rearing, identification and biology of parasitoids and predators

REARING, IDENTIFICATION AND BIOLOGY OFPARASITOIDS AND PREDATORS ASSOCIATED WITH

PISSODES WEEVILS IN CANADA

D.J.M. Williams, and D.W. Langor

INFORMATION REPORT NOR-X-386

Canadian Forest ServiceNorthern Forestry Centre

2002

© Her Majesty the Queen in Right of Canada, 2002

Catalogue No. Fo46-12/386EISBN 0-662-33396-9ISSN 0831-8247

This publication is available at no charge from:

Natural Resources CanadaCanadian Forest ServiceNorthern Forestry Centre5320 – 122 StreetEdmonton, Alberta T6H 3S5

A microfiche edition of this publication may be purchased from:

Micromedia Ltd.240 Catherine Street, Suite 305Ottawa, Ontario K2P 2G8

NATIONAL LIBRARY OF CANADA CATALOGUING IN PUBLICATION DATA

Williams, Daryl Joseph Michael, 1956-

Rearing, identification, and biology of parasitoids and predators associated with Pissodes weevils in Canada

(Information report ; NOR-X-386)Includes an abstract in French.Includes bibliographical references.ISBN 0-662-33396-9Cat. no. Fo46-12/386E

1. Parasitoids – Canada.2. Pissodes – Parasites – Canada. 3. Bark-weevil – Parasites – Canada.4. Pissodes – Predators of – Canada. I. Langor, David William , 1960- .II. Northern Forestry Centre (Canada)III. Series : Information report (Northern Forestry Centre (Canada)) ; N0R-X-386.IV. Title.

QL496.12W54 2003 595.7’17’857 C2003-980026-1

This report has been printed on Canadian recycled paper.

ii Inf. Rep. NOR-X-386

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Williams, D.J.M.; Langor, D.W. 2002. Rearing, identification, andbiology of parasitoids and predators associated withPissodes weevils in Canada. Nat. Resour. Can., Can. For.Serv., North. For. Cent., Edmonton, Alberta. Inf. Rep. NOR-X-386.

ABSTRACT Methods for collecting and rearing parasitoids of larvae and pupae of Pissodes

species are discussed. Illustrated keys are provided for the identification of 31species of Diptera and Hymenoptera, which are parasitoids, hyperparasitoids,and scavengers associated with Pissodes species. Twenty-six of these species areknown to occur in Canada, and another 6 are widespread in the United States andprobably occur in Canada. A list is provided of 33 other species which have beenrecorded as associates of Pissodes species, but whose records are suspect and havenot been confirmed. A brief discussion of the biology and abundance of eachspecies, which is included in the key, is given. A glossary is given.

RÉSUMÉ Les auteurs traitent des méthodes de collecte et d’élevage des parasitoïdes

des larves et des pupes des espèces du genre Pissodes. Ils présentent des clésd’identification illustrées de 31 espèces de diptères et d’hyménoptèresparasitoïdes, hyper-parasitoïdes et détritivores associées aux espèces du genrePissodes. Vingt-six de ces espèces sont présentes au Canada et six autres sontrépandues aux États-Unis et sont probablement présentes au Canada. Les auteursfournissent une liste de 33 autres espèces mentionnées comme apparentées augenre Pissodes, mais ces relevés sont douteux et restent à confirmer. Ils décriventbrièvement la biologie et l’abondance de chacune des espèces traitées dans la cléqui comprend également un glossaire.

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CONTENTS

INTRODUCTION........................................................................................................... 1

COLLECTING AND REARING PARASITOIDS AND PREDATORS.................... 2

KEY TO ADULT HYMENOPTERA AND DIPTERAASSOCIATED WITH PISSODES SPECIES IN CANADA ....................................... 4

NOTES ON BIOLOGY AND ABUNDANCE OF REARED TAXA ...................... 18Diptera ...........................................................................................................................18

Lonchaea corticis Taylor.......................................................................................... 18Lonchaea furnissi McAlpine................................................................................... 18Medetera apicalis (Malloch).................................................................................... 18Medetera vidua Wheeler......................................................................................... 18

Hymenoptera ................................................................................................................ 18Bracon nanus Provancher ...................................................................................... 19Bracon pini (Muesebeck) ....................................................................................... 19Calliephialtes comstockii (Cresson) ........................................................................ 19Coeloides pissodis (Ashmead) ................................................................................ 20Coeloides rufovariegatus Provancher..................................................................... 20Coeloides secundus (Dalla Torre) ........................................................................... 20Coeloides vancouverensis (Dalla Torre).................................................................. 20Cubocephalus occidentalis (Provancher)................................................................ 20Dolichomitus terrebrans nubilipennis (Ratezeburg) ............................................. 21Doryctes sp. or spp................................................................................................. 21Eubazus calyptoides (Martin) ................................................................................. 21Eubazus crassigaster (Provancher) ........................................................................ 21Eupelmus pini Taylor.............................................................................................. 21Eurytoma cleri Ashmead........................................................................................ 22Eurytoma picea Bugbee .......................................................................................... 22Eurytoma pissodis (Girault).................................................................................... 22Helcostizus albator rufiscutum Cushman.............................................................. 22Helcostizus contortae Townes................................................................................. 22Helcostizus subrectus Townes ................................................................................ 22Heydenia unica Cook and Davis........................................................................... 23Labena grallator Say ................................................................................................ 23Spathius brachyurus Ashmead .............................................................................. 23Spathius sequoiae Ashmead ................................................................................... 23Rhaphitelus maculatus Walker ............................................................................... 23Rhopalicus pulchripennis (Crawford).................................................................... 24Rhopalicus tutela (Walker) ..................................................................................... 24Trigonura tarsata (Dalla Torre) .............................................................................. 24

REFERENCES................................................................................................................ 25

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APPENDIXES1. Parasitoids, hyperparasitoids, and predators

associated with Pissodes species in Canada ......................................................... 28

2. Unconfirmed records of other parasitoidsassociated with Pissodes species ............................................................................ 30

3. Glossary .................................................................................................................... 34

FIGURES1. Lonchaea corticis Taylor wing .................................................................................. 4

2. Lonchaea corticis Taylor antennal area of head ..................................................... 4

3. Lonchaea furnissi McAlpine antennal area of head ............................................. 4

4. Medetera vidua Wheeler antenna ............................................................................ 4

5. Medetera vidua Wheeler wing ................................................................................. 5

6. Medetera apicalis (Malloch) wing ............................................................................ 5

7. Eupelmus pini Taylor forewing ............................................................................... 5

8. Coeloides rufovariegatus Provancher forewing ...................................................... 5

9. Trigonura tarsata (Dalla Torre) hind leg ................................................................. 5

10. Rhopalicus pulchripennis (Crawford) hind leg ...................................................... 6

11. Heydenia unica Cook and Davis forewing ............................................................ 6

12. Heydenia unica Cook and Davis front leg. A. Male. B. Female. ......................... 6

13. Eurytoma pissodis (Girault) antenna, setae omitted. ............................................ 6

14. Rhopalicus pulchripennis (Crawford) antenna of female. ..................................... 6

15. Eurytoma cleri Ashmead gaster. A. Lateral view. B. Dorsal view. .................... 7

16. Eurytoma picea Bugbee lateral view of gaster. ...................................................... 7

17. Eupelmus pini Taylor forewing................................................................................ 7

18. Eupelmus pini Taylor lateral view of thorax.......................................................... 7

19. Rhopalicus pulchripennis (Crawford) lateral view of thorax................................ 7

20. Rhaphitelus maculatus Walker hind coxa................................................................ 8

21. Rhaphitelus maculatus Walker antenna of female. ................................................ 8

22. Rhopalicus pulchripennis (Crawford) hind coxa. ................................................... 8

23. Rhopalicus pulchripennis (Crawford) antenna of female. ..................................... 8

24. Rhopalicus pulchripennis (Crawford) forewing...................................................... 8

25. Rhopalicus tutela (Walker) forewing. ...................................................................... 9

26. Dolichomitus terrebrans nubilipennis (Ratzeburg) forewing. ................................ 9

27. Calliephialtes comstockii (Cresson) dorsal view of gaster. .................................... 9

28. Bracon pini (Muesebeck) forewing. ........................................................................ 9

viiInf. Rep. NOR-X-386

29. Bracon pini (Muesebeck) dorsal view of gaster. ................................................... 9

30. Labena grallator Say dorsal view of propodeum and first two gastral segments. .............................................................................. 9

31. Labena grallator Say anterior view of head.......................................................... 10

32. Cubocephalus occidentalis (Provancher) dorsal view of propodeum and first two gastral segments................................................... 10

33. A. Cubocephalus occidentalis (Provancher) forewing. B. Helcostizus contortae Townes areolet. ............................................................... 10

34. Helcostizus contortae Townes lateral view of gaster. .......................................... 10

35. Dolichomitus terrebrans nubilipennis (Ratzeburg) forewing. .............................. 10

36. Dolichomitus terrebrans nubilipennis (Ratzeburg) lateral view of gaster. ......... 11

37. Cubocephalus occidentalis (Provancher) dorsal view of propodeum and first two gastral segments. .................................................. 11

38. Cubocephalus occidentalis (Provancher) lateral view of head............................. 11

39. Helcostizus contortae Townes dorsal view of propodeum and first two gastral segments . ..................................................... 11

40. Helcostizus contortae Townes lateral view of head. ............................................ 12

41. Helcostizus contortae Townes lateral view of gaster and hind leg.................... 12

42. Dolichomitus terrebrans nubilipennis (Ratzeburg) dorsal view of propodeum and first two gastral segments............................. 12

43. Calliephialtes comstockii (Cresson) dorsal view of anterior end of gaster. ....................................................................................... 12

44. Eubazus calyptoides (Martin) forewing. ................................................................ 12

45. Coeloides rufovariegatus Provancher forewing..................................................... 13

46. Coeloides rufovariegatus Provancher dorsal view of gaster................................ 13

47. Eubazus crassigaster (Provancher) gaster. A. Dorsal view. B. Lateral view. ........................................................................... 13

48. Eubazus calyptoides (Martin) gaster. A. Dorsal view. B. Lateral view............................................................................ 13

49. Spathius brachyurus Ashmead dorsal view of anterior end of gaster.............. 14

50. Coeloides pissodis (Ashmead) anterior end of gaster. A. Dorsal view. B. Lateral view. ........................................................................... 14

