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E < d&N'*- RAR 1 ES^SMITHSONIAN^ INSTITUTION^ NOliniUSNI^ NVINOSHJLIIrMS40 S3 I d Vd 8 . *5 /£%w&\ ? /^x | “ < q: '&TK?' “ ^ 5 5 LruiiSNI^NVINOSHIlWS ~S3 I a \/a 9 II "Tl B R AR I ES^SMITHSONIAN^INSTITUTIC r~ Z __ r~ Z ^ R I E S SMITHSONIAN to INSTITUTION NOlifUllSNI £0 NVIN0SH1IWS S3iavden £/? > LIJLSNI NVIN0SH1IIAIS S3ldVdail § - . LIBRARIES SMITHSONIAN 2 O INSTITUTION n 'iW&h UhHGnHh n Cover Design by Jeffrey Homar Title Page Design by Gail Mitchem School of Fine Arts University of Wisconsin-Milwaukee TRANSACTIONS OF THE WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS LVII—1969 Editor WALTER F. PETERSON mracTioNs of the wisdom academy Established 1870 Volume LVII GREEN POWER: THE INFLUENCE OF PLANTS ON CIVILIZATION 1 John W. Thomson THE RELATION OF HENRY JAMES’ ART CRITICISM TO HIS LITERARY STANDARDS 9 Donald Emerson VIOLENCE AND SURVIVAL IN THE NOVELS OF IRIS MURDOCH 21 Donald Emerson LIFE AGAINST DEATH IN ENGLISH POETRY: A METHOD OF STYLISTIC DEFINITION 29 Karl Kroeber JULIUS BUBOLZ FOUNDS AN INSURANCE COMPANY: A STUDY IN RURAL LEADERSHIP AND RESPONSIBILITY 41 Walter F. Peterson ANTI-GOLD RUSH PROPAGANDA IN THE WISCONSIN MINES 49 Watson Parker OSHKOSH GRADUATES VIEW PUBLIC SCHOOL TEACHING IN LETTERS TO RUFUS HALSEY FROM 1905 TO 1907 Edward Noyes GINSENG: A PIONEER RESOURCE 65 A. W. Schorger LAKE SIZE AND TYPE ASSOCIATED WITH RESORT LOCATIONS AND DENSITY IN NORTHEASTERN WISCONSIN: 1. ONEIDA-VILAS AREA 75 L. G. Monthey GLACIAL GEOLOGY OF NORTHERN KETTLE MORAINE STATE FOREST, WISCONSIN 99 Robert F. Black AGE AND GROWTH OF THE WALLEYE IN LAKE WINNEBAGO 121 Gordon R. Priegel REGULARLY OCCURRING FLUCTUATIONS IN YEAR-CLASS STRENGTH OF TWO BROOK TROUT POPULATIONS 135 Ray J. White and Robert L. Hunt DISTRIBUTION, STANDING CROPS, AND DRIFT OF BENTHIC INVERTEBRATES IN A SMALL WISCONSIN STREAM 155 John J. Peterka BIOLOGY OF THE COREIDAE IN WISCONSIN 163 T. R. Yonke and J. T. Medler HOST RECORDS AND PHENOLOGY OF LOUSE-FLIES ON WISCONSIN BIRDS 189 Nancy S. Mueller, Helmut C. Mueller, and Daniel D. Berger EDITORIAL POLICY The Transactions of the Wisconsin Academy of Sciences, Arts and Letters is an annual publication devoted to the original, scholarly investigations of Academy members. Sound manuscripts dealing with the state of Wisconsin or its people are especially welcome, although papers by Academy members on topics of general interest are occasionally published. Subject matter experts will review each manuscript submitted. Contributors are asked to forward two copies of their manuscript to the Editor. The manuscript should be typed and double spaced on 8V2 x 11" bond paper. 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Documentary footnotes should appear at the end of the paper under the heading “References Cited.” Supplementary or explanatory notes of material too specialized to appear in the text itself should be typed on a separate sheet entitled “Footnotes” and appended to the section entitled “References Cited.” Contributors should avoid unnecessary documentation wherever possible. Other matters of style should be in harmony with current practice in the subject matter area. Galley proofs and manuscript copy will be forwarded to the author for proofreading prior to publication; both should be returned to the Editor within two weeks. Papers received on or before July 15 will be considered for pub¬ lication in the current year. Papers received after that will be considered for publication the following year. Contributors will be given five offprints of their article free of charge. Additional offprints in sets of 100, 200, etc. may be ordered at the time galleys and copy are returned to the Editor. Price will vary according to quantity desired and the length of the article. Manuscripts should be sent to: Professor Walter F. Peterson Editor, Transactions of the Wisconsin Academy Lawrence University Appleton, Wisconsin 54911 JOHN W. THOMSON U7th President of the WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS GREEN POWER: THE INFLUENCE OF PLANTS ON CIVILIZATION John W. Thomson When considering the possibilities for this address tonight there was a strong temptation to entitle it “Flower Power”. But, such a topic presented by a professor from the University of Wisconsin might perhaps be unappreciated by many of our citizens who com¬ ment upon daily life on our campus. As a professional botanist my own field of research and teaching involves the science or art of classification and the naming of plants, a field known to the professional as plant taxonomy. In actual practise a plant taxonomist studies many aspects of the life of plants : their physiology, chemistry, ecology, cytology, uses, and many other things. Perhaps more than for any other botanist his tracing the history of the names of the plants leads him into an awareness of significant historical events which resulted from the intertwining of the life of man and the plants of his environment. The current geologist will point out a fact based upon his solid evidence that in the earlier days of its founding, the earth was surrounded by a different mixture of gases than that