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1 NAME OF AUTHOR/NOM DE L'AUTEUR avid-\. Gillespie

T<;EOF THESISITITRE DE LA THESE Classification of ~inal-instar Larvae of the

a (: )

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Ottawa, Canada CLASSIFICATION OF FINAL-INSTAR LARVAE OF THE ICHNEUHONINAE (HYHENOPTERA: ICHNEUNOWIDAE) . I

David Roy Gillsspie B. Sc., Simon Frassr University, 1976

A THESIS SUBHITTED IN PARTIAL FULFILLMENT OF

THE REQUIREHENTS FOR THE DEGREE OF HASTER OF SCIENCE in the Department

Biologics 1 Sciences

i-2 @ David Roy Gillaspie 1973

SIMON FBASER UNIVERSITY f'

- - Ma- rig h+s- reserved. - T hss--k&sisTtaay -neB be reproduced in whole or in part, by photocopy 02 other means, without permission of the author. PARTIAL COPYRIGHT LICENSE

I hereby grant to Simon Fraser University the right to lend my thesis, project or extended essay (the title of which is shown below) to users of the Simon Fraser University Library, and to ake artial or . P - !- single copies only for such users or in response to a requestb from the library of any other university, or other educational institution, on its own behalf or for one of its users. I further agree that permission d for multiple copying of this work for scholarly purposes may be granted by me the Dean f Graduate Studies. It is understood that copying orbublicationof 1is work for financial gain shall not beallowed without rnowritten permission.

Title of Thesis/Project/pnded Essay L Classification of Final-instar Larvae of the Ichneumoninae< *

(Hymenoptera: Ichneumonidae) 3 APPROVAL

Name : David R. Gillespie

Degree : Master of Science

Title :' Classification of Final-instar Larvae of the Ichneumoninae (Hymenoptera: Ichneumonidae) +

Examining Committee:

Chairman: . Robert C. Brooke

-7 - " Professor T. Finlayson -

-, 6.M. Mackauer

, I- Dr. B. ---P. Beirne

---.r. - *- " - Dr. H. R. warthy, Non-Supervisory Committee Examiner ,

#' Date ABS TRACT

Cephalic structures and skins of final-instar larvae- of 43 spscies of 24 genera of the subfamily Ichneumoninae 8, (Hpmenoptera: Ichneumonidae) are described and illustrated and keys are given f~rthe separation of tribas, genera, and

species. A further 2 species available for examination but . 5 previously described and illustrated elsewhere are included in the keys for a total of 45 species.

. Larval characteristics supported most of th-ibal, \ - gensric and specific entities defined on thi basis of adult 0 characteristics. In no case was there serious disagreement

between - the adult and larval classifications.

Larval characteristics place species of the tribe

Phasogenini as the most primitive of the subfamily and . suggest a close relationship between them and specibs of the tribe Ichneurnonini subtribe Cratichneumonina.

The tribe Listrodromini is considered' to be close1y

related to the tribe Ichneumonini, probably through a common

ancestor parasitizing butterflies. 1P

iii 0, &a a.' ACKNOWLEDGE BENTS

I thank my senior supervisor, Prof. T. Finlayson, for her advic~and consistent support during the course of my

research and writing of my thesis. I also thank the other . . memhers of my supervisory committee, Drs. J. P.44. Nackauer and B,P, Beirne for their advice acd helpful criticism.

I thank Dr. J.R. Barron, Biosgstematics Research

Institute, Ottawa, Ontario, Dr. H.G. Uylie, Research

Station, Agriculture Canada, Winnipeg, Manitoba, and Dr. H. Doganlar, Faculty of Agriculture, Ataturk University, Erzazum, Turkey, for the loan of specimens used in this study*

For advice and helpful information provided to me, I a thank Dr. J.R. Barron; Dr. H. Townes, American

a Entomological Institute, Ann Arbor, Hichigan; and ar. G.H. Beinrich, Dryden, Maine.

Last, but not least, I thank my wife, Beth, without

whoss constant support and encouragement, not hc mention

typing skills, this thesis would not be a reality. , - . -

TABLE OF CONTENTS Page . . EXAMILJIRG COMHITTEE APPROVAL.'. 0. 0. 00 .a. 0 0 0 0. 0 0.00.. 0.011

LIST OF TABLES...... xiii

Importance of Larval ...... 3 Ganeralized Final-instar !lorphologp of the Ichneumonidae .....~...... 5 Taxonomy of Ichneumoninae...... 10 Biology of Ichneumoninae...... 13

GENERAL HORPHOLOGY OF FINAL-INSTAR LARVAE OF THE ICHNEUHONINAE..o...o.eooooooo.ooooooooooo ooeoooooo22

KEY TO TRIBES OF ICHNEUClONINAE.. 0 0 0 0 .0 0 0 0 0 0 0 40

Key to genera of Phaeogenini...... 45 Genu-s Phaeogenes Wesmael...... 47 Key to species of Phaeogenes ...... 49 Genus Dicaelotus Wesmgel...... 56 Genus Diadromus Vesmael...... 58 Genus Bethecerus Uesrnael ...... 60 Summary of Pbaeogenini...... 62

Key to genera of Platplabini...... m.mmmm.mm63 Genus Platylabus Besmael ...... 70

Key to species of Platylabus...... m..m..m.m.m 70

Genus Cyclolabus he in rich...... ^ m.m.m.... 75

TRIBE LISTRODROHINIm.m.mm..m~mm~mm*mmmmmmm mmm mmmmmmmm.78 enu us Anisobas uesmael ...... 78

Key to genera of tchneuaonini ...... 85 Gsnus Hoplis~enusGravenhorst ...... 90 Key to subspecies of morulus ...... 91 . Genus Vulgicbneunon Heinrich ...... 96 Genus Thonson ...... '...... 99 Key to sp'ecies of Cratichneumon ...... 100 Genus.Pattodoides Beinrich ...... m...... I23 Genus Tischbein...... 126 Key to species of Aoplus...... '127 Genus Eutanyacra Cameron ...... m..m...... i 131 Genus Diphyns ...... 135

vii Genus Limerodops Heinrich ...... 138 + Genus Spilichneuaon Thomson ...... 141 Genqs Thyrateles Perkins ...... 144 Key to species of Thyrateles...... 145 Genus Linnaeus ...... em...m..J"0 . Key to species of Ichneumon ...... 151 Genus Orgichneumon Heinrich ...... 166

Key to genera of Protichneumonini ...... 182 Genus Coelichneumon Thomson ...... 184 Key to species of Coelichneumon ...... 185 Genus Syspasis Townes...... m189

Subtribe Callajoppi3a.~...... 197 Key to genera of Calla j oppina ...... 198 Genus Callajcppa Cameron ...... 198 Genus Conocalama Hopper ...... 201 Key to species of Conocalama ...... 201 s ubtribe Trogina ...... 206 Genus Trogus Panzer ...... 205 Key to specis of Trogus ...... 209 Z

DISCUSSION AND CONCLUSIONS. em em a216

1 viii FIGURE LIST Page

Fig. la. Diagram of the cephalic structures of a final-in star larva of an ichneumonid showing names of the sclerites...... a. -7

Pig. Ib. Diagram of the cephalic structures of a final-instar larv of an ichneumonine L showing names of th5 sclerites...... 24 Fig. 2. Left side of cephalic structure of Vulgichceumon brevicinctor (Say) showing accessory reticulate sclerotization (ARS) , accessory pleurostomal area (APA) , pleurostoma (P) , and hypostoma fH)aameaae~~~~a~m~~mm.mmaaam~~aa.maaa~aa~~m28

Figs. 3-4. 3, diagrana of hypostoma and inferior pleurost omal process of an ichneumonine. 4, spines on hypopharyngeal surface of an ichneumonine: A, single spines; B, spines in roirs...... 29 Figs, 5-7. 5,6, diagrams of mandibles of final-instar larva of an ichneumonine: 5, mandible in anterior aspect; 6, mandible in ventral aspect. 7, diagram of spiracle of final-instar larva of an ichneumonine~~...... 33

Fig. 8. Head capsule of first-instar larva of Trogus fulvipes Cress. MD, mandible; CL, clypeolabral plates; P, pleurostoma; AS/SS, fused hypostomal spur and stipital sclerite; ANT, antenna1 disc; : A, hypostoma; IS, labial sclerito...... m-...38 Pigs. 9-,10. Cephalic structures of final-instar larvae: 9, Phaeogenes hariolus (cress.) ; 10, -P. gaspesianus (Prov.) ...... 65 Figs. 1 1- 12. Cephalic structures of final-instar larvae: 11, Phaeogenes hebrus (Cress.) ; 12, Dicaelotus gelechiae (Ashm.) ...... 66 Figs. 13-20. 13,14, Ceph k ic structures of , final-in star larvae: 13, Diadromus sp. ; 14, Aethecerus sp. 15-20, Prothoracic spiracles of finql-instar larvae: IS, Phaeogenes Hariolus (cress.); 16, g.

aspesianus (Prov.) ; 17, _P. bebrus * ?Cress.) ; 18, dicaelotus gelechiae (Ashm.) ; 19, Diadromus sp.; 20, Bethec~rnssp...... 67 b Figs. 2 1-22. Cephalic structures of final-instar' larvae: 21, Platylabus ornatus Prov". ; 22, Pa- r ufipes ccnsors Cress...... B2 . Figs. 2 3-28. 23,24, Cephalic structures of final-inatar larvae: 23,-Cyclolabus dubiosus Perkins; 24, Anisobas luzerniensis (Brad. ) . 25-28, Prothoracic spiracles of f inal-instar larvae: 25, Platylabus ornatus Prov.; 26, P. rufipes ccnsors Cress. ; 27, ~~clGlabusDubiosus Perkins; 28, Anisobas luzerniensis (Brad.) ...... 83 Figs. 29-31. Cephalic structures of final-instar larvae: 2 9, Hoplismenus morulus morol us Say ; 30, He m. pacificus Say; 31, ~ulgichneu~on-brevicinctor(Say) ...... I69 Pigs. 32-33. Cephalic structures of final-instar larvae: 32, Cratichneumon + unif asciatorius vancouveriensis (Pro-v. ) ; 33, C. sublatus (Cress.) ...... ; ...... I70 Figs. 3 4-35. Cephalic structures of final-instar larvae: 34, Cratichneuaon anisotae. Heinr .; 35, g. varieqatus (Prov.) ...... 17'1 Figs. 36-37. Cephalic structures of fi~al-inshar' la-rvae: 36, ~ratichneumori.ritus Heinr. ; 37, Cratichneumon sp. nr. vescus (Prov .)...... a72 , - Figs. 38-40. Cephalic structures of ffnal-instar larvaer 3 8, Cra2iclineumon pteridis Tow. ; 39, Patrocloides perluctuosns fProv.) ; 40, Aopltts perrautabilis permutabilis Heinr...... 173 Pigs. 44-42. cephalic structures of final-instar larvae: 41, Aoplus velox velox (Cress.) ; 42, Eutanyacra suturalis (Say) ...... I74 Fig. 43. Cephalic structure of f inal-instar larva of Diphyns variegatus euoxae (Heinr.) .... .I75 Pigs. 44-46. Cephalic structures of final-instar larvae: 44, Limerodops belangeri (Cress.) ; 45, ~pilichne'umon subruf us (Cress.) ; 46, Thprateles lugubrakor {Grav.)...... em...... e476 Pigs. 47-43. Cephalic structures of final-instar larvae: 47, Thyrateleg'procax . (Cress.) ; 48, Ichneumon ambulatorius .Tab. ; 49, -I. caliginosus Cress...... -177 Pigs. 50-52. Cephalic structures of final-instar larvae: 50, Ichnen mon canaaensis Cress.; 51, -I, dioryctriae Heinr.; 52, -I, maius- Cress...... ,..,...... 178

Pigs. 53-54. Cephalic skructures of final-instar- larvae: 53, Ichneumon trizonatus Prov. ; 54, Orgichnenmon carcatorins (Thunb.) .... .I79 Figs. 55-67. Prothoracic spiracles of final-instar larvae; 55, Hoplismenus morulus morulus Say; 56, g. 5. pacificus Say; 57, Vugichneuaon brevicinctor Sag; 5 8, Cratichneumon unif asciatorius vancouveriensis (Prov. ) ; 59, C. snblatus (Cress.) ; 60, c. anisotae Heinr. ; 6 1, C. variegatus (ProV.) ; 62, s. situs-~einr.; 6,3, s rat ichneumon sp. nr. vescus (Prov. ) ; 64, s. pteridis Tow. , 65, Patrocloides perluctaosus (Prov. ) ; 66, Aoplus ...... 67, Pigs. 80. rot horacic spiracles of f inal-in stat larvae: 68, Eutanvacra sutyralis (Sa Y euoxae Hei,Dr. ; D (Cress. ) ; 71, (Cress.) ; 72, (Grav.); 7 3, T. procax (Cress. ) ; 74, Ichneuno n ambnlatorius Fab.; 75, g.' caliqinosus Cress. ; 76, g. canadens Cress.; 77, I. dioryctriae Heinr. ; 7 I,- naius cress.; 79, I. trizonatus Prov. ; 80, Orgichneamon calcatorius (Th~nb.)m~m~~m~~~~~ .*~aa~-a.aaam.-m.

Figs. 8 1-82. Cephalic structures of final-instar larvae: 81, Coelichneumon brunnori (Rohw.); 82, C.- orpheus (Cress.)...... 193 Pigs. 83-86. 83, Cephalic structure of final-instar larva of Syspasis tauma (Heinr.) . 84-86, Prothoracic spiracles- of f inal-instar larvae: 84, Coelichneumon brunneri (Ro~u.); 85, ~oelickneumonorpheus (Cress.) ; 86, Syspasis tauma (Heinr.) ...... I94 Pig. 87. Cephalic structure of f inal-instar larva of Calla joppa cirrogaster (Schrank) ...... 212 P Fig. 88. Cephalic structure of f inal-instar larva of Conocalama bolter i (Cress.) ...... 2 13 Figs. 8 9-9 1, Cephalic structures of final-instar - larvae: 83, Conocalama occidentalis Cress.; 90, Troqus fulvipes Cress.; 91, T.- psnnator Fab ...... 214

Pigs. 32-35. Prothoracic spiracles of ~ final-instar larvae: 92, cirro aster (Schrank) ; 93, &s Cress. ; 94, Trogus fu -- Cress. ; 95, T.- pennator Fab...... 2 15

xii LIST OF TABLES >

Table 1 The mean size in mm + SD of various morphological characters in th~ f inal-instar larvae of Cratichneumon , .pteridis Tow. reared from 3 different hosts. nPans for any character followed by. the same letter are not significantly different (Student- Neuman-Keul's multiple range test, pC0.05) . Sample size is in brackets.. .;. . .-115

xiii The subfamily Ichtleumoninae is one of the largest of the family Ichneumonidae (Hymenoptera). A11 species a parasitize larvae or pupae of Lepidoptera and emerge from the host pupa. Many are parasites of economically important hosts and a few have been used as biological control agents, I so their identification is important.

Relatively few studies on the taxonomy of the larvae of the Ichneumonidae have dealt extensively with any of the subfamilies of the Ichneumonidae. Finlayson (-1967a) dealt with the 'tribes of the Pimplinae and (1975) the genera of the subfamily . Short (1376, 197,7) described and classified the final-instars of the Hesochcrinae and

Anornalinae: Theriini respectively. Beirna (1 94 1) and Short

(1959, 1970, 1978) dealt with the subfamilies, tribes and genera of the Ichneumonidae. Gauld (1976) and Barron (1976) used larval characters to supplement classifications of the

~nomaloninaeand Eucerotinae respectively. Other workers have used immature instars to separate species attacking a singla hcst ' or grouAp,of hosts (eg. Brthur 1963 ; Cameron 1938; Pinlayson 1960a, l96Ob, .. 1962, 1963, 1967b, 1969; Gerig

1960; ,Hill and Pinckney 1940; Hackauer and Finlayson 1367;

Harris, Cameron and Jepson 1937; Sadava and fliller ' 1967; oppel 1965; Torgersen and Beckwith 1974; Torgersen and Coppel 1.965) . c' There are three reason to study larval forms: to

y provide information that may be useful in revealing

$ taxonomic relationships; to re-examine the clas'sification of the adults of the subfamily Ichneumoninae using information derived from their final-instar larvae; and to provide information that enables the identification of species in the absence of adult specimens. This study attempts to

combine all three approaches. As the keys and conclusions

are based on a limited sample of a large and complex subfamily, is is possible that some changes ma-y have to be made as additional information is obtained. They are

intended to be i beginking, and a basis for further study of the Ichneumoninae.

The keys will be of some practical value to separate parasites of Noctuidas as many of the common 0" noctuid-parasitizing ichneumonines are represented. *.&" Importance of Larval Taxonomy

Papers on the taxonomy of larval structures of the Ichneumonidae usually are of e ither of two types. The first deals with the recognition and identification of species parasitizing a specific host or group of hosts. There are numerous exam$les of this in the literature as mentioned previously. This type of work is of direct importance to those working in ecology and biological

control as the species that parasitized a particular host

can be identified in the'absence of the adult sp~cimen.

Field-collected host material may be dissected immediately

to determins the number and identity of species of parasites present, which speeds work considerably and eliminates space-consuming, and often unsuccessful, rearing. Also,

field collected material from which the adult parasite has

alrtady emerged can be identified to species. Adult \ parasite$ from a particular host can be identified from / their lahlremains and therefore *the adult parasite need not be mounted for identification brut can be kept

or experimental work. Hyperparasitic species can bs easily recognized because the primary and secondary parasite remains are both found within the phytophagous host, or within the primary parasite cocoon.

The second type of research is primarily taxonomic.

The members of a taxon are grouped according tc the affinities of their final-instar larval structures. The r, value of this work, is often underrated. bt the subfamily and tribal levels, classifications based on larval characters have often added weight to and supplemented classifications ,based on the adults. Finlayson and Hagen

(1977) and Short (1978) constructed keys to the subfamiles of the ~chnsumonidaethat demonstrate that the subfamilies,

as they are defined at present, ars more of less distinct 8 entities both as adults and as larvae, This r rob ably reflects natural, phylogenetic differences between the various subfamilies.

Characters that define many of the tribes within the subf smilies were noted by Beirne (1941) , Finlayson (l367a), and Short (1959, 1970, 1978) . The subfamily Campopleginae is a notable exception, as Finlayson (1975) and Short f 1978) 0 were unable to find larval characters which defined tribes

in a fashion congruent with the adult classification.

'studies of larval classifications at the cje&ric and specific levels provide insights into the grouping of species into genera. In one case (Finlayson 1375) a previously misplaced species, eruficinctus (Ualk.) , was placed in its proper genus as Sinophorous eruficinctus (Ualk.), on the evidence of larval characters.

It is possible therefore ,that cryptic species and

sub-species may be detected initially by larval characters. No naw species have been described on the basis of larval

structures and it is unlikely that this will be possibls P until the larvae of most of the species of any taxon have been described.

Generalized Final-i~star Morphology of the ~amilyIchneumonidae

The larval structures used in the classification of Ichneumonidae are the sclerotized cephalic structures

surrounding the oral region and the spiracles. Less C t '3 1

@ 1 6 ai P i -.r3 frequently, spines, processes or appendages on the anal S i i segment aid in classification. A11 of these structures are 42 part of the final-instar skin and are shed with it. They I are usually the only structures readily available for taxonomic study. T ? f 5 I The morphology and function of the generalized cephalic

structures ,(Fig. A) were described by Beirne ( 1941) , Finlayson (1960'a), Finlayson and Hagen (l977), Short (1952), and Hatsuda (1365), and are reviewed below.

. 8 The epistoma extends dorsally between the superior pleurostomal processes, may be complete or incomplete and

.separates the clypeus from the frons when -1ete. fhere . present, the labral sclerite separates ,the labrum from the clypau%; and the suspensorial sclerite, located behind the labral sclerite, may provide an attachment point for the dorsal hypoph%yngeal muscles of the hypopharynx.

The pleurostoma borders the mouth opening laterally betwaen the two pleurostoeal Frocesses and strengthens those Frocesses, which piovide articulation points for the mandibles. 8 Fig. la. Diagram. of the cephalic structures of a final-fnstar i larva of an ichneumonid showing names of the sclerites. After

inl laps on l96Oa.. Used by permission. Vacuole Labral scluita Swmsoriol Sclaita

I //+f--- Antenna Superior nmndibulor process Mondibh Pleurortomo / II Twh Lacinial xlrita I&iw mondibuh pocess

~i&taISCI& Dortol arm of labial Labial sclaite Labial ml~ I Each mandible has q lower articulation point (the condyle) that articulates with the inferior pleurostomal

process and an upper articulation point that articulates

wirh the superior pleurostomal process. The blade of the mandible may or map not have teeth.

The hypostcma extends laterally and/or ventrally from

the lower end of the pleurostoma across the head ca~suleand'

supports the maxillary region. The hypsstoraal spur (spur o"f

the hypostoma) extends into the maxilla, dividing it into 2 parts, and rests on the stipital. sclerite, which extends

along the lover margin of the maxilla.

1 The labial sclerite defines the borders od the prelabium; the postlabium is not defined. The silk orlfice

lies between the hypopharynx and prelabium, which are fused. The area surrounding the silk orifice is often sclerotized \ o form a silk press. A X-shaped prelabial sclerite is sometimes present on the prelabium. There are often spines on the dorsal surface of the hypopharynx, which may assist

in fesding. The maxillary and labial regions are most , complex 5n those species that spin cocoons outside the host

and .are gensrally less 'conplex in species -which pupate d f insida the host pupa, whether or not they spin cocoons. 95, j

h Antenna1 discs or papillae are scmetimes present dorsal to tha epistoma. Ocular lines are sometimes present and

ihdicate the position of the compound eyes in the adult.

The mature larvae of most Ichneumonidae have 9 pairs of

4 ,spiracles (Hagen 1964); the first and second on the pro- and metathorax (Clausen 1940) and the remainder on the abdomen. -4 The spiracls consists of an atrium and subatrium; the - subatrium usually has a muscular closing apparatus

-. C associated with it.

A few species of Ichneumonidae have caudal appendages and processes in final-instar larvae which are useful for

classification (Finlayson 1364,', e and Finlayson in press). The anal segment frequenQ lr has small spines which may be useful for classification.

. . (Pimplinae), which ars endoprasites of Lepido~tera,show soma structural convergcrnce with the Ichneumoninae. In Itoplsctis and Pimpla the hypostoma is reduced and the

hypostomal spur occupies the approximate posticn of the hypostoma in the Ichneumoninae. Pimpla and Itoplectis have

a labial sclerite and reduced stipital sclerites and the b Jchnaunoninae do not. The final-instar larvae of the .. 9 subfanil y Ortho pel natinae, which are endoparasites of

Mycetophilidae and Sciaridae (Diptera) (Short 1978) and of

gall- forming Cynipidae (H y inenoptera) (Barron 1 977;- Beir ne

1941), are almost identical vith those of the -Ichneumoninae,

but are distinguished from them by ho lightly-sclerotized cephalic structur e different shape of tha mandibles (Pinlayson and Hagen 1977; Short . 1353, 1 978) .

The adults of the Ichneumorinae fall into two major sub-groups on the basis of propodeal spiraclk shape: the Ichneumoninae Cyclopneusticae have circular or subcirclar propodsal spirac.% es; and the Ichneumoninae Stenopneusticae have elliptic to elcngate propodeal spiracles. In the latter group,. the tribe ~latylabini[=Pristocerotini of .riu TI W .p 0 (d U) rl -4 Q, kaa -rl s +,aao 4 a E-c 91 0 3 '44 w II 0 C u Y Ti .rl a ** u -4 U) -ri a @do SOB .rl a I H5al 4Ja'a u A ODAU U H UJ H II u U .rl s (d -4 TI ~auu -4 -r( o=u UOO I4 ia cuaa cdoo) OCIU .rl A +JGU U) .ri .,I IGlY a' ' 0% iLucs(U Biology of Ichneumoninas

1 d d The larvae of Ichneumoninae are, without exception, i i endopar asitic on Lepidoptera; T dults oviposit into , either host larvae or pupae but ys emerge from the pupa, , Many species of ~chheumoninaeoverwinter as adults,

Species of Stenopneusticae in which the female is amblypygous, generally oviposit into the aost larva whereas oxypygous females oviposit into the host pupa (Hinz 1973; Heinrich l96la) . h hi^-' general rule, established for the Stenopneusticae, also holds true for the Cyclo~neusticae, which are mostly oxypygous. Smith (1932) found that Pbaeogenes nkgridens Uesm. oviposits into the host pupa as does yerpestomus brunnicornis (Grav.) (Beirne 1943) .

The Ichneumoninae Cyclopneusticae utilize nicrolepidoptsra as hosts whereas the Ichneumoninae stenopneusticae generally parasitize Hacrolepidoptera

(Perkins 1953, Short 1978). The Platylabini parasitize ~eooetridae(Hacrolepidoptera) , a fact which places them biologically in the Stenopneusticae. The Listrodromini are

parasites of L ycaenldae (Lepidopter'a: Papilionoidea). 6(

The Trogini, subtribe Trogina, are parasites of swallow tail butterflies (Lepidoptera: Papilionidae:

Papilio) while members of the subtribe Calla joppina, as - defined by Heinrich (1962c), are parasites cf Sphingidao (a ich 1960a, 1962c) .

/ The host ranges of the Protichneumonini are more diverse. They do not utilize Papilionoidea as hosts although they have adapted to a wide range of other macrole pidopteran hosts, bein g recorded from Sphingidae , Arctiidae, ~ymantriidae, Notodontidae, Noctuidae, Geometridae, Pyralidae and Tortricidae (Heinrich 196Oa) .

The Ichneuntonini parasitize a wide range of Hacrolqpidoptara. The 4jority 05 tho genera parasitize Noctuidae and related groups (Heinrich 1960b). Recorded by Peck (1964) as parasites of Noctuidae are Spilichnea mon, Chas~ias,Limerodops, , ~utanyacra, Ctenichneumon, Esephanes, ~ctipomorpha, Patrocloibas, , Rubicunbiella and Vulqichneumon. The ma joritp of the smaller species of tlelanichneumon are parasites of Geometridae, as are speciss of Stenobarichneumon, Aoplus, and Platylabops. Orqichneumon parasitzes Liparidae; Anisopygus parasitizes Notodontidae; and Eupalamus parasitizes Lasiocampidae.