51. Spathius brachyurus Ashmead lateral view of propodeum............................... 14

52. Spathius sequoiae Ashmead lateral view of propodeum.................................... 14

53. Doryctes sp. anterior end of gaster. A. Dorsal view. B. Lateral view............... 15

54. Coeloides pissodis (Ashmead) base of antenna..................................................... 15

55. Bracon pini (Muesebeck) base of antenna............................................................ 15

56. Coeloides rufovariegatus Provancher lateral view of head. ................................ 15

viii Inf. Rep. NOR-X-386

57. Coeloides rufovariegatus Provancher forewing..................................................... 16

58. Coeloides secundus (Dalla Torre) lateral view of head. ....................................... 16

59. Coeloides secundus (Dalla Torre) forewing. .......................................................... 16

60. Bracon nanus Provancher lateral view of head................................................... 16

61. Bracon pini (Muesebeck) lateral view of head. ................................................... 16

62. A. Lateral view of habitus of adult male Hymenopteran. B. Lateral view of gaster of adult female Hymenopteran................................ 17

63. Lateral view of habitus of Dipteran..................................................................... 17

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The genus Pissodes includes 11 currently validspecies known to occur in Canada. These barkweevils attack the stems and terminals ofconiferous trees, and their larvae feed on phloemand pith. Most species attack weakened or dyingtrees, and they may be associated with primarypests such as bark beetles and root diseases.Some species attack living healthy trees,particularly the terminal-inhabiting Pissodes strobi(Peck) and Pissodes terminalis Hopping, which areknown to have significant economic impact onregenerating stands (Hiratsuka et al. 1995). Theliterature on the taxonomy, biology, and control ofPissodes is extensive (Langor 1998), and there iscontinuing study of economically importantspecies (Alfaro et al. 1994).

In the course of rearing Pissodes species fromattacked trees, a large number of individuals ofother insect species are inevitably recovered.Numerous species of the Hymenoptera familiesIchneumonidae and Braconidae and thesuperfamily Chalcidoidea and Diptera familiesLonchaeidae and Dolichopodidae have beenassociated with Pissodes-attacked trees. Thenature of the association of many of these speciesis not well understood, although the biology ofsome species has been studied and their status asparasitoids or predators of Pissodes larvaeconfirmed. There remain many gaps in ourunderstanding of the role of natural enemies inthe regulation of Pissodes populations, and hencethere are many opportunities to explore the use ofnatural enemies in controlling pest populations ofPissodes. Such studies could be aided by tools tohelp in identifying the complex fauna ofparasitoid and predator species.

We have developed an identification key toadults of the 31 species of Hymenoptera andDiptera that are confirmed or highly likelynatural enemies of Pissodes larvae or pupae (orboth) in Canada, according to literature recordsand examination of reared specimens in various

Canadian collections (Appendix 1). Biologicalinformation on the association of each specieswith Pissodes is summarized. Another 33 specieshave been cited in various literature sources asbeing associated with Pissodes (Appendix 2), butfor a number of reasons have not been included inthis key. Some of these records are probablyerroneous, as they involve species that do notnormally include Pissodes in their host range; inthese cases, the specimens might have beenmisidentified, or they might have been attackingother wood-boring species co-occurring withPissodes. Other records need confirmation; theseinclude single rearings of small numbers ofspecimens of species that have not beenencountered in other studies, rearing of speciesthat attack many host species and may haveattacked either Pissodes or some other host co-occurring in the reared material, reports ofunconfirmed data on specimen labels intaxonomic works, and records of species thatattack Pissodes in areas outside of Canada but thatare unlikely to occur here.

The taxonomic literature on natural enemiesof Pissodes is sparse. Most species belong togenera for which there are no identification keys,and the existing literature comprises isolatedspecies descriptions, which may contain a fewnotes on biology. For species with taxonomic andbiological literature, a brief summary and keyreferences are given here. For most species, thebest literature resources are taxonomic catalogsand overview syntheses. Catalogs for parasiticHymenoptera are available in book (Krombein etal. 1979) and electronic (Noyes 1998; Yu 1999)form, and taxonomic overviews exist forIchneumonidae (Townes 1969a, 1969b, 1969c,1971), Braconidae (Wharton et al. 1997),Chalcidoidea (Gibson et al. 1997), andHymenoptera (Goulet and Huber 1993). Ataxonomic review of the Diptera (McAlpine et al.1981) is also available, in which information on allDiptera taxa included here may be found.

INTRODUCTION

2 Inf. Rep. NOR-X-386

Most parasitoid and predator species thatattack Pissodes larvae and pupae are univoltine.They show a strong preference for mature larval orpupal hosts, overwinter as pupae or prepupae, andemerge as adults in late spring of the next year.However, some species may be multivoltine,particularly those with a wide host range, andthese may occur on different hosts in differentparts of the season. Most parasitoid species withinan assemblage may be recovered by samplingwhen Pissodes larvae have begun to construct chipcocoons and form prepupae. This stage may bedetermined by removal of small sections of bark onthe boles of trees or dissection of terminals. Torecover parasitoids that attack earlier stages, or todetermine which species may have more than onegeneration per year, samples should be takenearlier, when larvae are in middle instars.

Identification of Pissodes-infested hostmaterial is not difficult. Terminal leaders infestedwith P. strobi or P. terminalis wilt and discolor.Close examination of wilting terminals revealsnumerous clumped oviposition marks on thebark of the previous year’s terminal if attacked byP. strobi (Hiratsuka et al. 1995, Fig. 30C) andscattered oviposition marks on the current year’sterminal if attacked by P. terminalis (Hiratsuka etal. 1995, Fig. 31B). Pissodes species that infest bolesand branches attack only trees that are severelyweakened or recently killed by other agents.Infested boles and branches show no outwardsigns that they contain Pissodes, and the bark mustbe removed to confirm the presence of theseorganisms. Pissodes larvae are legless grubs withcreamy white, wrinkled bodies and reddishbrown heads. Superficially they resemble barkbeetle (Scolytidae) larvae. An easy,nonmorphological way to determine whether alarva fitting this general description is a barkbeetle or a weevil is to determine if the larvalfeeding gallery originates from an egg galleryconstructed by adults entering the bark (barkbeetle) or not (weevil). Furthermore, all bole- and

branch-inhabiting Pissodes larvae construct chipcocoons for pupation (Hiratsuka et al. 1995, Fig.30E), but bark beetles do not. Pissodes-infestedmaterials may be collected and transferred tocontainers that can accommodate single bolesections or terminals. Individual rearing ofsamples is the best approach for obtainingparasitoid information, as this reduces the chanceof co-occurrence of other potential host insects forany given rearing.

When terminals attacked by P. strobi arecollected, they must be snipped well below thefirst whorl of branches under the terminal. Mostparasitoids do not attack P. strobi until near orduring the time when host larvae are preparingchip cocoons for pupation, and this processusually occurs around the first branch whorl.Because P. terminalis usually confines its feedingto the current year’s terminal, it suffices to collectonly this part. Terminals are best rearedindividually in capped cardboard mailing tubes,hung vertically with a vial attached to a hole inthe lower cap to facilitate collection of emergingspecimens. Evidence of damage by otherterminal- or shoot-boring insects may be checkedby dissection of the terminal when rearing iscomplete or by observing the presence of hostsamong reared samples.

Insects in infested boles may be reared inlarger containers or cages fitted with collectiondevices. Many parasitoid species are minute, socare must be taken to seal larger containersadequately. Containers should be well vented toallow heat and moisture to escape and to retardfungal growth. Adult Pissodes emerge fromdamaged wood in late summer and autumn ofthat year, whereas most parasitoids do not emergeuntil spring of the following year, so provisionshould be made for ambient-temperature, long-term storage, to afford parasitoids theopportunity for diapause.

COLLECTING AND REARING PARASITOIDS AND PREDATORS

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As field-collected damaged wood maycontain a complex community of saproxylicspecies, confirmation of host identity is bestachieved by rearing the individual parasitizedhost larvae (although this approach has not beenused for predaceous Diptera larvae). With thehelp of the identification criteria provided aboveto discriminate between weevil and bark beetlelarvae, parasitized weevil larvae may be extractedcarefully from the wood for rearing. Hosts thathave been paralyzed by hymenopterousparasitoids are easily identified, as they areplump and immobile, and the dark scar of theoviposition puncture can be seen under low-power magnification. Care must be taken inremoving the host and any attached ectoparasiticlarvae or eggs. On occasion, parasitoid larvae andeggs may become detached from the host. Theseshould be placed gently back on the host in therearing containers. Larvae that are thought to beparalyzed but that have no ectoparasitoids shouldalso be reared, as they may containendoparasitoids. It is also wise to remove healthylarvae and preserve them in 70% ethanol; theseorganisms can be used to help in speciesidentification later, as the larvae of some specieshave been described (Williams and Langor 2002).

Healthy prepupae and pupae may be placed onmoistened tissue paper in petri dishes andallowed to continue development to adulthood.

Small (5- or 10-dram) plastic or glasscontainers or vials function well as rearingcontainers, with a thin layer of rearing medium(Robertson 1979) in the bottom to maintainhumidity and prevent fungal and bacterialgrowth. This medium contains an artificial dietthat will probably not be consumed, asparasitized larvae seldom return to normalactivity; the user may therefore decide to alter theformula to eliminate unnecessary ingredients.Vials may be placed in plastic bags or largercontainers with moist paper towels to maintainhumidity. Rearing containers should be stored inthe dark. For many species, diapause isfacultative, and parasitoid larvae will develop,pupate, and eclose to adulthood without coldtreatment. Species with obligatory diapause willrequire several months of cold treatment beforethey can complete development. Storage in arefrigerator or a cold room (5ºC) should suffice.In the field, parasitoid larvae occur in protectedhabitats, and hard freezing causes death.


lighter third segmentof antenna

setae

antennalseta

setae

antennalseta

lighter third segmentof antenna

For an on-line version of this key, see the Biodiversity web page of the NorthernForestry Centre Internet site, <http://nofc.cfs.nrcan.gc.ca/biodiversity/>.Figures 1-63 comprise the key to adult Hymenoptera and Diptera associatedwith Pissodes species in Canada. Figures are not necessarily numbered inthe order in which they appear in the key. In several instances figures areused more than once, to illustrate characters that help identify closelyrelated groups of species and to illustrate differences among these closelyrelated species. In these instances, the user is asked to refer to several partsof the key so that figures of closely related species may remain together.