with which we are familiar: The abundant volcanoes produced an atmosphere with a higher content of carbon dioxide than the 0.03% of today and less than the 20% of oxygen which sustains our life. The evo¬ lution of plants which used this high concentration of carbon dioxide gas in their photosynthetic activity, giving off oxygen as a byproduct, altered the atmosphere to the more liveable atmosphere on which we are so dependent today. The algae in the waters of the earth can take their carbon dioxide from the dissolved bicar¬ bonates, precipitating the unused portion as carbonates, usually lime carbonates. The thousands of feet thick of limestone deposits have locked up in them the carbonates from which the ancient algae released the oxygen in their life processes. The enormous coal deposits of the world also represent such an alteration of the atmos¬ phere in which the carbon dioxide of the air was taken out and changed into organic materials which were fossilized and the byproduct oxygen enriched the air. For some people, the earliest relationship of man and plants was perhaps that described in the delightful tale of the use of the apple by Eve in the Garden of Eden as related in Genesis. 1 2 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Studies of anthropologists tell us that man evolved as a predator, a hunter. His early weapons, his art, the artifacts of his civiliza¬ tion, attest to this. He had little to do with plants except when game grew scarce. Then he would gather nuts, berries, and roots of the plants which he found around him. He became a gatherer of plants as well. Probably his earliest use of plants on a consistent basis was as fuel for his fires. The early development of hand axes perhaps reflects the possibility of such use and yet these were mainly weapons. The use of fire dates back only some 20,000 to 30,000 years ago (Baker, 1965). Man may also have used plants in the making of shelters of wattle or of reeds and barks. Probably the grasses were early harvested in the wild as wild rice is still harvested in northern Wisconsin and Minnesota, or as small grains in the Sudan. At this point I should like to bring to your attention two pos¬ sible routes of the evolution or development of civilizations based upon plants. The tropics are large areas where a civilization based upon fruits and roots evolved, including products as breadfruit, the tubers of tara (corms) and “yams” of the banana type fruits or of Dioscorea (tubers). These are perishable foodstuffs. Fur¬ thermore their collection requires much hand labor. They cannot be laid up against starvation periods or crop failures, nor are they easily transported. A civilization based upon such foods as these is subject to periods of feasting or famine. How different is a civilization such as ours based upon the grains of grasses. The grass stores its food in a non-perishable easily handled package. Its stored food is on stems which are easily cut and processed and it lends itself to mass treatment, especially by machinery. Its prod¬ uct can be stored, protected from the depredations of insects for many years, circumventing the old cycle of inevitable feast and famine periods. Its ease of culture and of transport has led to a situation in which over 30 people may be supported in cities by one on a farm. This proportion, still increasing today, has made possible a civilization of immense cities in cold climates. It makes possible the feeding of one part of the world by the production in another. An equitable sharing of the products of the earth is more possible with the grains of this family of plants than with the products of any other plant family except possibly the legumes. The story of the rise of this allegiance of man to the grasses is a long story, perhaps of three chapters achieved quite independ¬ ently in three parts of the world ; one based upon wheat and origi¬ nating in Asia minor, one based upon corn and originating in the new world, and the third based upon rice originating in the orient. 1969] Thomson — The Influence of Plants on Civilization 3 Agriculture, according to Baker, had its inception in the Meso¬ lithic or Middle Stone Age, a period when the glaciers were melt¬ ing away in Europe. Around the communities of the “hunter- gatherers” were undoubtedly disturbed soils, midden heaps, areas rich in nitrogenous wastes. Some of the weeds may likely have developed there. They would have been derived from plants of dis¬ turbed habitats such as rock slides, cliff faces, sea shores, sand bars, cattle wallows, etc. Among these ancient weeds were species of Triticum, the wild wheats, three species of which are known from remains of a 7th millennium village in the Tigris-Euphrates valley. These grains were probably parched to assist in removing the husk and prepared in a gruel, as bread-making is a compara¬ tively recent art. In any case these weedy plants became crossed with other weeds in the genus Aegilops and the chromosomes mul¬ tiplied to give us the hexaploid bread wheats of today. The best of the progenies of these weeds were probably unconsciously selected and fell near the homes of our ancestors. Somehow, some¬ where, some genius among these ancestors decided that rather than to rely on finding the grains, a better way would be to scratch the soil and thereby promote their growth. Thus agriculture and the dependence of man on a sedentary life was born. A similar story can be told for the independent origin of indian corn or maize, in the new world. Its origin is less well known than that of wheat. It does appear to be related to teosinte (Euchlaena mexicana) but to also have characters derived from a weedy species of Tripsacum. One of the fascinating developments of such weeds into plants upon which we depend is the series of crops which European man developed from a nondescript weed of the chalk cliffs of England and France, a mustard, Brassica oleracea. From this unimposing start have been derived cabbage by developing the terminal bud, brussell sprouts by developing the lateral buds, cauliflower and broccoli by developing the flower buds, and kale by developing the leaves. Would that some genius among us find such promise in the pigweeds of our garden! One other food plant should be mentioned with respect to its influence upon our history. The potato originated in the high Andes where many varieties are grown. The conquerors of Peru intro¬ duced it to Spain during the 16th century and from there it spread to all of northern Europe. It has become one of the world’s most important crops. In Britain and Ireland it was the main crop fed to the peasantry with perhaps a pig or two per family per year allowed as a source of protein by the landlords who owned the land. Such dependence upon a single crop was dangerous and when the 4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 late blight fungus (a plant of course) hit the fields of Europe dur¬ ing 1845-1847 millions of the peasants across northern Europe starved to death. Many others, especially from hard-hit Ireland, migrated to North America, contributing to the large populations of Irish along the eastern seaboard and affecting the destiny of the United States. A very important development of this period was the repeal of the “infamous*’ corn-laws of England which pro¬ tected the small grain grower of Britain from competition of grains from the United States and Canada. John Peel (well known in a song) was influential in the British Parliament in securing this repeal. The increased import of grain from abroad made it neces¬ sary to secure ocean travel against privateers and pirates who then operated with “permits” from the nations, and Britain promul¬ gated the doctrine of the freedom of the seas to all trade, and built the tremendous navy necessary to enforce this doctrine. For many decades this navy was the guarantor of the freedom of the seas, and Britain was the world’s greatest seapower until the period of World War II. We must admit that this doctrine of the freedom of the seas has profoundly influenced the course of history, and the potato and potato blight initiated it. From one of our fiber plants, cotton, we have inherited some of the most serious issues of today. Let me tell you some of its back¬ ground. Cotton originated from wild plants, some of Asiatic, and some of North American origin. In India it has been cultivated for several millenia. The Asiatic strains were undoubtedly known in biblical times. The New World varieties had to await the discovery of the Americas for their introduction. In Europe, prior to the use of cotton, the main fiber used was wool. It was a warm fiber suited for use in a cool moist climate. Its original production was largely a home cottage industry, the sheep being sheared, the wool spun by the housewife and the thread woven either by the housewife or the itinerant weaver. Cotton altered this picture of domestic tranquility. The new methods of manufacture made possible a cheaper product. The cotton gin in¬ vented by Eli Whitney made it possible to get large quantities of the fiber separated from the seed easily and the spinning machines of Richard Arkwright (1768) and enormous weaving machines made mass production possible. The Industrial Revolution had begun with all its ills and abuses. It was a revolution which altered the whole way of life of man, taking him far from the land, putting him in cities and developing the immense urbanization of whole nations. It was with the fiber of cotton that all of this began. A further enormous sin may be laid to cotton. In the southern part of the United States a plantation system based on the produc- 1969] Thomson — The Influence of Plants on Civilization 5 tion of dye indigo had evolved. An important blue dye was obtained from this plant. The system was faltering in competition with the dye indigo obtained from India where labor was even cheaper than that of the slaves the planters of the south were importing from Africa. The rise of the mills of cotton manufactures made the demand for cotton so great that a rapid shift to cotton and tre¬ mendous expansion of the plantation system in the south occurred. The importation of the negro slaves from Africa and their use as labor for the plantation has led to a multitude of problems for our country. We can blame the Civil War upon