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

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

by Vicki H. Fedde

Agriculture

S ti7 t ;or1

(7

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

Vicki H. Fedde, Associate Entomologist Forestry Sciences Laboratory Research Triangle Park, North Carol ina

Monodontomerus dentipes (Dalman), a European parasite of cocoons of pine sawflies, has been a key factor in the control of the introduced pine , similis (Hartig), in the United States and Canada. Several other Diprion and spp. have also been reported as hosts, and the sawflies which occur in the southeastern United States are possible targets for control by E. dentipes. During a recent study of the development and emergence of the parasite, a comprehensive survey of the literature on M. was made. The purpose of this review is to aid others in studying o that its usefulness as an agent in the control of Neodiprion spp, can be enhanced.

The original description of M. dentipes is generally credited to Dalman (1820) who described both sexes of the adult. A number of sources (Hartley 1923; Faure 1925; Sitowski 1929; de Fluiter 1933; Mokrezecki 1933; Scheidter 1934; Gahan 1941 ; Bucher 1948) incorrectly 1isted the description by Boheman (1833), based on a single female as having priority, Dalman named the parasite * dentipedis and placed it inthe family Pteromal- idae, tribe Pteromalini, The species was transferred to Monodontomerus (Walker 18471, which at the time was in the family Callimomidae (Westwood i833). The name Call imomidae has been pemanentlj suppressed (peck 1963). Monodontomerus has also been listed in the family (Howard and ut is presently in the family (Gahan 1941 ). The is synonpous with Torymus vicicellae Forster (Walker neus Provancher (Burks 1963).

Several keys are available for the diaanosis of M, dentioes. Adults were separated fFom those of other ~onodontomerus spp.-b$-1 941 ) and Steffan (1952). A comorehensive 1ist of references by species of Monodon- tomerus was b) Burks (1963). Since there iLlittle overlapping osts of M. dentipes and other Monodontomerus spp.,.. - identification is simplified through the identification of the host, especially if the host is a diprionid sawfly (table 1). In North America, M. * Ashmead and Me denti~esare the on1 v Monodontomerus SDD. which attack saw- flies, The Gl af gLfe mkkican sawfly is in Canada, where it was released ayson and Finlaysen 1958b), In the abse --GUS on D, hercvniae, id ~rshablvhas not become established. M. dentioes \Has sepa- othbr paraiites of pine sawfl ies by Morri~e 371, Fin- layson (1960a, 1960b, 1962, and 19631, Mertinj (19671, and Mertins-and Coppel (1971 ) .

'~meno~tera: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 Braconidae Chalcididae Cyni pidae Diprionidae Ichneumonidae Megachil idae Sphecidae Tenthredinidae Vespidae

LEPIDOPTERA Arctiidae Lasiocampidae Liparidae Lymantri idae Olethreutidae Saturni idae Tortricidae Zygaenidae

D IPTERA

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 of the internal and external anatomy of both sexes.

GEOGRAPHIC DISTRIBUTION AND IPS RELATION TO HOSTS Generally, the distributi depends on the presence of sawflies of the family Dipri ther groups are repre- sented as host 2). Whe occurs, the hosts are most commonly spp. and ther Ifmenoptera, Dip- tera, and Lepidoptera are also parasitized, but far less frequently.

Table ?.--Recorded hosts of Monodontornerus

France Rosenburg 1934

Braconidae

SP a Poland Thompson 1944

-A, u-o'igmeratr;~Laa France Fawe 4925, 1926; Thompson 1944 Italy Thompson 4 444 --Meteorus versicolcr Wesmael HoI land de Flui ter 1933; Thompson 3944 Cynlpidae Skieden Balman 1820; Boheman 1833; Howard and Ashmead 1898 Digrianidae

SPP' Poland Mo krez ecki 1 933 USA: New York Viereck 1928; Gahan et al. 7928 --D. frutetorum [Fabricius) Canada: Ontario Raizenne 1957; McGugan and Coppel 7 962; Griffiths et al. 1971 C~echosloi~akia Finlayson and Finlayson 1958a Canada : New Brunswick Baird 1938 Nova Scotia Baird 1938