The gsnus Trogomorpha, which Heinrich (1962b) places close to the fiistrodroiaini, parasitizes Hesperidae

(Skippers). Hoplismenus, Thyrateles and the gracilicornis

group of Ichnaumon utilize Nymphalidae and Sat~pridaeas hosts (Heinrich 1960a) .

The majority of the species. of Ichneumon are parasites of Noctuidae while a few have specialized on such divers3 hosts as Arctiidae, Hepialidae and Pyralidae. The larger species of Cratichneumon parasitiza Arctiidae, Noctuidae, Lymantriidae, Notodontidae, Liparidae and Saturniidae, while the smaller species parasitize species of Geometridae. P Considering the large number of species of Ichneunoninae, relatively few papers are available that deal extensively with their biology. This situation is peculiar also because the species are mostly large and conspicuous, and many of them are important parasites .of pest

Lepidoptera.

Details of host finding are not knowp, but those species which have been observed appear to search randomly in suitable host habitats until a host is located (Abase and

Hathenge 1972; Smith 1932; Wishart 1948) ;-This suggests that no long-range host-finding mechanism exists, although the females seem to be efficient searchers, and readily recognize material with which the host had come in contact

(Smith 1932).

The overwhelming majority of the species investigated in the litarature parasitize hosts which are in a concealed . habitat during the stage attacked (abase and Hathenge 1972;

Beirne 1943; Cameron 1950; Eidt and Sippell 1961; ~oh~uddin 1972; Schaaf 1972; Sechser 1970; Smith 1932; Wylie and Bracken 1977) . One axception is Fubicundiella pertatrubix Heinr. for which host habitat was no-t reported by Wishart

(1948), although the species is a parasite of the European corn borer, ,Ostrir-ia nubilalis (Hbn.). It can ke assumed to

be a parasite of concealed hosts also, although it will attack non-concealed hosts in the laboratory. Another is

Thyrateles haereticus (Wesm. ) which - parasitizes Vanessa spp. (Nymphalidae) and apparently attacks the host in a non-concealed habitat, following a sudden dive on the larva *

Smith (1932) and Wishart (1948) have shown that all

instars of Phaeogenes nigridens and -R. pertatrubix feed actively with their mandibles on the hbst tissues. This is

contraky to the conclusions reached by Clausen (1940) that the early instars, especially the first one, feed on the

host haamolymph. The first- and second-instar larvae use

their mandibles to break apart the fat body:of the host, and \ .?, later instars feed directly on all the host tissues (Smith 1932; Uishart 1 948) .

There is some variation in the number of larval instars in various species of the Ichneumoninae. Smith (1932) reported 4 and possibly 5 instars in -P, nigridens (Phaeogenini) , and Wishart (1948) reported 3, in R. pertatrubix (Ichnsumonini) . . Gerig (1960) reported 5 larval instars in Phaeogenes osculator Thunb.; Clausen (1340) felt that most of the Ichneumonidae have 5. The number of larval instars may vary with host quality or may be a function of the taxon. J

Hany species of Ichneumonini and Phaeogenini overwinter as fertilized females (Heinrich 1360a; Hinz 1973; Claussn I 1940) although ~rotichneumonini,Trogini, Platylabini, and Listrodromini, as well as Cratichneumon, Ctenichneumon and related genera do not, According to Heinrich (1961a) those species which do not overwinter have 2 generations per year, -, vhile those that do have only 1. HETHODS AND MATERIALS *.

Host. of the specimens used in this study were received or loan from the Canadian National Collection, Ottawa Ontario. Many of them were reared by the Forest

rvey and the For~stInsect and Disease Survey (FIS); the

PIS number is quoted in the text. Additional - specimens were received on loan from the Agriculture Canada Experimental Station, Winnipeg, Manitoba; the remainder came from the personal collections of Dr. Hiktat Doganlar, Faculty of Agriculture, Ataturk University, Erzerum, Turkey, and myself.

' In this paper, uspecimenw refers .to a pinned adult ichneumonid (~chneumonidae: Ichneumoninae) , the host pupa from which it emerged and the larval remains inside the host

The pupa was rsmoved from the pin, all data from the specimen labels were transferred to a card and the slide label, and the specimen was given a number which keyed it to them. The parasite cocoon was examined and described as far as possible and the overall dimensions, shape, - coloar, and textur9 of the host pupa, position of the exit hole from the host pupa, and position of the parasite remains in the host pupa were noted.

Because of the fragility of the host pupae, some of which had been preserved for 70 years, a technique was evolved, similar to that subsequently reported by Short

(l978), for extracting the cast skins and cephalic structures of the final-instar larvae. The host pupa was transferred to a vial of 10% KOH and left to scak overnight (12- 16 h) . The softened host pupa was then slit longitudinally from the exit hole to the base of the abdomen, and the contents removed and placed in a dish of

10% KOH. The parasite remains were transferred to clean KOH and, if necessary, cleared further over low heat for 10 min. Head capsules of penultimate. and earlier instars were either washed in distilled water and stored in 70% alcohol or mounted as described below, along with the final-instar skin and cephalic structures.

The skin and cephalic structures were transferred to distilled water, rinsed, and mounted on a slide in dilute deFaure ' s leedium under an unsupported cover glass. Although an unsupported cover glass flattens the cephalic structure into a single plane, it gives constancy to mearurements by eliminating curvature, which makes illustrating easier.

Classification follows that of Heinrich (1960a, b,

1961a, b, 1962a, b, c) and Krombein and Burks (1967). Nomenclature of larval structures follows that of Pinlayson

(1960) and Short (1978) and nomenclature of adalt structures follows that of Heinrich (1961a).

For most of the species descriptions, only 2 to 5 specimens wsro available, and therefore only means and ranges are quoted in the text. For 1 species, Cratichneumon pteridis, of which a large enough sample was available for more detailsd analysis, the iaean, number of specimens (n) , range, sta~ldardeviation (SD) , and coefficient of va~iation

(CV) are given. Measurements of bilateral characters were averaged for each specimen. , The effects of host and location on various morphometric characters were analyzed using Oneway Analysis of Variance and Student-Newman-Keul's multiple range test. dOBPHOLOGY OF FINAL INSTAR LARVAE OF THE ICHNEUflONINAE

The final-irstar larvae of the Ichneumoninae always spin a partial cocoon inside the host pupa. This cocoon usually consists of a loose network of silk on, and attached , 0 at scattered points to, the intclrior surface of the host pupa. It usually ends at the hind margin of the wing pads of ths hcst pupa and is open. The front end of the cocoon is within the head capsule of the host pupa and is often closed, particularly in the Ichceumonini, Protichneumonini and Trogini. The head 2nd is usually the only art of the , cocoon of the Ichneumoninae that is solid and even here there are usually many small circular openings in its wall. r- Spreetimes a paiP of denser bands extends alcng the length of the cocoon which may serve to orient the emerging adult, as has been hypothesized for other groups (Salt 1977). The sxit hole from the host pupa is always, with the exception of the subtribe Trogina (Trogini) , through the host head capsule, and is formed by cutting away the head capsule of the host pupa, which may remain attached to the host pupa as a cap. The Trogina exit through the wall of the cocoon dorsally, laterally or ventrally through an opening . posterior to the head capsule of the host pupa. This ;e\ openidg oftsn has a burnt appearance, and does not appear to ,; have been cut. The difference in location of the exit hole may wueto the heavily-sclerotized structurs of +he head capsule of th2 hosts, - which are species of ~apilio (Papilionidae) , which couid make it difficult for them to cut through it.

The cast final-instar skin and cephalic structure is always found in the meconiurn in tLhe host pupa in the end furthest from the exit hole. Frequently, the resumed first-instar head capsule and intermediate cephalic structures are also found in the meconium.

The final-instar csphalic structures of the

Ichneumoninae (Pig. Ib) are distinctive and characterize the subfamily as readi'ly as do the adult structures. The distinctive characters of the cephalic structure are the lack of labial, prelabial and stipital sclerites, and , hypostomal spur, as also noted by Beirne (19419, Short

(1958) and Finlayson and Hagen (1977). Very rarely, the unsclerotized outlines of the labial and stipital sclerites Fig. Ib. Diagram of ths cephalic structuke of a final-instar larva of an ichneumonine showing naines of the sclerites. - -EPISTOMA

/----- PORE and hypostomal spur are visible. This is considered to be z derived character and does not indicate that these species .a .a are primitive as the adults are among the most advanced and specialized of the Ichneumoninae (Heinrich 1969~). Suspensorial and labral sclerites are alwayS absent, though a pair of sclerotized clypeolabral plates (Fig. 1) occupy their approximate position.

In the final-instar larva the epistoma is usually sclsrotized, though in some genera'or species it may be lightly sclerotized (some species of Cratichneumon and Aoplus) ', unsclerotizsd (Trogus, Platylabus) or absent (Spclolabus, Diphyus) . The epistoma is divided into'two sections by the anterior tentorial pits which are located latarally, just above the epistoma. According to Hatsuda (l965), the epist cma occurs medially between the anterior tentorial pits and the pieurostoma is on either side of the anterior tentorial pit* Short's (1952) deficition of the a epistoma is that sclerite found madially between the sup 1rlor pleurostomal processes. The term Itepistomal bridb is used to refer to the sclerite between the anterior tentorial pits, i.e. the epistoma sensu Hatsuda (1365). Pores in the epistoma vary in number and position within a species and are of limited taxonomic significance.

They are probably nerve canal tracks to sensilla on the epistoma that are lost or displaced during moulting, That part of the epistoma lateral to the anterior tentorial pits rarsly has pores.

The pleurostoma is always sclerotized and may have accassory areas outside its lateral margin which may vary in density and extent of sclerotization, The pleurostoma and accessory areas may also have pores. I

The overall width of the cephalic structure,includes the accessory pleurostomal areas because these areas con tribute significantly to the overall impression of size

in the specimens, and are of rslatively constant dimensions.

The hypostoma is always sclerotized, and never has pores, It varies from broad and truncate to narrow and pointed even within the specimens of a single spscies, and the distal- end nay be smooth or broken. The angle vhich the hypostom "\ forms with the pleurostoma is variable and is not used for classification except where constant and

exceptional. The hypostoma is measured from the inflection point of the curve joining the pLeurostoma and hypostoma on

the outside to the distal end of the hypostoma (Fig. 3).

Sometimes a second type of accessory sclerotization, (Fig. 2) associated with the distal end of the bypostoma,

'extends dorso-laterally from that structurs, b.ut is \un ssociat-di- with the accessory pleurostomal sclerotized area always has reticulate sculpture and is

to have accessory reticulate sclerotization.- This sclerotized area may represent the occipital suture in the larva or, possibly, reticulate sclerotization on the occiput, vertex and gena.

The superior pleurostomal process usually consists of

an anterior and a posterior strut, the latter often being reducad or absent. The inferior pleurostomal process also

consists of an anterior and posterior strut which may be fres from one another, fused ventrally to farm a trough, or fused dorsally and venteally to form a tube or pit. The

form of the process, the ratio of its length to its width

and the relative lengths and widths of the two struts (Fig. Fig, 2. Left side of cephalic structure of brevicinctor (Say) showing accessory reticulate sclerotization (ARS) , accesscry pleurostomal area (APA), pleurostoma (P) , and hypostoma (H) .

Figs. 3-4. 3, diagram of hypostoma and inferior . d pleurostogtal process of an ichneumonine. 4, spines on hypopha-ryngaal surface" of an ichneumonine: A, single spines; B, spines in rows. I PLEUROSTOMA I I I ACCESSORY PLEUROSTOMAL SCLEROTIZATION I 1 I I POSTERIOR STRUT OF INFERIOR PLEUROSTOMAL PROCESS \ 1 \ ANTERIOR STRUT OF INFERIOR PLEUROSTOMAL PROCESS \ C -D = LENGTH OF ANTERIOR STRUT ' \ . \ T2 - E= LENGTH OF POSTERIOR STRUT F - G = WIDTH OF ANTERIOR STRUT F - H = WIDTH OF POSTERIOR STRUT 3) are of taxonomic value.

. The maxillary and labial palps are disc-shaped, never papilliform, and bear sensilla which vary in number and

size. These have limited value taxonomically. The number

and shape of sensilla on the palps appears to be quite variable within a species and, in some cases, between the

right and left side of a single specimen. Where a large number of specimens has been available fcr com~arison, this usually ranges from cne fewer than 'average to one more than

average. It appears to have sevaral causes: two sensilla close together may appear as a single bilobed sensillum; a single large sensillum may appear as two due to abnormalities in the surrounding chitin; a single, small

sensillum may be absent; or a fracture in the disc may appear as a sensillum. The range and most common number is

given in ths text for species that have this variability.

The structure of the silk press varies between the species and genera and is of some taxonomic significance. ,The silk Fress is situated on the unsupported skin and is often distorted almost beyond recognition although one I specimen of each species was complete and suitable as a basis for illustration. \ Some of the illustrations are reconstructions based on examinations of more than one specimen. Ths lower margin of the silk press .sometimes projects ventrally as a spine-like process, rasulting in a bisinuate lateral margin, or as a lobe in 'which the lateral margin is evenly rcunded. There is usually a Fair of setae located on the silk press, the position of which has some taxonomic value.

Setation in the maxillary and labial regions is used as little as possible in classification due to the unstable nature of the surface on which the setae are located. They are therefore never illustrated.

Often, sculpture or spines occur on the hypopharyngeal surface, which are of taxonomic importance. The spines, when present, may be single or double (Fig. 4A) or may be in rows of 4 or more (Fig. 4B). When the spines are in rows of

4 or more they are usually on the lateral surfaces of the hypopharynx and the medial-spines are single or double. The mandibles (Figs. 5, 6) are aluayp heavily sclarotized, especially at the tip of the blade, and are -l .e uithout sculpture or 'teeth. They consist of two aa jor regions: the body and the blade. The body is enlarged and open at the base and has two articulation points: the dorsal one is a simple notch or groove which articulates with the superior pleurostomal process; the ventral condyle is bulbous and projecting and articulates with the inferior pleurostomal process. The -body narrows either gradually or

abruptly into a heavily sclerotized blade which may be .* Z slightly to strongly hooked in a posterior direction.

Depending on how the mandiblek- are turned during mounting, they may present one of two aspects. The ventral aspect (Pig. 5) is when the mandible rolls during mounting so that the ventral surface (i.e. the surface

which would be visible if the mandible were examined from

the ventral surface of the larva) is v This TIe* orientation reveals the curvature of the vnterior aspect (Pig. 6) is visibae when

pressed flat against the slide without rolling so that the h /- face-on surface is presented, which does not reveal the .-'Y

Figs. 5-7. 5,6, diagrams of mandibles of final-instar larva of an ichneumonine: 5, mandible in anterior aspect; 6, mandible in ventral aspect. 7, diagram of spiracle of f inal-instar larva of an ichnsumonine. -

1.

DORSAL ARTICULATION POINT OF MANDIBLE \ BODY OF MANDIBLE I - J - LENGTH OF MANDIBLE K - L = LENGTH OF BLADE M - N = WIDTH OF MANDIBLE 0 - P - WIDTH OF BLADE AT BASE

BLADE OF MANDIBLE

UL I I I I I

CONDYLE OF MANDIBLE I I

CONDYLE OF MANDIBLE

I - J = LENGTH OF MANDIBLE K - L = LENGTH OF BLADE M - N =WIDTH OF MANDIBLE 0 - P = WIDTH OF BLADE AT BASE

MANDIBLE

I b I BODY OF MANDIBLE 1 , I I M I I Ik w J

HAIRS IN ATRIUM SPINE IN ATRIUM ATRIUM

SPINES IN CLOSING APPARATUS

HAlAS IN CLOSING APPARATUS tlade~curvature. The maodible orientation used in

illustrations was determined by tho best specimen available

for illustration.

? In' Cratichneumon. 3teridis, the lengths, widths, and blade lengths of m andibles in each oriantation were

compared. In the anterior view, mean mandible length was

0.35 mia (n=21, range 0.32 To 0.41, SD=0.025, -12%); mean

nandibls width' was 0.27 rnm (n=21, range 0.24 to 0.32, SD=

0,013, CV=6,87%) ; and mean bgade length was 0. t 0' wt (n=21r ranga 0, 12 to 0.17, SD=O.O14, ~V=10.l6%). In td~evsntral view, msan ma-ndibie length was 0.37 rnm (n=12, rang@ 0.32 to

9.40, SD=0.026, CV=7,15%); Rean ~andiblewidth was 0.28 ma

[n=12, range 0.26 to 3.32, $D=O.OlU, CVs4.8996) ; and mean

blade length vas 0.13 mm (E= 12, range 0.11 to 0.14, SD=O. 0-11, CV=8.68%) . There was no significant difference betugen the mans of any of the above measurements (t-tes: p<0.05) . The mean ratio of rnandible length to mandible width was 1.30 (n=21, range 1.14 to 1.48, /- CV=7,66%) for the anterior- view and 1.29 (n=12, range I. 14 to 1.43, SD=0.077, CV=6.00%0 for the ventral vie.w, vith no significant difference between .the means (t-test p

43 3e v ?: The mean ratio of blade leagth to mandible length was 1 - 1 0.40 (n=21, range 0.30 to 0.47, SD=0.038, CV=9.45%) for the a anterior view and 0.35 (n=12, range=0.32 to 0.39, SD=0.020, ' f CV=5.58%) for the ventral view. Again there was no

significant difference between the means (t-test p<0.05)

although the blade length and ratio of blade length to

mandible length was consistently smaller in paired aandibles

(i1 in ventral view, 1 in anterior view]. The ratio of

Mads length to total length of mandible can be used as a

taxononic character although sxtreme care must be taken in

determining blade lengths.

As noted above, a pair of clypeolabral piates is always

pressnt on the clypec-labral region adjacent to the epistoma

abva the superior pleurostoaaf processes. The form and

position of these plates anrthe number of sensilla upon

then is cf. taxonomic significance. Sclerotiza hions of this

1--b sort are not unccmmon amongst final-instar larvae of the

Ichneuraonidae, although they do aot appear with such

consistency or complexity in any other group. The plates ,9

each aay be joined to the episto

pleit-ostomal processes, by sclerotized or maabranous regions - or may be unattached to the epistoma, They may als~be joined medially to each other.

Antenna1 discs are sometimes present 'bove the cephalic

structure and may be sclerotized or unscleratized. A pair of ocular lines, which are arAndicating the positions of

the compound eyes in the imago (Short 1978)' is also visible in some species and sometimes have reticulate sculpture on the surface. In scma species the above two structures may ( not be found in more than one or two specimens and, if so they are always lightly sclerotized and are not usually

Some spacies have spines at the bind end of and. these are us"ed frequently in classification.

The spiracles (Fig. 7) usually have an avoid atrium and may have spines within the atrium. The closing apparatus %t~suallyertsnds to the atri&n and often slightly into it:

there is no distinct neck bet ween the atrium and the closing

apparatus. There are always long, dense hairs within the

closing apparatus, and there nay be heavier interior spines in the distal end of the closing apparatus. There are 9 pairs of spiracles, 2 thoracic and 7 - abdominal. Short (1 978) has stated that "some authors have; described the spiracular atrium of certain species (of Ichneumonidae) as possessing spines, but these arg patterns of the cuticular spiral thickening of the tracheal lining

seen at various depths." Many species of Ichneumoninae have

definite spines inside the atrium of the spiracle and fine hairs are also frequently present. The closing apparatus

(subatrium of Hagcn 1964) aiso contains numerous spines and

no musculature is visible so the actual closing apparatus

may not be shed with the spiracles.

The cast cephalic structures and skins'of the first- and intermediate-instar larvae of Ichneumoninae found in the meconium within the host pupa cansist of a tubular head capsule, which is presumably the first-instar head capsule, and one or more .hemispherical head cqpsules, respresenting second and later instars.

- ?r. These head capsules, particularly the first-instar

(Fig. - 8), often have a labial sclerite as well as hypstomal

spurs and stipital sclarites. The stipital sclerite abuts b Fig. 8. Head capsule of f irst-instar larva of Trogus fulvipes Cress. HD, mandible; CL, clypeolabral platss; P, plenrosioma; HS/SS, fused hypostonal spur and stipital sclerite; ANT, antenna1 disc; H, hypostoma; LS, labial . sclerite . the labial sclerite, and the hypostomal spur arises from the hypostoma near the inferior pleurostomal process. The epist oma is indistinct, usually no more heavily sclerot ized than the dorsal aspect of the head capsule. The pores in the epistoma, visible in th2 final-instar larva, are present and have short trichoid sensilla associated with them.

There are two lobed structures which are sclerotized at the tip and bear s~nsilla. These are the clypeo-labral platas. The manditles have a strongly-hooked blade, which prob&bly aids in broaking up fat bodies in the host.

The skins of these larval instars are infrequently fouad. Their spiracles always appear to be non-functional, each having an attenuated closing apparatus and indistinct atrium. In many respects they resemble the last three pairs of abdominal spiracles the skin. cd m Id =I fa . u 0 .ri .rl . Cr . a 0 0 a Cc 0 0 0 H n 0 (d 0 0 rl . a. 0 rt cn . -4 0 cda,Cr- u . Urnfa. -4 0 -4 0 m Yrl 0 . A *@ 0.' 0 M . 4 .rtec,- 4' I3 0 -6-f &=lO(Uo d S -rlWb* 0 *+., . rn a, said,. U1Q,OvlF1. Id . ~aclcu . .a. ri.r(W4 L 4J.H. Urn .UW* (d-Id. I~CI-@a 03. Cc .rl . k .-r+ T3 4J. .ri 0.d U) u 01. am ad Id.0. wa, m.rl * 0 d +,-.41. -ww. 3 . om-On,. 0.a. U1 u .&I. vl U) .,-I h.@. I.)*-.a- d *& -84 b -4 U) . a-mc, (dcd -Id 0 LI. m aa, ah. cd .+,d A . b A* 0) w ma. v, $d L 0 0) A .4 a + 0 J A +J Y .ri wu .vl +' kO .U -E: .rl r-l -vi@ Id W w.a cr 0 d 41 E: W.43 OId moms a a ad -aa 31 .~+'a -,-I A * .rl -4 a kcrt -k5(tl c, +) d al 4. Blade of mandible 0.2 or less of total length of mandible...... a.a...... 5 Blade of mandible mora than 0.2 of total length of mandible...... a.m.....a...... 6

5. Epistoma complote ...... Ichneumonini (ir part) Epistoma incompletj...... Listrodromini: bnisobas

6. Epistoma incomplete and clypeolabral plates a small and much wider than deep; or opening of silk press half as wid? as deep...... Platylabini . Epistoma complets, or if incomplete,

clypeolabral plates large and about as

wide as deep; opening of silk press more than half as wid^ as deep...... 7

7. Lsngth of closing apparatus of prothoracic

spiracle more than 2.5 times width of atrium ...... ~xotichneumonini? Coelichneumon Lsngth of closing apparatus of prothozacic spiracle less than 2.5 times width of atrium...... 8 -1 8. Length of closing apparatus less than or equal

to width of atrium, and clypeolabral plates joined aedially.. , ...... ,Pratichn;umcnini: Syspasis Length of closing apparatus greater than ~idth

of atrium, or clypeolabral plates not jcined medially .. . , ...... Ichneumonini TRIBE PHAEOGENINI

The cephalic structures and spiracles of final-instar

larvae of 4 genera of Phaeogenini were examined and a key for their separaticn follows.

Genera and species of Phaeogenini described or illustrated in th~literature but not considered below are:

Herpestomus brunnicornis (Gravenhorst) by Beirne (1 943) and

Short (1978) ; Th~raeellacollaris (Gravenhorst)- by Given

(1944) and Short ( 1978) ; and Hemichnsumoc slongatus (Ratzeburg) , Oiorhinus pallipalpis Wesmasl, Centeterus opprimator (Graven horst) and Takamona vincibilis (Cresson) by Short (1978) .

Short (1378) separated the Pha~ogeninifrom the rcst of the Ichneumoninae by the presance of a "lightly sclerotized

plate.. .cn the clypaus ventral to and continuous with the

~pistoma,~~The form of this character was not clear or distinguishable in the specimens available. The alternate character which is us~dis: t~clyps~labralplates joinad to

epistoma near pleurostomal procasses by sclerotized areas, or, If area apparently un sclerotized, small islands of sclerotization visible in area, sometimes adjakent to

epist~ma.~~Difficulties arise in- the use of this character because Phaeogenes hebrus has a very .lightly sclerotized area the between the epistoma and clypaolabral, platqs, although there is some sclsrotization adjaceat to the . pleurostomal processes. This character is further ------confounded by the genera rati ichneumon, Vulgic-hneumon and

- Calla joppa (Trogin i) in which the clypeolabral plates are joined to the epistoma by unsclerotized, wrinkled areas.

Although the presence of some sclsrotization near the

epistoma sets -P. hebrus apart from these genera, the form of the clypeolabral platss suggests a close rslationship with

the above genera. It is possible that the 2 specimens

lxaminsd were the adults of .P. hebrus .which had been pinned

with the pupae from which another species of Icbneumoninae had emerged.

Other characters of the Phaeogenini are: mandibles with long blade which is slightly to strongly hooked; antenna1 discs and ocular lines not visible; and maxillary palps sach usually with 5 sensilla and labial $alps each usually with 4 sensilla. Short (1978) noted other species of ~haeogeniniwith different numbers of sensilla on their palps and with antenna1 discs, so some of the above characters may vary . \.,

The prothoracic spiracle varies considerably in form.

The atrium usually has small hair-like spines inside but . never has large heavy spines. Somstimes the closing apparatus has a few inner spines near its junctioa with the atrium, in addition to the- normal dens2 hairs found in the closirg apparatus.

