Y. Olume 39, No. L, August 15, 19 64
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TABLE OF CONTENTS Y.olume 39, No. l, Augu st 15, 19 64 Floral evolution in the Ranunculaceae. E.E. LEPPIK Volume 39, No. 2, November 15, 1965 Keys to the shore flies of Iowa (Dipte ra, Ephydridae). D. ;L. DEONIER 103 Phymatopsallus new genus, and new species of Phylinae from North America (Hemiptera, Miridae). HARRY H. KNIGHT 127 Residues of organophosphorus insecticides found on corn plants treated for European corn borer control. JACK E. FAHEY, R. D. JACKSON and R. T. MURPHY 153 List of Masters' Theses, 1963-64. 161 List of Doctoral Dissertations, 1963-64. 177 List of ISU Staff Publications, 1963-64 . • 187 Volume 39, No. 3, February 15, 1965 Periodicity of caryophyllaeid cestodes iz:i the white sucker, Catostomus commersoni (Lacepede). ROBERT L. GALENTINE and LEIGH H. FREDRICKSON 243 Cob morphology and its relation to combine harvesting in maize. S. M. SEHGAL and WILLIAM L. BROWN 251 Comparative morphology of the dormant plumule in some genera of Leguminosae. H.P. HOSTETTER 269 Taxonomy of the genus Sphenopholis (Gramineae). KIMBALL STEWART ERDMAN 289 The prediction of soil moisture for the winter period in Iowa. R.H. SHAW 337 Inheritance of quantitative characters in grain sorghum. G. M. BEIL and R. E. ATKINS 345 Volume 39, No. 4, May 15, 19 65 The mammals of Winneshiek and Allamakee Counties, Iowa. SHERMAN A. HOSLETT 359 Productivity and turnover relationships in native tallgrass prairie. MELVIN R. KOELLING and C. L. KUCERA 387 Estimation of field working days in the spring from meteorological data. R. H. SHAW 39 3 Combining biological data from European corn borer experiments over years. D. V. SISSON, T. A. BRINDLEY and T. A. BANCROFT 403 Life history of smallmouth bass, Micropterus dolomieui Lacepede, in the Des Moines River, Boone County, Iowa. JAMES B. REYNOLDS 417 Residues of chlorinated hydrocarbon insecticides found on corn plants treated for European corn borer control. JACK E. FAHEY, R. T. MURPHY and R. D. JACKSON 437 Residues of carbamate insecticides on corn plants treated for European corn borer control. R. D. JACKSON, JACK E. FAHEY and R. T. MURPHY 449 Effect of two legume and two grass crop root systems on Pythium graminicolum disease-inciting potential of field soil. W. L STAUDINGER and W. F. BUCHHOLTZ 453 IOWA STATE JOURNAL OF SCIENCE Vo L. 3 9, No. 1 August 1 5, 1 964 pp . 1 - 1 01 FLORAL EVOLUTION IN THE RANUNCULACEAE E . E . LEPPIK CONTENTS FOREWORD. 3 ABSTRACT. 5 INTRODUCTION. 5 HISTORICAL REVIEW . 9 Some Newly Ob served S e nsory Capabilities of Insects . 9 Experiments with Bees and Nerve Poisons. 10 Nerve Poisons are Disastrous to Bees and Other Pollinating Insects . 11 "The Struggle of Bees" . 11 Foraging Behavior of Honeybees 11 A Disaster with Nerve Pois.ons . 12 DEFINITIONS AND TERMINOLOGY 13 FLORAL MORPHOLOGY AND MERISTIC VARIATIONS IN THE RANUNCULACEAE. 19 Stamens and Carpels . 20 Bracts and Sepals. • . 21 Petals, Nectar-Leaves and Sernaphylls 21 PHYLOGENETIC CLASSIFICATION OF THE RANUNCULACEAE . • 22 Paeonioideae . 23 Anemonoideae 24 Helleboroideae 25 EVOLUTION OF THE FLOWER TYPES IN THE RANUNCULACEAE. 24 Pre-Ranalian Flower Types . 26 Early Rana Lian Flowers . • 26 Progression of Pleornorphism in the Ranunculaceae 29 The System of Iconic Numerals of Insects. , . 30 Origin and Development of Pleomorphism in Flowers. 31 Differentiation of Numeral Patterns in the Ranunculaceae . 32 Numerical Patterns in Pseudanthial Flowers 34 Stereomorphism 35 Zygomorphism . 35 2 E.E. LEPPIK NECTAR-PRESENTATION MECHANISMS IN THE RANUNCULACEAE. • • . 37 Secretion of Unconcealed Nectar . • . • . 38 Formation of Concealed Nectaries . 39 Conditioning of Insect Visitors to Concealed Nectaries 41 CLASSIFICATION OF FLOWER TYPES IN THE RANUNCULACEAE 4 2 I. Amorphic Type-Class . 42 II. Haplomorphic Type-Class . •. 42 III. Actinomorphic Type-Class. 44 IV. Pleomorphic Type-Class .. 45 V. Stereomorphic Type-Class. 45 VI. Zygomorphic Type - Class ----· 46 WINGED VISITORS OF THE RANUNCULACEAE 48 Coleoptera . 48 Hymenopte ra . • .· . • . • . 49 Lepidoptera . • . • . • . 50 Flower Visitors Observed on the Ranunculac eae 50 BIOCHEMICAL EVOLUTION OF THE RANUNCULACEAE . 52 GENERAL RETROSPECT • • 53 Primitive Flower Types in the Ranunculaceae .•.. 53 Differentiation of Numerical Patterns in Correlation with the Development of Concealed Nectaries • . • • . 54 Tropheclexis vs. Selection. • • . • . • • . .• 55 Evolutionary Correlation between Euanthial Flowers of the Ranunculaceae and Pseudanthial Flower Heads of the Compo sitae . • . • • • . • • . • . 56 Typological Grouping and Taxonomic Classification in the Ranunculaceae • . 57 Family Ranunculaceae. • 58 DISCUSSION . 60 ACKNOWLEDGMENTS. 62 LITERATURE CITED 63 PLATES •..• 73 AUTHOR INDEX. • • 97 SUBJECT INDEX • . 