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THE MEIl:MITHID ,NEMATODA) AND OTHERENDOf'A,RASITES OF SIMUl IDAE{DIPTERA\IN.N$ULAI! NEWFOUNDlAND

CENTRE FOR NEWFOUNDLAND STUDIES

TOTAL OF 10 PAG ES ONLY MAY BE XEROXED

IWithout Author'sPermlss,onl

BARRYALAN EBSARY

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3G2:: 7 0

, ,

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Pre6e~tedto'_ theDep~ri:ment~f 1l1~logy. He;"ori~~Unl v .!!uhy ofNewfoundl~d,:"

.":ofthe_Require~entsfor the Degree

Masterof'Science'

.( D" , Barry"Alan: Ebaary

C' "

Apdl. l91J

(6)

" f AESiRACT

.

.

In-1971,and 1972,198'at:~amsin in8ula r Newf ou n dl an dwere examfned for b1a,ckf1yendoparasLcee , For ty streamsyielded onecr

. .

mo re speciesofthe micros porida--tnelohania bl'Gcteata;,T. ,fibrata~

Plj.etpphora simutii,caudoepara simutiiandC.brevi cauda.Th efu~g.u8 Coe'LomycidilUTlsirrrn'Liiwasfound in14~tream s. ,ThebLackfLymermithids qastl'Oinernris viridis,'Leomemie.wieeoneineneie

a nd

Neomeeonermie [lumenaliswere recovered from 59 str e ams.Cen tralNew,found l a ndhad

"

" .

higherfrequen cies'of mermithid.infections (47.3-75 . 0%)tha n the more"

. ,

...J:oas ta 1 ar e as (11. 7-36. 3%).Thelongi tud inaldi strib uti on of mcndthid ", infe c t ed blackflies wi t h i nthest r eam.sh ce ed thatthi{'upper 40011'1. of thest r ea m were devoidofme itnithids. whP~th e lowe r rea chesofth e strealllharboure~mermithid s,.Current,depth 'an d'chemi c alParallle~e";-8 of thestr eam waterinetreamewithand withou t mermit h ids were examined.

~ 1thld .par,~s~tism .

dd.d..'00<

ap;",. ;~ b~. a'';' '1'<8n<1, .

affected by hese facto rs.Neomesclnernrie {lwnena'Liswas,rede s cribed in l ight of t .fLndfng of eggsand pre-pa r as ites.Adultmale s of, 'G.viri di sandI.uieconsine neiewereels aredescrdbed,Thelife

cycl~.~.f.N:./'L~ena.'Liswas des c r:i bed.F~llowing_emerge~ce.from'the

hoat,maleand femal eP.o·at-p a ra eitcs .mol t toadult~in 9-15day s'at

,r-

12oC .,10-13 day sa;

18o~.

37- 40 days at

6

oC

,: ~nd

failed to molt

.-a t 22_24° c.,Adouble

pre-~~it

mol t

an~ a -P~:·aSitic.O;Olt w~re not~d.

Matinglas t edfor 12- 24.hcu rs ,~ndus u al lyoc c u r r ed withina.f~w hour:s of molting to adults.Egg;- wer e laid 36-.59daysafter: mating . Fe mal es laid600-.65 0 cggE! over 3 daya,'The in cu b ationperiodwas 35-55 day sat12°C'iheav iest'egghat~hingocc urred 4-11 day s-after

the ~Sl!t ,of

hatching.No eggswerelai d at 3°, 70,18°,or

_2~_24oC .

(7)

: .

.~.

l're-paradt~s11Ye. 2-3'~llY$.The period fTo.eUler~encefrol:lthehost•

.

t~'

egg

_laYi~'g wa~

45- 74"day s ,.and'fr-.emergence

t~

'e gg hat ch ing ,80-

",129 days•.Mixe d infections invol vin glII1~ro5por lda .me.t'1Illt bl d s and

.

·<:oe.101D)'c1dl~nS

. ver e noted.Sea s on al"fluCCu at l oJ. i·a

~f ho~'t/paraslte

- -..

populationsInHalfMoonllr~okan d LongPon'dTrlbutllt)'-v~restudied.

N~omesominmiB

[lrirle11O.liS'

l~fect.ed·

the'P1'osimul:l ums4«Octobe randthe

"'earlysUIlllll~r5'im1lUWllSinlat eMay-early .nme•Carryove ~'of mermit hld s

to-adu'its1.lllul11d s,ba sedon

ext~~p~la~ed

dat afrolll

la~

larval'and

. . ,

"pupal infections.'l~dlc:at.edtha tN.fl.~n~Ji8'pI:Ob~ablYcarri\edove r

totheProaimul iuma

.

andSimuliUmG

.

ata rateof .0 26 and .062%·

'resp e c tively ",Gastromema,viridisandI~ rd8a8n~i~,eTlBi8probabl y in f e c

7

',,3-26:8%of

~~\e1r

" "si mul U d

h05t~/ H'o5t, s~edf1c':Lty

.: .,:.-;'

appeareJ to berelat ed }othe

a8ynchrono~8

H.fe orc,lell' of thehos tl parasitepopula t i on s. Sexrat i os,..o f1.4:11lale s tofema les,were

- ..

-noted forthe .erait hid. frot!! thePl'Oaitm.cliUlllBin Haff MoonBroo k.

Fe_aleto'mal emena ith i drat ios of1.33:1 and 9,,:2 : 1for the.,

.ProBimu_~ium8an~.Simu liWllain Long.! ondTri~ut.ry.·snd2.7:1·fo~;he

. SiIllUZi~:

in

f1~lf

Hoon

Br~~k ~re

alsonoteil :In

1~72

. ".,

I .

IJ

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\

.

<:>•,<

TABLE OF CONTENTS

INTRODUCTION

HISTORICALREVIEW •• • METHODS AND M.\TERIALS • RESULTS'AND

DIS~USSION

Dis tributfon-••

LongitudinalSt reamDistribution .• Physical and ChemicalParameters.'.

.T~xono,my •. • • ••-•• BIO~OMICS,' • • • .":• .' • • Life Cycle••• ..-••

.

.

Mix ed'Infections• • • •

Host/ParasiteSeasonal Fl:Uct uat: 1ons Sex Ratio

S\JMMARY ., .

REFE~NCES'CITW' 'APPENDICES

. . -

,r-,

Page'

s

"

,

15 15

"

as

2B.

33 33 42 44 sa- 62 65 76

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LIST OJ",TABLES TABLE:

'I.

,

. ,

DISTRIBUTION BY DISTRICT AND BLACKFLY SPECIEs OFi' ..

.PARASITESRECOVERED FROMNEWfOUNDLAND I,N1~71

AND1972 • • •• • •' • • ,,.• ,', • ••• ,

Page.

• ,16

* 2'. COMPARISONOF MEASUREMENtS OF NEWFOUNDLAND .MICROSPORIDANS Wlt,H THOSE OF.'W::ISER(1961)

AND.

JAMNEACK ,(1976}, .:·"'--':";'.' -~'•,:",' :.,'• 3.

~.

5 .

. .

'

'DI STRI CT DISTRIBUTION OF STREAMS llAI\BOlJRING P.FUSCUM/MIXTVMMERMI1lf;IDSINNEWFOOND~•'." "• COMPARISON OF WATER ANALYSES BETWEEN INFECTED AND UNINFECTEDSTREAMS IN DISTRICT,6• • ,';•~'. '• • RANGE OF ENVIRONMENTAL PARAMETERS

F~R'

:STREAMS

~'i:TH

.

AND,WITHOUTBLACKFLY HERHITHIDS .; •• •t • • •

MIXED PARASITEINFECTIONS RECOVERED FROM NEWFOUNDLAND."

BLACKFLIES••• • •• •••• .'. •• ••• • • •' ••• 7. SEASONAL FLUCTUATIONS.OF HOST SIMULIID/N.FLUMENALI S

POPlJLATIONS IN HALF MOON BROOK_' .:• • • •'.'• • • ••

8. .SEASONALFLUCTUATIONS· OF HOST SIMtlLlID/N.FLUMENALIS,

POPlJLATIONS IN LONG POND.TRIBUTARY• •; .'.•,• 9 • ~ARRYOVERPOTENTIAl,OF MERMITHIDS FROM INFECTED'

PRE-IMAGINES TO ADULTSIMUL~ IDS"INNEWFOUJID~

... 1 55

·.f"

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Figure FollOWing Page

"

lao,Re gu l a r collectiond tl-a.t·HalfMo on Brook, • lb. Upstreamview ofHalfMoon Brook.from re g ui ar collection

site •,'.: '•• • •• " • ',-:•,-...;~ •••..• ~ 2a. Regularcollec.tionsiteat LongPondTr lbut'ar;• 2b,' Upst relllll'vi"ew"Of Long Pond Tribu taryabove

the regul ar co.llectioneft.e ,•• •,.' . '•• .

