嵟怴偺尋媶徯夘

[ 儕儕乕僗丗 2019.02 ]
撧椙彈巕戝妛 壔妛惗暔娐嫬妛壢 娐嫬壢妛僐乕僗 嫵庼丂崅恵晇屽
http://gi.ics.nara-wu.ac.jp/~takasu

悢棟揑庤朄傪梡偄偰條乆側惗柦尰徾傪尋媶偡傞悢棟惗暔妛傪愱栧偲偟偰偄傑偡丅摿偵丄惗偒暔偺悢偺帪娫曄壔乮愱栧梡岅偱屄懱孮摦懺偲屇傇乯偵拲栚偟偨悢棟揑尋媶傪偟偰偄傑偡丅悢棟揑庤朄偲偼丄悢幃傗傾儖僑儕僘儉偲尵偭偨拪徾揑側傕偺偺尒曽傪捠偟偰尋媶懳徾傪儌僨儖壔偟偰夝愅偡傞庤朄偱偡丅惗柦尰徾偲尵偊偽堦斒揑偵偼惗暔妛偑埖偆尋媶懳徾偵側傝傑偡偑丄悢棟揑庤朄偲惗暔妛偑偳偺傛偆偵寢傃偮偔偺偐丄捈偖偵偼憐憸偱偒側偄曽偑彮側偔偼側偄偺偱偼柍偄偐偲巚偄傑偡丅偙偙偱偼丄悢棟揑庤朄偑擛壗偵偟偰惗暔妛偲寢傃偮偔偺偐傪愢柧偡傞偲偲傕偵丄巹偑庢傝慻傫偱偒偨尋媶帠椺傪偄偔偮偐徯夘偟丄悢棟揑庤朄偲惗暔妛偺怺偄娭傢傝偵偮偄偰抦偭偰捀偙偆偲巚偄傑偡丅娐嫬壢妛偼恖娫傪庢傝姫偔惗暔妛揑娐嫬乮惗懺宯乯傗暔棟妛揑娐嫬偵娭傢傞尋媶暘栰偱偡丅娐嫬壢妛偵偍偄偰傕悢棟揑庤朄偑栶偵棫偮偙偲傪抦偭偰捀偗傟偽偲巚偄傑偡丅

惗暔屄懱偑廤傑偭偨傕偺傪惗暔廤抍偲屇傃傑偡丅偁傞惗暔廤抍偺屄懱悢乮廤抍僒僀僘偲傕尵偆乯偼埲壓偺幃偵廬偭偰曄壔乮憹尭乯偡傞偙偲偼捈姶揑偵傕帺柧偱偟傚偆丅

[廤抍僒僀僘偺惓枴偺曄壔] = [弌惗悢] - [巰朣悢] + [奜晹偐傜偺堏擖] - [奜晹傊偺堏弌]

偙偺幃偼丄暔棟妛偵偍偗傞幙検曐懚偺朄懃偲摨偠偔丄拲栚偡傞惗暔廤抍偺屄懱孮摦懺傪乽尩枾偵乿婰弎偟傑偡丅

慡偰偺惗暔屄懱偼丄柍惈惗怋丒桳惈惗怋丄摦暔丄怉暔側偳斏怋條幃偺堘偄偼偁傞傕偺偺堦掕偺婯懃偵廬偄巕懛傪巆偟乮弌惗乯丄庻柦偑恠偒傑偡乮巰朣乯丅摦暔偺傛偆偵嬻娫傪堏摦偱偒傞惗偒暔傕偁傝傑偡丅偙偺幃傪梡偄傞偲丄弌惗偲巰朣乮偦偟偰堏摦乯偑偳偺傛偆偵婲偙傞偺偐偑傢偐傟偽丄偙傟傪婎偵偟偰廤抍僒僀僘偺惓枴偺曄壔偑寁嶼偱偒丄廤抍僒僀僘偺帪娫曄壔乮屄懱孮摦懺乯偑棟夝偱偒傞偙偲偵側傝傑偡丅

