Drosophila nigrospiracula: Difference between revisions
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'''''Drosophila nigrospiracula''''' is a fly species indigenous to the [[Sonoran Desert]], which spans Arizona, California, and part of the state of Sonora in Mexico. ''D. nigrospiracula'' share the Sonoran Desert with three other species of ''[[Drosophila]]'': ''D. pachea, D. mettleri'', and ''D. mojavensis''. ''D. nigrospiracula'' do not exhibit sexual isolation between the other species.<ref>{{cite journal | vauthors = Markow TA, Fogleman JC, Heed WB | title = Reproductive Isolation in Sonoran Desert Drosophila | journal = Evolution; International Journal of Organic Evolution | volume = 37 | issue = 3 | pages = 649–652 | date = May 1983 | pmid = 28563318 | doi = 10.2307/2408281 | jstor = 2408281 }}</ref> This fly breeds on the decomposing tissues of two species of cacti that are also endemic to the region: cardón ''(Pachycereus pringlei'' ) and saguaro ''(Carnegiea gigantea).'' |
'''''Drosophila nigrospiracula''''' is a fly species indigenous to the [[Sonoran Desert]], which spans Arizona, California, and part of the state of Sonora in Mexico. ''D. nigrospiracula'' share the Sonoran Desert with three other species of ''[[Drosophila]]'': ''[[Drosophila pachea|D. pachea]], [[Drosophila mettleri|D. mettleri]]'', and ''[[Drosophila mojavensis|D. mojavensis]]''. ''D. nigrospiracula'' do not exhibit sexual isolation between the other species.<ref>{{cite journal | vauthors = Markow TA, Fogleman JC, Heed WB | title = Reproductive Isolation in Sonoran Desert Drosophila | journal = Evolution; International Journal of Organic Evolution | volume = 37 | issue = 3 | pages = 649–652 | date = May 1983 | pmid = 28563318 | doi = 10.2307/2408281 | jstor = 2408281 }}</ref> This fly breeds on the decomposing tissues of two species of cacti that are also endemic to the region: cardón ''(Pachycereus pringlei'' ) and saguaro ''(Carnegiea gigantea).'' |
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== Taxonomy == |
== Taxonomy == |
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''D. nigrospiracula'' is part of the anceps [[species complex]].<ref name = "Alonso-Pimentel_1995">{{Cite journal| vauthors = Alonso-Pimentel H, Spangler HG, Heed WB |date= January 1995 |title=Courtship sounds and behaviour of the two saguaro-breeding Drosophila and their relatives |journal=Animal Behaviour |volume=50 |issue=4 |pages=1031–1039 |doi=10.1016/0003-3472(95)80103-0 }}</ref> This species is categorized as part of the |
''D. nigrospiracula'' is part of the anceps [[species complex]].<ref name = "Alonso-Pimentel_1995">{{Cite journal| vauthors = Alonso-Pimentel H, Spangler HG, Heed WB |date= January 1995 |title=Courtship sounds and behaviour of the two saguaro-breeding Drosophila and their relatives |journal=Animal Behaviour |volume=50 |issue=4 |pages=1031–1039 |doi=10.1016/0003-3472(95)80103-0 }}</ref> This species is categorized as part of the Mulleri subgroup, one of the five subgroups under the [[Drosophila repleta|repleta]] group of the ''Drosophila'' [[genus]].<ref>{{cite journal | vauthors = Durando CM, Baker RH, Etges WJ, Heed WB, Wasserman M, DeSalle R | title = Phylogenetic analysis of the repleta species group of the genus Drosophila using multiple sources of characters | journal = Molecular Phylogenetics and Evolution | volume = 16 | issue = 2 | pages = 296–307 | date = August 2000 | pmid = 10942616 | doi = 10.1006/mpev.2000.0824 }}</ref> The four ''Drosophila'' fly species inhabiting the Sonoran Desert are not phylogenetically related to each other but are all, under the repleta group. Each species came to inhabit this region independently, and they experience different resource availability. As such, they adapted uniquely to the environment of the Sonoran Desert and came to use different [[cactus]] species as a host.<ref name = "Frank_1992">{{cite journal | vauthors = Frank MR, Fogleman JC | title = Involvement of cytochrome P450 in host-plant utilization by Sonoran Desert Drosophila | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 89 | issue = 24 | pages = 11998–2002 | date = December 1992 | pmid = 1465429 | doi = 10.1073/pnas.89.24.11998 | bibcode = 1992PNAS...8911998F }}</ref> |
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''D. nigrospiracula'' is closely related, in a behavioral sense, to [[Drosophila mettleri]]. |
''D. nigrospiracula'' is closely related, in a behavioral sense, to [[Drosophila mettleri]]. |