3. Menn~thidwo rein-the~i~UIUd,·s•.tub"el'Oaum. • 11 .4. Microspori daninfecting the. simu liid .P.mietum ••.• 11 '5. Coelomyci rii wn lJimuliiinf ec t ingcbe simull1d,S. venus tum • 11

, ,

6. 'Dis t ; ibuti o n of streams ininsularNe~foundland

harbouri~gmeniith idinf.ectedsi mulUds. ·• ,:-...' 7.~ Longitudina.l stre:=d111t ri butionofN·.'fl lllflenal i s

.infected'sillluli ids·. ,• •, ,• •,,'. •,••,,

8.-.: .

Comp~ri8 onofmini mum andmaximummonthlydept hs for the tworegu1!ircollection sitesin19~1and1972••• 9 •.'.Comparisonofmini mumandmaximummonth lycurrentsfor the tW(l';r~gula.rcol 1ectio.n?t e s iI!.1971and..~972"• 10. seomeeonermie [Zwne naZio.. Taxonomic figures

20

25'

26

26 29 'II . Neom138omermisftume l1llZis• .Taxon o mi c figures ~.' 30

12. Seasonal fluctuat ionsofN.flume naii epop ul a t i ons

in Half MoonBrook - ~ 36

13. Seasonalfl uc.tuati onsofN. flwnel1lltiepopulati on s

inLong P.on dTribut ary 36

14. 'N. j"'LUlIienaUe!mer ging fromS.ve'nustum.-/. 38

, , ' ,"

,~

J15. Pre-a dultmoLt inN,flJ.Q71e~U8. 40

16. Simu 1Hd age classes f=', 44

(11)

"- .

'

/-",'

.Fig~r~

I, .

Following_,pa ge'

17a Se!!sona.leex-recr ce inthe aimul:l.:I,dsfrom

,t:'n~PondT~ ibuta t"~.'in.19 72.,.'._•: • • •-. • 59

17b. Seas o n a l·sex rati os in theaimu1i:l.asfrom

Half MoonBroo kin1972: -. ••• ••• .' .• • 59.,

"

18a Se ason alchanges ,in _averagenumbers of si mul 1i d

me~ thidsin Long, Pond Tributary in 1972••'•.,,--." • 60. ISb 'Se,aBon~lch a n ges in,averagen~bersof's i lilul1'i d .:,-

mennit_h~d,~:l.n Half-MoonBrookin

!In. . '. . ', ; ",' . . ..

60 19 Stre8 11lloc.at:l.onsw:l.th1n the Pf ek e 'van ce Cr.ee/t.

'<- "'""Y

area • • • • • .'••• '. .'• :7.'

\

\

I

\

.... "(

",', ,V~ ::..

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ACKNOWLEDGEMENTS

'rhert bywi s hto express my.stnce r.estapp reciation and'thanka to the,fol lowingfo~itheir IDOBtinvaluab l e assis tance'inhdp~ngmake

. ... ~ ,

thisst udy.Po~Bible ~'~., ;,~) .~

' ToDr.Gordon F. Bennett,~~~supervisor , forsuggestlngthls"

toplc;forhi'senc ourageme n t and guidancethroughoutthe study lIpd for

" .<,., . ' "j ' - . , ..'

i

bis patience incor.!ecting~~4:'manu~~r1Pt.. " , '

I

To Dr.Roger,Gordo~ftorhis manY,h.elpfulsuggestions throughout the course ofth~sstudy.

To Dr. WilliamThrelfa~'lfo'rhis helpful·cr i tlc ism of.the: manus~r1pt.

To Dr.Willla~R.Nic.k1e,United'St~tesDepar t-mentpf Agric ulture ,Agric~l~\!ralResearch Ser:vice,Be1tsville,'Maryland,for hie vetificatlonof.~E;t1Ilit hidideO,t i fi c Bt ion and helpfu l8ugge~tions.

To Dr;Jarosll!-v."Welser, cz'echo'sloyakA-;;adelllY of Sciences ,'

.

.

Prague,Czechos1ovakia ,fo~identifying',sollle~fthe,mlcros~ori.da~an d fUng,alparasites.

I . .

\ '." ,~oHr.DaVIdJ. Le~is ~:r ~heeX:.~i.~~.Ho"':.Of.his.b~a;:~lY m.a~e.ria1fromthe[Bona'{1sta:,an(portionsoI theAval on.Penfn au.las," snd fo;':h1S m?st

· invj~tiable'

assista ncein'thefieid.· ·

~":

: ....

.r cDr."Pe~erScot t fo;his'asa'iatancewith·t heidenti ficat io n ofthe pl a ntllUi~erial.

To the In t.erLib raryLoanSt8f.f af MemorialUn1vel"1li~,

~ , ' . ., " ",'1'.

·pa·r t lcularlyxisse aMarja~leKi n Kan,(~nler~e·Ja ckson,fo( , th e ir persia t en ce in cbtafnrogmuchneedell~r~ference sinthl s study.

T~Mias Mar ionBrake.f ?r.~bta1ning·wate rBo/.'lp l ea'.inDecember 1972f~~mtheport':"aux-BaBqUe~'area.

&• .

' .

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\

To Hr;Roy Fickenfor hisaSEf~stanc~ w~tht~ephoto'gr aphic reproductions •

.To,Technic:al Ty.pingforthe:l'I'

pati~nce.

lind

petBeverenl:~~ ~n

. the final'pres'ept_at1o~,~fth is thead s,

:Iothe Prl:iv1nc1alGoyernmentof Newfoundland and Labrador.

" "

and~to,Memor i al University ofN~foundlandfor their directf1nanc~al assistance,

" "

~Tothe Defence.Reaea r e:h Boardof Cana,da(Grantn~~ber6801-47),

! . . .

who~.effnancia.l assistance-supporte,d.t bepurchslI'1of~qulprnent'And the

•fieldwork.

And.1'1n a lly-to'my wife,'nonte,for.aiding1~thetedi ous job.

o~.proof-re.adlng,and forher.un~er8'tandlng'and encouragementthro~gh- out this study. :

.»,

( ."

",

" :>f';S" -

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"

) .

,.':

INTRODUCTION'.

Tb~' Si~Ul1ida.e

are·a

c:ofi1lCl Pol1ta~!g.roup

of

.he1lla.to~haloU;S

.

'Di pt e rawhich:.,31'i~ultl ,.•;enotot="io~ pe8t~,'ofwan-bl~odedanima1~,'".

andvectors

oi·

avari~tJo,fpatho ge n i c: organ18 . ...In can a da , the•.

. .

.,peeies,as vecccre. ofpath o geni c:ulI.cocytozoqn

.

ep,to ga"mebirda (Fallia 'andBennett ~'.19'58;Ben~e,ttand·Fal~i~,..19 60;'i enne ttand,Hac:In'n es,19.72), ... y hi nder de v elopment·ofwaterfowl;and game bird managementprograms• .

Exc:ea~~'ve" bl~~

feedingbyde use

pOPulatio~~'

ofSimll.t£wn,az:cti cwninfluence "'

. . ' . \" '

c:~ttle~a1.S.3:~.in,par~8of Alberta andsa8k.t~hevan(Cam~ro~ ,19.22;,HUlar an~Rempel,.19 44; Re mpel and Amas on,1947';rredeen ,.1~58,1969) and 18~ well

~oclll:len .t:ed

'caseoffi n an ci al'loss

to .th~ ~adisn' livest'ock

indust ry.

. .ra othe rpartsofthewoEjd .s1mu!iids a.r esorediuc:dy

'...ilivo1vedwlt~IlllIflo as veet~r8of serious~~thogens•.Ince~ta.inregions of Afr ic.i,Cen tra'IandS~ut~AlDerica,severalsimu lUdspec::1esar e rupona ib"leforthe epreedofOnchocerca volVII. Zua,I fil.;rialnematode whichcauses adebili tating hUllllu ddaease,Onchocerc1as1~'(Lewis ; ,19.53 ;,Dalmat , 1955 ;'Duke,1971),freq uen tl.>'Ieadfng't oblindness.' . Fro m at!ec~nomicand medica l'viewpoint!l.lackfliesrequi re

~

. . . '

atr1n~~.ntcon t.r~easures.Blackflycontr~lhasbe en.achie,,:~d.i nsome

loca~~1~,es '(Ga~~nd'HcMahOn~

1954{,notablywiththe use of

D~. ·

.-::....:.-;,:. -L

-'<'/:'"7:: '

(15)

.. -:».

' .\

'\ ,

.(

. .

,

"

ManyD~~~nsect1ctde8bavebee n"'developedtosti pp lant.DDT,but"all . .

~a~e

l,nherentdi f fi cu l tie s"I I'

.~~~t

1)th ;r

PO~'lut~

't h e

~eDvlrotlllle~ti

.:

;21,

the

~arget bl~c:kfly popubt:~on

maydev elop re s btanc e

~~

the

.pe·stl d deIn us e; 3) tbe y aay"elll11na tenOD-ta~~et,organ15l11S.

