Derocephalus angusticollis

Last updated

Derocephalus angusticollis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Neriidae
Genus: Derocephalus
Species:
D. angusticollis
Binomial name
Derocephalus angusticollis
Enderlein, 1922 [1]
Synonyms

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. [3] D. angusticollis flies found in the wild have accelerated speeds of development and age of mortality when compared to those in captivity. Derocephalus 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.

Contents

D. angusticollis practices polyandrous mating behavior. Males take part in the practice of mate guarding where they use their legs to keep hold of the female during copulation and to push competing males away. Sexual conflict can be found in this fly as females have coiled oviducts that lead to their spermathecae. In order for the male to reach the spermathecae to deposit their sperm, antagonistic coevolution of the males having a flexible aedeagus occurred.

Description

Like other flies within Neriidae, D. angusticollis has characteristic dorsocentral bristles located for the most part on their thorax as an identification marker. [4] The number of pairs of bristles varies within the genus; D. angusticollis usually has two pairs instead of one pair. [5] Variation has been observed on parts other than the thorax. Certain flies had a lack of bristles or fewer than the average due to genetic factors and environmental factors, such as the quality of diet. [4]

Sexual dimorphism

Derocephalus angusticollis flies demonstrated sexual dimorphism with males often presenting as larger and with more exaggerated traits than females. [5] Males, in particular, tend to have elongated heads, antennae, and legs than their female counterparts. [6] The extent of the dimorphism is dependent on genetics as well as parental diet as described below. The exaggerated features of males may assist them in male-male combat. [6]

Diet

The flies' diet consists of rotting vegetation, bark, mold, and sap. [7] One particular study found that a nutrient-rich diet for this fly was one of a higher protein and carbohydrate content. [8] The quality of the diets of parents, in particular, has been found to have effects on the phenotype of the offspring. Mothers with nutrient-rich diets were found to lay larger eggs that also developed faster. [7] Likewise, fathers that were raised on high-nutrient diets had larger adult offspring, improving the overall fitness of the offspring. [7] [9]

Researchers have demonstrated that early differences in nutrient content can impact D. angusticollis's ability to age successfully. [10] Male early life condition was manipulated by varying nutrient content of larval diet, and the interactions that adult males had later on in life with other adult males were also intentionally controlled and manipulated in the experiment. Males fed with high nutrient levels as larvae were able to develop more quickly, and reached their reproductive peak earlier on. [10] Males with a nutrient-rich diet also experienced more rapid reproductive aging and ultimately died sooner than their poorly fed counterparts. [10] High nutrient condition early in life is associated with rapid aging even in the absence of male-male conflicts that damage one's longevity. [10]

The effect of the quality of food having an effect on the fitness of the male is specific to their developmental diets . Where male D. angusticollis flies reared on nutrient-poor diets would develop smaller testes and accessory glands, while those that were already adults on a rich diet showing no effects. [11] The amount of protein as well as the age of the adult male can also affect the viability of their offspring. Where younger male adults with less protein in their diet had less viable offspring, yet as they grow older the viability of their offspring increases. [12]

Mating

Derocephalus angusticollis practice polyandrous mating behavior. [13] The possibility of a non-genetic mechanism of inheritance has been explored in this species. Researchers have investigated D. angusticollis to search for evidence of telegony, which suggests that previous partners might be able to influence traits in offspring raised by subsequent males mating with the same female. [13] Males are able to transmit environmentally obtained conditions through paternal effects on the offspring's body size. These effects were experimentally shown to come from a previous male that the female had mated with, not the genetic father of the offspring. [13] The presence of telegony was only observed when females were exposed to the first male after mating, implying that the mechanism is semen-dependent, and not correlated with female choice mechanisms. [13]