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== Distribution == |
== Distribution == |
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Typical population sizes range from 361 to 3,999 flies.<ref name = "Johnston_1976">{{cite journal| vauthors = Johnston JS, Heed WB |date= July 1976 |title=Dispersal of Desert-Adapted Drosophila: The Saguaro-Breeding D. nigrospiracula |journal=The American Naturalist |volume=110 |issue=974 |pages=629–651 |doi=10.1086/283095 }}</ref> These flies commute to other saguaro cacti at rates three times higher than any other ''Drosophila'' species; they live in short-lived yet widespread habitats, requiring constant relocation to look for resources. ''D. nigrospiracula'' travel up to 2 km per day to gain resources between the low density of cacti.<ref>{{cite journal | vauthors = Hurtado LA, Erez T, Castrezana S, Markow TA | title = Contrasting population genetic patterns and evolutionary histories among sympatric Sonoran Desert cactophilic Drosophila | journal = Molecular Ecology | volume = 13 | issue = 6 | pages = 1365–75 | date = June 2004 | pmid = 15140083 | doi = 10.1111/j.1365-294X.2004.02169.x }}</ref> |
Typical population sizes range from 361 to 3,999 flies.<ref name = "Johnston_1976">{{cite journal| vauthors = Johnston JS, Heed WB |date= July 1976 |title=Dispersal of Desert-Adapted Drosophila: The Saguaro-Breeding D. nigrospiracula |journal=The American Naturalist |volume=110 |issue=974 |pages=629–651 |doi=10.1086/283095 }}</ref> These flies commute to other [[saguaro cacti]] at rates three times higher than any other ''Drosophila'' species; they live in short-lived yet widespread habitats, requiring constant relocation to look for resources. ''D. nigrospiracula'' travel up to 2 km per day to gain resources between the low density of cacti.<ref>{{cite journal | vauthors = Hurtado LA, Erez T, Castrezana S, Markow TA | title = Contrasting population genetic patterns and evolutionary histories among sympatric Sonoran Desert cactophilic Drosophila | journal = Molecular Ecology | volume = 13 | issue = 6 | pages = 1365–75 | date = June 2004 | pmid = 15140083 | doi = 10.1111/j.1365-294X.2004.02169.x }}</ref> |
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== Home Range == |
== Home Range == |
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[[File:Pachycereus pringlei (5780799263).jpg|alt=|thumb|A cardón cactus (''Pachycereus pringlei)'' found in the Baja Peninsula of the Sonoran Desert.]] |
[[File:Pachycereus pringlei (5780799263).jpg|alt=|thumb|A cardón cactus (''Pachycereus pringlei)'' found in the Baja Peninsula of the Sonoran Desert.]] |
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[[File:Carnegiea gigantea or Sahuaro or Saguaro.jpg|alt=|thumb|A saguaro cactus (''Carnegiea gigantea)'' found in its natural environment - the mainland of the Sonoran Desert.]] |
[[File:Carnegiea gigantea or Sahuaro or Saguaro.jpg|alt=|thumb|A saguaro cactus (''Carnegiea gigantea)'' found in its natural environment - the mainland of the Sonoran Desert.]] |
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''D. nigrospiracula'' are only found in the Sonoran Desert |
''D. nigrospiracula'' are only found in the Sonoran Desert on cardón ''([[Pachycereus pringlei]])'' and saguaro (''Carnegiea gigantean)'' cacti.<ref name = "Johnston_1976" /> Cardón and saguaro are tall, columnar cacti with similar morphology, chemical composition, and low alkaloid composition (their main allelochemical). Saguaro cactus grow on the mainland, while the cardón species grows only on the Baja peninsula. ''D. nigrospiracula'' is restricted to these two forms of cacti because they emit the [[alkaloid]] [[gigantine]] and [[carnegine]], which ''D. nigrospiracula'' can [[metabolize]].<ref name = "Martinson_2018">{{cite journal | vauthors = Martinson VG, Carpinteyro-Ponce J, Moran NA, Markow TA | title = A Distinctive and Host-Restricted Gut Microbiota in Populations of a Cactophilic Drosophila Species | journal = Applied and Environmental Microbiology | volume = 83 | issue = 23 | pages = e01551–17, /aem/83/23/e01551–17.atom | date = December 2017 | pmid = 28939605 | doi = 10.1128/AEM.01551-17 | editor-first = Andrew J. | editor-last = McBain | name-list-format = vanc }}</ref> In general, the two cacti species confer ''Drosophila'' species specificity through their [[allelochemical]] composition. These allelochemicals can be harmful to a fly without the proper allelochemical detoxification enzymes – namely the [[cytochrome p450]] system.<ref name = "Johnston_1976" /> In general, formation of plant decay can come from injury or freezing of the cactus. Then, exudate flows down the plant, creating a place for ''D. nigrospiracula'' to congregate. |
