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The Development Stages of Scuttle Fly
Biomed Sci Letters 2018;24:125-129
Published online June 30, 2018;  https://doi.org/10.15616/BSL.2018.24.2.125
© 2018 The Korean Society For Biomedical Laboratory Sciences.

Hohyun Park

Department of Biomedical Laboratory Science, Mokpo Science University, Mokpo-si, Jeollanam-do 58644, Korea
Correspondence to: Hohyun Park. Department of Biomedical Laboratory Science, Mokpo Science University, Mokpo-si, Jeollanam-do 58644, Korea. Tel: +82-61-270-2745, Fax: +82-61-270-2745, e-mail: phh7082@hanmail.net
Received April 14, 2018; Revised May 14, 2018; Accepted May 17, 2018.
This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.
 Abstract

Megaselia scalaris, also known as the scuttle fly, has a humpbacked morphology and was categorized as an insect undergoing complete metamorphosis. Megaselia scalaris was reported to be found in diverse environments. Recently, the scuttle fly has been a focus of studies by forensic scientists, geneticists, and developmental biologists. However, detailed research into scuttle fly development is still unclear. Therefore, the current study has been performed to elucidate the development of Megaselia scalaris. The scuttle fly undergoes four distinct stages of development: egg, larva, pupa and adult fly. After mating, the scuttle fly eggs hatch within one day and become larvae. The newly generated larvae go through 1, 2, and 3 instar stages within a period of 5~10 days. The instar stage 1 takes 1~2 days, instar stage 2 takes 1~2 days, and instar stage 3 takes 3~5 days. Depending on the environmental temperature, the pupae stage takes approximately 10~15 days to complete. The pupa stage can be divided into five sub-stages: 1~3 days, 4~6 days, 7~9 days, 10~12 days and 13~15 days. After molting, the newly formed adult scuttle fly can fly after one day. The morphological characteristics of the scuttle fly during each stage of development was detailed within this study.

Keywords : Scuttle fly, Development stages, Megaselia scalaris
꽌濡

Scuttle fly쓽 븰紐낆 Megaselia scalaris씠硫 怨ㅼ땐媛(class Insecta), 뙆由щぉ(order Diptera), 踰쇰)뙆由ш낵(family Phoridae)뿉 냽븯怨 씪諛섏쟻쑝濡 “瑗쎌텛뙆由(humpbacked fly)”, “愿뙆由(coffin fly)” 씪怨좊룄 遺瑜몃떎(Triplehorn et al., 2005). 꽦異⑹쓽 紐멸만씠媛 0.5~5.5 mm濡 痢〓㈃紐⑥뼇 瑗쎌텛紐⑥뼇쓣 븯怨 엳쑝硫 寃깋, 媛덉깋, 끂瑜댁뒪由꾪븳 깋쓣 씈怨 엳뒗 寃 듅吏뺤씠떎(Peterson, 1992; Fig. 1).

Fig. 1.

The adult scuttle fly, Megaselia scalaris.


Scuttle fly쓽 꽌떇吏뒗 異뺤텞븯怨 遺뙣븳 쑀湲 臾쇱쭏씤 遺꾨씠굹 궡븘엳뒗 룞臾쇨낵 二쎌 룞臾쇱쓽 뜦 怨좉린肉먮쭔 븘땲씪 怨고뙜씠, 遺뙣븳 빞梨 諛 怨쇱씪, 떇뭹씠굹 諛곗닔濡, 븯닔援ъ쓽 쑀湲 臾쇱쭏 留 벑쑝濡쒖꽌 씠윴 怨녹뿉 븣쓣 궠쑝硫 쑀異⑹씠 릺뼱 꽦옣븳떎. 삉븳 뒿湲곌 엳뒗 벐젅湲고넻 諛붾떏, 뿕由щ쿋씠꽣떎쓽 諛붾떏, 뜑윭슫 臾쇨구젅, 젙솕議 벑뿉꽌룄 쑀異⑹씠 諛쒖깮븯硫, 솕遺꾩쓽 臾 諛쏆묠, 솕蹂, 븷셿룞臾 諛 媛異 궗쑁옣 諛붾떏쓽 異뺤텞븳 넗뼇 諛 諛곗꽕臾쇱뿉 씠瑜닿린源뚯 愿묐쾾쐞븯寃 떎뼇븳 솚寃쎌뿉꽌룄 깮쑁븷 닔 엳떎怨 蹂닿퀬릺뿀떎(Sukontason et al., 2003; Disney, 2008). 듅씠븳 寃껋 scuttle fly媛 援щ몢빟씠굹 럹씤듃 媛숈 怨듭뾽슜 臾쇱쭏뿉꽌룄 踰덉떇븷 닔 엳떎怨 븳떎(Lever, 1944; McCrae, 1967).

