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, , � , " , I ( t . ,f ,f " "',X1) \� I ,,1 ":; �.', A/ , ,� \ '\. I'") ,-.'\ " 1 '''{ I \ \ , t· J ::r' I'< /' '!� f 5(;' ", \ ).---- "I ... , "--':-/. , /;;' \';.;_:�;�,:� ;';<r:' >f UNIVERSITY OF SASKATCHEWAN This volume is the property of the University of Saskatchewan, and the literary rights of the author and of the University must he respected. If the reader oh tains any assistance from this volume, he must give proper credit in his own work. This Thesis by .. EIJ.,IS.McMILLA.N : . has heen used hy the following persons, whose signatures attest their accept ance of the ahove restrictions. Narne and Address Date ,.� A THESIS ent itled A PR3LIMINARY STUDY OF THE POLYEMBRYOUIC CUTWORM PARASITE BERECYNTUS BAKERI VAR. GEMMA GIRAULT BY Ellis McMillan presented April, 1930. to the Faculty of Arts and Science of the University of SasKatchewan, in part ful filment of the requirements for the degree of Master of Science. FOR E W 0 R D TOT H � THE SIS The field and laboratory worK involved in this study has been performed by the writer during the period 1927-1930 as a part of the investigational worK pertaining to the duties of Assistant Entomologist, Canada Department of Agriculture, and is used here by permission of the Dominion Entomologist, Mr. Arthur Gibson. The development of the full plan of the study, the worK in its entirety (except for certain routine phases), the assembling of data and the conclusions brought out, should all be credited to the writer. Ac Knowledgments in full are given of any assistance which has been received. In the following presentation, both on account of convenience and because of the large amount of data in volved, the worK has been divided into four principal sections. The first deals with the biology of the red baCKed cutworm (Euxoa ochrogaster), the chief host of the paraSite being studied, and is merely a summarized statement of previous worK done by K. :M. King and N. J. AtKinson. The second section deals with the systematic study of the paraSite consisting mainly of a treatment of larval morphology. The biology of the parasite in all its aspects is fully treated in the third section. while the fourth part outlines the pre- -�- liminar,y studies on the polyembryonic development of the parasite. Finally, credit for the original conception of the problem goes entirely to Mr. King, as also does some of the subsequent , supervision. -------...,_....".,......",...".......,,=---...=.��-�--- ACKITOWLEDGMEUTS An investigation of the nature of the present study, in parts so detailed and so intricate, requires the assistance of others who are fully aware of the dangers and pitfalls to be met with in polyembryony. It is with great pleasure, therefore, that the writer ac�nowledges his indebtednessto the following, from whom material help has been received. Chief among. those to whom the author wishes to ex press his gratitude in the'pre�ent �ork is Mr. K. M. King, Federal Entomologist for Sas�atchewan; to him belongs the initial conception of the problem, his permission and re commendation were necessary for the carrying on of part of the wor� in Branch hours and for the use of some mat erial presented herewith. Of equally great assistance was his continued interest in the problem, manifested by kindly criticism and many useful suggestions. To Dr. L. G. Saunders, under whose supervision the study has been conducted, I wish to express my apprecia tion of his help and consideration. It has been the �indness of Dr. W. P. Thompson and Dr. Saunders which has made possible the presentation of this work as a thesis. From H. G. Crawford, Entomologist in Charge of Field Crops and Garden Insects, Canada Department of Agriculture permission was obtained, upon Mr. King's recommendation, ---- _. -_. -----=��==""""=��=-==-��-- to use in the present thesis data and material obtained by 1fr. King and the author as officers of t:qe Entomological Branch, and to him the author wishes to express his sincere t hanks , The author has been fortunate in obtaining valuable advice from specialists in insect polyembryony. To Dr. H. L. Parker, European Parasite Laboratory, United States Bureau of Entomology, my most sincere thanKs are given for the advice received concerning larval morphology among the Chalcididae. From time to time,during my re search, considerable help in interpreting the polyembryonw development of the parasite,has been received from Dr. R. W. Leiby, North Carolina Department of Agriyulture, and I am exceedingly grateful for his sympathetic interest in my difficulties. To Dr. Patterson, University of Texas, grateful thanKs are due; this specialist in polyembryony brought to my attention relevant literature which proved invaluable. For laboratory facilities and supplies I would expre� my indebtedness to the Entomological Branch and to the Department of Biology of the University of Sask.atchev,-an. Messrs. A. Arnason, R. Glen and L. Paul, fellow officers at the Sask.atoon Laboratory, have materially helped at various times in the routine phases of the field and laboratory work.. Their friendly interest in the pro blem is also sincerely appreciated. -�- �- �- - --...;;:.,,::;,- ;.....