percent due to inbreeding depression. This suggests that the fitness of male ''D. montana'' is associated with the frequency of their courtship songs. The frequency of males' songs is also subject to considerable
outbreeding depression In biology, outbreeding depression happens when crosses between two genetically distant groups or populations result in a reduction of fitness. This is particularly likely if the subspecies have different habitats or if no genetic exchange has oc ...
.


Cuticular hydrocarbons

''D. montana'' produce cuticular
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ...
s, commonly abbreviated CHCs, which act as olfactory or gustatory cues for attraction. While more effective in other species of Drosophila, CHCs are a significant factor in mate selection. In particular, female ''D. montana'' CHC characteristics are correlated with rates of female rejection of songs. This suggests that females' CHCs act as pheromones, attracting more males and therefore necessitating more rejections.


Reproductive diapause

The typical longevity of adult ''D. montana'' in the laboratory is about 30 to 40 days, with females slightly outliving males. Adult female ''D. montana'' use
photoperiod Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light a ...
to determine seasonal changes and inform their reproductive
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...
for overwintering. This allows them to reduce their aging, increase their chances of survival over the winter, and increase their longevity. ''D. montana'' develop from egg to adult in about 26 to 30 days, depending on population and photoperiodic conditions. Individuals in the diapause state have arrested development and reproduction. They also become reliant on energy reserves in favor of feeding, so larger individuals typically fare better during diapause. ''D. montana'' develop quickly under shorter photoperiods, representative of autumn daylight conditions, to facilitate diapause. This development time is influenced by post-
eclosion A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
photoperiodic cues.


Parental care

The desired locations of
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
for ''D. montana'' coincide with their breeding sites. Observed substrates for oviposition include phloem and sap flux yeast growths on
birch trees A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech-oak family Fagaceae. The genus ''Betula'' contains 30 to ...
, as well as decomposing stalks of the plant ''
Nuphar lutea ''Nuphar lutea'', the yellow water-lily, brandy-bottle, or spadderdock, is an aquatic plant of the family ''Nymphaeaceae'', native to northern temperate and some subtropical regions of Europe, northwest Africa, western Asia, North America, and ...
'', the yellow water lily. The virilis group of Drosophila, to which ''D. montana'' belongs, is generally known to oviposit in sap. The species has overlapping preferences for oviposition with ''D. littoralis''. However, ''D. montana'' mate slightly earlier, and their larvae are more prevalent in drier substrates.


Commensalism

The collection of virilis group flies, including ''D. montana'', is significantly more successful when close to known
beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers a ...
residences. Beavers preferentially reside in bodies of water close to specific tree species including
birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' cont ...
and
alder Alders are trees comprising the genus ''Alnus'' in the birch family Betulaceae. The genus comprises about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few sp ...
, which they feed on in the winter. The beavers' feeding activity produce fallen trees and stripped bark that supply ''D. montana'', as well as other members of the virilis group, with their preferred sites of oviposition. Alder trees are specifically cited as a favored site of ''D. montana'' oviposition. These shared resources support commensalism between the two species. For ''D. montana'', this relationship is typically observed in North America, specifically with the beaver species ''
Castor canadensis The North American beaver (''Castor canadensis'') is one of two extant beaver species, along with the Eurasian beaver (''Castor fiber''). It is native to North America and introduced in South America (Patagonia) and Europe (primarily Finland and ...
''.


References

{{Taxonbar, from=Q13598262
montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columb ...
Insects described in 1942