98 3 FOREWORD One of the most puzzling chapters for the botany student through the years has been the origin and evolution of colored blossoms of flowering plants. The sharply contrasting colors and intricate forms are p eculiar to green plants, whose "fundamental task" in nature is photosynthe sis and whose characteristic color is due to the green pigment, c h lorophy ll. One can observe that showy blossoms function as attraction s for polli nating insects, but until very recently no one had proposed a satisfac t ory working hypothesis for the development of flowers to their presen t forms and colors. In 1941, R. J. Pool in his textbook, "Flowers and F Low e ring Plants, " characterized the situation as follows: "The esthetic appeal of flowers to the. human certai nly depends on their harmonious combinations of colors, their peculiar sym m e try, delicate constructions, and pleasing odors. But we cannot exp e ct a similar conception of beauty, based on a high order of intelligence, among pollinating insects. We know that insects do not "enjoy" flowers for their color or odor, but for the food that they s e cure. Further investigations may remove some of the "romance" from this alluring fie Ld, but they will probab Ly endow it with e ven a g reater wealth of definitely established scientific truth than is at pre sent a vailab Le . " To unlock this botanical strong box, a scientist was require d with unusually broad interest, a wide range of biological experience , a com mand of several Languages, uncommon powers of observation and a determination to publish his conclusions. In Dr. E. E. L eppik are com bined these traits and more. After many years of effort to bring order to the gray zones between several scientific disciplines and in spi te of interruptions from war and threats to persona L safety which force d him and his family over several years to move from one country to another, he continued his observations. His exploration into the mysteries of flowers has been both an exciting adventure and a successful scientific accomplishment. During numerous travels in Europe, Africa and North and South America h e gathered data on flower types and their pollinators from all geographic zones, arctic to tropic. This material enabled him to propose a new syste m for clas sifying flower types and to relate it to the development of insect groups that act as pollinators. The system revealed a sequence of floral evolu tion from the simple, elementary types to the complicated floral struc tures of the higher angiosperms which paralleled the sensor y d ev e lop ment of the pollinating insects. Implied is a reciprocal corre lation between the evolution of flower types and the sensory developme nt of their pollinators. As support for his hypothesis, Dr. Leppik presents evidence gathered from fossilized flowers and floral imprints in amber, compact clays, silts, muds, and very fine sands. In several papers (1960, 1961 b, l 963a, b) he discusses the fossil evidence for floral evolution avai Lab le at pre sent. 4 Particularly significant is the reconstruction of a Cretaceous mag nolia-Like flower collected from the Dakota sandstone. This is the oldest known floral structure of an angiosperm which shows the size, form and symmetry -of a haplomorphic flower of the Cretaceous period. This was reported in Advancing Frontiers of Plant Sciences (Leppik, l 963a). Under a research project entitled, "The Evolutionary Relationships Between Plants and their Environment," in the Department of Botany and Plant Pathology of the Iowa Agricultural Exp·eriment Station, Dr. Leppik has published a series of papers that brings light on the old mystery of floral evolution. Interest in his work is being expressed by scientists in several states and foreign countries. All provide encour agement for him to continue his efforts. -W . H. Bragonier 5 1 FLORAL EVOLUTION IN THE RANUNCULACEAE E. E. Leppik Iowa State University, Plant Introduction Station, Ames, Iowa ABSTRACT. S i x c l e a r l y s e p a r a b l e e v o l u t Lo n a r y l e v e l s i n the floral evolution of the Ranunculaceae were found to coincide with the six corresponding stages of sensory development of their pollinators as follows: amorphic - hap lo mo r phi c -a c tin o mo r phi c - p le o mo r phi c - .s t e .r e o - morphic-zygomorphic. This is a basic trend of floral evolution, fully recapitulated in a single family. Ex e e p t f o r t h e f i r s t ( a m o r p h i c ) , _a l l u p p e r l e v e l s a r e represented in the present-day floras as clearly sep arable type-classes. They are therefore accessible to direct observations and experimental study. Exten sive statistical data on flower visitors of the Ranun culaceae confirm the proposed theory of the mutual interrelationship between the evolutionary Levels of flower types and sensory stages of pollinating insects. The new picture, obtained by this study, gives us a better understanding of the evolutionary relationship between insects and plants and explains the extreme div er sit y in the fl o r a L st r u ct u r e of the Ran u n·q: u Lac ea e • INTRODUCTION Most taxonomists (Bessey, 1897, 1915; Hallier, 1905; Glo.ck, 1919; Hutchinson, 1923, 1926; Tippo, 1942; Cronquist, 1951; Eichler, 1958; Eames, 196la, b; Thorne, 1963) today consider the family Ranunculaceae to be a natural taxon with well-preserved primitive characteristics of a phylogenetically lowermost group among herbaceous dicots.