Morphology

Male

The male genitalia consists of two main components: the aedeagus and the epandrium. The epandrium, which is found at the distal end of the abdomen just before the aedeagus, is the structure used by males to raise the female oviscape in order to gain access to the reproductive duct and facilitate the insertion of the aedeagus. The male's aedeagus consists of three parts: a basal, middle, and distal section. The basal and middle section that are connected by a hinge that allows the aedeagus to be flexible. Its flexibility, in turn, gives males the ability to unfold their aedeagus once inside a female, maneuver through the coiled ducts of females in order to reach the site of the spermathecae, and release their sperm. Compared to T. lineolatus flies, D. angusticollis have a larger rigid spike at the end of their distal section. When genitalia is retracted, the distal section's flexibility allows it to be coiled up at the base of the epandrium. [5]

Female

The female reproductive anatomy consists of three main components: the oviduct, bursa copulatrix, and spermatheca. When not in use for copulation the bursa copulatrix is bent in an S-shape instead of straight. Derocephalusangusticollis females have three spermathecae surrounded by muscle and are spherical in shape: one at the anterior duct and the other two at the posterior duct. The ducts that lead to these spermathecae, the site where the sperm is stored, are winding and coiled which in turn has led to an example antagonistic coevolution as males, in turn, developed a flexible aedeagus. [5]

Sexual competition

Fighting

Males have been found to engage in fighting over territory, in particular, for aggregate sites where female mates are abundant. The dominant male that wins will continue to defend the territory after the fight is over, while the subordinate male moves to a less densely populated area. Competition between males begins by assessing the size of their opponent, and the decision of whether or not to challenge an opponent depends on size. [9] Only if the flies are similarly sized will they fight. Compared to other flies of the same family, D. angusticollis have been found to avoid fighting with males that are larger than themselves. [5] As such, if the condition that the opponent is of similar size or smaller is met, the male challenges his rival by raising the anterior portion of its body and forelegs in a steep angle. In battle, males will then use their head, antennae, and forelegs against their opponent . [6]

Courtship and mate guarding

Copulation is instigated by the act of the male mounting a female from behind. [5] Researchers have observed, however, that this fly does take part in mate guarding in which the male will use their legs to hold onto the female as well as use their forelegs to prevent other males from getting close to the female during copulation. [14]

Sexual conflict

Females have been observed to resist mating with a male both before and during copulation. Before copulation, a female may resist by running away, not raising her oviscape (thus preventing the male from having access to her reproductive tract), or using her back legs to kick the males that attempt to mount her. If a male is still able to mount and engage in copulation, females can continue to resist by using their legs to kick the male off. Another means of sexual conflict is cryptic female choice, in which the coiling of the female reproductive tract has coevolved in such a way that successful males would be those that are able to maneuver such coils in order for their sperm to reach the spermathecae. [5]

Social behavior

Females are less active and spend most of their time in the same area drinking sap from damaged trees. [15] Females may aggregate at the same oviposition site as well as act more aggressively with others. Their aggressive behaviors are different from males such as flicking their wings or using their forelegs to combat other females. [16] Conversely, males are more active in moving about aggregation sites in competing for, searching for, and copulating with mates. Some have argued that this more active behavior increases their visibility and probability of coming into contact with predators. [15]

Chemical signaling

Derocephalus angusticollis use chemical signaling by their levels of cuticular hydrocarbons (CHCs) during their social interactions. Both genders have been observed to alter the levels of CHCs depending on where they fall in the social hierarchy of dominance. Subordinates show similar patterns between sexes while dominant flies have different patterns of CHC levels. [16]

Enemies

The main predator that targets this species is skinks (Eulamprus tenuis). In the wild, skinks sneak up on the flies from below. During certain studies that required flies to be identified by markers, there was a concern that the markers increased risk of predation, as the markers were attached to the abdomen to make them visible from above. Due to predators sneaking onto their prey from below, however, researchers have speculated that identification markers on the top part of the flies as used in observations may not make them more visible, as skinks' line of sight is limited to the bottom half of the fly. [15]