Ontario Finlayson and Reeks 1936; Baird 1937, 1939; Peck 1951 ; Finlayson 7960b; McGuijan and Cappel 1962 Quebec Finfayson and Peeks 1936; Baira 7938; Briand 1949; McGugan and Cappel 7952 Ckiria Peck 1963 Germanj Pecr I963 Hull and Peck 1963

Poland Peck I 963 ESP: Maine Procter 9 938 Vermont Dowder; 7958 D. (Klurjj Czechosiavakia Morris et ai. 1937 continued

See footnotes at end cf table. (Dafmn) (continued)

Scheidter 7 934; Thompson '1 944

Holland Ritzenia Bas 1919; Thompson 1944

Pol and Sitowski 1429; Thompson 1944 Sweden Howard and Ashmead 1898

Gzechosl ovakia Molubajiv 1934; Morris et a?. 1937; Thompson 1944; Finlayson and Finlayson 1958b Gemany Morris et a?. 1937

Europe (general ) Gahan 1947; Dowden 1949

Canada: Ontario Baird 1939; Gahan 1941 ; Bscher 1948; Dowden 1949; Raizenne 1957; Sippeil et a?. 1960; Finlayson 7962; McGugan and Goppel 1962

Quebec Baird 1938; Finlayson 1962; McGugan and Coppel 1962

China Peck 1963

Germany Thompson 1944

Hol 1and Hart1ey 1923; Thompson 1944

Hungary Eahan 1941; Peck 1963

Poland Sitowski 1929; Finlayson and Finlayson 195Ba

Sweden Howard and Ashmead 1898

USA: Connecticut Britton and Zappe 1918; Gahan 1941 ; Dowden 1949 Maine Gahan 1941; Procter 1946

Michigan Gahan 1941

New Jersey Weiss 1917a, 1917b, 1918; Hartley 1923; Gahan 1941; Dowden 1949, 1962

New York Gahan 1941

Ohio Gahan 1941

Pennsylvania Hartley 1923; Gahan 1941; Oowden 1949, 1962 Wisconsin Shenefel t and Benjamin 1955; Dowden 1962; Mertins 1967; Mertins and Coppel 1971 (Harris) Europe (general J Baird 1944

Canada: Ontario McGugan and Coppel 1962

New Brunswick Baird 1944; McGugan and Coppel 1962 --N. excitans Rohwer USA: Florida Wilkinson, personal cmunication --N. lecontei (~itch)' USA: Fforida Wi1 kinson 7969 -N* (Norton) Canada: Ontario Raizenne 1957; McGugan and Coppel 1962

Rani toba Baird 1941 ; McGugan and Coppel 7962

continued

See footnotes at end of table. Taal e 2,--Recorded basts 04 ~odantmerus=I& (Dalman j icontirueci 1

I So~rce

USA: VSrginia &rwz 1971 --k, swainei Middleton USR: Wiscons~n B~cCerand Benjam*~7964

USP: Wisconsin Mertins 1367; Mertins ~ndCappel 1971 Tetrastickus xanthgps 7 Aiistrf a Seitner 1927

Exenterus anictorius JSA: Wisconsin Mertins 1957; Mertins and Coppel 1971 Mertins 1967; Mertins ano C~ppel7971

spa7 Austria 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

"c~or;ough) " GSA: Idaho Eeddrd 1938

Moll and de Fluiter 1933 P~~theLriadispar ii.j r;ew Enijlani? Burgers 4915 p- Pieridae

Fjer~~brass7cze L. France P--

See foetrotes at end of tati~, HOSES Source ---- Tortricidae

3' 3' L, France pose~%rg1534 ---Tortrix vsridana (Girard] Sweaen Loward ard Fshqeai 1898 Zygaenibae 1-aa i . Sweeien noward z~dffshead 9898 Z, occStansca de V.illers Fra~ee Hartley 1922

