Scathophaga stercoraria | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Diptera |
Family: | Scathophagidae |
Genus: | Scathophaga |
Species: | S. stercoraria |
Binomial name | |
Scathophaga stercoraria | |
Synonyms | |
Scathophaga stercoraria, commonly known as the yellow dung fly or the golden dung fly, is one of the most familiar and abundant flies in many parts of the Northern Hemisphere. As its common name suggests, it is often found on the feces of large mammals, such as horses, cattle, sheep, deer, and wild boar, where it goes to breed. [1] The distribution of S. stercoraria is likely influenced by human agriculture, especially in northern Europe and North America. [2] The Scathophaga are integral in the animal kingdom due to their role in the natural decomposition of dung in fields. They are also very important in the scientific world due to their short life cycles and susceptibility to experimental manipulations; thus, they have contributed significant knowledge about animal behavior. [1]
Scathophaga stercoraria is sexually dimorphic, with an average lifespan of one to two months. The adult males are bright golden-yellow with orange-yellow fur on the front legs. Females are a little duller in color, with pronounced green-brown tinges, and no brightly colored fur on the front legs. The adults range from 5 to 11 mm in length, and the males are generally larger than the females. [2] The physical features of separate S. stercoraria populations can vary greatly, due in part to the range of locations in which the species is found. [1] Generally, they are located in cooler temperate regions, including North America, Asia, and Europe. They may also favor higher altitudes, such as the Pyrenees and Swiss Alps. [2]
The adults mainly prey on smaller insects, mostly other Diptera. They can also consume nectar and dung as additional sources of energy. In a laboratory setting, adult S. stercoraria can live solely on Drosophila and water. [3] Females spend most of their time foraging in vegetation and only visit dung pats to mate and oviposit on the dung surface. Both males and females are attracted to dung by scent, and approach dung pats against the wind. [3] Males spend most of their time on the dung, waiting for females and feeding on other insects that visit the dung, such as blow flies. In the absence of other prey, the yellow dung fly may turn to cannibalism. The larvae are coprophagous, relying on dung for nutrition. [2]
Scathophaga stercoraria breeds on the dung of many large mammals, but generally prefers fresh cattle dung. The operational sex ratio on these pats is very male-biased and competition is high. Females are small and have limited precopulatory choice. [4] Copulation lasts 20–50 minutes, after which the male attempts to guard the female from other males. Both males and females often mate with multiple partners. [5] Reproductive success depends on a variety of factors, including sperm competition, nutrition, and environmental temperature. [2]
Females have paired accessory glands, which supply lubricants to the reproductive system and secrete protein-rich egg shells. Sperm is received in a large structure called the bursa copulatrix, and is stored in a structure called the spermatheca. Scathophaga species have three spermathecae, (one pair and one singlet), each with its own narrow duct that connects it to the bursa. Sperm can be stored in the spermathecae for days, weeks, or even years, and sperm from several males can be stored simultaneously. [6] Males have two projections, the paralobes, which are used to hold onto a female during copulation. Between the paralobes is the intromittent organ, the aedeagus, which transfers sperm into the female's bursa copulatrix. [7]
During copulation, sperm is not directly deposited into sperm-storing organs. Ejaculation occurs in the bursa copulatrix, and then females actively move sperm into the spermathecae using their muscular spermathecal invagination to pump sperm into transit. This gives females a level of control over which and how much sperm enters her system, an example of cryptic female choice. Although current results are inconclusive regarding whether or not females are cryptically selecting for a better phenotypic match, a female may benefit from having variable sperm fertilizing her offspring. Such adaptations are advantageous because females benefit from being able to control which sperm are successful in fertilizing eggs. The females may not be aware of which sperm are better suited for her offspring, but simply that being able to control the proportion of sperm from multiple mates can maximize the possibility of an optimal phenotypic match. It is to her advantage to have multiple males' sperm reach her eggs, rather than just one. After copulation, females prefer to lay their eggs on the small hills of the dung surface, avoiding depressions and pointed areas. This survival strategy aims to prevent desiccation and drowning so the eggs are placed where they have the greatest chance of surviving. [6]
Many studies have studied the many manifestations of sexual conflict, including post-mating sexual selection, in the yellow dung fly. Sperm competition occurs when a female mates with multiple males. Each male's sperm is then in direct competition to fertilize the eggs. Sperm mix quickly once they reach the female's stores. The goal of males is to displace the sperm of other males as much as possible. Larger males tend to have longer copulation times and greater rates of sperm displacement. The fertilization success of males that were secondary mates increased as their body size relative to the first male increased. [8]
Traits such as body size, testis size, and sperm length are variable, as well as heritable in S. stercoraria males. Larger sperm may be advantageous if they have greater propulsion along the female's spermathecal duct, resulting in higher fertilization success rates. When competition among males is high and females are mating with multiple males, those with the largest testes also have the most success in terms of proportion of sperm that fertilize a female's eggs. The resulting male offspring would then have a similar advantage.