Key to genera of Phaeogenini

1. Laqth of closing apparatus of prothoracic spiracle less than twica the width of atrium, or clypeolabral region completely scleroti~ed...... ~.~~~..~..~~.~.~~...... ~.~.~.~2

Langth of closing apparatus of prothoracic spiracle twice, or more the width of atrilum,

and ciypeolabraf region not completely 2. Length of inferior pleurostomal prbcess equal to, or less than, width of inferior pleuro- * stoma1 process...... Phaeogeaes Length of inferior pleurostomal process greater than width of infsrior pleurostcmal

3. Epistoma with 9 to 12 pores; each C~YP~O- 9 a labral plate with 4 to 8 sensilla; clypeo- la-bra1 plates sclerotized...... Aethecerus Epistoma with 6 pores; each clypeclabral plate

with one sensillurn; clypeolabral plates not or v?ry lightly sclsrotized...... Diadromus

*1 In one of th3 =YO specimens of Dicaelotus th~length of =he closing apparatus was less than twice the width cf the atzium and ir the other spscimen tha lsngth of the closing apparatus was more thsn twice the width of the atrium. The ---Fa clypeolabral region groups this genus with Phaeogenes. Genus Phaeogrrs Vesmael

Csphalic structuros ard spiracles of 3 species of

Fhaeogenes are described and illustrated h~re. -P. phycidis

(~s'hm.) and -P, laricellae 3ason were examlned but not described or i,mted ,as they had beer? described ard / illustrated ehsouhEze. A key io those 5 spscies follows.

Othsz species illustzated in ti19 literaturs ara: -P.

harioius {Cresscn) [ =Dirophanes] by Short ( 1953, 1378) and

Torgprsen and, Beckwith (1374) ; 2. laricellae Hasoc by Eidt

(1362) ; P. maculiccrnis (Stephsns) by 'Short (1970, 1978) ; --P.

3igridens Uesaael by Saith (1332) ; -P.- asculator------(Thunberg) by Gorig (1960) and by Short (I 378) ; P. phycldis Ashmead by .- - Pinlsyson (1967h); -P, suspicar Hsamael by BEii<(134I): and

-P. s:er Cressor, 3. haiussleri Uchida, 1. hebrus (Crsssori) ,

-P. invisor (Thunberg) , 2. ischiomal~usUssrael, end 1. \ ualshiss wslstiae Ashnead by Shcrt ( 1978) .

B Phaeogsnts larvae nay 52 distinguished by the 1e"qth of the closirg apparatus of the prcthoracic spiracle which is iess than twice tho widrh of the atriuu and by the short pl~urosroaalprecess, which is less than 1.5 times is long as -wide. Examination of illustrations of other species in the litorature snppozts this conclusion.

Despi'te the fact that illustrations in the litarature are often not sufficiently detailed or clear to allow

accurate conclusion-3, it appears that the species of - Phaeogenes fall irto two or three morphological groups on the basis of Eiral-instar cephalic structures. Group one consists oi species with th~c1jp~dabra1 region entirely -. sclsrotized, as in K. phycidis, and to a less,er extent, 2. Jaricallae. Group two consists of species with the clypeolabral regicn cot entirely sclerotizad as in -P. hariolus ard -P. hobrus. A possible third group, or a sub-group of eithar cr both of the above groups,has the distal end 05 the hypostoma extended up and around the aazgtns of' =hsZbead capsule as shown by Short (1978) for 2. -t -8 g. walshiae. --zter, haeussferi, and 2. ualshiae 'I ', '+a, Id. da Smaller species, cephalic structure 0.4 mm wide (range 0.38 to 0.44); depth between pleurostomal processes always more than half of length of hypostoma...... gaspesianus Prov. ~ar~erqpecies, cephalic structure 0.63 ma

wide (range 0.55 to 0.74); depth between pleurostomal Frocessss usually less than half of length of hypostoma...... hariolus (Cress.)

3 Phaeo genes hariolus (Cresson)

Figs. 9, 15

k The 15 specimens examined were raared from the spruce ;

J buduorm, Choristoneura funiferana (Clem.) (~ortricidae), i collected at Yale, Stanley, and Fort Frances (PIS 7759), B.C.

Parasite cocoon of a few strands of light tan silk with t two denser dorsolateral bands of silk inside host pupa. 1 ------Exit hole at head end of host pupa, slightly sub-terminal, about 2 x 2.5 ma wide. Cephalic structure of final-instar larva 6Fig. 9) 0.63 mm wide (range 0.55 to 0.74) . Epistoma complete and sclerotized, 'with 6 pores;- pleurostoma sclerotized, with very lighlg sclerotized dorsolateral accessory pleurcstomal area. Hypostoma sclerotized; usually broad and truncate, sometimes thinner and pointed. Clypeolabral plates joined

broadly and strongly to epistoma above the superior

pleurostomal processes, each with 2 to 6 sensilla. Anterior strut of superior pleurostornal process longer than posterior

strut; inferior pleurostomal process about 0.8 times as long as wide, trough-shaped. Silk press wider than deep, opening

about as wide as deep with 1 seta on each lateral margin.

Maxillary palps each with 2 large and 3 smaller sensilla. Labial palps each with 1 large an'd 3 smaller sensilla.

. Handibles 1.4 times longer7than wide, with blade strongly-curved, about one-third of total length of mandible.

Antenna1 discs ahd ocular lines not risible:

hypopharyngeal region with lateral spines in rows of 4 to 6, and medial spines single or paired. Accessory reticulate e sclarotization not visible; hind end of larval skin without

spines. , Atrium of prothoracic spiracle (Fig. 15) 0,043 mm wide 1 (rang2 0.034 to 0.056) with fine hairs inside. Closing apparatus narrower near atrium, 0.061 ram long (range 0.044 to 0.088) . Average ratio of closing apparatus length to atrium width 1.47 (range. 1. 11 to 1.78).

Phaeogenes gaspesia nus (Provancher) Figs. 10, 16

The 3 specimens examined were reared from the eastern black-headed budworm, Bcleris variana (Fern) (Tortricidae) , -G colLected at St. Alsxandre and Notre Dame de Portage, Que.

Parasite cocoon not visible inside host pupa. Exit b hole at head end of host Pupa, slightly sub-terminal, about

1.25 by 1.00 mm wide.

Cephalic structure of final-instar larva CFig. 10) 0.4 am wide (range 0.38 to 0.44) . Epistoma complete and sclarotized, with 8 pores; pleurostorma sclerotized with lightly sclerotized, dorsolateral accessory p~enrostomal areas; hypostoaa sclerotized, broad and truncate. Clypeolabral plates joined to epistoaa just above superior

pleurostomal processes, each with 2 sensilla. Superior

pleilrostomal process with posterior strut not xisible; inf srior pleurostomal process about as long as wide, trough-shaped. Silk press 1.3 times wider than deep, with opening shallow, about 1.8 timss wider than deep, with 1 -_ seta on each lateral margin. Haxillary palps each with 2 * largs and 3 smaller sensilla; 1 large sensillnm under palp.

~abial-".palps each with 1 large and 3 smaller sensilla. flandibles 1.4 times longer than de, blade st rcngly-curved, about one-third of tot31 length of mandible. . 4

Antenna1 discs and ocular lines not visible; hypopharyngeal surfaca with lateral spines in rows of V to 6 and single or dcuble medial spines. Accessory reticulate sclerotization not Gisible and no spines at hind end of 2 larval skin.

Atrium of prothoracic spiracle (Pig. 16) 0.028 rnm wide

(range 0.024 to 0.03tr) with fine hairs inside. Closing apparatus 0.050 an long in all specimens; ratio of closing apparatos'-'iength tc atrim width 1.83 (range 1.67 td 2.08). Phaeogenes hsbrus (Crssson)

Figs. 11, 17

The 2 specimens examined were reared from unidentified speciss of Lepidoptora collected at Ottawa and Simcoe, Ont. .'

Parasite cocoon not visible inside host pupa; exit

hole at head end of host pupa, fractured, no measurements tak sn.

Cephalic structures of final-instar larvae (Fig. 11) 0.65 rnm wide in on2 specimen and 0.68 ma wide in the other. Epistoea complete and sclerotized, with 10 pores; pleurostoma moderztely sclerotized, with accessory pleurostonal sclerotization surrounding entire cuter border.

Hypostoma sclaroti zed, narrow and long. Clypeolabral platas specimen narrowly, in, the other broadly, attached epistona above superior plenrostomal process by a msmbranous area but with sow scferotization near epistoata, each with 5

or 6 sansilla. hntsrior strut of superior plenrostoaal

r - pracess longer ths n posterior; inforior plenrostomal process trough-shaped, about 0.8 times as long as wide. Silk press 1.4 tiaes as wide as deep, opening about as. wide as desp and 1 pair of setae on silk press undsrneath opening. ~axillar~

* aach with 2 1 and 3 smaller sensillaf labial palps

each with 4 sensilla of approximately equal size. Handible 1.5 times ldnger than wide, blade strongly-curved, about one-quarter of total length of mandible.

Bntennal discs md ocular lines not visible;

hypopharyngeal surface with lateral spines in rows of 4 to

1 6, mdial spines single or paired. Light accessory

reticulate sclerotizstion present at distal end of

hypostoma; and large, single spines at hind end of larval

skin.

Azrioro of prothoracic spiracle (Fig. 17) 0.043 me wide

(ra2gs 0.040 to 0.051) and slightly cup-shaped. Closing

apparatus 0,054 ma long (ranga 0,048 to 0.0641, narrowed near atrium. Closing apparatus about 1.25 times longer than

iridth of atrium (ra~ge1.2 to 1.29) Ger?us Dicaelotus Uesmael

One specimen of one species, Dicaelotus gelechias (ashmead) is described and illustrated; a second specimon of an unidentified species of Dicaelotus was examined but not illustrated. Both differ from Phaeogenes in the length of the inferior pleuro~stomalprocessos which, in ~icaelotus, are l3ng (length/width greater than 1.5) as compared to Phaeoganes (length#widt h less than 1.0) . In Dicaelotus the struts of the inferior pleurostcnal processes are apparently fused dorsally and ventrally form a ftbe while in Phaeogenes the struts are free dorsally and form a trough. No setae are visible on the silk press which is exce

Dicaelotus gelechiae (Ashmead)

Figs. 12, 18

The I s fecimen examined was reared from the gal3 of ecori noscheea \s p. (Gelechiidae) on aster collected at Par~sboro~N.S, 'yn midentified species uas reared from 6.- gallaesolidaqinis (Riley] on solidago sp, ccdlected at aallqs Corners, On', J Parasite cocoon not visible inside host pupa. Exit hole at head end cf host pupa, terminal, not measured \ because fractured.

~inal-instarcephalic structure (Fig. 12) 0 .U2 mm wide.

Epistoma complete, sclerotized, with 6 pores; pleurostoma sclerotized, with narrow, sclerotized accessory pleurostomal area along outer edge. Hypostoma sclerotized, long and 7 Clypeolabral plates fill entire clypeclabral

region, each with 3 sensilla. Superior plrurostomal ~rocess apparently without pcsterior strut; inferior pleurostoma3 . process at least 1.5 times longer than uide; silk press 1.2

times wider thaq deep, opening about as wide as deep, no

setae visible on silk prsss. HaxFllary palps with t large

and 4 smaller sensilla; labial palps with 1 larger and 3

saaller sensilla. Bandibles 0.80 times longer than wide and

tlads 0.37 of total length of mandible.

~ntennaldiscs, ocular lines, and accessory reticulate sclzrotization not visible; lateral s~inescn hypopharyngeal

surface in rows of 4 to 6 and medial ones single or double;

hind and of larval skin with small spines, Atrium of prothoracic spiracls (Fig. 18) a. 030 mm wide,

ovoid, with fine interior hairs; closing apparatus 0.05 0

, 2- mm long with small interior spines near' junction with

' atrium.

Genus Diadromus Wesmael

Two specimens of an unidentified species of Diadromus were examined, and generic and specific descriptions are combined below. The characters that distinguish Diadromus

from the rest of the, Phaeogenini therefore may bs only

specific characters. :

Diadromus sp.

Figs. 13, 19

The 2 specimens examined were reared from the black-headed budworm, Acleris variana (Fern.) ffortricidae) , collected at Sasaginnigax, Man. (PIS 21.2836) .

-- - Parasite cocoon not visible inside -host pupa; exit hole at head end of host pupa, terminal, fractured, no 'C measurements taken.

Cephalic structures of final-instar larvae (Fig. 13)

0.35 ram wide in one specimen and 0.43 mm wide the other. Epistoma sclerotized and complete, with 6 pores; pleurostoma w&thout accesscry pl8urostomal areas. Hypostoma sclarotized, broad and truncate. Clypeolabral plates

lightlp-sclerotized, joined to cpistona by

light ly-sclerotLzed areas, each with 1 sensillurn. Superior pleurostomal process with posterior strut not visible; ,inferior pleurostomal process with struts fused ventrally to

form a trough. Silk press 1.14 times wider than deep, opening 0.9 as wide as deep, 1 pair of setas-located

ventrally below opening. flaxillary palps each with one - * large, 4 smaller sensilla; labial palps each with one large,

3 smaller sensilla. Handible 1.64 times lbnger than wide with blade 0.37 of total length of mandible.

1 Antenna1 discs, oculgr Iines and accessory reticulate scLe~o*izationnot visible; lateral spines on hypopharyngeal region in rows of 4 to 6, medial spines single or paired;

hind end of larvai skin with small spines.

- ~triumof pr~thoracicspiracle (Pig. 19) -0.024 mm wide I in both specimens, with fine hairs inside. Closing

apparatus 0.55 tun, long (range 0.050' to 0.058) ; fine spines inside closing apparatus at end nearest .atrium. Ratio of

length of closing apparatus to atrium width 2.4 or greater.

Genus Aethecerus Wesmael

Two specimens of one u&dentlfied species of

Aethwerus were examixied and therefore genaric and s~cific descriptions are combined below. The characters that define 4 the genus here / may in fact be only specific characters. '-7

Aethecerus sp.

Figs. 14, 20

One of the specimens examined was reared from Trichot aphe cir is=lla Py . (Gelechiidas) collecte-3 at Eobcaggeon, Ont., and the other from an un?&errefie& iorvdcollected at Finland, Ont. (PIS 849-4342) . Parasite cocosn not visible 'inside host pupa. ~lk;it . F .' , . hole at head end of host pupa, terminal, about 2 rrtm in diameter. 'L

The final-instar cephalic structures (Pig. 20) were \ 0.49 ram wide in one specimen and 0.48 mm in the other. .) Epistoma complete and sclerotizsd, with 3 scd 12 pores; pleurostoma sclerotizsd with lightly-sclerotized accessory pleurostomal areas almg outsf de edge. Hypost cma sclerotized, narrow, snd sharply angled away from mid-line. Clype olabral plates lightly sclerotize8 with V to 8 sensilla. Superior pleurostomal process with postprior strut short2r than antsrior szrut; inferior pleurostomal Frocess short with struts fused vsntrally tc form trough.

Silk press 1.2 times wider thac deep, opening 5.80 times

widar than deep, 1 stta on each side. flax~1lar~'palpsuith

1 large and 4 saaller sensilla; labial palps with 1 large

and 3 smaller sensilla. Mandibles 1.4 timts lcnger than

wid?, blade h~oked,0.40 of total length of mandibls.

Antenna1 discs, ocular lines, and accessory reticulate r- L- sclerotiza'rion not visible; lateral spines on hypcpharyngeal 4 id ' b b-4 id. 8-l foher-tribe. AS noted above. the genera Diadrcmus and - ?. Aethecerus of the Phaeogenini have lightly-sclerotized

cly peolabral regions. Perkins (1.959) hoteT that ,

~eterischnusadults were very distinct from the-rest of the ~ba;ogenini-and Tomes (196 parated this genus as a -- tribe, Heterischnini. Houe .the f inal-instar cephhiic-

structures of the Cyclopneu do not .damonstrate the *' divergence seen in -the adults of. the two tribes. On the - n* . - 'of bs basi-s larval structures it -would possi-bly- advisable to - recombine ths two tribes with as an anomalous - \

genus in the single tribe if further examinatien -of -- ,Beterischnus and adults anti larvae supports thfs . * - suggestion.

, + "- A. . .-r.. * . A - Rithin the final-inst& prothbra&c spiracles of the Phkeogenini there is an obvious dichotoiy in the ratio -of -- the length of apparatus to'ehe depth of th2 ,. atriarp. Diadroaus, Hisetus, and the length of the closing apparbtos'is sore than'

tvice the length of. the atrium whereas -in.the genera: atrium. The genu's ~eterischnusappears to be allied with . the former group, which supports the conclusion above. However, as much of this information comes from examination of illustrations ih the lite~ature,final conclusions will have to await a more comprehensive examination of the species.

Figs. 1 1 - 12. Cephalic structures of final-instar larvae: 11, Phaeoqenes hebrns (Cress.) ; 12, gelechiae

Figs. 13-20. 13,14, Cephalic structures of final-instar larvae: 13, Diadrolrns sp. ; 14, Bethecerus sp. 15-20,

Prothoracic spiracles of f inal-instar larvae: 15,

Phaeogenes hariolus (Cress.) ; 16, -P. gaspesianus (Prov,) ;

17, _P. hsbrus (Cress,) ; 18, Dicawiotps gelechiae (Ashm.) ;

19, Diadroaus sp.; 20, Aethecerus sp.

TRIBE PLATYLABINI

Characters have not pet been found to distinguish the tribe Platylabini from the other tribes of Ichnenmoninae.

Short (1978) defired the Platylabini as having the "clypeolabral region with tuo slender and lightly

C sclerotized raised areas appearing as bands or lines, each

with 3 slight dorsal arch; bands may bear sensilla

(Pristiceros) and setae or sensillae groupad beneath band or lineat* This character could not .be distinguished in the thzsa species described below. -- As far as is known all Platylabini are parasites of Geonetridae, with two known Palearctic exceptions, Platylabus tenuicornis Grav. and P. histrio Uesrnael, which feed an Drepana (Drepanidae) (Heinrich 1962b).

Perkins ( 19591, Heinrich (1962a) , and Tounes, Homoi and Townes (1365) placed the Plat ylabini close to the

Phasogenini becaqse of the circular or subcircular propodeal spiracla of the adult bat separated them on the basis of, -b - - aaong other features, the petiole structure. Biologically, 1 the Playlabini can be included in) ge Stenopnausticae

-' because they parasitize ~eoaecridse (Hacrolepidoptera) while aost known species of Cyclopneusticae parasitize flicrolepidoptera. They can also be placed with the

Stenopne usticae because of the st ructure of the clypeolabral plate described below.

-> Two genera were examined and a key for their separation 7- follows. Other genera and species have been illustrated in the literature as follows: &sthenolabus latiscapus

(Thomson) by Short (1370, 1378) ; Platylabus clarus (Cresson) by Short (1359, 1378) ; and Cyclolabus dubious Perkins, C. nigricollis (Wesmael), C_. pactor (Resmael), Dentilabus variagatus Wesmael , Eurylabus tristis (Gravenhorst) , Hypom ec us quadriannulatus (Gravenhorst) , Linycus exhortator fPabficins) , Pf at~fabus punctif rons Thomson, P_. yibratorius (Thulberg) , Poecilcstictus cothurnatus (Graven horst) and Pfistoceros infractorius (Linnams) by Short ( 1978) . Key to genera of Platylabini 7 1. Epistona complete; unsclerotized or

lightly sclerotized. ,, ,,, ,. , ,. .-, , . ., Pf aCglabns,f es~taef Genus Platylabus Wesmael

Two species of the gen* Platylabus are described and

ed and are keyed below. This genus can be i11us7-4 dist~nguishedfrom other gonera ,of Platylabini , except ! ~entilabbsand Enryfabus, by the complete, lightly sclerotized or unsclerotized, epistoma (Short 1978) . According to Short (1978), Eurylabus has the hypostoma 1.5 tims or more the length of the mandible while in Pfatylabus it is less. In both species of Platylabus eramined, the hypostoma is less than 1.5 tines the length of the mandible.

Dentilabus is distinguished from Platylabus by having two very small sensilla close together on both maxillary and labial palps (Short 1378) while both species of Platyla bus exaained lack then. The following key separates the species

Key to species cf Platylabus

1. Epistoaa ansclerotiztd; silk press with a

pair of setae on sclerotized protrusions - f roa its lateral margins; posterior strut - of inferior plenrostomal process much wider t har anterior stre-...... ornatns Prov. Epistoma lightly sclerotized; silk pzess utth a pair of setae located directly on its lateral mar gins; posterior strut of inf srior plenrostonal process about as wide as anterior strut...... rufipes ccnsors Cress.

Platylabus ornatus Provancher

Figs. 21, 25 Pi.>

The 6 specimens of P, ornatus examined were reared from the green spruce looper, Seaiothisa granitata (~uenge)

(Geometridae) , collected at Sa pard (PIS P5 1-2916-01) and '

--1 norsen (YS 55- 165 1-02A), B.C., Ilnisqake, Sask. (i1.1205), u2 Retigoache, N. B., and Victoria Co., N.S. ; and from Hacarip

, sp. (Geaaet dae) collected at Poangs Brook, N.S. t3

Parasite cocoon of sparse, cream-cbloured silk in a single, 7\narro bacd along the dorsnar or venter uithic host pnpa. Exit hole at head end of host pnpa, terminal, about 2 rr in diameter. -- Cephalic structure of f inal-instar larva EFig. 24)

with epistoma complete, nnsclerotized across bridge ; pleurostcma sclerotized, without visible accessory pleurostamal areas. Hypostoma short, about 0.20 am long

\ (range 0.15-0.23, trucate, twisted in some specimens. -4 Cly peolabral plates light ly sclerotizad, eack with 1 sensillurn, not joined to epistoma or joined mcdially. / Suptrior pleurosto~alprocess with only arterior strut visible; inferior plenrastolaal process trough-shaped, about as ,long as wide, pcsterior strut much wider than anterior strut; distance bstveen two processEs averages 0.20 ma (range 0.17 to 0.2 1) . Hypostoma about as lo~gas depth I between pleurostamal processes. Silk press very deep, about three-quarters wider than deep with opening abcut half as--

7' wide as deep> 1 seta on each lateral edge on two lightly- sclerotized protrusions. Maxillary palps usually each with

4 large and 1 saaller sensilla, sometines with only. 3 or 4 sensilla; labial palps sach with 3 larger and 1 smaller C -'% sensilla, sometimes with 3 or 5. Mandible about 1.2 to 1.3 hes ionger than rids; blade 0.33 to 0.40 of total length cf mandible. -- -

bntsnnal discs, ocular lines, and accessor1 reticulate \, scierotizations not ~isib~e;'~~~o~har~n~ealsurface without obvious spines. Hind end of larval skin with saall spines.

Atrium of prothoracic spiracle (Pig. 25) about 0,049 ram vide Irange 0.0&0 to 0,051r) with fine hairs icside, Closing spparatus about 0.077 rr long (range 0.070 to 0.030) and protruding into atrim.

Platylab us ruf ipes consors (Cresson) Pigs, 22, 26 .

The 2 specimens examined were reared from Hydriomena sp. f~sanetridaefcalfected at 8aide Creek B.C. (FIS

B.C. 5 1-2525-B) 3rd an unidenti fied species of Geometridae collected at Francis Lake 0.C. (PIS 51-30808) respectively.

Parasite cocoon of loose, cream silk along venter which appears to close off reconiam at rear of host pnpa. Exit bols at haad end of host pnpa, terminal, 2 to 2.5 mm in Ce~halicstructure of final-instar larva (Fig. 22) 0.48 , nr vide in one speciaen and 0.60 ma wide in the other.

Epistoma complete and lightly sclerotized with 6 pores. Plsurostoaa sclerotized, with accessory pleurastomal sclsrotization only on lowsr edges of pleurosto~a;hypostoma

abont 0.25 nB long, truncate, not twisted in either

specimen. Clypeol abral plates lightly sclerotized, each with 1 sensillura in sclerotiz'ed area and 2 sensilla on membrane below, not joined to epistoma or joined medially. Suparior pleurostomal process not visible, inferior

pleurostomal process abont 1.4 times as wide as deep,

trough-shaped, uith two struts of equal width. Depth

betusen the two prqcesses is about 0.20 mm and the hypostoma is about 1.25 longer than the dzpth between the processes.

Silk press very deep, amnt 0.7 times as wide as deep,

T. spening about 0.4 times as wide as deep; setae on silk press

more ventbally situated than in -P. ornatus and not located on lateral protrusions. Baxillary palps uith 5 sensilla of

abont equal size, 2 or 3 of these apparently setiform;

labial palps with 5 sensilla cf abont equal size, 3 of these setiform. Mandible about 1.2 times longer than wide, blade not distinct, abont0.39 of total length of mandible. Antenoal discs, ocular lines and accessory reticulate

scleratization not visible; h y popharyngeal slirface without

spines, Hind end of larval skin with small spines.

Atrium of prcthoracic spiracle (Fig. 26) 4.061 mm wide

(range 0.044 to 9.056) with fine hairs inside, beavy

sclerotization on surface. Closing apparatus 0.110 am long (rangs 0.090 to 0. 130). Q

Genus Cyclolabus Heinrich

Only one species of Cyclolabus was examined and genaric

and specific descf iptions are combined below. The -. incoaplete episto~asets this genus apart fron Platylabus. Additional characters which may be useful are: the

clypsolabral plates ars distinctly triangular; thq accessory

plearostomal sclerotization extends the whole length of the

pleurostoma; the posterior strut of the inferior mandibular / ? process is much longer than the anteripr &strut; there are no

sea4 on the silk Fress; and the openi& is about as wide as

deep. Cyclclabos dubiosus Perkins

Pigs. 23, 27

Th? 2 speciiens of G dubiosus exari3ed vere reared f roa Eopithecia sp. (Geoaetridae) collect@d at Squ asis5

Iiivsr allay, B.C. (PIS BC47-1959A) and ItIsland Core Brookw

Hfld. (PIS U3 L538).

Parasits cocoon of sparse, fine silk along venter inside

hcst pupa. Exit hcle at head end of host pupa, terminal, .j 4 1.5 to 2 mr in dianeter.

cephalic structure of final-instar larva (Pig. 23) with incomplete epistoma; pl~urostonasclerotizsd, wit& unscleratized accessory pleurostomal area along edgs of pleurostoma. Hypostoma broad and truncate, about 0.17 ma long (range 0.15 to 0, t 8) . Clypeolabral plates small and i triangular, not joined to epistoaa or joined mediallp, each it with one sensillnr , located dorsally on plates. Posterior

= > strnt of superior pleurosto~glprocess not visible, anterior strut protruding; ~csteriorstrut of inferior pleurostomal

- - - process much longer than aniorior strut, process about twice as l~ngas wide. Ceph between tuo processes 0.135 no fraage 0,12 t~o0.14) ; hypostora 1.07 to 1.38 tines longsr than 35pth bet uoeo p l~nrostomalprocesses. Silk Fress

about as vide as deep, wide as shallow, opening about. . as deep, uithcut setae. Maxillary palps with from 3 to 5 sensil3.a of equal size; labial palps pach with 4 setiform sensilla of equal size, Ma~dible1.0 to 1.2 times longer than vide; blade of mandible about 0.33 to 0.40 of tctal fen gt h of nandi ble ,

Antenna1 discs, ocular lines and acgessorp reticulate scltrotization not evident; no spines oi sculpture on hypopharyngeal surfact. * Hind end of larval skin without visible spines.