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=== Mechanism of metabolism of host plant === |
=== Mechanism of metabolism of host plant === |
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== Life History == |
== Life History == |
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''D. nigrospiracula'' eggs form in follicles that consist of 15 nurse cells, and a given female’s paired ovaries consist of hundreds of follicles.<ref name = "Frank_1992" /> The chromosomes of these flies are polyploidy and are actively transcribed to RNA, which is then passed to oocytes to help with embryogenesis. In the final stages of oogenesis, nurse cells dispose of their nuclear components into growing oocytes, providing them resources for embryonic development. |
''D. nigrospiracula'' eggs form in follicles that consist of 15 nurse cells, and a given female’s paired ovaries consist of hundreds of follicles.<ref name = "Frank_1992" /> The chromosomes of these flies are [[polyploidy]] and are actively transcribed to [[RNA]], which is then passed to [[oocytes]] to help with [[embryogenesis]]. In the final stages of oogenesis, nurse cells dispose of their nuclear components into growing oocytes, providing them resources for embryonic development. |
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=== Role of Phosphorus in Egg Formation === |
=== Role of Phosphorus in Egg Formation === |
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''D. nigrospiracula'' exhibit sexual dimorphism through differing ratios of phosphorus concentration in adult males and females.<ref name = "Frank_1992" /> Females have about 3 times as much phosphorus in their gonads as males. This inequality is due to the importance of phosphorus in the synthesis of nucleic acids during oogenesis. The cacti on which ''D. nigrospiracula'' breed and feed have low concentrations of phosphorus, but research has shown that the higher concentration in females comes from the phosphorus transferred from the male to the female during copulation. The phosphorus from the male ejaculate is incorporated into the female ovaries and traceable in RNA and DNA. Inside the female, phosphorus is then used to synthesize nucleic acids necessary in egg production. Reduced phosphorus levels in mature female diets slowed oogenesis. |
''D. nigrospiracula'' exhibit [[sexual dimorphism]] through differing ratios of phosphorus concentration in adult males and females.<ref name = "Frank_1992" /> Females have about 3 times as much phosphorus in their gonads as males. This inequality is due to the importance of phosphorus in the synthesis of nucleic acids during oogenesis. The cacti on which ''D. nigrospiracula'' breed and feed have low concentrations of [[phosphorus]], but research has shown that the higher concentration in females comes from the phosphorus transferred from the male to the female during copulation. The phosphorus from the male ejaculate is incorporated into the female ovaries and traceable in RNA and [[DNA]]. Inside the female, phosphorus is then used to synthesize [[nucleic acids]] necessary in egg production. Reduced phosphorus levels in mature female diets slowed oogenesis. |
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== Mating == |
== Mating == |
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=== Courtship === |
=== Courtship === |
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''D. nigrospiracula'' exhibit varied mating behavior.<ref name = "Alonso-Pimentel_1995" /> Males |
''D. nigrospiracula'' exhibit varied mating behavior.<ref name = "Alonso-Pimentel_1995" /> Males instigate courtship first by physical contact with females. They then perform wing displays while they approached the female from behind, often circling the female multiple times. The wing display consisted of quickly opening and then closing of one or both wings, and the males also paired this act with bursts of sound. Males were also observed to lick the female genitalia if the female stopped moving. Females showed willingness to mate by spreading their wings, while uninterested females relocated away from the male. Male ''D. nigrospiracula'' are persistent in courtship but do not mount an unwilling female. |
