Migra+º+úo de insetos Williams

19
INSE MIGRION C. . WILLIAMS Ad nh, inaig, Snd Te word "graon s consdered or e purposes o e presen ar ce o reer o oeens o anas n a drecon and or a dsance oer wc ey ae conro and wc resu n a eporary or peranen cange o aba. In nsecs suc oeens ay coer a ousand es or ore bu n oer cases e dsances are sorer and e ower o a rue graon s dcu o deerne. Te nubers o ndduas n a s ng e oeen ay range ro a ew undred u p o ousands o ons In soe groups o anas e wrd graon as been consdered o app y on y wen ere s a regu ar o and ro oeen o e popu a on beween wo areas a deren seasons o e year; s pon w be ds cu ssed ore uy aer bu a presen a drecona oeens n nsecs are aken no consderaon weer ee s ye any edence o a reurn or no Pracca a edence reers o oeens o wnged adus ere are records o ass spreadng o arae o epdopera and young ocuss oe seady across counry bu apar ro s s doubu wngess nsecs u nderake oeens wc coud be consdered as ruy gra ory Edence o graon coes ro arous sources wc ay be brey suarsed as oows a Obseraons by ed naurass o arge nu ber s o nsecs ng sead y n one dene d rec on. Suc g s ay as or nues, ours das or een or weeks on end Tere ay be on a snge speces noed or e g a ncude any speces and een a es nsecs o deren orders () Very sudden appearaces o wged nsecs n ubers n an area were ey were no known o be presen preousy and w no edence o oca breedng or eergence. coroary o s s e sudden dsappearance o nsecs wen ue abundan wou any reason o suspec sudden dea. (c) Te presence o ng nsecs ar ou a sea or on oceanc sands and on e snow a g audes n oun an ranges (d) In counr es were nsecs ae been under con nuous sudy or any yea rs obs era ons sow a cera n nsecs on y occur a cera n seasons and canno be ound n any sage a oer es o e year. Edence o s naure as o be crcay exaned e In recen years ere as been edence o graon ro recapures o arked nsecs As a resu o accuuaed obseraons o e aboe ypes parcuary n e as 50 years we now know a graon o ccurs reguary n ocu ss dragones bueres and os soe eeropera soe beees (espe c a y e Cocc ne dae and soe D pera (par cu ar y e Syrp dae bu ere s scaered edence a e ab s een ore wdespread an s HITORICL oug e eares reerence o graon o ocuss goes back o bca es (1 was no un e rs a o e nneeen cenury    A   n   n   u  .    R   e   v  .    E   n    t   o   m   o    l  .    1    9    5    7  .    2   :    1    6    3   -    1    8    0  .    D   o   w   n    l   o   a    d   e    d    f   r   o   m    w   w   w  .   a   n   n   u   a    l   r   e   v    i   e   w   s  .   o   r   g    b   y    U   n    i   v   e   r   s    i    d   a    d   e    F   e    d   e   r   a    l    d   e    G   o    i   a   s   o   n    0    9    /    2    0    /    1    1  .    F   o   r   p   e   r   s   o   n   a    l   u   s   e   o   n    l   y  .

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INSE MIGRION

C. . WILLIAMS

Ad nh, inaig, Snd

Te word "graon s consdered or e purposes o e presen arce o reer o oeens o anas n a drecon and or a dsance oerwc ey ae conro and wc resu n a eporary or peranencange o aba. In nsecs suc oeens ay coer a ousand esor ore bu n oer cases e dsances are sorer and e ower o a rue graon s dcu o deerne. Te nubers o ndduas na snge oeen ay range ro a ew undred up o ousands o ons

In soe groups o anas e wrd graon as been consdered oappy ony wen ere s a reguar o and ro oeen o e popuaonbeween wo areas a deren seasons o e year; s pon w be dscussed ore uy aer bu a presen a drecona oeens n nsecsare aken no consderaon weer ee s ye any edence o a reurnor no Pracca a edence reers o oeens o wnged adus ereare records o ass spreadng o arae o epdopera and young ocussoe seady across counry bu apar ro s s doubu wngessnsecs underake oeens wc coud be consdered as ruy graory

Edence o graon coes ro arous sources wc ay be breysuarsed as oows a Obseraons by ed naurass o arge nubers o nsecs ng seady n one dene drecon. Suc gs ay asor nues, ours das or een or weeks on end Tere ay be on asnge speces noed or e g a ncude any speces and eena es nsecs o deren orders () Very sudden appearaces o wgednsecs n ubers n an area were ey were no known o be presenpreousy and w no edence o oca breedng or eergence. coroaryo s s e sudden dsappearance o nsecs wen ue abundan wouany reason o suspec sudden dea. (c) Te presence o ng nsecs arou a sea or on oceanc sands and on e snow a g audes n ounan ranges (d) In counres were nsecs ae been under connuoussudy or any years obseraons sow a ceran nsecs ony occur aceran seasons and canno be ound n any sage a oer es o e year.Edence o s naure as o be crcay exaned e In recen yearsere as been edence o graon ro recapures o arked nsecs

As a resu o accuuaed obseraons o e aboe ypes parcuaryn e as 50 years we now know a graon occurs reguary n ocussdragones bueres and os soe eeropera soe beees (especay e Coccnedae and soe Dpera (parcuary e Syrpdaebu ere s scaered edence a e ab s een ore wdespread ans

HITORICL 

oug e eares reerence o graon o ocuss goes back obca es (1 was no un e rs a o e nneeen cenury

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   b  y   U  n   i  v  e  r  s   i   d  a   d  e   F  e   d  e  r  a   l   d  e   G  o   i  a  s  o  n   0   9   /   2   0   /   1   1 .

