USDA Forest Service General Technical Report S E- 6 December 1974

Current Knowledge of Monodontomerus dentipes (Dalman), A Cocoon Parasite of Pine Sawflies

by

Vicki H. Fedde

Agr icul ture

S ti7 t ;or1

(7

Current Knowledge of Monodont omerus dentipes (~alman),' A Cocoon Parasite of Pine Sawflies

V i c k i H. Fedde, Associate Entomologis t Fo res t r y Sciences Labora to ry

Research T r i a n g l e Park, Nor th Carol i n a

Monodontomerus den t ipes (Dalman), a European p a r a s i t e o f cocoons o f p i n e saw f l i es , has been a key f a c t o r i n t h e c o n t r o l o f t he in t roduced p i ne sawf ly , D i p r i o n s i m i l i s (Ha r t i g ) , i n t he Un i t ed S ta tes and Canada. Several o t h e r D i p r i o n and Neodipr ion spp. have a l s o been repo r t ed as hosts, and the s a w f l i e s which occur i n t h e southeastern Un i t ed S ta tes a re poss ib l e t a r g e t s f o r c o n t r o l by E. dent ipes. Dur ing a r ecen t s tudy o f t he development and emergence o f t h e pa ras i t e , a comprehensive survey o f t he l i t e r a t u r e on M.

was made. The purpose o f t h i s rev iew i s t o a i d o the rs i n s tudy ing o t h a t i t s usefu lness as an agent i n t h e c o n t r o l o f Neodipr ion spp, can be enhanced.

The o r i g i n a l d e s c r i p t i o n o f M. dent ipes i s g e n e r a l l y c r e d i t e d t o Dalman (1820) who descr ibed bo th sexes o f t h e a d u l t . A number o f sources ( H a r t l e y 1923; Faure 1925; S i towsk i 1929; de F l u i t e r 1933; Mokrezecki 1933; Sche id te r 1934; Gahan 1941 ; Bucher 1948) i n c o r r e c t l y 1 i s t e d t h e d e s c r i p t i o n by Boheman (1833), based on a s i n g l e female as hav ing p r i o r i t y , Dalman named t h e p a r a s i t e

* den t i ped i s and p laced i t i n t h e f a m i l y Pteromal-

idae, t r i b e P te roma l i n i , The species was t r a n s f e r r e d t o Monodontomerus (Walker 18471, which a t t h e t ime was i n t h e f a m i l y Cal l imomidae (Westwood i 833 ) . The name C a l l imomidae has been p e m a n e n t l j suppressed (peck 1963). Monodontomerus has a l s o been l i s t e d i n t h e f a m i l y Cha lc id idae (Howard and

u t i s p r e s e n t l y i n t h e f a m i l y Torymidae (Gahan 1941 ) . The i s synonpous w i t h Torymus v i c i c e l l a e F o r s t e r (Walker neus Provancher (Burks 1963).

Several keys a r e a v a i l a b l e f o r t h e d iaanos is o f M, dent ioes. Adu l ts were separated fFom those o f o t h e r ~onodontomerus spp.-b$-1 941 ) and S t e f f a n (1952). A comorehensive 1 i s t o f re fe rences by species o f Monodon- tomerus was b) Burks (1963). S ince t h e r e i L l i t t l e over lapp ing

o s t s o f M. den t ipes and o t h e r Monodontomerus spp., i d e n t i f i c a t i o n . . - i s s i m p l i f i e d through t h e i d e n t i f i c a t i o n o f t h e host , e s p e c i a l l y i f t h e hos t i s a d i p r i o n i d saw f l y ( t a b l e 1 ) . I n Nor th America, M. *

Ashmead and M e d e n t i ~ e s a r e t he on1 v Monodontomerus SDD. which a t t a c k saw- f l i e s , The Gl a f gLfe mkkican saw f l y i s i n Canada, where i t was released ayson and F in laysen 1958b), In t h e abse -- GUS on D, hercvniae, i d ~ r s h a b l v has n o t become es tab l i shed . M. den t ioes \ Has sepa-

o t h b r p a r a i i t e s o f p i n e sawf l i e s by M o r r i ~ e 371, F in - layson (1960a, 1960b, 1962, and 19631, M e r t i n j (19671, and Mer t i ns -and Coppel ( 1 971 ) .

