Teleopsis dalmanni

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Teleopsis dalmanni
Teleopsis dalmanni.jpg
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Diopsidae
Genus: Teleopsis
Species:
T. dalmanni
Binomial name
Teleopsis dalmanni
(Wiedemann, 1830)
Synonyms [1]
  • Cyrtodiopsis dalmanni(Wiedemann, 1830)
Eyestalk close up Teleopsis dalmanni Teleopsis dalmanni 2.jpg
Eyestalk close up Teleopsis dalmanni

Teleopsis dalmanni, synonym Cyrtodiopsis dalmanni, [1] also known as the Malaysian stalk-eyed fly, is a species of fly in the family Diopsidae. T. dalmanni flies possess lateral elongations on their head capsules called eyestalks. These eyestalks play an important role in mate selection and as a result physical characteristic of the fly has been the subject of several studies on sexual selection, natural selection, and mating behavior.

Contents

Distribution

Teleopsis dalmanni is found in Malaysia as well as other parts of Southeast Asia. [2] Flies used in studies are usually collected in Malaysia.

Habitat

Teleopsis dalmanni are predominantly found at the edge of forest streams, where their mating rituals occur. [2] They have been observed resting on dried leaves or undergrowth at the water’s edge, and occasionally even on the sand. [3]

Life history

As with other Dipterans, T. dalmanni flies undergo a larval phase before transitioning into adults. There is little information available on the systematics and behavior of the larval stage in T. dalmanni. As holometabolous insects, the flies have dividing cells restricted to the gut and gonads. [4] Due to this, the adult size is fixed by the nutrition received in the larval phase. Following the larval stage, further nutrition is required for the development of internal reproductive structures to reach sexual maturity.

Breeding experiments typically assign three to six weeks for these flies to reach sexual maturity, but a value for wild fly populations has not been determined with confidence. Sexual maturity can be indicated by accessory gland growth, maturing of sperm bundles, and sperm motility. Accessory gland growth was found to have the largest effect on sexual maturity, but it is not the sole effect. [4]

Food resources

Adult flies exhibit solitary foraging behavior, feeding on rotting vegetative matter in the daytime. [5]

Mating

Female T. dalmanni roost on root threads overhanging streams at dusk. Males compete to gain control of these root hairs. [2] Upon gaining control of a root hair they form harems, and females decide which male’s harem to join. Studies have shown that females prefer to roost with males with larger eye-spans. [2] Mating occurs at dawn, and usually multiple mating events take place. [5] Research shows that increased mating frequency correlates with a higher proportion of fertile eggs. This is supported by the idea that multiple matings do not reduce the receptivity of a female fly to copulation, regardless of the characteristics of the mating male. Studies showed that the number of copulation events rather than the number of mates led to an increase in hatching success. Additionally, a large number of copulations are unsuccessful, with a study finding that females that copulated once had a hatching success rate of only 10%. [2] Copulation specifics are not entirely understood, but some copulation events are rapid and last less than 60 seconds. [5]

Spermatophores

Male flies produce gourd-shaped spermatophores that occupy only a part of the vaginal capacity, as compared to the spermatophores of certain other diopsid flies that occupy the entire vaginal cavity. [6] It has been suggested that the smaller size of the spermatophore is an adaptation to reduce investment in sperm due to a high mating frequency in a short amount of time. [7] Accessory glands are responsible for forming the spermatophore that transports sperm during mating. [4]

Morphology

Eyestalks

The eyestalks of T. dalmanni, as well as other Diopsidae flies, are frequently used to study the fields of sexual selection and mating preference. Eyestalks have been known to range from 4 mm in length up to 17 mm. [8] Eyestalks can even be longer than body length. [9]

Hyperencephaly of the type that resembles the eyestalks in T. dalmanni is seen is other Dipteran organisms, but flies of the family Diopsidae are unique in that both males and females display some degree of eyestalk elongation. In T. dalmanni flies, males have larger eye-spans relative to body length compared to females. This feature of the fly is subject to studies that investigate the genetic background of sexual selection, female mating preferences and reproductive consequences. [5]

Mate choice

Eyestalks play an important role in mating behavior, as studies have shown that females prefer males with larger eye spans relative to body length. It was found that female preference for large eye-spans is evident when the difference between eye-spans of available males is large, not moderate. In cases of low and moderate difference, a female with a large eye-span is more likely to pick the male with larger eye-span. [8] There are multiple theories posited to explain this, including the idea that female preference for male eye-span may have a ceiling, or that female choice is linked genetically to female eye-span.