   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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6 WIAM

that aythig i th atu of a iti tuy lop I 185 DS (2) wot a boo o "D Cau Migatio Di Aimaux,i whih h ot about 30 pag to migatio of it A itt lat

Va Bmm (3) i 185 a ag () i 1861 ga ummai of olo th fom hiy of agoi th att hiy of pioptah t ummay of ifomatio o migatio of Lpiopt i thtopi wa by Pip i 1890 (5). Btw 1898 a 1901 utt (6) pubih a uy of migatio a itibutio of it of all o. I1930 I publih () a fu aout o migatio i butti with a bibiogaphy of about 800 f I 1932 Fal (8) pou a gauy of th migatio of it of a o I 192 I pubih ioaboatio with oth (9) a futh iuio o pobm of migaio

i th Lpiopta with 50 mo , a at pt I ha i thp a o boo (10) giig a aout of migatio i al o ofit

h ha b thoua of pap publih about out aligh a th with migatio, but o ummay latig patiulaly tothi pobm a a who. Uao ioy of th "pha of out wat gi i 1921 (11) a hi boo of 1928 ( 12) i a mot impotat gar 4 reeree

MMRY O VENCE I IFFERE RDERS

Orthoptera.-T i i of migatio i tai ga of thAiiia (iluig th out) a light i i om outia(logho gahopp) a i om Matia

h wo "lout ha b u fo may ya fo tai pi ofhotho gahopp (Aiiia) whih ha a habit of agggatigi omou wam a migatig o log ita h a oly io pi i th whol wol whih titly om u thi atgoyalthough amag i o at tim by ma outba of oth Aiiia h "out a a folow: Dociostaurus maroccanus (hubg)hiy i th Mitaa outi a Nothwt Aia, a it oot appa to ma migatio o y og ita Locusta migratoria(iau) with aiou a o ubpi oig at tm outhEuop aly al Afia, Maagaa South Aia, th Eat IiNw Zaa, a pat of Autalia; Schistocerca gregaria (Fo) hiyi th oth half of Afia, a i outhwt Aia a fa a WtIia chistocerca paranensis (Bumit) thoughout mot of SouthAmia Cta Amia, outh Mxio, a oaioay i om of thWt Iia ia; Locustana pardalina (Wa) a Nomadacris septemfasciata (Sill) i th outh half of ia a Melanoplus exicanumexicanus (Sauu) [= Melanoplus spretus (Walh)] whih u to oui omou umb i th Miwt tat of th Uit Stat.

No iuio o th migatio of lot a b attmpt without mtioig ao "pha thoy, whih h t put fowa i 192111). a wi a " of igatoy out th i aoiat a

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   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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INET MIGTN 5

oe olitay oe etay gahoppe ieig oft i olou tutu phyiology a bhaiou but atually a fom o "pha of th ampeie Th olitay fom ha be piouly ow but oiee tobe itit pi o xample octa daca Liau a olitay ga hopp wa fou to be th oltara phae of L mgratora, oe of the tuelout It i ow ow that th "phae i whih the pei ou plagely but ot tily o th eg of owig to whih th iet haebee ubete If gg lai y a igle grgara femal ae ii ito twobath i o of whih th eultig laa a pt ow togeth ath oth i whih eah youg hopp i emo to a epaate ag a ooa it hath, the fome will gow ito grgara lout a th latt itooltara gahoppe At t it wa thought that all migatio wa i thegrgara pha but mo t wo (3 ) ha how that log itaemigatio a uta by th oltara pha. iial of the olitaypha of L mgratora ae oaioally aptu i the Bitih Il afom maumt of th it ha b tablih that hy mut haow om t eeig gou i te la Sea aa a ita of about1500 ml

The migatio of lout wam appa to b moe igula a u peitable ta tho of om oth iet Thi ha tly be howto be pobaly ue to th fat that at the gat height that may wamy th wi pe i geat tha th maximum p of ight of th i t Thu the etemiig fato i thei itiutio may be the wiitio a ot ay powe of oietatio i the lout thmle (516). Th i i of a faily lag egula eaoal moemet i grgara i Noth Afia a Aia Mio e the a uually two boop yea th wit eig bg moe otat oth of the etlt I the pig th i a outhly moemt o ift, a t ummeo "ooo ig ao i hiy outh of t Sahaa Thi i followby a etu othwa to hei wit quat

O th othe ha a xti of migatio of L mgratora iCtal a Southe Afia twee 198 a 193 how o uh aoalpatt (17). Th wa a oigial outba t i th mil Nigeegio of h Sua i the t ya follow i the eo ya y apa to th wt outh a eat I th two followig ya th eatwamomet ahe ight ao Afia a i th ext the ya xteto th outh a outhwt, ahig ba agai to th wt oat i AgolaThi e ya outba wa oughly a lowi pea ou, but aoi ig th wet topial foet aea of the Cogo bai

I the ae of M mxca i th Uit Stat it i ow belie tat

the ol oy Moutai lout M prt) i the ggaiou fom a thpet wily itibut M

Imxan, the oltara fo of the ae

pie Some hag i th eiomet poibly the pea of lag aleagiultu, a pt h lot of the ggaiou fo i te of the lat tuy Although M mca i at time y etutieit oe ot appa the a tti wa M rt ue to

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  s  o  n   0   9   /   2   0   /   1   1 .

   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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66

Thr wa ome vidnce of eglar o and fo saonal movemns inM ru and Rily disingised wn "invading swams wiccame fom h nohwes in Jly o Sem and ening swam whic

ew o he norhwe in May and Jn An inring afemah of hga migraions of M ru in h pas is he occrnce of millions ofindividals fozn ino glais in monains of Monana (18)

T general hory of obreas of locs a psen i ha he aaas, ofn compaaively small, where rding ae lace slowly yaaf ya, chi�y wih e incs in he oliay hase Th aeas arally in dirics of low rainfall A invals som chang aes place,osibly an incrad ainfall prodng moe lian vgaion wicca a graly incad a of mliplicaion, an incad habi of

aggrgaion, h dvlopmn of e rara pae, and nally a spadfom e mo pmann reding aras ov sronding aas, nofen conining for ndred of mil I is cios, and no y rolyndrood, ha h imp of incrad filiy and mliplicaion ap peas o pesis fo sveal gneaions, sling in eensiv damag inareas whr h inec ar no ally known Th oblem of oinaionand wind will b rfrrd o again la on