' ~meno~ te r a : Torymidae.

Table 1,--Host families of Monodontomerus dentipes (Dalman) and some other Monodontomerus spp, with those parasites having host families in comon with M, dentipes indicated by horizontal 1 ines

Order and family

HYMENOPTERA

Apidae

Braconidae

Chalcididae

Cyni pidae

Diprionidae

Ichneumonidae

Megachil idae

Sphecidae

Tenthredinidae

Vespidae

LEPIDOPTERA

Arctiidae

Lasiocampidae

Liparidae

Lymantri idae

Olethreutidae

Saturni idae

Tortricidae

Zygaenidae

D I PTERA

Tachinidae X X

Description of Adults

The most complete treatment of the morphology of the adults of M. dentipes is a detailed study by Bucher (7948). He included numerous zraw- ings with descriptions o f the internal and external anatomy of both sexes.

GEOGRAPHIC DISTRIBUTION AND IPS RELATION TO HOSTS

Generally, the distributi depends on the presence of sawflies o f the f a m i l y Dipri ther groups are repre- sented as host 2). Whe occurs, the hosts are most commonly spp. and ther Ifmenoptera, D i p - tera, and Lepidoptera are a l so parasitized, b u t far less frequently.

Table ?.--Recorded hosts of Monodontornerus

France

Braconidae

SP a Poland

A , o'igmeratr;~ L a a - u- France

I ta ly

Meteorus versicolcr Wesmael HoI land -- Cynlpidae Skieden

Digr ian idae

SPP'

D. frutetorum [Fabricius) --

Rosenburg 1934

Thompson 1944

F a w e 4925, 1926; Thompson 1944

Thompson 4 444

de Flui ter 1933; Thompson 3944

Balman 1820; Boheman 1833; Howard and Ashmead 1898

Poland Mo krez ecki 1 933

USA: New York Viereck 1928; Gahan e t a l . 7928

Canada: Ontario

C~echosloi~akia

Canada : New Brunswick

Nova Scot ia

Ontario

Quebec

Ckiria

Germanj

Hull and

Poland

E S P : Maine

Vermont

D. ( K l u r j j Czechosiavakia

Raizenne 1957; McGugan and Coppel 7 962; Griffiths e t a l . 1971

Finlayson and Finlayson 1958a

Baird 1938

Baird 1938

Finlayson and Reeks 1936; Baird 1937, 1939; Peck 1951 ; Finlayson 7960b; McGuijan and Cappel 1962

Finfayson and Peeks 1936; Baira 7938; Briand 1949; McGugan and Cappel 7952

Peck 1963

Pecr I963

Peck 1963

Peck I 963

Procter 9 938

Dowder; 7958

Morris e t a i . 1937

continued

See footnotes a t end cf table .

(Dafmn) (continued)

Scheidter 7 934; Thompson '1 944

Holland Ritzenia Bas 1919; Thompson 1944

Pol and S i towski 1429; Thompson 1944

Sweden Howard and Ashmead 1898

Gzechosl ovakia Molubajiv 1934; Morr is e t a ? . 1937; Thompson 1944; Finlayson and Finlayson 1958b

Gemany Morr is et a?. 1937

Europe (general ) Gahan 1947; Dowden 1949

Canada: Ontar io Baird 1939; Gahan 1941 ; Bscher 1948; Dowden 1949; Raizenne 1957; S ippe i l e t a?. 1960; Finlayson 7962; McGugan and Goppel 1962