' ", der~tspr;; ns nypErparac?t7c cr wcf~r~~ell;~ i. thrailzb t ttaqicd, 5. dent~p~raIS -yperpa%iizit-c 01 ?~er~sbrass-cae L. kpr3:gh 5, $&T?)"KYL~- " E, Cent~pesis kyperparasit~cGP z_?l-ps& re?s I. "c~r3,qF., z*SC~IW- (de FIUIt2~ 4333,. Earlj !North drner~cankeq~rerces t:, g, =cjnme r-rstakenl; -iden?f;eC less f;, @;jt~~urn(#P~L!IB~ and Copre: 19621,

------s g, a~d COP~UISI~O~. was released agalns't theze ~~st;lr a bl~;03l~alcclbro; effort of the iiSDC csrest ve beer made to dale. is hyperparasi"c-;c: on De~drol-a "srrcug$ tb cles {Seitre. 19211, was reported as a prima;-;. paras~teof 4, ~r)l?as~t:ngempty cciicuo~s of 12,

M, is native to the Palearctic regioq of the world and dis- tri butzd ly in the Holarctic region (table 31, blthcugk -M, --den- * Is found th~oughsestthe Palearctic region, only a single locality exists for China (Gahan 1947), -M, has been reported throughout northern and central Europe, R om France and Austria 1 ist nonsawfly hosts exclusively. Peck (1961) be1 ieviid these hait: suq- gested ite was identified in~orrociila*.: ti 5. aer~ijshzlke! ,

not E, In Europe, Dip~i~~pii,l (1 2 -- .AA."-~zolvfowlr;i:am---- (&rtiqIi P are the most frequently pawas-itlzed hests,

In the Bearctic region, Ee occurs in the United States ana Canada, primarily east of the Gre M, &isJ$j= was s, acr;Sdenta" !Y introduced into the United Stater as a parasiTe 6-f the sawfly, ). &m""l;s, which subsequenuy became the most frequefitly attacked host in this coun- dry. gLg, was fa"rst reported ip Co~necticutaaoucd 5910 (Britton 19159 Fr-i Zappe 1918) and shortly after i-~New Jersey {Us'ss 11%17a, 19]7b, 19m; Hartley 132% &hac "GdI: EI.~U:<.:. i44;), z:f 1322z U, w-- ss"mi1-8"~was effectively parasitized cj -"i"", denljpes---- T-.i a aik;,or f-Jeslaej3~ near Philadelphia, Pennsyl va:ila (t;ar::!i.j { i27j~~~~-ct~vn2rkeC the rs~ehern- most extension of the parasite's distribution at that time, in later years, the parasite was reporte e same host ifi !4atng, t-!

Ccsntineot and Hosts Source country

ASIA

China D~prionhercyniae jiiartigj Peck 1963 --O, sinilis (~artig) Peck 1963

EUROPE (general 1 (I--1 Steffan I952 --3, sirnilis Gahan 1441 ; Giowden 1749 Sreffan 1452 --a, sertifer Geoffror Austria Dendral imus pini L > Seitner 1916, 1927 Tetrastichus (~atzebur~)' Seitner 1921 sp." Seitner 1927 Therion (~ravenhorst)' Seitner 1927 --Theronia atalanta (~oda)' Seitner 7827 Rhogas sp." Seitner 1927 Finlayson and Finlayson Czechoslovakia e. frutetorum (F.) 295% -8. Kl ug Morris ei al. 1937 D. (Hartig) Kolubajiv 7934; Morris et al. 1947; Thompson 1944; Finlayson and Finlayson 1958a France Faure 1925, 1926; Thompson 1944 Rosenburg 1934 Cydia pornonella L. " Rasenburg 1934 --Pieris brassicae L." Faure 1925 acci tanica de Yillers hart1 ey 1923 Germany Peck 1963 Diprion Scheidter 1934; Thompson 7 944 D. Harris et al. 1937 Hol 1and -D. hercyniae Peck 1963 --D. sirnilif Hart1 ey 1423 ; Thmpson 1944 Diprion pinj Ritzema Bos 1919; Thompson 1944 Meteorus ~ersicolorMesmel de Fluiter 1933; Thompson 1944 Stitpnotia salicis t. de Fluiter 1933 continued