cotton, the exhaustion of the lateritic soils of the south, the social ills of the south, and the riots of today upon cotton. Truly this plant opened to us a Pandora’s Box containing much evil. From time immemorial the spices have been articles of trade, sometimes of use only to the wealthy as a status symbol of greater value than any commodity even the precious metals. Cloves, pepper, and cinnamon found their way from the Orient during the Middle Ages via long voyages, camel trains and arduous journeys. They were prized for their flavors, yes, but in addition for the promotion of perspiration, for aid to digestive processes, for preservatives, and because of lack of refrigeration, to cover up the flavor of par¬ tially rotting meat, and were even used as deodorants by people who believed that baths were dangerous and unhealthy. To avoid the inevitable losses and tribute attrition of profits caused by the long overland trade routes, the great voyages from Spain, Portugal and Italy were undertaken. These may have commenced with Marco Polo’s epic 24 year journey in the mid 13th century and continued with those dates which every school child can recite for you: Columbus 1492, Vasco da Gama 1497, and Magellan 1519-22. Many were the wars and naval engagements fought in our history to secure monopolies of the spice trade. Portugal, Spain, England, and Holland fought for and sometimes won, sometimes lost, colo¬ nial empires whose products were the fragrant and profitable spices. Let anyone think that we have not inherited a legacy of problems from these plants and we shall let the names of Oriental and south seas trouble spots roll from the tongue — Indonesia, Macao, Ceylon, Malaysia, Hong Kong, to name a few. The development of the overseas routes discovered during the great period of exploration led to the need for navies to keep open the routes and protect the trading vessels from piracy. That was the day of wooden vessels, for it was not until the battle of Hamp¬ ton Roads in 1862 when the Monitor met the Merrimac that the era of wooden ships began to end. The need for forests of oak for the ships and of conifers for spars led to many problems as the 6 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 forests of Europe became exhausted. Naval operations around the Baltic sea were necessary to insure Britain a supply of masts and spars. Naturally this type of operation became objectionable. Finally during the latter part of the 17th century North America became a source of naval timbers and supplies. During the Napoleonic Wars of the 19th century it was possible for Britain to win the victories of Aboukir Bay and Trafalgar only because she had access to the timbers of Canada for naval construction. Lest we think that the movement of armies and navies caused by plant resources is a phenomenon of only the past, something which ended in the Victorian Period, let me hasten to call to your attention some of the strategies of World War II. These were determined in the Pacific Theatre of Operation in large part by plants. In central America and on the Amazonian slopes of the Andes and the Amazonian slopes of Colombia and Venezuela are trees from which rubber is obtained. This product remained mainly a curiosity until 1839 when Charles Goodyear discovered the proc¬ ess of vulcanization. This discovery makes it possible to use rubber in the manufacture of tires for vehicles including those on which our vast and mobile civilizations and armies are now completely dependent. The rubber trees were eventually imported to Ceylon, Malaya, and Indonesia where the principal plantations of the world are located. A second plant genus of the same region of northern South America, Cinchona, supplies the world with quinine, the drug nec¬ essary to control malaria. Its introduction to the East Indies led to the source of 90% of the world's supplies being the plantations of Java. Without quinine, no army can effectively operate in the tropics. Without the wheels to roll upon, and quinine to protect against the scourge of malaria, our great armies would have been impotent. The strategies of Japan in the Pacific were to cut us from these vital supplies coming from the far reaches of the Pacific. Our strategies had to include campaigns to regain control of these vital plantations. The quest for plant materials to treat the ailments of man is an old one. Probably our ancestors in testing the edibility of plants when they were in transition from hunters to hunters and gatherers sampled everything in their environment and in some cases found surprising effects ; laxative, emetic, hallucinatory, soporific etc. The body of lore they accumulated was passed on as an oral tradition by their medicine men; later the first books on plants dealt mainly with their medicinal values. This traditional background of botany still finds use today although many drugs of today are now synthesized rather than extracted. 