Quebec Baird 1938; Finlayson 1962; McGugan and Coppel 1962

China Peck 1963

Germany

Hol 1 and

Thompson 1944

Hart1 ey 1923; Thompson 1944

Hungary Eahan 1941; Peck 1963

Poland Sitowski 1929; Finlayson and Finlayson 195Ba

Sweden Howard and Ashmead 1898

USA: Connecticut B r i t t o n and Zappe 1918; Gahan 1941 ; Dowden 1949

Maine Gahan 1941; Procter 1946

Michigan Gahan 1941

New Jersey Weiss 1917a, 1917b, 1918; Har t ley 1923; Gahan 1941; Dowden 1949, 1962

New York Gahan 1941

Ohio Gahan 1941

Pennsylvania Har t ley 1923; Gahan 1941; Oowden 1949, 1962

Wisconsin Shenefel t and Benjamin 1955; Dowden 1962; Mert ins 1967; Mert ins and Coppel 1971

Neodiprion a b i e t i s (Har r is ) Europe (general J Bai rd 1944

Canada: Ontar io McGugan and Coppel 1962

New Brunswick Ba i rd 1944; McGugan and Coppel 1962

N. exci tans Rohwer - - N. leconte i ( ~ i t c h ) ' - - N* - (Norton)

USA: F l o r i da W i l kinson, personal c m u n i c a t i o n

USA: F for ida W i 1 kinson 7969

Canada: Ontar io Raizenne 1957; McGugan and Coppel 1962

Rani toba Ba i rd 1941 ; McGugan and Coppel 7962

continued

See footnotes a t end of table.

Taal e 2, --Recorded basts 04 ~ o d a n t m e r u s =I& (Dalman j icontirueci 1

I So~rce

USA: VSrginia &rwz 1971

k , swainei Middleton - - USR: Wiscons~n B~cCer and Benjam*~ 7964

U S P : Wisconsin Mertins 1367; Mertins ~ n d Cappel 1971

Tetrastickus xanthgps 7 Ai i s t r f a Seitner 1927

Exenterus anictorius JSA: Wisconsin

spa7 Austria

"c~or;ough) " GSA: Idaho

Moll and

P~~theLria dispar ii. j p-

r;ew Enijlani?

Pieridae

Mertins 1957; Mertins and Coppel 1971

Mertins 1967; Mertins ano C ~ p p e l 7971

Seitner 7421

Seitner 1927

Seitaer 1327

paward and Ashmead 1898

Sei tner 7 94 6, i 427 ; Mart1 ey i 923

Thompson 9 943

Peck 1963

Eeddrd 1938

de Fluiter 1933

Burgers 4915

F j e r ~ ~ brass7cze L . P -- France

See foetrotes a t end o f t a t i ~ ,

HOSES Source ----

Tortricidae

3' L , France p o s e ~ % r g 1534

Tortrix vsridana (Girard] --- Sweaen Loward a r d Fshqeai 1898

Zygaenibae

1-aa i . Sweeien noward z ~ d ffshead 9898

Z, occStansca de V.illers Fra~ee Hartley 1922

' ", der~tspr ; ; ns nypErparac?t7c c r wc f ~ r ~ ~ e l l ; ~ i . thrai lzb t t t a q i c d , 5. d e n t ~ p ~ r a IS -yperpa%iizit-c 01 ? ~ e r ~ s brass-cae L. kpr3:gh 5, $&T?)"KYL~- " E, Cent~pes i s kyperparasit~c GP z_?l-ps& re?s I. "c~r3,qF., z* SC~IW- (de F I U I t 2 ~ 4333,.

