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<h1>LIFE CYCLE</h1>
<p>The biology of each species of sand fly is unique and complex, including reproduction, feeding, dispersal and other activities, and being of importance for the epidemiology of transmitted diseases and vector control (WHO, 1990).</p>
<p>Sand flies undergo complete metamorphosis, passing through four distinct stages during their life cycle: egg, larva, pupa, adult/imago.</p>
<p>In arid and temperate regions, larvae may remain in a state of diapause for many months. In temperate countries, sand flies overwinter as eggs or larvae in a so-called diapause. For this reason, adults can appear early in summer, and <a href="https://campaigns-dev.azurewebsites.net/en-us/diseases/sand-fly-borne-diseases/leishmaniosis/">leishmaniosis</a> –&nbsp;as one the sand fly-borne diseases&nbsp;– can exist in places with a cold winter (Lewis, 1971). Similarly, diapause might also be the means by which many tropical sand flies survive periods of drought or heavy rain (Ward and Killick-Kendrick, 1974).</p>
<p>Palearctic sand flies overwinter as fourth instar larvae (Killick-Kendrick and Killick-Kendrick, 1987; Ready and Corset, 1980). From laboratory studies, a complete gonotrophic cycle normally does not last less than 6 weeks, but can differ between different species (Killick-Kendrick and Killick-Kendrick, 1999).</p>

<h2>EGG</h2>
<p>Eggs are oblong-oval, minute (average 0.3-0.4 mm, according to Shevchenko, 1929) and difficult to find in nature. Newly-laid eggs are soft, light yellow, with a shiny surface. After several hours, they darken, become hard, dark brown and show characteristic thin ridges (Perfiliev, 1968).</p>
<p>They are laid in cracks and crevices in moist ground, among leaf litter, between buttress roots of trees, at the bases of termite mounds, on stable floors and in poultry sheds, where adult flies rest and the microclimate is high.</p>
<p>The site and type of habitat is species-dependent. Between 10/15 and 100 eggs are laid in a single batch (40-70 usually in one oviposition cycle [WHO, 1990]). Under optimum conditions, they hatch within 6-17 days. Longer periods are necessary in cooler conditions.</p>

<h2>LARVA</h2>
<p>Sand fly larvae possess one respectively two pairs of caudal bristles and characteristic 'matchstick' hairs on each segment.</p>
<p>Four instars of larvae follow over a period of 21-60 days (Peters, 1992), respectively 16-90 days (Service, 2001). Larvae are mainly scavengers. In laboratories, they are fed on semi-rotting vegetable matter and decomposing arthropod bodies, chow or even faeces of vertebrates (e.g. aged rabbit faeces [Singh, 1985, based on Endris et al. 1982]). The developing larva is cutting an exit hole with a sort of 'egg tooth' into the egg shell.</p>
<p>The final-instar larva sheds its skin, and the pupa is formed. Larval breeding sites are seldom found. Bettini and Melis (1988) suggested pre-imaginal stages associated with a comparatively stable, cool, humid environment, protected from rain and sunshine and in the special analyzed site, rich in clay and organic nitrogen.</p>

Larva

<h2>PUPA</h2>
<p>The pupae of sand flies are 3 mm long and club-shaped. The narrow part is curved and the head and thorax are curved backwards. The posterior segments are covered with the exuviae of the fourth-stage larva. With it, the pupa is attached to the substrate in a usually vertical position (Perfiliev, 1968). After approximately 5-10 days the adult emerges.&nbsp;</p>

Pupa

<h2>ADULT</h2>
<p>Adults can be identified from other insects by their behaviour, appearance, size etc. (see under <a href="https://campaigns-dev.azurewebsites.net/en-us/vectors/sand-flies/morphology/">Morphology</a>). Both sexes feed on plant sugars (Lewis and Domoney, 1966) and possibly also honeydew of aphids and coccids (Killick-Kendrick, 1979). The type of sugar and the frequency of uptake may be a factor in the insect's ability to transmit <em>Leishmania</em>. Plant sugars may be essential for parasite survival, and promastigotes require carbohydrates for their development (Killick-Kendrick, 1978). Only females feed on blood, which is suspected as a source of protein for egg development. When seeking food, sand flies normally have a characteristic short hopping flight.</p>
<p>Females require the blood meal to mature about 80-100 eggs, and the time from engorgement to oviposition is not less than 6 days (demonstrated in Mediterranean species) (Killick-Kendrick and Killick-Kendrick, 1999). Males are attracted to females, when those are feeding, and will mate with them even while the females are taking a blood meal.</p>
<p>The life expectancy has hardly been determined for wild female sand flies, but Killick-Kendrick et al. (1984) recaptured marked <em>Phlebotomus ariasi</em> 28 days after release.</p>
<p>In nature, more than one gonotrophic cycle is considered normal (Dolmatova, 1942), whereas in the laboratory, most gravid females lay eggs and die either as they lay or shortly afterwards. For some subgenera of <em>Phlebotomus, </em>the females are gonotrophically concordant and take one blood meal each gonotrophic cycle (Killick-Kendrick and Killick-Kendrick, 1999).</p>
<p>Most <em>Leishmania </em>species complete their life-cycle within a single ovarian cycle of the sand fly, and parasites may therefore be transmitted at the first blood meal after an infective feed. But in some species (<em>Leishmania donovani donovani</em> and some strains of <em>L.</em>&nbsp;<em>donovani infantum</em>), the development is not complete until after the second blood meal.</p>