A positive correlation was found between sperm length of males and spermathecal duct length of females. The size of male testis was also positively correlated with female spermathecae size. Additionally, females with larger spermathecae are better able to produce spermicidal secretion. This cryptic female choice betters their ability to influence paternity over their offspring. These covariances are an example of an "evolutionary arms race". This suggests that each sex evolves certain traits to undermine the beneficial traits of the other, resulting in the coevolution of male and female reproductive systems of S. stercoraria. [9]
The eggs that the female lays on the dung hatch into larvae after 1–2 days, depending on temperature. The larvae quickly burrow into the dung for protection and feed on it. At 20 °C, larvae undergo three molts over five days, during which they grow exponentially. [3] After growth, larvae spend another five days emptying their stomachs before pupation, where no additional body mass is gained. After 10–20 days, the larvae burrow into the soil around and beneath the dung and pupate. The time needed for the juvenile flies to emerge can vary from 10 days at 25 °C to 80 days at 10 °C or less. The smaller females typically emerge a few days before the males. The fitness of the resulting juveniles is greatly dependent on the quality of the dung in which they were placed. Factors affecting dung quality include water content, nutritional quality, parasites, and drugs or other chemicals given to the animal. [2]
Yellow dung flies are anautogenous. To become sexually mature and produce viable eggs or sperm, they must feed on prey to acquire sufficient proteins and lipids. [2] Females under nutritional stress will have higher rates of egg mortality and less survival of offspring to adult emergence. [10] S. stercoraria females can then produce four to 10 clutches in their lifetimes. The adults are active throughout much of the year in most moderate climates. [5]
Yellow dung fly viability depends strongly on the environment. In warmer climates, a sharp population decline often occurs in the summer, when the temperatures increase above 25 °C. [3] Meanwhile, no population decline is seen in colder climates, such as Iceland, Finland, and northern England, and high elevations. Additionally, the number of generations per year varies with altitude and latitude, typically between two and four overlapping generations. [3] The end of winter synchronizes the first emergence in March, and the overwinter generations are produced in the fall. In northern Europe, where the mating season is shorter, only one or two generations can be expected. [3]
Yellow dung flies have extremely variable phenotypes – body size and development rate in particular. Proximate causes of variation include juvenile nutrition, temperature, predation, and genetic variation. [3] Much phenotypic plasticity in yellow dung flies is a result of food (dung) availability in the larval stage, which is often mediated by conspecific competition. Less dung results in more competitors, and more drying results in decreased growth rate and adult body size. [3] Additionally, when exposed to constant temperatures in a laboratory setting, higher temperatures during growth yield smaller flies. [3] Egg volume, but not clutch size, also decreases with increasing temperature. [3] Giving merit to the hypothesis that constraints on physiological processes at the cellular level account for temperature-mediated body size, studies have also shown that S. stercoraria body size varies via gene-by-environment interactions. Different cell lines vary significantly in growth, development, and adult body size in response to food limitation. [3]
Scathophaga stercoraria's phenotype has been shown to vary seasonally, latitudinally, and altitudinally as a result of an adaptive response to time constraints on development due to temperature changes. In the fall, as the temperature cools, the flies are able to increase development rate, so they can achieve the necessary, albeit smaller than average, size. [3] Furthermore, S. stercoraria development rate increases with increasing latitude. [3] This is likely an adaptive response to shorter mating seasons. Body size, but not development rate, vary with altitude. Dung flies are larger at higher altitudes as a result of colder temperatures. [3]
Larger yellow dung flies have a competitive advantage. Therefore, body size plasticity must be a survival mechanism. Offspring of large adults still survive under food limitations, despite needing more nutrients for a longer development. [3] Thus, the observed growth plasticity is a result of altering body chemistry and not differing survival rates of offspring from small and large parents. [3] Plastic development rate and body size are effective at avoiding premature death, meaning S. stercoraria adopts a strategy of being small and alive over large and dead. Smaller flies have an advantage in stressful environmental situations, due to larger dung flies needing more energy. [3] Additionally, low genetic differentiation exists between yellow dung fly populations, likely due to extensive gene flow, as S. stercoraria is able to travel great distances. [3] When species are unable to adapt through genetics, phenotypic plasticity is the most viable option to adjust to changing environments. Yellow dung flies develop in extremely variable environments, with pat drying, dung availability, and larval competition hindering survival. Therefore, phenotypic plasticity allows S. stercoraria to adjust development according to unpredictable ecological situations without genetic adaptation. [3]
Since S. stercoraria is a synanthropic fly, it does carry the risk of passively contaminating human food with various pathogens, molds, or yeasts. [11]
Some sexually transmitted diseases of insects are known, particularly in Coleoptera. Similar diseases have also been studied in S. stercoraria. Many of these sexually transmitted diseases are from multicellular ectoparasites (mites), protists, or the fungus Entomophthora muscae . These are frequently responsible for either sterilizing or killing the host fly. [12]
As well as being an easy meal for a great many bird and bat species, these flies are also preyed upon by other insects. Much competition exists between larvae of different species within a dung pat. Other insect species may also use the pats as hunting grounds. These include robber flies and clown beetles. [13]
Like Drosophila melanogaster , the yellow dung fly is an ideal model organism due to its short lifespan and susceptibility to various experimental manipulations. Initial interest in yellow dung flies came from their potential as biocontrol agents against pest flies around livestock. In the past 40 years alone, many studies have used S. stercoraria to research topics such as sperm competition, mating behavior, sexual conflict, reproductive physiology, thermal biology, and genetics. In particular, research on yellow dung flies has contributed greatly to understanding of multiple mating systems and sperm competition.
