Phormia


Phormia regina, the black blow fly, belongs to the blow fly family Calliphoridae and was first described by Johann Wilhelm Meigen. It is the only species in the genus Phormia.
The black blow fly's wings are specialized with a sharp bend. These flies also have well-developed calypters. Blow flies generally are about the size of a house fly or a little larger, and many are metallic blue or green in color. Key characteristics of this species include black gena, mostly white calypters, and anterior thoracic spiracles that appear to be orange yellow due to being surrounded by bright orange setae.
Phormia regina is especially important to forensic entomology. Female P. regina, like other blow flies in the family Calliphoridae, oviposit their eggs on carrion, where they hatch. The larvae develop through three instar stages until pupation. Adult black blow flies aggregate on feces in order to mate. The success of these mating interactions has been studied and appears to be related to size and diet of the adult flies. Following successful mating encounters, the adult females are then attracted to decaying material rather quickly for oviposition, allowing forensic entomologists to use development stages of larvae as a clue in determining an approximate time of death. P. regina are extremely common in the United States and other areas in North America. Combined with their importance in forensic entomology, these flies have been relatively well studied, particularly for variables affecting their development.

Taxonomy

Phormia regina was described by the German entomologist Johann Wilhelm Meigen in 1826. Its specific epithet is derived from the Latin word regina meaning 'queen'.

Description

Phormia regina adults are metallic blue or green in color and have a distinctive set of orange setae near the anterior thoracic spiracle. They are also recognizable by their black gena, mostly white calypteres, and the distinctive bend in their wings.

Distribution

Phormia regina is commonly found across the entirety of the United States as well as across other regions in North America

Habitat

Phormia regina can be found on human corpses, animal carcasses, and feces. Researchers have observed these flies on a wide variety of animal excrement, but at least in the northwestern United States, they exhibit a preference for human, mink, and swine feces. P. regina is predominantly found in the northern United States during spring and summer months, but in the winter they are localized to more southern regions. This localization is tentatively linked to the importance of temperature on their development. These flies prefer rural areas, especially near water sources, as opposed to urban spaces—at least in the San Francisco Bay Area.

Life history

The life cycle and development of Phormia regina is similar to that of most other Dipteran species, in which females oviposit their eggs onto a nutrient substrate. Then, after hatching, the larvae feed throughout three instar stages until they have stored up enough calories to commence pupation and finally emerge as adult blow flies. Each transition from first, second, and third instar is marked by a molt, and eventually the third-instar larvae develop sclerotized casings which envelop and protect them throughout metamorphosis.

Larval development

Relatively few studies have been conducted on the adults of this species in comparison to those on larval development, mostly due to the importance of blow fly larvae in determining the post mortem interval of corpses during investigations by forensic entomologists. For this reason, many researchers have conducted experiments to investigate the effect of various environmental factors on the duration time of larval development in this species.
Because the larval life cycle of this species is dependent on a climate with temperatures ranging from 12.7°C to 35 °C, it tends to inhabit the northern regions of the United States during summer months and southern regions in the winter. Researchers have discovered that at 40 to 45 °C, larval development occurs normally until the prepupal stage, at which point a majority of the larvae die. The few able to pupate at these high temperatures do not emerge as adults. The lowest temperature threshold for this species was found to be 12.5 °C below which females will not oviposit. The highest rate of development was seen at a constant temperature of 35 °C, where the average time of adult emergence was 265 hours. Flies at constant temperatures between 15 and 30 °C developed slower, with flies at the coolest temperatures taking the longest. Cyclic temperatures ranges of 25 to 35 °C and 15 to 25 °C proved to decrease the rate of development when compared to constant temperatures.
Also, studies have been conducted to assess the effects of light exposure on developmental variability in larvae. Larvae exposed to cyclic photoperiods have higher rates of development than larvae exposed to constant photoperiods. These findings suggest that darkness may be a stimulus for larval growth. However, these variations in light photoperiods failed to influence pupal duration times.

Adult development

While this species is in the wild, dung constitutes a majority of the nutritional intake used for sexual development in both male and female adults, but diets with higher protein intake better facilitate mating ability of both sexes. Female sexual maturity requires the completion of 10 stages of follicle development in the ovaries to produce eggs that are completely mature, and ovaries in females which have been deprived of a high-protein diet do not develop fully. Although it is possible for females on a diet of only dung to reach the final stage of sexual maturity, it takes much more time than if they were to feed exclusively on beef liver; even then, a lower percentage of those feeding on dung will have fully developed. An experiment by Stoffolano demonstrated this, revealing that 100% of females feeding exclusively on beef liver were able to reach the final stage of sexual development after 13 days, while only 78% of females were able to do so when feeding exclusively on pig dung over a 20-day period. Although some Dipteran species oviposit on dung, P. regina females exclusively deposit their eggs on carrion.
The neuroendocrine system in adult males, which controls their mating behavior, must be stimulated before they will mate with a female. Protein in the male diet is not necessarily needed for this stimulation, but Stoffolano also saw that higher percentages of females were successfully inseminated by male specimens which had been fed either dung or beef liver versus specimens with a diet of only sugar. Protein is not generally necessary for spermatogenesis in male flies, but it is paramount for accessory reproductive gland development, higher rates of copulation, and the capability of impregnating females. An additional study found a positive correlation between male head size and the size of the aedeagi, which has been proposed as a possible reason for lower percentages of insemination between small males and large females within this species.

Food sources

Feeding habits vary greatly throughout the life cycle of P. regina. The feeding habits of adult flies depend both on sex and reproduction status of the fly. This is particularly important with respect to their protein intake, which literature suggests has an important role in sexual maturation and copulation ability in adult P. regina. Like most other flies, the black blow fly feeds via sponging. Though adults require various types of nutrients, in the wild they are typically able to procure all their nutrition from feces.

Larvae

Adult females oviposit eggs on carrion. These eggs hatch into larvae that feed on the decaying tissue.

Adult females

Virgin adult females, when given access to protein and carbohydrate sources, utilize both for food. However, after their emergence as adults, they will gradually increase their protein intake until they are approximately 8 to 10 days old. From here they will continue to intake protein at a relatively constant rate that is lower than their required carbohydrate intake. Once a female has mated, either for the first time or all subsequent times, a spike in protein intake occurs following oviposition of her eggs.

Adult males

Protein intake in adult male black blow flies follows similar trends to that of virgin adult females; however, they require less total protein than females. The males will gradually increase their protein intake for the first 8–10 days of adulthood. They will then drop their protein intake to a relatively low and constant amount.

Mating

Adult black blow flies aggregate on feces, particularly animal dung pats, not only because they act as a food source, but also in order to mate. Some research has suggested a preference for settling on human, swine, and mink feces, but this may be localized to the northwestern region of the United States. P. regina is a panmictic species. Research supports mating success of adult males positively correlates to their size. Larger males have larger aedeagus which leads to more successful insemination of females. The mechanism of how and why larger aedeagus lead to better insemination is unknown.

Female reproductive system

Female black blow flies each have a pair of ovaries and lateral oviducts, one common oviduct, a vagina, and a bursa copulatrix, which is simply a sac-like cavity within the female that is roughly equivalent to a uterus. Each ovary has a lateral oviduct; these two lateral oviducts meet to form the common oviduct and connect the ovaries to the burs copulatrix. Three spermathecae are connected to the bursa copulatrix via spermathecal ducts, and two accessory glands attach to the bursa copulatrix near the spermathecae entries. The vagina is connected to the bursa copulatrix on the opposite end of the ovaries and allows the male aedeagus access to the bursa copulatrix and spermathecae.