Stalk-eyed fly

Stalk-eyed flies are insects of the fly family Diopsidae. The family is distinguished from most other flies by the possession of "eyestalks": projections from the sides of the head with the eyes at the end. Some fly species from other families such as Drosophilidae, Platystomatidae, Richardiidae, and Tephritidae have similar heads, but the unique character of the Diopsidae is that their antennae are located on the stalk, rather than in the middle of the head as in all other flies.

The stalk-eyed flies are up to a centimeter long, and they feed on both decaying plants and animals. Their unique morphology has inspired research into how the attribute may have arisen through forces of sexual selection and natural selection. Studies of the behavior of the Diopsidae have yielded important insights into the development of sexual ornamentation, the genetic factors that maintain such a morphological feature, sexual selection, and the handicap principle.

Stalk-eyed flies
Diopsid2
A diopsid from Cameroon
Scientific classification
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Diopsidae
Genera
Synonyms
  • Centrioncidae[1]

Distribution and habitat

More than 100 species in the Diopsidae are known, with the greatest diversity found in the Old World tropics.[2] They are distributed throughout the region, with the best-known species being from Southeast Asia and Southern Africa. Also, two species in North America have been described and a European species has recently been found in Hungary.[3]

Adult diopsids are typically found on low-lying vegetation in humid areas, often near streams and rivers, where they feed on fungi and bacteria, which they scavenge from decaying vegetation. The larvae are saprophagic or phytophagous, eating decaying and fresh plant matter. Diopsis macrophthalma Dalman, 1817, is a pest of rice and sorghum in tropical Africa.

The peculiar morphology of stalk-eyed flies makes it easy to identify their fossils (e.g. in amber); one prehistoric genus that has been discovered in that form is Prosphyracephala.[4]

Morphology

The Diopsidae are small to medium-sized flies, ranging from about 4.0 to about 12.0 mm in length. Their heads are subtriangular, with transverse eye stalks in all genera except the African genus Centrioncus and Teloglabrus. The head is usually sparsely haired, with vibrissae (whiskers) absent.[5]

The posterior portion of the fly's metathorax, or scutellum, has a pair of stout processes, and often the laterotergite (one of a number of lateral flanges) of the postnotum (a small dorsal sclerite on the insect thorax posterior to the notum) has a dome-like swelling or spine-like process. The anterior femora of the legs are stout, with ventral spines. Adult males have lost tergites seven and eight, and the seventh sternite forms a complete ventral band.[5]

Stalk-eyed flies, as the name implies, typically possess eyestalks (in all but the two genera listed above). Their eyes are mounted on projections from the sides of the head, and the antennae are located on the eyestalks, unlike stalk-eyed flies from other families. Though both males and females of most species have eyestalks, they are much longer in males, a sexual dimorphism thought to be due to sexual selection.[6] A rather remarkable feature of stalk-eyed flies is their ability, shortly after they emerge from their pupae, to ingest air through their oral cavity and pump it through ducts in the head to the tips of the eye stalks, thereby elongating them while they are still soft and transparent.[7]

Taxonomy

True stalk-eyed flies are members of the family Diopsidae, first described by Fothergill[8] and named by Carl Linnaeus in 1775.[9] The family Diopsidae is contained within the order Diptera and suborder Cyclorrapha, and features some 150 described species.[10]

The African genus Centrioncus (once placed in Sepsidae, but then moved to Diopsidae) was once recommended to be treated as a separate family, Centrioncidae, a sister group of the diopsids,[11] but since then this lineage has usually been treated as a subfamily.[12]

Behavior

Vision

Despite the unusual morphology of the eye, each compound eye sees a region of space extending over more than a hemisphere in all directions. Thus, extensive binocular overlap occurs, with about 70% of the ommatidia of each eye having a binocular partner ommatidium in the opposite eye which views in the same direction.[13][14] The binocular field is most extensive in the frontoventral quadrant, where it reaches over 135°, and is smallest in the dorsal region. The behavior of stalk-eyed flies is very much determined by vision. During the day, temporary territories may be defended by threatening behavior. At dusk, the animals gather in small groups on selected thread-like structures, returning to the same site each day. When males of about equal size encounter one another within such a group, they may engage in ritualized fights (or occasionally contact fights). Competitors are driven away by the dominant male. Conspecifics are most likely to elicit a threat or flight reaction when they are at a distance of about 50 mm, and reactions to model flies and reflections in a mirror also occur at about this distance.[13]