屄懱孮摦懺偺嵟傕恎嬤側椺偲偟偰抧媴恖岥乮抧媴忋偺恖娫偺悢乯偑嫇偘傜傟傑偡丅抧媴恖岥偼2018擭偵76壄傪挻偊偨偲悇掕偝傟偰偍傝丄崱屻傕憹壛偟懕偗傞偙偲偑妋幚偲偝傟偰偄傑偡丅抧媴恖岥偑憹偊懕偗偰偄傞偺偼丄枅擭偺弌惗悢偑巰朣悢傪忋夞偭偰偄傞偐傜偵懠側傝傑偣傫乮抧媴撪奜傊偺堏摦偼柍帇偱偒傞乯丅堦曽偱丄擔杮偺恖岥偼夁嫀10擭偺娫尭彮偟懕偗偰偍傝丄偙偺孹岦偼摉柺懕偔偲梊憐偝傟偰偄傑偡丅偁傞擭偺弌惗悢偼弌惗棪乮彈惈堦恖偑嶻傓愒偪傖傫偺悢乯偲彈惈恖岥偺愊丄巰朣悢偼巰朣棪乮堦恖偑昦婥傗帠屘側偳偱巰朣偡傞妋棪乯偲恖岥偺愊偲偟偰梌偊傜傟丄偙傟傪尦偵恖岥偺惓枴偺曄壔乮僾儔僗偐儅僀僫僗乯偑寁嶼弌棃傑偡丅弌惗棪偲巰朣棪偼擭楊偵埶懚偟傑偡丅嬶懱揑偵偼丄奺擭楊偺恖岥傪儀僋僩儖偱昞偟丄擭楊偵埶懚偡傞弌惗棪偲巰朣棪傪梫慺偵帩偮峴楍傪梡偄偰梻擭偺恖岥儀僋僩儖傪寁嶼偡傞儌僨儖偲側傝傑偡乮慄宍戙悢偺抦幆偑昁梫乯丅

偙偙傑偱偼惗暔屄懱傪扨埵偲偟偰屄懱悢偺曄壔偵拲栚偟偰偒傑偟偨偑丄乽嵶朎乿傪拲栚偡傞扨埵偲偡傟偽丄忋幃傪梡偄偰丄懱撪偺摿掕偺嵶朎乮椺偊偽偑傫嵶朎乯偺摦懺傪悢棟揑偵婰弎偱偒傞偙偲偵側傝傑偡丅傑偨丄杻怾傗僀儞僼儖僄儞僓偲偄偭偨姶愼徢偺傛偆偵丄惗暔屄懱偑條乆側忬懺乮枹姶愼偱崱屻姶愼偟偆傞姶庴惈忬懺丄姶愼忬懺丄夣暅屻偺柶塽妉摼忬懺側偳乯傪帩偮偲峫偊傞偲丄姶庴惈屄懱偼姶愼幰偲愙怗偡傞偙偲偱姶愼忬懺偲側傝乮姶愼徢偺弌惗乯丄姶愼幰偺夣暅乮姶愼徢偺巰朣乯傪梡偄偰丄姶愼幰偺屄懱孮摦懺偲偟偰偺姶愼徢偺棳峴偑悢棟揑偵婰弎偱偒傞偙偲偵側傝傑偡丅

屄懱孮摦懺偺悢棟偱庢傝埖偆嵟傕扨弮側儌僨儖偼丄旝暘曽掱幃乮楢懕帪娫偺応崌乯傗嵎暘幃乮棧嶶帪娫偺応崌乯傪梡偄偰婰弎偝傟傑偡乮夝愅妛偺抦幆偑昁梫乯丅拲栚偡傞廤抍偑擭楊乮壗嵨偺屄懱偑偳傟偩偗偄傞偐丠乯傗嬻娫暘晍乮偳偙偵偳傟偩偗偄傞偐丠乯丄偦偟偰姶愼徢偺傛偆偵條乆側懏惈傪帩偮応崌乮偙偺傛偆側廤抍傪峔憿傪帩偭偨廤抍偲屇傃傑偡乯偼丄峴楍儌僨儖傗曃旝暘曽掱幃丄愊暘曽掱幃側偳丄偝傑偞傑側悢幃傪梡偄偰屄懱孮摦懺傪婰弎偡傞偙偲偵側傝傑偡丅懳徾偲側傞屄懱孮摦懺傪堦扷乽悢棟儌僨儖乿偲偟偰婰弎偟偰偟傑偊偽丄屻偼悢妛偺抦幆傪梡偄偰偙偺儌僨儖偺怳傞晳偄傪夝愅偡傞偙偲偵側傝傑偡丅儌僨儖偑暋嶨側応崌偼丄寁嶼婡傪梡偄偰悢抣揑傕偟偔傗僔儈儏儗乕僔儑儞偵傛偭偰怳傞晳偄傪夝愅偡傞偙偲偵側傝傑偡乮僾儘僌儔儈儞僌偺抦幆偑昁梫乯丅傑偨丄嵶朎傗屄懱偺弌惗巰朣偲偄偭偨抁偄帪娫僗働乕儖偱偼側偔丄撍慠曄堎偺弌尰偲偄偭偨傛傝挿偄帪娫僗働乕儖偵拲栚偡傞偲丄偁傞宍幙偺恑壔偵娭偡傞恑壔摦懺傪埖偆偙偲偵側傝傑偡丅