In~iudlng would~be

preda.to n ,wh i ch

~elp 'natu'rallY~ O_ cootro'i-blaC~~Y

~~~& .

.

~ .

:

.~lthsuc:.hpo~ent lalprob l(Nlls,nlter l\4tc!<:.oDtrolmeasu resshoul d beinvestigat ed an dperhaps one.,of-t hemO.8 t promt.liIlogme t hodsb~ugh .theuse-ofblo1o~lcalcon tr ol:'Bl~iogl~81co~t~olms ybe defin ed 8Sthe . reduction

In"'.~umbers

of a

pe:t 8pe~les

bymean s'of

anot~e[ Organ~8m" :

(t h at'hasbeen in someway mana gedor

Int.er;ere~ ~il~b b~, ~anj ~o .J/.

lev e'l.

'a,t':iWhich;:t:ll e pe! t

" IiIP~Cl~1

ceasestobe

~\i. econoDli~.' lIIe4i~al o~

vete rinary probl em(!,8kew.•1971~,

Biologica l

...

contro l,

.

using: mass rea r ed' organisms. has" be en. dealOns trated'tobean effe ctive_"~h~of'contro llingSOliepeatinse'ct Pop u lati ons,Bacil-Z.U8popiUieu:thltky'provi ded"t h e hratreal

,J

, " , ', ., " .

. "

encourage~nt{or'th lUlil of ldc rob l,u,orga nilDls.1,bio l ogi c a'lcon trol

'\ . " ,

, . , .

'J apa nesepe et lellr u ba inHar)'l an dfor at lu st nine yeari post

::-;.- . I

:app l ication (Corya~d LangfOrd~.19S5),~'NUClearpolyhedros ll'vi? s es

(NPV!J)hav,e

als~'"~d1ucc,e8~fullY: ~n

con trolli n g'

~opul~,t10na

of

'-':_'. 'J'roichoP:l.Uf.ia

~i (Hubne~) , th~

ca bba geloo p er

o~

Brassica,crepe

th;O!:lghou'tNorthAmet:i~a(fetzi n.196~),.p.oln~r(1971) revlewe~the ,fieldtrla!~involvingNBoaplect~

;raseroi

'an d the'ODlJ6':s't r a l n'o f

.N•.,';:'~cap8~;it wa ademons tratedt~tthe l ene.~t,od~scoul d'

..'

~ara81t:ize

I'awide.

var~et~

of pest" ,inse c ta wi th

aig~iicant r~duct'ion:

.

In pe s tnualbersoc cu r ring"

" ,

(16)

" . '.

,,.' -

1 .,

( "".

Many~ra61te6'and.pre~ator6'o fblackfl~~shave bee nreported ._"...,....,..k:i.tl.t heworldHterature (J'e nk in s, 1964 ) , bu.tnone havebe~e;1,'tnaS6re a r ed

>:-~ ..';..;,... . ' • ; .. .- ",".

,~'!'-"inthe laboratoryfor use as biologicalcont rolagen ta ,'s eve r a t...

...

f'0\~

..

para:altlc'·"gro~ps.

. 'lncfud l n g" , .

~Ungi

.

a~'d

,

prot~zoa. hav~

'c ontrof'

P~:~lblllt'i~S

~ . .

, ,~:'t'"ye:t;.hap~~;~ ca~didat~

gro up.

wi~ .

thegr ea\ e s t:pot,el\tial

i~

the. :.

: ::?

-,~etna~Odefam!lY;'.Menll1thld,!e (Wel~~,,1~6~b"1964,1.965;We~ch~~d Poflla~-i';

1965;Weiaer,1963;Poinar,1971; Gofdonet at:.I 1973),

"

.

" .

'The

M~~1thldae Br~ t~get~er w1t~

the

Tetradonematida~

Cobb'

<fti!a~ha.Sm1dhnn~matdde;.of.~h~.Supe r f amU y' Mermlthoidea'Wu1ker.

'Mermith1d sare' filifo rw~orins',attdiningle ngths of

up

to·'50 ems ,that

.f'- . " n ,. ' .. •.,~•.;.,

are parasites.as juvenileso! lnverteb~ates, l~~~U~ifig:-l~ehes.. api der s , .

~Oll~S~.~, .e.rus t .acean~ .

and other;emato(tea,.bu't are"prillla1"lly

co~f1ned '

to about"15 order's 0'£aquatic~ndti'rrestrilil"intnsecra.(Welch ,1962bj0

I " i ·. · . . "

Hermithids.areuniq ueamong neeatc deain.tha tt~eesophagus,a non-"

.

~ . "

muscular,cut ic ularizedtube , often runs posteriorlyfor overhal fthe

~OdY.len'gth.but,doesnotent.er the.inte8~:1~e ,althoughadhering toi t.. .The

i~tes~ine ,

is

~iso peculi~r

in thatit lacks·..an'anus and lumenand

1; h1ghlymod~f).ed'~oact a;a food st orageorgan (t rophOSOllle)for the free-l1v.ingad,ults which probablydonot'feed.(BayUs,1944 ;Rubtiaov , 1965,

.i9 6,6d.:

". Go~don

.et

~l.~

, •.1973). --,..,; "

.4lthoughstudi e s by l'ickavance·et

aJ..•

(1970).Frost_(1970).

Fr 08t :ndo

~nie"r

(;971),'Bradbur:y (1972)

a~~ Fr~8t a~d

NOlan.'Ci972)

t ," .

de fi ned~he.0 ~ CUrI"enCeof cer ta insi mul l1 d species andtheir micro- sp~ri1ianorfungal ••~ymbiontsitl.Ne~foundland , ~o'thingwaskn~

.i::oncenhngthe prevalence'a nd d1stril!:ution ofth e meI1llith:l.dparasites '. :0 £ b1ac'sfl iesonth e

'i~lar\(jo

ThiS

, at\id~."

ther e f or e,wa s in itiatedto

. . ...

./

I~~~~;;

.

-"

(17)

.'i-~.

:' - ~;'.

det ermine

N1e" s'p~~1'~S

ofmerm.1t hids'prese nt'.i n"bll\ckfl1 es•.

th~iX:

distribut:ioo''"~

.'

in,'insul a r

Newf~~~dl~'[\d,'as':well . '

as'

.

longitud'inal

' ~tream . .

d:C$tri bution , season~l'flu~tuationsofhostlpar~s1te',populat i ons ,'the

.b;·~.oi~~~.:

'.

,a~.d.:eco·l~g.ica1,,~a~t?rs ~ffec~i~; ~ar~~iti8~. ~~~ t~~ '!.~~'~ors.

af~ect~rl;',:i~iig; ~t1ng~gg,,la Yi ng~ndeg~:ha~c~in~.-~n.additio~!·

miscellaneous data on.the distributionof fungal andrnicros pori dan.

~

.-.

~ , ,

.

- '. :. : .'

.

par~s1tesrecovereddur~ng" the'blackflYcOl,l.ec,tton.s vereabo coll at ed .

,.~..••Three',sp~cies:of·bl'a c kf lies , theProsimulium'[us ewn/mixt Um.

'syme:and

DaVi~.~·"COlllple~~ ~'nd simuti~ ~enu8tUm s~y.

.. both

or Wh~~~:'are

seriousllIan biters andcommon pests'du ri n g theear lyspr i n g and,summe.r

.. '

.

.

... .

mon t hsin Newfoundl an d, were,lIIostfrequ e ntLy 'enc oun te re dandpara~ it ize~, .and

~r~vided

the'basis fo r this

~tl1dY,.

'The

'~~rmithld

Neome:omermis .f1.umena1.io" occur red in 99%of:indivi dua l infectionsan d th iS st udy fd

~r1marl1yb~sedon't h is~pecies.

,';'

\ .

.

~

.

' /

(18)

/

HISTORICALREVIEW'

~efirstrefere n cetoabl a ck fl yUletllli~hid ,~~ras itewa sth a t of Siebo ld(1848),whofound aMennispara s i t e'.inSirgu~~um,r'epum ein, G~rmany,Diesing(185~ )ll-aming this'wormMomie simuZiael'eptantis.I ,Uns e ow~lB89)r-e ctrr de dMennis crasea,.also fro m,S.l'eptans,but La ter

su~ges ted

(Linsfsw ,1,898) that this specieswas

sy~~nymous

wit hM.

simuUae~eptal1tie.,.M1l1·1e r (1931) described another species,Mennie

·si muZi aefrom alar v alSimUUqsp.,from the Alps. ~e8cr1ption8ofall t.he s e species are Incompletean d'for any futuretaxonQIDic work are to be~onsideredaanomina dubia(We~ch,1964).,

Thei~ent;f1catio nof-Me~ithidaeisdependent principa llyon adult characters ,and sincethema j ori t y of simuliid worke r s only

,

.

reco~eredtheparasit icjuvenilestages,specific determinationswe~e notgen~ta11ymade. Hence"wo r k ers recc rded thcpr c s cn c e ofMermis, often a repositoryfor questionablemermithida(Strickland ,1911,1913 ;

-W'b.,t~i ,

1914,M.ll.,h. 1914,Edworde , 1921,

.

O'Kane,-1926,

B'~"-";t.

,~

.1934;Twi nn,1936, 1939; Smart,.1944;Crisp,1956~•..0th~Two.r~c:~s..

assignedmerlllithids to diffe r entgenera,Agamomenm..~(Lutz , 190 9), Limnomsmis(Sommermanst

az.