1969] Thomson— The Influence of Plants on Civilization 7 For the alleviation of pain we still may use cocaine from the leaves of the cocoa tree and morphine from the latex of poppies. Digitalin from the foxglove is important in the treatment of cer¬ tain heart diseases; atropine from belladonna finds many impor¬ tant medicinal uses; precursors of cortisone are obtained from yams of the genus Dioscorea; and Rauvolfla yields reserpine which has proven so effective in the treatment of mental ailments. The tremendous effects on world health by the antibiotics obtained from the mold fungi such as Penicillium must also be recognized. Some of the extracts from plants contain addictive alkaloids as morphine or its derivative heroin in the latex of the opium poppy or nicotine in the tobacco plant. What have been the effects of such plant products on man? It is not necessary here to do more than remind one of the tremendous economic and social effects of these addictions. It is an ironic twist of events that the addiction of the nicotine user is the chief source of funds for the Outdoor Recrea¬ tion Act of Wisconsin, or that a major source of revenues for the government is this same addiction. Many of us also certainly are habituated, if not addicted to the milder alkaloid caffeine contained in coffee. Sometimes too, we may be enlivened by the alcohol pro¬ duced by the yeast plants when carrying on anaerobic fermenta¬ tion in obtaining their energy for life processes. There are not many other plant products, or even animal products, that have not been utilized by man in seeking to harness the yeasts to his desired aims. Tonight I have spoken of but a few of the examples of the power of plants in influencing man’s course of action. I could have spoken of many more, of cacao and coffee, of tea, and of the forests and the influence of paper, other foods and fibers, things to chew, an amazing array of plants from far corners of the earth. But, I should like to conclude with some thoughts concerning our future relationships with plants. In the newspaper this past weekend was a discussion of a vast project which was proposed for South America. A dam would form an inland sea as large as western Europe. The newswriter with remarkable insight commented that he wondered what the significance could be of removing so much of an area of trees from the oxygen regeneration system of the world. This is part and parcel of the same problem in which we use the fossil fuels of the earth, coal and petroleum, taking oxygen from the atmosphere and releasing carbon dioxide by the millions of tons, a process which Prof. Reid Bryson, one of our academy members and a climatologist will tell us portends serious altera¬ tions in the climate of the earth. This too, Prof. Bryson tells us, is correlated with problems caused by the removal of the vegeta- 8 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 tion of large areas of the earth by man and his domesticated ani¬ mals, especially the goat, leading to atmospheric dust pollution which causes greater reflection of the radiant energy of the sun, in turn thus cooling the earth and its climate. Continuation of this process will certainly force man to alter much of his way of life in the northern hemisphere. To reverse this process and to revege¬ tate the large areas of the world which once were green and sup¬ ported huge cities as those of the Mycenians and the Sumerians in the region of Asia Minor seems well nigh impossible. The evi¬ dence seems incontrovertible that the loss of the vegetation of these regions caused the decline of great civilizations. A hitherto unsuspected influence of plants is only now making itself felt. Henry Thoreau’s statement of it was borrowed for the title of a recent publication of the Sierra Club “In Wildness Is The Preservation Of The World”. Perhaps we can say that this is an influence upon the soul of man ! Man was a creature of the edge of the forest, perhaps of a savannah-like landscape. He feared the deep forest, he feared the open prairie. Does he not cut down the forest around his home to open it to sunlight? When travelling in the Dakotas does he not seek a tree under which to picnic? Can he bear the solid expanse of concrete and asphalt of the urban horrors which have arisen today? Certainly he is never happy in them, he escapes to the greener expanses of suburbia when he can and is happiest when camping in what he can call the “wilds”. He has come a full circle. From a native home in a world clothed in plants, he came to a utilization of plants which led to vast cities in which man scarcely felt any relationship to plants, so far was he from the land. And now he has returned to a realization that plants are necessary not just for economic values but merely because he feels he needs them and is unhappy without them. This is the age when the pressures upon the plants are so severe that the conscience of man feels the need to grant the plants sanctuaries, when the Nature Conservancy and the Sierra Club become inner needs felt by so many people. They are an expression of the power of plants upon people, a green power whose influence will continue to be needed no matter how far we progress in achievement. When we reach for the stars, plants will ride with us, exchanging oxygen for the carbon dioxide produced by the astronauts, a cycle with which we started our thoughts tonight, the