1. Adult with two pairs of wings, forewings often much biggerthan hindwings. Larva spins silken cocoon upon emergencefrom host or emerges as adult (Hymenoptera) ............................. 5

Adult with one pair of wings, densely hairly on most of body.Larva hardens into a brown capsule upon emergence from host,with pupa inside (Diptera) ............................................................... 2

2. Body smooth and shiny. Forewing with costal and subcostalveins separate, subcostal cell extending to wing margin (Fig. 1).Lonchaea species. These species are very difficult to identify,especially the females........................................................................ 3

Body with matte microsculpture. Forewing with costal andsubcostal veins fused except near base; subcostal cell reduced(Fig. 5). Medetera species (key by Bickel 1985)............................... 4

KEY TO ADULT HYMENOPTERA AND DIPTERAASSOCIATED WITH PISSODES SPECIES IN CANADA

subcostal cell

costa subcosta

Fig. 1. Lonchaea corticisTaylor wing.

Fig. 2. Lonchaea corticisTaylor antennal area ofhead.

Fig. 3. Lonchaea furnissiMcAlpine antennalarea of head.

scape

pedicel

Fig. 4. Modetera viduaWheeler antenna.


3. Third antennal segment of male dark brown with orange orlight brown area extending over most of length. Setae on area ofhead between antennal insertions thinner than large seta onsecond antennal segment (Fig. 2) .............. Lonchaea corticis Taylor

Third antennal segment of male with orange or light brownarea near base of segment only. Some setae on area of headbetween antennal insertions as stout as (although shorter than)seta on second antennal segment (Fig. 3)........................................................................................................... Lonchaea furnissi McAlpine

4. Antenna with scape and pedicel yellow (Fig. 4). Vein dm-cu ofwing about as long as vein CuA. (Fig. 5) ................................................................................................................. Medetera vidua Wheeler

Antenna with scape and pedicel dark brown to black. Vein dm-cu about half as long as vein CuA (Fig. 6) .......................................................................................................... Medetera apicalis (Malloch)

dm-cuCuA

dm-cu

costa + subcostasubcostal cell

CuA

Fig. 5. Medetera viduaWheeler wing.

Fig. 6. Medetera apicalis(Malloch) wing.

Fig. 7. Eupelmus pini Taylorforewing.

Fig. 8. CoeloidesrufovariegatusProvancher forewing.

femur

tibiateeth

Fig. 9. Trigonura tarsata(Dalla Torre) hind leg.


5.(1) Body usually < 4 mm long. Forewing with a single pigmentedvein along anterior margin, (Fig. 7), some species with variousfaint veins or creases near base (Fig. 24). Chalcidoidea............... 6

Body usually > 4 mm long. Forewing with numerous veinsenclosing several cells (Fig. 8). Ichneumonoidea........................ 14

6. Femur of hind leg enlarged, toothed ventrally. Hind tibiacurved, fitting into groove in ventral side of hind femur (Fig. 9)............................................................ Trigonura tarsata (Dalla Torre)

Hind femur and tibia similar to those of middle leg (Fig. 10)..................................................................................................................... 7

Fig. 13. Eurytoma pissodis(Girault) antenna, setaeomitted.

Fig. 14. Rhopalicuspulchripennis(Crawford) antenna offemale.

femur

A

B

femur

tubercle

tubercle

Fig. 12. Heydenia unica Cookand Davis front leg.A. Male. B. Female.

Fig. 11. Heydenia unica Cookand Davis forewing.

femur

tibia

Fig. 10. Rhopalicuspulchripennis(Crawford) hind leg.


7. Forewing with a large darkened area consisting of pigmentspots and stout, dark setae (Fig. 11). Femur of front leg en-larged, with a ventral tubercle (Fig. 12A, B).............................................................................................. Heydenia unica Cook and Davis

Forewing clear (Fig. 17), or with small darkened spot on vein(Fig. 24). Femur of front leg similar to that of middle leg(similar to Fig. 10) .............................................................................. 8

8. Body black. Head and thorax covered with coarse punctures.Antenna of male with segments constricted on ends (Fig. 13).Gaster of female somewhat laterally compressed, narrower indorsal view than lateral view (Fig. 15A, B). Eurytoma species.These species are very uniform in appearance and difficult toidentify, especially the males. (key by Bugbee 1967) ................... 9

Head and thorax metallic green, usually with granular or striatesculpture. Antenna of male with segments cylindrical (similar toFig.14). Gaster of female not laterally compressed, about as wideor wider in dorsal view as in lateral view (see Fig. 47) ............................................................................................................................... 11

setal patch

marginal vein

submarginal vein

seta row

seta row

A

B

Fig. 15. Eurytoma cleriAshmead gaster. A.Lateral view.B. Dorsal view.

Fig. 16. Eurytoma picea Bugbeelateral view of gaster.

Fig. 17. Eupelmus pini Taylorforewing.

mesopleuron

forewing

mesopleuron

forewing

Fig. 18. Eupelmus pini Taylorlateral view of thorax.

Fig. 19. Rhopalicuspulchripennis(Crawford) lateral viewof thorax.


9. Front and middle legs uniformly light brown, some specimenswith middle femur slightly darkened dorsally.................................................................................................. Eurytoma pissodis (Girault)

Front and middle legs with dark patches on femora and tibiae............................................................................................................ 10

10. Female with a single irregular row of setae on the fifth gastraltergum (Fig. 15A,B) ................................... Eurytoma cleri Ashmead

Female with several rows or an irregular patch of setae on thefifth gastral tergum (Fig. 16)....................... Eurytoma picea Bugbee

marginal vein

submarginal vein

stigmal vein

Fig. 24. Rhopalicuspulchripennis(Crawford) forewing.

Fig. 23. Rhopalicuspulchripennis(Crawford) antenna offemale.

setae

Fig. 22. Rhopalicuspulchripennis(Crawford) hind coxa.

apical projection

setae

Fig. 20. Rhaphitelus maculatusWalker hind coxa.

Fig. 21. Rhaphitelus maculatusWalker antenna offemale.


11.(8) Forewing entirely clear, with marginal vein longer thansubmarginal vein (Fig. 17). Thorax of female elongate, withlarge, evenly sculptured and convex mesopleuron (Fig. 18)............................................................................ Eupelmus pini Taylor

Forewing with a small darkened spot on vein (Figs. 24, 25),marginal vein shorter than submarginal vein (Figs. 24, 25).Thorax of female with mesopleuron divided by areas ofdifferent convexity and sculpture (Fig. 19) ................................. 12.

Hind coxa bare dorsally or with a few erect setae (Fig. 20).Antenna of female short and club-like, with an apical finger-likeprojection (Fig. 21) ............................ Rhaphitelus maculatus Walker

Hind coxa with scattered reclining setae dorsally (Fig. 22).Antenna of female more elongate, without apical projection(Fig. 23). Rhopalicus species ............................................................ 13

marginal vein

stigmal veinparastigma

Fig. 25. Rhopalicus tutela(Walker) forewing.

T1 T2 T3

r-m vein

2r-v (absent)

2r-v

Fig. 26. Dolichomitusterrebrans nubilipennis(Ratzeburg) forewing.

Fig. 27. Calliephialtescomstockii (Cresson)dorsal view of gaster.

Fig. 28. Bracon pini(Muesebeck) forewing

T2+3T1

Fig. 29. Bracon pini(Muesebeck) dorsalview of gaster.

T1 T2

propodeum

cells

carinae

Fig. 30. Labena grallator Saydorsal view ofpropodeum and firsttwo gastral segments.


13. Forewing with marginal vein slightly longer than stigmal vein,with dark pigmented spot near wing margin (Fig. 24)..................................................................... Rhopalicus pulchripennis (Crawford)

Forewing with marginal vein much longer than stigmal vein,with dark pigmented spot around parastigma (Fig. 25).................................................................... Rhopalicus tutela (Walker)

14.(5) Second recurrent vein (2r-v) of forewing present (Fig. 26).Second and third gastral terga (T2 and T3) separate (Fig. 27).Ichneumonidae................................................................................. 15

Second recurrent vein (2r-v) of forewing absent (Fig. 28). Secondand third gastral terga fused (T2+3), joint marked by a groovein some species (Figs. 29, 47A, B). Braconidae ............................ 21

A

B

areolet

ovipositor

Fig. 33. A. Cubocephalusoccidentalis(Provancher) forewing.B. Helcostizuscontortae Townesareolet.

Fig. 34. Helcostizus contortaeTownes lateral view ofgaster.

areolet

Fig. 35. Dolichomitusterrebrans nubilipennis(Ratzeburg) forewing.

eye

notch

Fig. 31. Labena grallator Sayanterior view of head.

propodeum

T1 T2

Fig. 32. Cubocephalusoccidentalis(Provancher) dorsalview of propodeumand first two gastralsegments.


propodeum T1 T2

length wid

th

ovipositor

Fig. 36. Dolichomitusterrebrans nubilipennis(Ratzeburg) lateralview of gaster.

Fig. 37. Cubocephalusoccidentalis(Provancher) dorsalview of propodeumand first two gastralsegments.

T1 T2

eye

Fig. 38. Cubocephalusoccidentalis(Provancher) lateralview of head.

Fig. 39. Helcostizus contortaeTownes dorsal view ofpropodeum and firsttwo gastral segments.

15. Propodeum with well-developed carinae enclosing several cells(Fig. 30). Inner margin of eyes notched near antennae (Fig. 31).............................................................................. Labena grallator Say

Propodeum with few carinae and without closed cells (Fig. 32).Inner margin of eye straight or sinuate........................................ 16

16. Areolet five-sided, with one side shorter than the others (Fig.33A) or partially faint or missing (Fig. 33B). Ovipositor shorterthan gaster (Fig. 34) ......................................................................... 17

Areolet four-sided and diamond-shaped (Fig. 35). Ovipositorlonger than gaster (Fig. 36)............................................................. 20


17. First gastral tergum (T1) much longer than apical width (Fig.37). Eye narrower than posterior part of head in lateral view(Fig. 38)................................ Cubocephalus occidentalis (Provancher)

First gastral tergum (T1) slightly longer than apical width (Fig.39). Eye wider than posterior part of head in lateral view (Fig.40). Helcostizus species (key by Townes 1983) ............................. 18

18. Thorax with large brown area midventrally ................................................................................ Helcostizus albator rufiscutum Cushman

Thorax black ..................................................................................... 19

19. Ovipositor about 1.5 times as long as hind femur (Fig. 41).Tegula white. Hind tibia uniformly dark brown ............................................................................................ Helcostizus contortae Townes

Ovipositor more than twice as long as hind femur. Tegula whitewith brown apex. Hind tibia darker apically than basally ........................................................................... Helcostizus subrectus Townes

r-m vein (absent)

Fig. 44. Eubazus calyptoides(Martin) forewing.