Atrium of prothoracic spiracle (Fig. 27) 0.036 mm wide

{ranga 0.034 to 0.040) with no spines or hairs inside. Closing apparatus 0.052 ma long (range 0.050 to 0.056) . TRIBE LISP ROD ROHINI

Genus Anisobas Wesmael

on4 species, hnlsobas luzerniensis, of the tribe Listrod romini has been examined and illustrated. Therefore tri bal, generic, and specific descriptions are combined below.

Anisobas sp. and Listrodromus nyctheaerus (Gravenhorst) have baen illustrated by Short (1953, 1978).

I Both species illustrated by Sho~tand -A. lnzerniensis show the character which defines the tribe according to

Short (1378) : nthe blade of the mandible is one-third or less the length of the mandible and the base of the eaandible is very broad. Species of Hoplismenus, Orgic hneumon, and

9. Trogus have very similar mandibles with blades less than one-quarter but greater thm one-fifth the total mandible length. In Bnisobus sp. (Short 1959, 1978), and 8. luzerniensis the blade of the mandible is less than one-Mfth the length of ths aandible which provides a reliable distinguishiag character for the genus. However,

Listrodtomus nycthermPrus (Short 1959, 1979). bas a mandible blade which is about 0.3 the total length of the mandible.

Eo~lismenushas a ventral protrusion on the silk press,

uhich .Listrodroiitus does not; Orgichneuaon has 6 sensilla on each maxillary pal^ while Listrodromus has 5; and Trogus has ) characteristically bent hypost cmal sclerites, which

Listrodromus does not. The most characteristic feature of

Listrodromus as illustrated by Short (1355, 1908) is that the blade of the mandible is bent sharply upvards in the anterior aspect. This is possibly an artifact of mounting.

. . lnisobas luzerdre~sis(Bradley)

Figs. 24, 28

The three specimens examined were reared from Incisalia niphon (Hbn.) (~ycaenidae)collected at Hoar Lake, Ban. (PIS

1-1654-02) and Canterbury, I. B. ; and from Lycaonopsis pseudargiolus Boisd. & Leconte (L ycaenidae) collected at

Bart, B.C. (PIS 50-1001C). -- Parasite cocoon of sparse silk except for a solid mat on ma side of interiar of host pupa. Exit hole about 3 mm in diaaeter.

Cephalic structure of final-instar larva (Pig. 24) 0.59 in me specimen and 0.67 EH ufcle in the other, un~easurable in the third. Zpistona lightly sclerotized, partially incoapiete medial1 y, with- two pores; pleurostoaa sclsrotized, with light accessory pleurostonal sclarotization. Hppostoaa short, about 0.21 nm long (range

0.18 to 0.24) and atout same length as distance between plsnrostoaal processes. Clypeclabral plates large, almost filling clypeolabral region and joined medial1y, tach with fron 8 to 10 sensilla, two of which are raised. Soperior pLeurostcma1 process not visible, inferior plearostoaal process wider thaa long, trough-shaped. Silk prlss aDout

0.6 timss as deep as wide and opening about 0.B times as uide as deep, a pair of setae under vsntral edge of opening.

Yaxillary palps each with 4 to 5 sensilla of equal size; labial palps each with 4 sensilla of about equal size. In the specimen from Hart, 3,bath pafrs of pafps are elliptic, with the sensilla c&ustered a* ow? em3 its il2!1szrated (Pig. 24); in the cther two spscizsns all palps

are r~nndwith the sensllla-distriboted evenly over snzfacs.

This imp1ies the existence of geographic vari3tion that

iiga: uarrant fnrthsr investigation. Handible about 1.3

times longer thz~uidz, blade 0.13 to 0.20 cf total lsngth

of sandible. Oze specisen had deformed mandibles bu'., could

still be recognized as Bnisobas by the fcrs of the epistoma and clypeolabral plat?s.

Antenna1 discs md ocular lines present, but only

lightly scler~tize6. Ocular lines about 7.75 times long21

than depth between pleurastomel processes and withcut

reticulate sculpture on their surfaces. Accessory

reticulate sclerctizatian not visible; hypopharyngeal

surface without spines. Rind end of larval skin with sm311 spines.

Atriu~lof prothoracic spiracle (Pig. 28) 0.055 mm wid*

(rangs 0,052 to 0,056) with fine hairs insi3e. Closing

apparatus 0,085 am long (rangt 0.080 to 0.030) and * co~strictednsar its junction with> atrium. Figs. 21-22. Cephalic structures of final-Fnstar larvae:

21, Plat ylabus ornatns Prot. ; 22, P. rufipes ccnsorz Cress.

Pigs; 23-28.!' 23.24, Cephalic structures of final-instar

't* '?P larvae: -23, Cyclolabu~dubiosus Perkins; 24, Anisobas lozerniansis (Brad.). 25-28, Prothorac2c spiracles of final-instar larvae: 25, Platylabus ornatos Prov. ; 26, g. rofipes consora Cress. Cyclolabus dubiosus Perkins;-28, - - Anisobas lazerniensis (Br d. ) . far their s*parstiaa foZlors. 'i.hs tribe fchoeu~n'iaiseers

maera of fclncbomrrin~not discussed beloo but calamagrostidkis Beinrich, Em ischioranthus (Gravenhorst) , B, ulbrichti Hinz, Hepiopelmas mlanoqaster (Gmlin) , 1 - i;* Platylabops puchellatus (Bridgaan) , Probolas culpatorius ," -

Cameron, Bobicundiella perturbatrix beinrich, Stenichneumon

I capator scbrank, and Tricholabus strigatorins '(~raienhorst) by Short (1978) .

. Rep to Genera of Ichneumonini

Cljpeolabral plates joined medially or- each as wide as deep and filiing the clypeo- labra1 regicznanUsnall~t~uchin-g- m#iallg...... ~...... 2

plates-not foined madial.Jp, _each - - - -p - wider than deep, not f ilfidg clypeolabr a1 region and not usually touching msdially...... 9

,

~o-uermar gin of silk press extends ventrally into a lobe or spine-like process,. ...,...... 3 Lover aargin of silk press dues not extend - ventrally into a lobe or spine-like process...... 6 3. pair of setae on silk press located on lateral margins above ventral margin of opening ...... mwHoplis~e~us . Pair of setae on silk press may or may not be on Xateral margins but are al~aysbelow # ventral margin of opening...... m...... V

- -- - - p------U. tength of ocularplines more than trice depth

between plearostoaal processes and width of a- ocnlar lines about equal to depth between pleurostoaal processes...... I,imerodops ? Length of ocular lines less than twice depth

--a --a ?IFwidth of ocular lines less than depth between

5. Bedial join between clppealabral plates very indistinct, almost invisible; antenna1 discs and ocular lines aluaps present., ...... Thyrateles aedial 'join between clypeolabral plates

distinct; antenaal discs and ocular - \ lines sometimes not ~isible,...,..,~..~ .....Ichneumon Cbypsolabral plat$s about as wide as deep ...... 7 Cly peolabral plates much wider than deep. ..Orgichnenmon .

Length of ocular lines greater than 2 times depth between pleurostomal

-

Length of ocular lines less than 2 times . depth between.- ple~rostomalprocesses...... Entan yacra

Episto~aincomplete and/or unsclerotized...... Diphytis pist to ma complete andLsclerotized...... Spilichnenmon