怨쇨굅뿉뒗 scuttle fly 꽦異⑹씠 鍮꾩쐞깮쟻씤 옣냼뿉꽌 蹂묒썝꽦 꽭洹 諛 諛붿씠윭뒪 벑쓣 紐몄쓽 몴硫 遺쐞뿉 遺李⑹떆궎嫄곕굹, 蹂묒썝泥대 癒뱀씠 븿猿 꽠痍⑦븯怨 냼솕湲곌쓣 넻빐 遺꾨怨 븿猿 諛곗텧 삉뒗 癒뱀 臾쇱쭏쓣 넗빐궡뒗 벑쓽 諛⑸쾿뿉 쓽빐 궗엺뿉寃 쟾뿼떆耳 떇以묐룆怨 옣뿼쓣 씪쑝궎뒗 빐異⑹쑝濡쒕쭔 씤떇릺뿀怨, 삉븳 씤媛 援щ뜑湲곗쬆쓣 씪쑝궎뒗 怨ㅼ땐쑝濡 蹂닿퀬媛 릺뿀떎(Benecke et al., 2004; Hira et al., 2004; Mazayad and Rifaat, 2005). 씠윭븳 씠쑀濡 슦由щ굹씪뿉룄 쑀븘湲 븣 二쇰 씠猷⑤뒗 諛붿씠윭뒪뿉 쓽븳 湲됱꽦꽕궗吏덊솚怨, 뿰졊씠 넂븘吏먯뿉 뵲씪 諛붿씠윭뒪 꽭洹좎씠 썝씤씠 릺뒗 湲됱꽦꽕궗 吏덊솚 벑쓣 쑀諛쒗븷 닔 엳뒗 썝씤씠 맆 닔 엳떎怨 깮媛곹빐 蹂 닔 엳떎(Lee et al., 2017).

븯吏留 理쒓렐뿉뒗 씤媛꾩씠 留뚮뱺 빀꽦 臾쇱쭏濡 씠猷⑥뼱吏 솚寃쎌삤뿼썝쓣 젣嫄고븯怨 젙솕븯뒗 怨ㅼ땐쑝濡 二쇰ぉ諛쏄퀬 엳쑝硫, 궗썑 媛꾧꺽쓣 異붿젙븯뒗 踰뺤쓽븰뿉꽌 궗슜릺뒗 벑 뿬윭 媛吏 쑀씡븳 痢〓㈃뱾씠 遺媛곷릺怨 엳떎(Greenberg and Wells, 1998; Disney, 2008). 듅엳 쑀쟾븰 諛 諛쒖깮깮臾쇳븰뿉꽌 뿰援ш 吏냽쟻쑝濡 吏꾪뻾릺怨 엳떎(Willhoeft and Traut et al., 1990; Traut, 1994; Willhoeft and Traut, 1995; Sievert et al., 1997; Disney, 2008; Wotton, 2014).