Lepdoptera.-Th bi mgin s fd all famles f e Ropalocra o eris is paiclaly fn in e ierida

anaidae, and e ympalidae, and lss common in e yanida anHidae In h moh, he hai is widepad in h Singida andoccr frnly in h Nocida b appa o les common in Gomerida In all nighying Lpidopa i i of co dicl o gdirc evidence of migaion, and i is inesing o no ha in om dayying mos (sch as Pua amma (inna) and pcis in e smallopical family aniida) hr a more cords of diecional ighshan in many of i

As an ample of migaion in anaida, Danau xu innas, anes drin smme mons ove ra a of nia and ohn anaa n h amn h bi mov oh wad, a in all gops ofen congeaing lae ino ands ofmany hosands, and nally ac sae odring on lf ofMico and sorn Califonia He y el on e, vey ofn loso h ea, and pnd he win in a ae of mihibnaion n h pringh inaing ands bra p pairing akes plac, and o s moveadily norh, laying ggs on h mlkw wich is j appaing aovh gond. Th hy rpolae aa deed in he prvio amnAccoding o h laide hr a one, wo, or phaps occasionally hgnaions efoe he amn movemn s in Te ig ay b ove1500 mile in ach dicion, and e sam individals wich hav gone soin amn n o no in h pring, he is no evidencha any of e n so fo second joney In si of is same go of s may sd ya afe ya y h inaing in scs, as fo amle a Mony on coas of Califonia The mos

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  a  s  o  n   0   9   /   2   0   /   1   1 .   F  o  r  p  e  r  s  o  n  a   l  u  s  e

  o  n   l  y .

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NSEC MIGRTON

othey hibenting e kown is in the neighbohood o n ancisco(titde 38° bt mny thosnds of btteies y sothwads ech tmthogh Teas ito Meico ad wht hes to these is t esent

kowThee e thee aces of D.  ppps in Ameica distigishabe bythei coo ad mkings The Noth Ameic migt om pxipps)is od s soth s t o the Cet Ameic enis d someo the othe West India Isds he sothen om ripp) is fondsoth of the Amazo ad is obaby so a miat Betwee the two intoic Soth Ameica is a ce  rpps which es to be aeysedenty bt hs been ecoded migtig in Veee Intemediatese kow between the Noth ad Cet oms bt ot betwee the

ea ad Sothe foms The eistece of these foms ssists i gthe imits o the migiosVss r (ies is bttey o wodwide distibtio

i te othe hehee seds eh se oten to qite hihtitdes, d eteats i the wite to dy sbtoic aeas. It beedsaog the edes o the Noth Aic set i the wite nd i sigmoe oth acoss the Mediteane into Eoe sometimes i miionsIt egay eaches the titde of the Bitish Ies whee it beeds digthe smme nd i some yes may go beyond this atitde to Icead oFind whee it h been seen beyod the Actic Cice hee is someeidece o etn ight i the atm The distance om Noth Aicto sothe Britin is abot 1500 mies nd on to Iced amost othethosand Thee is eidece of simia migatios oe the whoe width ooe ad some ecods o mss ights coss the motis i othwestIdi I Noth Ameica the secies aes to beed dig the winte ithe d es o othweste Meico nd Baj Caonia ad oesothwd in the sing sometimes i hdeds of miios ito the neihboing states o the nited States he tota amot o immigtio ndsed ies eaty om ye to yea bt it ot ieety oes noth wd into Cada nd nothestwad to the New Eand States theGf of St Lawence d ee occasioy to Newodand hee isno eidence of winte sia ywhee i the Uited Sttes ecet ehs in the e teme sothwest and o eidece of y etensie imigatio thoh Foid o aywhee ese aong the G of Meico hee aea few ecods idicatig sothey moements in the atmn bt they codot t esent be cosideed as cocsie The distce om NothwestMeico to the moth of the t Lwence ie is oe 3000 mies

These two secies ae tyia of the smme migtios o btteies

ito teet nd codtemete eions and they cosey esembe themoements o my mit bids The ootion of mint secies isch climtes cn be ey hih In the Bitish Ises whee the btteyfa is obaby bette kow th aywhee ese in the wod bot seies ot o a tot of 6 e tiay o coetey deedent o immigo o thei occ ece the Ute tat o 0 migt seis ae

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  a  s  o  n   0   9   /   2   0   /   1   1 .   F  o  r  p  e  r  s  o  n  a   l  u  s  e

  o  n   l  y .

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6 WILIAMS

alady nown t th lit will proaly conidraly xtndd withfth tdy

In th topic ttry miation a on a vry xtniv cal and in

om ara ar o nt and conpico to com oc of ptition to ndcatd rac In yon 0 ot of a tota of aot 250 pci ofRhopaoca hav n codd a takn pat in what ar localy knowna "ht hich omtim conit of notom lik ma of milionof individal all in tadily in on dirction omtim for day on ndThi oiin and dtination a nnown t thi aonal occncappa to latd to th monoon with a pa of occrnc at taot th innin of th nothat monoon which in th ain (19,20)

In tropical Afica thr ar many rcord of ma iht in a dnitdiction chiy atn to th pci of Piida and on of th familyLiythida. In th form family atopsiia orella (aici) ha npotd a a a miant paticlay in th atn haf of th continnt fom th Union of Soth Aica to a fa north a Eypt whr it ian occaional mirant In Sothn Rhodia it mirat in nm to thothwt in cmr and Janary and thr i om vidnc of a rtniht tds th thst t sx k t Lby bdWtood mo to th oth in th Gold oat and Niia aot March

and Apil which i toward th nd of th dy aon and thr i om vidnc of a cond miation aon lat in th ya (22In topical Amrica miation occr on a iantic cal pcialy in

th n  hoebis ieridae hoebis seae (inna) and   hoebis statir(am) hav n cordd in ma iht ov mot of topical AmicaTh fom mirat ach prin into th othn nitd Stat and thi dnit vidnc of a rtn to th oth in th atmn (23 In thePortachlo Pa in th coatal an of montain in Vnzla B(24 ha codd contl million of tti, onin to ov 250

pc pain throh from north o oh amot withot a ra romMay to Sptm vy ya. ay and nht yn moth and many othod of inct w alo incldd in th iht Nothin i known of thioriin o dtination o of any rtrn movmnt.