Lifespan in captivity versus in the wild

In a study that compared wild and captive D. angusticollis flies, researchers observed differences in both the speed of development as well as the age of mortality. Flies in the wild had accelerated development and did not live as long as those in captivity. This is likely due to varying conditions in the wild (harsh temperatures, risk of predation) as opposed to controlled conditions in the lab. [15]

Related Research Articles

<span class="mw-page-title-main">Mating</span> Process of pairing in biology

In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. Fertilization is the fusion of two gametes. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization. Mating may also lead to external fertilization, as seen in amphibians, fishes and plants. For most species, mating is between two individuals of opposite sexes. However, for some hermaphroditic species, copulation is not required because the parent organism is capable of self-fertilization (autogamy); for example, banana slugs.

<span class="mw-page-title-main">Sperm competition</span> Reproductive process

Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and variety of mates increases a female's chance to produce more viable offspring. However, multiple mates for a female means each individual male has decreased chances of producing offspring. Sperm competition is an evolutionary pressure on males, and has led to the development of adaptations to increase male's chance of reproductive success. Sperm competition results in a sexual conflict between males and females. Males have evolved several defensive tactics including: mate-guarding, mating plugs, and releasing toxic seminal substances to reduce female re-mating tendencies to cope with sperm competition. Offensive tactics of sperm competition involve direct interference by one male on the reproductive success of another male, for instance by physically removing another male's sperm prior to mating with a female. For an example, see Gryllus bimaculatus.

<span class="mw-page-title-main">Reproductive success</span> Passing of genes on to the next generation in a way that they too can pass on those genes

Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves.

<span class="mw-page-title-main">Sexual conflict</span> Term in evolutionary biology

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.

<span class="mw-page-title-main">Neriidae</span> Family of flies

The Neriidae are a family of true flies (Diptera) closely related to the Micropezidae. Some species are known as cactus flies, while others have been called banana stalk flies and the family was earlier treated as subfamily of the Micropezidae which are often called stilt-legged flies. Neriids differ from micropezids in having no significant reduction of the fore legs. Neriids breed in rotting vegetation, such as decaying tree bark or rotting fruit. About 100 species are placed in 19 genera. Neriidae are found mainly in tropical regions, but two North American genera occur, each with one species, and one species of Telostylinus occurs in temperate regions of eastern Australia.

<span class="mw-page-title-main">Sexual cannibalism</span> Practice of animals eating their own mating partners

Sexual cannibalism is when an animal, usually the female, cannibalizes its mate prior to, during, or after copulation. It is a trait observed in many arachnid orders and several insect and crustacean clades. Several hypotheses to explain this seemingly paradoxical behavior have been proposed. The adaptive foraging hypothesis, aggressive spillover hypothesis and mistaken identity hypothesis are among the proposed hypotheses to explain how sexual cannibalism evolved. This behavior is believed to have evolved as a manifestation of sexual conflict, occurring when the reproductive interests of males and females differ. In many species that exhibit sexual cannibalism, the female consumes the male upon detection. Females of cannibalistic species are generally hostile and unwilling to mate; thus many males of these species have developed adaptive behaviors to counteract female aggression.

<i>Phormia regina</i> Species of fly

Phormia regina, the black blow fly, belongs to the blow fly family Calliphoridae and was first described by Johann Wilhelm Meigen.

<i>Scathophaga stercoraria</i> Species of fly

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. The distribution of S. stercoraria is likely influenced by human agriculture, especially in northern Europe and North America. 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.

Bateman's principle, in evolutionary biology, is that in most species, variability in reproductive success is greater in males than in females. It was first proposed by Angus John Bateman (1919–1996), an English geneticist. Bateman suggested that, since males are capable of producing millions of sperm cells with little effort, while females invest much higher levels of energy in order to nurture a relatively small number of eggs, the female plays a significantly larger role in their offspring's reproductive success. Bateman's paradigm thus views females as the limiting factor of parental investment, over which males will compete in order to copulate successfully.

