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The University of Manitoba in Partial Fulfillment Of SOME ANT-APHID ASSOCTATTONS IN MANITOBA I^TÏTH OBSERVATIONS ON INTERACTIONS BETI^IEEN Formica oreas comptula I^IIIEELER AND APHIDS AT BTRDS HTLL PARK, MANITOBA A Thesis Submitted to the Faculty of Graduate Studies The University of Manitoba by Mary Catherine Anne Madder In Partial Fulfillment of the Requirements for the Degree of Master of Science Department of Entomology October L978 f/,9R¡rnlçs SOMT ANT.APHID ASSÛCIATIt)NS IN MANÏTOBA I.IiTH OBSERVAT I(]NS CN INTTRACT IC)NS BET!,.ITEN Formica orç!q comp'bula l,lHETLER AND APHIDS AT BIRDS IIILL PARK, MANITOBA BY MARY CATHER.TNT ANNE il4ADDER A <lissertution submitted to the Faculty of Cra¿åuate Stuclies <¡f ttre Urriversity of Manitobu in partiir! fulfillment ol' tfue requirernents ol' tl¡e degrce of MASTER OF SCIENCT o" 1978 Fer¡nission hi¡s ber:¡r grurttcd to the LlllRAll'Y OF'TCI!: [JNIVUÈì- StTY OF MAN¡TO!ìA to lcnd or scll copies of tltås dissertatiol¡. to the NAT¡Ol"¿AL LIBRARY OF (IANADA to nlicrol'ilsn tltis rlissertatio¡r and tt¡ lend or sell cr.rpies of tl¡e filnl, und UNIVUIIStTV M!CIì,ûFILMS to publlsh at¡ ubstruct of this dissertation. The autltr¡r reserves other publieutit¡n rigfits, a¡ld neither tlte dissertation ¡ror extensivc cxtructs f rom it ntuy be printerJ or r-¡thcr- wise reproclucecl without thc autl¡or's writtctr irertttlssitltt. DEDÏCATIOT{ Dedicated to Gladys and Jimmie Carter of Corkery, Ontario (for all the happy summers) and to my Mother ]-I ACKNOI'TLEDGEMENTS r wish to express my gratitude to Dr. Grant Robinson for his patience, guidance and encouragement throughout this . study; for his assistance in the collection of samples and identification of aphids, and for stimurating and informative field trips. I am very grateful to Dr. David p. Scott of the Freshwater rnstitute, Environment canada, Inlinnipeg for his advice and assistance. My husband's patience, encouragement, advice and constructive criticism throughout the study and preparation of this thesis \ùere very much appreciated. The second phase of this investigation would not have been possible without the assistance of Mrs. Li-yen Inlang, Ms. sandy venus and Mr. Jim Madder who, in addition to being excellent technical assistants in the f ie1d, \¡¡ere able to relieve the long hours of monitoring with their light-hearted senses of humor. r wourd also like to thank chief Ranger Don Emes, John Nickrin and all the orher people ar Birds Hill park for their many courtesies extended me during the study. My thanks to Dr. André Francoeur for identifying many samples of fusca ants and to Mr. Gordon Ayre for often l_Il- conf irrning my ant identif ications. I am indebted to Ms. C.M. Baker of the University of tr^linnipeg for her translation into English of works by Fossel (L972) and Horstmann (L972). I very much appreciared rhe assistance of Mr. John Serger of the Department of Entomology, university of Manitoba, in reviewing the works by Atanassov .(L974), Egger (1973), Herzíg (1937), Klofr (f959) and Scheurer (1e 7r) . Many of my fellow graduate students and friends, both at University of Manitoba and McMaster University, have contributed in so many viays to the successful completion of this investigation and thesis and I sincerely thank them all. I am grateful to the Department of Entomology, university of Manitoba, for the financial assistance provided through a National Research council operating Grant awarded to Dr. A.G. Robinson. I appreciated the time and accommoda- tion provided to me during the final stages of writing this thesis by McMaster University and Dr. Douglas M. Davies. The time taken by ry supervisory committee to review this thesis and their advice concerning the thesis' improve- ment \^/as very much appreciated and I thank them f or their contributions. l_v TABLE OF CONTENTS Page ÏNTRODUCTTON I CHAPTER I LTTERATURB REVIEI^I. 3 Defining the Relationship. 4 Behavioral Modifications of Ant-Attended Aphids. -7 Absence of Predator-Avoidance Behavior 7 Modification of Excretory Behavior I Behavioral Modifications of Attendant Ants 9 Absence of Aggression Toward Aphids. 9 Drop let- Inducing Behavior . 10 Active Defense and Protection of Aphids. l1 Construction of Shelters L4 Prevalence of Associations Between Ants and Aphids L6 Formicidae l6 Aphido idea L7 Intimacy and Dependence on the P'elationship. 18 Polyethism, Route and Task Fidelity. 23 Species Preferences. ¿-u Factors Affecting Foraging Activities. 27 Temperature and Relative Humidity. 27 Light-Dark Cycle 29 Attendance Time. 29 Number of Attendants 30 Honeydew Foraging Trails 31 Interspecific Competition. 32 Reproductive Behavior of Ants. 33 Brood-Rearing Period 33 Queen and Male Ants. 33 Biology and Ecology of the Experimental Species. 34 Formica oreas comptula Wheeler J+ eñãïEophorus sãIfõlñÏger (Ihowlton) . 35 ell) 3s Stmtãõfus ñerTcanus Baker. 