Simia miation hav n cord in Atalia, and on pci ofHprida Baamia excamatiois Farici) mirat alon th at coatof Qnland with vidnc of a movmnt to th oth in Dcm anJanay i.e prin and ary mmr) and a tn ht to th noth inMarch It ha n aid t withot any xact vidnc that wam of thittry co th a fom Nw ina to Qnlan (2

In th moth vry xtniv miration occ in many pci of thSphinida hich a nally la in with a powfl iht Otof 7 pci of th family nown to occr in Britain a complt miant an ldom o nv viv th wint in th latitd On of thmo inttin pci i Ceerio ieaa (Farici) which ha on pci eata (aici) in Amica and anoth, C.  or

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   b  y   U  n   i  v  e  r  s   i   d  a   d  e   F  e   d  e  r  a   l   d  e   G  o   i  a  s  o  n   0   9   /   2   0   /   1   1 .   F  o  r  p  e  r  s  o  n  a   l  u  s  e

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  C MRT 6

(Epe in Afi  nd Eope The e beed in oh Afi ding hewine nd invde Eope in m o ge nmbe in he ping Theve feed on ine nd e of he immigion hee my be ex

enive dmge o vineyd in Spin Iy nd Sohen ne In omeye he moh y i fhe noh nd my eh he ohen o ofEngnd Hee howeve hey e iie nd in he p hnded yeony bo 7 moh hve been epoed by oeo of whih 53 weeped in 3 hee hve been mny ye wiho inge eod InNoh Amei hee i vey imi e of i. The ine beed ding he wine omewhee in Cen o Soh Amei mo einyin he emiid ime of he ween o Thene hey move noh inhe ping ino he ohen e whee in Eope hey beome

mino pe of vie In dy of he p hioy of obe of hi ineGn 2) hoed h hee w iy ignin evidene fo yeof hevy o of igh invion o o mneoy on boh oninen

Among he oide hee e mny nown min of whih hioi y he oon efwom laama ariaca (Hbne) i of ei inee being he e oid Eope in whih migion w ggeed fo moh Thi epnion w p fowd in 8 by Gohm 2) o expin i io eo oene b i w no ni bo 5 yee h he heoy beme geney eped Thi peie h ome moe

o e pemne beeding gond in he nohen hf of Soh Amei,fom whih wm mige nohwd ome by wy of Cen Ameind Mexio) o invde he oon be o oh Amei. Thee i pideie of geneion foowing one nohe ding he gowing eon ndhen he ine ddeny dippe nd nno be fond in ny ge nihe new immigion in he foowing ye Thee i vey pobby enigh of moh o he oh b he ine i non nd he nbe oied obeve in Soh Amei i i ey nd o f o obev ion o h movemen hve been bogh fowd i howeve e

be nd expined ee h in he f when one wod ex pe h ohey igh moh my ppe in he noheen e ndeven in ohen Cndix o even hnded mie noh of he oonbeoeime in h nmbe ond he eei mp in iie o be poiive nie A he me peiod he moh my beome mino peby pieing nd ing ipe fi h gpe nd pehe 28).

A mo embe e of migion in noid moh i h of rotfs (Bidv he oed Bogong moh o Ai I h beennown fo ove eny h he moh ongege in ge nmbe inve nd ef n o high id in he monin o New SohWe Thei nmbe wee o ge h he boigine of hi e ed ome oney o e ve whee hey peed he moh wih moe nded hem oe of ood whih i vey ih in f Reeny Common) h nveiged he pobem nd nd h moh whih hve beenbeeding ove vey ge e in he ownd ding he wine, move pino he hi in e Oobe nd ovembe ie png nd go ino e

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   b  y   U  n   i  v  e  r  s   i   d  a   d  e   F  e   d  e  r  a   l   d  e   G  o   i  a  s  o  n   0   9   /   2   0   /   1   1 .

   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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WAM

of almo coml domancy in cav and imila lac Th olaonah mamm y mdDm ofn a many a 500 inc qa foo of fac and h i no dclin n nm nil h nd of

Mach (amn) whn all aidly diaa All h moh in hi a ofavaon a ally mma Th acla cav nvgad yommon w in h Bindalla Rang in h aial Tioy ondana a high of wn 4000 and 6000 f A omwha mila cai fond in orzros ars (o in oh Amica which ha cnly n fond o aiva nd on on c a alid of ao000 f n h monain of Monana 0

n h Amican oi h a wo maal dayying mohUr es (inna) and U fes Wal oh of which a gla

migan hy a ghly colod wih long al and a qnly mian fo wallowail i U fes ang fom Mico hoghnal Amca o olomia and Ecado and cod a fqnly cvd of lag nm coing h anama anal h igh aa o mo fqn wn Mach and Sm and in om of h nalAmcan con h vidnc of gh n wo dicon a di nim of h ya A long ago a 57 Fidich a qod y VanBemmeen (3). se is secies wngl ienie a  emga vy ya in oha Mco gong o h oh fo ao h

w in Ail Ao v o i w la h i a n igh o hoh in mall nm h fmal wiho gg Thi cla c amn ha no y n conm may wll coc

Odo-W hav val hndd cod of ma gh of dagoni moving adily in on dicon h infomaion i ally o incoml and o fqnly h ci i nidnid ha i dcl on h daa In Eo h chif migan a Le depressinna qdrm Linna and mperm sroam (hani) many oh c a alo involvd Of h 4 ci of dagon

nown inG

iain0

a aial o coml immigan In4 h wa a v xtnv imrin o in u

of and fo wo o h day and i i amo cain ha hy had madan ova igh of ao 500 mil fom h Sanih ninla Tha aid o qi gla annal mgaion in a ohwly dctionalong h ohan ho of h Balic Sa and ma migaiona fqnly n fh noh in nland 45