J 1955; Ab.de 1nu r f, 1968) ..llydromsl'I'Ilis Ande rsonand Dicke,1960; Fred een andSheman~huk,1960;_Pe te r s on,1960 ), and

?arame~8 (M~Comb

and

Bic~l~y,

195;) .

Welch (1962a)."!asthe firstworke r to describethre e ';"al1 d b La ckfLyme rml t h i d species. Thespe cie sbelonge dtothree generaarid weredeJ-ign~tedasGastromermis viridi.s,Isomernrts wiseDnsinensis.,and N~eomenni~j7..umenaZis.Phe1ps~and'DeFol la r t (1964; re de sc r i b e d the s e '

(19)

)

6

~hree ~pedes.whileNickl-e.·(1,972)reasB~gnei'M(J8omermi~fluirJen;:;,is to a new genusNeomesomerwrisflumenalis":The genericchange was .,necessitated.e Ince~he

""?"

ge~er,ic'd,e s c r i p ti onforneeonemteW8.S

basically incompleteandcouldfi t'several other common mennithid"

.gcnc r~..JNickle(op;eit.~)alsoreflg~~edG".viridis:

'R,ubtaov(1965)'d e s c r i b e d 1n detail r.heanatomy'ofseveral genera of.juvenilebla~k'flymermith~~8'~dproposedthe adoptionof various larval

chara~ters--the

eurccccre of the

·intea~inJ

••lon git ulfinal

~ords .

esophagusand-: BBo c l a t e dgiant

cells--a~

valid criteria

f~r

the

ide.neification of .mennlthld spectee, These characterswere~statedto be ecreconstant

than

the.-a ,fu lecharac'cers,and permittedquicker and ,mote reliable'idimtif1cationof-species which would~ot'no rma lly be

\

recognr eed in the adult stage. This paperill the basis for much of his later work.

Welch and Rubtsov· .(1965)described a new species of caetromermie plus six varieties. basedmai~lyonanatomical .differencesin the free- - living poat-parasiticjuvenile stages.

.

.

Rubtsov (1966a) described a nev spedes ofMesomennis.whUe Rubtso.,v(l966h,C,1967a,b , 1968)and RubtBOV and Doby (1970)deSC(jed new speciesandsub~peciel:or varietiesfrom the generaMermithonem~

. '

.

Apl'Oqtonema~ Te tradom~rmis~'Me~ome1'f!1i8~Limnome1'fl1ia,Gaat romemri s, Iaomermis andllyd:l'omermi s .

Simul1id mermithids arewidely dlstrlb.utedandhave been recorded'from: Nearctic- -Webo t er(1914).Malloch (1914).alKane (1926), Ja me o (1950). Davies (1958), 'Wolfe and-Peterson(1959),Fredeenand Shemanchuk'(1960). Pe t e r oo n'andDavieo (1960),Abdelnur(1968) and Fr edee n

(20)

(1969);P.al~arc tic"';-Edwar d s(1921),Smart (1934,19 44) , Grenier(19 4 3) , Rubt s ov(1950, 1963,196 4 1 , Carlsson (196 2) andWeiser (1963,196 4); Ethiopiari-Lewis(195.1, 1958 , 1965),Crosskcy(1954),Crisp (1956) and Carlsson(1968,19 70) . and Neotr·opiCal.re,giOnS--Bequllert(1934), Dalmat (1955 )and Gamham al)d Lewis (1959). To date these merrnithidshav~not been reportedfrom either the Orien tal'orA~stralianregions. There vi ews of Welch (1964) andGordo~et: al.~ (1973)contai n mor e detaileddiscussions of distribution .

Thelif e cycle of simulii dmermithidahaa beendetailedonly fo?

G.viT'idi~andI.vieaoneineneie (PhelpsandDeFol:art ,196 ~ ) ,but is ol.a t ed observations~notherblackfl ymermithidswere p rovdded by Strickland

~911 ),-Ande rsonandDicke(1~60 ) , Peter~on(1960)" Andersonand DeFoliart (19 62),andWelc h and Rubtsov (1965).

Strickland (1911 ) . thefirstworkerto detail thepathological effects of's i mulii dMermis Lnfec t.Lc n e,rec o rd edtha'ttpupuLhis ~oblaa ts failed to develop,therebypr even tingpupa tion. In latela r v al life,fat bodies'we r e observedto be muchred uc e d or abaent; , and infect edlarvae tended to be~uch_longe~thanuninfec tedindividuals: Othr:'.~worke r s have si nceCOrroboratedthes ef1nd ~ ng s ~ndersorrand Dicke,.1960;Pe te rson, 1960;Ander s o n and DeFoliart, 196 2).

Wu(1930),Grunin(1949):Ho cki ng and Uckering (1954),Davie.s (1958) , Pet e rson(1960), and.Phef pa~ndDeFoliart (196 4 ) , repc rted tnermithids from pupa eand/o radults'of vario ussimul1id species. Hoc kin g and Picke ring (1954) re cord e d that

f-

uenuetum adult females in fe ct e dwit h mermithids.we r e devoidof the alimentarycanal,fat bodie s , reprod u c t i ve and nervoussystems withinthe abdomen. Pet erson(1960).

(21)

"' ,," r'

.and Phelps andDeFOl1~rt. ~1~~.4).r~.pcrtedsimi,larfindln.~s. ..

.p.!lrasit.lze d ,,a dult . s ilnllliidsareinvariab lY,sterile(Lewisand Iba n ez

. - "

de Aldecoa , 196 2; Shir.~:~.ina:1963;Leaerre., 1966 ) . Externa:l indications

\f

simu111dmermithidparas itismareev ddenced by o;:l~w abnormalities'(Peterson ,~960). g~nandromorphism(Edwards ..,19 31 ).snd

.

.

,

intc~scxfonnntion(Rubtsov,1958; Fredeen , 1970 ).

Since'me~ithidinfectionsin"bla ckfly populationsarealways f~ta l[0"t hehostitisect,the per cen t a ge ofinfection isanindicat~n;.

alb eIt an appn ox 'lmat e one, ot'mor t a l i t y incurred •.Welch(1964)and

. .

Gordonet al.,(1973) rev i ewedmuch of th e liter a tur eon.inf e ct ).on

)

.

.

percent ages

.

inwildsimuliids, a nd.co.ncluded tha t rnermithids

. .

have defi ni t e~otenti a ~'~sccne refegenreofblackflies. Other wo r ke r s

" ~~t\el;S

and Deiol1.art.,1964;,Welch

a~d, p~ina~. 1965~

were

al~~

of ' "

I

same opin i on. , " '.• ,

' ~.

(22)

..~

',~

-

. " " /

METHODSANDMATERIALS

i I '

Apr elimi n ary surveyof ro a ds id eetreeee, selected for ease of

access, occur ring0'1)the'Avalon Peninsulavaa init i a t edin Apri l19 7 1 ecLc ca rebLackf Iy'populat i onswi t h a high levelofmertlli~hidpara si t is' m•

Fouratreams were origina ll ysel ec t edforcontin~~~gstudy,butthe numb e r wasLacer reduced.(la t e io 19 11) totwo_st~esms'as adeq ua te qcvet-age of allfourst rep-1llS was notfeas ib le.

The two streams selectedfordetailedinvest i ga t i onwe.rs both ofth eyoung str e a mclass (Andersonand'Di cke,1960). The fir:t,Half

"HoonBroo k , wa s aroads i destr eam cross ingthe highway"abou t"2

- .

"

.kilomet,ersNNWof,th~N entrance to thevillage ofFl~tRock, '(l N/1 0G'6998 74j' ,1 Ha lf MoonBro,~korig1natea at~alfMoon-Pcnd and~"

flow-s2.8 kilOJilete rsbefo reempty i nginto the sea.

The streamand collectionsi t e, (Fi g.la, b), were chauac- teei aed by an eme rgentflora consisting la r gely ofMyriaacai e-L.,"

Ftcea mariana (Hi ll.)BSP.,Alnus rugosa"(DuRoi) Spreng ,Junt:Us

•ettueue

t:,

andChamaedaphne caZyauZata(L.)Moe nch'.,whilethe

"

s ubmer geot flora wa s dominated,byPotamogetoi1"ameriaanuec. an d

s.,

SparganiWTl sp.sridmosses.

, Thesecon detre ae cross e.d thehighway3kilometersN'ofthe villa geof-'Bay'Bull:s,(IN/7 63345 4),and dra in e da distanc eof300 M.