See footnotes at end of table. Table 3.--Palearetic distribution of Honodcntornerus (Dalman) (continu&)

Continent 1 and Hosts country

Hungary --D, simrlis Gahan 1941 ; Peck 1963 Italy Thompson 1944 P~Iisnd T~O~OSOG1944 Mokrerecki I933 Mcikrerecki 1333 Pack 4 963 Sitowskr" 1929; Thompson 1944 --D. sirnilis SStowskl 1924; Finlaysop and Ffnlayson 1958h P. brassicae Mokrerecki - -"- 1993 Sweden Cynipid wasps Balmn 3820; Bobeman 4833; Howard and Askmead 1898

5 Howard and Ashmead 1898 --13. sirnilis" Howard and Ashead 1898 e." Howard and Ashead I898 --Yortrix vtridana (Giraudj Howard and Ashead 7898 L. Howard and Ashead 1898

In,- is a hyperparasite of Dendroli~us@ ha through these species [Seltnsr 9927)- is a nyperparii-stke of P. 3 - E. is a hyperparasite of ge;, 414. is a hyperparasite of 5. g, ve r jde Flutter 19331, 5~s " AS sirnilis Wartig, " AS L, Table 4,--Neitrctic distributlcnn of Panodentmerus dentipes (Dalnan)

Source

NORTH PMERICA

(general j Peck 1957 Peck 7951

--D. sisilis Idaraig] Peck "91 Cjiptera Peck 7951 j~e~onnoughjZ Peck 1957 Peck 1951

Burgess 1915

Ganzda (genera? j -P. sfrntlis MrGdgan and Coppel 1962 (Geocfroy 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) Ba~rd7944; McGugan and Cappel 196% Nova Scotia B. Ontiar~c -G. --frutetorum (F*) 6zirenne 1957; McG~gan and Co~pel7962, Griffiths et ai. 1971

Finlayro~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 et al. 1960; Finlzysan 7962; McGugan aqd Cappel 1962 -N. -- abiet-is k. pineturn (Norton:

N. banksianae Baird 19dl Finlayson 7962; -N. ser"iifer 1960a; MsGugaq and Coppel Lyons -- 4454; Grirfiths ~t a:. 1477 7959; --a -buoliana ~SchifFerm~ll~r) Miatsop and Arthur McGugar: and Coppel 1062 Quebec Finlayson and Seeks 1930; Baird 193e, 1939; Briand 1949; McGugan and Coppel 1962

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

See Pcotnotes at end of table. Table 4.--learctic d~sdribud;an of ?"ksnodontorre~ (0almnj (continued)

Local r ty Hosts Source i

Connecticut f. sfmilis Britton and Zappe 1918: Britton 1926; Gakan 1941: 5ouden 79d9

Maine

Meiss 1917a, 1937b, 1978; Hartley 1923; Gakan 1943; Dowden 1949, 1962 Mew '{orb: sop, Viereek 1928; Gahan et al. 1928 --D. simills Ohio --D. sjrnilis Gahan 1941 Pennsyivan~a -D. --s-imi'iis Hartley 7923; Gakan 1941 ; Dowden ?945, 1962 "imont n,

Virginia N a W.isconsin --Ancistrocerus (Causs~~re;"

Shenefelt ard Benjamin 1955; Dcwden 7962; Yerti~s 1967; Wertins and Coppel 1971

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

'AS Henerscamn ~seudotsugata. 9 -- Early North Amerfcan refererces to &, and Coppel 1962). 3~,dent-i~es was released against these hosts ir 1965-1979 through the bltilogrcal control program of the USDA ForesT ~xeiit has nut been recovered to date, *M.-- was listed as a pr-inary parasite tigr.rs cornon Irrhaafeants of cocoon; cf -6, --s~milis,