In h nohan nid Sa Shannon 6 ha codd glaohly o ohwy movmn of dagoni in h amn alongh ho of ong Iland in oan wih migaing Monach ie

pps h c concnd w Lbe pe y j y and Trme er Hagn On on occaion h tmad ha 50000 inc ad along h coaal i wihin wo hon gnna esk orss Ram aa a m in conlmillion ying o h noha a ho m fo h aival of a v

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   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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INECT MGION

vioen souh-wesery wind ocay known as he "Pamperio Non arseen o reurn (37)

as ben sugges parcuary n wesrn Europ hat mass

ighs of dragonis ar assoiad wi h drying up of ir breednggrounds, u h vidnce sems incocsive

ClTer ar quie a nmber  of recors of sady gs ofbs in one irecion, bu i s ony in e occinia ha he evncis sucin for us o begin o see any paern In some of h species of isfamiy hre appear o e rguar sasona migraions beween arge breding areas and smar winer or summr uarrs wher hr s a dormantpriod hse rsing paces ar ofen a hig aiuds In aifornaHippdamia  r Gurn breeds during he summer n e coasa

pains and owr vaeys and in e auumn he bs y a undred mesor mor o he mounains, were hy hirna undr sones dad eavesc a eighs of abou 5000 fee (38) n e spring e bac o bdng areas For bernaon y of mas togeher in ily packdaggrgaions of housands of inscs an a on im s wre cocand kp in coo sorage, o e irad during he summer wen requiro conro ouras of aphids I was sad a wo men coud coec ovr  mion bes a day Simar masses of bees ave bn found in isin h midde of h summer, bu wer se ar eary comers or a

sayrs or some nw phenomenon, does no appear o be propery undrsood he is evidenc of smiar winr aggrgaions in mounains inmany pars of he Unid as bu no gnera sudy of migraions as aprobm of is own has ye bn mad

In man pars of Europe or Africa and Asia Mnor spces of adbe have ben found n smar aggegons usuay n wnr bu somimes   n misummr, ofn on mounain ops bu aso a sa v herare man records of suddn appara f gra swarms n nganofn aong he coass u somims w nand In 1925 Marriner (39

cocd records of grea swarms of daa paa (Linaeus) wic adsuddeny appeard, and e considerd a y had moved in a norweser direcion from h sou midands o exrem noreas ofngand (a disanc of or  200 mis) an a crossed e cenra rdg of nnine His in wo paces n Egyp in Apr 1939 Over  0) rporvas numes of ia dmpaa innaeus coming in from e seon e derranan coas n addiion o he miions ing er was adrf ine for a eas fouren mis aong coas wi an sima nur of ovr  50000 es per foo

hr rdrhr ar man observaions suggsing mraion among

e Heropera bu ony one in any ay propery sudied is a of hsma pnaomid rar rp uon in a Mnor wr i is srious pe of barey he insecs pass on gneraion n e sprng fedng oncreas in vays, as for eampe in zksan a a eig of aou 2000 o000 fee e aus engorge w food eave e vas bou June and

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  s  o  n   0   9   /   2   0   /   1   1 .

   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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WAMS

t a et f aut 7000 t 8000 feet n te untans wee teaestvate n te autun te ece actve aan and descend t aut5000 feet wee te enate. e n Mac p nt avn taen

an fd snce te evus spn t ack t te eve f cutvatnfeed ate vst and de Tee s tus n eac ea a sne feccewc as ne feedn ped tw peds f danc and tee pedsf atn 4

In te ptea t f up t sevea nded es ae nwnn te eet eafe Circulir nllu Bake n Nt eca 42wc n ste f te sa sze f te nsect aeas t e ae nde endent f wnd dectn

n te enptea tee s a eaae case f setn cse e

sen atn n ne f te wn was phx agypiacueeete wc uses custs as fd f ts un In ast fca n te ava f a swa f hioa geaia was at nce fwed teaeaance f tsands f te px nt ne f wc ad een seen n tedstct n te we f te eed ea Te wasps uwed n teund and caut aased ad ed custs f daeak t sunsetf tw das en te cust swa depated and wtn an nt neSPx ws   be s A  e  wss w fes 4 ae sevea tesa cases wee epted and e seve saw ac ees n a ass te

se f a tee n n te dst f a swa f custs Tee s n dttat ts speces as sew deeed a at at n de t keepu wt te wandens f ts e It sees ae tat n suc a casean entatn s n a questn f tatn te csts and s nt de tened ndependent te wasp.

n te up f te sawes entedndae tee s as se evdencef atn and Ahalia roa  naeus as unduted cssed fte Cntnent t Btan n enus nes ut at n ntevas 44

n te ptea tee ae nce e scatteed ecds f ss ts nne dectn n an faes n a ats f te wd ut at pesentte evdence st dente n te f Spdae. n tese ve tee appea t e eua seasa ent t n tn u andn te easten Unted States ad pa esewee In te autn f880 e (45 ecded tuands f Eisali ylaiu Meen and yphuslanula  Meen vn t te sutwest up ne f te vaes n suteasten Swtzeand Sa sutwad autun atns ave eenfund ecent t f eua ccuence t an f te passes f tePenees 46 47 48

un te se nts eat swas f Sdae ften apea nte sut an east casts f Enand ut as te n s unknwn t can nt e assued tat te epesent a etun veent f te st.Qute ecent sute autun veents f ve es ave een epted t e f ea ccuece te east cast f nand at Sunead 5 In eca Sannn 4 as seen autun veents f alsn sutwad an te ses f New Jese It s eakae tat te

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   b  y   U  n   i  v  e  r  s   i   d  a   d  e   F  e   d  e  r  a   l   d  e   G  o   i  a  s  o  n   0   9   /   2   0   /   1   1 .

