Experimental Pollination; an Outline of the Ecology of Flowers and Insects
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EXPERIMENTAL POLLINATION AN OUTLINE OF THE ECOLOGY OF FLOWERS AND INSECTS BY Frederic E. Clements and Frances L. Long QK926 | Q £ Published by the Carnegie Institution op Washington Washington, 1923 Oty* B. 31 Mi ffitbrara Nortlt (Carolina &tatp Inineraitij olina State Library ^["CAROLINA STATE UNIVERSITY L THIS BOOK IS DUE ON THE DATE INDICATED BELOW AND IS SUB- IECT TO AN OVERDUE FINE AS POSTED AT THE CIRCULATION DESK. 19 1980 MAY /yyy CLEMENTS & LONG Edith S. Clements I Painted flowers of Aconitum and Delphinium, showing the method of experiment. North Ca(J$i* S'aLe Library # Raleigh EXPERIMENTAL POLLINATION AN OUTLINE OF THE ECOLOGY OF FLOWERS AND INSECTS BY . a ca Frederic E. Clements and Frances L. Long North Q£M!ina State Librarv Published by the Carnegie Institution of Washington Washington, 1923 CARNEGIE INSTITUTION OF WASHINGTON Publication No. 336 Copies of this feoek NOV 6 1923 ... .. CONTENTS. Page 2. Normal and Experimental Pol- 1. Introduction and Methods 3 Introduction 3 lination {Continued). (Continued). Objectives 4 Delphinium scopulorum Methods 4 Addition of odor 25 25 General principles 4 Perfume 26 Normal pollination 6 Summary 26 Experimental pollination 6 Rubus deliciosus 26 Organization of experiments 7 Normal pollination structure 26 Change of position or place 7 Habit and 26 Concealing or disguising flowers. Behavior 27 Mutilation Variation in visits 29 Artificial and painted flowers .... Experiments Mutilation 29 Addition of parts or substances . Competition Petals split or shortened .... 29 29 Manipulation of insects Stamens covered Life-history methods and records Artificial and painted flowers . 30 30 Life-history record of a represen- Crepe-paper corollas 30 tative species Painted corollas Addition of honey or odor .... 31 2. Normal and Experimental Pol- lination Honey 31 Aconitum columbianum Summary 31 32 Normal pollination 15 Rubus strigosus 32 Structure 15 Normal pollination Behavior 15 Habit and structure 32 32 Experiments 1(3 Behavior 33 Change of position 16 Experiments Horizontal racemes 16 Mutilation 33 Racemes inverted 16 Floral envelopes or stamens Mutilations 17 removed 33 Cotton plugs 17 Rosa acicularis 33 Stamens removed 17 Normal pollination 33 33 Hood split 17 Habit and structure Hood removed 17 Behavior 33 Hood and nectaries removed 17 Experiments 34 34 Lower sepals removed 17 Mutilation 34 Side petals removed 18 Corolla shortened Competitive relations 18 Stamens masked 35 Artificial and painted flowers . 35 Artificial and painted flowers. 18 Normal colors 18 Crepe-paper corollas 35 . 36 Artificial flowers 19 Addition of honey and odor. Painted flowers 20 Honey Honey and talcum powder. Addition of honey and odor. 21 Honey 21 Camphor Perfumes 22 Summary Summary 22 Geranium caespitosum Delphinium scopulorum 22 Normal pollination Normal pollination 22 Habit and structure Habit and structure 22 Behavior Behavior 22 Calendars Experiments 23 Experiments Change of position 23 Change of position Inverted racemes 23 Flowers vertical or inverted Horizontal racemes 23 Mutilation Mutilation 23 Cotton over nectaries Cotton plugs 23 Excision Petals removed 23 Artificial and painted flowers. Spur removed 23 Crepe-paper corollas Addition of honey and odor. Landing-platform removed. 24 Honey Artificial and painted flowers . 24 Crepe-paper corollas 24 Summary Crepe-paper corollas with Chamaenerium angustifolium spurs 24 Normal pollination Painted flowers 25 Habit and structure ... .. IV CONTENTS Page Page 2. Normal and Experimental Pol- 2. Normal and Experimental Pol- lination (Continued). lination (Continued). Chamaenerium angustifolium (Cont.). Pentstemon glaber 65 Behavior 45 Normal pollination 65 Experiments 48 Habit and structure 65 Change of position 48 Behavior 65 Racemes inverted 48 Calendars 66 Mutilation 48 Experiments 68 Floral envelopes removed. 48 Change of position 68 Upper petals and the in- Racemes inverted 68 cluded sepal removed .... 49 Racemes horizontal 70 Stamens and style removed 49 Mutilation 70 Artificial and painted flowers. 51 Cotton at the corolla mouth. 70 Crepe-paper corollas 51 Styles, stamens, and stami- Painted corollas 51 node removed 70 Addition of honey and odor. 52 Corolla split 70 Honey 52 Corolla lips separated and Odor 52 staminode raised 71 Perfumes 53 Upper lip removed at the Flavoring extracts 53 throat 71 Summary 54 Lower lip shortened half. ... 71 Pachylophus caespitosus 54 Lower lip removed 71 Normal pollination 54 Lower lip and part of the Habit and structure 54 tube removed 71 Behavior 55 Lips removed except lower Experiments 56 lobe 71 Artificial flowers 56 Corolla tube shortened half. 71 Crepe-paper flowers 56 Petals separated 71 Petals obscured 56 Lower lip split into three Mentzelia multiflora 56 petals 71 Normal pollination 56 Tube end of a corolla slipped Habit and structure 56 over the staminode 72 Experiments 56 Comparative relations 72 Crepe-paper flowers and com- Artificial and painted flowers. 