35 CHAPTER II MATERTALS A\]D MBTHODS . JO Collection and ldentification of Ant-Aphid Associations JO Experimental 0bservations of Ant-Aphid Interactions. J/ Site Selection J/ V Page Preparation of the Site. 38 Marking Experiments. 4L Choice of Experimental Aphid Species 49 Experimental Observation period and Procedure. 50 Statistical Analysis 57 CHAPTER ITI RESULTS AT{D DISCUSSTO}T 60 collection and rdentification of Ant-Aphid Associa t r_ons Formicidae 60 Aphidoidea 60 B2 survey of Associations at Birds Hill park Experi mental Site. B3 Experimental observations of Ant-Aphid rnteraction S 86 Attendance Time. 86 Marking Experiments. B6 Erratic Forager Behavior B8 Analysis of Data B8 Bivariate Correlations Between Environmental Parameters 96 Factors Tnfluencing the Number of Aphids in the Colony 101 Day of Season. 101 Ambient Temperature. L02 Number of Ants Present at the Aphid Colony L02 Factors rnfluencing the t{umber of Aàts present a t the Aphid Colony, Stem Activity Up, and Stem Activi ty Down 103 Day of Season. 103 Ambient Temperature. 105 Relative Humidity. r0B Barometric Pressure. 110 Rain 110 skv. LL2 Number of Aphids in rhe Aphid Colony 113 Stem Activity To and From the Aphid Colony 115 Number of Ants Present at the Aphid Çolony . 116 Coefficient of Multiple Dererminatiòn (RZ) LL7 Adjusted Group Means 119 Behavioral }fodifications of Aphids 119 Absence of Predator-Avoidance Behavior r19 Modification of Aphid Excretory Behavior rt9 Change in Aphid Colony Appearance L20 Behavioral Modifications of Attendant Ants ].2L Absence of Aggression Toward Aphids L2L Active Defence and Protection of Aphids by Ants L2L Predators and Parasites I2L Shelters . T26 vi Page Honeydew Foraging Trails t26 Interspecific Competition. L29 CONCLUSIOI.IS. 131 Recommendations for Further Study. 136 LIST OF REFERENCES 138 APPENDICES L45 vll LIST OF FTGURES Figure Page 1. Composition and Distribution of Ant Nests and Plants within Each Quadrat of the Experimental Site at Birds Hill Park, 30 M"y, T977. 40 2. üIest Boundary of Study Area Ïllustrating String Grid, and Hiking Trail to the Right of the l,üoodãn Stakàs. 43 3. Stevenson Screen Containing Device to Monitor Daily Ambient Temperature, Relative Humidity and Barometric Pressure. 43 4. Aspiration of Aphids from a Colony Being Tended by For1icg oreas comptula on Salix bebbiâna inEo a Glas s Vial ContãiñlngÃ% ETOE . 44 - 5. Nest of Formica oreâs comptula- Located in Quadrat l-C of Srudy Areã-ãE-BirAs-EiTf-Þã¡E; Maniroba on 8 June, L977 46 6. Nest of Formica oreas comptula Located in Quadrat 6-8 of Study Area-ãt eiñs Fiff-ÞãiEl Manitoba on (a) I June, L977 and (b) 15 July- L977 48 7. Collection of Honeydew by Formica oreas comptula from Symydobius americanus: (a) ApñIã feedïng-Sire Locatü-ãÏone-thtBranãE-along of Betuia slanduloã" rr"r. landulifera and (b) Aphid FeeãIng*SiEã-Io cated on the Terminal Growth of Þ. glandulosa var. glandulifera 52 B. Collection of Honeydew by Formica oreas comptula from Chaitophorus saliciniger Feeãïr¡g on SãTix 6-ãFÏana. 54 9. Collection of Honeydew by Fognica oreas comptula from Pterocomma smithiae Feeding . 54 10. Daily Mean lTumber of Ants and lTumber of Aphids in Birch Aphid Colonies (a) 3-E-2 and (b) 3-E-38 from B June, L977 at Birds Hill Park. 9l 11. Daily Mean ltTumber of Ants and Number of Aphids in I¡,lillow Aphid Colonies (a) 5-D-9 and (b) 5 -D-347 from 8 June, L977 at Birds Hill Park. 93 viii Figure Page L2. Daily Mean I'lumber of Ants and ]Tumber of Aphids in Balsam Poplar Aphid Colonies (a) 9-C-14 and (b) 5-E-5 from B June, L977 at Birds Hill Park. 95 13. Syrphid Larva Preying on Aphis viburniphila on Viburnum rafirleeggianum while Formica oreas comptulal¡Jcrker eõTlects @the Aphîd-Tdlõnr . r23 L4. Root Chamber at the Base of a Poplar Sapling wherein Many Formica oreas comptula T¿'Iorkers tr'lere Obs erved 15. A Typical Trail Used by Formica oreas comptula ro Reach Piänts upon uihich aphiã-TõlfoniãFÏãre-TõããEéã L28 IX ABSTRACT Madder, Mary Catherine Anne. M.Sc., The University of Manitoba, October , L978. Some Ant-Aphid Associations in }{anitoba with Observations on Interactions between Formica ofeas comptula Inlheeler and Aphids at Birds Hill Park, Maniloba. Major Professor: A.G. Robinson. A survey of Manitoba ant-aph.id associations \,üas con- ducted by collecting and identifying samples of ants ancl aphids found together on aphid host plants. Qualitative and quantitative monitoring of the interactions of Formica oreas comptula with Symydobius americanus, Chaitophorus saliciniger and Pterocomma spithiae was carried out during L977 at BirCs Hill Park. Data were analyzed by stepwise multiple linear regression analyses. Twenty-seven species (seven genera) of ants tending 62 species (28 genera) of aphids on 39 host plants T^iere collected in Manitoba during I976 and 1977. Formica species were the dominant aphid-attendants and Aphis species \^7ere the most frequently attended aphids .
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