   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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INSECT MGTON 3

bsrvd mgrns n Eurp n nud n h sm gh sps gnr h ly drn d hbs n h lrvl sg: yrphuhh r phdrs nd Erisis hh h mlr rld rv

ds dyng vgnn h grns Srphd hr hv bn ssd lrge

nubs hr Dpr nludng sps h gnr Cllipo ndLuili s s grups (n h my Clphrd ms s b nsdrd s pssbl mgrns

CU

hs bn nssry g n s dl n srvyng h vdn mgrn n nss bus unl brd mgrn h s ppr

b s ll nn vn mng ngss Evn h h grly nrsd nubr bsrvrs nd bsrvns n h ps 2 yrs sllh nly vry supr nldg bs h nrm nmbr sps n h ss ns nd dly png ld dt hot spemn f

W n n s hvr h h hb grn s dspdmng mny' grups ngd nss hugh dnly mr mmn nsm mls hn hrs n s sps h mgrns r s dn nds rgulr s hs mny brds; n hrs n sp mny bsrvns

h prn s sll ndsn n h jry sps h mun nrmn s q nsun r ny hrsng Th ghs my nludhundrds llns ndvduls nd n xnd vr hundrds mlsThr q rqunly ghs n drns dn ssns ns ss h gh n n drn bng n rgr nbs hn h p ps gh n h pr rgns h vns r nry ysy rm qur h sprng nd rds n h uun sr sly rld mprr hs h s l r ns r r unrn bu n yln nd shrn nd hy d nly rd h hngs h nsn.

n sm ss brdng s p bh nds h rng n hrs hmgrns r bn brdng rs nd rs r hbrnn nd svn n sm ss s n h hlls h Hls h ghs sm b up hll h bgnnng h h nd dn hll hn h pr urs ll. Mnn rngs 1 r vr y b rssdd rss ghs hndrds ls rgulry ndrkn Eh yrmllns nss sy rss h Mdrrnn nd r n pprnly nd s nrrr prs n nrly l h ndrds mss ghs pdpr h hv bn bsrvd h nss hv bn yng h

nrl gh r rdnry vy nry ll bl rm h grndh prhps snl srggrs r 2 vr xpnsr nn D  peippus hs rqnly bn rprd "s r n h r sh y n s nd durng vry lrg rn Asci mout (n ns n Suh Amr nubrs rrd by rplns p svrlusnd Thr s vdn h bs a wn

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  s  o  n   0   9   /   2   0   /   1   1 .

   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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4 WLAMS

the wnd ehnd them nd lower when there hed wndThe few ht of Dpter tht hve een oeved hve ll een

te cloe to the rond nd n the tmn ht cro the renee

the re often jt mmn the rond In he loct on the ontrr lthoh ome wrm on dtnce lke lnet over therond follown ll t ontor mn other hve een reported thondof feet n the r nd t ommon to ee them lke rown moke hh ovewth the ndvd nect ndtnhle even wth eld le

Hvn otlned the ft one ome to the prolem of how nd wh?Of thee perhp the mot fndmentl the ntre of the oentton o hmrtn ndvdl, wthot renttn n dni dretn htwod e mle nd wthot n form of relrt

There no dot tht mrtn nect re le to eep for d fterd nd mle fter mle to ht n dente dreton n pte o hnen the tpe of ontr nd t ontor whether the re on ld or e ndn pte of mn chne n the wether Th pple not onl to dnmrnt t to the thond of moth tht nht nd to ndn nect tht hve to contne to nht when on lon eron e ontn of drectn t let n ttere mrtn nerthe rn, v a aka If th obsacl h hoe or tree the wll nerl lw re nd oer em rher th

trn to left or rht nd o rond Mrtn ttere hve een nown to throh rlw tnnel nd nto wndow t one de o he nd tt the other Another nteretn pnt tht f the re n over the topof thk foret nd come to ern the wl deend t the ner eded re n t the fr de t do not trht over t the ornl level :the me ht h een noted n ttere cron rrow vlle orcnon On the other hnd the dreton of ht n e teporrl dverted horelne or the n of llys, ut n h ce oonero lter the net wll te p once n ir orrct orentton

Tere re mn e o derent pee o net ll toether nte me ht the me dreton t o fw rvs on deentpee n t the me tme nd pe n derent drecton For exmpen t Afr n 1929 there w for nerl month a northeterl htof Catopsiia forella Fr) nteln wth othweter ht ofTera enegale Bodvl th erd whle for two d of thmonth there w thck ht f lct geara, on to the othetThe three trem were ll nder ten feet frm the rond, nd the orentton of ec pece hd no pprent relton t the other

Mn fctor hve een eted externl de to orentton the

n te wnd rdent of tempertre or motre or rometr prere,the erth mnet ed, the movement of retnl me nd o on Then d the moon t nht) n thnk e rled ot nl de nlewe re wn to ow nect ene of tme whh wold enle them tolow for the movement of the n not onl drn the d t for t et e we d n net ornn n the et rthermore

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  o  n   l  y .