72 petition 57 Crepe-paper corollas 72 Crepe-paper corollas 57 Corolla painted with water- Summary 58 colors 72 Frasera speciosa 58 Honey and odor 72 Normal pollination 58 Sirup added 72 Habit and structure 58 Pentstemon gracilis 73 Behavior 58 Normal pollination 73 Experiments 59 Habit and structure 73 Mutilation 59 Behavior 73 Types 59 Calendars 74 Artificial and painted flowers. 59 Experiments 75 Crepe-paper corollas 59 Change of position 75 Painted corollas 60 Racemes inverted 75 False corollas 61 Racemes horizontal 75 Mutilation Addition of nectar and odor. 61 75 Nectar 61 Landing-platform removed. 75 Summary 61 Brush of staminode removed 75 Mertensia pratensis 62 Anthers and recurved por- Normal pollination 62 tion of upper lip removed 75 Behavior 62 Upper lip partly removed. 75 Experiments 63 Lobes of upper lip separated 75 Mutilation 63 Petals split to the base 76 Changes of corolla 63 Pentstemon glaucus 76 Castilleia miniata 63 Normal pollination 76 Normal pollination 63 Normal behavior 76 Habit and structure 63 Experiments 76 Behavior 63 Mutilation 76 Experiments 64 Upper lobes split to base ... 76 Mutilation 64 Lower lip removed 76 Upper lip removed, spike in- verted 64 Outer lobes of lower lip re- Painted flowers 64 moved 76 Painted bracts 64 Staminode removed 76 CONTENTS Page Page 2. Normal and Experimental Pol- 3. Competition and Constancy (Con- lination (Continued). tinued) . Pentstemon glaucus (Cont.). Chamaenerium angustifolium 110 Middle lobe of the lower lip Comparison HO removed 77 Experiments HO Upper lip removed 77 Summary HO Pentstemon secundiflorus 77 Pentstemon HO Normal pollination 77 Comparison HO Behavior 77 Experiments 113 Calendar 77 Summary 113 Experiments 77 Monarda fistulosa 115 Odor 77 Comparison H5 Powders and extracts 77 Experiments 115 Cotton wads sprinkled with Summary 115 peppermint 78 Weight and composition of pollen Summary 78 loads 119 Monarda fistulosa 79 Value and methods 119 Normal pollination 79 Discussion 127 Habit and structure 79 Constancy 128 Behavior 79 Definitions 128 Experiments 80 Early observations of constancy 129 Mutilation and competition. 80 Bennett's studies of constancy 130 Plan 80 Christy's studies of methodic Summary 88 habits 130 Resume 89 Mueller's results 130 Variation in number of visits. ... 89 Bulman's studies 131 Changes of position 90 Ord's conclusions 131. Masking with cotton 90 Plateau and Perez 131 Mutilation 91 Constancy in Bombus 132 Artificial flowers 92 Lovell's conclusions 132 Painted flowers 92 Kranichfeld's observations 133 Honey and odor 93 Origin of oligotropism 133 3. Competition and Constancy 94 Resume 134 Significance 94 Experimental results in compe- Competition 94 tition 134 General plan 94 Effects of competition 134 Rubus strigosus 95 Constancy as shown by pollen General relations 95 loads 135 Rubus strigosus and Rubus deliciosus 95 4. Principles and Conclusions 136 Comparison 95 Introductory 136 Experiments 95 Early experiments of Plateau and Summary 97 others 136 Rubus strigosus and Rubus deliciosus 97 Artificial flowers 136 Comparison 99 Nectaries 137 Summary 99 Color sense of bees 138 Competition of Rubus with Frasera, Response to detached petals. ... 139 Cleome, etc 99 Color preferences of nocturnal Comparison 97 moths 140 Experiments 99 Response to color without anten- Summary 99 nae '. 140 Rubus strigosus and two or more Perception of form 140 competitors 99 Response of wasps to color 143 Comparison 99 Response of wasps to odor 145 Summary 100 Main researches of Plateau 145 Rubus deliciosus 101 Masked flowers 145 Experiments 101 Removal of corolla 147 Summary 101 Response to different colors 148 Rosa acicularis 101 Addition of honey to vivid nectar- Comparison 101 less flowers 149 Experiments 102 Anemophilous flowers 150 Summary 102 Entomophilous flowers of dull Competition with normal and color 150 mutilated Aquilegia 105 Artificial flowers, second series . 150 Geranium 105 Artificial flowers of green leaves . 151 Comparison 105 Conclusions as to artificial flowers 152 Experiments 106 General summary 152 Summary 109 Role of vexillary organs 153 VI CONTENTS Page Page 4. Principles and Conclusions {Con- 4. Principles and Conclusions (Con- tinued), tinued) . Main researches of Plateau (Con- Recent investigations (Continued). tinued). The pollination of green flowers . 195 Choice of colors by insects 154 Conspicuous flowers rarely visited Errors made by Anthidium 155 by insects 196 Admiration of syrphids for bright Response of honey-bees to colored flowers 156 artefacts 197 Attraction of colored cloths and Pattern vision in the honey-bee. 198 brilliant