<span class="mw-page-title-main">Female sperm storage</span>

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, but not in humans. 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:

Sexual antagonistic co-evolution is the relationship between males and females where sexual morphology changes over time to counteract the opposite's sex traits to achieve the maximum reproductive success. This has been compared to an arms race between sexes. In many cases, male mating behavior is detrimental to the female's fitness. For example, when insects reproduce by means of traumatic insemination, it is very disadvantageous to the female's health. During mating, males will try to inseminate as many females as possible, however, the more times a female's abdomen is punctured, the less likely she is to survive. Females that possess traits to avoid multiple matings will be more likely to survive, resulting in a change in morphology. In males, genitalia is relatively simple and more likely to vary among generations compared to female genitalia. This results in a new trait that females have to avoid in order to survive.

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.

<i>Sepsis cynipsea</i> Species of fly

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.

A nuptial gift is a nutritional gift given by one partner in some animals' sexual reproduction practices.

<span class="mw-page-title-main">Sexual selection in amphibians</span> Choice of and competition for mates

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.

Sexual coercion among animals is the use of violence, threats, harassment, and other tactics to help them forcefully copulate. Such behavior has been compared to sexual assault, including rape, among humans.

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.

Drosophila nigrospiracula is a fly species indigenous to the Sonoran Desert, spanning Arizona, Baja California, and part of Sonora, Mexico. D. nigrospiracula share the Sonoran Desert with three other species of Drosophila: D. pachea, D. mettleri, and D. mojavensis. 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).

Glyphidops flavifrons is a member of the Neriidae family of the order Diptera. This fly is found in the southern United States, Central America, and South America. Historically, it has also been called Oncopsia seductrix Hennig or Oncopsia mexicana.G. flavifrons live, reproduce, and lay their eggs on the bark of trees in the early stages of decay. In this species, it is common to see the male flies to exhibit aggression in the presence of the females. These males may attack the copulating pair to help decrease the chances of other males mating and increase their own chances.

<i>Prochyliza xanthostoma</i> Species of fly

Prochyliza xanthostoma, the waltzing fly, is a species of carrion-feeding cheese skipper, insects in the family Piophilidae and the order Diptera. P. xanthostoma is a member of the genus Prochyliza, which contains eleven species. The adult flies are found through North America and are brown-bodied, with orange and black coloring. Mating occurs on animal carcasses and male perform mating rituals; females engage in ejaculate feeding. The waltzing fly is known for its exaggerated sexual dimorphism and has thus become a prominent model for sexual dimorphism and larval behavior. These organisms are known as cheese skippers because when startled, the larvae can leap several inches into the air. P. xanthostoma is an important model organism for sexual selection, larval behavior, and adult reproductive success and survivability.