T1 T2

propodeum

pitgrooves

Fig. 42. Dolichomitusterrebrans nubilipennis(Ratzeburg) dorsalview of propodeumand first two gastralsegments.

T1 T2

Fig. 43. Calliephialtescomstockii (Cresson)dorsal view of anteriorend of gaster.

ovipositor

femur

Fig. 41. Helcostizus contortaeTownes lateral view ofgaster and hind leg.

eye

Fig. 40. Helcostizus contortaeTownes lateral view ofhead.


20.(16) Gastral tergum 1 (T1) longer than tergum 2 (T2), with a pair ofdorsal carinae surrounding an apical pit, tergum 2 with a pairof lateral oblique grooves at base (Fig. 42)...................................................................... Dolichomitus terrebrans nubilipennis (Ratzeburg)

Gastral tergum 1 (T1) shorter than tergum 2 (T2), withincomplete carinae, tergum 2 without lateral oblique grooves(Fig. 43) ........................................ Calliephialtes comstockii (Cresson)

21.(14) Second submarginal cell open, r-m vein absent (Fig. 44). Gasterwith first three terga forming a carapace that covers most ofdorsal surface (Figs. 47A, 48A). Eubazus species ........................ 22

Second submarginal cell closed, r-m vein present (Fig. 45).Gaster with at least six large terga visible dorsally (Fig. 46)............................................................................................................ 23

22. Body dark brown, with gaster lighter apically. Fourth and fifthgastral terga (T4 and T5) wide in dorsal view, together about athird to half as wide as third gastral terga (Fig. 47A), withabundant hairs (Fig. 47B).......... Eubazus crassigaster (Provancher)

Body black. Fourth and fifth gastral terga (T4 and T5) hiddenbeneath third gastral terga or visible as narrow strips in dorsalview (Fig. 48A), with few, scattered hairs (Fig. 48B) ....................................................................................... Eubazus calyptoides (Martin)

B

A

T4 T5

Fig. 48. Eubazus calyptoides(Martin) gaster. A. Dorsal view.B. Lateral view.

r-m vein


T1 T4 T5 T6 T7 T8T2+3

Fig. 46. CoeloidesrufovariegatusProvancher dorsal viewof gaster.

B

T4

T1 T2+3

T5

A

Fig. 47. Eubazus crassigaster(Provancher) gaster.A. Dorsal view.B. Lateral view.


propodeum T1

T1

T2+3

T2+3

B

A

Fig. 50. Coeloides pissodis(Ashmead) anteriorend of gaster. A. Dorsalview. B. Lateral view.

Pt

T1thorax

Fig. 51. Spathius brachyurusAshmead lateral viewof propodeum.

thorax

T1

Fig. 52. Spathius sequoiaeAshmead lateral viewof propodeum.

23. Body uniformly light golden brown. First gastral tergum (T1)narrow and elongate (Fig. 49). Spathius species (key byMatthews 1970) ................................................................................ 24

Body dark brown, or with dark and light patches, or gasterlighter than head and thorax. First gastral tergum (T1)rectangular (Fig. 50A) or triangular (Fig. 53A) ........................... 25

24. Propodeum with a pair of lateral tubercles (propodeal tubercle,Pt) (Fig. 51). Ovipositor slightly longer than hind tibia............................................................. Spathius brachyurus Ashmead

Propodeum without tubercles (Fig. 52). Ovipositor nearly twiceas long as hind tibia .............................. Spathius sequoiae Ashmead

25. First gastral tergum (T1) covering entire surface of segment 1 indorsal view (Fig. 53A), convex in lateral view (Fig. 53B). Gastralterga 1–3 mostly striate (Fig. 53A)................................ Doryctes sp.

First gastral tergum (T1) with membraneous areas of segment 1visible laterally in dorsal view (Fig. 50A), platelike, narrow inlateral view (Fig. 50B). Gastral terga with striate area limited tomiddle of second tergum or absent (Fig. 50A)............................ 26

T1 T2+3propodeum

Fig. 49. Spathius brachyurusAshmead dorsal viewof anterior end ofgaster.


B

A

T1 T2 T3

Fig. 53. Doryctes sp. anteriorend of gaster. A. Dorsalview. B. Lateral view

26. Antennal flagellum with 1–3 basal segments concave ventrallyand with flared rim, clearly different in shape from otherflagellar segments (Fig. 54). Coeloides species (key by Mason1978) .................................................................................................. 27

Antennal flagellum with basal segments sometimes a differentlength than other flagellar segments, but otherwise similar inshape (Fig. 55). Bracon species ....................................................... 30

27. Coxa and femur of hind leg, and in some specimens middle leg,red or lighter in color than remainder.............................................................................................................. Coeloides pissodis (Ashmead)

Legs uniform dark brown to black ............................................... 28

28. Head mostly or entirely orange–red, with variable darkmarkings on or near the occelli .......................................................................................................... Coeloides vancouverensis (Dalla Torre)

Head black or brown, some specimens with pale markingsaround eye margin (Figs. 56, 58) ................................................... 29

Fig. 55. Bracon pini(Muesebeck) base ofantenna.

Fig. 56. CoeloidesrufovariegatusProvancher lateralview of head.

modified flagellar segments

Fig. 54. Coeloides pissodis(Ashmead) base ofantenna.


3RSa

second submarginal cell

2RS

Fig. 59. Coeloides secundus(Dalla Torre) forewing.

Fig. 60. Bracon nanusProvanchr lateral viewof head.

Fig. 61. Bracon pini(Muesebeck) lateralview of head.

Fig. 58. Coeloides secundus(Dalla Torre) lateralview of head.

29. Head with distinct pale markings around eye, including innermargin between antennae and mandibles (Fig. 56). Secondsubmarginal cell of forewing long, 3RSa vein longer than 2RSvein (Fig. 57) ............................ Coeloides rufovariegatus Provancher

Head without pale markings, or with faint markings arounddorsal eye margin, markings absent between inner marginbetween antenna and mandible (Fig. 58). Second submarginalcell of forewing short, 3RSa vein as long as or shorter than 2RSvein (Fig. 59)................................... Coeloides secundus (Dalla Torre)

30.(26) Head bicolored, with approximately equal areas of light anddark patches (Fig. 60) .............................. Bracon nanus Provancher

Head uniformly dark brown to black, some specimens withpale markings around dorsal eye margin (Fig. 61)................................................................................................ Bracon pini (Muesebeck)

3RSa

second submarginal cell

2RS



A

head thorax gaster

ovipositor

gastral tergites

propodeum

Hind wing

forewing2R1 cell

tegulaantenna

eye

mandible

coxa

trochanter

f emur

tibia

tarsus

tarsomere

areolet

B

Fig. 62. Adult HymenopteraA. Lateral view of habitus of adult male. B. Lateral view of gaster of adult female.

wing

antenna

eye

ocelli

tarsus

tibia

femurcoxa

M-vein

abdomenthoraxhead

T2 T3T1

Fig. 63. Lateral view of habitus of adult dipteran parasitoid.


Diptera

Diptera associated with Pissodes are predaceous,and they may also act as scavengers. Eggs are laidin adult weevil feeding and ovipositionpunctures, and fly larvae search for prey. Eachlarva remains free ranging, probably attackingseveral living or recently dead Pissodes larvaebefore development is complete.

Lonchaea corticis Taylor

Also as Lonchaea polita Say, Lonchaea rufitarsusMaccquart, Lonchaea hirta Malloch, Lonchaealaticornis Meigan

Host: P. strobi

The biology of this species has been studied inBritish Columbia by Alfaro and Borden (1980) andHulme (1989), and specimens have been rearedfrom terminals attacked by P. strobi and collectedfrom across Canada and the northern UnitedStates. It is a common predator of P. strobi larvaewherever they occur in Canada. It is recovered inalmost all rearing studies (Langor 1998), and itmay have a significant role in regulating weevilpopulations at some sites (Nealis 1998). The flylarvae attack weevil larvae gregariously, usuallypenetrating the posterior of the host, and eat theentire internal contents, leaving an empty skinwith a posterior hole. They may also feed ondead larvae. Predation by this species can destroymost of the larvae in a terminal.

Lonchaea furnissi McAlpine

Host: Pissodes fasciatus LeConte

This species has been reared only once fromPissodes (Deyrup 1978), but the record isconsidered confirmed. Specimens have beencollected from the western United States andBritish Columbia, and it may attack Pissodesspecies there, although this occurrence isprobably rare. No information on the biology ofthis species was given by Deyrup (1978), but it isprobably similar to that of L. corticis.

Medetera apicalis (Malloch)

as Medetera arctica Van Duzee

Host: P. fasciatus

This species has been reared only once fromPissodes (Deyrup 1978), but the record isconsidered confirmed. Specimens have beencollected or reared from the boles of coniferousand deciduous trees throughout North Americaand Europe. Little is known about the biology ofthis species, but like other species in the genus itis probably predaceous on a variety of bark- andwood-boring beetle larvae (McAlpine et al. 1981).

Medetera vidua Wheeler

Host: P. fasciatus

This species has been reared only once fromPissodes (Deyrup 1978), but the record isconsidered confirmed. Specimens have beencollected from across Canada and the UnitedStates in coniferous forests. This species isprimarily associated with bark beetles (Bickel1985), and attack on Pissodes appears rare.

Hymenoptera

Hymenoptera associated with Pissodes areparasitoids or hyperparasitoids. Pissodes larvaeare attacked directly by ovipositing femaleparasitoids. The female parasitoid stings the hostlarva with the ovipositor to inject venom, whichimmobilizes the larva. Eggs are laid in, on, orvery near the host. There is no or minimal hostsearching by the parasitoid larva, and it remainsassociated with a single host larva. This pattern isalso the behavior of hyperparasitoids, whichattack the parasitoid species associated withPissodes. There are two types of parasitism:Idiobiont and Koinobiont. In idiobiosis, adultfemales attack later larval host instars or pupae.Immobilization of the host is protracted and maybe permanent or result in death. Idiobiont species

NOTES ON BIOLOGY AND ABUNDANCE OF REARED TAXA


are usually ectoparasitic, with eggs deposited onor near the host and parasitoid larvae feedingexternally through a rupture in the hostintegument. If the host regains mobility afterbeing stung, it quickly becomes moribund againas a result of parasitoid feeding. It is unlikely thathost larvae will resume normal development.Most Hymenoptera associated with Pissodesexemplify of this type of parasitism.