Earlj !North drner~can keq~rerces t:, g , =cjnme r-rstakenl; -iden?f;eC l e s s f;, @ ; j t ~ ~ u r n (#P~L!IB~

and Copre: 19621,

s g, a ~ d C O P ~ U I S I ~ O ~ . - -- - - -

was released agalns ' t theze ~ ~ s t ; l r a bl~;03l~al cclbro; effort of the iiSDC csres t ve beer made t o da le . i s hyperparasi"c-;c: on D e ~ d r o l - a "srrcug$ tb cles {Seitre. 19211, was reported as a prima;-;. paras~te o f 4, ~ r ) l ? a s ~ t : n g empty cciicuo~s o f 12,

M, is native t o the Palearctic regioq o f the world and d i s - tri bu tzd l y in the Holarctic region ( t a b l e 31, b l t h c u g k - M, -- den-

* I s found t h ~ o u g h s e s t the Palearctic region, only a single locality e x i s t s for China (Gahan 1947), - M, has been reported

th roughou t northern and central Europe, R om France and A u s t r i a 1 i s t nonsawfly hosts exclusively. Peck (1 961) be1 ievi id these hait : suq- gested i t e was i d e n t i f i e d in~orrociil a*.: ti 5. aer~ijs hzlke! , not E, In Europe, D i p ~ i ~ ~ pii,l ( 1 2 -- .AA."-~z olvfowlr; i : am---- (&rtiqI i P

are the most frequently pawas- i t l zed h e s t s ,

I n the Bearctic region, Ee occurs i n the United States ana Canada, primarily east o f the Gre s , M, &isJ$j= was acr;Sdenta" !Y introduced i n t o the United S t a t e r as a parasiTe 6-f the sawfly, ). &m""l;s, which subsequenuy became the most frequefitly at tacked hos t i n th i s coun- dry. gLg, was fa"rst reported ip Co~necticut aaoucd 5910 ( B r i t t o n 19159 Fr-i Zappe 1918) and shor t ly after i - ~ New Jersey {Us'ss 11%17a, 19]7b, 19m; Hartley 132% &hac "GdI: EI.~U:<.:. i44;), z:f 1322 z - U,

w-

ss"mi1-8"~ was effectively p a r a s i t i z e d c j i"", denljpes T-.i a aik; ,or f - J e s l a e j 3 ~ -" ---- near Philadelphia, Pennsyl va:ila (t;ar::!i.j { i 2 7 j ~ ~ ~ ~ - c t ~ vn2rkeC the r s~ehern- most extension of the parasite's distribution a t t h a t time, i n l a te r years, the parasite was reporte e same hos t i f i !4atng, t-!<chigan, Gh-is, 6334 Wisconsin, and on other spp, in Maine, Ver*nsnt, and t4ew Yoork { t ab le 4). West o f Wisconsin, a single record e x i s t s n f M. dcnli_ijes, and two --- -- -- lepidopterous hosts are I isted (Beaard 4 938j,

Table 3.--Palearctic distribution of lulonodontonterus (Calman)

Ccsntineot and

country Hosts Source

A S I A

China D~prion hercyniae jiiartigj

O, sinilis (~artig) - -

EUROPE

(general 1 (I--1 3, sirnilis - - a, sertifer Geoffror - -

Austria Dendral imus pini L > Tetrastichus (~atzebur~)'

sp."

Therion (~ravenhorst )'

Theronia atalanta ( ~ o d a )' -- Rhogas sp."

Czechoslovakia e. frutetorum (F.)

8 . - Kl ug

D. (Hartig)

Germany

Hol 1 and

France

Cydia pornonella L. " Pieris brassicae L." --

acci tanica d e Y i l lers

Diprion

D.

D. hercyniae - D. sirnilif - - Diprion pinj

Meteorus ~ersicolor Mesmel

Stitpnotia salicis t.

