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THE BIOLOGY AND VECTOR CAPABILITY OF TRIATOMA SANGUISUGA TEXANA USINGER AND TRIATOMA GERSTAECKERI (STAL) COMPARED WITH RHODNIUS PROLIXUS (STAL) (HEMIPTERA: TRIATOMINAE) By WARREN FLOYD PIPPIN (f Bachelor of Science Colorado State,University Fort Collins, Colorado 1941 Master of Science Oklahoma State University Stillwater~ Oklahoma 1958 Submitted to the faculty of the Graduate College of the Oklahoma State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY May, 1968 OKLAHOMA STATE UNIVERSfl'Y Li8RARY OCT :& ·1 1968 THE BIOLOGY AND VECTOR CAPABILITY OF TRIATCJJ.MA,M,,._.,N,-,,---~, ••.. , . SANGUISUGA TEXANA USINGER AND TRIATOMA GERSTAECKERI (STAL) COMPARED WITH RHODNIUS PROLIXUS (STAL) (HEMIPTERA: TRIATOMINAE) Thesis approved: Thesis Adviser 688686 ii PREFACE The presence of Trypanosoma ~. the causative agent of Ameri can trypanosomiasis, has been co:n.firmed and well documented in the United States. '1,'he biolo~y and vector effectiveness of many species of Triatoma that transmit this organism are not well known. In particular, studies that directly compare South American an.d United States species have been limited. It was decided that a comparative study of two Texas species with th.at of a h,ighly effec;tive South American vector would be desirable. Data were taken during the period November, 1964 through November, 1967. I should like to express my appreciation to Sgt. M. J. Gaylor, USAF, and MSgt A, T. O'Neal, USAJ!\ who directly assisted in all aspects of the stuo.y an<;i to Dr. R. L. Usinger, Chairman, Division of Entomology and Acarology, University of California, Berkeley, California, who kindly checked the initial identification of the triatome bugs. Dr. Paul F. Law, Southwest Research Institute, San Antonio, Texas, gave valued assistance in the collection of bugfl. Dr. Irving G. Kagan and Miss Lois Norman, Laboratory Branch, Communicable Disease Center, Atlanta, Georgia, pro vided serological support, furnished !· cruzi antigen, and offered valued suggestions, Lt. Colonel Gregory Skafidas, USAF, BSC, provided certain serological and parasitological support and for this I am grateful. Rosalie Watkins, Hooper Foundatic;m, University of California Medical School, San iii Francisco, California, ldndly furnished specimens of Rhodnius prolixus for colonization. Pr. D. E. Howell, Head, Depi:J.rtment of Entomology, Oklahoma State University, served as chairman of the writer's graduate committee and gave valued assistl;lnce on the manuscript preparation. I am indebted to the members of the thesis committee, Dr. R.R. Walton and Professor Q, B, Graves fQr their review of the manusqript. Lt. Colonel Wesley R. Nowell, USAF, BSC, · pffe;ed valued suggestions on all phases of the study. My since;re acknowledgement is made to the United States Air Force and in par... ticular to Colonel Elmer V. D;:ihl, USAF, MC, Commander, USAF Epidemio logical Laboratory (AFSC), Lacldand Air Force Base, Texas, who made this study possible, iv TABLE OF CONTENTS Chapter ·Page I. ~NTRODl,TCTION • • • • • • • • . .. 1 n, REVIEW OF THE LITERATU;Jl.E. 3 Historical r • • • • • • • • . 3 Distributio:p. and Incidence of tb,e Disease . 5 Mammalian Reservoirs • . • • . 7 Vectors . • • • • . • • • • • 7 Infection. Rate in Triatome Bugs . 8 Bionomics of Triatome Bugs • 9 General . • • • , . 9 '.!:· sanID1!,s.uga. • • • . · 10 . 11 T. gerstaeckeri • ' . F.eeding Habits • • , • • • • . 11 Size of Blood Meal. 13 Host Relationships. ' . ' . 14 Geographical Distribution . • • • . 14 Dispersl;l.l. • . • • , • . , . 15 . 15 In Vitro Feedi:Q.g • • •. , • ~ . Transmission, and Vector Efficlency 16 u;c. M.A.TE:&IALS AND METHODS • . ' . 19 Laboratory, Procedures 19 Re~ring Bugs . • , . • 19 Feeding . , . · 20 Maintenance and Handling of I· cruzi Cultures . ~ . ·. 21 Infection 1:1,nd In(ectivity of Experimental Anim~ia . ~ .. 23 Infecti(i)n and Infectivity of Triatoma . • . 26 Membrane Feeding • . • · • . · . 28 v Ch~pter Page Field Stqdies ·• ... .. 29 Outdoor Cage . • . , 29 Field Collection of Bugs and Wood, Rats 30 ·. Nocturnal Flight Habits. 32 IV. RESULTS •...•... 34 . Laboratory Studies . 34 · Mating .. .. ' ,. 34 Eggs .... 35 · Ovipos~tion • , . 35 ' . ' ' Fecundity and Longevity of Adults . 35 Egg Production of Virgin Females . • .. 37 Number of Eggs Laid After One Copulation 37 Fecundity and. Longevity of Unfed Adults 38 . Pevelopment Period from Eigg to Adult . , •• 38 Feeding Habits. • . • , • • . , 39 Feeding and, Defecation. Times . 41 Coprophagy . • . , . 43 Amoqnt of Blood Ingest~d at One· Feeding . , . 43 Infection of Experimental Animals • . 44 SusceptibiU.ty of Bugs to Infection ' . 46 . Membrane Feeding . , . .. 47 Field Studies , . ., 'I • ' • • . ·, 48 Developm~ntal Period Egg to Adult. , 48 Comparative Fecup,dity and Longevity of Adults , . 