Scuttle fly뒗 怨ㅼ땐쓽 4媛吏 諛쒖깮 떒怨꾩씤 븣(egg), 븷踰뚮젅(larva), 踰덈뜲湲(pupa), 꽦異(adult fly)쓽 怨쇱젙쓣 嫄곗튂뒗 셿쟾 蹂깭 怨ㅼ땐씠떎(Fig. 2). 씪諛섏쟻쑝濡 븣 븯猷 留뚯뿉 遺솕릺뼱 븷踰뚮젅濡 꽦닕릺怨 븷踰뚮젅뒗 깮쑁맆 븣 떎瑜 諛쒖깮 떆湲곕낫떎 솚寃쎌쟻씤 긽깭뿉 뜑 쓽議댄븯寃 릺뒗뜲, 蹂댄넻 22~24°C뿉꽌 媛옣 옒 꽦옣븳떎. 븷踰뚮젅 떆湲곗뿉뒗 꽭 媛쒖쓽 졊(instar)엳怨, 젣 1졊 떆湲곕뒗 1~2씪, 젣 2졊 떆湲곕뒗 1~2씪, 젣 3졊 떆湲곕뒗 3~4씪 젙룄쓽 깮쑁 湲곌컙쓣 嫄곗튂吏留, 寃쎌슦뿉 뵲씪 1~2씪 젙룄 뜑 異붽릺뼱 諛쒖깮씠 씠猷⑥뼱吏꾨떎. 3졊 떆湲곕뒗 씪諛섏쟻쑝濡 븷踰뚮젅뿉꽌 踰덈뜲湲 떆湲곕줈쓽 洹뱀쟻씤 蹂솕媛 씠猷⑥뼱吏湲 븣臾몄뿉 1, 2졊蹂대떎 뜑 삤옒 吏냽릺뒗 듅吏뺤씠 엳떎. 踰덈뜲湲 떆湲곕뒗 13~15씪 젙룄濡쒖꽌 씠썑 꽦異⑹쑝濡 蹂깭媛 씠猷⑥뼱吏꾨떎(Disney, 1994).

Fig. 2.

The life cycle of scuttle fly undergoing complete metamorphosis.


吏湲덇퉴吏 吏꾪뻾맂 뿰援щ뱾쓣 醫낇빀빐꽌 留롮 怨ㅼ땐 以묒뿉꽌 scuttle fly 諛쒖깮뿉 븳 뿰援ш 留롮씠 씠猷⑥뼱吏吏 븡怨 엳湲 븣臾몄뿉 諛쒖깮븰쟻 듅吏 벑쓣 궡렣蹂닿린濡 븯떎.

옱猷 諛 諛⑸쾿

궗쑁

떎뿕뿉 씠슜븳 scuttle fly뒗 Megaselia scalaris씪뒗 븰紐낆쓣 媛吏 fly씠硫, 珥덊뙆由ъ 룞씪븳 癒뱀씠瑜 씠슜븯뿬 궗슜븯떎. 궗쑁쓣 쐞븯뿬 利앸쪟닔 250 mL뿉 dextrose 16 g, yeast flakes 6.5 g, cornmeal 20 g, agar 2.25 g쓣 援먮컲湲곕 씠슜븯뿬 샎빀븯怨 쟾옄젋吏뿉꽌 10遺 젙룄 걪뼱 꽆移섏 븡쓣 젙룄濡 媛뿴븯떎. 洹몃━怨 methylparaben 6.25 mL瑜 泥쒖쿇엳 뼥뼱쑉젮꽌 샎빀븯怨 諛곗 넻뿉 遺 썑 떇엺 떎쓬 꽦異⑹쓣 씠룞떆耳 궗쑁쓣 븯떎. 궗쑁 1二쇱씪 媛꾧꺽쑝濡 떎瑜 諛곗뿉 꽦異⑹쓣 삷寃⑥꽌 吏냽쟻쑝濡 떆뻾븯떎. 젣옉맂 궗쑁 諛곗뒗 4°C 깋옣怨좎뿉 蹂닿븯쑝硫 궗슜 쟾 떎삩뿉 留욎텛湲 쐞빐 誘몃━ 爰쇰궡 넃 썑 궗슜븯떎.