f,om a mars h"int~LongPon d. The collec tionsite,(Fig.2s , b) was chara cterizedby"n domi nanteD,K'rgent.~loraofMyricaet:l e L.,

lCana dianNationalTopogr a phicse'ftea.

(23)

" ..

Figure la.

Regularcol lec tion.site atHalfMoon,Brook"•

.

~.

'

.•..

F.igurelb".

Upstreamviewof Hal f Moon'Brcb k fromre gular'colle ction site •.Note the he avy shading by shrubbery.

(24)
(25)

:J

" j

!

'"\. F1gure2a•

•" \ . .ie • • •

'''''~

" " . . - . ; . . ". . . .

\

t

I

UpstreBlll viewof Long Pond Tribu tary above.the regular collection,s i t e.

(26)
(27)

" .~/.

.., ...,10 'I CaPe;;

ro8t~ta

Stokes,-J w1cUBeirusua'Lol':1mixtu re

~f gra~e~ and~I ,_

\lI0SGCS. and a subeeegentflora; ofJuncus bu.fonius L.

Mcnn1thids or otherbl~ckflyparas ite,a we r e collected~tle4~~"

,..~ once frommany otheraee ee e localit1esInNcwfou!l.d.land endthese are

"detailed.hi'Appendices.1-4.

Slmuliids wct:,ecol,leeredon aw~_l:!,Ic ~YbasiB from Hayto July and biweeklyduring August;'Ind.t hr oughthewi nt er monthsfrOmHalf

. ,Mo on Br;;okand Long Pond Tr'lbutllry. ,Col l ec tions weremade1~theIlIOrning

and wh erepossib le..attempts werema d eto collecton thesa.meda y atth e

;"eeklyor 'biweekly int;rvaL

A

udnimum sampleof l~Ola r v a ea~d/orpupae were1:I8u811y

collect~d~

The temperature (DC.) anddepth (in em,s.)we r e

-

recorded enea ch collection day for ea ch a i t e . The.re t.e of water flow

. f,-- . ~ . 'I

andthewate rconten tanalyses were conductedbiweeklyin1971 and

> " " "

monthlyin 1972: The velocityof th e current wa sd~termineJ"withan' Ott,

~mall

Cur r ent

Meter~. w~ile

the'waterana lyses

o(.~.1i7cblo;id~S,

silica.sulfides,nit rat,es.phosphates ,dissolvedoxyge~.~amrt'lclni a ,'.

c~rbondioxideand totalhardnes~ w~reconducted in thefielddur in g th e sumser ecn the usinga....LaMotte water.~alysisKit. 3Durin g thefall. and~in.ter,watersamples were brought backto the laboratoryin• sealed. 250 ml ,jars~filledto overhowi ng,andanalysed.1n1':.4 hours afte rcol lectio n;

.

Simul1i d larvae andpup a ewe re collecte drand~lIl1y1n the field

,by scraping th em fromrocksand vegeta t i on in to10 em. diamet er Pe t ri

dishes lined'wi t h saturated(c~veredwith a film'ofwater) filterpape r.

2A•Ott. Kempten-Allgaeu! HydraulicLaboratory•

~L4Motte

.ch e mical

Prod~cts

Company, cneseeeecvn,Maryl a nd

(28)

---

~.

~.-;-:...

" . .

"','.,

11

this_t hod.

-c-:

,-;....-.. andeaea r aofth e.coel Olll1ceee ce a t e~de. Smearsvere then airdr le~,

fixed in absol;te ethy l al coho l for20sec.and stainedwith a1:9

.·· :. ,.··dilu ti on_ofGtems~· 1.stain for,.15.minutes. Microspori dsnlind fungal

par asites wereidentiUed using the read tllof.Debalsieux(1920),

"'''.~

Thefdlsh~erecarr ie'dinround,stalnles ~eeee r,Petri dis h8~orage

c::ontaine~s'(capa ci tyIS dishn) ,10 whichke p t. the siia.liliid sllOderately

-,.

.

.cool.mi:I.,,'outofd~rectsunligh t. Surv iva l rat e was exee f .tec e~slng

»>:

:All larv.lle~ndpupal!!were examine d inthela borat orywi th in 1- 4 hours followi ng collec tion ,and t.he species deteI8 1 ~~dusing keys by Sto ne andJamrW8ck"(1 955).Da vi e set·'at..~ (1962).and Wood"et.aI.•

(~96]).

. Parasiti:l;edlarvae'wet;e.usual lyre?Ogn l U ~bythe presenceof .mermlthidnelll:lt odcacoiledin'the abdo men (Fig.J). large wn i teorre d

cheesete xt ured cysts (1ll1crospo r lda) (FiS:4),o~ama ll,white, ro und

• sporangiaof CoeZomyci.diWIIaimu.l.ii8cattere<llnthe coelomicfluidof

> - . . , ' .

th e body cay-ity.(Fig.5). Non- pa r a s.ithe dlarvse vere se pa r a t ed frOll para sitizedIndivi dual a andatored.i~-95%ethyl.dcoho lfor future

~. ...

. . . ..

dis s ection and closerex_tna t ionfor,paras i t espossi bl yove r looke d duringth e

.. .

in it talaeparation•

WheRaicros po rl danand fungal infe cti on swererec ogniz ed ,the abdcee ueof liVi nginfectedhrvaewerepi~rcedwithj~el~er 'sforceps

Weiser (1961), Rubtsov(1969),and Jsmn1;lac:k(1970 ). Someof the

'm1cros pprldan andCoe]..orrrycidi Ullliden tifica tionswereconfirme d by

.'."."

,,;-"';"?

r: _ _ -- . - ~- .

-H.-s.-, .-entandCo.t.ea.,

To~t.~ , Oni:ar1.o~·

.'}

.

(29)

~

-

.

.

.

'\

..

..

~.

' ~0~~

~ ..

f;-.

(30)
(31)

, .

Mi~roapOr1da:n" ·infe~tfng·the 8:t:mui1id~·P.'..',r;~tU'n

.

., .. (Linearm.agnif1.cat10~_,X10.0~·:..

(32)
(33)

, ,: ~ ; " .' : :, ' ','. F ig".

'5.

: j ' ' : ' , , .

' ~oeZOlliycidiW1l

simu.z-t.i"in f ecti ng th'

~~mUl1id.

,S."

ven'~Btum.

. .(L.inear,magrtif1catlonX 12.0) .,. . .

(34)
(35)

12

Dr.J.Weisel:.

Living firl!'tendse co nd,in s tar.aimu 111ds were equnehedin

,

'

Ringer' 8PhySiol ogicafIn s ect Salineforexaminationfor slPa lltnetlllithld parasitesnot.visiblein,th e inta ct"la rvawhe nviewedund e r the c\isaectingmicros cop e. Parasites froll!fixe dU!ste r i a l were mountedin

, "

Rubin'sflu i d(Rubin,1951) forex amina t i o n andi~entificati'on.

Livingdmuliidlarvae parasitized byme r mi t h i d s:','ie r eplace~in 5001Il1~graduated cyl1t:'dersfilled.with'st~amwater•.A wa tercurre:nt

.

~.

of0.3 -0.5 fe e t/s ec.wliscre a ted by pa s ai.nga st r e amof fine air. bubbles,de rf ved from aqu

,

ariaair pumps,throughthe water., The st ream' wateerepparenc.Iy contained su ff ici en t nutrients.to mainta ingrow~h'of.

.

.

silllull1dsfor'pe ri od s, ofupto on e week,and no foo d additivessuch as yeastor algae~ereeuppfted, Forextended rearing'periodsofup ~o onemonth ,100-200ml',of water were,removed from't~e.r e a r tngcYli~ders weekly and replacedwith'f re s hwate~ w~ichpresumablyaugmentedthe 'f ooi supply. Thespr i ng~ndsUlIlID.ercollectedsim uliidswere rear edat

twote mpe r'a t u r e s (1 2°,and l8° e.)',while therearing of species colle cted durfpg th e winter month swas conducredat.telllp~rsturesequal to those

~xistingin the,field at thetim"eof coll e c t ion,

Post-parasitic'mermithids were sexed(onth e baeds ofsize).· sepa rated'and placediIj\St e n d e rdishes~'~~.:.2;5ee , diame,te r) filled wit h wa t e r; to a dept,hof about1 ee, Stream waterfro~'the co ll e c t ion s,ites was regu larlyused b,ut tsp and distilledwaterwe realsout il iz e d without noticeably increasingmortality. Several substratesce reused duringth~cou rseof'the study. The lIIostsatisfac t orysubstrate.f or obaervll~lonof the~rmB(al thoughnotfor oviposition) was found to