悢棟惗暔妛偱梡偄傞悢棟揑庤朄偼丄奆偝傫偑偍側偠傒偺暔棟妛乮僯儏乕僩儞椡妛丄揹帴婥妛丄摑寁椡妛丄検巕椡妛側偳乯傗壔妛乮斀墳懍搙榑側偳乯偺暘栰偱偼昁恵偺庤朄偵側偭偰偄傑偡偑丄惗柦尰徾偵傕摨條偵墳梡壜擻偐偮桳梡偱偁傞偙偲偑偍暘偐傝偵側偭偰捀偗偨偐偲巚偄傑偡丅悽娫堦斒偱偼丄惗暔妛乮摿偵崅峑傑偱偺惗暔妛乯偼悢棟揑庤朄偺懳嬊偵偁傞傛偆偵偲傜偊傜傟偰偄傞愡偑偁傞偺偱偡偑丄寛偟偰偦偆偱偼側偄偺偱偡丅

巹偼偙傟傑偱悢棟揑庤朄傪梡偄偰條乆側惗暔妛偺栤戣偵庢傝慻傫偱偒傑偟偨丅偙偙偱偄偔偮偐偺帠椺傪徯夘偟偨偄偲巚偄傑偡丅

傑偢嵟弶偵丄捁椶堢帣婑惗偺恑壔摦懺偵娭偡傞悢棟揑尋媶偑嫇偘傜傟傑偡丅堢帣婑惗偼戰棏偲傕傛偽傟傞斏怋宍懺偱偁傝丄懠屄懱偺堢帣楯摥偵婑惗偡傞斏怋曽朄偱偡丅戰棏傪峴偆捁偲偟偰偼僇僢僐僂 Cuculus canorus 偑桳柤偱偡丅僇僢僐僂偺戰棏傪庴偗擖傟偨廻庡偑帺暘偲偼帡偰傕帡偮偐偢戝偒偔堢偭偨僇僢僐僂悧偵塧傪傗傞幨恀偼丄帺慠奅偺晄巚媍偺堦偮偲偟偰峀偔抦傜傟偰偄傑偡丅偟偐偟丄捁妛幰偺尋媶偵傛傞偲丄僇僢僐僂偺戰棏傪擣幆偟偰攔彍偡傞擻椡傪帩偮廻庡偺懚嵼偑抦傜傟偰偄傑偡丅僇僢僐僂偺戰棏傪庴偗擖傟傞偲廻庡偼帺暘偺巕懛傪巆偣側偄偨傔丄帺慠慖戰偵傛傝丄廻庡偼僇僢僐僂棏偲帺暘偺棏偺怓傗柾條偺堘偄傪庤偑偐傝偵偟偰戰棏傪擣幆攔彍偡傞戰棏懳峈庤抜傪恑壔揑偵妉摼偡傞偲峫偊傜傟傑偡丅傑偨丄廻庡偺戰棏懳峈庤抜偵懳峈偟偰丄棏偺怓傗柾條偑廻庡棏偵偦偭偔傝側棏媅懺傪傕偮僇僢僐僂偺懚嵼偑抦傜傟偰偄傑偡丅廻庡偲戰棏捁偑屳偄偵懳峈庤抜傪恑壔偝偣傞夁掱偼嫟恑壔偲屇偽傟丄嫟恑壔摦懺偺寢枛偑偳偆側傞偐偵娭偟偰偼條乆側悢棟揑尋媶偑峴傢傟偰偄傑偡丅偙偺嫟恑壔摦懺偵娭偟偰捁妛幰偲嫟摨偟偰堦楢偺悢棟揑尋媶傪峴偄丄戰棏懳峈庤抜偺廻庡廤抍拞偱偺恑壔丄棏媅懺偺恑壔丄偦偟偰丄廻庡偲婑惗幰廤抍拞偺棏柾條偺懡宆堐帩偵娭偡傞悢棟揑尋媶傪峴偄傑偟偨乮Takasu 2003, Takasu et al. 2009, Fossøy et al., 2016, Liang et al. 2017側偳乯丅