green power which is indispensable to man. THE RELATION OF HENRY JAMES’S ART CRITICISM TO HIS LITERARY STANDARDS Donald Emerson Professor of English University of Wisconsin-Milwaukee Henry James’s use of the representational arts as material for fiction extends from the early “A Landscape Painter” of 1866 to The Outcry of 1911. In the Preface to The Tragic Muse he speaks of the fascination of the artist-life as “a human complication and a social stumbling-block,” the conflict between the claims of art and of society being, he feels, one of “the half-dozen great primary motives.”1 He exploits this subject also in Roderick Hudson and in many of the short stories. Although in fiction he treats problems of the painter, the actor, and the sculptor, his art criticism and his discussions of the problems of representation deal most frequently with painting. The terms of painting enter into his criticism of literature, a great deal of metaphor is drawn from this art, and he alludes frequently to specific canvasses. But as art critic James is an amateur, a lover of painting who could never become the rigorous professional he made himself in his proper field. Mr. John Sweeney, who has collected much of the art criticism, emphasizes James’s reliance on personal impressions in his role of an attentive spectator interested in questions of rep¬ resentation. He delighted in “shows,” and he made his investment of time and interest yield a return for his developing critical sense. As Mr. Sweeney points out, he never concealed the fact that he found literary as well as plastic values in pictures, and uread pic¬ tures with an eye for their possible lurking donnee.”2 James’s criticism of painting is to be found in art reviews and accounts of travel. The reviews appeared from 1868 to about 1882 and again, briefly, in 1897 ; the travel accounts which deal with painting were written in the early 1870’s. Mr. Sweeney has col¬ lected the bulk of the reviews, and Transatlantic Sketches includes most of James discussion of painting in his travel writings. The great difference is that the travel accounts record personal, imme- 1 The Art of the Novel (New York, 1934), p. 79. 8 Henry James, The Painter’s Eye, ed. John L. Sweeney (Cambridge, Mass., 1956), p. 12. 9 10 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 diate responses and discoveries which enriched James’s visits to new places. The reviews are more reserved, but without being impersonal. All the accounts of painting are best seen in relation to James’s criticism in general. In his career James is at first more the re¬ viewer and the critic than the writer of fiction, and the art review¬ ing begins some years later than the book reviewing. One notable difference follows : Where the reviewing of books gives place to extended critical essays in which James eventually speaks with the authority of “a man of the craft,” as he later styles himself, the art notices remain always the appreciations of an intelligent ob¬ server, and eventually cease. James disclaims interest in technical criticism of painting; this is a matter which is to be left to others. There is a certain sort of talk which should be confined to manuals and notebooks and studio records; there is something impertinent in pretending to work it into literary form. ... It is narrow and unimaginative not to understand that a very deep and intelligent enjoyment of pictures is consistent with a lively indifference to this “inside view” of them. It has too much in common with the reverse of a tapestry.3 The “inside view” is precisely what makes James’s later literary criticism uniquely valuable, yet the record of his enjoyment of pic¬ tures has its own interest, for his views on painting are related to his inseparable concerns for the role of the critic and his assess¬ ment of the imagination of the producer in all artistic performance. In his earliest literary criticism, which antedates any of the art criticism, James takes the position that the critic must be opposed to his author, bound to consider the work within the limitations of subject imposed on him, without reference to extraneous theory or critical dogma. He distinguishes between “great” criticism, which touches on philosophy in the fashion of Goethe, and the prac¬ tice of Sainte-Beuve, which at this time he considers productive of “small” criticism. It is, he maintains, the duty of the critic to “com¬ pare a work with itself, with its own concrete standard of truth,” and to rely on his reason rather than his feelings.4 He makes a Coleridgean distinction between imagination and fancy. Imagina¬ tion enables the writer to present recognizably living figures, to whom the imaginative reader can respond; the merely fanciful writer seeks cheap and easy effects because he recognizes no stand- 3 Review of Eugene Fromentin’s Les Mditres d’ Autrefois : Belgique-Hollande, Nation , 23 (13 July 1876), p. 48. 