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
3G2:: 7 0
, ,
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" 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 at6
oC,: ~nd
failed to molt.-a t 22_24° c.,Adouble
pre-~~it
mol tan~ 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 .
: .
.~.
l're-paradt~s11Ye. 2-3'~llY$.The period fTo.eUler~encefrol:lthehost•
.
t~'
egg_laYi~'g wa~
45- 74"day s ,.and'fr-.emergencet~
'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 afrolllla~
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 dh05t~/ 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~:
inf1~lf
HoonBr~~k ~re
alsonoteil :In1~72
•. ".,
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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
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"
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 figures20
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
"- .
'/-",'
.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.'\
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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.
lindpetBeverenl:~~ ~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. :
.»,
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,.':
INTRODUCTION'.
Tb~' Si~Ul1ida.e
are·ac: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 usepOPulatio~~'
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'lossto .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 ofD~. ·
.-::....:.-;,:. -L•
-'<'/:'"7:: '
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ManyD~~~nsect1ctde8bavebee n"'developedtosti pp lant.DDT,but"all . .
~a~e
l,nherentdi f fi cu l tie s"I I'.~~~t
1)th ;rPO~'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 ape:t 8pe~les
bymean s'ofanot~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 ero~
Brassica,crepeth;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 thaig~iicant r~duct'ion:
.In pe s tnualbersoc cu r ring"
" ,
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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\tiali~
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~
theTetradonematida~
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"llyco~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 lumenand1; 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~~~~;;.
-"
.'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 rNewf~~~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.
.. bothor 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.
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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 specieswassy~~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
andBic~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 '
)
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.Sbasically 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 dgiantcells--a~
valid criteriaf~r
theide.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
(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)."' ,," 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;,Welcha~d, p~ina~. 1965~
wereal~~
of ' "I
same opin i on. , " '.• ,
' ~.
..~
',~
-
. " " /
METHODSANDMATERIALS
i I '
Apr elimi n ary surveyof ro a ds id eetreeee, selected for ease ofaccess, 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.
" ..
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.
:J
" j
!
'"\. F1gure2a•
•" \ . .ie • • •
'''''~
" " . . - . ; . . ". . . .\
tI
UpstreBlll viewof Long Pond Tribu tary above.the regular collection,s i t e.
" .~/.
.., ...,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:I8u811ycollect~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 entMeter~. w~ile
the'waterana lyseso(.~.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 micalProd~cts
Company, cneseeeecvn,Maryl a nd---
~.
~.-;-:...
" . .
"','.,
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~·
.'}
.
~
-
..
.
'\
....
~.' ~0~~
~ ..
f;-.
, .
Mi~roapOr1da:n" ·infe~tfng·the 8:t:mui1id~·P.'..',r;~tU'n
.
., .. (Linearm.agnif1.cat10~_,X10.0~·:..
, ,: ~ ; " .' : :, ' ','. F ig".
'5.: j ' ' : ' , , .
' ~oeZOlliycidiW1l
simu.z-t.i"in f ecti ng th'~~mUl1id.
,S."ven'~Btum.
. .(L.inear,magrtif1catlonX 12.0) .,. . .
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
. '
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;~
eecont~~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.
.
. .
··,i
"
Measurements'
~f p~_ep8raS1tes
wer;m~de
en a CarlZeissMic.ro-~
.yideomat6 at a 3200]1:
lI1ag~1fic.at:ibil. ~Bennet~
andCamPb~~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
,
weretakenon
a Carl ZeissTeSBovar•.
60b erkochen, We s t
G~rmanY.
"i
RESULTS ANDDISCUSSION
-.'Distribution
TWo_
au~veys'
for•.•biackfly endoparas iteswe.~,~'Cdii'd~~'~~~
1nN~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 onal1 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'iyp~~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
;-
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-
7S.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 1S.•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
:;:
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 rJidiumExamined 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.
</
:;
. . .
,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 ree~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 andca.u~al
len gt h.'Measur~mcnts
werebased'on20 sp or e af~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
)'
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- 24C,,b(e~cauda 4.8- 6•.0 S.O -6.0 i.9--3~9
.~'~ ' -· ·~:,S . ,
5.6'- 103.5 -\, 2.
,,~
.
0
,
..
:;;
\,.
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.5Of~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 .
Tholohaniabl'act~ata
andT•.fi bra t aare'n ew reco r daforNewf~undl;.and .The"fungus
Co~Zomycidi:.m's~lii. . .
&1so.
' 's newNewfo·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.';lreconsideredlat 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 ehsr~~ur.:i~t·;
....
" q
..•'
"/
"+ . ""
".,~".."
~-.:
..
".". t
t;..;
l"
\
. v
nmerm~tblds.·Gas t l'Ome:rnn. s viridis.~elch.1962 W8S'pr esentin 2
o f
59 .(3~3%)andIeomermiamscol1sinensiawelch~1962 in 4of
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'idisW~8 fOU~ln
S. cOl'bisand 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.rwinter~.
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)!
~~
~:I 1 ~
~ ~ N 0 4 ~ 0l'i
N!
~.
~ .
-r:o":£:l~::: '"
~. ~ . .~ ~
~ ~.',
22
...
-«:
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 .
~-~---,
e ~: '
~~ ~
~ ~ ~ ~ ~ ~ ~~~
~i:
'0~
~
~. ~i;j
,.-.
~t:~f}
~tl~~
o~
j
~~ 0 0 0 0 0 0 0
' ) ~ll ~~
J
! ~ :::
~:::
~ ~~ ~~
.I:*
25
« ...r
penn8YIV~~ca~:~d
MyricaGate
Iwhe re the b'Ulkof thec::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
Figure"7. : Lo~itudinalstre"smdistr ibu tionof
N. fZumena'Li8infeet~d sima.l1ids.
Note:Station 1.headwat er s of stream.
" .
-:..
'J
,
)
-
, { r
'\ ~
\ ' .
--. '<
~' \
.
\
, ....l•
•~Q1J.:JUIl lN3':ltGd:
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, whenth(
streams. . "
,.~··r
were ice-cove red,'8sl!ght·d ecrea8~:·in dissolv..!4'oxy genand asligh t
·
,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
:','
....
r.,.
I
t•
-c Ii
~'"
iil
0.!!
.~:>
0
§ "
~R
~e
~
~.••0
9
(t'O)1Ud 3Q
."
. ...
::.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 ' "
1.10
~ ~
.o~ .>;
;
: ~
I:SO·u.·
qO.
>:oJ
LONG RINDTRIBUTARV
.M J0-.J A SON 0 J F.M0A M ~ J A
. .
::"
~GE
OFENYrRO~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-.
gig2 - 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:''-';.' .
' .
. ;,,'~~'
. .-
.-...