In Canada, E, has been released against pine sawflis in biological control p n Ontario, Quebec, Manitoba, Nova Scodia, Mew Brunswick, and Prince Edward Island (table 4). In Ontario, E. is camon and more frequently at$acked than any ather potential host, How- ever, in Quebec and the Haritiine Provinces, D, is the most abun- dant sawfly and the most heavily parasftlzed: Generally, in the Nearctic region, M. is concentrated in coastal regions, Yke distribution sf the pa tends along the Atlan- tic coast from Pennsylvania to Quebec, and westward along the Great Lakes ta Wisconsin and Manitoba. Recent releases against sawflies in the south- eastern United States (Wil k 9; Morris 1971 ) have provided an oppor- tunity for the range of Me to be extended southward as far as Florida, however, estebl7shment has not yet been determined. HOSTS

Included among the known hosts of M. are six families of , one family of Diptera, and five farnil~esof Lepidoptera (table 2). Among the various farnil ies that have been reported are: Diprionidae, Braconidae, Ichneumonidae, , Vespidae, and Cynipidae in the order Hymenoptera; Tachinidae in the order Diptera; and iiparidae, Lpantriidae, Zygaenidae, Tortricidae, and Pieridae in the order Lepidoptera. throughout the distribution and Pive species of as potential hosts, are

In the Hearctic rion species and Pive

United States, In Nor niae are the most fre-

the parasite has been reported iKfr as we11 (table 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 of nonsawfly hosts are insignificant when compared numerically to records of sawfly hosts portant factor in the host relatia~shipsof H. been recorded on three species of Braconidaedn Ichneumonidae, Similarly, only a few cases of

tera in &o~LhAmerica are from Id

"risr to 1941, when B was correctly identified by Snitk (1941) 2s the species involved in Canadian infestations of this sawfly were believed to be conspecific wth those of the Eur ies D. s such, several early sources incorrectly refer to 6, as %). {MsGuga Morris et a1 , (1937) and Mertins (3967) are the only authors who have touched on a variety of topics related to the life cycle sf the parasite, In their studies, they included mating and oviposition, the eggs, maturation of larvae, pupation, emergence from the host cocoon, longevity of adults, and host relationships of M. Mertins and Coppel (1971) revised the material pertaining to ho jon- ships that Mertins (4 provided, They omitted much sf the detailed information about E* that Mertins (1967) had included,

The stages in the 1 ife cycle of M, ere described initially by Morris et a1 . (1937). These workers s" n the laboratory so their description was Id and on field-collected material, The fe five or six (maximum, 15) eggs in a sing externally on the prepupa sf the sawfl vouring it cofipletely, The larvae matured after 3 to 4 weeks, M, overwi ntered as prepupae, or, if weather were favorable, develope tely into pupae, In April, when overwintering individuals pupated, the pupal stage lasted 3 weeks, During the summer, when other individuals pupated, the pupal stage was shorter, Four or five (maximum, 12) adu9 ts of M, emerged per host cocoon. Emergence of the first generation of a&l ts occurred in May, al- though adults were found in the field into September. A full month elapsed between the emergence of the First and last from one cocoon,

Mertins ' (1 967) observations agree essential ly with those of Morris et al. (1937). However, Mertins provided additional information about the Wisconsin parasites of -0, sirnilis. He reared Ma from field csl- lected cocoons of Dm and studied the bgha longevity of the para er unspecified laboratory conditions. M, took 28 days to develop from eggs to adult% Larvae hatche8 with- in 3 days of oviposition, fed on the host for 8 days, then were quiescent Psr 3 additional days before defecating* Afterwards, the larvae tended to position themselves in a single layer around the host cadaver and cocoon walls, a71 facing in the same direction, They pupated within 2 days after defecation and emerged 9 to 13 days later* No cocoons were formed, Mean emergence from 100 parasita'zed cocoons was Te74 (range, 4-22), The sew ratio was $,I6 Females per male* Males emerged up to a day before the fe- males. A male would cut an exit hole in the host cocoon, emerge, and wait near the opening for a fernale to emerge, Then, beat-ing his wings intermit- tently, he would fallow her, climb on her back, tap her head with his an- Semae, and eventually attemp% copulation, Males usually remounted until successful. Copulation lasted 15 to 20 seconds. Adult males lived 9.8 days while females lived 30.6 days in the laboratory when fed sugar water.