Peck 1963

Peck 1 963

Steffan I952

Gahan 1441 ; Giowden 1749

Sreffan 1452

Seitner 1916, 1927

Seitner 1921

Seitner 1927

Seitner 1927

Seitner 7827

Seitner 1927

Finlayson and Finlayson 295%

Morris e i a l . 1937

Kolubajiv 7934; Morris e t al. 1947; Thompson 1944; Finlayson and Finlayson 1958a

Faure 1925, 1926; Thompson 1944

Rosenburg 1934

Rasenburg 1934

Faure 1925

hart1 ey 1923

Peck 1963

Scheidter 1934; Thompson 7 944

Harris et a l . 1937

Peck 1 963

Hart1 ey 1423 ; Thmpson 1944

Ritzema Bos 1919; Thompson 1944

de Fluiter 1933; Thompson 1944

de Fluiter 1933 continued

See footnotes at end o f table.

Table 3.--Palearet ic d i s t r i b u t i o n o f Honodcntornerus (Dalman) (continu&)

Continent 1 and

country Hosts

Hungary D, simrlis - - I t a ly

P ~ I i s n d

D. sirnilis - - P. brassicae - -"-

Cynipid wasps

5

13. s i r n i l i s " - -

e." Yortrix vtridana (Giraudj --

L.

Sweden

Gahan 1941 ; Peck 1963

Thompson 1944

T ~ O ~ O S O G 1944

Mokrerecki I933

Mcikrerecki 1333

Pack 4 963

Sitowskr" 1929; Thompson 1944

SStowskl 1924; Finlaysop and Ffnlayson 1958h

Mokrerecki 1993

Balmn 3820; Bobeman 4833; Howard and Askmead 1898

Howard and Ashmead 1898

Howard and Ashead 1898

Howard and Ashead I898

Howard and Ashead 7898

Howard and Ashead 1898

In, - i s a hyperparasite of Dendroli~us @ h a t h rough these species [Seltnsr 9927)-

3 - i s a nyperparii-stke of P. E. i s a hyperparasite o f ge;,

414. is a hyperparasite of 5. g, ve r jde Flutter 19331, 5 ~ s " AS s i r n i l i s Wartig, " AS L,

Table 4,--Neitrctic distr ibutlcnn o f Panodentmerus dentipes (Dalnan)

Source

NORTH PMERICA

(general j Peck 1957

Peck 7951

D. s i s i l i s Idaraig] - - Peck "91

Cjiptera Peck 7951

j~e~onnough jZ Peck 1957

Peck 1951

Burgess 1915

Ganzda

(genera? j P. sfrntlis MrGdgan and Coppel 1962 - (Geocf roy j Finlajson 7560a

l ex McFugan and Coppel 1962

Manitoba banksianae ReRwer Baird 1941 ; McGugan and Coppel 1962

Nek Brunswick C;. Balrd 1938, 7939

N. abietis (Harris) - - B a ~ r d 7944; McGugan and Cappel 196%

Nova Scotia B.

Ontiar~c - G . -- frutetorum ( F * )

Quebec

N. abiet-is - -- k. pineturn (Norton:

N. banksianae

N. ser"iifer - --

6zirenne 1957; McG~gan and C o ~ p e l 7962, Griffiths e t a i . 1971

F i n l a y r o ~ and Reeks 7936; Baird 1937, 1939: Feck 1951; Finlayson 1960b; McGugan and Coppel 1962

Baird 1939; Gaban 7941 ; Sucher 7968; Gowden 1944; Raizenne 1957: Sippel1 e t a l . 1960; Finlzysan 7962; McGugan aqd Cappel 1962

Baird 19dl

Finlayson 1960a; MsGugaq and Coppel 7962; Lyons 4454; Grirfiths ~t a:. 1477

a buoliana ~SchifFerm~ll~r) Miatsop and A r t h u r 7959; McGugar: and Coppel 1062 -- - Finlayson and Seeks 1930; Baird 193e, 1939; Briand 1949; McGugan and Coppel 1962

Raird 1938, 7439; Fdnlaysan 1962; MeGuyan and Ccppel ;462

See Pcotnotes a t end o f tab le .