Koinobiosis requires a longer and more intimateassociation between parasitoid and host and isconsidered more specialized or derived thanidiobiosis. Adult females often attack earlier-instar larvae. Immobilization is temporary, andthe host usually returns to activity after attack, insome cases resuming feeding and growth for anextended period. Koinobiont species are usuallyendoparasitic, with eggs deposited inside thebody of the host. The parasitoid larva remainsinside the host for an extended period, eitherdeveloping slowly to allow the host to remainactive or feeding on nonessential organs such asthe fat body. The host may remain active untilparasitoid emergence is imminent and may not bekilled until it is ruptured by the mature parasitoidlarva. Most hyperparasitoids and someparasitoids are of this type.

Bracon nanus Provancher

Also as Microbracon nanus Provancher

Host: P. strobi

This species has been recorded as an uncommonidiobiont parasitoid of P. strobi larvae in theeastern United States (Harman and Kulman1967). Specimens of this species have also beencollected in Quebec, and it probably also occurs inOntario and the Maritimes. Although it has beenmisidentified as the more common Bracon pini(Muesebeck) (Harman and Kulman 1967), therecord is confirmed. Little is known about thebiology of B. nanus, but it is probably similar tothat of B. pini.

Bracon pini (Muesebeck)

Also as Microbracon pini Muesebeck

Hosts: P. fasciatus, Pissodes schwarzi Hopkins, P.strobi, P. terminalis

The biology of this common idiobiont species wasexamined by Deyrup (1978) and Taylor (1929). Itis recovered in most studies of Pissodes parasitoidsthroughout the United States and Canada(Langor 1998). It attacks middle- to late-instarhost larvae and occasionally pupae. Several eggsmay be laid per host, with one to five parasitoidlarvae per host feeding gregariously. Severallarvae can successfully mature from one hostbecause of the small size of the adult parasitoid.Most individuals overwinter as prepupae andemerge as adults in the spring. About one-thirdof individuals emerge the same year and mayattack new hosts. Note: Identification of Braconspecimens should not be considered definitive.This genus contains many species of commonparasitoids, described and undescribed.Confirmation of the identity of Bracon specimensshould be made by a specialist.

Calliephialtes comstockii (Cresson)

Also as Exeristes comstockii Cresson

Host: P. strobi

This species is a common idiobiont parasitoid ofnumerous species of shoot- and terminal-boringLepidoptera and Coleoptera across the UnitedStates and Canada (Yu 1999). Parasitism ofPissodes is rare but has been confirmed byStevenson (1963) and Williams and Langor(unpublished) in Alberta. References toparasitoid biology are numerous forlepidopterous hosts (Arthur 1963), and thisspecies has been the subject of experimentalresearch on host–parasite interactions (Thompsonand Barlow 1972). Several eggs may be laid perhost, but usually only one larva develops. Mostindividuals overwinter in terminals as prepupaeand complete development the following spring.


Coeloides pissodis (Ashmead)

Hosts: Pissodes nemorensis Germar, Pissodesstriatulus (Fabricius), P. strobi

This species is an uncommonly encounteredidiobiont parasitoid of Pissodes species and somebark beetles. It has been recovered several timesfrom Pissodes species in the eastern United Statesat low incidence, often below 1% (Taylor 1929;MacAloney 1930). Higher attack rates have beenobserved for some bark beetle hosts. Specimenshave been collected from eastern Canada as farwest as Manitoba, although this species has notbeen recovered from Canadian Pissodespopulations to date. Females attack maturelarvae and prepupae, laying one egg per host.Parasitoid prepupae overwinter in a cocooninside the host gallery, and there is one generationper year.

Coeloides rufovariegatus Provancher

As Coeloides dendroctoni Cushman

Host: P. strobi

This species is a common idiobiont parasitoid ofwood-boring Coleoptera throughout NorthAmerica. It is associated with various bark beetlespecies of economic importance, and it may be afactor in regulating host populations. Parasitismof Pissodes is rare but has been confirmed byStevenson (1967) and Williams and Langor(unpublished) in Alberta. Little is known aboutthe biology of this species in Pissodes hosts. It isprobably similar to that of C. pissodis, although theincidence of C. rufovariegatus is significantlyhigher in bark beetle hosts.

Coeloides secundus (Dalla Torre)

Host: P. striatulus

This species has been reared several times from firlogs containing Pissodes dubius Randall andPissodes fraseri Hopkins (synonyms of P. striatulus)in the eastern United States and Ontario, Quebec,and the Maritimes. Specimens have beencollected as far west as Michigan. Records arelimited to a list of specimens examined by Mason(1978) in his taxonomic treatment of the genus.This species is rarely collected, but the repeated

occurrences support its inclusion in the key. Littleis known about the biology of this species, but itis probably similar to that of other Coeloidesspecies.

Coeloides vancouverensis (Dalla Torre)

As Coeloides brunneri Viereck

Hosts: P. fasciatus, P. strobi

This species is a common idiobiont bark beetleparasitoid in the western United States, BritishColumbia, and Alberta. There have been someinvestigations of the biology of this speciesattacking bark beetles (Ryan and Rudinsky 1962[as C. brunneri]), for which it may be a factor inregulating host populations. Parasitism ofPissodes is rare but has been confirmed by rearingfrom individual P. fasciatus larvae (Deyrup 1978)and from terminals attacked by P. strobi(Stevenson 1967). In bark beetles, a single egg islaid per host, and parasitized host larvae arepermanently immobilized. Larval developmentis complete within 1 month, and adults mayemerge and begin another generation if suitablehosts are still available. Otherwise, prepupaeoverwinter in a cocoon inside the host gallery,completing development to pupae and adults thefollowing spring.

Cubocephalus occidentalis (Provancher)

As Phygadeuon nitidulus (Provancher)

Host: P. strobi

Little is known about the biology of this species.It is a rare primary parasitoid of P. strobi in theeastern United States (Taylor 1929; MacAloney1930; Mott 1930), and specimens have beencollected across Canada. The authors cited havelisted this species as a potential hyperparasitoidbecause other Cubocephalus species attackHymenoptera species, but these hosts are free-living, usually defoliators, not parasitic.Cubocephalus is a large genus, which attacks awide range of free-living and concealedColeoptera and Lepidoptera. The species forwhich biology is known are solitary koinobiontendoparasitoids.


Dolichomitus terrebrans nubilipennis (Ratzeburg)

Hosts: P. fasciatus, Pissodes rotundatus LeConte, P.striatulus, P. strobi, P. terminalis

This species is a common idiobiont parasitoid of awide range of bark- and shoot-boringLepidoptera and Coleoptera throughout thenorthern hemisphere, and there are numerousreferences to its taxonomy, biology, and hosts(Taylor 1929; Stevenson 1967; Alfaro et al. 1985;Langor and Williams 1998; Yu 1999). It is one ofthe largest of the Pissodes parasitoids, and femalesshow a preference for attacking fourth-instar hostlarvae, tending to delay attack until later thanother parasitoid species. When parasitizing P.strobi and P. terminalis, this species is one of fewthat successfully attack larvae that haveburrowed into the pith, which females can accesswith their very long ovipositors. A single egg islaid on the host larva. The mature larva spins adistinctive, semi-translucent cocoon, in which itoverwinters. This species can occur in largenumbers and may be a factor in regulating localhost populations.

Doryctes sp. or spp. (undetermined)

Hosts: P. schwarzi, P. strobi

This record is considered confirmed, although theidentity of this rare species is unknown, and it isnot certain that specimens reared from P. schwarziin California (Stevens 1966) represent the samespecies as those reared from P. strobi in NewEngland (Plummer and Pilsbury 1929;MacAloney 1930). The New England speciesprobably also occurs in Quebec and theMaritimes. It is clear from details in the recordsthat one or several species of Doryctes are solitaryidiobiont parasitoids of Pissodes species, althoughthese hosts were not recorded in a taxonomicstudy of the genus by Marsh (1969). There is littleinformation on the biology of this genus asidefrom fragmentary observations and host data onspecimen labels, but observations by Stevens(1966) indicate that incidence can be moderatelyhigh locally.

Eubazus calyptoides (Martin)

As Allodorus calyptoides Martin

Hosts: P. schwarzi, P. strobi, P. terminalis

We have repeatedly recovered this species as acommon parasitoid of P. strobi, P. terminalis(Langor and Williams 1998), and P. schwarzi in thePrairie provinces. Alfaro et al. (1985) mentionAllodorus sp. nr. crassigaster, which may be A.calyptoides, as a parasitoid of P. strobi in BritishColumbia. Specimens have also been collected inWisconsin. Nothing is known about the biologyof this species, but it is probably similar to that ofEubazus crassigaster (Provancher).

Eubazus crassigaster (Provancher)

As Allodorus crassigaster (Provancher), Eubazidonstrigitergum (Cushman)

Hosts: P. fasciatus, P. striatulus, P. strobi

This species is a common and importantkoinobiont parasitoid of Pissodes speciesthroughout North America (Stevenson 1967;Deyrup 1978; Alfaro et al. 1985) and has beeninvestigated as a potential biological control agentfor P. strobi (Hulme 1994). The genus Eubazusbelongs to a tribe of braconids that is noteworthyfor oviposition in host eggs. Parasitoid larvaeremain associated with the host for a protractedperiod after the egg hatches, allowing them to gothrough most of the larval period of feeding andgrowth with minimal interference. Eubazuscrassigaster apparently follows this habit, asDeyrup (1978) observed adult female parasitoidswaiting near ovipositing female weevils. Note:There is currently no key to species of Eubazus,and the two species discussed here are difficult toidentify. The identity of specimens of Eubazusshould be confirmed by an expert.

Eupelmus pini Taylor

Host: P. strobi

This species has occasionally been recovered inlow numbers in rearings of P. strobi in NewEngland (Taylor 1929; MacAloney 1930; Harman


and Kulman 1968) and may occur in Quebec andthe Maritimes. Little is known about its biologyexcept that mature larvae and pupae have beenrecovered from host chip cocoons. Ovipositionprobably occurs on middle-instar host larvae, asadults are active mainly in June. Most otherspecies of Eupelmus are endoparasitic, andevidence shows that host larvae remain active,continue to feed, and construct a chip cocoon forpupation after being parasitized by E. pini, whichindicates that this species is probably akoinobiont.