Width of atrium of prothoracic spiracle about

~~~t~y~ar . .eA*a width of atrium of prothoracic spiracle less e

than length of closing apparatus,. .r .., ...... It3 - -

Clypeolabral plates not apparently joined to epistoma laterally,...... I1 Cly peolabral plates apparently joined to 11, Rithont sclerotized antenna1 discs,. .. , . . . . Patrocloides With sclerotized antenna1 discs...,...... , ..Thyrateles

'12. accessory reticulate sclerotization heavy and in a perpendicular line leading auay from &&& &&&

Accessory set iculate sclerot ization light, usually not in such a definite perpendi-

------cular line at distal end of hypostoma.. ,Cratichneumn

~einrich( 1967a) divided the tribe Ichneumonini into 5 '.r subtribes: Ichnen monina, Aab lytelina, Hoplismenina, Wthioplitina, and ~ratichnenmonina. The congruence of the final-instar larval class th the adum classification is total, but some of the species can be

- ifif ferentiated only with aifficnlty. Specfaens of ' Aethioplitina were not available.

Heinrich ( 1967a.c; l968a, b) designated the ~abtribesof 1chneawnini.on the basis of African fauna. Most of the

I: miah h*r+a are -= s b~ ~einrich(1963, 1371, 1972. 1375). Those not

placed by Heinrfch can be assigned to subfamilies using

- Hainzichts (1972~) key to the subtribes of the Ichnea8onfni. The cratichnenmonina, represented here by

Cratichneuaon, Vulgichnenmon, and Aoplns, have the

b clypeol&bral plates joined- to the epistoma. This character i~ indistinct in AOP~US; the Aeblptelina and Ichneurotrina do not have it,

The Amblytelina, represented here by Diphyas, Limsrodops, Spilichnennon and Ent an yacra, seem to have

------specialized as parasites of iloctuidae. The final-instar larvas have large clypeolabral plates which are as deep as vide or deeper, filling- the clypeolabral region, and which * may, or map not, be joined tedially, w-- ,

represented hare by patrocloides, Th~rateles,Icbneomon, and

respects, internediate betueen those of the other two subtribes, In noctuid-parasitizing _I[chnenmon the

clypeolabrql plates are lafge, bat nct as wide as deep, and

joined mediallp, similar to the Amblytelina. In the other species of Ichneumon and, in Org%chaennon they are sraller,

not as deep as wide bat joined mediallp, which is distinct from tip .other tuo sabtribes, Species. of hyratele? and ~atrocldideshare clypeolabral plates which are small, .not,

joined medially, and wider than deep; sometimes in Thyrateles they are indistinctly joined medially, These 2

species have in common the amblypygous apex of the abdoasn,

The Hoplismenina, represented here by Hoplismenns, have ------clypeolabral plates which are not as deep as wide and are.

joined medially. The blade of the mandible is short and

hooked sharply. This combination of characters sets the

snbtribe apart.

--- -- anus Hoplismenns Gravenhors-

Two sub-species of Hopl-islenus moralas ate-describ=& --- and illustrated and a key for their separation follows. Hoplismenus bidentatus (Gmelin) was figured by Short (1 978) . . . .

The characteristics of the genus" are; clypleo-labral and strongly hooked in ventral silk press greatly

elongated ventrally; and a pair of setae on silk press - d located on- ~ateralmargins of silk press above ventral. -. I < margin of opening of silk press. -

-- -. bidam€amw as nlost-raten~short(i9';rT8) does not complete1y agree with the above description, t bough the

clypeolabsal plates, though not figured as being joined

medially, are broadly triangular, each with 13 sensilla,

The silk press is not illustrated as ventrally produced and

I the setae on the silk press are located below the ventral marcjin of the opening.

Key to subspecies of Hoplismenus morulns

- -

-% -

1, Anterior strnt of inferior plensostomal

-J' process longer than posterior strut * ...* ...... e.....m~rulu~ pacificus Say Anterior strnt of inferior pleutostomal - ing cloak butterf lg. Symphalis antiopa (I.) (Tymphalidae) . collected at Candy Lake flan., and f roa an unidentified species of Bymphalidae collected at an unknown location in 1

Parasite cocoon of loose but relative1y dense silk. I extending as far back as the aeconium and all arcund the

inside of the host pupa. Silk more yellow and darker than that of H. & gacificus. Exit hole at head end of host - ! pupa, terminal, 5 to 6. mm in diameter.

Cephalic struc'turs of final-instar 'larva (Fig. 29) 1. I6 t mm ride in oneppecimen and 1.24 mm in the other. Episto~a I sclerotized, with 12 or 14 pores; plearostoma sclerotized with narrou accessory pleurostomal sclerotization on outer margins. Hypostoma long and sharply bent in one speciaen, as figured, and shorter and not as sharply bent in the other.

- - more like -8. a. pacificus (Pig. 30). Clypeolabral plates broadly triangular and joined Mially, each with 9 to 12 sensilla. Snpeh ior pleurostoaal process with posterior process not visible; inferior plenrostomal process with anterior strnt ,shorter than posterior strnt and pointed, struts free. Hypostoma 1.5 to 1.9 times longer than depth between pleurosfoaal processes, depending on specimen. If

times depth between pleurostoaal processes, is measured to

the point where it bends, the hypostona is 1.5 times longer than the depth between the pleurostoral processes. This is . the s3me as the ratio in the specimen with the shorter hypostoma, which indicates that the section distal to the

bend in the one specimen map be an abnormal extension of the hypostoma. Silk press abont as wide as deep and projecting

-ventr&lyxra-krmnd sp ar , dar52[-~argin-s--af-str~-~res~ projecting up and over opening which is about one-quarter

deeper than wide, sets on each lateral margin abcve ventral margin of opening. Baxillary palps each with 2 large and 3 saaller sensilla; labial palps each with 1 large, long

I sensillum and 3 ssaller sensilla or with 5 sensilla of equal sizs. mandible abont 1.4 times longer "than wide, blade Antenna1 discs lightly sclerotized and apparently

0 irregularly shaped. Ocular lines sometimes visible, lightly

sclorotf zed, with no obvi-ous reticulate sculpture.

Hypopharyngeal surface with single medial spines, and

lateral spines in rows of 4 *to 6; light accsssorg reticulate

% -- ~irotization visible, originating f roi distal end of hypostoma. Hind end of cast larval skin withost visible

- -- - - spines. ------

Y

Atrium of prothoracic spiracle (Fig. 55) 0.075 mm wide (range 0.070 to 0.080) vith some heavy spines but no fine

hairs insida. Closing apparatus 0.110 to 0.130 mm long

(mean 0. 123).

Hoplismenas morulus pacificus Say

- -. - Figs. 30, 56

The 2 specimens examined were reared .from Ilgmphalis caliiornica (Bo~s~uQ~~)(lymphalidae) collected at

ootischentts B.C. (PIS 62-6616-12 i) and- &OR a species of

-- --~~~eeallected at Langtord, B.C. (FIS 250-2458}. Parasite cocoon of coarse, light-colored silk, around

inside of host pupa, Exit Bole at head end of host pupa, I terminal, abont 4 rr in diameter.

Cephalic structure of final-instar larva tfig. 30) 0.99

-vlae~~%S~~a an6 i/TT"mrin the other, Epistora sclerotized, narrower than that of H, &, aornlos, with 11

or 12 poEes: plearostora Sclerotized, with light accessory ------pleurostomal sclerotization around lateral margins of

(* pleuostoma. Hypostoma shorter in relation to depth betwew 4 pleurostoral process than in 8, 1. *ornlus; not bent sharply but with some f rsgnentarp sclierotization at the distal end

in one specimen, Clypeolabraf. plates joined mdially, not as .

deep as those of -H. -a. mornlns, each with 9 or 10 sensilla. Supsrior pleurostomal process with posterior strut not

risible ; inferior -process with anterior strut lopger and

much narrower than posterior strut, struts free, Silk press

1.33 tims deeper than wide, .with ventral margin produced 1- into a long, narrow process, opening of silk press abont as

wide as deep, seta on each lateral margin above ventral sensillnm is setifor.. ~andible1.4 times longer than wide - with blade about one-quarter of total mandible length.

Antenna1 discs not apparent; ocular lines lightly scl8rotized. not always visible, with no reticulate -- sculpture apparent on surface. Hypopharyngeal surface with

single spines only; light accessory reticulate i

- sdarat ization originatim~atm M--- - 1

end of cast larval skin without- visible spines. -

* ~trinrrof prothoracic spiracle (Pig. 56) -0.076 mm wide (range 0.070 to 0.080) with heavy spines and a feu fine

hairs inside. Closing apparatus 0.124 to 0.133 mm long

Genus Valgichnenaon Heinrich

~ulgichneu.on bkvicinctor (Say) is described and

ill ~stratedand generic and apecif ic descriptiens are

coabined,' Short (1453, 1978) also figured Vulaichnen.on -- * @evicinctos and Rothschild (1374) figured Vnlichneamon sp. nr. leacaniae (Bchidaf , plates not joined redially, saaU and elongate-owdl-; length of closing epparatas aliost twice as great as uidth sf - ,+ ,+ atrinr; with saaL1 spfngs in raws on hind end of larval

skin; and the ~zui3far-ypips each usttall~with 6 seasi3;Xa -fi

-- LC*' ---63, 2I .

+ The 2 specimens examined were reared fro. the bertha . + , -

- Parasite cocoon not apparent inside host ppa. Exit

- -- - - hoie at head end of host pupa, tekminal, about 5 ;D in *

Cephalic structure of firtabinstar iarta *pig. 31) 1.0

mm vide irt both specimns, Bpistora sclerotized, with tO. or sclerotized, ritb .many scnsilla. outer aargin of plaurostoma

indistf nct; hypostoaa sclerotised hnd short. Clypeolabral plates not joined ~ediaXly.small and elongate-oval. each ritk 7 to 11 smailJa. Soptior pleurostomal grucess uith poster;iat strat not risible, anterior strut slightly

half as wid8 as deep, strots abqtzt same length and hot fused P renttally, Jmse of anterior strut narrower than base, of

apning. Llsxilfarf paps each with 5 or 6 sensilla, if 5, I sh3: labial paps each uith 3 large and 3

wida, blade ahat half of total length of mandible. -

em-this6 longer than depth betueen pleorcstolal processas. - - rratiutlat e scolptoxe on surface, Bypapharyngeal

ef ELjpu~tum.' Spi-s OB hind end of cast larval skin small Atrium of prothoracic spiracle (Fig. ' 57) (I .OS7 ma* vide (range 0.056 to 0,060) with some short heavy spines but no fine hairs inside. Closing apparatus 0.084 mm long (range

0.076 to 0.090)

Genus Cratichneumop Thoason

Seven species of the genus Cratickneumon are described

------p - - and-illustrated and a key for their separation follows. The

variation in -C. pteridis, of which 31 specimens were

availible, is analyzed as an example of intraspecific variation in the ~chnenmo@iinae.

-C. coler (~hlar)by Sechser (1970) and Short (1378) ; . Cnttichneunon mifascsatarfns (5q) By Short f t6S9, 1378) ;

and -C. corrnscator (Linaaeas) C. dhplicatus (Say) C. , , -- fabricator (~abricius)and -C, varipes f6rarenhorst) by Short Generic characters are: clypeolabral plates not -joined medially and usually narrowly oval; closing apparatus of prothoracic spiracle longer than width of atrium; spines at hind end of body: and accessory reticulate scleroti zation light.

The clypeolabral lates of all species of Cratichneumog 'p have vestiges of a jci to the epistona, This raises the ------possibility of confusing some of the ~ratichneamonwith species of Phaeogenini, and ~haeogeheshebrns vith species of rati ichneumon, as discussed previonslp..

L

Key to species of -Cxatichneumon

1. Epistol~albridge joined ta lateral sections

- - of epistoma by evenly sclesotized areas...... 2 Epistomal bridge joined to lateral sections of epistoma by lightly sclerotized or spines on hypopharyngeal surface.. .gtnisotae ~einr. '* Y without spines on hypopharyngeal surface...... m..~.sublatus (Cress.) a 4 d pnif asciator ios vancouveriensis (Prov. ) t rmspines on hypopharyngeal surface...... 4 Hitboilt spines on hypopharyngeal *, *,

surface.. .,c..sp,-nrrse(Prorr.l------,...... - - --

4. Hypopharyngeal surface with fewer than 40 spines, all single and large; only one pair of setae on silk press...... pteridis Tow. Hypopharyngeal snrface with more than 40 spines; or sowe in rows, or 2 pairs of setas on silk press...... m...... m.m.~...... 5

5. Blade of mandible more than 0.40 of total length of mandible; I pair of setae on silk press; cephalic structure more than k 1.0 mm ride...... varioqatus (Prov. ) 0 Zixt5e of aanaibfe less than 0.40 of total

setae on silk press; cephalic structure less than 1 .O mm vide...... ritss Eeinr. The larvae of Cratichnenmon separate into 2 mofphological groaps: those with a separate epistomal bridge

joined to the lateral sections of the epistoma by

nnsclerotized or lightly sclero~izedareas; and those with a

a ~~t~~ OL a unsclerotized areas. These two groups correspond roughly to geometrid parsi tes and non-geonet rid parasites, ------respectively, although -C. sublatns is an exception in the lattsr group am3 C. varieqattls in tlie former. Separation

of the species is difficult past this point.

lon-overlapping characters were not found for 5. y.

yanconveriensis and C. . snblatus, although a large saaple of these two species might reveal distinctive characters.

------Cratichneunon trni fasciatorius vancouveriensis (Provancher). Pigs. 32, 58

-" The 2 specimens examined were reared from Syngrapha sp.

(loctnidae) collected at Kitimat, B. C. (PIS 62-843- 021) and

C fa -- fro UL LL

D Parasite cocoon not visible inside host pupa. Exit

hole at head end of host pupa, terminal, about 5 mm in dianeter.

Cephalic structure of final-instar larva (Fig. 32) 0.85

measurable, about one-f onrth to one- third larger than the specimen measured. Epistoaa sclerotized, with 5 pores in

* ------pppp ------one specimen and 11 pores in the other, with sclerotized connect ion to epistomal bridge. Pleurostoaa sclerotized, with accessory pleurostomal sclerotization around lateral * margins and extending more than half-way down hypostoma; hypostoma sclerotized, with slight upward projection at

-tEstz&umf- -~~p~mra-lp~at-e-~S~~~eachwlth 4 or 6 sensilla, not joined to epistoaa. Superior pleurostomal process with anthior strut prominent, posterior strut concealed; inferior pleurostomal process with struts free,

posterior strut about twice as long as anterior strut. Silk

press abont 0.9 times wider than deep, with brogd ventral projection about twice as deep as depth of opeqing, opening

tit one fm+-hwi* 1 =+a 0-h - '. labial palps each vith 5 sensilla, ~andible1.13 ti'mes longer than wide and blade about one-third bf total "length of mandible, vith a slight bulge in the middle ,of .the blade after which itMnarrousrapidly to the tip.

hypopharyngeal snface without spines. AcCessOr~reticulate sclerotization very light and originating from the distal ------end of the hypostoma. Large, single spines on hind end of larval skin.

* Atrium of prothoracic spiracle (Fig 58) 0,064 or 0.084 am wide with short and long hair-like spines inside,

cratf chneu~onsnblatus Cresson 0

Pigs. 33, 53

he 8 specimens examined were reared from Erranis Y, 9 tiliarlh+~arr.) (Geosetridae) collected at st. Raphael Co., @iarton (FIS 695-0040-0 1- 13), Burks Palls (PIS

695L0037-01-3) , Dorset, fleaford (PIS 68-5-0058-0 1) , and

\-

Orville (PIS 635-0064-01-I), Ont. 3 --

Parasite cocoon of dense, coarse but loose silk around

----ent-ire-ias~nf~U-sg-ba&~-.+)ln.

Exit hole at head end of host pupa, te inal, about 4 rnm by 5 mm in diameter. f ------

Cephalic itructure of final-instar larva (Pig. 33) 1.11

em wide (range 1.00 to 1.25). Epistoma sclerotized, broad, usually with 8 pores (7 or 9 exceptional) , with distinct epistonal bridge which is ccnnected to sides by sclerotized

-meas- - P~-~rostom-a~s-c~erOt~~e-d~wlthsclerotlze d accos sor y

pleurostomal area around lateral margins; h ypostoma

sclerotized and broad. Clypeolabral plates lQhtly -

sclerotized, not joined medially, each with 3 to 9 (usually

5) sensilla. Superior pleurostomal process projecting with anterior strut longer than posterior strut; inferior

plearostomal process with struts of about the same length, F

and projecting ventrally, opening about as vide as deep, dorsal margins of silk press extending above opening and one

.?a, L seta . on each lateral margin of silk press, about parallel "h with dentral margin of opening. Maxillary palps each with 5

or 6 sensilla, 1 or 2 very small; labial palps each with 5

sensilla. Mandible about 1.1 times longer than wide, blade

--_ahout ~.br~~O.J4& 0 t6ta3l-.length-o-nf- mandible.

------~ntennaldiscs and ocular lines not visible,

hypopharyngeal surface without visible spines. Accessory

reticulate sclerotization not visible. Spines on hind end of larval cast skin large and mostly single.

- --kt ~h~--~f--prOthO~5 9 8 6 8-&B

(range 0.060 to 0.080)' with fine hairs inside. Closing apparatus 0.106 mm long (range 0.084 to 0.340) .

Cratichneumon anisotae Heinrich

Figs. 34, 60

The 4 specimens exaained were reared from

rabicanaa (F.) (~itheronidae)collected at Echo Bay, B. C.

(PIS 549-26-2,4,5) and Jocelyn Tp. Div. 36, [?B.C. ] (PIS Parasite cocoon not visible within host papa; exit hole at head end, terminal, about 5 mi in diametsr,

Cephalic structure of final-instar larva wig. 34) 1.15

------am - wide (range 1,tQ to 1.74). Xphtalaa- scleratizan--3 to 13 pores, connection between bridge and lateral sections of epistoma sclerotized. Pleurostora sclerotized, with

------accessory pleuro~~omalsclsrotization along lateral margins; hypostoma broad, Clppeolabral plates not joined medially, oval and curved, each with 2 to 7 sensilla. Snperior pleurostomal process with anterior strut projecting and longer than posterior strut; inferior plenrosto~alprocess

*tk st-rtrts-ap ~~~~rus-a~l- - fyi-t-g*md, anterior strut wider and deeper than posterior strut. Silk press with indistf nct projection ventrally, opening about twice as deep as wide; 1 seta on each lateral margin above ventral margin of opening. ~axillarypalps each with 5

sensilla, usually 4 in vertical row; labial palps with 5 . - larger palps and 1 very small palp. Bandibles about 1.5

--- times longer than wide, blade of mandibleab 0.47 of

total length of mandible, with slfsht bnlcre near tip. - antenna1- discs and ocular lines sometimes visible; if 2- HSF2 -L *. they are, then very indistinct. Hypopharyngeal surface with scattered, single s~ine3; vithout accessory reticulate

sclerotization at distal end of hypostoma, Spines on hind

end of larval skin large and mostly single. -- --

Atrium of prothoracic spiracle (Fig, 60) ~0.072 mm wide ult6 tine liarss-inside,- Closing - [range 0.m to 0,088) .

apparatus 0.135 am long (range 0.110 to 0,160).

Cratichnenmon varieqatus (Provaneher)

Figs, 35, 61

.:+-, The 5 sppcimens examined were reared from the saddled prominent, Heterocampa gnttivitta (Ulk.) (Mctodontidae) , collected at Orillia, (PIS 635-0060-01-2) and ~igrton.Ont., (FIS 635-0040-01-1-,14; 695-0665-01-3) and from Anisota

rubiconda (P.) (Citheronidae) collected at HGdemeya 167,

f ?Ont, f (PIS 248-6871-5). i Parasite cocoon represaGd by a few strands of fine, clear silk usthin host pupa; I exit hole at head end cf host pupa, terminal, 5 to 6 ma in diameter.

Cephalic structure of final-instar 1.(Pig. 35) 1.14

A-p-- ma ride lraage l,t2 tCL__1_*2L_3$,EaistMhscLaraAize&,-with_tO to 14 pores, bridge distinct but connected to lateral areas by sclerotized areas. Pleurostoaa sclerotized., surrounded

------lateral1y by accessory p+urostomal sclerotization; hypostoma sclerotized, Clypeolabral plates distinctly

rectangular, each with 4 to 8 sensilla. superior pleyostomal process with posterior strut not visiblb; in•’erior plenrostomal process with anterior strut wider than

--Ypst~i;-s t-rwtT--Stf-tfts-f-&hb~~3.pngt~~e.

Silk press indistinctly extended ventrally, opening about 0.20 tines teeper than wide, uith a seta on each lateral margin above ventral margin of opening. aaxillary and labial palps each with 1 large and 4 smaller sensilla. Mandibles about one-eighth Xonger than wide, blade about 0.42 of total mandible length, Antenna1 discs and ocular lines not visible; hypopharyngeal surface with, single spines in transverse rows, Accessory reticulate sclerotization not visible; spines on bind end of cast larval skin large and mostly single.

Atrium of prothoracic spiracle (Fig. 61) 0.081 mm wide

(range 0.076 to 0,090) with long, heavy spines inside, fine

Cratichnenbon ritns Heinrich

Pigs. 36, 62

The 3 specimens examined ' were reared from Erranis

tiliaria (Harr, ). (Geometridae) collected- at St, Bnbert (PIS

362-347-01, #l9,#32) adSt. , Raphael (PB 363-682-01), Qne.

Parasite cocoon not visible within host pupa, exis hole at head end of host pupa, tetrinal, about 4 by 3 mm wise. Cephalic structure of final-in&ar larva {Pig. 36) 9.80

m~ vide (range 0.75 to 0.85j. Bpistoaa rclerotized, rith 8 ,

or 3 pores, epistumal bridge joined to lateral portions of

epistoma by lightly sclerotized, alnost menbranous area. Flewostoma sclerotized uitb accessory pleurosto~al

xferotization extending around f atera1 margins; hypostoma sclerotf zed and broad, Clype~labralplates irregularly

,rhoabic, each with 7 sensilla plus* 1 just below plate.

tisible, anterior strnt protrading; inferior pleorostomal process wfth anterior strat triangular and shorter than posterior strut, strnts either free cr trough-shaped. Sf lk /- j press blunt rentrally, not produced, aboat>as wide as deep, s opening about one-third deeper than vide, two pairs of setae

on later'al margins of silk press in tuo of the speqimens, &e pair in tbe other. Saxillary palps-each with 5 04f sensi&&la; labial palps each with 5 or 6 sendlla. landibles . -. r about 1.2 tiae%t'longer than uide and blade about 0.36 of

Aatennal discs and ocdar lines not risible;

8 hypopiaryngeaf surface rfth scattered angle senes. .- &ccessery reticelate scleratization not visible; spines at

hind end of cast larval skim large and mostly single. -

Atrim of pretbotacic spiracle (Pig. 62) 0.046 mm wide frange O.WO *p 0.058) dth-fine hairs inside. closing agparatas 0.070 rm long (range 0.050 to 0.090) .

The 2 specimens exaained were reared from' the lesser

maple spanworm, Itase pastofat'ia (Gven. ) (Geonettidae) e- collected at Tay Creek, Turk County, B.B.

at head end of host paps, terriaal, about .2 ms in diameter.

C cephalic structure of final-fnstar larva @kg. 37) O.&9

u wide in one specimen and 0.52 mm in the &Her. bitpistoma

4 scl.ro$ized with 9 pores, epistoml brvge separate4 from -4 lateral sections of gpistoma by membsadb48 area. Pleurostom sclerutieed with dorsolateral and- ventrolateral accessory pleurostomal sclerotization or with narrow 2 accessory pleurcstomal sclero tization around entire lateral _P margins; hypostol~asclerotized. Clypeolabral plates broadly

triangular, each with 4 to 6 sensilla and with' double and setiform sensilla located on skin of top of plates. - i Supsrior plenroslto ma l process projecting, posterior strut not visikle; inferior pleurostomal process with anterior

strat shorter tha~posterior strut, and triangnlar. Silk press indistinctly projecting ventrally, about one-quarter wider than deep, 1 sqta on each lateral margin above ventral margin of opening. ~axillarypalps each with 5 sonsilla; * - labial palps with 4 sensilla, or, if with 5, 1 extremely

small. Mandibles about 1.30 times longer than wide, blade.

about one-third of total length of mandibls, with no bulge near tip.

Antenna1 discs and oculkr lines not risible; no risible

spines on hypopharyngeal surface. ~ccessory'reticulate scl2rotization not visible; spines on hind end of larval skin large and mostly single. - '. atrium of prothoracic spiracle (Fig. - 63) 0.036 rnm wide, with fine hairs inside, Closing apparatus 0.050 mm long.

All specimens identical.

Cratichneumon pteridis Townes

'P Figs, 38, 65

The 28 specimens examined were reared fsom Semiothisa sp, (Geometridae) colled at ~pruc'eac~odfian. Jk-ls 6301) ; from the dotted line looper, Protobo+ ramia porcelaria (Gusn. ) (Geornetridae) , collected at Treesbank, Han. (PIS 62988) ; and from Belanolophia imitata (Flalker) (Geometridae) ij collected at Port Clements (PIS 64-50-018-E) and Kyuquot ii /L - (PIS * +J i 3 60-2454-41Y), B.C.

The effect of "host speciesn and on some of the characters used to classity &.larvae of Ichneuaoninae was examined using a one-way analysis of variance and Student-Beusan-Kenl's multiple range test (p<0 .O5) . Table I ** - s shows the results of this analysis. chlr3cters in the finll-lnstlr l3rvae of ~ratichn6umon ptzrtais TOW. roared frm 3 difflrent hosts. nezns f3r znv ch3rlster. falt~we3by ths .same latter arc, not signifcactly di,f f?ren+ (Student-lenmsn-Keull s multlple range tpst, p<3.35). Sample size is in brackets.

Pr~toboaramil Semiothisa sp. p~rcslaria(Ran.) (Clan. )

Intsrpr~cess 0.22+0.014 5 lsngth ( 1 6) -% Hypostoma 1.5+0.16 3 length/intar- ( 16) pncess depth

bhae length 0. 14+0.014 a 0.14+0.014 s 0.1 1+0. 020 b (1 6) (12) (3 1

Closing app- 0.08+0.009 a 0.07+0.015 3 0.07+0.004 a a=tns length (36) (11) (3)' Cl~singapp- 11 5+0.22 a 1.5+0.33 a 1.6+0.30 s - antas le~g+h/ (16) (1 r) ( 31, ' atrinm width - L + The specimens reared from SemXotXisa sp. were significantly smaller than the specimens reared from 5 imitata an2 P. porcelaria with respect to the overall width of the cephalic structure, the length of the hgpostoaa, the depth between the pleurostomal processes, and the length of the mandible blade. In the specimens reared from Semiothisa : - - - sp., the ratios of length of hypostoma to depth between pleurostomal processes, and of mandible length tc lsandible j width were also significantly smaller than in specimens from 3 4 i the other two hosts. The specimens reared from -M. imitata J2

i and -P. porcelaria were not significantly different from each other with respect to the above characters. The hypostoma is disproportionatly shorter in specimens reared from

Semiothisa sp. and the mandible width, in relation to the

------mandible length, is much greater in these specimens.

The length of the aandible differed significantly , between larvae from all 3 hosts; those fron & - imitata were ' h - largest and those from Semiothisa sp. smallest. Specimens reared from & iaitata had wider mandibles than those reared $ from -P. porcelaria and ~emiothisasp., which did not differ i from-ezhother. The similarity between larvae f roq porcelaria and Semiothisa sp. appears to be due to the proportionatly wider mandible of specimens reared from

Semiothisa sp. Although the length- of the blade of the -

t ' mandible was significantly shorter in specimens reared from

Semiothisa sp., the length of the blade relative to lengtK P of the mandible was not significantly different between

- A- L A- A larvae from any of the 3 hosts.

There were no statistically significant differences between larvae from any of the three hosts with respect to b , the width of the atrium, the length of the closing C apparatus, or on the ratio of length of closing apparatus to width of atrium. It is of importance that the spiracular measurements were not significantly smaller in larvae reared

- - pP------P - from Semiothisa sp. This possibly suggests-that the host does not influence parasite body size; however, this seems unlikely as the parasite adults are much smaller than those reared from P. porcelaria and g. imitata.

The number of pores in the epistoma was significantly * greater in larvae reared from &. imitata than in larvae reared from -P. porcelaria and Semiothisa sp., which were not - 4 * - - - -- significantly different from each other. One other obvious i

\ difference is that larvae reared from g. imitata and g.

.porcelaria had 40 or fewer large, single spines on the

hypopharyngealu surface while larvae reared frole Semiothisa

s6. hGd 60 or more small spines in rows across the = hypopharyngeal surface.

d The source of the differences between larsae reared from 1

1

the 3 different hosts cannot be reliably attributed to ebher - i host or location as prasites reared from all 3 hosts at

both locations, from all 3 hosts at 1 location or from 1 .8 host at both locations were not analyzed and were not available.

------The I. I larvae of pteridis reared from imitata in British Columbia and g. porcelaria in hanitoba differ significantly from each other only In the number of pores in

the epistoma and in the overall lengths and widths of the *mandible and do not differ significantly in proportions of

sclerites measured. The adult parasites do not vary to any large degree from the description of the species by lieinrich (1960b) . Therefore, pending analysis of more material, they are treated as members of the same population.

Conversely, specimens reared from Semiothisa sp. differ from specimens reared from the other 2 host species both in size and in prvportim. M particular significance are th-e- differences in the ratios of the hypostoma length to the depth between the pleurostomal processes and the length of the mandible to the width of the mandible; and the differences in the number and form of spines on the hypopharyngeal surface, The lack of significance in the ratio of the length of the blade to the length of the mandible is not particularly i~portantas most species of

- -- - - Cratichneamon examined fall int6 the same gmeral range.

The adult parasites reared from Semiothisa ep, are much smaller (length 7 to 8 mat) than those reared from -M. imitata and P.- porcelaria (length 10 to 11 am). Also, the normal yellow coloration in -C. pteridis adults is repaaced by a - red-orange color; the black of C, pteridis is re~lacedby a ------A adult speciuens agree structurally, except in size, with Heinrich's (1960b) description of C,- pteridis, and will key to that species. The differences between the adults, as noted above, are consistent with the magnitude of differences between closely-related species in the