Mating

Teleopsis dalmanni
Male Teleopsis dalmanni

Stalk-eyed flies roost at night on root hairs hanging by streams. Mating usually takes place in the early morning in the vicinity of their roosts. Females show a strong preference for roosting and mating with males with longer eyestalks, and males compete with each other to control lekking aggregations through ritualized contest. This contest involves males facing one another and comparing their relative eye spans, often with the front legs spread apart, possibly to emphasize their eye-stalk lengths.[15] Male stalk-eyed flies with long eyestalks gain mating advantages both because of female choice and because they are better able to compete with rival males.[2][16]

Sexual selection

Though the evolution of exaggerated male traits as a result of female mate choice was at one point controversial, the Diopsidae are now regarded as a classic example of animals that exhibit sexually selected traits.[6][17][18] One view maintains that male ornaments co-evolve with female preferences. The selection of an ornamented mate causes genes that influence expression of the selected male trait and genes coding for female preference for this trait to be passed on to offspring[19][20] This process creates linkage disequilibrium between selected alleles, with the magnitude of resulting genetic correlations influencing evolutionary outcomes. If the genetic correlation is high relative to the heritability of the male ornament, then a runaway process can occur leading to extreme sexually selected traits, such as the incredible eye spans observed in male stalk-eyed flies. Otherwise, the trait and preference for the trait increase until natural selection against further trait elaboration balances sexual selection.[19]

Teleopsis dalmanni 2
Close-up of a male Teleopsis dalmanni

The extreme morphology exhibited by stalk-eyed flies (especially males) has been studied in an effort to support the hypothesis that exaggerated male traits could evolve through female mate choice and that the selection on male ornaments should cause a correlated response in female preferences. Researchers noted that the flies roosted along stream banks in peninsular Malaysia and that the males with the largest eye spans were accompanied by more females than males with shorter eye spans. From January to October, the researchers counted males and females on 40 root hairs along a single 200-m stretch of stream bank to confirm this observation.[21]

Sexual selection experiments

Researchers collected stalk-eyed flies and observed their behavior under laboratory conditions. In the lab, each individual was scored for eye spans, body length, age, and fecundity.[22] Pairs of males differing in eye span but matched in body length were conducted to quantify mate choice in the presence and absence of male interactions. Test males with the longest or shortest eye-span to body-length ratios were mated with 25 randomly chosen females. Wilkinson and Reillo then tested female choice in the presence and absence of male competition and in the presence of males with abnormally long and abnormally short eye spans.[21]

Males dispersed themselves, while females clustered in certain areas of the cage. As observed prior to the study, researchers found that the average number of females per male increased with male eye span in field collected aggregations of stalk-eyed flies. Under laboratory conditions, researchers found that female preferences for male characteristics changed as the males sexual characteristics changed. After 13 generations of artificial selection, they found that long eye-span male line females (i.e. females whose fathers had long eye spans) preferred long eye spans in both the selected males and in males that were not bred through artificial selection, while short eye-span male line females (i.e. females whose fathers had short eye spans) found short eye spans to be the most attractive, even over males with long eye spans. Because researchers kept the females separate from males prior to mate selection, the finding supported the hypothesis that the change in female mate choice was genetically based and not learned. Thus, stalk-eyed flies have been able to evolve sexual trait in males that corresponds directly to traits that affect mating choices made by females.[21]

Stalk-eyed fly (Diasemopsis) (4561140578)
Stalk-eyed fly (Diasemopsis)

Handicap selection

However, the evolution of extreme morphology in male flies and the corresponding evolution of female preference for these characteristics as an effect of sexual selection is only half the picture.[23] Handicap models of sexual selection predict that male sexual ornaments have strong condition-dependent expression, and this allows females to evaluate male genetic quality.[24][25][26][27][28]

Genetic variation underlies the response to environmental stress, such as variable food quality, of male sexual ornaments, such as the increased eye span, in the stalk-eyed fly.[23] Some male genotypes develop large eye spans under all conditions, whereas other genotypes progressively reduce eye spans as environmental conditions deteriorate. Several nonsexual traits, including female eye span and male and female wing length, also show condition-dependent expression, but their genetic response is entirely explained by scaling with body size. Unlike these characteristics, male eye span still reveals genetic variation in response to environmental stress after accounting for differences in body size. Thus, it could be inferred that these results strongly support the conclusion that female mate choice yields genetic benefits for offspring as eye span acts as a truthful indicator of male fitness. Eye span is, therefore, selected not only on the basis of attractiveness, but also because it demonstrates good genes in mates.[23]