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== Enemies == |
== Enemies == |
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=== Macrocheles subbadius === |
=== Macrocheles subbadius === |
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''[[Macrocheles subbadius]]'' is an ectoparasite also indigenous to the North American region of the Sonoran Desert that inhabits both saguaro and cardón cacti. It reproduces on moist soil, animal feces, and rotting plant tissue. They obtain nutrients by biting into the integument of flies and intaking the [[hemolymph]] found in the body cavity of the fly, using it for nutrients. This parasite reduces the life span of ''D. nigrospiracula'' in a manner proportional to the amount of parasites found on the body. Reduction in overall health is due to physical trauma caused by ''Macrocheles subbadius'' and also due to the loss of nutrients. This parasite was also found to hinder female fertility and development of offspring.<ref name="Polak_1998">{{cite journal|vauthors=Polak M, Starmer WT|date=November 1998|title=Parasite-induced risk of mortality elevates reproductive effort in male Drosophila|journal=Proceedings. Biological Sciences|volume=265|issue=1411|pages=2197–201|doi=10.1098/rspb.1998.0559|pmc=1689506|pmid=9872007}}</ref> |
''[[Macrocheles subbadius]]'' is an [[ectoparasite]] also indigenous to the North American region of the Sonoran Desert that inhabits both saguaro and cardón cacti. It reproduces on moist soil, animal feces, and rotting plant tissue. They obtain nutrients by biting into the integument of flies and intaking the [[hemolymph]] found in the body cavity of the fly, using it for nutrients. This parasite reduces the life span of ''D. nigrospiracula'' in a manner proportional to the amount of parasites found on the body. Reduction in overall health is due to physical trauma caused by ''Macrocheles subbadius'' and also due to the loss of nutrients. This parasite was also found to hinder female fertility and development of offspring.<ref name="Polak_1998">{{cite journal|vauthors=Polak M, Starmer WT|date=November 1998|title=Parasite-induced risk of mortality elevates reproductive effort in male Drosophila|journal=Proceedings. Biological Sciences|volume=265|issue=1411|pages=2197–201|doi=10.1098/rspb.1998.0559|pmc=1689506|pmid=9872007}}</ref> |
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=== Effects of Parasitism on Sexual Reproduction === |
=== Effects of Parasitism on Sexual Reproduction === |
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== Microbiome == |
== Microbiome == |
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As mentioned, ''D. nigrospiracula'' use the necrotic tissues of the saguaro cactus for mating and also as their food source.<ref name = "Alonso-Pimentel_1995" /> However, the microbiota of ''D. nigrospiracula'' is significantly different from the microbiota found on cardón and saguaro cacti and actually is more similar to flies from different cacti from other regions. The most abundant of the bacteria does not come from the food on the cacti.<ref name = "Martinson_2018" /> These bacterial colonies were colonized through host-to-host transmission by other species and settings. |
As mentioned, ''D. nigrospiracula'' use the necrotic tissues of the saguaro cactus for mating and also as their food source.<ref name = "Alonso-Pimentel_1995" /> However, the microbiota of ''D. nigrospiracula'' is significantly different from the microbiota found on cardón and saguaro cacti and actually is more similar to flies from different cacti from other regions. The most abundant of the bacteria does not come from the food on the cacti.<ref name = "Martinson_2018" /> These bacterial colonies were colonized through host-to-host transmission by other species and settings. |
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== Genetics == |
== Genetics == |
||
This species is behaviorally most common to [[Drosophila mettleri|Drosophila ''mettleri'']]. A major difference between these two fly species, however, that serves to limit the amount of competition between them in nesting site selection is more genetically than behaviorally explained. D. ''mettleri'' contains an upregulation in the CYP28A1 gene in the P450 gene family. Upregulation in this gene enables D. ''mettleri'' to breed in both the toxic soil surrounding rotting cacti and on the tissues of rotting cacti patches. D. ''nigrospiracula,'' however, lacks this upregulated gene and can only nest on the tissues of rotting cacti patches. This particular genetic difference enables both species of Desert Drosopholids to use the same host cacti in the same geographic region without competitive pressures. |