Table 4.--learctic d~sdribud;an o f ?"ksnodontorre~ (0almnj (continued)

Local r ty Hosts i i Source

Connecticut f. sfmilis

Maine

Mew '{orb:

Ohio

Pennsyivan~a

"imont

Virginia

W.isconsin

sop,

D. simills - -

D. sjrnilis - - D. s- imi ' i is - -- n, N a

Ancistrocerus -- (Causs~~re;"

Britton and Zappe 1918: Britton 1926; Gakan 1941: 5ouden 79d9

Meiss 1917a, 1937b, 1978; Hartley 1923; Gakan 1943; Dowden 1949, 1962

Viereek 1928; Gahan e t a l . 1928

Gahan 1941

Hartley 7923; Gakan 1941 ; Dowden ?945, 1962

Shenefelt a r d Benjamin 1955; Dcwden 7962; Yerti~s 1967; Wertins and Coppel 1971

Fi, swiaicei Mfddleton -..-.- Becker and Benjamin 1364

' A S Henerscamn ~ s e u d o t s u g a t a . 9 --

Early North Amerfcan refererces to &, and Coppel 1962).

3 ~ , d e n t - i ~ e s was released against these hosts i r 1965-1979 th rough the bltilogrcal control program o f the USDA ForesT ~ x e i i t has n u t been recovered t o date ,

*M. -- was listed as a pr-inary p a r a s i t e tigr.rs cornon Irrhaafeants of cocoon; cf 6, s~milis, - --

In Canada, E, has been released against pine s a w f l i s i n biological control p n Ontario, Quebec, Manitoba, Nova Scodia, Mew Brunswick, and Prince Edward I s land ( t ab le 4). In Ontario, E. i s camon and more frequently at$acked t h a n any ather potential host, How- ever, i n Quebec and the Haritiine Provinces, D, i s the most abun- d a n t sawfly and the most heavily parasftlzed:

Generally, i n the Nearctic region, M. i s concentrated i n coastal regions, Yke distribution s f the pa tends along the Atlan- t i c coast f rom Pennsylvania to Quebec, and westward along the Great Lakes ta Wisconsin and Manitoba. Recent releases aga ins t sawflies i n the south- eastern United States ( W i l k 9; Morris 1971 ) have provided an oppor- tunity for the range o f M e t o be extended southward as f a r as Florida, however, estebl7shment has not yet been determined.

HOSTS

Included among the known hosts o f M. are s i x families of Hymenoptera, one family o f Diptera, and f i v e fa rn i l~es of Lepidoptera ( t a b l e 2 ) . Among the various farnil ies t h a t have been reported are: Diprionidae, Braconidae, Ichneumonidae, Eulophidae, Vespidae, and Cynipidae in t he order Hymenoptera; Tachinidae i n the order Diptera; and i i p a r i d a e , L p a n t r i i d a e , Zygaenidae, Tortricidae, and Pieridae i n the order Lepidoptera.

t h r o u g h o u t the dis t r ibut ion and P i v e species o f

as p o t e n t i a l hosts, are

In the Hearctic r ion species and Pive

Uni ted States , In Nor niae are the most fre-

the parasite has been reported iKfr as we11 ( t a b l e 41,

Throughout Europe and the Nearctic region, E* has been reported on bepidsptera and Hymenoptera other than DI e (tables 3 and 4 ) . Collectively, records o f nonsawfly hosts are insignificant when compared numerically to records o f sawfly hosts portant fac to r i n the host r e l a t i a ~ s h i p s of H. been recorded on three species of Braconidaedn Ichneumonidae, Similarly, only a few cases o f

tera in & o ~ L h America are from Id

"risr to 1941, when B was correctly identified by Sni tk (1941) 2s the species involved i n Canadian infestations o f this sawfly were believed to be conspecific w t h those of t h e E u r ies D. s such, several ear ly sources incorrectly refer to 6,

as %). {MsGuga

Morris e t a1 , (1937) and Mertins (3967) are the only authors who have touched on a variety o f t o p i c s related t o the l i f e cycle s f the parasite,