Ichneunoninae, as are the differences between the larvae. A- - Therefore, pending analysis of further material, the specimens of "C,- pteridisw reared from Semiothisa sp. in Manitoba are considered to represent a new species of

Cratichneumon, or a subspecies of C.- pteridis, and are not considered in the following description of C.- pteridis.

Parasite cocoon of loose whdte si-lk inside host pupa.

Exit hole at head end-of hostLpupa, terminal, 3- to04 am-* diameter,

Cephalic structure of final-lnstar larve (Pig. 38) with , mean width of 0.72 em (n=24, range 0.62 ma to C.86 am, SD= d 0,064, CV=8.38%) . Epistoma complete and lightly sclerotized, epistoaal bridge joined to lateral portions of epistoma by membranous regions, and with an ayerage of 9

------PP -- -- pores (range 7 to 9, median=9, Mean=9) ; ple trostoma ------sclerotized, with lightly sclerotized accessory pleurostomal 7 areas around outer borders; Hypostoma sclerotized, mean length 0.34 ma (n=28, range 0.29 to 0.39, SD=0.023,

CV=6,8O%) and length of hypostoma -1.51 tines depth between pleurostosal processes (n=28, range 1.26 to 1.75, SD=0.0184, ,

CV=8, Xi@. Cly pedahral plates joined lightly to epistoma ky unsclerotized areas not joined rnedially, each with from 4 to 3 sensilla (usually 8) , some of which are located on unsclerotized areas below the sclerctized areas of the plates, Superior plaurostomal process projecting, posterior strut not visible; inferior p leurostomal process with anterior strut reduced and postsrior strut longer than anterior strut, struts not fused ventrally, mean depth b~tueenp~eurostomal processes 0.22 mm (n=28, range 0.20-to-

0.24, SD=0.012, CV=5.53%). Silk press about 1.4 times wider than desp, extending slightly ventrally, opening about as vide as Keep, 1 seta on each lateral margin above ventral margin of opening and some slight areolete sculpture on dorsolateral corners. Maxillary palps each ussally with 1 large, 3 medium and 1 small sensilla, or sometimes with 1 sensillla, one of which is bilobed and 2 smaller sensilla.

Mandibles 0.35 an long on average (n=28, ran@ 0.32 to 0.40,L fl SD=0. 023, CV=6.54%), 0.28 mn vide on average (n=28, range

0.24 to 0.32, SD=O.017, CV=6.06%), and 1.29 mm longer than lids OR average (n=28, range 1.14 to 1.48, ~~=0.099,

CV=7.634). %fadeof mandible 0.14 mm long on average -

(n=28, range=Oell to 0.17, SD=O.O14, CV=9.34%& and 0.33 of total length of mandible (n=28, range 0.30 to 0.47, SD=O.O41, CV=10.49%).

Antenna1 discs never visible; ocular lines rarely visible, or if so then extremely lightly sclerotized.

Hypopharyngeal surface with between 10 and 40 large, single

------spines, accessory reticulate sclerotization very light, soaatiaes not visible, 'and originating at distal end of hypostoma. Hind end of larval skin with many small distinct spines in rows.

Atrium of prothofacic spiracle (Fig. 65) with mean width of 0.52 ma (n=27, range 0.046 to 0.060, SD=O.OOU, 0.1 I, SD=0. 012, CV=l5.52%) . Length of closing apparatus on average 1.48 times greater than width of atriua (n=17, range -_

Genus Patrocloides Heinrich

One species of Patrocloides is described and

illustrated and generic and specific descripticns are fi combined. Species of this genus are not described or

illustrated in the literature.

I Key characteristics are: clypeclabral plaRes not

joined medially, distinctly teardrop-shaped; width of atrium

- -- of prothoracic spiracle less than length of closing

apparatus; hind end of cast skin without spines; a pair of

large vacuoles "is usually present on the epistcaa, and when

present, constitutes a diagnostic character for the genus.

A Patrocloides perluctnosus (Provancher) ' Figs. 39, 65 ------The 5 specimens exaained were rearea from the angulated -z cutworm, Syngrapha rectanqula (Kby. ) (Poctuidae) fi collected at Pass Lake Rd ., B.C. (PIS 63-5950-0 1-i) ; from the spruce climbing cutworn, g. alias (~ttolengui),collected at Sequodie Creek (FIS P53-468-01) and Big Tree Creek (PIS . P53-389-03) B.C., and aoonbeam, Ont. fFIS S60-0803L-0t-t) .

Parasite cocoon not visible within host pupa; exit hole at head end of host terainal, about 3 by 4 mm wide.

.I

Cephalic strnctnre of final-instar larva [Pig 39) 0.81 am wide (range 0.77 to 0.86). pist to ma sclerotized, with 2 to 5 pores and usually 2 large vacuoles; pleurastoma around lateral Bargins. Hypostoma sclerotized, two-thirds to three quarters longer than depth between pfenrostoaal procasses. Clypeolabral plates not joined medially, distinctly teardrop-shaped, each with 2' to 5 sensilla. Superior plenrostoral process pro jecting, posterior str at > not visible; *inferior pleurostomal process ver 9 long and

trough-shaped,------struts - -- trough-shaped. Silk press slishtly -- V projecting ventrally, about one quarter wider than deep, ------b

- - - - - t2-5 -j - f i Y

------b ~4thareolate scsiprnre dorm-laterally, opening aboqks

U uido as deep, 1 seta on each lateral margin above vwral

d margin of openizg. Earillary paps each uith * lar4e and 1 /

-- -- - hntennal discs not visible; ocular lines lightly sclerotizer3, without reticulate sclerotization . Bypopharyngeal sarfece uith a few scattered spines, the -

nedial ones single, lateral ones in rcus of 4 to 6.

Accsssory reticulate sclerotization not visible in most

spec2ens: ne visi32 t qrirros-at h ix CF~caJf Tarwar ------skin. .

btriom of prothoracic spiracle (Pig. 65) 0,063 mi vide

[range 0.058 to 0,070) vith short heavy spiresbido, Closing apparatus O.O83 a. long (range 0.070 to 0.110) . Genus Aoplas Tischbein

Two species of Aoplas axe described and illustrated and a key for their sepration follows, Other species described and illustrated in the literature are: Aoplas ratzehnrgii (lartig) by ~bort(1978) and -A. velox (Cresson) by Short f 1959, 1978:.

not joined aediallf; and u5dtb of atriua of prothoracic spiracle abour the same as length' of closing apparatus. --

L

-a, ratzebnrgif, as illnsrrated by Short (1978) closely approaches the ahre key character, while &, velox, as

-ppp -- -- illustrated by Short (I 378, 1359) does not agree with the width of atrium versos length of closing apparatus character, Bouerer -8. velox [Short 1978) agrees uith the speciltens exdmined fa characters of the mndible, described I below. Key, to species cf Aoplns

1. Baadible 1 .U2 or more longer than wide and blade 0.37 or more of total length of mandible...... velox velox (Cress.)

blade 0.32 or less of total length of mandible...... permutabilis perautabilis Eeinr.

d (1462) a Heinrich considered Aoplns and Cratichnenmon closely related on the basis of adult morphology. They are also closely related on the basis of immature morphology,

and would be virtually inseparable were it not for the

-- - -- ef ference %n spiracle aorpfrcrf ogy, *or the - geometrid-parasitizing Cratichneumon. In addition, both

gerxsra have large spines on the hind end of the larval skin and the characteristical1.y loose1y-assembled epistoma which

seems to be associated with parasitism of geometrids.

h Bei~ichdivides Aoplns into the confirmatus group and

--the vel ox group on thebasis of adult mo~pholoqv; &. e.

-- lox to the latter, The observed differences in nandi3le morphology ray provide justification for this grouping when further material becomes available.

\

0 P Aoplus permuta bilis permutabilis ~einrich

The 3 specimens exaained were reared grom Filament aria * [=llenatocampa ] limbata (Haworth) (Geometridae) ccllected at Trout Lake, B.C.

Parasite cocoon of very sparse strands of silk inside host pupa: exit hole at head end of hcst pupa, terminal,

Cephalic strncture of final-instar larva fFig. 40) about 0.63 mrn wide (range 0.62 to 0.63). Epistoma lightly sclerotized, with 7 to 11 pores, epistomal bridge joined to lateral portions of epistoma by- sclerotized area.

- Pleurostoaa lightly sclerotized with even lighter accessory pleurostomal sclerotization around lateral margins; - - -

-

- -- - hypostoma lightly sclerotized . Clypeolabral plates lightly sclerotized, e longa te-oval with slight dorsal arch; each with 3 to 5 sensil$a. Superior pleurostomal process with both struts visible, anterior sWut larger than posterior

strut; inferior plenrostomal process trough-shaped, abont . one-third longer t ban wide. Silk press about as wide as deep with a slight ventral projection, opening abont

parallel with ventral margin of opening. M,axillary palps

each with 5 sensilla, all of equal size or"with 1 larger and - - 4 smaller as in 8. 5. velox; labial palps each uith 5

sensilla, 4 larger and 1 smaller, or 1 large, 3 medium and 1 small. Mandible about one-third longer than wide, blade slightly hooked in ventral aspect in some specimens, about 0.30 of total length of mandible.

Antenna1 discs and ocular lines not visible, hypopharyngeal surface uith a few single spines. Bccessory reticulate sclerotization not visible, large single spines

on hind end of cast larval skin. a

Atrium of prothoracic spiracle (Fig. 66) 0.041 mrn wide

width and closing ap~aratnslength about equal. 2 P Aoplus velox velox (Cresson) Figs. 41, 67

The 3 specimens examined were reared from the hemlock

looper, Lambdina f iscellaria (Guen. ) (~eometridae) ,

Parasite cocoon of single strands of silk dcrsally and - a solid sheet of silk laterally and ventrally within host papa. Exit hole at head end of host pupa, terminal, about 3 - an in diameter.

cephalic structure of final-instar larva (Pig. 41) 0.66

ma wide(range 0.64 to0.77). ~pistomascl~otl%d,with8 -

to 10 pores and with lightly-sclerotized area separating epistoaal bridge and lateral areas of epistoma. Pleurostolsa sclerotized, with accessory plenrostomal sclerotization - surrounding lateral narghs; hypostoma sclerotized. t Clypeolabral plates elongate-oval, each with 2 to 5 sensilla. Supe~iorplenrostoaal process with posterior *~W--kmgee t h-~tz rior st*MWz+mee&em&L--- 1.4 times wider than deep and opening about 1.3 times deeper than wide;- 1 seta on each lateral margin of silk press and- silk poess opening. Haxillary palps and labial palps each with 1 large and 4 smaller sensilla. Handible about 1.5 times longer than-de, blade Long and tapering, about 0.43

Bntennal discs and ocular lines not apparent; hypopharyngeal surface with mostly single sp$nes, a few lateral cnes in rows. Some faint accessory reticulate scleroti-zation visibls near distal end of hypostoma; large, single spines present on hind end of cast larval skins.

- - --- Atrium of prothoracic spiracle (Pig. 67) 0.023 a wide (range 0.020 to 0.027) with fine hairs and some heavy spines inside. Closing apparatus 0.021 nm long (range 0.018 to

0.025). iiidth of atrium approximately equal to length of e closing apparatus.

Gemg Eutanyacra Cameron

and generic and specific descriptions are combined below. Mhar species figured in the literature are; g. sataralis (Say) by Schaaf (1 972) and' Short (1978) ; and -E. glaucatorius (Fabricins) and -E. picta (Schrank) by Short (1 978) .

Key characters are: clypeolabral plates broadly -. triangular, as deep as wide, and joined medially; lower

margins of silk press not produced ventrally; and length of .

ocular lines less than 2 times depth between pleurostomal

processes. B - -

~6eshapes of the clype.olabra1 plates of E.- suturalis

as illustrated by Schaaf ( 1972) and of E_. qlaucatorius and -E. picta as illustrated by Short (1978) agree aith the present description though the medial join is not shoun.

------A- The base of the blade of E.- suturalis, illustrated by Short (1978) is shown as very narrow, which is not corroborated by

the present description. The shape of the silk Fress is similar in all three species to that illustrated but the inferior pleurosto~alprocess is not clearly illustrated.

Even so, Eutanyacra seems to be a clearly-definable genus. Eutanyacra suturalis -(Say)

Figs. 42, 68 -_ The 3 specimens examined were reared from Euoxa

flavicollis (Smith) (Noctuidae) collected at S askatoon,

- Sask. ; from a species-of -Noctuidae collected at Hission

Flats, B.C. ; ard f ram the clover cutuorm, ~cotoqramma trifoLii (Toss, ) (Noctuidae) , collected at Herscbel, Sask.

Parasite cocoon of sparse, white silk on venter inside host pupa; exit hole at head end of hcst pupa, terminal,

about 4 mrn in diameter.

------Cephalic structure of final-instar larva 4Fig. 42) 1.10 mm wide (range 1.04 to 1.15). Epistoma complete and \ sclerotized, with 6 to 9 pores and reticulate sclerctization at d~rsolaterafiaar~insof apistomal bridgo, lateral portions of epistoma between pleurostcmal processes and bridge very narrou, Pleurastoma sclerotized, with scl3rotiied accessory pleurostomal area around lateral margins; TypOpostoPa usually shoTPandTrved; lenq'th0.73to - ppppp

large and broadly triangular, about as wide as deep and

joinsd nediallp, each with 5 or 6 sensilla. Superior

pleurostomal process vith posterior strut not visible,

antsrior strut pro jecting ; inferior, pleurostomal process

about one-third deeper than wide, struts fused ventrally, adan-riur &%EX& ahat one-fi-EtB as wide-as po&er&er--- -*ut. Silk press not produced ventrally, about' 1.7 times wid ar than deep, depth of opening about three- quarters of width, 1 seta on each lateral margin, below ventral margin of opening. Haxillary and labial palps each with 1 larger,

3 medium and' 1 small sensillurn or labial palps each with 4

sensilla of equal size. Bandible 1.2 to 1.3 times longer than wide, blade 0,38 to.O.42 of total length of mandible.

Antenna1 discs visible, with sclerotized rim; ocular

lines visible, length 1.2 tc 1.6 times the depth of pleurostcmal processes, with reticulate sclero tization on

s,urface. Hypopharyngeal snrf ace with spines mostly in rows \ of 4 to 6 and a feu single aedial spines reticnlate sclerotization extending from distal end of Atrium of prothoracic spiracle (Fig. 68) 0.062 mm wide (ranga 0.052 *to 0.072) with fine hairs and a Ker short spines inside. Clcsing apparatus 0,097 mm long kange 0.080

Genus Diphyus. Kriechbauner --v gs

One species of Diphy us is described and illustrated. The specimens examined were in the 'Canadian Natiwal collection as Diphyus Ott. spec. 1. By examination of the specimens and comparison with descri~tionsof species of

Riphyus they were determined to be Diphyus variegatus euoxae

- - -- Heinrich (1 969) . -

Species of Diphyus figured in the literatare are:

~iphyussp. by Schaaf (1972); i). indocilis (Flesmael) by

Short ( 1370, 1978) ; D. nuncius (Cresson) by Short (1959, D. D. 1978) ; and - animosus rubellus (Cresson) , - palliatorius (Grasenhorst), -D. gnadtipunctorius (Hneller) and Generic characters are: clppeolabral plates large and broadly triangular, although not joined nedially, near1y touching pedially; lower ~arginof silk press not extending *- ventrally; length of ocular lines greater than 2 times depth bet ween pleurostornal processes; and epistoma i nconplete .-' and/br unscleroti zed. In dddition, the base Q f the-.~an&ible!__r narrows rapidly to the blade, so that the blade is well defined. A11 spedies of piphyus illustrated bq Short (1959) aqd Schaaf (1372) agree to sene extant with the above description.

Diphy us variegat us euoxae Heinrich

Figs. 43, 69

Ths 4 spscimens examined were reared from the red-backed cutworn, Euoxa ochrogaster (Guen. ) (Noctuidae) , collected at Beaver Lodge, Alta.; and from a species of Poctuldae. collected at Saskatoon, Sask., and Lilloet, and

Bission Flats, B.C.

#

-Parasite------cocoon --- of fine silk along venter, inside host .

pupa; exit- hole-- - at head end of host pupa, terminal, about 4 - mm in' diameter. P / / Cephaic structure of final-instar larva (Pig. 43) 0.74

am wide (range 0.67 to 0.81). Epistora incomplete or lightly sclerotized and with some sclerotization over bridga, no visible pores. Plenrostoma sclerotized, with light narrow accesscry pleu2ostonal sclerotization around ------latsral margins. Hypostoma broad and truncate, about '. one-third longer than depth b etreen haorcstcmal processes.

- Clypeolabral plates about as wide as deep, filling 1: .+ clypeolabral region, alsost touching aedially, each with 5

or 6 sensilla. Superior plenrostonal process rith posterior -.. strut not visible, anterior strut protruding; inferior pleurostomal process about4 as wide at base as long, anterior strut shartar -thar-posterior strut, Silk press bbtt* as - - - wide as deep, not produced ventralli, opening about as wide

as deep and 1 seta located below ventral margin of opening and msdial to lateral margins of S&Yk press. aaxiliary and

labial palps each with 4 large and 1, smaller sensillnm. Mmdibles 1.2 to 1.3 times longer than vide and blade narrow

at base, length of klade about 0.35 of total length of ------antenna1 discs not visible; ocular lines with reticulate sculpture on surface, length ;ore than tuice

depth between p leorostomal processes. Hypopharyngeal?& - sur fece with singla spines; accessory reticulate sclerotization very light and extending around lateral

~arginsor cephalic st ru-nre t~e~istoia,- Slallspines.... present or; hia6 end of larval cast skin,

- - - Atria. of prothoracic spiracle (Pigs 63) 0.064 mn vide

(raags 0.056 to 0.070) with fine hairs and a feu short,

heavy* spines inside. Closing apparatus 0.03~BR long (range

0.072 to 0.110).

One sp%ies of Litercdops is describe6 and ifl~strated

and s secmd, Limerodops subsericans (Gr%venhorst), uas

illustrated by Short (1978).

Key characters are: clypeolabral plates large and joined sediagfy; loser aarqfn of silk press prodacea

ventrally and I seta on each lateral margin below ventral - -- -- =gin of opening; and langth of ocular lines more than

twice the depth between pleurssto~alprocesses;, and width of

.ocular . lines eqaal to depth between pleorostmal processes.

- The clypeolabral plates of &. sobsericans (Short 1978) are siailar to these of L. belangeri and--_the mandible~impe------is also si8ilar. Mher characters in Short's illnstration are either not shovn or not clear,

Liisradups belangeri (Cresson)

Pigs, 44, 7 .

ParaSite COCOOG ~f SPZKSB~ loose, whits silk around

anterior one-third of inside of host pupa; exit bole at head M erd of host pupa, tarminal, about 3 mm in dtametet,

Cephalic stractare of f inal-instar larrae (Fig. 44)

snsclerotized accesscry pleurosrosal *area, and sensilka -at fa6 tires the ilepth bet ween processes, Clypeolabral

1 plates rectangalar, joined medially and not attached to epistosa, each uitk 5 or -6 sensflla. Superior pleurostomal process vith postesiar strut not visible, anterior strut

~odisti E&, not protruding ; inferior p~urostomalpzocess uith anteribr strat shorter and narrower than ~osterier

strat, and struts apparently free. Silk press 1.3 times

wider than deep,apeningabout timas deepexthan uide, a --* ------gair of setae located redfalfy on -silk presE below ventral

margin of opening, Maxillary palps nsnally each uith 6 sezsilla of about equal size; labial palps each uith from 4

ta 6 sensilla (if 4, then one larger than others, if 5, then

all of sqnaZ size, if 6 then one of the sensflla very ssall) . Mandible 1,3~to T .5 times longer than wide, yde 0.42 to 0. St of total fengtb of msnilible,

lateanal discs not risible, length of ocular lines

about trice the depth betseen plaarostonal gircicosses and C

s~lrfacewkth single spfses; accessory reticulate diszal end of hypostoma around lateral aargins of A plastostoma to epistcm~ A very few spines and ~uch. areolate sculptuse at hind end of larval skin.

Atrium of prothoracic spiracle (Fig. 70) 0.059 ma wide ~~z), with~zZ'iti&insidQ. - apparatus 0. @76 mm long (range 0.070 to 0.084).. C

Genus Spilichneumon Thomson

One species of Spilichneumon is described and. illustrated here. Other species described and/or illustrated in the literature are; -S, brontens (Cresson) by Short (1959, 1978) ; -S. superbus (Provancher) by Schaaf

(1972); and Spilichnenmon spa by Short (1978)

Generic characters are: clypeolabral plates joined r&faZly, about as deep as uide; lower margin of silk press nut produced ~entrallf;anterior strut of - inferior t plearostoadl process deeper and wider than posterior strut; The species of Spilichneumon figured by Schaaf (1972)

aad Short (1952, 1978) agree with this description, although details of the inferior pleurostoaal process are not clearly illustrated.

S pilichneumon subruf us (Cresson)

Pigs. 45, 71

The 2 specimens examined were reared from an unknown species of Lepidoptera, probably of loctuidae, collected at

-Hemmirigford, Que.

Parasite cocoon iot visible inside host pupa; exit hole

------

at head add host pupa, terminal, about 4 mm by 6 ma vide. ,

Cephalic structtue of final-instar larva tFig. 45) 1.0

mm wide in one specimen and 0.93 ma in the other. Epistoma sclerotized, with 12 pores; epistomal bridge connected to lateral sections of epistoma by sclerotized area. Pleurostoma sclerotized with nnsclerotized accessory pleurostonal area around lateral margins; hypostoma sclerotited. Clypeolabral plates large, about as deep as wide, joined medially, each wits 7 sensilla. Superior pleurostomal process protr6ding, posterior strut not

visible; inferior pleurostomal process with anterios strut

uider and deeper than posterior strut, concealing posterior

strut. Silk press about 1.9 times uider than deep, uith areolate sculpture on dorsolateral corners; opening about 1.4 times wider than deep, a pair of setae. located mediclly

oo silk press beleu verhraL arrnqi~of opening. Hhxikhtry------palps each with 5 large and 1 smaller sensilla, labial palps

each with 4 sensilla of equal size. Handibles about 1.3

ti~eslonger than wide, blade about 0.37 of length of & mandible.

2.5 times longer than depth between pleurcstcrnal processes.

uith reticulate sclerotization on surface. Hypopharyngeal surface without visible spbes; hind end of cast skin without visible spines.

Atrium of prothoracic spiracle (Fig. 71) 0.061 mm wide

--- (range 0.060 to 0.064). with a few short spines and no fink/ hairs inside. apparatus am Closing -- 0.072 long (range 0 .O6 Genus Thyra teles Perkins

Two species of Thyrateles are described and illustrated

here. Species illustrated in the literature are: 2,

camel inus (Uesmael) , -T. lugobrator (Gravenhorst) and T.

This genus presents some difficulties in

- -- - classification. according GHainrica 1361) and Perkins (Qg6O) , the adults are virtually indistinguishable from species of Ichnea.cn. The larvae are readily distinguished by the indistinct aedial join between the clypeclabral

plates in Thyrateles, which is much more distinct in

Ich ne 80 -specAs the ------u n,- -In sera-specirewz&&t&t joip-k not apparent at all and these key to near Patrocloides and Cratichnaumon.- They are distinguished frem Cratichneumon by the clypeolabral plates not ha ring a semi-apparent join to the epistoma, and from Patrocloides by the presence of antenna1 discs, Key to species of Thyratelas.

1. Silk press with areolate sculpture over entire surf ace...... , ...... ,...... procax (Cress. ) silk prsss with areolate sculpture restricted to sages, - -or not- A sculptured...... ,. ,lnqubrator [Gray* 1

Th yrateles lugnbratar (Gravenhorstj

Pigs. 46, 72

The 4 specimens examined were reared f ram the painted lady, Vanessa cardni (L.) (Hymphalidae) ,'collected at

1 Sterling, Alta. and Port fiaaaond, BrC.; from the American

pain- IaLy, C ynt hia-TFizqLffisasi-s fDmzy- ~y@&li&a+), collected at -Black- Sturgeon, Littlefield Stn. Camp '32 [ lprorirrca 3; and frq8- an oniaentif ied species of ilyaphalidae

< .A collected at st. rloois Ravignon, Que.

Parasite cocoon not ,visible inside host pupa; exit hole

* at hsad end of host papa, terminal, about 5 ma in diameter.

- Cephalic stnrcture of final-instar larva (Fig. 46) 0.91

nn wide (range 0.83 to 0.93). Epistoina sclerotized, usually . vith 9 to 1t pores (18 exceptionally), epistomal bridge joinad to lateral sections of epistoma by sclerotized area. Pleurostoma sclerotized, with sclerotized accessory pleurostonal area around lateral margins; hypstoma ------sclsrotized and narrou. Clypeolabral plates appear to have the vestiges of a medial join in some specimens and not in

------others, each with 5 to 10 sensilla. superior pleurostomal

- . process with broad base, posterior strut not visible: - infarior pleurosto~alprocess with ,struts trough-shaped .

- Silk press without areolate sculpture or with restricted

C sculpture along edges, about 1.2 times longer than wide,

- ventral 1-p prckradisg, ~psnin+abs+~Be+pas *i-,-me seta on each lateral margin about parallel with ventral

margin of opening, Baxillary palps each vith 1 large and 4

smaller sensilla, sometimes t large, 3 medium, and t small

sensilla; labial palps each vith 2 large, 3 aedium and 1

small sensilla. Handible usually 1.2 to 1.3 times lager than wide (I specimen 1.5 tines longer than wide), blade 0.24 to 0.28 of the total length of mandible. ------Antenna1 discs with heavily sclerotized rim, greatest diameter about 1.3 ti~esthe depth between pleurcstomal processes; ocular lines about 1.8 times longer than depth bet ween pleurostomal processes, without reticulate sclarotization on surface. Hypopharyngeal surface with

sclerotization originating from distal end of hypostoma.

Hind end of larval skin without visible spines.

Atrium of prothoracic spiracle (Pig. 72) 0,055 mm wide (rangs 0.050 to 0.058) , vithout visible spines or hairs inside. Closing apparatus O.08Y arm long (range 0.062 to 0.036).

The specimen of T, lngubrator reared from -C. virginiensis coflected at Black Sturgeon is exceptional in tuo respects: the epistona has 18 pores as opposed to 9 to 11 in the others; and the mandible is 1.5 times longer than uide, as opposed to 1,2 to 1.3 in the others. These moralies may be due to host effects, i.e. the other

description of -T. lngubrator. Th yrateles procax (Cresson) C Figs. 47, 73

The 4 specimens exanined were reared from the mourning

cloak butterfly, l y~phalisantiopa (L. ) (Nymphalidae) ,

collected at Sault stah Barie, Qnt. (PIS-YSJ(3--159-9j ,---

Parasite cocoon not visible inside host pupa; exit hole

------at head end of host pupa, terminal, about 5 am by 3 ma in diameter.

cephalic structure of final-instat larva (Pig. 47) 0.97 nm wide (range 0.95 to 0.98) . Epistoma complete and

~kretfbed,uM- 5 to 7 p~~~s;-~Itnr0~~6rnaBscZferotize~~ pp --- uith ride, lightly-sclerotized acc3ssory pleurcstomal area:

hypostom scleroti zed. Clypeolabral plates indistinctly joined medially in seme specisens but not in others, each

uith 5 sensilla, 2 or 3 of those located just belou the plate on an associated membranous area. Superior

plearostomal process uith base somewhat narrower than in -T. pgubrator, posterior s+ret not visible; irrfer - - - inr pfearootomal process with struts trough-shapnd, posterior - strut much deeper than anterior strut. , Silk press uith areolate sculpture over entire surface, about as deep as

rids and protruding ventrally, opening about as deep as

uide, one seta on each lateral margin, about parallel with

ventral margin of opening. Haxillary palps each with 5 sans5lla of abo- _equ-al size: labial- palps each -~~i.t4~-4-la&

and one very small sensilla or with one large and 5 smaller sensilla. ~andiblo1.3 to 1.6 times longer than vide, blade

-- - - 0.30 to 0.40 of total mandibla length.

2 II antenna1 disc with heavily-sclerotized rim, greatest diaaeter about equal to depth between pleurostomal A processes: 'ocular lines about 1.4 times longer than depth

sclerotization on surface. EIypophaiingeal surface with scattered, sing1e spines; accessory reticulate sclerotization extending from distal end of hy postoma a round .4 lateral mprgins cf cephalic structure to epistoma. Bind end of larval skin wikh small spinas in rows.

atrium ofprothoracic-- spiracC1eBtPi-g&) 4Imn7;,ar wih -- enge 0.066 to 0.080) , with short heavy spines and hairs - - -- inside. Closing apparatus 0.118 mm long (range 0.100 to Six species of Ichneuaon are described and illustrated

here. Species of Ichneumon figured in the literature are:' . anbulatori'us Fabricius by Short (1959, 19743) ; 1.

ext ernso rins-y-ISho~t ( 193-0, t 928)-; -I + - supiciosus- Uesnael by Cameron (1350, 1351-) ; and -I. caliginosus Cresson -I. caloscelis Besmael, I. conf usor Gravenhorst, L.

------deliratorius Linnaeos, I. dioryctriae ~einrich,1. laetus 0rulle; E. melanotus Eolmgren, z. sarcitorius Linnasus, and -I. stramentarius Gravenhotst by Short (1 978) .

~enericcharacters are:. clypeolabral plates usually

- large and slways fhne-&al-*r-ber-aargb-of-si-fk-- press- s produced ventrally, setae on silk press located beneath . ventral margin of silk press opening; and ocular lines, when present, with length -less than twice depth bet ween pleurostomal processes and width less than depth bet ween pleurostonal processes,

I

- - Some-- --- of- thed2Anstrations ., by Short (1978) ng-, T. . L confusor, I. dioryctriae, I. melanotis, I. sarcitorius,and -I. stra~entarias,agree particularly well with the above 0" *r( l-4 0 4 +, +, Id 4 4 JIW C1 cU rs cd d . .rl . U n . i-4 . cd $I 0 . 3 ul -d d a. o Q) .--I +, Wcd 0 N 4 a+' U 0 a, I4 d u W rl 0 U 0 cn - -- -- 5. length-of hypostoma more than 1.5 tiaes depth

bet ween plenrostomal processes; antenna 1 discs not visible...... maius Cress. /' Length-bf h~pcsromless than 1.5 tines depth between pleurostomal processes; antenoa 1 discs usually visible...... trizonatus Prov. up - --- -A a