Sexual selection

Sexual selection posits that structures that are perceived as reliable indicators of individual quality are selected for. Teleopsis dalmanni eyestalks are an example of sexual dimorphism that has developed for mating purposes; larger eye-span is selected for as evidence indicates that it is preferred by females. [10] A study also found that female individuals with larger eye-spans had significantly more mature oocytes than those with shorter eye-spans. [10]

Furthermore, eye-span independent of body size or mass has been identified as the key determinant in the outcomes of contests between males. [11] This suggests that longer eyestalks developed in males as a result of mating success pressure, and hence are an example of sexual selection. However, it has been shown that females that mated with males with large eye-spans did not receive short-term fecundity benefits and fertility benefits, and in fact recorded a lower fertility rate than females mating with males with short eye-spans. [2]

Handicap hypothesis

Male eye-span is strongly dependent on conditional factors, comparatively more than other non-sexual traits, [11] and eye-span becomes variable under conditions of increased stress. Similar patterns are not seen in comparable nonsexual traits, and are more closely correlated with body size. The hypothesis drawn from these results is that eye-span provides additional information about the condition beyond that provided by body size. Hence eye-span could be an alternative indication to females of the male's greater ability to survive. This relates to the handicap theory of evolution of ornamental male traits. [11]

However, studies cite a lack of evidence to support the idea that the extended eye-stalks are a handicap to male Diopsid flies. While the eye-stalk increases the moment of inertia of male flies, they were not found to suffer a flight performance decrement. Specifically T. dalmanni male flies were found to perform better than females in free-flying turning behavior. [10] If longer eyespans do not pose a handicap to male flies, an alternative explanation may be more successful in explaining the development of this trait.

Eyes

Teleopsis dalmanni has been used as a study subject in experimental subject in studies on data processing in insect optic centers, as it is easy to record impulse activity in the eyestalk.

Related Research Articles

Handicap principle Hypothesis in evolutionary biology

The handicap principle is a hypothesis proposed by Amotz Zahavi to explain how evolution may lead to "honest" or reliable signalling between animals which have an obvious motivation to bluff or deceive each other. It suggests that costly signals must be reliable signals, costing the signaller something that could not be afforded by an individual with less of a particular trait. For example, in 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 then know that the signal indicates quality, because inferior quality signallers are unable to produce such wastefully extravagant signals.

The Coolidge effect is a biological phenomenon seen in animals, whereby males exhibit renewed sexual interest whenever a new female is introduced to have sex with, even after cessation of sex with prior but still available sexual partners. To a lesser extent, the effect is also seen among females with regard to their mates.

Speckled wood (butterfly) Species of butterfly in the family Nymphalidae

The speckled wood is a butterfly found in and on the borders of woodland areas throughout much of the Palearctic realm. The species is subdivided into multiple subspecies, including Pararge aegeria aegeria, Pararge aegeria tircis, Pararge aegeria oblita, and Pararge aegeria insula. The color of this butterfly varies between subspecies. The existence of these subspecies is due to variation in morphology down a gradient corresponding to a geographic cline. The background of the wings ranges from brown to orange, and the spots are either pale yellow, white, cream, or a tawny orange. The speckled wood feeds on a variety of grass species. The males of this species exhibit two types of mate locating behaviors: territorial defense and patrolling. The proportion of males exhibiting these two strategies changes based on ecological conditions. The monandrous female must choose which type of male can help her reproduce successfully. Her decision is heavily influenced by environmental conditions.

Sperm competition

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 males' chance of reproductive success. Sperm competition results in a sexual conflict of interest 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.

Spermatophore Packet containing sperm in invertebrate reproduction

A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores may additionally contain nourishment for the female, in which case it is called a nuptial gift, as in the instance of bush crickets. In the case of the toxic moth Utetheisa ornatrix, the spermatophore includes sperm, nutrients, and pyrrolizidine alkaloids which prevent predation because it is poisonous to most organisms. However, in some species such as the Edith's checkerspot butterfly, the "gift" provides little nutrient value. The weight of the spermatophore transferred at mating has little effect on female reproductive output.