The hast relatiaansh9"ps of E, are often complex and potenti- ally confusing since the range of i ities spans that of a primary parasite to possibly that of a semi-scavenger, On its mast important hosts, the pine sawflies, it can act directly as a primary parasite, or indirectly as a hyperparasi te and Coppel (1 960) reported M. parasite of the sol itary wasp, Ancistrscerus- habiting empty cocoons of D, In ssm S, it can even compete successfully in a <iparasitic situation. 1 reported "&hat & Mertins (1967) re is isc cons in collee 1s However, from a group o itized D. sirnil is cocoons he exaniined, M. was at least associated with another parasite species in seven cases,

Mertins (1967) also reported that E* could compete success- fully in a multiparasitic relationship on g, -- During the sum~er months, adults of ye emerge constantly with the fernal es oviposi t- ins in new sawfly cocoons, so that the parasitism of sawflies is cumulative ashe surnmer prbgresses (Morris et al. 1937). If tbe population of M. increases sufficiently by the time the second generation of saw- ms cocoons, M, sawfly cocoons a%ready para- other indiFfdua or by some other species of parasite (MertJns 19671, rn, muStiparasitism, or super- parasitism may occur, M, rvives in either case, Mertins (1967) seldom fzun in cocoons that had also been attacked by one ins served both vesicularis (Retzius) and M, the same host cocoon, In addition, Mertins (1967r a rom the same host moon ~i&hE, arriictgz, t&g %Sohard, was a

In addition, M, may be a sewi-scavenger, Herfins (1967) re- orled, in his work-on the Wisconsin parasites of D. 1 fs that E. den- developed once on the remains of a sawfly IeFt after the de ichneumon-% parasdte, He was uoakrle to discern whether ipi, ttaeked the sawfly cocoon during cr after the deve:spment s ichneumonid, She the time o$ parasitism by " was the crucial factor, he was unable to eliminate the possibility that the relalionrhip had been multiparasitic origina7ly,

ile in the host relationships it can assume, ationship proves to be, the extent to which the an successfu79y attack pine sawflies is determined 7 or duff where the h forms its eocaon~ D. sirnilis, and E, are heavi l y para- ey occurinthe same h k, sertifer in ut N. sertifer is seldom attacked on either con- tinent, In the first record of on N. sertifer in Canada, only one host cocoon produced the parasi layson d subsequent re- ports are similarly meager (Griffiths et al. 1971 ; ~~onk1964). M, was not effective against g. sertifer because the para- site caul ch the cocoons of 5. sertiferinthe mineral soil (Lyons 1964). Lyons (1 964) reported that fewer than 3 percent of the sertifer at Ckadsworth, Ontario, coeocnel above the mineral soil, a1 though a flicting report exists (Morris and Cameron 1935). in contrast, D. and -D, sirnil is a1 l cocoon above mineral soil ; --ilis, on-st tree and nearby shrubs; and gT in The surface l i tker (McGugan and Coppel 1962), The extent to whkh E, parasitizes a given host may also de- pend on physical and physiological characderist-ics sf the host cocoon or host prepupa, The parasite is able to distinsuish between the larch saw- (Hartig), which is not known to be a host of M. hich is successfully parasitized nd Fedde 1972), She ability of M,

e hosts, which are very similar Tn aoDearance,* n is evident at both the cocoon and prepupal levels,