fr\ Heinrich (1361a) dividOd the speciei of %hneu~on into

rmbulatorius and &. canadensis are in the laetus group, 1.

dioryctriae and -I. caliginosus are in the feralis group and

-I. - maius and -I. trizogatos ara in the gracilicornis group. Z Bepreseatatives of the inurbanus and arteais groups were not examined. - - pp ------

Larvae of the 3 groups in- which representatives were sxsmined can be separated as follows: thp laetus group do not have spines on either the hypopharyngeal surface or the hind

end of the larval skin; the feralis group have spines on the hjpopharyngaal surface but not on the hind end of the larval sfin: and the gracilicornis group has spines on both the hypopharyngeal surface and the hind end of the larval skin. Irr b-douk=tful that this classification ot thelarob ~tlr-~ t f, show caaplete congruence uitk that of the adults as Heinrich

f 196 la) noted the presence of :iany linking and intermediate forms between these two gtoaps,

Beinrich ' f f 96 la) establishsd a borderline betueen the

specias are -1. cafiginosns and T.- caliginops Beinr., which

are distfngnished by the forrer- - being oxyppgoog- and the- - - -pp -- - latter, seniamblypygous. Hithin Thyrateles there are tuo groups with ?respect to tk.e nature of the hypoppgium of the female: one is se~iamblypygoos (including 2.- cafiqinops and

7 -9, lugubrator) and the other is a~blypygousf incPading__T. procar). he forser gronp is vex). similar tc the feralis arid T. lagabrator also has-spines on the bat not on the hind end of the larval. skin as in the ferafis ,group, which suggests a close relationship betueen the tuo groups. T. procax has spines an bath the hypopharyngeaX s~rface'andthe hind end of the larval skin, bat the adult is not similar to any neshrs of the gracificornis gronp so no relati ip f s suspected, r

/ -- 1% s; ', -1 > t . 4 +r 14 a d4J, 4 d1 i.l r * i a , 4 4-01 k ~arlr( VUd pa^ 2 0 k X * 4 W, 4.CJ 0 u a +r e, MU+, mead ra 9,' 1 w(Paw d aim a40 Old Q).- 0 a 9 8 Y E4? u k E) w

a .g @ tQ 4 a ,#+* "rd fl.Qb 2 g1 . IJ -$ trs3 PI . a M a a dl w $8 ~14.1q+1-4 +, Me I 9Q %, g. 4 opening about as deep as uide, one seta on each lateral

C margin below ventral margin of opening. ~andiblesaboat 1.3 - b the= wider than deep, with blade 0.38 to 0:40 of total

length of mandible. . ..

/~ntennal discs with sclerotized ria, their greatest ------* - diaeater ,about- equal to depth between plenrostcmal -, r . %, -< . -. .- , - - processes; kular lines' not visible. Hypopharyngeal s:arf ace

s-rotization originating from distal end of hypostom and - I erten@bg. around 1ate;al masgins of cephalic structure to - ,- -, . epistcha. Hind. end of larval skin without risible spines.

. - &trio; uf prothoracic spiracle (Pig. 74) aboat 0.054 mm pp-ppA ------

a -..-' "- + , &thoat risible spines or hairs inside. Closing apparatus

j=aboot 0.069 -am long. -

n3 The 4 specbens erarined were reared from iyaphafis spa 4 (wtphal3daet collected at SBashuap Lake, 0-C. (?IS BC E 4 19-1465); from- a species of Irmphalidae collected at Kaslo, - Bate ;and Zroa Polygon a satyrus (Edw.) (Byrphalidaej

collected at Ottaua, Ont.

Parasite cocoon of sparse, loose silk around inside of hast papa and extending back to meconina at rear of host papa, Exit hole at head end, terminal, about 5 to 7 mm in diameter, andTth a slimly scorzed'-appearaace around edges.

Cephalic st4? ctura of final-instar larva {Fig. 49) 1.06 ma ride (range le$0 t~ 1.16) . Epistoaa complete and sclerotized, with 10 to 12 pores usually, 20 pores exceptionally; plenrostoaa sclerotized, with scierotized accessory plenrcstamaf area, Hppostoma sclerotized, length to depth t.2 1.5 khes between plearosto~lprocesses. I Clypeolabral plates jof sed medially, each uii3er than deep and with 8 or 9 sensilla. superior plenrostomal process protruding, posterior strut not visible; inferior pXearostosal process with struts free, ttpterior strut triang~lar, psterfor strot trwcate, Silk press produced e rentrally, sligbtly rfdsr than deep, opening marginally rider than dwp, une seta on each lateral nargfn af silk - press about parallel with ventral margin of silk press . \ open id^. Maxillary palps each with 5 or 6 sensilla. (if 5, \ \ all about equal size; if 6, then one very small) ; labial palps

aach with 4 large and 1 sadsensillun (if 6, then 2 very \ small). fian&bles 1.2 to 1.5 times longer than wide, with

Antenna1 discs rgth heavily sclerotized rim, greatest

mete+& 4+ t. 3 tfws processes; ocular lines scXerotized with reticulate

sclerotization on surface, 1.3 to 1.8 times longer than"

depth between processes. Accessqry reticulate

sclezotization originating from disSal end of hypostoaa ; hypopharyngeal surf ace uith media 1 s~inessingle or double, ------lateral ones in rows, Hind end of larval skin without visible spines,

Atrium of prothoracic spiracle (Pig, 75) a.074 mm vide

, (range 0,066 to 0.080) , uith fine hairs and a feu short spines fnsf de. Closing apparatus 0,038 mm long (range 0.07 to 0.13oj. - fchaeumon -canadensis Cresson - - ec ~iGs.50, 76 -i

The 4 specimens examined were reared from the, bertha

armyvor a, Hamestra conf ignrata Wlk. (loctuidaeif , collected at Glenlea, Man,

Parasite cocoon of dense, coarse silk inside host pupa;

- &-ke- h6f)CC*e-t-a 1 am------#

Cephalic structure of final-instar larva pig, 50) 1.03

mr, wida (range 0.98a to 1 .O5) . Epf stosa complete and scfarotized, with fro. 6 to 10 pores, Pleurostoma, accassor y plenrust omal area, and hypostom sclorotized.

Clypeolabral plates joined medially, each wider- - than- deep, with 5 or 6 sensill%. Superior pleurosto&al process protruding, posterior straz not risibh; inferior ph~rctstomalprocess with struts free, anterior strut much

narrower at free ead than pcsterior strut, Silk press produced rentrall~. slf &tly ddar than deep. opening abmt 3s vide as &eep, a sea on each lateral rstgb about parallel with ventral margin of opening. ~axillarypalps .

each uith 5 large and f small sensilla, labial palps each

uith 5 sensilla. Handibles 1.2 to 1. U times lcnget than wide, blade indis+,inct, 0.3 to 0.4 of total length of mandibles.

* Antenna1 discs with scleroti zed rims, greatest diaaeter

0 0,9 to 1. I times depth between pleurostoaal prccesses;

ocular lines sclerotized, t,3-to-1,7 *~ll6nger thandepth-

between pleurostomal processes. H y popharyn gea 1 surface

uithoat visible spines; accessory reticulate sclerotiza tion originating from distal end of hypostoma. Hind end of larval skin without visible spines.

(range 0.054 to '0.060f. uitheut risible spines or hairs

-inside. Closing apparatus 0.082 BE long (range 0.070 to Pigs. 51, 77 o

The 4 spechens exasined were reared from ~ior~ctria

reniculella ' (Grate) (Pyralidae) collected at Gainer Lake, -&a,.and fro. amknown sqecies of Tortricidae GO-llected'at

Uchilk (PIS 547-3372Q and Iloberts (PIS 3Uf-2848A). Ont.

------Parasite cocoon of sparse yellow silk inside host pnpa; exit hole at head end of host' pnpa, terminal, about 2 na in diameter.

Cephalic strnctare of final-instar larva Wig. 51) 0.5%

Pt ~*+he&St&.wr-~~&-wIwW

and nasrow, vith 6 to 9 pores; pleurostoma sclerotized with fightly-scf erotf zed accefso~ppXenrostora1 aress B ypstuwa * sclerotized, length more than twice depth betveen processes. Clypedabral plates each wider than deep, joined aedially, each with 5 to 7 sensilla, Saperior pleorostoLa1 process protruding, posteriar strut not visible; inferior

--- plearosto~fprocess with struts apparent1 y fused tentrally, trough-shaped, anterior strut not triangular, posterior - strnt deeper than anterior strat. Silk press about

three-quarters as vide as deep, protruding rentrally,

opening abont as deep as vide, 1 seta on each lateral

margin, abont parallel with ventral margin cf opening.. '

Maxillary palps each with 1 larger, 2 medium and 3 smaller

sensifla; labial palps each with 1 large and 4 small

- - -A------' / sensilla, Handlbles 1.3 to 1.5 times longer than wide, base L, x' of blade very broad uith blade distinct y wedge-shaped, 0.31

,*-8.37 tf*s t-2 *Gh-&d&f&s-- tb z --

Antenna1 discs, ocular lines, and accessory reticulate sclerotization not visible; hypopharyngehl surface with large, single spines, hind end of larval skin without visibls spines.

Atrium of prothoracic spiracle (Pig, 77) 0,038 mm wide * (range 0,044 to O.OSUf, with fine hair, no spines,

i.ns3.de. Closing appztratas 0,056 mr long

0,060) - -pp I chnenmdn mains (Cresson)

C The Q spechens examined were reared from Crambidia

casta Sanborn (Arctiidae)- collected at Hestbank, B.C.

Parasite cucoon not visible within host pupa; exit!' hole

at haad end of host papa, terminal, about 2 mm in diameter.

Cephalic mtrrtct* of final-instar larva (Pig. 52) 0.-6.3-d

RS vide (range 0.62 to 0.71). Epistosa complete and lightly filerotized, w itd to 10 pores: pleurostoaa sclerotized, uith light lysclerotized accessory pleorosto~alares.

Bygost-oma scles~tized,length p--~-pp 1.5 to 1.9 times depth between -- I plonrostcmal processes. Clypek abral plates joined ~edially.each wider than deep with 5 or 6 seneilla, one of vhich is bilobed. Superior plenrosto~lprocess

pro j+cting, posterior strut not visible; inferio~ ple~rostomalprocess uith anterior strut concealing psterior strut, apparently fused and trough-shaped. Silk press produced w~trally,about as vide as deep, opening

~rginallywider than deep, 1 seta en each lateral murgin + ------about parallel with ventral margin of opening. Maxillary

$alps each with 5 sensilla of equal size or, if 6, one very

small; labial palps each with 4 large and .one small sensilla. Mandible 1.4 to 1.5 tines longer. than wide, blade 0.37 to 0.41 af total Length of mandible.

,-7

- - -- -A- --- A - ---LL -- Antenna1 discs not visible, ocular lines usually not visible, or if visibla very ,lightly scleroti zed, length

------+beat f .8 times +h bet- ~ustos&lgroeesses. - Bypapharyngeal surface with medial s~inessingle or double, 3 kateral spines mostly in rous of 10 or more; light accessory

Gulatexlerotization so~etimesvisible, originating from distal end of hypostoma. Hind end of larval skin with

large spines in rous.

Atrium of prothomcic wracle (Pig. 78) 0.044 mm wide

frasge 0,040 to 0,048). without visible spines or hairs * inside, Closing apparatus 0.064 ma long frange 0.052 to \ Ichneumon trizonatus Provancher figs. 53, 79

The 3 specimsns examined were reared from an unknown species of Aoctuidae collected at at qnknuwn location in

Canad*?

------A- -

Parasite, cocoon of loose, white silk around anterior

host pupa, terminal, about 4 mr in diameter.

Caphalic structure of final-instar larva wig. 53) 0.97 43 ma wide (range 0.8 2 $0 3.13) . Epistoaa sclero tized and

complete,------with --~~ ~ 7 to- 3 pores; pleurostoma and-- accessory

pie urostomal area scleroti zed. Hypostoma scletotized,

length 1.3 to 1.4 times depth betwsen pleurostomal

processes. Cly peolahgal plates joined mediallq, each wider than deep and each with frcm 6 to 8 sensilla. Superior plearostomal process projecting, base quite narrou,

pasterior strn t not vksible; iaferior pleurostomal process "

uith struts apparently free, anterior strat deeper than -

- - - - pearior strut, Sf lk press slfghtly projectkng ventrally, about 1.2 times wider than deep, opening about 1.6 times * wider than deep, 1 seta on each lateral margin qbout parallel vith wentral margin of opening, Harfllary palps each with 5 or 6 sensilla, when 6 sensilla present, 5 of =% equal size and 1 very small; labial palps each with 5 or 6

rery small, if 6, & of egbal size and 2 rery small. Uandible abont 1.3 times lodger than vide, blade 0.3 to 0.4

------oftotal IreagtZ 6f maii&iBls. -

Antenna1 discs with part of margin scletoti zed, great ast diaaeter about equal to depth between plenrostomal

processes; ocnlar lines sclerotized, length 1,-0 to 1,8 times

--depth b et Y eea _plenrosto=~lprocesses ,uittLret_kc~late--~ sculpture on sar face, Hypopharyngeal surf ace with medial , spinss single ar b~nbltaand lateral spines in raws -4 to * 6; accessory reticulate sclerotization originating from hist31 end of hypastowa. Bind end of larval s Yh smll, single or double spines.

Atrior of pmthoracic spiracle wig. 73) 0.055 mm long (langs 0,052 ta 0,066) . without risible spines .or hairs inside. Closing apparatus 0.085 mm long (range 0.068 tp-' One species of Orgichneumon is described and illustrated, blo larvas of this +genus have been previously described,

T6eLge n3s iXveTyclose t to Ehneu miX-onnt3eIba slsVp- adult morphology and is in many ways similar tc Coaj$chneumon of the Ptotichneumonini, The -generic - -4------characters are: clypaolabral plates joined aedially; lower aargin of press not produced into a lobe; and clypeolabral plates much wider than dgep. The one character in which this species differs greatly ' from Ichneumon is that the lover ~arginof the silk press is not pro3 meed

Or ichne mon calcatorius (Thnnberg) *Pigs. 54, 8 Parasite cocoon not visible inside host pupa;* exit hole C

2 to 3 nm in diameter. + T a- Cephalic structure of final-instar larva (Pig. 54) 0.83 P

ma vide (range 0.76 to 0.33). Epistoma complete and lightly -

------a - a- sclerotized, vith 10 to 12 pores; pleurostoma scleIOtized, with sclerotized accessory pleurostomal area which extends

C Balfuay down hypostoma. - Hyp-ostoma sclerotized, &mut --- , length 1.6 to 2.0 tims depth between pleurostcmal - processes. Clypeolabral plates joined mediallp, each wider than deep, vith 7 to 9 sensilla. Superior pleurostonal

process projectjag, posterior strut present but concealed by - , anterior Strut; inferior plenrostamal process with strut

------p------free, posterior strut deeper than anterior strut, Silk

press not produced ventrally, about 1.4 times uider than ** >

dl - deep, opening about as vide as deep, 1 seta on each lateral

# margin about parallel witi9 entral rargin of opening. iuxillary palps each with 1 large, 9 mediun'and one very smll sensilla; labial palps each with 1 large, 3 medium a r- f md 1 refy s~llsensilla. Baodibles 1.3 to 1.5 -times ' / \ - -- - lager than ride* blade 0.31 to 0. 3sdal lengt b af mandible* Antenna1 discs,' ocular lines. and accessory reticulate

4 scl3rotization not visLble; hypopharyngeal surface with medial spines single or &able. lateral spines fn zows of 4

6. Bind'end 6f cast larval skin without visible spines.

(range 0.052 to 0.060) ith short heavy spines, no fine 3 t, 'T i hairs inside. with discernible sclerot?ization on surface. - * Piqs. 29-31, Cephalic strnctnres of f inal-instar larvae:

23. Ho~lismenusmorulus morulos Say ; 30, 8. r. pacif icas - Say; 3 1, Vulqichneaion brericinctor (Say) .

Pigs. 32-33, Cephallc str~uctnr.esof final-instar larvae:

32, Cratichqeuaon unif asc$atorius va~converiensis(Prov. ) ; f 33, g. snblatus ' (~ress,&. 7

Pigs. 34-35. Cephalic structures of f inal-instar larvae:

34, Cratichneuaon anisotae Heinr.; 35, variegatus (Prowm}

Pigs. 36-37. cephalic structures of final-instar larvae:

36, Cratfchnsuron zitus Heinr. ; 37, Cratichneumon sp. nr. s Vf2SCOS fPfOV.) .

Pigs. 3 8-40 .cphalic structures of final-instar larvae: - b pteridis Tow. ; 39, Patrocloides - po~luctoosus(Prov.J; 40, Aoplus perautabilis perautabiliq Heinr.

f -- Figs. 41-42. -.~k~halic *- stsactares of final-instar larvae:

4 I, aoplns velox relox (Cress.) ; 42, Eutanyacra suturalis *

Pig. 43. Cephalic stractnre larva of

Figs. 44-46. i Csphallc structures of final-instar larvae:

44', Linerodops belangeri (Cress.) ; 45, Spilichneumon subrufus (Cress .) 46, Thyrateles luqubrator (Grav. ) . Ll-

B Figs. 47-49. Cephalic structwes of f inal-instar larvae:

47, Thyrafeles procax - (Cress.) ; 48, Ichneumon- ambulatorins

Fab. ; 49, I. caliginosus Cress.

Pigs, Cephalic structures larvae:

50, Ichneumon canadensis Cress. ; 5 1, I,- dioryctriae Heinr. ;

52, -I, maius Cress.

Figs. 53-54. Cephalic structures of final-instar larvae:

53, Ichneumon trizonatus Prov. ; 54 j O~chneumoocalcatorins .. Y (Thnnb, ) .

Figs. 55-67. Prothoracic spiracles of final-instar larvae:

55, Hoplisnenus aoralns e~rulusSay; 56, H. a.. pacificus

Say; 57, Vugichneumon brevicinctor Say; 58, Cratichneumon . unifasciatorius vanconveriensis (Proa .) ; 53, C. sublatus i (cress.) ; 60, anisotae Hsinr. :k~,C. varieqatus

(Prov.); 62, C. ritns Heinr.; 63, ~ratichnenmonsp. nr. ,

? vescus (Prov. ) ; 64, g. pteridis Tow., 65, Patrocloides

~einr.; 67, Aoplas velox velox (Cgess. 1 .

Figs. 68-80. Prothoracic spiracles larvae:

68, Eutanyacra suturalis (Sap) ; 69, ~iphyusvarieqatus euoxae Heinr. ; 7 0, Limerodops belanqeri (Cress.) ; 7 1,

Spilichneumon snbrnfns (Cress.) 72, Thpratejes luqubrator

(Grav.) ; 73, x. 74, Ichneumon ambulaturius

Pab. : 75, 1. caliginosns Cross. 76, L. ,canadensis Cress.;

77, -I. dioryctriae Heinr.; 78, L. maius Cress.; 73, T.

TRIBE PEIOTICHNEURONIHI

Three spocies of 2 genera srct describ~dand Illustrated. Species of Protichnensonini figured in the

- pepticus Ciesson by Short

Itpress~rZetterstedt by Shcrt (1 970,i / 1373) ; and -b~~lpjo~~a - pzotens Christ, Coelichneuron

nigsrriaus ? Ste?C~ns,C. subquttatus Gravechor,*,

Zratto joppa rcbusta Cameroa, Protichnonno~lsp., Syspasis I ------tgua (HeinrLch) , and 2. fineator (Fabrfcius) by Short

(1378) e

Kay tc GeDera of Protichneunonini

1. Zlosing sppzrl-tus of prothoracic spiracle more

rhan 2.5 tines ES long as vidth of atrium /--1. / u...... ','\ Coelichneumon ,/' Slosaq*- a2paratu.s of pzotnoracic spiracle much

12ss :tin 2. 5 tiles as long as vidth of

C , atrLni...... Syspasis TBIBE PBOTICHIEUMONIII

Thzec species of 2 genera are describsd and illuszrate3. Species of Prozichnenmonini figured in the

pepticus Cresson by Short

tterstedt ky Short A1 970% / 1378) ; and hbfyjogpa proteas Christ, Coel$chneumon

nigsrrimus ? Stephens, C, subquttatus Gravenhorst, :

Crattojoppa-- robasta Caasron, Protichneamon sp. , Syspasi s ,' -=#a (fleinr'ch) , and 5. lineator (Pabricius) by Short

Key to Genera of Protichnenaonini

1. Closing app~ratusof prothoracic spiracle more

i f10s-&g+aeparatus - - of protnoracic spiracle much less thm 2.5 tiles ;s long as width of * . atrinm...... p ...... ,.. . .Syspasis Heinrich (1360a, 1367a) placed the Protichneumonini * phy logenetically botveen the Ichneanonini and the Trogini. .-"-.&- Bart advanced species show affinities with the Trogini, . while the less advanced- have definlte similarities with the Ichneu~onini. The f inal-instar larvae are easily separated from thcse of the more specialized Trogini; while the~~~cannot - -- - - w d be separated as a group from t%eCIchneumonini, the 2 genera can be separated individually.

The ~orphologyof the clypeolabral plates in both Syspasis and Coelichneunon strongly resembles those in Ichnen~onand Orgichneumon (Ichneumonini: Ichneumonina) , - which suggests a close rolat

Syspasis tanma has a prothoracic spiracle with a very short clcsing apparatus, completely unlike that of * ------Coe lichnenmon. aoplns (Ichneuaonini: Cratichneuaonina) has ------I a similar closing apparatus but not the medial join between the clypeolabral plates, which places Syspasis near

Ichneumonini: Ichneomonina. -S. lineator, illustrated by Short ( 1978) conforms to this description. Callajoppa and Canocolama (Trogini: Callajoppina) likewise have short closing apparatuse s.

------. - The larvae of other species of Protichneuaonini illustrated by Short have elongate closing apparatuses,

- although often not as long as those illustrated here.

The two extremes of closing apparatus length within

this one tribe suggests either a polyphyletic crigin for the

tribe, which is not supported by the adults, or diverging

-- --- evolutionary trends. ------

Genus Coslichneumon Thonson

Two species are described and illustrated and a

key for their separation follors. The generic , character, the closing apparatus of the prothoracic

spiracle with length more than 2.5 times the width of ------the atrium, is a relatively constant one, though some

------185

------

of the species illustrated by Short (1978) do not show it.

Key to species of Coelichneuinon.

1. &axi;llaxy palps with gfeatest diatlheter- laore - - -A- than half of depth of inferior plenro-

stoma1 process...... a a. a a .. . a .a a brunneri (Roh.) raxillary palps with greatest diameter less

than half depth inferior pleuro- stoma1 processa.a..a.a.aaaaaa.aa~a.a~orpheus(Cress.)

a, Coelic hnenlaon brunneri (Rohuer)

The 2 specimens examined were reared from 4 Dioryctria spa (Pyralidae) collected at Hammette Lake

fload, BaC. (PIS 55-6854-02-1) and frona Dioryctria ancanticalla (Groteq collected at Elko, B.C. ($IS

at haad end of host pupa, terainal, about 3 me in diameter.

* pleurostoaal processes; ocular lines lightly sclerotized,

- not always visible, length about 1.6 times depth between pleurost cmal processes, with reticulate sculpture on

surface. Hypopharyngeal surface a>d hind end of larval skin, without visible- spines. Light accessory reticulate sclerotization originatin g near distal end of hypostoma.

atrium of prothoracic spiracle (Pig. 841 C.046 mm vide

(tasge 0.044 to 0.048), with short, heavy spines, no fine

hairs inside. Closing apparatus 0,123 mm

long (range 0. 124 to 0. 136) . ,

Coelichneneon orpheus (Cresson)

Figs. 82, 85

The 2 specimens 2xamined were reared from the

American dagger moth, Acronicta americana (Harr .) (locrnidae) , collected at Hount Lowette, aue. 8. americans is a new host record for g. orphens.

about 7 in diameter. Caphalic structures of f inal-lnstar larvae (Pig. 82)

1.63 am wide in one specimen an8 1.65 ms in the other.

Epistaaa complete and sclerotized, with 9 pores in one specimen and 12 in the other; plert~ostonasclerotized with heavif y-scf erotized accessory pleurostoaal area, Hyposto~a sclerotizeb, between 1,s anA 1,B +Alaes longer than ~3ept-h - betusen pleurostamal processes. Clypeolabzal plates heavily-sclerotized, joined inedially by heavily-sclerotized area, each wider than deep with 5 to 9 sensilla. Superior pleurostomal process projecting, postclrior strut not visibf e; inferior pleurostoraf process fused ventrally, trough-shaped, struts of approximately eqnsl width and depth. Silk press protruding ventrally in bib-like process,

%boat 1.3 times deeper thbn wi-de, openihg-sfig-htlywk3er - - \ than deep, seta on each lateral margin bslov ventral aargirr gf opening, ltariflary and labial palps each with 5 - serisilla, greztest diameter cf raxillsry palps much less than half of depth of inferior plenrostomal process, palps dispr~portionat €1 y srall in relation to cephalic structare. t I d $ Iadible about 1.2 tims longer than wide, blade 0.35 to

0 of ~ ~- - --- i !8 30talLLeen2thof aartdibAe,~-- P *i

-- - ~ - -- - ~~ - - - -

hntennal discs vith heavily-sclerotized rims, greatest A diaaarer between 0.90 to 1.0 of depth betreenpleurostomal processes; ocular lines heavily sclerotized, length 1.4 to *

1.5 tia~sdepth betmen inferior pleurostonal processes, with reticullte sculpture on surface. ~ypophargngeal

surface without visi Ole spines; heavj accessory reticdate- - sclerotization originating at distal end of hy postora. Hind end of cast larval skin with small spines in rows,

-

Atrium of prothoracic spiracle (fig. 85) 9.074 am vide

(rango 0,066 to 0,082), with short heavy spines, no visible fine hairs inside, sari ace with heavy, irreglar lines of scl3r~tization. Closing apparatus 0.223 am long (racgb

Only 1 species of Syspasis is described and ilius?rated. This ganas, as characterized by -5. taura (Heinis) , is quite diff srent frro~the genus Coelichoeuaon.

of the prothoracic spiracle, which is less than or equal to ------the uidth of the atrium. ------4 The postion of ths genus is in some dispute, as Townes \ ; 4. 4 (1965) placed it in the Ichnenmonini and Hsinrich (1969)

placed it in the Protichnzumonini. The adults show some " +. 2 similarity to speci~sof Ichneumon but differ in biological ; !

attributes (3. Bairon, !3iosystematics Research Institute, j 1 1 4 it be Ottawa Ont., pers coma.) . Therefore should retained -- -- I 1 4 in the Protichneumcnini . 9i

Syspasis tauma (Heinrich)

Pigs. 83, 86

T~P4 specimens examined were reared from species of Gaonetridae collected at Meddle Bra, Man.

CFIS H2808) and Plaaders, Ont, (PIS - LC071 g and from - unidentified species of Lepidoptera collected at

Constance Bay, Ont. (PISD46 4788) and Curtis Twp., N.S. '\ (PIS S46-1490P) .

Parasite cocoon not visible inside host pupa; exit-

hole at head end of host pupa, terainal, about 3 nm in Cephalic structure of final-instar larva (Fig. 83) 0.68 =. =. mn wida (range 0.65 to 0.70). Epistoma complete and sclerotized, with 8 to 10 pores; pleurostoma sclerotized,

with narrow, light ly-sclerotized accessory pleurostomal

area. Hypostoma sclerotized, length 1.4 to 1.8 ti~esdepth between pltrarust6~af processes. Clypeolabral plates *i-u

aedlally by indistinct membranous area, each wider than

deep, each with 5 to 7 sensilla of which 2 or more are on

- membranous areas appended to plates. Superior plenrostomal process projecting sightly, anterior and posterior struts

about aqua1 size; inferior pleurostomal process -with

equal-sized struts fused ventrally tc form trough. Silk press slightly extended ventrally, about as wide as deep, i opening about as wide as seep, T seta on each- lateral \ \' 1 margin, seemingly variable in location, always above ventral margin of opening. Haxillarp palps each with 1 large and 5

saall sensilla or with 1 large and 4 small sensilla;

labial palps each with 1 large and 4 small-,. sensilla.

andibles normally 1.4 tiaes longer than wide, sometimes

about 1.2 times longer than wide; blade 0.35 to 0.43 of Antenna1 disc lot visible; ocular lines lightly

sclerotized without reticulate sculptuqe,.-A. A

4 > *- times the depth between p leur cstoaal processes. Hypopharyngeal surface with medial spines single 6; double and lateral ones in rows of 4 to 6; light accessory ret iculte _sclerxttization originating f roa_ di~tale_nd__of hypostoma. Hind end of larval skin without visible spines.

Atriu~of prothoracic spiracle (Pig. 86) 0.050 ma wide (range 0.044 to 0.058) , without visible spines or hairs inside. Closing apparatus 0.041 an long (range 0.030 to 0.050) and iongth always less than width of atrium. Pigs. 81-82. Cephalic structures of f inal-instar larvae: C 8 1, Coelichneunas brunneri (Rohw.) ; 82, -C. orpheus (Cress.) .

Figs, 83-86. 8 3, cephalic structure of f inal-instar lasta of Syspasis taura {Heiar.) . 84-86, Prothoracic spiracles of flnal-knstar larvae: ' 84, Coelichneumon branqari (Bahv.) ;

85, Coefichneouon orFheas (Cress.) ; 86, Syspasis tauma

final-instar larvae suggests,that a great deal of divergence has occurred between the larvae of the two groups.

The Trogina are parasites of swallowtail butterflies (Papilionidae: Papilio) , while the Callajoppina are garasites of sphingids (Sphingidae) The larvae of Trogina . ------have a distiictive hypostoma, w,hich is bent at about its i midlength, and the .Callajoppina do not. Other features of dif ferenc= between the two subtribes are: cly~eolabral f plates joined medialy izh~d~op~ina,not joined in the \ Trogina; blade of mandible distinct, long and narrow-a in t e Calla joppina, indistinct, short and relatively broad in thu

Trogina; and closing apparatus short in Callajoppina, long

Areoscelis rnfa as illustrated by Short (1978) is a a

taxonomic anomaly in the Trogini. It has all of the above R L - characterstics of the callajoppina, including a sphingid \

" host, but Townes and Townes (1966) placed it in the Trogina ?g .. .- -x (sensu Heinrich 1962c, l967a) , closely aalied with Trogns.

>+ = Dr. B. Townes, Eserican Entomological Institute, Ann Arbor; ------'$T* Michigan, (pers.*aa.-- f said its hosts were Papf lio spp. - - - -5.

3-9 - %. - -. < One of two possible situations exists with respect to A* -rufa. One is that the adnlt has been placed in the wrong subtribe by Fovnes and that it truly belongs in the Calla joppina, the adnlt of A. ruf a resembling Trogus through convergence. I! kncwledge of the hosts would be valuable in assessing this possibility: if -A. -rufa is parasitic on

Sphingidae it belongs in the Callajoppina, whils if it is - parasitic on pilio it is more. likely to b~longin the Trogina.

, The secorid, and less likely of the possible situations is that -A. is truly in the Trogina and therefore the classification of ths larvae presented here is coincidental and has no relationship to-the adults.

Sub-tribe Calla joppina

>

The species in two genera are described and illustrated. The subtribe is characterized in the larvae by a the presence of abundant, long heavy spines inside the Conocalama occidentalis (Cress .) has the unsclerotized outlines of labial and stipital sclerites and a hypostomal

spur which Conocalama bolteri (Cress.) and Callajoppa

cirrogaster do not. However, C.- cirrogaster, as illustrated by Short (1378) , does have these sclerites as d~es

Areoscelis- --ruf a. ---The presence of these scleriA- hes, which -

.L further define and isolate the subtribe,. may be dependant on either the quality of the mount or on some host effect.

Key to genera of Callajoppina

1. Epistoaa with more than 10 pores...... Conocalama Epistoma with less than 10 pores...... Callajoppa

Genus Calla joppa Cameron

one species is described and illustrated. -C.

cirrogaster was illustrated by Short ( 19781 and agrees with the specimens examined. Callajoppa cirrogaster (Schrank)

Pigs. 87, 32

The 3 specimens examined were reared from Dilina tilias

(L.) (Sphingidae) collected at weiden Obpf., Germany.

-