Adult

<h2>REFERENCES</h2>
<h4>Introduction</h4>
<p>Killick-Kendrick R, Killick-Kendrick M: The laboratory colonization of <em>Phlebotomus ariasi </em>(Diptera: Psychodidae). Ann Parasitol Hum Comp. 1987, 62, 354-6</p>
<p>Killick-Kendrick R, Killick-Kendrick M: Biology of sand fly vectors of Mediterranean canine leishmaniosis. In: Killick-Kendrick R (ed.): Canine leishmaniasis: an update. Proc. Int. Can. Leishm. Forum, Barcelona, Spain, 1999, Intervet Int., Boxmeer, The Netherlands, 1999, 26-31</p>
<p><a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2427821/pdf/bullwho00201-0064.pdf" data-new-window="true" target="_blank" rel="noopener noreferrer">Lewis DJ: Phlebotomid sandflies. Bull. WHO 1971, 44, 535-51</a></p>
<p>Ready PD, Croset H: Diapause and laboratory breeding of <em>Phlebotomus perniciosus</em> Newstead and <em>Phlebotomus ariasi</em> Tonnoir (Diptera: Psychodidae) from southern France. Bull Ent Res. 1980, 70, 511-23</p>
<p><a href="https://apps.who.int/iris/bitstream/handle/10665/39337/WHO_TRS_793.pdf?sequence=1&amp;isAllowed=y" data-new-window="true" target="_blank" rel="noopener noreferrer">WHO: Control of the leishmaniases. Report of a WHO Expert Committee, Tech. Rep. Ser. No. 793, WHO, Geneva, 1990</a></p>
<h4>Egg</h4>
<p>Perfiliev PP: Fauna of U.S.S.R. Diptera. Phlebotomidae (sandflies). Acad. Sci. U.S.S.R. Zool. Inst. New Ser. No. 93, Vol. 3 No. 2, Israel Program for Scientific Translations, Jerusalem, 1968&nbsp;</p>
<p>Shevchenko FI: Vneshnyaya morfologiya lichinok moskitov.<em> P.&nbsp;papatasi, P.&nbsp;chinensis, P.&nbsp;sergenti </em>(The external morphology of sandfly larvae (<em>P.&nbsp;papatasi, P.&nbsp;chinensis, P.&nbsp;sergenti</em>)). Meditsinskaya Mysl' Uzbekistana I Turkmenistana, 1929-1930, 67-83</p>
<p><a href="https://apps.who.int/iris/bitstream/handle/10665/39337/WHO_TRS_793.pdf?sequence=1&amp;isAllowed=y" data-new-window="true" target="_blank" rel="noopener noreferrer">WHO: Control of the leishmaniases. Report of a WHO Expert Committee, Tech. Rep. Ser. No. 793, WHO, Geneva, 1990</a>&nbsp;</p>
<h4>Larva</h4>
<p>Bettini S, Melis P: Leishmaniasis in Sardinia. III. Soil analysis of a breeding site of three species of sandflies. Med Vet Entomol. 1988, 2, 67-71</p>
<p>Endris RG, Perkins PV, Young DG, et al.: Techniques for laboratory rearing of sand flies (Diptera: Psychodidae). Mosq News. 1982, 42, 400-7</p>
<p>Peters W: Leishmaniases. In: Peters W (ed.): A colour atlas of arthropods in clinical medicine. 1992, Wolfe Publish. Ltd., London, pp. 115-34</p>
<p>Singh P: Multiple-species rearing diets. Diptera: Psychodidae. In: Singh P, Moore RF (eds.): Handbook of Insect Rearing. Vol. 1, 1985,&nbsp;Elsevier, Amsterdam, pp. 33-4</p>
<p>Service MW: Phlebotomine sand-flies (Phlebotominae). In: Service MW (ed.): The encyclopedia of arthropod-transmitted infections. 2001,&nbsp;CABI Publish., Oxon, New York, pp. 395-7</p>
<h4>Pupa</h4>
<p>Perfiliev PP: Fauna of U.S.S.R. Diptera. Phlebotomidae (sandflies). Acad. Sci. U.S.S.R. Zool. Inst. New Ser. No. 93, Vol. 3 No. 2, Israel Program for Scientific Translations, Jerusalem, 1968&nbsp;</p>
<h4>Adult</h4>
<p>Dolmatova AV: Life cycle of <em>Phlebotomus papatasi</em> (Scopoli). (in Russian) Med Parazit. (Mosk.) 1942, 11, 32-70</p>
<p><a href="https://www.e-periodica.ch/cntmng?pid=act-001:1978:35::460" data-new-window="true" target="_blank" rel="noopener noreferrer">Killick-Kendrick R: Recent advances and outstanding problems in the biology of phlebotomine sandflies. Acta Trop. 1978, 35, 297-313</a></p>
<p>Killick-Kendrick R: Biology of <em>Leishmania</em> in phlebotomine sandflies. In: Lumsden WHR, Evans DA (eds.): Biology of the Kinetoplastida. Vol. 2, 1979,&nbsp;Academic Press, London, New York, pp. 395-460</p>
<p>Killick-Kendrick R, Killick-Kendrick M: Biology of sand fly vectors of Mediterranean canine leishmaniosis. In: Killick-Kendrick R (ed.): Canine leishmaniasis: an update. Proc. Int. Can. Leishm. Forum, Barcelona, Spain, 1999, Intervet Int., Boxmeer, The Netherlands, 1999, 26-31</p>
<p>Killick-Kendrick R, Rioux JA, Bailly M, et al.: Ecology of leishmaniasis in the south of France. 20. Dispersal of <em>Phlebotomus ariasi</em> Tonnoir, 1921 as a factor in the spread of visceral leishmaniasis in the Cévennes. Ann Parasitol Hum Comp. 1984, 59, 555-72</p>
<p>Lewis DJ, Domoney CR: Sugar meals in Phlebotominae and Simuliidae. Proc R Ent Soc Lond. A 1966, 41, 175-79</p>

References

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