This species is behaviorally most common to [[Drosophila mettleri|Drosophila ''mettleri'']]. A major difference between these two fly species, however, that serves to limit the amount of competition between them in nesting site selection is more genetically than behaviorally explained. D. ''mettleri'' contains an upregulation in the CYP28A1 gene in the P450 gene family. Upregulation in this gene enables D. ''mettleri'' to breed in both the toxic soil surrounding rotting cacti and on the tissues of rotting cacti patches. D. ''nigrospiracula,'' however, lacks this upregulated gene and can only nest on the tissues of rotting cacti patches. This particular genetic difference enables both species of Desert Drosopholids to use the same host cacti in the same geographic region without competitive pressures. |
Revision as of 01:45, 4 December 2019
Drosophila nigrospiracula | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Diptera |
Family: | Drosophilidae |
Genus: | Drosophila |
Species: | D. nigrospiracula
|
Binomial name | |
Drosophila nigrospiracula Patterson and Wheeler, 1942[1]
|
Drosophila nigrospiracula is a fly species indigenous to the Sonoran Desert, which spans Arizona, California, and part of the state of Sonora in Mexico. D. nigrospiracula share the Sonoran Desert with three other species of Drosophila: D. pachea, D. mettleri, and D. mojavensis. D. nigrospiracula do not exhibit sexual isolation between the other species.[2] This fly breeds on the decomposing tissues of two species of cacti that are also endemic to the region: cardón (Pachycereus pringlei ) and saguaro (Carnegiea gigantea).
Taxonomy
D. nigrospiracula is part of the anceps species complex.[3] This species is categorized as part of the Mulleri subgroup, one of the five subgroups under the repleta group of the Drosophila genus.[4] The four Drosophila fly species inhabiting the Sonoran Desert are not phylogenetically related to each other but are all, under the repleta group. Each species came to inhabit this region independently, and they experience different resource availability. As such, they adapted uniquely to the environment of the Sonoran Desert and came to use different cactus species as a host.[5]
D. nigrospiracula is closely related, in a behavioral sense, to Drosophila mettleri.
Distribution
Typical population sizes range from 361 to 3,999 flies.[6] These flies commute to other saguaro cacti at rates three times higher than any other Drosophila species; they live in short-lived yet widespread habitats, requiring constant relocation to look for resources. D. nigrospiracula travel up to 2 km per day to gain resources between the low density of cacti.[7]
Home Range
D. nigrospiracula are only found in the Sonoran Desert on cardón (Pachycereus pringlei) and saguaro (Carnegiea gigantean) cacti.[6] Cardón and saguaro are tall, columnar cacti with similar morphology, chemical composition, and low alkaloid composition (their main allelochemical). Saguaro cactus grow on the mainland, while the cardón species grows only on the Baja peninsula. D. nigrospiracula is restricted to these two forms of cacti because they emit the alkaloid gigantine and carnegine, which D. nigrospiracula can metabolize.[8] In general, the two cacti species confer Drosophila species specificity through their allelochemical composition. These allelochemicals can be harmful to a fly without the proper allelochemical detoxification enzymes – namely the cytochrome p450 system.[6] In general, formation of plant decay can come from injury or freezing of the cactus. Then, exudate flows down the plant, creating a place for D. nigrospiracula to congregate.
Mechanism of metabolism of host plant
The cytochrome p450 system is responsible for processing Pheromones and Steroids and also metabolizing insecticides, drugs, Mutagens, and Carcinogens. The p450 system allows Drosophila to metabolize and detoxify the secondary compounds that the cacti use as a function of herbivore defense in order to viably use the plant as a host.[6] One such other species of Drosophila that uses the p450 system to detoxify allelochemicals is Drosophila mettleri.