Eurytoma cleri Ashmead

Host: P. strobi

Hopkins (1899) collected specimens of this rarespecies directly from chip cocoons produced by P.strobi larvae in Virginia and West Virginia, and wehave examined specimens reared by Alfaro et al.(1985) from British Columbia, so the record isconsidered confirmed. Little is known about thebiology of this species, but it is probably similar tothat of Eurytoma pissodis (Girault).

Eurytoma picea Bugbee

Hosts: P. strobi, P. terminalis

This rare species has been collected fromterminals attacked by P. strobi in British Columbia(Bugbee 1967; Alfaro et al. 1985; several series ofspecimens the authors have examined) and P.terminalis in the western United States (Stevensand Knopf 1974). Little is known about thebiology of this species, but it is probably similar tothat of E. pissodis, and it may act as ahyperparasitoid of other Pissodes associates.

Eurytoma pissodis (Girault)

Hosts: P. strobi, P. terminalis, Ceoloidesvancouverensis, Coeloides sp.

This species is an abundant and commonidiobiont parasitoid of terminal-inhabitingPissodes that is recovered in most surveysthroughout North America (Taylor 1929;Stevenson 1967; Harman and Kulman 1968;Alfaro et al. 1985; Langor and Williams 1998).Adults of this species are active in July, andfemales prefer to attack host prepupae after

construction of the chip cocoon, laying one to fourdistinctive black eggs on the host. Only oneparasitoid larva per host survives, overwinteringas a prepupa and completing pupation and adultemergence the following spring. Eurytomapissodis apparently attacks any species present inchip cocoons, acting as a hyperparasitoid on otherspecies of parasitoids that have killed the hostlarva. Note: Species of the genus Eurytoma arevery similar to one another in appearance. Eventhough a key to species is available (Bugbee 1967),identification is difficult and should be confirmedby an expert.

Helcostizus albator rufiscutum Cushman

Host: P. strobi

Little is known about the biology of this rarespecies. The genus Helcostizus belongs to a tribe ofIchneumonidae for which there is little host data.Those species of the tribe for which hosts areknown are parasitoids of the prepupae and pupaeof Coleoptera larvae. Specimens have beenreared from conifer boles or collected inCalifornia and Arizona. Stevenson (1967) rearedthis species from individual hosts in BritishColumbia, a record that is considered confirmed,but provided no other details about life cycle.

Helcostizus contortae Townes

Host: P. terminalis

See comments for H. a. rufiscutum, above.Stevenson and Petty (1968) and Langor andWilliams (1998) have reared small numbers of thisspecies from lodgepole pine tips containing P.terminalis larvae from several different localities inAlberta. Townes (1983) also examined specimensof this species reared from lodgepole pinecontaining P. terminalis in British Columbia andOregon.

Helcostizus subrectus Townes

Host: P. terminalis

See comments for H. a. rufiscutum, above. Stevensand Knopf (1974) recovered this species in severallocations in the western United States, and a fewspecimens have been collected from Arizona toAlberta.


Heydenia unica Cook and Davis

Host: P. strobi

This species is a common idiobiont parasitoid ofnumerous species of bark beetles. It has beenrecovered from conifers or collected throughoutthe United States and probably occurs in southernCanada, at least in British Columbia and Ontario.Parasitism of Pissodes is rare (Taylor 1929) but hasbeen confirmed by Hopkins (1899), who obtainedspecimens from P. strobi chip cocoons. There havebeen some investigations of the biology of thisspecies attacking bark beetles (Dix and Franklin1983), but little is known about attack on P. strobi.This species prefers to attack late-instar larvaeand frequently attacks pupae. A single egg is laidper host, and the parasitoid larvae overwinter,completing development the following year.

Labena grallator Say

Also as Labena apicalis Cresson

Host: P. strobi

This species is a common parasitoid of a widerange of wood-boring Coleoptera, primarilyCerambycidae, in the southern and easternUnited States. It probably also occurs in Ontarioand Quebec. Little is known about the biology ofthis species, except that it is rare in Pissodes(Barnes 1928; Taylor 1929; Harman and Kulman1968) and seems to have low incidence on allknown hosts.

Spathius brachyurus Ashmead

Hosts: P. nemorensis, P. strobi

This rare idiobiont parasitoid species has beenrecovered several times in studies of Pissodes inthe eastern United States, at low incidence(Britton 1920; Peirson 1922). Specimens have alsobeen collected in Quebec, and this species isprobably also present in Ontario and theMaritimes. Parasitoids attack prepupae, usuallylaying several eggs, with the number dependingon the size of the host. Larvae feed gregariously,and several can successfully mature from one hostbecause of the small size of the adult parasitoid.Matthews (1970) recorded as many as seven S.brachyurus cocoons associated with the remains of

a single host larva. Parasitoids overwinter asprepupae in cocoons spun in the host gallery.Adults emerge in late spring and are activethroughout the summer. There may be severalgenerations per year if suitable hosts areavailable, either Pissodes species or several speciesof bark beetles that are also attacked by thisparasitoid.

Spathius sequoiae Ashmead

Host: Pissodes sp.

This idiobiont parasitoid species attacks primarilybark beetles in the western United States, andparasitism of Pissodes is rare (Matthews 1970).Specimens have also been collected from Alaska,British Columbia, and Alberta, and it probablyoccurs in the Yukon and Northwest Territories aswell. There are few records of attack on Pissodes,and available specimens have been reared fromhost larvae in boles. The biology of this speciesattacking bark beetles resembles that of S.brachyurus, although it is not known if the larvaefeed gregariously.

Rhaphitelus maculatus Walker

Hosts: P. strobi, Pissodes sp.

This species is a common and abundant idiobiontparasitoid of bark beetles and other wood-boringColeoptera occurring worldwide (Yu 1999).Parasitism of Pissodes is rare, although it can belocally common in the Prairie provinces (Williamsand Langor, unpublished). There has been someinvestigation of the biology of this speciesattacking bark beetles (Campos and Gonzalez1993), but little is known about attack on P. strobi.This species is flexible in both host range andbehavior. It attacks both prepupae and pupae andwill facultatively act as a hyperparasitoid on otherP. strobi parasitoids, presumably those occurringin chip cocoons. Adults are active in July, andfemales will deposit one or several eggs on a host.A single parasitoid larva survives per host,overwintering as a prepupa. Development iscompleted the following year.


Rhopalicus pulchripennis (Crawford)

Hosts: P. strobi, P. terminalis, Coeloidesrufovariegatus

This species is a common and abundant idiobiontparasitoid of Pissodes, bark beetles, and otherwood-boring Coleoptera throughout NorthAmerica (Plummer and Pilsbury 1929; Taylor1929; Stevenson 1967; Harman and Kulman 1968;Langor and Williams 1998). Adult females attackmiddle- to late-instar host larvae and may havemore than one generation per year. In areaswhere studies have been conducted, there are twoperiods of adult activity, in early June and lateJuly. Females that have parasitized middle-instarhost larvae early in the season may emerge ontime to attack later instars. A single egg isdeposited per host, and the larva, if itoverwinters, does so as a prepupa. This specieswill facultatively act as a hyperparasitoid on otherP. strobi parasitoids.

Rhopalicus tutela (Walker)

Also as Rhopalicus tutela suspensus (Ratzeburg)

Host: P. strobi

This species is a common and abundant idiobiontparasitoid of bark beetles and other wood-boringColeoptera worldwide. Parasitism of Europeanspecies of Pissodes is common, but uncommon inP. strobi, although the parasitoid has been

recovered several times (Britton 1920; Plummerand Pilsbury 1929). Some information is availableon the biology of this species attacking barkbeetles (Berisford et al. 1970), but little is knownabout attack on P. strobi. Adults attack middle- tolate-instar bark beetle larvae and are active overmuch of the season, and there might be more thanone generation per year. A single egg is laid perhost, and the larva, if it overwinters, does so as aprepupa. In studies of parasitism of Ips, larvaeand pupae of this species have been foundseparate from host remains, having moved fromone part of the host gallery to another. It ispossible that individuals of this species attackmore than one host, facultatively acting aspredators.

Trigonura tarsata (Dalla Torre)

Host: P. strobi

Little is known about the biology of this rarespecies. Specimens have been collected from theeastern United States as far west as North Dakota,and this species is probably also present inOntario and Quebec. It was reared by Barnes(1928) and Taylor (1929), and these records areconsidered confirmed, although this species hasnot been recovered since. Other hosts includeseveral species of the genus Magdalis, which arebark-boring weevils with similar biology to thatof Pissodes.


Alfaro, R.I.; Borden, J.H. 1980. Predation by Lonchaea corticis(Diptera: Lonchaeidae) on white pine weevil, Pissodesstrobi (Coleoptera: Curculionidae). Can. Entomol.112:1259–1270.

Alfaro, R.I.; Hulme, M.A.; Evans, J.W.E. 1985. Insects associatedwith the Sitka spruce weevil, Pissodes strobi (Col:Curculionidae) on Sitka spruce in British Columbia,Canada. Entomophaga 30:415–418.

Alfaro, R.I.; Kiss, G.; Fraser, R.G., eds. 1994. The white pineweevil: biology, damage, and management. Proceedings ofa Meeting, Richmond, British Columbia, 19S21 January,1994. Nat. Resour. Can., Can. For. Serv., Pac. For. Cent.,Victoria, BC, and BC Minist. For., Victoria, BC, Can–BCPartnership Agreement For. Dev., For. Resour. Dev.Agreement II Rep. 226.

Arthur, A.P. 1963. Life histories and immature stages of fourichneumonid parasites of the European pine shoot moth,Rhyacionia buoliana (Schiff.) in Ontario. Can. Entomol.95:1078–1091.

Barnes, T.C. 1928. A biological study of the white pine weevilwith special reference to anatomy, flight, phenology,parasitism, behaviour, and injury to young trees. D.Sc.thesis, Harvard Univ., Cambridge, MA.

Berisford, C.W.; Kulman, H.M.; Pienkowski, R.L. 1970. Notes onthe biologies of hymenopterous parasites of Ips spp. barkbeetles in Virginia. Can. Entomol. 102:484–490.

Bickel, D.J. 1985. A revision of Nearctic Medetera (Diptera:Dolichopodidae). US Dep. Agric., Agric. Res. Serv.,Springfield VA, Tech. Bull. 1692.