諛쒖깮 떒怨꾨퀎 쇅삎 愿李

諛쒖깮 떒怨꾨퀎濡 삎깭븰쟻 듅吏뺤쓣 궡렣蹂닿린 쐞빐꽌 scuttle fly쓽 븷踰뚮젅(larvae) 떆湲 2졊 2~3씪, 3졊 3~5씪, 踰덈뜲湲(pupae) 떆湲 1~3씪, 4~6씪, 7~9씪, 10~12씪, 13~15씪, 꽦異(adult) 떆湲곕퀎濡 5媛 씠긽쓽 媛쒖껜瑜 꽑깮븯뿬 떎泥댄쁽誘멸꼍(stereomicroscope, LEICA EZ4HD, Germany) 긽뿉꽌 삎깭瑜 愿李고븯떎. 꽦異⑹쓽 寃쎌슦뒗 궇븘꽌 룄留앷湲 븣臾몄뿉 씠궛솕깂냼(CO2) 媛뒪媛 遺꾩궗릺뒗 뙣뱶(pad)뿉 삱젮넃怨 留덉랬떆궓 썑 삎깭瑜 愿李고븯떎.

寃곌낵

Scuttle fly 꽦異⑹씠 援먮몃 븳 떎쓬 븣쓣 궠寃 릺뒗뜲 씠븣 癒뱀씠媛 맆 닔 엳뒗 怨녹뿉 븣쓣 궠뒗떎. 븣(egg) 븯깋쓽 썝삎쑝濡 릺뼱 엳怨 긽떦엳 옉湲 븣臾몄뿉 쑁븞쑝濡 듅吏뺤쓣 愿李고븯湲곌 옒뱾怨, 1씪 젙룄 썑뿉 遺솕릺뼱 1졊쓽 븷踰뚮젅媛 맂떎.

븷踰뚮젅뒗 빟 5~10씪 젙룄쓽 1, 2, 3졊(instar)쓽 諛쒕떖 떒怨꾨 嫄곗튇떎. 븷踰뚮젅 1졊 떆湲곕뒗 뮘履쎌뿉 湲곕Ц씠 븘吏 諛쒖깮븯吏 븡쑝硫댁꽌 議곗쭅씠 誘몄셿꽦 긽깭濡 1~2씪 젙룄 꽦옣븳떎. 癒뱀씠瑜 꽠痍⑦븯뒗 빟媛꾩쓽 吏곸엫留 엳쑝硫댁꽌 嫄곗쓽 諛곗 諛붾떏뿉 臾삵 엳湲 븣臾몄뿉 븣 떆湲곗 留덉갔媛吏濡 쑁븞쑝濡 듅吏뺤쓣 愿李고븯湲곌 돺吏 븡븯떎. 븷踰뚮젅 2졊 떆湲곕뒗 깮쑁 湲곌컙씠 1~2씪 젙룄濡쒖꽌 諛붾떏뿉꽌 癒뱀씠留 꽠痍⑦븯怨 엳뒗 寃껋쓣 愿李고븷 닔媛 엳뿀떎. 洹몃━怨 븷踰뚮젅 2졊 떆湲곕꽣 湲곕Ц(spiracles)쓣 愿李고븷 닔 엳뿀뒗뜲 듅엳 닲쓣 돩뒗뜲 씠슜릺뒗 湲곌(trachea)씠 몢 以꾨줈 슌졆씠 蹂댁씠뒗 寃 듅吏뺤씠뿀떎. 븷踰뚮젅 2졊 떆湲곗쓽 寃쎌슦뒗 3졊蹂대떎 궗씠利덇 몢 諛 젙룄 옉怨 1졊뿉 鍮꾪빐꽌뒗 겕떎뒗 寃껋쓣 솗씤븯떎. 삉븳 몴뵾媛 씛깋씠硫 닾紐낇빐꽌 궡遺湲곌쓣 뼱뒓 젙룄 愿李고븷 닔 엳뿀떎(Fig. 3A, Fig. 3B). 븷踰뚮젅 3졊 떆湲곗뿉뒗 깮쑁 湲곌컙씠 빟 3~5씪 젙룄濡쒖꽌 癒뱀씠瑜 꽠痍⑦븯怨 3졊 留먭린뿉뒗 솢諛쒗븯寃 쐞瑜 뼢빐 吏곸씠뒗 寃껋씠 듅吏뺤씠떎. 븷踰뚮젅 3졊 떆湲곕뒗 몴뵾媛 2졊 떆湲곗뿉 鍮꾪빐 몢爰쇱썙吏硫댁꽌 빟媛 遺덊닾紐낇븳 긽깭濡 蹂솕媛 씠猷⑥뼱吏怨 엳뿀쑝硫, 湲곕Ц룄 슌졆씠 蹂댁씠怨 몢 以꾩쓽 湲곌룄 옒 愿李고븷 닔 엳뿀떎. 洹몃━怨 븷踰뚮젅쓽 삎깭룄 留롮씠 鍮꾨빐졇 엳뒗 寃껋쓣 蹂 닔媛 엳뿀怨(Fig. 3C, Fig. 3D), 쟾泥댁쟻쑝濡 蹂댁븯쓣 븣 븷踰뚮젅 2졊怨 3졊 떆湲곗뿉뒗 꽦異⑹씤 scuttle fly 鍮꾧탳뻽쓣 븣 겕湲곌 썾뵮 겙 寃껋쓣 愿李고븷 닔 엳뿀떎.