In their studies, they included mating and oviposition, the eggs, maturation o f larvae, pupat ion, emergence from the

host cocoon, l ongev i t y o f a d u l t s , and host relationships o f M. Mertins and Coppel (1971) revised the material pertaining t o ho jon- ships t h a t Mertins (4 provided, They omi t ted much sf the detailed information about E* t h a t Mertins (1967) had included,

The stages i n the 1 i f e cycle o f M, ere described initially by Morris e t a1 . (1 937). These workers s" n the laboratory so their description was I d and on field-collected material, The fe f i v e o r s i x (maximum, 15) eggs i n a sing externally on the prepupa s f the s a w f l vouring i t cofipletely, The larvae matured af ter 3 t o 4 weeks, M, overwi ntered as prepupae, o r , i f weather were favorable, develope t e l y i n t o pupae, In April, when overwintering i nd iv idua l s pupated, the pupal stage l a s t ed 3 weeks, During the summer, when other individuals pupated, the pupal stage was shorter, Four or f ive (maximum, 1 2 ) adu9 t s o f M, emerged per host cocoon. Emergence o f the f i r s t generation o f a&l ts occurred i n May, a l - though a d u l t s were found i n the f ie ld i n t o September. A f u l l month elapsed between the emergence of the F i r s t and las t insects from one cocoon,

Mertins ' (1 967) observations agree essential l y w i t h those o f Morris e t a l . (1937). However, Mertins provided additional information about the Wisconsin parasites o f - 0, sirnilis. He reared M a from f i e l d cs l - lected cocoons of D m and s tud ied the bgha longevity o f the para er unspecified laboratory conditions. M,

took 28 days t o develop from eggs to a d u l t % Larvae hatche8 w i t h - i n 3 days o f oviposition, fed on the host f o r 8 days, then were quiescent Psr 3 additional days before defecating* Afterwards, the larvae tended t o position themselves i n a single layer around the host cadaver and cocoon walls, a71 fac ing i n the same direction, They pupated w i t h i n 2 days a f t e r defecation and emerged 9 to 13 days later* No cocoons were formed, Mean emergence from 100 parasita'zed cocoons was T e 7 4 (range, 4-22), The sew r a t i o was $ , I 6 Females per male* Males emerged up t o a day before the f e - males. A male would cut an e x i t hole i n the host cocoon, emerge, and wait near the opening f o r a fernale t o emerge, T h e n , beat-ing h is wings intermit- t e n t l y , he would fallow her, climb on her back, tap her head w i t h his an- Semae, and eventually attemp% copu la t ion , Males usually remounted u n t i l successful. Copulation lasted 15 t o 20 seconds. A d u l t males l i ved 9.8 days while females l i v e d 30.6 days i n the laboratory when fed sugar water.

The hast relatiaansh9"ps o f E, are often complex and potent i - a l l y confusing since the range o f i i t i e s spans t h a t o f a primary paras i t e to possibly t h a t of a semi-scavenger, On i t s mast important hosts, the pine sawflies, i t can act d i rec t ly as a primary pa ras i t e , or indirectly as a hyperparasi te and Coppel (1 960) reported M. parasi te o f the sol i tary wasp, Ancistrscerus - h a b i t i n g empty cocoons o f D, I n ssm S, i t can even compete successfully i n a & l t i pa ra s i t i c situation.

1 reported "&hat & Mer t ins (1967) re

1 s However, i s isc cons in collee from a g roup o i t i z e d D. sirnil i s cocoons he exaniined, M. was a t l eas t associated w i t h another parasite species i n seven cases,

Mertins (1967) a l so reported t h a t E* could compete success- Dur ing the sum~er f u l l y i n a multiparasitic relationship on g, --

months, adults of ye emerge constantly w i t h the fernal es oviposi t- ins in new s a w f l y cocoons, so t h a t the parasitism of sawflies i s cumulative a s h e surnmer prbgresses (Morris et a l . 1937). If t b e population of M.