Parasite cocoon of fine silk spun over entire inside of + hosc pupa; exit hole at head end of host pupa, terminal,

about 8 mm in diameter.

Cephalic structure of final-insta larvas (Fig, 87) 1.42

mm vide in ons specimen, 1.44 in second and not measurable

in third. Epistoma complete and heavily sclerotized, with 6 - =%

to 8 pores; pleufostosa fteawily scIerokize4, with hsa+ifg -

sclarotized accessory* p rostomal reas, margins of pleurostoma indistinct rflypostoiaa f%heavily sclerotized, length 1.3 to 1.5 tines depth between pleurostcmal

processes. Clypeolabral platss each joined by membrane to ,spistoaa well above superior plenrostoaal process,

d indistinctly joined &iallJ, each with 8 to 10 sensilla.

Saparior plenrosto~al- process- -strongly protruding, ------.5 triangular, posterior strut not visible; inferior ------Q ------ple urostomal process with struts fused, trough-shaped.

Siik press 5 qnared v entrally but not extended, about 1.3

times vider than4deep, opening about as wide as deep, a seta

on aach ventrolateral corner of silk press below ventral

margin bf opening. Maxillary palps and labial palps each

with 4 Xarger and I very small sensilla; usually one

'sclarotized sensillun adjacent to each maxillary palp.

klandibles about 1.2 times longer than wide, blade narrow and

-- - distinct, 0.32 to 0.38 of total length of mandible.

Bntennal discs with wide, heavily-sclerotized rims,

/ greatest diameter about equal. to depth bet wee^ pleurostomal

processes; ocular linss heavily sclerotized with reticulate

scnlptnre on surfaces, -length I. 3 -to 1.6--times degkh-bet wee%

pleuostomal processes. Hypopharyngeal surface without

spinss, soms areolate senlptnre visible; accessory

reticulate sclerot izatio~ilight and restricted. Hind end of

larval skin with small spines in rows.

Atriua of pro thoracic sp iracle (Fig. 92) 0. ide

(range- 0,110 to- 0.140)--- - with- - abundant ------long -- heav.1 - spines- inside. Closing apparatus 0.116 ma long (range 0.090 to ------0.1 SO) . Genus Conocalaaa Hopper

Two species of Conocalama are described and illustrated. Conccalaaa has 13 or more pores in the - -- - - epistoaa, which distinguishes it from Calla joppa, which has

6 to 8 pores In the epistoma.

Key to speciers'cf Conocalana.

1. Less than 15 pores in epistoma; 5 or 6 sensilla on clypeolabraf plates; length

of hypostoma less than 1.5 tines depth ------between plenrostomal processes...... bolteri (Cress.) nore than 15 pores in epistona; 40 or 12 sensilla on clypeclabral plates; length

of hypostona more than 1.5 times depth -between plenrostomal processes.. ..occidentalis Cress. Conocalama bolteri (Cresson) Fig. 88

The 1 specimen examined was reared from a species

of Sphsngidae collected at landerhoof, B.C. ' (PIS B.C.

Parasite cacaon ~f spars2 fine silk inside host . pupa; exit hole at head end of host pupa, tersinal, not measured because host pupa broksn.

Cephalic strncture of final-instar larva (Pig. 88)

sclerotized, with 23 pores; pleurostoma heavily scisrotized, with heavily-sclerotized accessory

ple urostonal area, lateral margins of pleurostoma

distinct. H~~OS~OE~heavily sclerotized, length 1.25 : to 1.30 times depth between pleurostcmal processes. 5

Clypeolabral plates not joined to a epistoma, joined aediallp, each wider than deep and each with 10 or 12

------sensirla. Superior plenrostural process large and

------protruding, posterior strut concealed by anterior strut; inferior pleurostornal process with struts

apparently not fused ventrally, and anterior strut a about same size as posterior strut. Silk press extended vsntrally , with aieolate sculpture over entire --- surface, about 1.2 times wider than deep, opening about

.1.2 times wider than deep and with a ssta on each lateral margin below ventral margin of opening. Plaxillary

paps aach with 6 sensilla; labial palps each uith S or 6 sensilla. Mandible about 1.1 times longer than

wids, blade narrow and distinct, 0.45 to 0.50 of total length of mandible.

Antenna1 discs with heavily-sclsrotized rims,

- - - gre3test dianeter a little less than depth bstueen pleurostomal processes; ocular lines heavily \ sclsrotized, with reticulate2 sculpture on

surface; length about I .9 times depth between

pfeurostoeal processes. Hypopharyngeal surface without '

~isiblespines; accessory reticulate sclerotization light. Hind end of larval skin with small spines in Atrium of prothoracic spiracle 0.11 6 ma wide on one side and 0.130 me wide on the other, with abundant, long heavy spines inside. Closing apparatus about 0.lUO mm long.

The prothoracic spiracle is not illustrated as neither spiracle was suitabls. They are nearly identical to those of C. occidenta1Ls 5ut are slightly larger on aoQrage.

rr Conocalama occidentalis Cress.

'T* t

The 1 specimen examined was reared from Smerinthus cerisyi Kirby {Sphingidae) collected at Salmon River, B .C.

------Parasite cocoon hot visible inside host pnpa; exit hole at head end of host pupa,- terminal, no measurements taken because host pupa broken.

Cephalic structure of final-instar larva (Fig. 89) 1.32 rm wide. Epistoma complete and heavily sclerotized, with 13 - paras; plenrostona heavily sclerotized with scilerotized timas depth bet ween pleurostonal processes. Clypeolabral plates net joined to epistoma, indistinctly joined medially, each with 5 sensilla, 2 of which are appended on 4 unsclerotized azea beneath plates. Superior pleurostoaal process protruding, rounded, posterior strut not visible; fused ventrally, posterior strut slightly longer than anterior strut. Silk press aDout 1.1 times wider than deep, not sxtended ventrally, with areolate sculpture on lateral surfaces, opening about 2.4 times deeper than wide, 1 seta on each lateral margin about parallel with ventral margin of f opening. Maxillary ,pslps each with 5 sensilla, 1 of which appears bi-lobed; labial palps each with 1 larger and 3 smaller sensilla, orpwi€h 5 orequalr size. Ussclerotiied outlines of labial sclerite and what is either the hyposronal spur or lacinial sclerits, 0.r both fused, are visible, Mandibles, about 1.2 times longer than wide, blade narrow and distinct, about 0.36 of total-length of mandible.

Bntennal discs with heavily- sclerotized rims, greatest diaacter abo~lt-klt=~>~aatfers-aL&g&Lhe~eu~ostamal pgocesses; ocuiax ILine~heavipscLerotizedcwithticulae ------sculpture on surface, length about 1.4 timas depth betveen pleurostomal processes. B ypopharyngeal surface without visible spines; accessory reticulate sclerotization light and indefinite. Hind end of larval skin with, small spines in. short parallel rous. .

------Atria. of prothoracic spiracle (~ig.93) 0.1 10 mm vide on one side and 0.120 am wide on the other, with abundant long heavy-spines inside. Closing appaxatus abhlLtOAY&m long.

Subtrlbe Trogina

Two species of Trogns are described and illustrated.

The subtribe can be defined with some confidence by the bent hypostoaa because Short (1378) showed another genus in the

Trogina, Holcofoppa, to have the same character. The genus ~o~lismenosof the Tchneumonfni. which has a hypostoma similar to that of Trogas in one specimen of & rornlus

-- .ornlus~can= separated from ~rbginaby the following: Bopliswenns has the clypeolabral plates joined medfally,

> species cf Trogina apparently do not; the spisto~a sclerotized, end cosplete in Hoplismenus, while it is

light ly-sclerotized, nnscleratized or incoaplete in Trogina; and the silk press is extended ventrally in Elo#ismeaus

- vhile it is not in Tragina. Ir. is possible- - -- - that- - - the long, bent byposto~ais adaptive for parasites of Papilionoidea,

as Fioplis~eansis parasitic on Wy mphalidae. 4

Key to species of Trogus. r

1, Labial palps each with 6 ssnsilla, one of which is apparently appended to the margin

- - of the -pala,*--,**-- **-,,,-

Labial palps each with 5 sensilla, one of . which lay be apparently appended ts the margin of the pulp. .,...... folvf pes Crsss.

?ragas fDfripes Cresson

Figs. 30-, 34

md 91ack starq& lake, Oat. and fro. an unidentified species cf Papifio at an unspecified loca,tion in Canada.

Parasite cocoon 'of fine silk i~side, enclosing

anterior end of host pnpa about 2 to 4 am frcilr head.- Exit . at end hols head of host pnpa, subterminal, located eithera - - -BartSxXZg7 -veXtr=aISry, or:Ta€eTxlZy ab6u t 5 mifrO~tlp o t- ' 2.' bead; 3 to 4 ai h diaaeter, with a distinct burnt

. Cephalic structare of final-instar larva (Fig. 90) 0.85

wide (rangs 0.84 to 0.86). Epistoma complete and *.

ansclerotized, withaat pores; pleurostoma scle&tized with , i '\ lightly-sclerotizsd accessory pleurostomal area. Hypostoma

plates plenrostosal processes. Clypeolabral - lightly sclerotized, not jcined redially. or to epistoma, each with 2 -- to 6 sensilla. Superior phnrostomal proce ES protruding, uizh both strnts visibls and anterior strut slightly longer

than posterior str nt ; Pnf erior pleurostomal process with

struts lrot fused ventrally, posterior strut longer than

p- -- - b C~L10 I: bT;~-press marginally via- t*mn ampr not vide, 1 seta on each latkral margin, slightly above ventral

margin of opening. Maxillary and labial palps each with 1 t

very saall and 4 larger sensilla. ' Handible 1.2 to 1.3 ti& B longer than wide, blade short, broad and indistipct, 0.30 to

0.33 of total length of mandible.

P Antenna1 discs usnally not visible, but if so, then 4 4a unsclerotized, greatest diameter about 1.5 ues depth F . between pleurostomal 9 racesses; oculB r lines usual1y t visible, lightly sclerotized, with faint, reticulate t sclorotization on surface, length 2.0 to 2.7 times depth

bdtween pleuro,stomal processes. Hypopharyngeal surface and

reticulate sclerotization light and indistinct.

Atrium of prothoracic spiracle (Fig. 94) 0. CL79 mm wide

(range 0..072 to 0.086). with some short heavy spines inside. \ Closing apparato&40, 189 ma long (range 0,158 to 0.200). \ Tro gns pennator Fabricius

Figs. 91, 95

The 3 gpecimens examined were reared from -Papilio ajax L. (Papilionidaej collected at Baddeck, B. S.

Parasite coccon of fine light silk in nearly-entire* sheet around inside of host pupa, closing off front egd about 2-4 mm from head. Exit hole at head end, subterminal,

located laterally about 8 mm back from head, about 5 -ma by 4

ma wide, margins with distinctive black burnt appearance.

Cephalic structure of final-instar larva {Fig, 911 0.94

en wi-ds [range 0,9CLtu0.37). Zpistuma cersp2et+-anil- - - unsclerotized, without visible pores; pleurostcma

sdarotized, with lightly-sclerotized accessory pleurostomal areas. Bypastoma sclerotized, length 2.2 to 2.8 times depth L between pleurostoral processes. Clypeolabral plates lightly sclerotized, small and tapering, not joined medially nos to epistoma. Superior pleurosto~alprocess protr uding, with struts about equal length, posterior strut mostly concealed ------p-L -

\ by anterior strut; inferior pleurostomal process with struts ------not fused. ventrally, posterior strut slightly lcnger than anterior strut. Silk press about 1..5 times deeper than wide, opening about 2.3 times deeper than wide, 1 seta on - sach lateral margin, about parallel with ventral margin of opening. Haxillary palps each with 3 large and 2 small sensiUa, labial palps ~5th3 largo and 2 small sen-sills-an3

1 sensillurn appended to margin of each palp. dandibles about 1.25 times longer than wide, blade short, broad and indistinct, about 0.27 of total length of mandible.

Antenna1 discs usually not visible, if so then lightly sclerotized, and greatest diameter about equal to depth between pleurostomal processes; ocular lines aluays visible,

------lightly sclerotized, with reticulate sculpture on surface, length 2.6 to 3.3 times. depth between pleurostcmal processes. Hypopharyngeal surface and hind end of larval skin without visible spines; accessory reticulate sclerotization light and indistinct.

I Atrium of prothoracic spiracle (Pig. 95) 0.031 ma wide Pig. 87. Csphalic structure of final-instar larva of Callajoppa cirrogaster (Sc hrank) .

Figs. 92-95. Prothoracic spiracles of final-iastar larvae:

92, Callajoppa cirrogaster (Schrank) ; 93, Conocalama occi dentalis Cress. ; 94, TroquS fulvipes Cress. ; 35, -T. pennator Fab.

- - - - - DISCUSSIOI AN? COUCLUSIOIS w

conclusions on the .classification of the Ichneumoninae reached in this work are based primarily on characteristics of the f inal-instar larvae. Information provided *by adult aorphalogy is used to snppfe~sntmany of these conclusions, - - - - - LL - -- especially those concerned with relationships between taxa.

As relationships between genera and species are discnssed

* antler generic a& tribal head3ngs, onZp bmad - -- - - considerations and conclusions are discnssed here.

Characters were fonnd which distinguish all of the

gerera dnd species establiskea or the basis cf adnlt characters. According to van Emden's (1957) concept of - - L------taxonoaic congruence between larval and adult '--. '-. classifications, the classification of species and genera

considered in this vork appears to be a natural one. In addition, the larval and adult classificati.06~of the tribes and subtribes almost coincide. The few differences that exist

are apparently due to the nature of the subfamily and not to Heinrich f 33603, b, 1361a ,b, l362a,b,c, lgEt7a, b,c,

1968a,b, 1363, 137 1, 1972, f 973, 1975) has consistently noted the existence of species which are intermediate in form between related taxa, and which could easily be placed in either taxon, Eeirtrich - (1 367a) considered the Ichnenmnina

and diversifying at a rapid rate. He also hypothesized that ancestral species of many taxa are still present today in a

- -- - - relatively unchanged state, and are reprosehted by forms intsraediate between taxa. In this sf taation, charactets fonna in other fife stages, p&ticnlarly the final-instaz larvae, can contribnte enorronsly to the placement of apparent intetmedf ates in* one taxa or the other ar to the

allows the *tightecingu of brderlines bet ween taxa ,* and resalts in an faherentfy tore stable c2assiffcatiun in which the relationships betreen taxa are known.

The form and structure of the clypeolabral plates has proven to be the aost useful larval character for higher classification of the Ichoeomntnae. Case 8-t be taken not to over-~rocessthe srrecimens in KOH as this can result in ' destruction of that character, An ander-cleared specimen is aach preferable to an over-cleared ma,

The adults of the Phaeugenini are the least I specialized, and presombly the most primitive of the

_7__------A A A - -- - - Xc3iEii~on1nae [G. Heinrich, Dryden, Haine, U.S. A., pers..

coin,), This can also be seen in the finaf-instar larvae

where the clppecfabral plates are usual. foined rigidly txx --= - - the epistoma and apparently not free to extend and probe into the entironment ahead of the larva, as they are in other groups, blso, in the final-instar larvae of the Phaeogenini, the blade of the mandible is strongly-carved, siailaz to the early-instar larvae, while other tribes of

condition and hare Bore or less straight blades.

The adults of the genns Cratichnenaon (~chnenrdnini: Cratichztenmxiina) are closeXy related to the tribe * Phaaogeaini (Heinrich 1347~);so much so that Beinrich

cu~sideredthe pfaceaent of so8e of the smaller species of Cratkchneatonin~erthat genus, or in the Phaeogenini, The larvae of Cratichneumon, and the c osely-related ff genera Aoplus and Vnlgichneunon, are also closely related ta

7- the Phaaogenini. In these genera, the clypeolabral plates are joined loosely to the epistoma by an unsclerotixed .area of cuticle, l!hiiss character-is not_seen in other _gener~_oL--_- .2 Ichnaumoninae, barring Callaj oppa cirrogaster (Trogini: Callajoppina) where the area of cuticle joining t=he

- clyp+olabral plates to the epistoma is located much more dorsally than in the above genera, which suggests that it is a secondary development in L' cirrogaster.

A Phaeogenes hebrus (Phaeo genini) has a primarily

+em ~E~EK,us -afea--of-csnt i&e johing-t he ef gpeokabra-2 -ea tes to the epistoma. ~lthoughsmall p ts of sclerotization

in this cuticle allow pfaceaent of the species ~iththe Phaeogenini, other larval characters suggest a slxonger tie with the ~ratichd~nmon-likegenera. These include the presence of conparatively large spines on the hind end of the' larval skin, which other Phaeogenes do not have, and

nany Cratichneamon do; and a relatively straight mandible - -- blade. ------The final-ihstar larvae do not provide any evidence of relationships between the ~ratichneumonina, ~rnblytelina, and

, Ichneumonina. Houever, no species known to be intermediate - between these groups were examined. The adults-of the two 4f gen+ra, ~h~ratslesand Patrocloides, in uhich the larvae

unrelated to rati ichneumon and more closely related tb 0 Ichneumon.

The clypeolabral plates of Hoplismenus (Hoplismenina) are similar to those found in ~ch~eumon(Ichneomonina) which suggests a-relationship between the t wo~_groops.

Similarities in the adults are the distinct gastrocoeli and

------the oxypygons apex of the abdomen.

Of the Platylabini, Heinrich (1962b) has said, "This is P the most clearly defined tribe of the .subfamily in structure as well as biology. The borderlines between it and othex , tubes are clear cut and muddled nowhere by linking form^.^^ The larvae of the two genera examined have distinctive - - dlar tc those of Cratichneumon, although they ars not ?joined laterally to the epistoma. The round to semi-oval propodeal spiracles of the adult suggest a relationship to the smaller Gratichneumon or tbe

Phaeogenini, but, in the absence of linking forms, there is - IK) direct evidence for this. The examination and analysis of further spacies will be required to place -their

relationship to the ether tribes of the ~c~~aonini,

The Listrodromini is a well-defined tribs, .both

biologically and morphologically. A11 species of the tribs are parasites of Lycaenidae (Papilionoidea) , Beinrich . - (1962a) thought the tribe to be related to 'the Ichneumonini ------through the genus ~rogohrpha, the adults of which have many structural similarities with the Listrodromini, The hosts \ of Trogoaorpha, skippers (Hesperidae) , are also closely related to those of the Listrodromini (Heinrick (1962a) . t Latoas of species of Trogomorpha were not available for study. The short mandible blade is distinctive to the ~istrodroaini. Similar mandibles are found in Eoplismenns and Thyrateles (Ichneuronini) , and in Trogns (Trogini) ,

-- - -- which are also parasites of butterflies (Papilionoidea) . The adults ate quite dissimilar, so larval resemblance is probably due to coavergenca.

The Protichneunonini are difficult to assess,' partly because a limited number of species were available for

The character of the propodeum used to se~aratethe adults of Protichneumonini from other tribes of the

Ichneuaoninae is an intergraded one (Beinrich 1960a) ; the

boandar ies intergrade at one end with the Ichneumonini and

at the other with the Trogini, A secondary character of wing venation which distinguishes the Trogini $as eliminated \ ---..A

their confusion wf th the ?rot i&nenmoni;ni;- (Hebrieh 496-7~tF.-- Bowaver, much potential for confusion still exists between the Protichneumonini and the Echneuaonini. tlany features of the adults of Protichneumonini, including the deep gastrocoeli, the oxy pygous abdomen, and the r< - longitudinally-striated postpetiole suggest deir close relationship to Ichneumon and ~rqichne~bmon(Icbneumonini: Host 'of the larvae of Protichneumonini, including those illustrated by Short (1978), can be distinguished fron the

Ichneumonini by the very long closing apparatus of the

prothoracic spiracze. The clypeolabral plates of most A species are joined ~bediallyand are wider than deep,

a ------suggesting a r eflationship with ,Ichneumon ant% brqichneumon.

Syspgsis tauma, illustrated here, and .S. lineator,- * - Z illastrated by Short (1978), are exceptions: both species have the length of the closing apparatus about equal to the width of the atrium, which is diqmetrically opposed to the

- situation in other Protichneumonini. The clypeolabral plates of 2. tauma are also similar to those bf Ichneumon + ------and Orgichneumon. Shor? (n1978) does not show clypeolabral * . plates in -.S. lineator.

The evidence fron the larvae suggests that Syspasis may

be more closely related to the Ichneumonini than to the Pretichneumonini. Borarer, examination of the Gts in * J light of this information will be required to determine if a mnge3n postionoT;ypsisf r6iTProtichneunonini to * , I /

nmmEmi*jo~tiriea. rheotberspecles* - ---- t Protichneunonini form a fairly well defined unit. ------Larvae of the two subtribes of the Trogini are divergent in form, while the adults are more similar to each

other. The larvae and the adults both indicate that each of

the two subtribes, Calla joppina and Trogina, are natural groups, quite distinct from each other and from other tribes

in the Ichneumoninae. Heinrich ( l962c, t967a) placed the ------tribe Trogini at the peak of the evolutjonary hierarchy of the Ichneumoninae, as the most advanced and specialized

groap in that s~bfem5fp. Therefore it is surprising to - - find, in a supposed&y ~onophyleticgroup, more differepces between the larvae of the two subtribes than there are

1 between any of the tribes or subtribes in the subfamily. This suggests that the ancestors of the two snttribes were different, or that the tuo groups diveqed early in their ------evolutionary history. In either cqse, the larvae show that the two subtribes are snffiaigntly different to warrent consideration of tribal rank for thea.

The final-instar larvae prpvide a reliable means of identifying the genera and species of the subfamily

Ichnennoninae. In addition, eride nce from the f inal-in star.

-- - larvae has confirmed the boundaries between many of the . ,- snpar-specific groupings and indicated relationships Xetween the groups. Where the classifications of the adults and lkrae have not agreed, the differences have been spurious,

I i,e., diffexences between larvae were greater than those between adults or vice versa,

This degree of taxonomic congruence indicates that the taxonomy of the group is in good order, and that the groupings proposed in Heinrich's works and used here are natural ones, Abasa, R.O. and U. H. nathenge. 1972. ~bservationson the biology and host-ssarching behaviour Of Af ronelanichneunon sporadic~sand Cryptns niqropictus (Hpmenoptera: Ichneuaonidae) , ~upalparasites of Ascot is $elenaria reciprocaria (Le ido opt era: Geoaetridae) . Entomophaga. 17 (1) : 93-97. Arthur, a.P, 1363. life histories and immature stages of four ichneumonid parasites of European pine shoot moth, Rhyacionia buoliana (Schif f. ) , in Ontario. Can. Ent - -- L - . -- 3 -IUT8-ImI,-

\ Barron, J. Re 1 976. Systematics of Nearctic Eaceros * - (Hymenoptera: Ichneumonidae : Eucerotinac) . Natur. Can. 103: 285-375. - -

Beirne, B.P. 1941. A consideration of the cephalic structures and spiracles of the final instar larvae of the Ichneumonidae (Hym.). Trans. Soc. Br. Ent. 7: 123-190.

4 Beirns, B.P. 1343, The biology and control of the small ermine moths (Hyponomeuta spp. ) in :Ireland. Econ. Proc. Be Dubl, Soc. 3: 191-220. ..' .. 1

Camsran, E. _ 1318. A rfa&&at~e-natuLc~raU_the - -4 pea moth, cyt3i.a nigricana Steph. Bull. -ent. Res. 23: 277-313.

Caaeron, E. 1950, The biology and economic imqortance of ii 4 Aloapa dsbellator (Fa) a relrarkabl~para site of the , d swift moth, Hepfalns lapnlinus (L.) ..Bull, ent. Rss. I * 2 41: 423-438. i! i Clausen, C.P, 1940. Ento.ophagonslinsects. leu York and - 4 ~on&nf McGr au-Bill. a3 : 2 D.C. Eidt, , 1962. Distinguishing the larvae of three pupal 21 parasites of Arqyresthia laricella Kft. (Lepidoptera: 1 .Yponoa~-utidae) . Can. Ent. 94: 32- 34, - p~~-~~~p: &&t,Bee. a& eL-.Sfgpelf.- tmt. TEe-I?3%7ristory, parasites, and economic status of larch shoot motFi, Argyresthia 1aricella Kft. (Lepidoptera: Iponomeutidae) , and comparisons with 8. laevigatella Ha-S. Can. Ent. 93: 7-24. Pinlagson. T. 1960a. ax on on^ of coc6ons and poparia, and their contents, of Canadian parasites of Jeodiprion sertifer (Geoff .) (Hymenoptera: Diprionidae) . Can. - Ent* 92: 20-47.

~inla~son,T. 196Ob. Tixxonomy of cdcoons and _puparia,' and - - -their contents, of C-adian parasites .of Dl~rz0n - hercyniae (Ht g. ) (Hymenoptera:- ~iprionidae). Can. Ent, 92: 922-9415 4

Pinlayson, T. 1962. Taxonomy of cocoons and puparia, and their contents, of Canadian pa'rasites of Dipion sirnilis (~tg.) (Hymenoptera:. Diprionidae) Can. Ent. 94: 271-282.

linld$soa, T. 19d3. Taxonomy of cocoons knd puparia, and their contents, of Canadian para,si$es. of some native Diprionidae (Hymenoptera) . Can. ~nt'. 95: 475-507, - # A Pinlapson, T. 1964. The caudal appendage of final-instar * la~vaeof some Porizoatinae (Hymenopt sra: Ichneq~onidae) . ~b~~~nt.36 :1-155-115 8. ------# Pinlayson, T-. 1967a. A classification of the subfamily Piaplinae (By menoptera: Ichneumonidae) based on final-instar larval. - characteristics. Can. Ent. 39:3 1- 8 . - Pinlapson, -T, l967b. Final-instar larvae of. the hy menopterons and dipterous parasites of acrobasis spp. (Lepidoptera: Phycitidae) in the Ottawa region. Can.

I Ent, 99: 1233-1271. Pinlayson. T. 1969. Pipal-instar larvae of tuo hysenopterous parasites of a wood-boring beetle, Tetropiula vel ntinn~Leconte (Coleoptera: -- --- Cerasby&&w ~~Ex~%~~~,IT~TcGIxIQ~,~~=62- 6 3. - / ------<------0 I) - --FfrrLaym, T. tW5. 3%e q hali~~dna~iracles~ of f inal-inst ar Jarvae of the subf?arnily dampopleginae, Tribe Campoplegini (Hymenoptera: . Ichneumonidae) . Hem. ent, Sac. Can., NO. 34.

Pinlagson. T, and K. S. Hagen. 1977. Final-instar larvae of parasitic Hymenoptera. Pest Banagement Papers NO. 107 Simon Praser University, Burnaby, B.C. Canada. ~rediabi,D. 1957. Note sulla Thyraeella collaris Grav. (Hymenoptera: Ichneumonidae) parassita dell1 Acrolepia assectella Zell. in Toscana. . Boll. Lab. Bnt. Agr. 15: 2Tt-245 - - -- A A A - - - mu - --*

* Gauld, 1.D. 1976. The classification of the .8nomalinae (Hymenoptera: Ichneumonidae) . Bull. Br. nus. (Nat. Hist,) Entomol. 33: 1-135. ------

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Gillespie, DeRe and T. Finlayson. The function of the caudal appendage 'in cocoon jumping in Phobocampe sp. (Hymenoptera: Ichneumonidae: Ca apoplegj.nae), J. ent. Soc. Br. .Coluab. In press.

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Heinrich, G. l96lb. Synopsis of Nearctic Ichneamoninae Stenopneusticae yith particular reference to the northeastern region (Hymenoptera) . Part IV. Can. Ent. suppl. NO. 23.

Bei BE ic h, G , 4362a- Spepsii-s-of- Nearctic Lchneltnoninae- Stenopneusticae with particular reference to the 1' . northeastern region (Hyaenoptera) Part V. Can. Ent. suppl. NO 26*.

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Heinrich, 6. 1962~. Synopsis of ~eascticIchneunoninae Stenopneusticae with particular reference to the northeastern region (Hymenoptera) . Part VII. Can. Ent. suppl. NO. 29.

Heinrich, G. 1367a. Synopsis and reclassification of the -- -- - Iiclinenrsaninae St~rop~~e-~st-l'ca~-off~frica-scuttr-of-th-e-- Sahara (Hymenoptera). Volume I. Farmington State College Press, Fanrington, Maine, pp. 1- 250.

Heinrich, G. l967b. Synopsi-s and reclassification of the - Ichneumoninae Stenopneusticae of Africa south of the ~aahara'(Hymenoptera) . Volume 11. Parmington State College Press, Parmington Maine. pp. 251-480.

Heinrich, G. 1367~. Synopsis and reclassification of the Ichnenaoninae Stanopneusticae of Africa south of the Sahara (Hymenoptera) . Volume 111. Farmington State College Press, Farmington Maine. pp. 481-692. -

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Heinrich G. 1973. Synopsis of Nearctic Ichneu~oninae Stenopneusticae with particular reference to the northeastern region (Hynenoptera). Suppl lo, 4. Carlsonia, Cratichneuaon .f laschkai, Helanichneuron aargaritae. latur, can. 100: 461-465.

- ---minr kli, 6. T3?5: SyjopSls ~f~ea~ticIc6ineu~oninae Stenopneusticae with particular reference to the northeastern region (Hymenoptera). Suppl. lo. 5: Ichneumoninae of the Island of Newfoundland. latur. can. 102: 753-782.

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-- - --w- --BULL--%~L-u-€k 341-393.

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I Perkins, J.P. 1360. Handbooks ?for the identification of British , Bymenoptera ~chneumoeoidea, Ichneu monidae, sabfa nilies Ichneuwninae 11, Alomvinae.- kgr*F@nzrsp i-=m-tywrininae . R.. SW

solitario delle larve di papilid hobiton-6ene (tepidoptera, Papilionidae). Boll. Ent. Bologna 26: 283-3 18.

Bothschlld, G.E.L. 3374. Parasites of the armyuorm Spodoptera manrikia acronyctoides in 8alaysia. Entomophaga 3 3: 293-233,

Sadava, Dm and C.D,F. Bflfez 1367. Saxanoty of last-instar

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schiaf, A. C. 1972. The parasitoig co~plorof Euxoa oc hreo~aster(~nenee) (Lepidoptera: * Noct nidae) . ~uaestiones- entomologicae 8: 8 1-420, '"~echssz,won B. 1370. Der Parasitenkoaplex des Kleinen Prostspanriers (Operophtera brumata k) (L>~., ~somtrfdde) nnter besonderecksichtigung der

- Xokonparasiten. 1, 2. angew, Ent, 662 1-35,--- I. .' * '. I. Short, 3J,B.T. 1952. The norphology of the head of larval Hf~enoptera vith special reference to the head of Schnennonoidea, inclnding a- classification of the final

------af Tr&-B-ant,Sb~--- - instar hrrae the dzh=nid-d~& Lond. 103: 27-84.

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