Fig. 3.

Gross morphological changes during larval development. stereomicroscopy, x25. A, B; 2-nd instar C, D; 3-rd instar.


踰덈뜲湲(pupa) 떆湲곕뒗 蹂댄넻 諛쒖쑁 삩룄뿉 뵲씪 10~15씪 젙룄쓽 깮쑁 湲곌컙쓣 嫄곗튂뒗뜲 1~3씪, 4~6씪, 7~9씪, 10~12씪, 13~15씪濡 깮쑁 湲곌컙쓣 援щ텇븯뿬 삎깭쟻 蹂솕瑜 愿李고빐 蹂댁븯떎. 踰덈뜲湲 1~3씪 떆湲곕뒗 湲곕Ц씠 諛뽰쑝濡 룎異쒕릺湲 떆옉븳 긽깭濡 愿李곗씠 릺뿀怨, 븷踰뚮젅뿉꽌 踰덈뜲湲곌 諛쒖깮븯뒗 1씪吏몄뿉 遺뱶윭슫 씛깋쓽 걧떚겢쓣 媛吏 몴뵾濡 蹂솕媛 씠猷⑥뼱吏硫댁꽌 젏李 떒떒빐吏怨 3씪씠 릺硫 솴媛덉깋쑝濡 蹂븯寃 릺뒗 寃껋쓣 蹂 닔媛 엳뿀떎. 몴뵾뿉뒗 誘몄꽭븳 룎湲곕뱾씠 굹굹湲 떆옉븯怨 泥댁젅 蹂꾨줈 빟媛꾩쓽 留뚭끝遺媛 以꾩씠 洹몄뼱졇 엳뒗 삎깭濡 愿李곗씠 릺뿀떎. 삉븳 3씪 썑遺꽣 誘몄꽭븳 肉붽린뫁씠 留뚮뱾뼱吏湲 떆옉븯뒗 寃껋쓣 솗씤븷 닔 엳뿀떎(Fig. 4A, Fig. 4B). 踰덈뜲湲 4~6씪 떆湲곕뒗 걧떚겢쓽 깋源붿씠 젏젏 吏꾪븳 媛덉깋쑝濡 蹂븯硫댁꽌 泥댁젅쓽 留뚭끝遺 以꾩씠 뜑슧 뜑 슌졆븯寃 吏꾪빐吏怨 룎湲곕룄 슌졆븯寃 룎異쒖씠 릺뼱엳뒗 寃껋쓣 솗씤븯떎. 삉븳 肉붽린뫁룄 뜑슧 뜑 諛뽰쑝濡 湲멸쾶 룎異쒕릺뼱 엳뒗 寃껊룄 愿李고븷 닔 엳뿀떎(Fig. 4C). 踰덈뜲湲 7~9씪 떆湲곕뒗 걧떚겢쓽 깋源붿씠 媛덉깋쑝濡 吏꾪빐吏硫댁꽌 泥댁젅쓽 留뚭끝遺쓽 以꾩씠 紐낇솗엳 뱶윭궗怨, 룎湲곕룄 슌졆씠 룎異쒕릺뿀떎. 