Furthermore, some populations of stalk-eyed fly females carry a meiotic drive gene on their X chromosomes that causes female-biased sex ratios.[29] In these populations, males which carry a gene to suppresses X chromosome meiotic drive have longer eyestalks. Thus, females that mate with these males gain a direct genetic benefit by producing male offspring in a female-biased population. In other words, the gene for long eye-stalks is linked to a gene that makes males sire more male offspring.[30] Alternatively, long stalks may signal fertility, perhaps by encouraging females to use the sperm of a long-stalked male so as to produce more fertile sons.[30]

References

  1. ^ Feijen, Hans R. (1983). "Systematics and phylogeny of Centrioncidae, a new afromontane family of Diptera (Schizophora)". Zoologische Verhandelingen. 202: 1–137. Retrieved 30 October 2016.
  2. ^ a b G. S. Wilkinson & G. N. Dodson (1997). "Function and evolution of antlers and eye stalks in flies". In J. Choe & B. Crespi (eds.). The Evolution of Mating Systems in Insects and Arachnids. Cambridge: Cambridge University Press. pp. 310–327. ISBN 978-0-521-58976-5.
  3. ^ L. Papp, M. Földvári & P. Paulovics (1997). "Sphyracephala europaea sp. n. (Diptera: Diopsidae) from Hungary represents a family new to Europe". Folia Entomologica Hungarica. 58: 137–146.
  4. ^ Schumann, H (1994). "Diopsidae from Saxon Amber (Diptera, Diopsidae)". Deutsche Entomologische Zeitschrift. 41 (1): 141–145. doi:10.1002/mmnd.19940410111.
  5. ^ a b Peterson, B. V. (1987). "Diopsidae." In McAlpine, J. F., B. V. Peterson, G. E. Sherwell, H. J. Tekey, J. R. Vockerorth, and D. M. Wood, (cords.). Manual of Nearctic Diptera. Vol. 2: 785–789.
  6. ^ a b Davies N, Krebs J, and West S. (2012). An Introduction to Behavioral Ecology, 4th Ed. Wiley-Blackwell; Oxford: pp. 196-198.
  7. ^ Buschbeck, E. K., Roosevelt, J. L. and Hoy, R. R. (2001). Eye stalks or no eye stalks: A structural comparison of pupal development in the stalk-eyed fly Cyrtodiopsis and in Drosophila. J. Comp. Neurol., 433: 486–498.
  8. ^ Shillito, J. (1976). Fothergill and Linnaeus: The background of De Bigis Insectorum, 1775. Biol. J. Linn. Soc., 8; 75-86.
  9. ^ Shillito, J. (1976). Bibliography of the Diopsidae-II. Journal of the Society for the Bibliography of Natural History. Volume 8, Page 65-73.
  10. ^ Allaby, M. (1999). "Diopsidae." A Dictionary of Zoology. Encyclopedia.com. (December 15, 2012). http://www.encyclopedia.com/doc/1O8-Diopsidae.html
  11. ^ Feijen, H.R. (1983). "Systematics and phylogeny of Centrioncidae, a new afromontane family of Diptera (Schizophora)". 202 (1): 1–137.
  12. ^ Meier, R. & Hilger, S. (2000). "On the egg morphology and phylogenetic relationships of Diopsidae (Diptera: Schizophora)". Journal of Zoological Systematics and Evolutionary Research. 38 (1): 1–36. doi:10.1046/j.1439-0469.2000.381128.x.
  13. ^ a b D. Burkhardt & I. de la Motte (1983). "How stalk-eyed flies eye stalk-eyed flies: Observations and measurements of the eyes of Cyrtodiopsis whitei (Diopsidae, Diptera)". Journal of Comparative Physiology A. 151 (4): 407–421. doi:10.1007/BF00605457.
  14. ^ I. de la Motte & D. Burkhardt (1983). "Portrait of an Asian stalk-eyed fly". Naturwissenschaften. 70 (9): 451–461. doi:10.1007/BF01079611.
  15. ^ Wilkinson, G. (2001). Model Syestems in Behavioral Ecology: Integrating Conceptual, Theoretical, and Empirical Approaches. Ed. Lee Alan Dugatkin. Princeton University Press; Princeton: pp. 84-91.
  16. ^ T. Chapman, A. Pomiankowski & K. Fowler (2005). "Quick guide: stalk-eyed flies". Current Biology. 15 (14): R533–R535. doi:10.1016/j.cub.2005.07.015. PMID 16051154. Archived from the original on 2006-10-18.
  17. ^ Kirkpatrick, M and M.J. Ryan. (1991). “The evolution of mating preferences and the paradox of the lek.” Nature, Lond. 350: 33-38.
  18. ^ Maynard Smith, J. (1991). “Theories of sexual selection.” Trends Ecol. Evol. 6: 146-151.
  19. ^ a b Lande, R. (1981). “Models of speciation by sexual selection on polygenic traits.” Proc. Natn. Acad. Sci. U.S.A. 78: 3721-3725.
  20. ^ Kirkpatrick, M. (1982). “Sexual selection and the evolution of female choice.” Evolution 36: 1-12.
  21. ^ a b c Wilkinson, G. and P. Reillo. (1994). “Female choice response to artificial selection on an exaggerated male trait in a stalk-eyed fly.” Proc. R. Soc. Lond. B. 255: 1-6.
  22. ^ Wilkinson, G. (1993). “Artificial sexual selection alters allometry in the stalk-eyed fly Cyrtodiopsis dalmanni.” Genet. Res. 62: 212-222.
  23. ^ a b c P. David, T. Bjorksten, K. Fowler & A. Pomiankowski (2000). "Condition-dependent signalling of genetic variation in stalk-eyed flies". Nature. 406 (6792): 186–188. doi:10.1038/35018079. PMID 10910358.CS1 maint: Multiple names: authors list (link)
  24. ^ Andersson, M. (1986). “Evolution of condition-dependent sex ornaments and mating preferences: sexual selection based on viability differences.” Evolution 40: 804–816.
  25. ^ Pomiankowski, A. (1987). “Sexual selection: the handicap principle does work—sometimes.” Proc. R. Soc. Lond. B 231: 123–145.
  26. ^ Grafen, A. (1990). “Biological signals as handicaps.” J. Theor. Biol. 144: 517–546.
  27. ^ Iwasa, Y. and A. Pomiankowski. (1994). “The evolution of mate preferences for multiple handicaps.” Evolution 48: 853 –867.
  28. ^ Rowe, L. and D. Houle. (1996). “The lek paradox and the capture of genetic variance by condition dependent traits.” Proc. R. Soc. Lond. B 263: 1415–1421.
  29. ^ G. S. Wilkinson, D. C. Presgraves &. L. Crymes (1998). "Male eye span in stalk-eyed flies indicates genetic quality by meiotic drive suppression". Nature. 391 (6664): 276–279. doi:10.1038/34640.
  30. ^ a b Zimmer, Carl (2008). "The Evolution of Extraordinary Eyes: The Cases of Flatfishes and Stalk-eyed Flies". Evolution: Education and Outreach. 1 (4): 487–492. doi:10.1007/s12052-008-0089-9.