Britton, W.E. 1920. The white pine weevil, Pissodes strobi Peck.Pages 144–155 in 19th report of the state entomologist ofConnecticut for the year 1919. Conn. Agric. Exp. Stn., NewHaven, CT. Bull. 218.

Bugbee, R.E. 1967. Revision of chalcid wasps of the genusEurytoma in America north of Mexico. Proc. US Natl. Mus.118:443–552.

Campos, M.; Gonzales, R. 1993. Effect of parent density onfecundity of two parasitoids (Hymenoptera:Pteromalidae) of the olive beetle, Phloeotribus scarabaeoides(Coleoptera: Scolytidae). Entomophaga 36:473–480.

Dasch, C.E. 1979. Ichneumon-flies of America north of Mexico:8. Subfamily Cremastinae. Mem. Am. Entomol. Inst. 29.

Deyrup, M.A. 1978. Notes on the biology of Pissodes fasciatusLeConte and its insect associates (Coleoptera:Curculionidae). Pan-Pac. Entomol. 54:103–106.

Dix, M.E.; Franklin, R.J. 1983. Behaviour of four braconidparasites and one pteromalid parasite of the southern pinebeetle, Dendroctonus frontalis. J. Ga. Entomol.Soc.18:125–138.

Gibson, G.A.P.; Huber J.T.; Woolley, J.B., eds. 1997. Annotatedkeys to the genera of Nearctic Chalcidoidea(Hymenoptera). Natl. Res. Counc. Res. Press, Ottawa, ON.

Goulet, H.; Huber, J.T., eds. 1993. Hymenoptera of the world: anidentification guide to families. Agric. Can., Res. Branch,Ottawa, ON. Publ. 1894/E.

Harman, D.M. 1966. Biology and control of the white pineweevil, Pissodes strobi (Peck), in Virginia. PhD thesis,Virginia Polytech. Inst., Blacksburg, VA.

Harman, D.M.; Kulman, H.M. 1967. An annotated list: parasitesand predators of the white pine weevil, Pissodes strobi(Peck). Univ. Maryland, Natl. Res. Inst., College Park, MD.Contrib. 323.

Harman, D.H.; Kulman, H.M. 1968. Biology and control of thewhite-pine weevil in Virginia. Ann. Entomol. Soc. Am.61:280–285.

Hiratsuka, Y.; Langor, D.W.; Crane, P.E. 1995. A field guide toforest insects and diseases of the prairie provinces. Nat.Resour. Can., Can. For. Serv., Northwest Reg., North. For.Cent., Edmonton, AB, Spec. Rep. 3.

Hopkins, A.D. 1899. Report on investigations to determine thecause of unhealthy conditions of the spruce and pine from1800 to 1893. WVa. Agric. Exp. Stn., Morgantown, WVa.Bull. 56.

Hulme, M.A. 1989. Laboratory assessment of predation by Lonchaeacorticis (Diptera: Lonchaeidae) on Pissodes strobi (Coleoptera:Curculionidae). Environ. Entomol. 18:1011–1014.

Hulme, M.A. 1994. The potential of Allodorus crassigaster for thebiological control of Pissodes strobi. Pages 294–300 in R.I.Alfaro, G. Kiss, and R.G. Fraser, eds. The white pineweevil: biology, damage, and management. Proceeding ofa Meeting, Richmond, British Columbia, 19–21 January,1994. Nat. Resour. Can., Can. For. Serv., Pac. For Cent.,Victoria, BC, and BC Minist. For., Victoria, BC, Can–B.C.Partnership Agreement For. Dev, For. Resour. Dev.Agreement II Rep. 226.

REFERENCES


Krombein, K.V.; Hurd, P.D.; Smith, D.R.; Burks, B.D., eds. 1979.Catalog of Hymenoptera in America north of Mexico.Volume 1. Symphyta and Apocrita. Smithsonian Inst.Press, Washington, DC.

Langor, D.W. 1998. Annotated bibliography of North andCentral American species of bark weevils, Pissodes(Coleoptera: Curculionidae). Nat. Resour. Can., Can. For.Serv., North. For Cent., Edmonton, AB. Inf. Rep. NOR-X-355.

Langor, D.W.; Williams, D.J.M. 1998. Life cycle and mortality ofPissodes terminalis (Coleoptera: Curculionidae) inlodgepole pine. Can. Entomol. 130:387–397.

MacAloney, H.J. 1930. The white pine weevil (Pissodes strobiPeck)—its biology and control. Bull. N. Y. State Coll. For.Syracuse Univ. 3:1–87.

Marsh, P.M. 1969. The Nearctic Doryctinae, VII. The genusDoryctes Haliday (Hymenoptera: Braconidae). Trans. Am.Entomol. Soc. 94:379–405.

Martin, J.C. 1956. A taxonomic revision of the triaspidinebraconid wasps of Nearctic America (Hymenoptera). Can.Dep. Agric., Ottawa, ON. Publ. 965.

Mason, W.R.M. 1978. A synopsis of the Nearctic Braconini, withrevisions of Nearctic species of Coeloides and Myosoma(Hymenoptera: Braconidae). Can. Entomol. 110:721–768.

Matthews, R.W. 1970. A revision of the genus Spathius inAmerica north of Mexico (Hymenoptera: Braconidae).Contrib. Am. Entomol. Soc. 4:1–86.

McAlpine J.F.; Peterson, B.V.; Shewell, G.E.; Teskey, H.J.;Vockeroth, J.R.; Wood. D.M., eds. 1981. Manual of NearcticDiptera. Part 1. Agric. Can., Res. Branch, Ottawa, ON.Monogr. 27.

Mott, P.B. 1930. An annotated bibliography of the white pineweevil, Pissodes strobi (Peck), for white pine blister rustworkers and others. N.J. Dep. Agric., Trenton, NJ. Circ.177.

Nealis, V.G. 1998. Population dynamics of the white pineweevil, Pissodes strobi, infesting jack pine, Pinus banksiana,in Ontario, Canada. Ecol. Entomol. 23:305–313.

Noyes, J.S. 1998. Catalogue of the Chalcidoidea of the world.UNESCO Publishing and Expert Center for TaxonomicIdentification, Paris, France.

Peirson, H.B. 1922. Control of the white pine weevil by forestmanagement. Harvard For., Petersham, MA. Bull. 5.

Plummer, C.C.; Pilsbury, A.E. 1929. The white pine weevil inNew Hampshire. Univ. of New Hampshire Exp. Stn.,Durham, NH. Bull. 247.

Robertson, J.L. 1979. Rearing the western spruce budworm. USDep. Agric., For. Serv., and Canada/United States SpruceBudworm Program, Washington, DC.

Ryan, R.B.; Rudinsky, J.A. 1962. Biology and habits of theDouglas-fir beetle parasite, Coeloides brunneri Viereck(Hymenoptera: Braconidae) in western Oregon. Can.Entomol. 94:748–763.

Stark, R.W.; Wood, D.L. 1964. The biology of Pissodes terminalisHopping (Coleoptera: Curculionidae) in California. Can.Entomol. 96:1208–1218.

Stevens, R.E. 1966. Observations on the Yosemite bark weevil inCalifornia. Pan-Pac. Entomol. 42:184–189.

Stevens, R.E.; Knopf, J.A.E. 1974. Lodgepole terminal weevil ininterior lodgepole forests. Environ. Entomol. 3:998–1002.

Stevenson, R.E. 1963. Insects associated with the Engelmannspruce weevil, Pissodes engelmanni Hopkins. Can. Dep.For., For. Entomol. Pathol. Branch, Ottawa, ON. Bi-mon.Prog. Rep. 19:2–3.

Stevenson, R.E. 1967. Notes on the biology of the Engelmannspruce weevil, Pissodes engelmanni (Curculionidae:Coleoptera) and its parasites and predators. Can. Entomol.99:201–213.

Stevenson, R.E.; Petty, J.J. 1968. Lodgepole terminal weevil(Pissodes terminalis Hopping) in the Alberta/NorthwestTerritories region. Can. Dep. For. Rural Dev., For. Branch,Ottawa, ON. Bi-mon. Res. Notes 24:6.

Taylor, R.L. 1929. The biology of the white pine weevil, Pissodesstrobi (Peck), and a study of its insect parasites from aneconomic viewpoint. Entomol. Am. 10:1–86.

Thompson, S.N.; Barlow, J.S. 1972. Synthesis of fatty acids by theparasite Exeristes comstockii (Hymenoptera) and two hosts,Galleria mellonella (Lepidoptera) and Lucillia sericata(Diptera). Can. J. Zool. 50:1105–1110.

Townes, H. 1969a. Genera of Ichneumonidae, Part 1. Mem. Am.Entomol. Inst. 11.

Townes, H. 1969b. Genera of Ichneumonidae, Part 2. Mem. Am.Entomol. Inst. 12.

Townes, H. 1969c. Genera of Ichneumonidae, Part 3. Mem. Am.Entomol. Inst. 13.


Townes, H. 1971. Genera of Ichneumonidae, Part 4. Mem. Am.Entomol. Inst. 17.

Townes, H. 1983. Revisions of twenty genera of Gelini(Ichneumonidae). Mem. Am. Entomol. Inst. 35.

Wharton, R.A.; Marsh, P.M.; Sharkey, M.J., eds. 1997. Manual ofnew world genera of the family Braconidae(Hymenoptera). Int. Soc. Hymenopterists, Washington,DC. Spec. Publ. 1

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Yu, D.S. 1999. Interactive catalogue of world Ichneumonidae1998 [CD-ROM]. Taxapad: scientific names for informationmanagement, Natural History Musuem, London, UK.