. '

(36)

13

. , 5 ' .

be-15 mm. x 5~.atdps of gcet.c hbrtte scouringpads, i~itill.l lyboiled in three changes of water for fifteen llIin uteseachlind placed in the Stender dishes.Them~nlithids(lipto~Operdish) actively moved

. . . . . . J

above, under andthrough th e aubatra teand rIOlted withinit.Sand substrates.about 5 mm. thick,similarto'thoseus e d , byMuspratt (19~7).

Phelps and,DeFol1ll.rt-( 196 4 ) and Petersenand Willis (1972),wereused in bne experiment onoviposition. ,

Phelpsand.DeFoliart (1994) andGo~don(peracne L c01lllllunic.ation) found "thatfungalii?fections increased themOr t ali tyrate'of the,i r mermithids,but in this study fungi werenota problem and-no stepa

SUl;~

ee

cont~~r s ter~liz~tion

ordaily water

~hangell

werefoilowed.

Fungalin f ec ti ons did occur1? iso1ated Lne tancea 1nthis s tudy,but, oni y when theWOl1llS were reared at te mpe r at ur e sh~gherthan laoe.or . whenthe wonns.wer e obviouslydamaged or in1ured .

Adult WOl1llS were'rela'xedfor".ide~tificationby gently heating unti l extended.for 4-6 secondS, a drop of water onaglass Slide.

-The specimens were

the~

fixe;

-s-: (c~urtney

etoJ ... 1955). ".

-I dent if ication,was made followi.ngthe scheme ofWelch (1962a) and Nic~le'

(19-12);the latterconf Lrmed the identiUcation of·me rnd t h t da in the study;Preparasiticjuvenilesvereidea cr fbed from materiallIIOunt ed,.in , Rubin 's'f~uid"Relaxation of preparaiHtesby gentle.heating was ' attemp te d,but~est.re eur te were obtai ned by directlyplacingthem into

,

.

Rubi n ' s fluid.Eggswe~edescribed frolllfreti!hmater~alheld 1nvaee r

5Minneso[&Mining and Manufac t uringCompany.

.

. .

(37)

··,i

"

Measurements'

~f p~_ep8raS1tes

wer;

m~de

en a CarlZeissMic.ro-

~

.yideomat6 at a 3200]1:

lI1ag~1fic.at:ibil. ~Bennet~

and

CamPb~~l

,<1972)

providedecalls 00.the opersti.oo"Of.theMicro-Videomatfor,the measu r ementof length andwi d thof bi r dblood parasites;th i s procedure was fol lowed,cl o sely-.

Al ldrawings weremadewi t h the aid-ofa cameralucida,attach- mentfor "theCarlZeiss ResearchMicroscope. Photographs

,

weretaken

on

a Carl ZeissTeSBovar•.

60b erkochen, We s t

G~rmanY.

"i

(38)

RESULTS ANDDISCUSSION

-.'Distribution

TWo_

au~veys'

for•.•biackfly endoparas ites

we.~,~'Cdii'd~~'~~~

1n

N~Wfoundlf~.~ {hi971

and,1972,

Fi~.t.~::,~,ight: "~·~reams

on tbe Avalon Peninsu~~.wn'hina 40kilo~.eter'·r~diusof se.Jobn'a wereexa.mi~~din thef1rs't''Su'tveY''~fiil'~an7ddi U onal

1 3'

streams wereexamined in.19.12 to furtherdelimit.thedistribu t ionof blackflyendopar.,!S1t~son.the

; - '. .

.

.Aval on."The·second survey was concerned primari lywi t bdetermining the'-

dis~ribution

of bla ckf'iy

p~~asltes .

aCr OS9.

i~sular

Newfoundland,. and th e r ef o re;atran~,-iSlandsurveywas conduc tedalongthe.Trans, CanadaHighway(TCH)·frOlll Wbitbourne(SO kilometersW.~f"se.John's) toPort-aux-Baaque·s . and~heGreatNorthern Peninsu la as far~orthas Bellburn's ;·l Ot.ee eeeesweresampleddu rfng Hay 23 to June8,1972•

",",. Theen dof May-earlyJune.samplin g perd.cd was selected as data' collected in1971indicate~thatme rmi t h i d a vereprescntin the, simuliidhostsduringthistilllCof year. Bla ckflieswere~la o collected fromi3additi~nalatr~ams.cn the'Avalon and Bonaviata

.

.

Yen i n s ula s during1970-1972 (Lewis). A total of19Bst r ea ms, f!OIIIa~l sour c esIwereexami n ed. (Fig ,6).

. .

.Tab le I,details_on a districtbasis ,th e dist ributionof mermithid, mi crospo r i dsn and flingal parasitesof the blackfliesP.

. .

fusc um/mi:ct umS)'Il1e andDavie~.C.mu.tata'Malloc~,.-S,corbisTwinn,S.

Zatip e s·(Mei g en) .s.tu.bero aum(Lundstrom ),S. venustamSay,andS•.'

. .

.

vitt a t iulrZe t t ers.t edt. ~istric t1consists of.ch eAvalon Penins ula,E~

of Whi ~bourne ,'Districts'2-6comp.risetheTCH frolll Whitbourne to

(39)

;-

TABLE1

DISTRIBUTIONBY DI$TRI CTANDBL.\ CKFLYSPECIES OF PAlU\SITES RECOVEREDFROM NEWFOUNDLAND IN".1971AND-1972

I

Total Bla ck fl y . To t al Tota l Infe ctions IBlac kflySpeci es· -'.

DtBt ri ct ·Str e p.ms Species Presence

Examined in-St re amsMe mi thi ds MicrosporidaCoel omyC!i diwn simul.ii

1.Aval onPenins ul a,E.

r-

[ueaumlmixtum 33 12 8

-

of WIlltbourne 78 C.mutata 11

-

1

-

S.corMs in 1 1 1

, S.l.atipes 18 5 5

-

.

,

S.S.tuberoeumvenUll t~ 5.23' i

-

161

-

7

S.vittatum 15 1 3 1

2.Whi tbo urneto Ter ra

..

P.fus cwn/miztum 18 .5 3

-

NovaPark S.carb'is" 4

- -

1

'He adquarter sAccess 25 S.tuberosUT11 io

-

1

-

S.venustum.> 12

,

4 1

S.•vittatum 3

-

1

-

3.Te; ra Nova Park. to

. Ip.

[ue""""mix","

.-/', ,

1

-

East of Bishop ' s

Fa lls . -11 . '

4.Bishpp's Falls.rc P.j"uscum/miztum 19 9

., , ,

BirchyLakeNarrows 19

:;:

(40)

TABLE 1(CONTI NUED) .

Tot al Blaclt fl y Tot a l Tota l In fe ctt"".,..IBlackf1ySpecies.

Di8 tri ct St re a ms Specie s Pres en ce

MermithidsMit;rosporida

I

CoeZomy rJidium

Examined in Streams

simuUi

5.BirchyL~KeNarrows P;fusCWIl/mixtwn 16 5 1 1<

to Stephenville ~':"

Cros sin gAccess 16

6.Stephenville P.fus:unz!miJ:tum 17 2 J

-

Cross ingee'

Fort -aux-Rasq u es 17

7~.Great Northern P.fuscum/mixtwrl l~" J 1 1

Peninsu1.a,Northof Deer Lake /

" ...

8. Bonavista Peninsula 17 P.fuscum/mktum .4 1

- -

I

Tot a l s 198 68(59) 52(40) 15(14)

*Th~to~a1number

,;c:..

tr~alllShar b o u r ingmeIlll-ithida,.!l1crospor ida orCr:Jelomycidiwn simuliii..

reeethanthetotal par a s ite'in fe c t i ons indicated . si n c e aome of the,atresl'IIS har b ou r e dmor e thanone infe ct ed blackflyspecies. Ac:tualstream tot als for each"paraSite group in bracke ts.

</

:;

(41)

. . .

,Port~aur-B!l8ques.deli mi tedatappro xima te 160kllolIletar InteN sls . District s7an d8 are of theGreatNorthernandBon aviBt a Penl nau1s s res pectivel y.

. Twelve'othersped es,ofblackfly ,fr o m thr e egenera , recovered

I

in thi s

8Urv~y

we re

e~1IIined

fo r thei rendop a-r ss.l t e s during,

1~71 "

_

and 1972.butwer~uni rtf ected. .~espec ies.are. as fo llows: P.p~eul'llle Malloch;C.dacotena{sDyar andShan non ;S.atireum.Fries ; S.c't'(J:~toni