49 Survival with One Feeding; . , . 51 Intraspecific Parasitism • , . , 51 Field Collections • . 52 ·• Noc;turnal Flight Habits, 54 v. SUMMARY A.ND CONCLUSIONS 57 LITERATURE CITED . ' . 66 APPENPJX "A'' . .. 74 A.PPENOIX "B" , . ' 95 vi LIST OF TABLES Table Page I. Fecundity and Longevity of Adult Triatominae at 65-82F ;ind 20-65%R.H ...••. , ..••.•.•.. , . 75 II, Fecundity and Longevity of Adult Triatominae at 80 F and 65% lt, H. • • . , . • . • . • • . 76 III. Maximum Number of Eggs Laid After One Copulation . 77 IV. Reproductive Capacity and Longevity of Unfed Male and Female Triatominae • • . • , . • , . • 78 V. Developmen.tal Period, in Days, at Two Different Tempera- tures . • • • . • ,00,,,, •• ,. ••••• 79 VI. Feeding, a:nd. Molting of First Instar Nymphs Fed on Various Host Animals . • • . • 80 VII. Defecation ancl Feeding Times of l'riatominae . • 81 Vlll. Mean Weight and Amount of Blood Ingested by Triatominae • 82 I IX. Virulence c;,f Two Strains of '.!'.. cruzi to Laboratory Animals 83 x. Infection of Experimental Animals by Various Routes of Inoculation . • • . , . , , , 84 XI, The Susceptibil,i.ty of Triatominae to Two Strains of T.-~ cruzi 85 :XII. Susc;ieptibility of Triatom.inae Fed on Infected Animals With no Observable Circulating Parasitemia . • . • 86 · XIII. Relative Number of Trypanosomes (Per 100 Fields) Passed in. Adult Bug Feces . , , . • . 87 XIV. Infection and Feeding of First Instar Nymphs and Adult Triatominae l'nrough Membranes . • . 88 vii Ta.bl~ Page xv. Developmenta~ Periodi in Days,. Under Field Conditions .• • • 89 XVI. Comparative Fecundity and Longevity of Triatoma Under Field Conditions . , , . , . , . , 89 XVIJ. Number and Viability of Triatoma Eggs Under Field C9n1;Utions • . , . 90 XVIII. Survival of Triatoma Under Field Cond~tions With One Feeding "' , . , . , · . ~ . 91 XIX. Compa:rison of Adult ! . gerstaeckeri Collected in· Light Traps at Two J,)ifferent Sites . • , , . , . 92 }p{. Compadson of Adult !· gerstaeckeri CollectE;ld During Successive Years in. Light Traps Located in the Same Area . , , , , . , " . , . 93 X:XJ. temperature and Light Trap Collection Correlations for .!., .~e~st~eokeri on. Lackland l\.FB . , , . , , 94 x:xu. 'rime and Temperature Correlations for T. gerstaeckeri Collected in, Light Traps . , , . ~ : . , · . ' 94 viii LIST OF FIGURES Figur~ Page 1. Reartng Container for· B1,1gs . ' . 96 Iqtt Confined i:Q. Feeding Chamber . 2. ' • ft • . ' . 96 3. C~ge Holder with Restrained, Rat and Feeding Bug. 97 4. Bug Susp~nded o:n Glass Rod . , . ' ' 97 5. SQsper;i.ded Bug Feeding on R~stra,ined :a.at . 98 Bu,gs ·Suspended Over Defecation Chambers . ' . ' . 98 7, Membran~... Feeding Apparatus. ' . 99 8. Bug Poi;;i,tioned .Under ·Membr::i,ne . " . 99 9. Outdoor Cage • . • ~ . • . 100 10. Simulated Wood Rat Nest in Outdoor Cage 100 11. Clump of Cactu,s Containing Several Wood Rat Nests. 101 12. · Wood Rat Nest at Base of Cactus Plant, . 101 13. Standard Blacklight Trap Used in Survey . .. ' . 102 Adults of ! ... l?ierstaeckeri (A), '!.. ~ .. texana (B), and ~·. prolix:us (C). ' Females on the Left . • • • . ' . 103 19. First Instar Nymphs of !· gerstaecke:ri Feeding cm Emerging Adl,llt of&· prolixus . • . • . • • 103 ix CHAPTER I INTRODUCTION . A parasitic hemoflagellate, Tr,manosoma cruzi, Chagas, is the causative agent of Ch,aga~' disease or Ap.1.erican trypanosomiasis. Thi.s :parasite has been recovered in nature frolt'l' ~evera1 species of. mammals. found in the. south and southwestern United States. In addition, this same organism occurs naturally in m~ny speoies of indigenous blood sucking insects belonging to the order Hemip- te;t;"a 1 subfamily Tria.tominae. In 1909 1 Carlo~ Chagas, working in Brazil, some..:. what clarified the epidemiology o~ the disease by tracing the life cycle of the try- p~oso;rne from mammal reservoir to man through infected triatome bugs. Despite this natural reservoir-vector relationship, and the relative pre- valep.ce of Chagas' disease throq.gJ:iout South and Central America and Mexico, no infections were reported in the United States until 1955. In that year, two indi~nous cases were re9orded from south Texas. Subsequent serological studies in Texas have shown complement fixing antibodies in the blood of nine of 500 huma:r;is tested. It is not c;:ompletely kn.own why: this disease is not more prevalen{ in humans in the southwestern·part of the United States. There is considerable evidence that humans frequently contact blood sucking triatomid bugs that harbor the protozoan Trypanosoma cruzi; One of several reasons . -- suggested for the Qomparatively· low human incidence of Chagas' disease in the United States is difforences. in the ~bits of the ini;Ject vectors . 1 2