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orienion kes lce nigh in sie o he oiced oeens he oon nd is eriods o drkness Also orienion is uie denie iddy in he roics when he sun is so ner he zenih s o be rialy

useess or horizon orenions. I s on he oher hnd, ery noiebleh igrion in bueries es oen eses or is uh reduced whenhe sun is clouded bu so so o ossibl siir exen is norl nonnigring ciy The se criicis cn be eeled gins he use ooried igh (suggesed s n id o he hoing o insecs bu no sor s I wre, or heir igrion . The direcion o lne o olrizionreie o he insec deends on he osiion o he sun nd o couds or ohe oon nigh l o which ry ridy

The wind direion is uch ore like cor, bu in he Leidoer

es I he coe o he oncusion he eidence is srongy ginsny rey obligory ssoiion (0 Alhough we know o ew ses oigh direcion ssocied wih chnge o he wind (soeies dierenebeween orning nd ernoon o he se ere re nes o oserons o ghs coug sedy in one direcion dy er dy insie o chnges in wind direcion soeies los round he ossor exe in y 197 Pers brsse (Linneus ws ying o he norhner London or bou six ds during whih he wind ws ro he souhsouhes wes es nd norhes The se secies ws ying in he se

oiy in Juy nd Augus 190 owrds he souh or oer wo weeks wihwinds ro he souh souhwes wes norhwes nd norhes Asmouste ws igring o he souh on he Anic os o· orid in hesring o 19 nd 199 wih winds ro eer oin o he oss, buhiey ro he souhes nd es Le in he suer o ech er wsing o he norh wih os excy he se disribuion o winds Ees ike his cold be ed os ineniel I is ion oreeber howeer h l hese gs o eidoer were king ewih os indiidus ess hn en ee ro he ground.

n recen yers Kennedy Rine nd ohers (15 1 he sressed hec h s he xiu seed o gh o ocuss in sil ir is no uhoer en ies er hour nd s swrs reueny ise o heigh o 1000ee or ore hey wil reueny be in condiions where he wind elociyis geer hn heir own. As resu e drecion o oee o he swrus be deerined by he direcion o he wind los irresecie ony eor o he inses o y in ny oher direion A deied orisono swr oeens in S. gregri wih synoi weher chrs hssrongy suored his conceion In he edierrnen nd Norh Arihe prevag winds he srong norhery oonen: in enrl Aricahere is sed souher coonen Where he wo sres ee isknown s he Inerroi Conergen Zone nd i is long his be hhe swrs end o ongrege. Chnges in he liude o he conerenon re ssocied wh chnges in he disribuion o lous srs ndbnor winds re oen ollowed b bnorl swr oeens. Sinin ny ccloni ir sse he winds re bowing sirll in he enre i

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   F  o  r  p  e  r  s  o  n  a   l  u  s  e  o  n   l  y .

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WMS

follows ha there is also a tendency for locus swarms o be moe fequenn areas of ow pessue This was noed n Egyp y Gough n 1915 51

Since rainfa ends o e moe euen owards he cene of a cyconic

sysem we ge he ineresing resu hat igh down-wnd w brng heinsecs no aeas whee rain s moe ikey o fa As ocuss ae n genealnhabans of semaid aeas hei oemen wih he wnd will auomacay resul in hei eaching aeas where egeaon is liely o e moe luxuran

The orienation of locuss when near the gound is sll not properyundesood In wndy condons hey ae o and land agans he wind aswil mos ohe nsecs Somemes hee may e simulaneous ghs n differen drecons a dieren lees My own experience n Eas Afica showed

no eidence of a seady reaion beween igh decion and wind, pacuay if one eemes ha hee is a pevailing wind diecion and peaiing gh diecion, a eaion of cause and eec s ey dcul oproe unless changes n he one ae eeced y changes n he ohe

Locuss and some ueries) when migaing in a gusy wind hae eenoseed o ale he body aes dung a sde gus, so ha he diecon ofovn elae o he gound tac wa ept contt by yng hediecion of he gh houg he a couse 43 This would ndicat

a dene means of oienaion indpenden of any eaon o he wnd and

aso of any image of he ground in he reina of he eyes I is mporan orememer ha he mechansm of orenaion mus enabe he nsecs oesume hei euied igh diecion afer hey hae een dieed fom eihe momenaily y guss o wnd, o y isiing owes o small osaces,or fo longer peiods y darness o bad weahe

adiens of emperaure mosure or pressue al fail o account fo oenaon, as none of hse ae consan a one spo Wih he elaelyshot disance ha an insect can moe n one hou he changes along hegradien oaned y such a moemen excep possiy uphi woud

scarcey e dieren fom he aerage cange epeience y tyin tilWhche dco an nsec moes and een i i does no moe in healy moning o a summe day, he emeae is iey o ise

There s as ye no edence of any inuence of he eah's magnec edon he oenaion of insecs 2) nor any suggesed ogan n he nsecody ha could e supposed o e sense o magneic poaiy muse aditted hat we are a prese whou any saisfacory epanation ofthe mechansm of oenaion n mgran nsecs u, consde ha n spieof he nown eect of wnd on locus swams such an oientaion exsts asa fundamental facor in ther migraton

second mos mpoan proem s ha of he socaed "eun ghtFor many yea i ws admied ha inec coud y uie arge disancesin a consan diecion i was consideed ha hese wee only dspesalghs of a populaion oeowng fom an ercowded pemanent breedng area lighs of his type were though o e deren n orgn andmeanng from the tue mgraon of rds n whch there was a oandfo

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NSET MGTN

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WLAM

Som mgrton undoutdy trt tr n x uld up o poul ton ut n n mgrtory p grt undn nd not rut nmgrton. Two r known o normou o bundn o th r

o th prd uttry Gyes uro (ru (=mee) on nnd nd on n t Ar whn th ood pnt wr mot dotdIn nthr w thr ny dn o mgrton nd th rutng mono buttr r gglyng on t ot h ompxty o th rtonbtwn grgton nd mgrton w hown n Ds pZeppus nth th r n ummr r wdy ttrd or lrg r dng onnt o kwd (ses In th l th buttr trt to mo toth outh nd grdully ggrgt nto rgr nd lrgr group untl whnthy rh th outhrn tt thr my ggrgton o mny thound

o nddu Hr thy o th mgrtory ntnt but rmn ggrgtddurng thr ntr dormny. In th rng th rr hng hnthy o t o thr grgrou ntnt ut rum thr mgr ton k to th north.