External links

Biological ornament

A biological ornament is a characteristic of an animal that appears to serve a decorative function rather than a utilitarian function. Many are secondary sexual characteristics, and others appear on young birds during the period when they are dependent on being fed by their parents. Ornaments are used in displays to attract mates, which may lead to the evolutionary process known as sexual selection. An animal may shake, lengthen, or spread out its ornament in order to get the attention of the opposite sex, which will in turn choose the most attractive one with which to mate. Ornaments are most often observed in males and choosing an extravagantly ornamented male benefits females because the genes that produce the ornament will be passed on to her offspring, increasing their own reproductive fitness. As Ronald Fisher noted, the male offspring will inherit the ornament while the female offspring will inherit the preference for said ornament, which can lead to a positive feedback loop known as a Fisherian runaway. These structures serve as cues to animal sexual behaviour, that is, they are sensory signals that affect mating responses. Therefore, ornamental traits are often selected by mate choice.

Diasemopsis

Diasemopsis is a genus of stalk-eyed flies in the family Diopsidae. They are known from sub-Saharan Africa.

Diopsoidea

The Diopsoidea are a small but diverse cosmopolitan superfamily of acalyptrate muscoids, especially prevalent in the tropics. Some flux exists in the family constituency of this group (e.g., the Megamerinidae are sometimes placed here, but may also be placed in the Nerioidea), and the final classification is likely to require the use of molecular systematics.

Eyestalk

In anatomy, an eyestalk (sometimes spelled as eye stalk or known as an ommatophore) is a protrusion that extends the eye away from the body, giving the eye a better field of vision. It is a common feature in nature and frequently appears in fiction.

Handicap principle

The handicap principle is a hypothesis originally proposed in 1975 by Israeli biologist Amotz Zahavi to explain how evolution may lead to "honest" or reliable signaling between animals which have an obvious motivation to bluff or deceive each other. The handicap principle suggests that reliable signals must be costly to the signaler, costing the signaler something that could not be afforded by an individual with less of a particular trait. For example, in the case of sexual selection, the theory suggests that animals of greater biological fitness signal this status through handicapping behaviour or morphology that effectively lowers this quality. The central idea is that sexually selected traits function like conspicuous consumption, signalling the ability to afford to squander a resource. Receivers know that the signal indicates quality because inferior quality signallers cannot afford to produce such wastefully extravagant signals.