Family Species Host Rolea Distribution

Diptera

Lonchaeidae Lonchaea corticis Taylor P. strobi Pred Transcontinental

Lonchaeidae Lonchaea furnissi McAlpine P. fasciatus Pred Western North America

Dolichopodidae Medetera apicalis (Malloch) P. fasciatus Pred Western North America

Dolichopodidae Medetera vidua Wheeler P. fasciatus Pred Western North America

Hymenoptera

Braconidae Bracon nanus Provancher P. strobi Par Eastern North America

Braconidae Bracon pini (Muesebeck) P. fasciatus, Par TranscontinentalP. schwarzi,P. strobi,P. terminalis

Braconidae Coeloides pissodis (Ashmead) P. nemorensis, Par TranscontinentalP. striatulus,P. strobi

Braconidae Coeloides rufovariegatus P. strobi Par TranscontinentalProvancher

Braconidae Coeloides secundus P. striatulus Par Transcontinental(Dalla Torre)

Braconidae Coeloides vancouverensis P. fasciatus, Par Western North America(Dalla Torre) P. strobi

Braconidae Doryctes sp. P. schwarzi, Par TranscontinentalP. strobi

Braconidae Eubazus calyptoides (Martin) P. schwarzi, Par Western North AmericaP. strobi,P. terminalis

Braconidae Eubazus crassigaster P. fasciatus, Par Transcontinental(Provancher) P. striatulus,

P. strobi

Braconidae Spathius brachyurus P. nemorensis, Par Eastern North AmericaAshmead P. strobi

Braconidae Spathius sequoiae Ashmead Pissodes sp. Par Western North America

Chalcididae Trigonura tarsata (Dalla Torre) P. strobi Par Northeastern NorthAmerica

Eupelmidae Eupelmus pini Taylor P. strobi Par Eastern U.S.A.

APPENDIX 1

PARASITOIDS, HYPERPARASITOIDS, AND PREFATORSASSOCIATED WITH PISSODES SPECIES IN CANADA


Family Species Host Rolea Distribution

Eurytomidae Eurytoma cleri Ashmead P. strobi Par TranscontinentalHyp

Eurytomidae Eurytoma picea Bugbee P. strobi, Par British ColumbiaP. terminalis Hyp

Eurytomidae Eurytoma pissodis (Girault) P. strobi, Par TranscontinentalP. terminalis Hyp

Ichneumonidae Calliephialtes comstockii P. strobi Par Transcontinental(Cresson)

Ichneumonidae Cubocephalus occidentalis P. strobi Par Transcontinental(Provancher) Hyp

Ichneumonidae Dolichomitus terrebrans P. fasciatus, Par Transcontinentalnubilipennis (Ratzeburg) P. fasciatus,

P. rotundatus,P. striatulus,P. strobi,P. terminalis

Ichneumonidae Helcostizus albator P. strobi Par Western North Americarufiscutum Cushman

Ichneumonidae Helcostizus contortae Townes P. terminalis Par Western North America

Ichneumonidae Helcostizus subrectus Townes P. terminalis Par Western North America

Ichneumonidae Labena grallator Say P. strobi Par Transcontinental

Pteromalidae Heydenia unica P. strobi Par TranscontinentalCook and Davis

Pteromalidae Rhaphitelus maculatus Walker P. strobi, Par TranscontinentalPissodes sp. Hyp

Pteromalidae Rhopalicus pulchripennis P. strobi, Par Transcontinental(Crawford) P. terminalis Hyp

Pteromalidae Rhopalicus tutela (Walker) P. strobi Par Holarctic (EasternPred North America)

aPred = predator, Par = parasitoid, Hyp = hyperparasitoid. Species with more than one designation may take either role or both.


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Abdomen (see also Gaster): The most posteriorbody section of an insect, containing digestiveand reproductive organs in adult insects (Fig.63).

Antenna: A sensory structure on the front of thehead, which is thread-like on Hymenoptera(Fig. 62A), and club-like on Diptera (Fig. 63).

Anterior: Pertaining to the front or headwardedge or surface of a body section orappendage.

Areolet: The first radial sector cell in the forewingof Ichneumonidae adjacent to the large 2R1cell (Fig. 62A).

Carapace: A hard dorsal shell formed by thefusion of several abdominal tergites, whichcovers and protects the gaster of someHymenoptera.

Carina (pl. carinae): A ridged type of sculpture.Carina length and placement can be highlyvariable among species.

Cell: An area of clear wing membrane that issurrounded on all sides by pigmented veinsor the wing margin (Fig. 62A), or an area ofsclerite surrounded and defined by carinae.

Chip cocoon: A structure prepared by Pissodeslarvae from wood shavings taken from thehost tree in which they pupate.

Cocoon: A construction of silk spun by the larvaeof some insects, in which they pupate.Parasitic Hymenoptera larvae emerge fromthe host and spin white, oblong cocoonscovered with loose fibers. Some spin cocoonsinside the host, which are not visible unlessthe host cadaver is dissected.

Coxa: The first section of the leg, attached directlyto the thorax (Figs. 62A, 63).

Diapause: A period in the life cycle of an insectduring which development is temporarilysuspended.

Dorsal: Pertaining to the upper surface of a bodysection or appendage.

Ectoparasitic: Pertaining to a parasitoid larva thatlives outside the host’s body, but feeds on thehost’s internal tissues and body fluids.

Endoparasitic: Pertaining to a parasitoid larvathat lives entirely within the host’s body.

Eye: The large, multifaceted visual organs oneither side of an adult insect’s head (Figs.62A, 63).

Femur (pl. femora): The third section of the leg,between the trochanter and the tibia (Figs.62A, 63).

Flagellum: The third section of the antenna ofHymenoptera, comprising the third to endsegments, which resemble one another in sizeand shape and are different from the first twosegments (see Scape and Pedicel).

Forewing: The larger, anterior pair of wings inHymenoptera (Fig. 62A), or the only visiblepair of wings (commonly referred to assimply the wings) in Diptera (Fig. 63).

Gaster (see also Abdomen): The most posteriorbody section of Hymenoptera, which consistsof all abdominal segments except the first.The first abdominal segment is fused to thethorax and forms the propodeum (Fig. 62A).

Granular: A pebbled type of microsculpture thatimparts a matte surface to a sclerite.

Generation: The time required to complete onefull life cycle, including all stages.

Habitus: A drawing or photograph intended todisplay the general shape and feature of aspecies, upon which specific morphologicalcharacters may be highlighted.

Hair: Filaments that occur all over the bodies offlies and come in a variety of sizes. Setae areenlarged hairs, but for this purpose the term

APPENDIX 3

GLOSSARY


hair is used to refer only to the smallest sizeclass.

Head: The most anterior body section of an insect,containing sensory organs and mouthparts inadult insects (Figs. 62A, 63).

Hind wing: The smaller, posterior pair of wingsin Hymenoptera (Fig. 62A). The hind wingsin Diptera are reduced to small clubs and noteasily visible (Fig. 63).

Hyperparasitoid: A parasitoid that attacksanother parasitoid inside a host, not the hostitself.

Idiobiont (Idiobiosis): A type of parasitism inwhich later host stages are parasitized, hostactivity is arrested following attack, andassociation of the host and parasitoid isrelatively brief (see Koinobiont).

Instar: A stage in an insect’s development.

Integument: The hard outer skeleton of an insect.

Koinobiont (Koinobiosis): A type of parasitismin which earlier host stages are parasitized,host activity continues following attack, andassociation of the host and parasitoid isprolonged (see Idiobiont).

Larva: An immature stage in the life cycle of aninsect.

Lateral: Pertaining to the side surfaces of a bodysection or appendage.

Mandible: The chewing mouthpart of manyinsects (Fig. 62A).

Mesopleuron: A sclerite of the side of the thoraxof Hymenoptera.

Microsculpture: Very fine surface texture of asclerite that is visible only under very highmagnification.

Multivoltine: Having several generations peryear.

Occelli: A group of light-sensing organs on thetop of the head of some insects (Fig. 63).

Oviposition mark: A small puncture in the barkof a terminal of a host tree left by an egg-laying Pissodes female.

Ovipositor: The egg-laying structure visible at theapex of the gaster in Hymenoptera. It isusually covered by a hairy sheath (Fig 62B).

Parasitoid: An insect that feeds on the internaltissues and body fluids of another insect,eventually killing the host.

Parastigma: An enlarged bulb on the end of thestigmal vein in some Hymenoptera.

Pedicel: The second segment of the antenna,which is often small, cylindrical, anddrumlike, between the scape and theflagellum.

Posterior: Pertaining to the rear surface of a bodysection or appendage.

Predator: A free-living insect that attacks andconsumes another or several other insects.

Prepupa (pl. prepupae): A stage of the last larvalinstar, just before molting to the pupa.

Propodeum: The first segment of the abdomen ofHymenoptera, which is fused to the posteriorend of the thorax and which appears to be athoracic sclerite (Fig. 62A).

Punctate: Referring to a part of the integumentwith distinct pits or circular impressions,often with a hair attached in the middle.

Pupa (pl. pupae): A stage (usually quiescent) inthe development of an insect during whichlarval structures are lost and adult structurescreated.

Puparium (pl. puparia): An oblong, cylindricalstructure composed of the skin of the maturelarva of flies, which hardens and turns brown.The pupa is formed inside the puparium, andthe adult fly emerges from this structure.

Scape: The first segment of the antenna, whichattaches directly to the head and which isusually larger and more rounded than theremaining segments.


Sclerite: A distinct section of the body integumentthat is distinguished from other such sectionsby a ridge or groove.

Sculpture: Fine surface structures on theintegument. These may be fine or coarsewrinkles, pebbling, pits or punctures, largeridges, or other features that otherwise alterthe smooth, shining surface.

Seta (pl. setae): Very large, dark hairs that areoften arranged in specific patterns.

Striate (pl. striae): A type of sculpture whichcomprises numerous parallel fine wrinkles.

Tarsus: The fifth and end-most section of the leg,which is attached to the tibia and whichcomprises four or five segments calledtarsomeres (Figs. 62A, 63).

Tegula: A flap-like sclerite in Hymenoptera thatcovers the base of the wing where it attachesto the thorax (Fig. 62A).

Tergum (pl. terga or tergites): The dorsal scleritesof body segments. The posterior edge of atergum is also called the apex or the margin,and the anterior edge is called the base.

Terminal: The topmost growing point of the mainstem of a conifer tree.

Thorax: The middle body section of adult insectswhere the wings and legs are attached. InHymenoptera the apparent thorax iscomposed of the true thorax plus the firstabdominal segment (propodeum) (Fig. 62A).In Diptera the thorax is a true thorax (Fig. 63).

Tibia (pl. tibiae): The fourth section of the leg,between the femur and the tarsus (Figs. 62A,63).

Trochanter: The second section of the leg,between the coxa and femur (Fig. 62A).

Tubercle: A swelling or bump on the surface of asclerite.

Univoltine: Having only one generation per year.

Vein: The thickened, pigmented lines of thewings, which provide a supporting frame forthe thin, membraneous remainder. Thearrangement of veins and the cells theysurround may be distinct among species(Figs. 62A, 63).

Ventral: Pertaining to the lower surface of a bodysection or appendage.

rearing, identification and biology of parasitoids and predators

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