恾1: 廻庡偲婑惗幰廤抍拞偺棏懡宆偺昿搙摦懺偺堦椺乮Liang et al. 2017傛傝敳悎乯丅椉廤抍拞偱4偮偺棏宍幙昿搙乮攚宨怓偑敀丒敄惵丄斄揰偑桳傝丒柍偟偺4捠傝乯偑怳摦偟偰嫟懚偡傞乮嵍偑廻庡丄塃偑婑惗幰乯丅廲幉偼昿搙丄墶幉偼帪娫乮悽戙乯丅

師偵丄庽栘儅僣偺昦婥偺乽儅僣屚傟乿乮儅僣嵽慄拵昦乯偵娭偡傞悢棟揑尋媶偑嫇偘傜傟傑偡丅儅僣屚傟偼儅僣僲僓僀僙儞僠儏僂偲偄偆僙儞僠儏僂偑儅僣偵姶愼偡傞偙偲偵傛偭偰堷偒婲偙偝傟傑偡丅僙儞僠儏僂偼偦傟帺恎偱偼姶愼儅僣偐傜姶庴惈儅僣偵堏摦偱偒側偄偺偱偡偑丄儅僣僲儅僟儔僇儈僉儕偵傛偭偰攠夘偝傟傞偙偲偑柧傜偐偵側偭偰偄傑偡丅寬慡側徏偼徏帀偺偨傔僇儈僉儕儉僔偺嶻棏偵偼揔偝側偄偺偱偡偑丄僙儞僠儏僂偑姶愼偟偰悐庛偟偨儅僣偼徏帀偺燌弌偑掆巭偟丄僇儈僉儕儉僔偺奿岲偺嶻棏応強偲側傝傑偡丅偮傑傝丄僙儞僠儏僂偼僇儈僉儕儉僔偵怴偨側嶻棏応強傪採嫙偟丄僇儈僉儕儉僔偼僙儞僠儏僂傪寬慡儅僣偵攠夘偡傞偲偄偆丄憡棙嫟惗偺嫮屌側娭學偑宍惉偝傟丄偙偺偙偲偑儅僣屚傟偺媫懍側奼戝偵偮側偑偭偨偲峫偊傜傟偰偄傑偡丅儅僣屚傟偼丄偍偦傜偔姶愼偟偨儅僣嵽偺杅堈偵傛傝丄尰嵼偱偼娯崙傗拞崙偲偄偭偨傾僕傾抧堟丄偦偟偰儓乕儘僢僷偵傑偱奼戝偟偰偍傝丄偄傑偩怺崗側旐奞傪傕偨傜偟偮偮偁傝傑偡丅儅僣屚傟偺奼戝偵娭偟偰丄扨弮側嵎暘幃傪梡偄偨尋媶傪旂愗傝偵丄僇儈僉儕儉僔偺嬻娫堏摦傪峫椂偟偨尋媶傊偲丄儅僣僈儗偑偳偺傛偆偵嬻娫揑偵奼偑傞偐偵娭偡傞堦楢偺悢棟揑尋媶傪峴偄傑偟偨乮Yoshimura et al. 1999, Takasu et al. 2000, Takasu 2009乯丅



恾2: 儅僣僈儗偺嬻娫揑奼戝僔儈儏儗乕僔儑儞偺堦椺乮Takasu 2009傛傝敳悎乯丅嵍忋偐傜塃壓傊偦傟偧傟 t = 0, 10, 20, 30, 40, 50擭屻偺屚巰儅僣丒姶愼儅僣偺嬻娫暘晍丅奃怓偑姶愼屻偵屚巰偟偨儅僣丄崟偑摉擭偵姶愼偟偨儅僣丅