The generality of the phenomenon is the matter of some debate and disagreement, and Zahavi's views on the scope and importance of handicaps in biology has not been accepted by the mainstream. Nevertheless, the idea has been very influential, with most researchers in the field believing that the theory explains some aspects of animal communication.

Liwagu River

The Liwagu River (Malay: Sungai Liwagu) is a river in West Coast Division of Sabah, Malaysia, flowing eastwards off the southern slope of Mount Kinabalu into the Labuk River in Sandakan Division. Most parts of the river are covered by primary and secondary forests.

Meiotic drive

Meiotic drive is a type of intragenomic conflict, whereby one or more loci within a genome will affect a manipulation of the meiotic process in such a way as to favor the transmission of one or more alleles over another, regardless of its phenotypic expression. More simply, meiotic drive is when one copy of a gene is passed on to offspring more than the expected 50% of the time. According to Buckler et al., "Meiotic drive is the subversion of meiosis so that particular genes are preferentially transmitted to the progeny. Meiotic drive generally causes the preferential segregation of small regions of the genome".

Prosphyracephala

Prosphyracephala is a genus of flies in the family Diopsidae. The genus is presumed to be extinct, and is known from specimens in Baltic amber. Among extant taxa, Prosphyracephala most closely resembles the genus Sphyracephala. It was originally described as a species of the latter genus, but was recognized as distinct by Willi Hennig in 1965.

Sex

Organisms of many species are specialized into male and female varieties, each known as a sex. Sexual reproduction involves the combining and mixing of genetic traits: specialized cells known as gametes combine to form offspring that inherit traits from each parent. The gametes produced by an organism define its sex: males produce small gametes (e.g. spermatozoa, or sperm, in animals; pollen in seed plants) while females produce large gametes (ova, or egg cells). Individual organisms which produce both male and female gametes are termed hermaphroditic. Gametes can be identical in form and function (known as isogamy), but, in many cases, an asymmetry has evolved such that two different types of gametes (heterogametes) exist (known as anisogamy).

Physical differences are often associated with the different sexes of an organism; these sexual dimorphisms can reflect the different reproductive pressures the sexes experience. For instance, mate choice and sexual selection can accelerate the evolution of physical differences between the sexes.

Among humans and other mammals, males typically carry an X and a Y chromosome (XY), whereas females typically carry two X chromosomes (XX), which are a part of the XY sex-determination system. Humans may also be intersex. Other animals have different sex-determination systems, such as the ZW system in birds, the X0 system in insects, and various environmental systems, for example in crustaceans. Fungi may also have more complex allelic mating systems, with sexes not accurately described as male, female, or hermaphroditic.

Sexual selection

Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (intrasexual selection). These two forms of selection mean that some individuals have better reproductive success than others within a population, either because they are more attractive or prefer more attractive partners to produce offspring. For instance, in the breeding season, sexual selection in frogs occurs with the males first gathering at the water's edge and making their mating calls: croaking. The females then arrive and choose the males with the deepest croaks and best territories. In general, males benefit from frequent mating and monopolizing access to a group of fertile females. Females can have a limited number of offspring and maximize the return on the energy they invest in reproduction.

The concept was first articulated by Charles Darwin and Alfred Russel Wallace who described it as driving species adaptations and that many organisms had evolved features whose function was deleterious to their individual survival, and then developed by Ronald Fisher in the early 20th century. Sexual selection can, typically, lead males to extreme efforts to demonstrate their fitness to be chosen by females, producing sexual dimorphism in secondary sexual characteristics, such as the ornate plumage of birds such as birds of paradise and peafowl, or the antlers of deer, or the manes of lions, caused by a positive feedback mechanism known as a Fisherian runaway, where the passing-on of the desire for a trait in one sex is as important as having the trait in the other sex in producing the runaway effect. Although the sexy son hypothesis indicates that females would prefer male offspring, Fisher's principle explains why the sex ratio is 1:1 almost without exception. Sexual selection is also found in plants and fungi.The maintenance of sexual reproduction in a highly competitive world is one of the major puzzles in biology given that asexual reproduction can reproduce much more quickly as 50% of offspring are not males, unable to produce offspring themselves. Many non-exclusive hypotheses have been proposed, including the positive impact of an additional form of selection, sexual selection, on the probability of persistence of a species.

Extant Diptera families

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