Th xtrm thnn o olton tht n undrtk ut dntmgrton prh t obrd or th In th dtrrnn Ih n tdy northwrd momnt o Vess ru whn wth tho wh, l tn a dzn burf hor r n 57. hdnty mut h bn o th ordr o thr or our pr ur m On th

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r r grty rom yr to yr n th o V cru t omtmmot nt rom rtn durng ummr nd omtm ry bundnth m rton o ound n th Untd tt. A tttl tudyo th yr o 'unuul undn n th two ontnnt h hown n th o Ceero e rdy mntond gnnt tndny orth p to thr mor omon or l ommon multnouy onoth d o th Ant. Th uggt tht th ondton (proylmt dng to unuuy rg mgrton mut tnd to ry n thm drton or r orng prhp onthrd o th rth rum rn n nrly ll mgrtory ght oth x r prnt n prox mty u proporton though thr r rortd whr on xonly ourrd. n h to b rul n rgung rom th numr o turdmn omtm on x mor y ught thn th othr ut o33 drt out tht ught n t Ar 2 wr m nd 337 ml n mgrtory kppr buttry dromus eder Pt thptur wr xty 97 o h x n th othr hnd th phe mgrtngth th lout n th m r r m A ttmnt runtymd th only th ml monrh buttry mgrt to th north n thprng norrt

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SCT AT 7

hd thr r udoutdy ss o grto hh th sts ropty tur d y ggs durg th ght Ths hs osd Pri baica Egd d othr prds Cyo, Est Afr

d ustrFor out thrty yrs thr h rous prts o r

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s osdd ssry to out for th osrd fts

TERATURE CTED

Te B xdu ha. 0 erses 2-1 ( 200 B.C.)2 D Seres, ., De aue de Mgain de Die Animaux (agy F

Pars, Frace, 626 pp 843 Va A A Hand. Nied. nm e 23 (8574 He, H Se n eiung 7383 (86 Pper, M. C Nau Tijd Ned nd (Baa , 9827 ( 890 ; 7,

076 (897

6. T, ]. W., Migain and Dipea ne (Ello Sock Lodo, glad,32 pp, 902 reped wh addo rom ere o acle hEn. Mny Mag (898902)

7 llam C B Te Migan Buie (ler ad Bod dbuhScolad, 73 pp 930

8 Fraekel, S ge Bi 9, 238 (9329 Wll . . Cockll, . F, ad Dow J A an Ry

En. Lndn 92 0280 (9420 Wllam C e Migain ne (Coll odo lad re) Uvarov P Bu nm Reea 12 363 (92

2. Uvov, B Lu and appe (mpral Bureau o Eooloodon Enland 32 p 928

3 Ramchadra o ad Bhaia, D. ndn J Ag. Si 9 7907 (939. Dave T N 72, 720 (93) . Keedy, J. S Tan Ry. S (Lnn [B 63290 (9)6. Rainey R C Nu 168, 070 (9 7 rov u oo Rr \ 0

Statoer Oc,   odon glad 67 9

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180 WILLIAMS

8 Guney, A B, Smionia Ins Rep for 5 302 92)9. Mande, N. Tran Enomo Soc London, 708 904)20 William, C Tran Enomo Soc London, 33 927)

2 Cockbill, G F, Proc Roy Enmot Soc Lodo, [A]26 328 9 22 William, C B, Nigerian Fid 6 29 9)23 William, C B, An Enomo Soc America 2 39 938)24. Beebe, W, e a, Zoooga 439 947) 070 92 949) 35

768, 8996 90) 36 6 2433 266 9)2 Pxle, W L, Gee Iand n lierig Sea p 29 George llen & Unwin,

Ltd, London, England, 3 6 pp, 92)26. Gan, K J , Tan Roy Enomo So London 86 347 937)27 Goham, D B, De Bow' Revw, 3 343 847)28 Wolo, G N., Am Naurai 6 8287 929)

29 Common, F B, Auaian 1 Zoo, 2 22363 94)30 Chapman, J A, Rome, J I and tak, J, Eoogy 6 68 9)3 . Williams, C B, Pro Roy Eomo Soc Londo, [2 447 937)32 Longeld, C, Irih Na I 9 33 948)33 Williams, C B, Eomoog, 62 448 929)34 edee, H, ca Soc Fauna Foa Fenn , 38 908) Vale, K J, Enoo Fnii, 2, 4 96)36 annon, H. Hap' ag 62 9)37 Hdson, Hi i Rim k, d d 0-8 ( Lnn Eg

n 0 pp 9)38 Cane, E K on Bu Se Cmm Saameo Ca 778 9)39 Maine, T F Enomo Reod 046 939)40 Olive, F , o Roy oo So odo, A) 888 943)4 eedelski, A A Rev Ap Enoo, 40 309 92)42. Romne, E, U S Dep Agr Circ, 18 4 p 939)43 illiams, C B, Ann App Bio 20 46397 93344 Ja, G, and oeton, D Enomo Mony Mag 4243 948)4 Eime, T are Ve Vaeand Nauu WUemb 38 03 882)46. Lak, D. and Lak, E, 1 Anma Eo 20 637 9)47 Ga, J H, Loke, , and Pnam, C D Eomoogi, 86 687 93)48 Wiliam, , ommon, , nh, R A, Ma, , an lliam,

M C Ta Roy oo o odon (n e)49 hannon, , 1 Y oo So 34 9920 926)0. William, C , Po Roy Eomo So Lodo U 784 949) Gogh, L H, Reo o e rea Ivao o Lou n Egyp i 5 p 7

ovenmn Pess, Caio, Ep, 72 pp, 96)2 Keewe, H D Proc. Roy nomo So London, ClI2 43 947)3 ettewell, H B D Naue 6 83233 92) Fe, J c Si Bu Dep Ag Fo S Ara, No 234 943) Fae, J. C, ag S Ai, 18 6978 943)6. Long, D. B Ta Roy Eomo Soc London 48 93)

7 William, C B, Tan Roy Eoo Soc London, 20733 923)8. Uqhat, F A anadn Eomoog, 7 222 94)9 Owen, D F, Enomo Moy a 92 4344 96)60. awad, K J, Pro Roy Enoo Soc London [)28 6373 93)6 . Uhat, F A, Repo o Movemen o e Moar Buey (Roal Onio

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