븳렪 씠 떆湲곗뿉 꽦異⑹쓽 떎由ш 留뚮뱾뼱吏湲 떆옉븯뒗 寃껋쓣 蹂 닔 엳뿀怨, 뜑遺덉뼱 궇媛쒖 빟븯寃 븞援щ룄 留뚮뱾뼱吏뒗 寃껋쓣 愿李고븷 닔 엳뿀떎(Fig. 4D). 踰덈뜲湲 10~12씪 떆湲곕뒗 몴뵾쓽 걧떚겢 깋源붿씠 吏숈 媛덉깋쑝濡 뜑슧뜑 吏꾪빐吏怨 泥댁젅쓽 留뚭끝遺쓽 以꾨룄 뜑슧뜑 吏꾪빐졇 엳쑝硫댁꽌 踰덈뜲湲 쐵遺遺꾩쓽 떗옄紐⑥뼇쓽 留뚭끝遺媛 젏젏 깉씫씠 릺湲 떆옉븯떎. 몴뵾瑜 踰쀪릿 踰덈뜲湲 삎깭瑜 궡렣蹂댁븯쓣 븣 肉붽린뫁 2媛쒓 븘吏곴퉴吏 슌졆씠 룎異쒕릺뼱 엳怨 궇媛쒖 떎由ш 嫄곗쓽 셿꽦릺뼱 엳뒗 寃껋쓣 蹂 닔 엳뿀떎. 洹몃━怨 뼇履쎌쓽 븞援ъ쓽 삎깭媛 슌졆븯寃 媛덉깋쑝濡 愿李곕릺뿀떎(Fig. 4E). 留덉留 踰덈뜲湲 13~15씪 떆湲곗뿉꽌뒗 몴뵾쓽 걧떚겢 깋源붿씠 吏꾪븳 쓳媛덉깋쑝濡 蹂븯硫댁꽌 踰덈뜲湲 쐵遺遺 泥댁젅쓽 留뚭끝遺 씪遺媛 떗옄紐⑥뼇쑝濡 몴뵾媛 媛덈씪졇 뼥뼱졇 엳뒗 寃껋씠 愿李곕릺뿀怨, 踰덈뜲湲 몴뵾瑜 踰쀪꺼궡뼱 삎깭瑜 궡렣蹂댁븯쓣 븣, 몴뵾 諛붽묑履쎌쓽 肉붽린뫁씠 븞履쎄낵 遺꾨━릺뼱 엳뿀怨 癒몃━ 媛뒾, 諛곌 遺꾨━릺뼱 愿李곕릺怨 궇媛, 떎由, 븞援 벑씠 셿꽦맂 삎깭濡 뻼 留 븞뿉꽌 留뚮뱾뼱졇 꽦異⑹쓽 삎깭瑜 셿쟾엳 媛뽰텛怨 엳뒗 寃껋쓣 솗씤븯떎(Fig. 4F).

Fig. 4.

Gross morphological changes during pupal development. stereomicroscopy, x30. A; 1 day, B; 3 days C; 4~6 days, D; 7~9 days, E; 10~12 days, F; 13~15 days.