References

  1. Enderlein, Günther (1922). "Klassifikation der Micropeziden". Archiv für Naturgeschichte. Abteilung A. 88 (4): 140–229. Retrieved 29 January 2021.
  2. Cresson, E. T. Jr. (1926). "Descriptions of new genera and species of Diptera (Ephydridae and Micropezidae)". Transactions of the American Entomological Society. 52: 249–274.
  3. Cassidy EJ, Bath E, Chenoweth SF, Bonduriansky R (February 2014). "Sex-specific patterns of morphological diversification: evolution of reaction norms and static allometries in neriid flies". Evolution; International Journal of Organic Evolution. 68 (2): 368–83. doi: 10.1111/evo.12276 . PMID   24111624.
  4. 1 2 Bonduriansky R (April 2009). "Condition dependence of developmental stability in the sexually dimorphic fly Derocephalus angusticollis (Diptera: Neriidae)". Journal of Evolutionary Biology. 22 (4): 861–72. doi: 10.1111/j.1420-9101.2009.01686.x . PMID   19220651.
  5. 1 2 3 4 5 6 7 Bath E, Tatarnic N, Bonduriansky R (2012-12-01). "Asymmetric reproductive isolation and interference in neriid flies: the roles of genital morphology and behaviour". Animal Behaviour. 84 (6): 1331–1339. doi:10.1016/j.anbehav.2012.08.025. ISSN   0003-3472. S2CID   53191020.
  6. 1 2 3 Bonduriansky R (January 2007). "The evolution of condition-dependent sexual dimorphism". The American Naturalist. 169 (1): 9–19. doi:10.1086/510214. PMID   17206580. S2CID   17439073.
  7. 1 2 3 Bonduriansky R, Head M (November 2007). "Maternal and paternal condition effects on offspring phenotype in Derocephalus angusticollis (Diptera: Neriidae)". Journal of Evolutionary Biology. 20 (6): 2379–88. doi: 10.1111/j.1420-9101.2007.01419.x . PMID   17956399.
  8. Runagall-McNaull A, Bonduriansky R, Crean AJ (June 2015). "Dietary protein and lifespan across the metamorphic boundary: protein-restricted larvae develop into short-lived adults". Scientific Reports. 5: 11783. Bibcode:2015NatSR...511783R. doi:10.1038/srep11783. PMC   4484247 . PMID   26119686.
  9. 1 2 Adler MI, Bonduriansky R (2012). "Paternal Effects on Offspring Fitness Reflect Father's Social Environment". Evolutionary Biology. 40 (2): 288–292. doi:10.1007/s11692-012-9211-6. S2CID   9765190.
  10. 1 2 3 4 Hooper AK, Spagopoulou F, Wylde Z, Maklakov AA, Bonduriansky R (March 2017). "Ontogenetic timing as a condition-dependent life history trait: High-condition males develop quickly, peak early, and age fast". Evolution; International Journal of Organic Evolution. 71 (3): 671–685. doi: 10.1111/evo.13172 . PMID   28067402. S2CID   16594401.
  11. Macartney, Erin L.; Nicovich, Philip R.; Bonduriansky, Russell; Crean, Angela J. (2018). "Developmental diet irreversibly shapes male post-copulatory traits in the neriid fly Telostylinus angusticollis". Journal of Evolutionary Biology. 31 (12): 1894–1902. doi: 10.1111/jeb.13384 . ISSN   1420-9101. PMID   30267554.
  12. Macartney, E. L.; Crean, A. J.; Bonduriansky, R. (2017). "Adult dietary protein has age- and context-dependent effects on male post-copulatory performance". Journal of Evolutionary Biology. 30 (9): 1633–1643. doi: 10.1111/jeb.13087 . ISSN   1420-9101. PMID   28386961.
  13. 1 2 3 4 Crean AJ, Kopps AM, Bonduriansky R (December 2014). "Revisiting telegony: offspring inherit an acquired characteristic of their mother's previous mate". Ecology Letters. 17 (12): 1545–52. doi:10.1111/ele.12373. PMC   4282758 . PMID   25270393.
  14. Bonduriansky R (May 2006). "Convergent evolution of sexual shape dimorphism in Diptera". Journal of Morphology. 267 (5): 602–11. doi:10.1002/jmor.10426. PMID   16477603. S2CID   15548020.
  15. 1 2 3 4 Kawasaki N, Brassil CE, Brooks RC, Bonduriansky R (September 2008). "Environmental effects on the expression of life span and aging: an extreme contrast between wild and captive cohorts of Telostylinus angusticollis (Diptera: Neriidae)". The American Naturalist. 172 (3): 346–57. doi:10.1086/589519. PMID   18710341. S2CID   14750605.
  16. 1 2 Wylde, Zachariah; Adler, Lewis; Crean, Angela; Bonduriansky, Russell (2019-12-01). "Perceived dominance status affects chemical signalling in the neriid fly Telostylinus angusticollis". Animal Behaviour. 158: 161–174. doi:10.1016/j.anbehav.2019.10.013. ISSN   0003-3472. S2CID   208048582.