Life History
D. nigrospiracula eggs form in follicles that consist of 15 nurse cells, and a given female’s paired ovaries consist of hundreds of follicles.[5] The chromosomes of these flies are polyploidy and are actively transcribed to RNA, which is then passed to oocytes to help with embryogenesis. In the final stages of oogenesis, nurse cells dispose of their nuclear components into growing oocytes, providing them resources for embryonic development.
Role of Phosphorus in Egg Formation
D. nigrospiracula exhibit sexual dimorphism through differing ratios of phosphorus concentration in adult males and females.[5] Females have about 3 times as much phosphorus in their gonads as males. This inequality is due to the importance of phosphorus in the synthesis of nucleic acids during oogenesis. The cacti on which D. nigrospiracula breed and feed have low concentrations of phosphorus, but research has shown that the higher concentration in females comes from the phosphorus transferred from the male to the female during copulation. The phosphorus from the male ejaculate is incorporated into the female ovaries and traceable in RNA and DNA. Inside the female, phosphorus is then used to synthesize nucleic acids necessary in egg production. Reduced phosphorus levels in mature female diets slowed oogenesis.
Mating
Researchers observed statistical significance with regard to size of copulating males in nature.[9] Larger males tended to be more successful in mating than smaller males, as they win the male-male competition for females. Females approach males to mate at non-resource-based territories controlled by a male.
D. nigrospiracula has a sex ratio that favors females, as females are seen to mate multiple times in one morning. Additionally, females also are ready to mate 3 days after birth, but males do not reach sexual maturation until they are 9 to 10 days old. These two factors greatly increase the operational sex ratio in favor of females.
Courtship
D. nigrospiracula exhibit varied mating behavior.[3] Males instigate courtship first by physical contact with females. They then perform wing displays while they approached the female from behind, often circling the female multiple times. The wing display consisted of quickly opening and then closing of one or both wings, and the males also paired this act with bursts of sound. Males were also observed to lick the female genitalia if the female stopped moving. Females showed willingness to mate by spreading their wings, while uninterested females relocated away from the male. Male D. nigrospiracula are persistent in courtship but do not mount an unwilling female.
Enemies
Macrocheles subbadius
Macrocheles subbadius is an ectoparasite also indigenous to the North American region of the Sonoran Desert that inhabits both saguaro and cardón cacti. It reproduces on moist soil, animal feces, and rotting plant tissue. They obtain nutrients by biting into the integument of flies and intaking the hemolymph found in the body cavity of the fly, using it for nutrients. This parasite reduces the life span of D. nigrospiracula in a manner proportional to the amount of parasites found on the body. Reduction in overall health is due to physical trauma caused by Macrocheles subbadius and also due to the loss of nutrients. This parasite was also found to hinder female fertility and development of offspring.[10]
Effects of Parasitism on Sexual Reproduction
Despite energetic costs to courting and mating, D. nigrospiracula increased reproductive efforts when perturbed by parasites.[10] Younger males devoted even more time to courtship when affected by more parasites. The same phenomenon was also observed in a study with older males when lightly affected by parasites, but courtship levels dropped to below control levels when heavily parasitized. Researchers attribute this behavioral characteristic of D. nigrospiracula as an evolutionarily adaptive advantage, as the flies attempt to pass on their genes when lower life expectancy is detected.
However, while parasite-infested Drosophila look to mate at higher frequencies, another study shows that males and females are less likely to find mates.[11] Males have an even harder time finding mates than females even if infected by the same amount of M. subbadius. Furthermore, the intensity of female choice is directly proportional to the depth of parasitic infection. Researchers attribute these observations to reduced male-male competitive ability, decreased courtship vigor, and even physical prevention of copulation.
Microbiome
As mentioned, D. nigrospiracula use the necrotic tissues of the saguaro cactus for mating and also as their food source.[3] However, the microbiota of D. nigrospiracula is significantly different from the microbiota found on cardón and saguaro cacti and actually is more similar to flies from different cacti from other regions. The most abundant of the bacteria does not come from the food on the cacti.[8] These bacterial colonies were colonized through host-to-host transmission by other species and settings.