I

Nichol s on andMi.ck e l; S.eury adni ni cu l WIIDavi es ;S. ezaieumDavi e s.

Peters on andWoo d; S.gouZdingiStone;S. tongistyZ.p..tum(1) Shewe ll;

'? "

p14getenseDyarandSha nnon ;S.quebec~n8e Twinri'; S.rivuz..t (1) Twinn,

and'B.tJerecu,rniwnSto ne andJamnhack.

Micr osporidans rec~vered1I"8; eThelo hci,IVZbr~cteataStri ck l an d ; T. fibrataStr ic; k land ;FlieteophorasimuUiLutzandSple.n d o re ; Caudoapora.simu.ZiiWeis er ,an dC.breuiooudaJ8lllI'1back. The fungal-

•parasiteCo~tomItaidiwne.i~tii·Deba feLeuxwas also,"fou nd. The numbe r' of blsckfl1e se';"Wnined andinf~';tedwith mierospor l dansor the fun g us aredeta i led 1~Appendi c e s2and3. TSb~e2 e0ll!Ps r e smea~ureme n tsof knDlinNewf oundlandmiero s.poddan species with thos e of.Yeis e r(196 1 ) andJamnba ci·(1 970) . No signi f i cant differenceswerenoted :exceptin

. ',

,

C.simu"Liiand

q..

breoi-oauda whichvaried sU. gh tly 1.nspor esize and

ca.u~al

len gt h.

'Measur~mcnts

werebased'on20 sp or e a

f~o;

e"achof5

".infect ed hosts .

M:i.cr osp o rid ansareof scat te r e d occurrence1.nNewfo~dla~d'

·b lackf l1 e s (Table.l) ,rith40 of198 ecreeea (20. 2%)harbouring on e or. IDDre specie s. Prosimu"LiUJ1/fU8c~mi:r;t;umlarv'a e were'found'Infect~d

~1y"ithC.oimu Ziiin 19of 40(47. 5%) Infected st r e8lll6jC.mutata

(42)

)'

r

..

""

,

...

!

,,1\"

,)

.

\~

"< "

1;" .:'

·i.I.1

!

TABLE 2

COHPARlSONOF HEAStJREMEirrS-()F,NEllPOtmD LAND HICROSPORIDANS WITIITHOSEOF WEISER(1961) ANDJAMNBACK(19 70)

.'f;;;":J;lr...,"1te SporeLetl gt h Spo r.Wi dth Cauda1.au gth

'~"'Spe$:1e s·

I

A' ,S .A

- s

'A

, 's I)

.T. br act eata 2-.5-4.0 3.5-4.0 .2.3- 3.6 2.8...3.'

" - .. -

T~ fib~a 5.0-7.0 4.5- 6. 0 3.0- 4.0 3. 5 - 4.S

-

,\,

p'."si muUi -4.0:-5.5 4.0-4.5

.. 2 .,' -"3.,

3. 0-: 3.5 ~

-

5.0',6.~

.

c.

simutU 4.5- 5.0 5.0' J.O-.5.0 14- 24

C,,b(e~cauda 4.8- 6•.0 S.O -6.0 i.9--3~9

.~'~ ' -· ·~:,S . ,

5.6'- 10

3.5 -\, 2.

,,~

.

0

,

..

:;;

\,.

(43)

20·

·withC.brevicau da in 1 of 40 (2.5%);S.cormswl~h-T.bl"'ao.~eata-in 1or 40 (2.'5%)

i

S.

la:Upes

withT.i?rdcteata~T.fib~ata'or P.~imuZii In.5

Of~12

..

5%·)·;~;

tube7'OsWn wi thT.librate.

o~

P.simulii.in'2 of

, ' " .-

. .

.

100(5.0%);S.',.venus

tum

withT.bJ'aCt~ata.T•.fibrata.or P.s,imulii tn 20

' of 4 0 ~50. 0,%);

andS.,

~i-t;tatum

withT.

bl'~cteatq

.. T. fibrota!orP.

sinrnZii in"of40 (10.0:0

of

lnf ee,t ed streams. Le ve l s" of para sitism .

(APpen d ix2) for all recorded spec i es werege~e~allYles sthan5.0%.

altho ugh l~fec:tionsof u'pto.I OO.a%'wereobtainedfrom cer tai n tndlcatedbias e d

~,~ples .

Tholohania

bl'act~ata

andT•.fi bra t aare'n ew reco r daforNewf~undl;.and .

The"fungus

Co~Zomycidi:.m's~lii. . .

&1so

.

' 's new

Newfo·undlan~··

.

-

·reccrd,wasrecover.edfrom1" atreeme(Table~),and infectedthe

•blackfl1esP.fusewn/mi:r.~~S. cor bi ea~d·S. venustum" The.in cidence

, . . ;.

of parasiti smwas"gen e r a llY'·.l e s s,t.han5.0%~but20.9;was.reco~ded~m one occasion(APplen?ix3).

Mermithids'areof

wldespr~ad,'but

scatte r e ddis t.r i1Jutlonin.:, Newfoundland (Table ,I , Fig.6)and.were recov er ed.f r? m 59~f1?8 I

.(29.7%) streamsYie~~~_~!:~.:~f.l1~S .The.t...:oregula~cOllei:tio~si tes, Half~~~__~.r;ook-1fii.QLongPO~dTr i b uta ry.ar e~ncluded,b:-,'Fig.,6 { but' '"': •.

~~xcl~ded

(romtabulationin Tablesl.andsppendtx 4,and.';lreconsidered

lat er~nmuchmor~detail. The averagemermithidinfe cti~mracewas u8ual l l(l e s s t'han10.0%,but;'heavily infe cted~carteredpopul ati ons of 20:0%ormore'occur~ed.

~ . -ro ee e spect.ea ofblac~f1ymerm1thids.we r e recovered ;namely NeomesomeI;'llli sflume11i:li is(Welch,-1962) NdckIe,1972,which was th emost '.ceea on,and

~resent,;~'n

S6of59(94.9%)of\ h estre ea e

hsr~~ur.:i~t·;

....

(44)
(45)

" q

..•'

"/

"+ . ""

"

.,~".."

~-.:

..

"

.". t

t;..;

l"

(46)

\

. v

n

merm~tblds.·Gas t l'Ome:rnn. s viridis.~elch.1962 W8S'pr esentin 2

o f

59 .(3~3%)andIeomermiamscol1sinensiawelch~1962 in 4

of

59 (6~7%)o~

the~tr~am8havingsim~lild lIIemi~h~dB. The~e'th'reemennithide ar,:

new recordsfor Nevfoundland.

.'NeomesO:neroria fLwnena lisregul a rly intects P.[uscwrt( mi:rtl;Dll and."S.venu8t~in Newfoundland ,butwasa18~r~cord edfrolllS.Zatip ea andS.tuberosWII.Iaomermial.>i8con8i~n8i8wasrecc ve e ed from

S':~ 've~u'8'~um

and,S.

~vi~ta~>

whileG.viI'idis

W~8 fOU~ln

S. cOl'bis

and S.-vitt atum.

Table)'showsth~~n~ldel'i'cecSfmerml thld'~nfeetl?nin,P . fU8CUJ!l/ mixtum,theon l ysl muUi dspecieswhich. were commonto all.

'S~H9trlcts. The data clearl!showt~the centra'l-regionsof Newfoundland (Districts)and 4) had a higher incidenceof Infe ctio~

with mernithids(75 . 0 and 47. 3%)than the more coastaldistricts.

District 6 had the lowest incidenceof'infection (11.7%' whiiethe

.

.

~.rema dnde r of th:districts.had.s i rnul1 i d s infected:o1thin the range of 20.6-36.3~.

Thereas o,n for the"h i gh e r prevalen ceor'm'erini t h idne~todes in th~int.eriorregionsof Newfoundlandis lat'gely speculativeat 't h i stime . Howeve r,it may

.~~

,th'at'

C~imatiC

factors,(cofde.r

winter~.

endwormer summers)are decisive.in influencingmermi~hid paras1ti~~

in Centra-lNewfoundlan~.It is also~onceivabl~tp a tth'~si mulii d . populations,which are generally conceded to

be 'morar..proliH~··i~·

Centra l Newfoundland'than in thecoa s t al areas,

wa~rant'-th~

cay-ryover' IIImcrep~raa1ticmermithids froml<8i-.i~e.to adults •than in.o the r.a r e a s)

(47)

!

~

~

~:

I 1 ~

~ ~ N 0 4 ~ 0

l'i

N

!

~

.

~ .

-r:o":£:l~

::: '"

~

. ~ . .~ ~

~ ~

.',

22

...

-«:

(48)

23

thet-ebypemit tingwider and easierdlB~era 81oftheme rmi t hida to new localit i e s.

" .

.

Thedat'sfrom Distr l(:[6,appea r stobe anomal ouswhen compa r ed with theothe r7dis t ri cts . To test possible reasons fo rlower paraslt& prevalence inDist r ic t6, wa t e ranaiyses of sevenstr e ams., .s{xWith ou t andone withblackfly'm;nnit hidinf ections , we r eca r r i e d out:inlateDecemb e r,19 72.

There sul t s (Table4) in d ic a tethat th e:r;e wereno aignifica n t