Recently, S. stercoraria was approved as a standard required test species for ecotoxicological testing. This includes evaluating the residues of veterinary drugs in livestock dung. [2] Yellow dung flies are a key part of decomposing waste in pastures, which is key to preventing the spread of endoparasites and returning nutrients to the soil. The species’ diet also serves to reduce the abundance of pest flies. [14] To test a chemical's toxicity, the chemical is mixed with bovine faeces, to which yellow dung fly eggs are added. Then, endpoints, such as sex and number of emerged adult flies, retardation of emergence, morphological change, and developmental rate, are measured and analyzed to determine toxicity. [15] A great deal of research has been done on the effects of avermectins on populations of S. stercoraria. Avermectins are used to control endoparasites in livestock. The resulting dung contains drug residues that can have unintentional adverse effects on yellow dung fly populations, such as increased mutations and decreased offspring viability. If the use of such drugs in agriculture is not carefully monitored, considerable economic losses could occur. [14]
Sex is the biological trait that determines whether a sexually reproducing organism produces male or female gametes. During sexual reproduction, a male and a female gamete fuse to form a zygote, which develops into an offspring that inherits traits from each parent. By convention, organisms that produce smaller, more mobile gametes are called male, while organisms that produce larger, non-mobile gametes are called female. An organism that produces both types of gamete is hermaphrodite.
Drosophila melanogaster is a species of fly in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly", "pomace fly", or "banana fly". In the wild, D. melanogaster are attracted to rotting fruit and fermenting beverages, and are often found in orchards, kitchens and pubs.
Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For internal fertilization to happen there needs to be a method for the male to introduce the sperm into the female's reproductive tract.
Sexual conflict or sexual antagonism occurs when the two sexes have conflicting optimal fitness strategies concerning reproduction, particularly over the mode and frequency of mating, potentially leading to an evolutionary arms race between males and females. In one example, males may benefit from multiple matings, while multiple matings may harm or endanger females due to the anatomical differences of that species. Sexual conflict underlies the evolutionary distinction between male and female.
Phormia regina, the black blow fly, belongs to the blow fly family Calliphoridae and was first described by Johann Wilhelm Meigen.
Nicrophorus orbicollis is a nearctic burying beetle first described by Thomas Say in 1825. It is a member of the genus Nicrophorus or sexton beetles, comprising the most common beetles in the family Silphidae. This species is a decomposer feeding on carcasses of small dead animals. N. orbicollis can be used for scientific research both medically and forensically.
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.
Bicyclus anynana is a small brown butterfly in the family Nymphalidae, the most globally diverse family of butterflies. It is primarily found in eastern Africa from southern Sudan to Eswatini. It is found mostly in woodland areas and flies close to the ground. Male wingspans are 35–40 mm and female wingspans are 45–49 mm.
Female sperm storage is a biological process and often a type of sexual selection in which sperm cells transferred to a female during mating are temporarily retained within a specific part of the reproductive tract before the oocyte, or egg, is fertilized. This process takes place in some species of animals. The site of storage is variable among different animal taxa and ranges from structures that appear to function solely for sperm retention, such as insect spermatheca and bird sperm storage tubules, to more general regions of the reproductive tract enriched with receptors to which sperm associate before fertilization, such as the caudal portion of the cow oviduct containing sperm-associating annexins. Female sperm storage is an integral stage in the reproductive process for many animals with internal fertilization. It has several documented biological functions including:
Interlocus sexual conflict is a type of sexual conflict that occurs through the interaction of a set of antagonistic alleles at two or more different loci, or the location of a gene on a chromosome, in males and females, resulting in the deviation of either or both sexes from the fitness optima for the traits. A co-evolutionary arms race is established between the sexes in which either sex evolves a set of antagonistic adaptations that is detrimental to the fitness of the other sex. The potential for reproductive success in one organism is strengthened while the fitness of the opposite sex is weakened. Interlocus sexual conflict can arise due to aspects of male–female interactions such as mating frequency, fertilization, relative parental effort, female remating behavior, and female reproductive rate.