Genetics
This species is behaviorally most common to Drosophila mettleri. A major difference between these two fly species, however, that serves to limit the amount of competition between them in nesting site selection is more genetically than behaviorally explained. D. mettleri contains an upregulation in the CYP28A1 gene in the P450 gene family. Upregulation in this gene enables D. mettleri to breed in both the toxic soil surrounding rotting cacti and on the tissues of rotting cacti patches. D. nigrospiracula, however, lacks this upregulated gene and can only nest on the tissues of rotting cacti patches. This particular genetic difference enables both species of Desert Drosopholids to use the same host cacti in the same geographic region without competitive pressures.
References
- ^ Description of new species of the subgenera Hirtodrosophila and Drosophila. JT Patterson and MR Wheeler, 1942
- ^ Markow TA, Fogleman JC, Heed WB (May 1983). "Reproductive Isolation in Sonoran Desert Drosophila". Evolution; International Journal of Organic Evolution. 37 (3): 649–652. doi:10.2307/2408281. JSTOR 2408281. PMID 28563318.
- ^ a b c Alonso-Pimentel H, Spangler HG, Heed WB (January 1995). "Courtship sounds and behaviour of the two saguaro-breeding Drosophila and their relatives". Animal Behaviour. 50 (4): 1031–1039. doi:10.1016/0003-3472(95)80103-0.
- ^ Durando CM, Baker RH, Etges WJ, Heed WB, Wasserman M, DeSalle R (August 2000). "Phylogenetic analysis of the repleta species group of the genus Drosophila using multiple sources of characters". Molecular Phylogenetics and Evolution. 16 (2): 296–307. doi:10.1006/mpev.2000.0824. PMID 10942616.
- ^ a b c Frank MR, Fogleman JC (December 1992). "Involvement of cytochrome P450 in host-plant utilization by Sonoran Desert Drosophila". Proceedings of the National Academy of Sciences of the United States of America. 89 (24): 11998–2002. Bibcode:1992PNAS...8911998F. doi:10.1073/pnas.89.24.11998. PMID 1465429.
- ^ a b c d Johnston JS, Heed WB (July 1976). "Dispersal of Desert-Adapted Drosophila: The Saguaro-Breeding D. nigrospiracula". The American Naturalist. 110 (974): 629–651. doi:10.1086/283095.
- ^ Hurtado LA, Erez T, Castrezana S, Markow TA (June 2004). "Contrasting population genetic patterns and evolutionary histories among sympatric Sonoran Desert cactophilic Drosophila". Molecular Ecology. 13 (6): 1365–75. doi:10.1111/j.1365-294X.2004.02169.x. PMID 15140083.
- ^ a b Martinson VG, Carpinteyro-Ponce J, Moran NA, Markow TA (December 2017). McBain AJ (ed.). "A Distinctive and Host-Restricted Gut Microbiota in Populations of a Cactophilic Drosophila Species". Applied and Environmental Microbiology. 83 (23): e01551–17, /aem/83/23/e01551–17.atom. doi:10.1128/AEM.01551-17. PMID 28939605.
{{cite journal}}
: Unknown parameter|name-list-format=
ignored (|name-list-style=
suggested) (help) - ^ Markow, Therese (1988-07-01). "Reproductive Behavior of Drosophila melanogaster and D. nigrospiracula in the Field and in the Laboratory". Journal of Comparative Psychology (Washington, D.C. : 1983). 102 (2): 169–73. doi:10.1037/0735-7036.102.2.169.
{{cite journal}}
: Unknown parameter|name-list-format=
ignored (|name-list-style=
suggested) (help) - ^ a b Polak M, Starmer WT (November 1998). "Parasite-induced risk of mortality elevates reproductive effort in male Drosophila". Proceedings. Biological Sciences. 265 (1411): 2197–201. doi:10.1098/rspb.1998.0559. PMC 1689506. PMID 9872007.
- ^ Polak M, Markow TA (August 1995). "Effect of Ectoparasitic Mites on Sexual Selection in a Sonoran Desert Fruit Fly". Evolution; International Journal of Organic Evolution. 49 (4): 660–669. doi:10.2307/2410319. JSTOR 2410319. PMID 28565138.