踰덈뜲湲곗뿉꽌 깉뵾빐꽌 꽦異⑹씠 릺硫 1씪 썑뿉 궇븘떎땲湲 떆옉븯怨, 꽦異⑸뱾씠 癒뱀씠瑜 癒밴굅굹 븣쓣 궠뒗 떆湲 쇅뿉뒗 궗쑁 諛곗 쐞履쎌쓽 슌猿묒쓣 뼢븯뿬 臾대━ 吏뼱 엳뒗 紐⑥뒿씠 듅吏뺤쟻 씠뿀떎. 꽦異⑹쓽 紐몄 겕寃 꽭 遺遺꾩쑝濡 굹늻뼱吏뒗뜲 븳 뙇쓽 뜑벉씠, 늿, 엯쑝濡 援ъ꽦맂 癒몃━ 遺遺꾧낵 궇媛쒖 떎由ш 엳뒗 媛뒾 遺遺, 洹몃━怨 諛 遺遺꾩쑝濡 援ъ꽦릺뼱 엳뿀떎. 꽦異⑹쓽 닔而룰낵 븫而룹쓣 鍮꾧탳빐蹂대㈃ 닔而룹 븫而룹뿉 鍮꾪빐 겕湲곌 옉怨 蹂듬媛 寃怨 걹씠 뫁洹 듅吏뺤쓣 媛吏怨 엳뿀떎. 洹몃━怨 븫而룹 겕湲곌 닔而룹뿉 鍮꾪빐 2~3諛 겙 렪씠怨 蹂듬쓽 걹씠 寃吏 븡怨 빟媛 毓곗”븯怨 湲 삎깭쟻 듅吏뺤쓣 愿李고븷 닔媛 엳뿀떎(Fig. 5).

Fig. 5.

Gross morphological comparsion of the male and female scuttle fly stereomicroscopy, x30.


怨좎같

Scuttle fly뒗 씠由꾩씠 븫떆븯벏씠 留ㅼ슦 룆듅븳 뻾룞 듅꽦쓣 媛吏꾨떎. 슫룞 以 媛묒옄湲 젙吏븯떎媛 삉 鍮좊Ⅴ寃 씠룞븯怨 吏㏃ 씠룞 뮘뿉 삉 떎떆 젙吏븳떎. 肉먮쭔 븘땲씪 scuttle fly뒗 遺뙣븳 쑀湲곕Ъ씠굹 떖吏뼱 쑀湲곗슜留ㅺ 샎빀맂 怨듭뾽슜 臾쇱쭏뿉꽌룄 踰덉떇븯뒗 벑(Lever, 1944; McCrae, 1967), 씠뙆由ш 珥덊뙆由 벑쓽 쑀궗븳 삎깭쟻 듅吏뺤쓣 媛吏뒗 샎뿰醫낅뱾怨 李⑤퀎릺뒗 룆듅븳 듅꽦쓣 媛吏怨 엳떎. 씠윴 뻾룞 洹쇱쑁怨 떊寃쎄퀎뿉 쓽빐 議곗젅릺誘濡 scuttle fly쓽 떊寃쎄퀎 洹쇱쑁怨꾨뒗 떎瑜 醫낅쪟쓽 뙆由щ뱾怨 李⑤퀎솕맂 룆듅븳 듅吏뺤쓣 媛吏怨 엳뒗 寃껋씠씪怨 삁긽븯硫댁꽌 씠 遺遺꾩뿉 빐꽌 뿰援ш 吏냽쟻쑝濡 씠猷⑥뼱졇빞 븷 寃껋씠떎.

ACKNOWLEDGEMENTS

蹂 뿰援щ뒗 怨좎씤씠 릺떊 紐⑺룷븰援 깮臾쇳븰怨 꽭룷븰 떎뿕떎 源옱留 援먯닔떂쓽 吏룄뿉 쓽빐 닔뻾릺뿀뒿땲떎.

CONFLICT OF INTEREST

None.

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