SUMKARY AND SCGGEST%ONS FOR FURTHER STUDY Most pub1 ications referring to & pertained primarily to its hosts or distribution until 1937, when Morris ed al, described the ~arasite along with athers of Ee in Europe. When the importance o? E. in the control af 9, --sirnilis in the United States and Canada was real ized, the number of host and local ity records of this mushroomed. MerLins (3967) recently made %Re second major contribution to kn~wledse w about the life history and host relationships sf -M, is potentially useful Par bidsgical control of ocoon above ground, in the southeastern United nal studies are needed on the parasite" sa$ility to function in such a capacity. There is a need for more understanding of the physio- 4sg$caB responses do environmental factors for this parasjte, as well as s$ natural food of the adults and the parasite's behavior and ability to select is to be employed suceessfu%1.y against sawflies of uch more about the bielogy sf the parasite must be understood,

LITERATURE CITED

7937. Biological coctrol of the spruce sawfly Hartig, Pulp Pap, Mag, Can, 32: 311-312, 314-375,

Bafrd, A, B, 4938, Parasite and predator liberations in 1937, Can, Insect Pest Rev, 16: 77-754,

Baird, A, B, 1939, Parasite and predator liberations in 1938, Can. Sfisect Pest Rev, 17: 401-128,

Baird, A, B, 1944, Parasite and predator liberations in 1946, Can, Insect Pest Rev, 79: 94-725,

Baird, A. B, 1944, Parasite a~dpredator liberations in i943, Can, Insect Pes? Rev, 22: 147-138,

Becker, G, C,, Jr,, and Benjac~~,3, M, 4464* Notes on insect parasites a~dpredators cf the Swaine jack-pine sawfly in His- consin, J, Ecsn, Eastoatcl, 57: 355-357,

Eedard, W. 5, 1936- An aonotated list cf tke insect fauna of Ceuglas fir (?seud@tsuga mucrnnata Rafi~esq2e)in the nwthern Pocky Mcuntain region. Can, Entorn~l,70: 196,

tiviand, k, J, 4 Spruce sadfly parasite 1 rvestigaticns, Park Reserve, St, FI exzndre, Kawouraska, Que~ee. Annu. Rep. Ouekec Sot, Prot, Piants 3Q: 54-59, Britton, W* E. 1915, A destructive pine sawfly introduced from Europe. hartig, J. Econ, Ento~ol,S: 379-382.

Britton, W, E, 1920, Check-list of the insects of Connecticut, Conn. State Geol, & Rat. Hist. Surv. Bull. 31, 397 pa

Britton, W. E,, and Zappe, M. P, 1918. The imported pine sawfly ) s-im'ile Hartig. 77th Report of the State Entonolaqist of Ccnaecticut for the year 1917, Csnn. Agric. Exp, Stn.

Bucher, G. &. 1948, The anatow sf Boh., an entomophagous Chalcid. Can, d, Res, 26: 230-28,.

Burgess, A, F. 1915, Report on the gipsy moth work in New England, U-S, Dep. Agric., Agric. Full, 204, 32 p.

Burks, Be D. 1963. The Provancher species of Chalcidoidea (Hymenoptera), Can, Entom01 . 95: 7 254-1 263. Cappel, H, 6, 1960, Empty cocoons of the introduced pine sawfly, sirnilis, as habitat niches For , Ann. Entamol, Sac. Am. 53: 847-848,

Dalman, J, V. 1820, FOrs6k till Uppstatlning af Insecx-familjen Pderomalini, i synnerhet med afseende pa de i Sverige Punne Arter. Mongl, Sven. Vetensk, Acad. Handl. 47: 123-182.

Dowden, P, B. 1949, , wasps, and parasites, p. 591-636, &F. 6, Craighead fed.], Zn- sect enemies of eastern forests. U,S, Dep. Agric., Misc, Publ. 657, 679 p,

Dowden, P, Be 1962, Parasites and predators of forest insects liberated in the United States through 1960, U,S, Dep, Agric,, Agrie. Handb. 226, 70 g.

Droom, A, S., and Fedbe, V. He 1972, Discriminate host selection by kiionodontonerus Environ. Entomol. 1: 522- 523.

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Faure, 3, C. 1926. Contibu"eion $ S"6tude dhn eomplexe biologique: la pigride du chou (~iecris brassicae, L. ) e% ses parasites hp&noptdres, (Abstr,) Rev, Appl , E Ser. A, 14: 516,

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