Coelopa frigida is a species of seaweed fly or kelp fly. It is the most widely distributed species of seaweed fly. It can be found on most shorelines in the temperate Northern Hemisphere. Other species of seaweed flies include Coelopa nebularum and Coelopa pilipes. C. frigida feeds primarily on seaweed, and groups of C. frigida flies tend to populate near bodies of water. Climate change has led to an increase in C. frigida blooms along shores, which creates a pest problem for human beach-goers. C. frigida is also an important organism for the study of sexual selection, particularly female choice, which is influenced by genetics.
Sepsis cynipsea is a European species of fly and member of the family Sepsidae. It is a coprophagous fly that feeds on dung. These flies are most commonly found around freshly laid cattle dung where they eat and reproduce. Due to human agricultural practices involving cows, these flies are now common in other areas of the world.
Sexual selection in insects is about how sexual selection functions in insects. The males of some species have evolved exaggerated adornments and mechanisms for self-defense. These traits play a role in increasing male reproductive expectations by triggering male-male competition or influencing the female mate choice, and can be thought of as functioning on three different levels: individuals, colonies, and populations within an area.
Sexual selection in amphibians involves sexual selection processes in amphibians, including frogs, salamanders and newts. Prolonged breeders, the majority of frog species, have breeding seasons at regular intervals where male-male competition occurs with males arriving at the waters edge first in large number and producing a wide range of vocalizations, with variations in depth of calls the speed of calls and other complex behaviours to attract mates. The fittest males will have the deepest croaks and the best territories, with females making their mate choices at least partly based on the males depth of croaking. This has led to sexual dimorphism, with females being larger than males in 90% of species, males in 10% and males fighting for groups of females.
Polyandry in fishes is a mating system where females mate with multiple males within one mating season. This type of mating exists in a variety of animal species. Polyandry has been found in both oviparous and viviparous bony fishes and sharks. General examples of polyandry occur in fish species, such as green swordtails and Trinidadian guppies. Specific types of polyandry have also been classified, such as classical polyandry in pipefish cooperative polyandry in cichlids and convenience polyandry in sharks.
Cryptic female choice is a form of mate choice which occurs both in pre and post copulatory circumstances when females in certain species use physical or chemical mechanisms to control a male's success of fertilizing their ova or ovum; i.e. by selecting whether sperm are successful in fertilizing their eggs or not. It occurs in internally-fertilizing species and involves differential use of sperm by females when sperm are available in the reproductive tract.
In behavioral ecology, polyandry is a class of mating system where one female mates with several males in a breeding season. Polyandry is often compared to the polygyny system based on the cost and benefits incurred by members of each sex. Polygyny is where one male mates with several females in a breeding season . A common example of polyandrous mating can be found in the field cricket of the invertebrate order Orthoptera. Polyandrous behavior is also prominent in many other insect species, including the red flour beetle, the adzuki bean weevil, and the species of spider Stegodyphus lineatus. Polyandry also occurs in some primates such as marmosets, mammal groups, the marsupial genus' Antechinus and bandicoots, around 1% of all bird species, such as jacanas and dunnocks, insects such as honeybees, and fish such as pipefish.
Chionea scita is a species of crane fly in the family Limoniidae. C. scita is known as a type of snow crane fly because it is commonly seen walking on piles of snow during the winter months. These flies are also often observed in caves and heavily wooded areas. C. scita flies are small, hairy, wingless, and somewhat spider-like in appearance, unlike other flies.
Derocephalus angusticollis is a fly in the family Neriidae. They are typically found on the east coast of Australia near rotting vegetation. Aggregating on the rotting bark of trees such as Acacia longifolia and other trees in New South Wales and southern Queensland. Derocephalus angusticollis flies found in the wild have accelerated speeds of development and age of mortality when compared to those in captivity. One characteristic of the neriid fly is that it demonstrates sexual dimorphism. Males have a larger build as well as exaggerated physical characteristics such as wider heads and longer limbs. Certain phenotypic characteristics are dependent on the diet of the parents.
Sepsis thoracica, more commonly known as the black scavenger fly, a species of fly from the genus Sepsis and the family Sepsidae. It was discovered by Robineau-Desvoidy in 1830. It resembles a small flying ant. The fly is most commonly found inhabiting cow dung.
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