4 Notes and Review by Henry Janies, ed. Pierre de Chaignon la Rose (Cambridge, Mass., 1921), p. 102f. 1969] Emerson— Relation of James's Art Criticism 11 ard of truth or accuracy. “As in the writing of fiction there is no grander instrument than a potent imagination/' James declares, “so there is no more pernicious dependence than an unbridled fancy/'5 Fancy alone may convey the impression of physical sur¬ roundings; the reconstruction of feelings and ideas requires imagination. Within a very few years James modifies this stand and takes a sterner view of the function of the imagination, which he now maintains should “hold itself responsible to certain uncompromis¬ ing realities.”6 Beyond this respect for fact, the imagination, by sympathetic penetration of its subject, can convey the very color of reality. He shortly adds that the working of the imagination is connected with questions of both realism and morality; analytic imagination, presenting a scene with “hard material integrity,” can leave behind a certain moral deposit.7 At the same time he softens the tone of critical authority and calls now for justness of characterization. The day of critical dog¬ matism, he holds, is over, and with it the ancient infallibility and tyranny of the critic. It now seems to him his duty as critic to de¬ tach from a work under discussion “ideas and principles appreci¬ able and available to the cultivated public judgment.”8 At this point James begins his discussions of painting, and it is at once clear that the principles he has formulated for literature are to be ap¬ plied also to painting; justness of characterization of a canvas re¬ quires, quite as much as does analysis of a novel, an estimate of the imaginative force behind its creation. The bases of James's responses to painting and painters are: A distinction between imagination and fancy; a concern for reality; a search for justness of characterization; a fondness for the nar¬ rative or literary aspects of a canvas; and a demand for morality and taste. They all appear in one of the earliest of his reviews of an exhibition of paintings in which he begins a discussion of Alex¬ andre Decamps by naming this painter as representative of the gifted class of artists who pursue effect without direct reference to truth whether it be in literature, music, drama, or painting. Yet he goes on to acknowledge Decamps' “penetrating imagination” as warrant for a background much resembling, so far as it relates to reality, “some first-rate descriptive titbit of Edgar Poe or Charles Baudelaire.” And then James finds that, in default of reality, the somewhat arbitrary and ambiguous air of grandeur and lustre is 6 Ibid., p. 32. 6 “Novels by the author of Mary Powell,” Nation , 5 (15 August 1867), p. 126. 7 Review of Gustave Droz’ Around a Spring, Atlantic, 28 (August, 1871), p. 251. 8 Review of Rebecca H. Davis’ Dallas Galbraith, Nation, 7 (22 October 1868) p. 330. 12 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 the conception, rather, of “a supremely vivid fancy.” In compari¬ son, the juxtaposed canvasses of Eugene Delacroix reveal “a gen¬ erous fallibility which is the penalty of his generous imagination . . . he is a painter whose imaginative impulse begins where that of most painters ends.” It is not that Delacroix selects grotesque or exceptional subjects, but that he sees them rather in “a ray of that light that never was on land or sea — which is simply the light of the mind.” James goes on to describe a picture of men around a campfire, and he finds great fault with the drawing; but in the picture, he feels, Delacroix has shown an eye for the “mystery” of a scene which fuses expression and details into the harmony of poetry. And when it comes to morality and taste, James can describe a Daubiguy canvas as “a little blank and thin; but . . . indefinably honnete” in the fashion of one of George Sand’s rural novels.9 With variations this is the pattern of the early art reviewing, in which James respects the definitions and discriminations he has already made clear in his general critical effort. He deplores, in a picture admittedly painted with precision and skill, the total lack of what may be called “moral atmosphere.”10 He notes that Fox- croft Cole’s pictures have rather less of “an imaginative or reflec¬ tive germ” than suits his taste and finds it a pity that a painter should ever produce anything without suggesting its associations, its human uses, and its general “sentimental value.” He discovers that art is thoroughness and intelligent choice, and that beauty is sincerity; that the artist who would avoid superficiality must deal with the simple and the familiar ; that superficiality is the only vul¬ garity ; and that to be broadly real is to be interesting.11 Returning to Decamps, he finds that his work is rich in “skill . . . invention . . . force . . . apprehension of color . . . and insincerity ,” since his prime warrant is his fancy.12 He feels that Winslow Homer “not only has no imagination, but he contrives to elevate this rather blighting negative into a blooming and honorable positive. He is almost barbarously simple, and, to our eye, he is horribly ugly; but there is nevertheless something one likes about him.”13 He finds an extraordinary impression of “imagination, vigor, and facility,” in the work of Gustave Dore.14 A comparison of the art reviews with the reviews of books and the criticism of authors which James was multiplying at the same 9 “French Pictures in Boston,” Atlantic, 29 (January, 1872), p. 115-118. 