Anisogamy Sexual reproduction involving a large, "female" gamete and a small, "male" gamete

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Stalk-eyed fly Family of dipteran insects with antennae located on eyestalks

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.

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Courtship display

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References

  1. 1 2 "Teleopsis dalmanni". Taxonomy Browser. National Center for Biotechnology Information (NCBI}. Retrieved 2020-10-15.
  2. 1 2 3 4 5 6 Baker, Richard H.; Ashwell, Robert I. S.; Richards, Thomas A.; Fowler, Kevin; Chapman, Tracey; Pomiankowski, Andrew (2001-11-01). "Effects of multiple mating and male eye span on female reproductive output in the stalk-eyed fly, Cyrtodiopsis dalmanni". Behavioral Ecology. 12 (6): 732–739. doi: 10.1093/beheco/12.6.732 . ISSN   1045-2249.
  3. Burkhardt, Dietrich (1972-06-01). "Electrophysiological studies on the compound eye of a stalked-eye fly,Cyrtodiopsis dalmanni (Diopsidae, Diptera)". Journal of Comparative Physiology. 81 (2): 203–214. doi:10.1007/BF00696633. ISSN   1432-1351.
  4. 1 2 3 Baker, Richard H.; Denniff, Matthew; Futerman, Peter; Fowler, Kevin; Pomiankowski, Andrew; Chapman, Tracey (2003-09-01). "Accessory gland size influences time to sexual maturity and mating frequency in the stalk-eyed fly, Cyrtodiopsis dalmanni". Behavioral Ecology. 14 (5): 607–611. doi: 10.1093/beheco/arg053 . ISSN   1045-2249.
  5. 1 2 3 4 Wilkinson, Gerald S.; Reillo, Paul R. (1994-01-22). "Female choice response to artificial selection on an exaggerated male trait in a stalk-eyed fly". Proceedings of the Royal Society of London. Series B: Biological Sciences. 255 (1342): 1–6. Bibcode:1994RSPSB.255....1W. doi:10.1098/rspb.1994.0001.
  6. Kotrba, Marion (1996-07-01). "Sperm transfer by spermatophore in Diptera: new results from the Diopsidae". Zoological Journal of the Linnean Society. 117 (3): 305–323. doi: 10.1111/j.1096-3642.1996.tb02192.x . ISSN   0024-4082.
  7. Kotrba, Marion (1996-07-01). "Sperm transfer by spermatophore in Diptera: new results from the Diopsidae". Zoological Journal of the Linnean Society. 117 (3): 305–323. doi: 10.1111/j.1096-3642.1996.tb02192.x . ISSN   0024-4082.
  8. 1 2 Hingle, Andrew; Fowler, Kevin; Pomiankowski, Andrew (2001-03-01). "Size-dependent mate preference in the stalk-eyed fly Cyrtodiopsis dalmanni". Animal Behaviour. 61 (3): 589–595. doi:10.1006/anbe.2000.1613. ISSN   0003-3472. S2CID   53154862.
  9. Ribak, Gal; Egge, Alison R; Swallow, John G (2009-05-07). "Saccadic head rotations during walking in the stalk-eyed fly (Cyrtodiopsis dalmanni)". Proceedings of the Royal Society B: Biological Sciences. 276 (1662): 1643–1649. doi:10.1098/rspb.2008.1721. PMC   2660984 . PMID   19203925.
  10. 1 2 3 Husak, Jerry F.; Ribak, Gal; Wilkinson, Gerald S.; Swallow, John G. (2011-06-01). "Compensation for exaggerated eye stalks in stalk‐eyed flies (Diopsidae)". Functional Ecology. 25 (3): 608–616. doi: 10.1111/j.1365-2435.2010.01827.x . ISSN   1365-2435.
  11. 1 2 3 Cotton, Samuel; Fowler, Kevin; Pomiankowski, Andrew (2004). "Condition Dependence of Sexual Ornament Size and Variation in the Stalk-Eyed Fly Cyrtodiopsis Dalmanni (diptera: Diopsidae)". Evolution. 58 (5): 1038–1046. doi:10.1111/j.0014-3820.2004.tb00437.x. ISSN   1558-5646. PMID   15212384.