__ __;..._��===--_�=_=_O'"........._=__------ T _\.B2::,E 0 Fe\) UT �UT S LIST OF FIS-TIRES I IUTRODUCT Io:r 1 II BIOIOr'jy OF THE CHIEF EOST. :::t1!O_\. OCHROGASTER ( SUI��L\';ty ) 3 II I SYSTEMAT IC 11 IV IITORPHOI.OGY 19 1. The egg 19 o ..... The La rva (a) Larval types in the chalcid oidea. 21 (b) External anatonzy 28 (c) Internal anatomy 29 3. Prepupa 32 4. Pupa 32 V BIOLOGY OF THE PARASITE 1. Emergence of the adults 35 2. Longevity of the adults, their habits and reactions. 39 3. potential progenitiveness ,45 4. Copulation 47 5. Oviposition 49 6. Parthenogenesis 55 7. Appearance of parasitized e�gs 60 8. Feeding of parasitized larvae and non-parasitized larvae. 61 9. Comnarison of the amount of time spent Lu the different stages of development by no rmal and para- sit ized larvae. 66 10. Hosts. 69 11. Super-parasitism of Euxoa ochrogaster. 72 1·::>..... Interrelationship of dise�se and parasitism by Berecyntus. 75 13. The general effects of parasitism on the hast and on the parasite. 77 14. The of food with reuro- relationship � ductive activity and longevity. 82 15. The relationship of abund�nce of the parasite to weather conditions. 86 16. Seasonal history. 107 17. Percentages of parasitism and economic importance. 116 VI PO 1YEliIB�YO BY 1. General considerations on polyernbryony132 ( a) thea ries 132 (b) polyembryony in animals 136 2. Historical review of literature on polyembryony, chiefly insect poly- embryony. 144 3. No t e s on the host and parasite 154 4. Development of the egg (fall period) 156 (a) preoviposition 156 (b) ovipoeitio� to cleavage 157 (c) egg after oviposi�ion 158 ( d) d e ve Lo pine rrt in the host eg;s 162 1. maturation and fertiliz- ation. 163 Z. cleavage to overwinter- ing stage. 164 (e) EuffiLaryand discussion of the development' of the para.site egg to the overw int er ing st age. 168 5. Develo:pment of pazas Lt e body in host larva (spring and e arly euumer ) 169 (a) development of the parasite body after �tching of the host but previous to any indication of polyembryony. 170 1 • po 1ygerma 1 rlJ9..SS 17�� 2. f 0 rrna t Lo n 0 f po1ygermal mass 173 (b)' first indications of polyembry- ony. 1. polyembryonal mass 175 2. the Lnt e r etub ryo na L matrix 176 (c) pseudogerms, pseudomoru1as� pseudo embryos and pseudolarvae 178 (d) .independant life of par�site in the lymph of the host 183 1. the larvae 183 2. prepupa1 stage and pupa- tion 156 6. Relationship betv.een specific poly embryony of parasitic hymenoptera and experimental polyembryony� 187 7. Rela.tionship be twe e n specific poly embryony of parasitic hymenoptera and other methods of agamic reproduction. 188 8. F:.wtors conditioning or aiding specific polyembryony in par:3.sitic hymenoptera. 194 9. Rearing da.t� on the sexes. 200 r!umber of adult s emerging from host. 203 10. The origin of mixed broods in poly- e mb ry o n i,c hymenopt era. 207 VII T:::CH�Trr�UE 215 -- -� ----- -------------------�====------------ 1. Rearing methods. 215 2. Histological technique. 222 VIII 'llHE CO:::Et�:::::'ATIO�r OEl STRUCTU� AND HOST �ATIOU -'i.HOnG TES EHCYRTHL\E 225 IX SUI!I:U.RY 229 X BIBLIOGRA.EHY 235 LIST OF lIT GtJRFS Atter page l5-----PlateI: Antenna, head, mandibles and ovary of Berecyntus. l8-----Plat e II : Adult, pupa and larva of 1!. baker! • 20-----Plate III: Eggs of B. baker!. 27-----Plate IV: Larval types in the Chalcidoidea. 4l-----Chari I: Length of life of adults of !.bakeri. 52-�---Chart II: Duration of oviposition and feeding � puncture. 63-----Chart III: Amount of tood consumed by healthy and parasitized '. larvae of Euxoa ochrogaster. 66-----Chart IV: Length ot ttme spent in the different stages of development by nor.mal Eu�oa ochrogaster larvae and by larvae parasitiaed by Berecyntus. 76-----chart V: Mortallty of Euxoa ochrogaster from insect parasitism and "diseases" of larvae and pupae at Saskatoon during 1925. , . 82-----Plate V: :'Diagramatic representations of three shapes aaaimed by larvae of Euxoa ochrogaster parasitized by Berecyntus. se----Chart VI: Ntunber of Berecyntus adults emerging from Euxoa ochrogaster larvae, the eggs fram wbidt these larvae hatched having previously bee� kept at different temperatures. .. -_ _._- --"""-"--':;;..;._-.........__....;..---------------__.............. 90----Chart VII: Nmnber of adults of Berecyntus emerging fram Euxoa tristicula larvae, these larvae having previously been reared together under laborator.y conditions from. the egg to the fourth moult and then subjected to different temperatures tor a period of 30 days.