偙偺懠偵傕丄擔杮偱偼摿掕偺崅嶳懷偺傒偵惗懅偡傞愨柵婋湝庬偺儔僀僠儑僂 Lagopus muta 偺懚懕壜擻惈暘愅傪峴偭偨悢棟揑尋媶偑偁傝傑偡乮Suzuki et al. 2013乯丅擔杮偺儔僀僠儑僂偼尰嵼偱偼偛偔彮悢偺崅嶳懷偺傒偵惗懅偟丄憤屄懱悢偼2000塇傪壓夞偭偰偄傞偲偝傟偰偄傑偡丅懚懕壜擻惈暘愅偲偼丄婓彮庬側偳屄懱悢偑彮側偔側偭偨惗暔廤抍偺愨柵儕僗僋昡壙傪峴偆夝愅庤朄偱偡丅忔埰妜偺儔僀僠儑僂廤抍偵娭偟偰偼丄儔僀僠儑僂屄懱偺擭娫惗懚妋棪丄帗偺堦暈棏悢乮堦搙偵嶻傓棏偺悢乯丄奺棏偺泎壔妋棪丄憙棫偪妋棪側偳偺屄懱孮僷儔乕儊乕僞偑挿擭偺栰奜挷嵏偵傛傝悇掕偝傟偰偍傝丄偙偺抣傪梡偄偨僔儈儏儗乕僔儑儞傪峴偆偙偲偱忔埰廤抍偑彨棃愨柵偡傞妋棪側偳傪寁嶼偡傞偙偲偑弌棃傑偡丅杮尋媶偱憐掕偟偨屄懱孮僷儔儊乕僞偺壓偱偼丄忔埰妜廤抍偼埨掕偟偰偍傝30擭屻傕懚懕偡傞壜擻惈偑崅偄偙偲傪帵偟傑偟偨丅偟偐偟尰幚偺宯偱偼奺崅嶳懷偵惗懅偡傞儔僀僠儑僂偼偁傞昿搙偱堏摦暘嶶偡傞偙偲偑帵嵈偝傟偰偄傑偡丅暋悢廤抍娫偺堏摦暘嶶偑宯慡懱偺懚懕壜擻惈偵梌偊傞塭嬁偼柍帇偡傞偙偲偑弌棃偢丄崱屻偙偺岠壥傪峫椂偟偨尋媶傪恑傔傞昁梫偑偁傝傑偡丅

惗暔妛偼儈僋儘偐傜儅僋儘僗働乕儖偵偄偨傞條乆側惗柦尰徾偵娭偟偰悢棟揑尋媶傪峴偆忋偱尋媶帠椺偺曮屔偲偝傟偰偄傑偡丅悢棟揑庤朄傪梡偄偨尋媶偑崱屻傑偡傑偡拲栚偝傟丄嫽枴怺偄尰徾偺夝柧偵偮側偑傞偙偲偑婜懸偝傟偰偄傑偡丅

嶲峫暥專

1. Wei Liang, Canchao Yang, and Fugo Takasu (2017) How can distinct egg polymorphism be maintained in the rufescent prinia (Prinia rufescens)-plaintive cuckoo (Cacomantis merulinus) interactions- a modeling approach. Ecology and Evolution 1-8. DOI: 10.1002/ece3.3090.
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5551090/)

2. Frode Fossøy, Michael D. Sorenson, Wei Liang, Torbjørn Ekrem, Arne Moksnes, Anders P. Møller, Jarkko Rutila, Eivin Røskaft, Fugo Takasu, Canchao Yang and Bård G. Stokke (2016) Ancient origin and maternal inheritance of blue cuckoo eggs. Nature Communications 7, Article number: 10272. doi:10.1038/ncomms10272
(https://www.nature.com/articles/ncomms10272)

3. Ayaka Suzuki, Atsushi Kobayashi, Hiroshi Nakamura, and Fugo Takasu (2013) Population viability analysis of the rock ptarmigan Lagopus muta japonica in Japan. Wildlife Biology 19:339-346.
(https://bioone.org/journals/Wildlife-Biology/volume-19/issue-4/13-021/Population-viability-analysis-of-the-Japanese-rock-ptarmigan-i/10.2981/13-021.full)

4. Takasu F. (2009) Individual-based modeling of the spread of pine wilt disease: vector beetle dispersal and the Allee effect. Population Ecology 51:399-409.
(https://link.springer.com/article/10.1007/s10144-009-0145-5)

5. Takasu F. (2003) Co-evolutionary dynamics of egg appearance in avian brood parasitism. Evolutionary Ecology Research 5: 345-362.
(http://www.evolutionary-ecology.com/issues/v05n03/eear1470.pdf)

6. Takasu F., N. Yamamoto, K. Kawasaki, K. Togasi and N. Shigesada (2000) Modeling the range expansion of an introduced tree disease. Biological Invasions 2(2):141-150.
(https://link.springer.com/content/pdf/10.1023/A:1010048725497.pdf)

7. Yoshimura A., K. Kawasaki, F. Takasu, K. Togashi, K. Futai and N. Shigesada (1999) Modeling the spread of pine wilt disease caused by nematodes with pine sawyers as vector. Ecology, 80(5), pp. 1691-1702.
(https://esajournals.onlinelibrary.wiley.com/doi/full/10.1890/0012-9658%281999%29080%5B1691%3AMTSOPW%5D2.0.CO%3B2)