"dif f er en ce s between~nfectedand un inf e c te d str eamsfor pH,~orldes andammonfa, No eXplanationsare cu rren tlyavailable'forth~anomaly not edinDistrict6,b~tit' maybe rela tedt~reducedmenni thid ca Tr~overtotheadult,simuliidsor the,coastalclimaticconditions inthisre gi on.The possi bilityth at.menn ith id inf ec t e d simu111 rls

__cverem1s s ~ddur ingcollec tions~houldals ono t1J.~xcluded.

:Lon gi t ud ina lSti.:eamDist ributio n

The!iis~ribUtionofbl a ck fl y~rm1thid8,in.a streamfrom source to mout h va s stud ie d to determine ifinfections were mainta i nedat ,c'oRs tsn t lev els in all'areas orwex e confi ned toloca l iz~dportions of th e st reaill.HalfMoon Brook (2.8 kill.lon g) andPickav anceCreek' (1. 2 km.long)we r e examinedat int ervalsof approximate ly 200m, Bo th streamsori~inatedfrom ponds,·pr oce e ded through open scrub - marshy' 'a rea s to mixedforestofPiceamarit:::ma~A~nu8.rugo8a~Prounu8 .

(49)

~-~---,

e ~: '

~~ ~

~ ~ ~ ~ ~ ~ ~

~~

~

i:

'0

~

~

~. ~i;j

,.-.

~t:

~f}

~tl

~~

o~

j

~~ 0 0 0 0 0 0 0

' ) ~ll ~~

J

! ~ :::

~

:::

~ ~~ ~

~

.I:

*

(50)

25

« ...r

penn8YIV~~ca~:~d

Myrica

Gate

Iwhe re the b'Ulkof the

c::o llect1n~

vee

.PickavanceCreekwa sexami n ed pn7-e. 27/vt/72. while.Half,Moon BrookWIiSexamined,9/ V/7 2 and 15/VI/12.The_nn .Ly 1I'lermit!)idspecies rec?vered from bo thstreamswas N. fl.WT1ena,Z~,8•ThemefDIithidsfrom

.

,

Half.~o~nBr.:ookwerecollec tedfromP. fU8cum/mi;r;tWTI(9/V/72)._S.

venustuma~dS.tube1'C6WT1(one speHmenonlyat station 10, 15/VI/72),

. -

while Picka vanceCreekyielded onlypara'sitize.d larvae of S.venus,tum:.

Figure..,Bho~sthat,the first40Om.,or theopen~crubare a s', of"t heup'pe rportionsofthestreamswe r ed~voldofmerm1th1d parasites, butno tth ehost sp ecies.Merml thl dsoccurr ed in iti al lywherethe st reamsenteredshaded,mixed.fores t'areas,andthe'lev el of parasitiBm~

deciinedinadownst~~amdirection ,rising aga i nsli gh t l yat th e stre ammout h, The reason forthisdistri butionpatt~rnisobs c ure , atp reaent;,andadditiona l deta i l e~rese a rchshouldbe under taken t.ohelp clarify this prob l em.

Phys i c a l and ChemicalPa rame te rs

inanefforttod~finethe.ebeeucstand physicalenvironme nt al ,reqUi~e!ll~ntsof simuliid'mermithids.and thuspossiblyprovide ,8 means 'of pred ic tingthe su itabil ityof ~stresmfor ebee, a cpmparisonwas mad.::of suchfactor~existi.ngin two streams·wi.t h•.and one at reaewith out

(51)

Figure"7. : Lo~itudinalstre"smdistr ibu tionof

N. fZumena'Li8infeet~d sima.l1ids.

Note:Station 1.headwat er s of stream.

" .

-:..

(52)

'J

,

)

-

, { r

'\ ~

\ ' .

--. '<

~' \

.

\

, ....l

•~Q1J.:JUIl lN3':ltGd:

(53)

26

black!l)'mermlthids.Ten chemicalparameters (dl ssoI vc:doxy gen , carb ondioxide,eeeona,e ,ni t ~ate s. phosphate e,chlo ri des , totalhard- nes s,.,silico n di oxide, hydro gensul f i de'a nd pHl,..an d' th ephytdt!al fa ctorsofwatercu r r e n t , depthand temperatureweremeasured .Thest r e ams stu di e d were HalfMoon Brook "and LongPond Tribut a ry , bothst re a ms

.

.

/

withslmul 1i~sInfec.tedwi thmennlthids,and PiecosBrook'(Append ix1).

inloIh i c hon ly asin.gI e.mermithld 1nfec t l onwa s fo un<(April,1972).i n twoyearsofsampiing. In all the s,c r e ams,al!mlQn~a.nitrates.'phoBphat~s

·and hy dro ge nsulf i de were virtuallyabs en t ~ndar eomitt e d fromfu~ther.

• consideration .

There aul t s ofthecurrent . depth and watercon t e n t analy se s over theper iod23/VI/71 to,17/VII/72'are presentedinFigures8 and9

·and Appendix5;the effe cts oftemperaturewill becons i de r e d.inmore detsil inthe seasonal fluctuat i,ons sec ti on.I twillbe. no te d (Table 5) J thatallrang esofth e.var doua par ameter soverlap, and nocha~~c~eri9tiC

veefoundwhich coul d best a t e d,,"8being a speci fi crequ iremen tfor mertllith l ds.

t· : .

Isol a t e d observationson.wa t e r anaiysesfrom,otherNewfoundland streams ha r bou rin grne rmith l d infectedsitnul iids provide stbebasi sfor' theestablis hme ntof ...guidelin eto the chemicaltolera n celimi tsfor

~ . ' I '

N.fluihena.lia.Maximumandminimumchemi: alparameterli~itsforcarbon d·lox.lde of2-15ppm, chlorides of15 -250 ppm,eoeaihardnes of 20-25~p'pm 'a nd:pHof4.5- 7.5were noted.Tofu~thereIucfdat.e the to l e ran c.e limitsforN. j"ZwnertaU a.,addit i o na l detailedinve s tigati ons

·

·

of wat er-fromst r e ams with and withoutmermithids,. should be undertaken

. '

~ .

.~

During'thewi nte rmont hs ,Jan u ary to MSl;ch, when

th(

streams

. . "

,.~

··r

were ice-cove red,'8sl!ght·d ecrea8~:·in dissolv..!4'oxy genand asligh t

(54)

·

,

Figure 8.'

.,- ' ., "

. . . .

"t oiopa r1 s on of mi ni muriiand maxlmUIIImont h lydepths.for, the.tworegula'!:co llec tion:sitesin 19 71 and19.12.

~ • J

(55)

:','

....

r.

,.

I

t•

-c Ii

~

'"

iil

0

.!!

.~

:>

0

§ "

~

R

~

e

~

~.

••0

9

(t'O)1Ud 3Q

."

. ...

::.

(56)

r

' f

' '\

"

Figure 9.

, . ,

"

.~,Col1lp arh onpfm.in~ml1mand lIlB.)I:imummont hl ycurr ent sfor thet~.re ~ lar'col1ee t ~onB~.te9 1~ ~971and:\.972.

":. '

".1 ' "

(57)

1.10

~ ~

.o~ .>;

;

: ~

I:SO

·u.·

qO.

>:oJ

LONG RINDTRIBUTARV

.M J0-.J A SON 0 J F.M0A M ~ J A

. .

(58)

::"

~GE

OF

ENYrRO~AL:'P~TERS ~R':~~

.

' WITH ANI1 WITHOtrt BiAcur.yHERMITRlns

', . -, '- .' . z

._~.".:.

:'

.'."2: g·~·.5.5'. :.:

. : :;t+;:~·,

".·S.~-

'6'-'

~.O -

i:) ,,' o

-21,

,'';'-

-r-

.,;:.

, ~'. '.

- ~'p:l,c:cos. .".':'

."Br ook . '

,~

i

,'j:,

",2.0 7.0

6 .8 9.'" ;.

~:.

1-5 25 45.-:70.-. .\~,

" 4 .S .' - '6.0 : ~. ,

IS-'Z. S

o "--22.. 0 .32':1.56>

1 -32

Long Pond: .Tr l bu.t a r,.!l;.!!"

TABLE5

":' .-

;

....

, ' . \ ,

2.0-.S.O.

6;0 - 9.0..

.'20'.'-'

30 .

,20 .."30 4~5

- 6. 5

·l .~ :~ 2 .;:'

O·.•'';'20.0•

.

,~.2i-

.

gig

2 - 40

.HaitHoon: : .". Bro?k····:....

\",::,-.

",., "

Carbo n dioxide·

·Di..oiV~d·,oXyg~.

Chlor ides'"

Totalhardneu *

•pH..'- ' . • ,

Si liconiio xi de""_

.Tempeu t u r e.(~C~)•' '. , Cu~~~nt(met ers/sec•.)

~tK'(em. ) .--,?

"I

... ..

,,~

.~

.

:.'.

" ....

...

i,"

',.:",

1- "

...

. "

.

"

:"

".:': . , '.

'

*in p arts',

'per-'lIli1ii~n . . il!:.*hs:rbo~ r cen:dthi~Inf~ ctedblac kfl1ea:'

'-';.' .

' .

. ;,,'~~' ­

. .-

.-...

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