increases sufficiently by the time t he second generation of saw- ms cocoons, M , sawfly cocoons a % ready para-

other indiFfdua o r by some other species of p a r a s i t e (MertJns 19671, rn, muStiparasitism, o r super- parasitism may occur, M, rvives i n e i t h e r case, Mertins (1967) seldom fzun i n cocoons t h a t had a l s o been attacked by one ins served b o t h vesicularis (Retzius) and M, the same hos t cocoon, In addition, Mertins (1967r a

rom the same hos t m o o n ~ i & h E, a r r i i c t g z , t&g %Sohard, was a

In addi t ion, M, may be a sewi-scavenger, H e r f i n s (1967) re- or led , in his work-on the Wisconsin parasites o f D. 1 f s that E. den-

developed once on the remains o f a saw f l y IeFt a f t e r the de ichneumon-% parasd te , He was uoakrle to discern whether ipi,

ttaeked the sawfly cocoon d u r i n g c r a f t e r the deve:spment s ichneumonid, S h e the time o$ parasitism by " was t he crucial f ac to r , he was unable to eliminate the possibility t h a t the relalionrhip had been multiparasitic origina7ly,

i l e in the host relationships i t can assume, a t ionsh ip proves t o be, the extent to which t h e

an successfu79y a t t a c k pine sawflies i s determined 7 or d u f f where the h forms i t s eocaon~ D. sirnilis, and E, are heavi l y para-

ey o c c u r inthe same h k, sertifer i n ut N. ser t i fer i s seldom attacked on e i t h e r con-

t i n e n t , In the f i r s t record of on N. sert ifer i n Canada, only one h o s t cocoon produced the parasi layson d subsequent re- ports are similarly meager ( G r i f f i t h s et a l . 1971 ; ~ ~ o n k 1964).

M, was n o t effective against g. sertifer because the para- s i t e caul ch t he cocoons o f 5. s e r t i f e r i n t h e m i n e r a l soil (Lyons 1964). Lyons (1 964) reported that fewer than 3 percent o f t h e sertifer a t Ckadsworth, Ontario, coeocnel above the mineral so i l , a1 t h o u g h a flicting report e x i s t s (Morris and Cameron 1935). i n contras t , D.

and - D, sirnil i s a1 l cocoon above mineral soi l ; i l i s , on-st tree and nearby shrubs; and gT - -

i n The surface l i tker (McGugan and Coppel 1962),

The extent to w h k h E, parasitizes a given host may a l s o de- pend on physical and physiological characderist-ics sf the host cocoon o r host p repupa , T h e parasite is ab le to distinsuish between the larch saw-

(Har t ig) , which i s not known t o be a host o f M. hich i s successfully parasitized

nd Fedde 1972), She ability o f M, e hosts, w h i c h are v e r y similar Tn aoDearance, is * n

ev ident a t both the cocoon and prepupal levels,

SUMKARY AND SCGGEST%ONS FOR FURTHER STUDY

Most pub1 ications referring to & pertained primarily t o i t s hosts o r distribution until 1937, when Morris ed a l , described the ~arasite along w i t h athers o f Ee i n Europe. When the importance o? E.

in the control af 9, sirnilis in the United States and Canada was -- real ized, the number of hos t and local i ty records o f t h i s insect mushroomed. MerLins (3967) recently made %Re second major contribution t o kn~wledse

w

about the l i f e history and host relationships sf - M,

is potentially useful Par b i d s g i c a l control o f ocoon above ground, i n the southeastern United

nal s t u d i e s are needed on the parasite" sa$ility t o function i n such a capacity. There i s a need for more understanding o f the physio- 4sg$caB responses do environmental factors for t h i s parasjte, as well as s$ natural food o f the adu l t s and the parasite's behavior and ability t o select

is t o be employed suceessfu% 1.y against sawflies of uch more about the bielogy s f the parasite must be

understood,

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