10 “Pictures by William Morris Hunt, Gerome, and Others,” Atlantic, 29 (February, 1872), p. 246. 11 “Art,” Atlantic, 29 (March, 1872), p. 372 f. ia “The Wallace Collection in Bethnal Green,” Atlantic, 31 (January, 1873), p. 72 f, is “On Some Pictures Lately Exhibited,” Galaxy, 20 (July, 1875), p. 93, 14 “London Sights,” Nation, 21 (16 December 1875), p. 387, 1969] Emerson— Relation of James's Art Criticism 13 time shows how closely connected are his theories of criticism and of the imagination in all fields. By 1872 he expresses a preference for the method of Sainte-Beuve, whom he once slighted as a “small” critic, over the supposedly scientific method of Hippolyte Taine. While Taine attempts to knock loose chunks of truth with a blow of his critical hammer, Sainte-Beuve rather disengages its diffused and imponderable essence by patient chemistry, by dissolving his attention in the sea of circumstances. James now considers Sainte- Beuve’s provisional empiricism more truly scientific than the pre¬ mature philosophy of M. Taine.15 He begins to revise his own critical practice, and the sympathetic essay on Turgenev of 1874 reveals something of the critical empiricism he praises in the French critic. He finds Turgenev a searching observer, but even more a man of imagination, universally sensitive, who surpasses the French realists in appreciation of sensuous impressions and at the same time appreciates impulses outside the realists’ scope. He discusses Turgenev’s imagination, which he cannot praise too highly for its “intensity and fecundity.” No novelist seems to James to have created a greater number of living figures, to have had so masterly a touch in portraiture, or to have mingled so much ideal beauty with so much unsparing reality.16 Thus it is hardly surprising that at very nearly the same time he can note Winslow Homer’s “perfect realism” while remarking that although Homer is a genuine painter it is not his practice to think, imagine, select, refine, or compose. He goes on in the same review to say of another painter that he lacks intellectual charm, a thing which James finds precious even to its being the only thing of deep value in a work of art, since imagination or intellectual elevation cannot be studied or acquired, whereas everything else can.17 And just as he expresses fatigue that his self-respect re¬ quires his being analytical in observing pictures, he experiences revulsion from literary criticism as he has practiced it. Examina¬ tion of paintings in Italy has persuaded him that the whole history of art is the conscious expression of a single mysterious spirit. He has worked off his juvenile impulse to partisanship, and he now perceives a certain human solidarity in all cultivated effort. “There comes a time,” he confesses in 1874, “when points of dif¬ ference with friends and foes and authors dwindle, and points of contact expand. We have a vision of the vanity of remonstrance and of the idleness of criticism.”18 Within the year he speaks of criticism as “deep appreciation.” 16 Review of Taine’s English Literature, Atlantic, 29 (April, 1872), p. 469. xo French Poets and Novelists (London, 1878), pp. 275, 318. 17 “On Some Pictures Lately Exhibited,” Galaxy, 20 (July, 1875), pp. 91, 93. 18 Review of Victor Hugo’s Quatrevingt-trieze, Nation, 18 (9 April 1874), p. 238. 14 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 During this same time he enlarges his conception of the imagina¬ tion, and of the imaginative force behind artistic construction. Flaubert in Madame Bovary reveals what the imagination can accomplish under a powerful impulse to mirror the unmitigated realities of life.19 Emile Montegut’s “cultivated imagination” gives out in his work “a kind of constant murmur of appreciation — a tremor of perception and reflection.”20 The “true imaginative force” enables Howells to give his readers not only the mechanical structure of a dramatic situation, but also its atmosphere, mean¬ ing, and poetry.21 James cites also such negative examples as Charles Kingsley, whose imagination died a natural death when Kingsley turned didactic historian,22 and Bayard Taylor’s, which was so cold it could not kindle the reader’s.23 When in 1875 he dis¬ cusses Balzac extensively for the first time his chief concern is the quality of Balzac’s imagination, and in later essays he returns to it again and again. It becomes for James the great explanatory fact behind Balzac’s reality, his vividness, and his systematizing of the Comedie Humaine. Its deficiences explain Balzac’s failures of por¬ trayal whenever he attempts to touch the moral life.24 This discussion of literary matters, which deliberately departs from James’s concern with painting, serves two ends : It shows the inseparable connection of his changing conceptions of the critic’s role in his responses to both books and pictures, and it underscores his developing sense of the crucial role of the imagination in all artistic production. And since it deals with the formulations which are most explicit in his writings of 1872 and 1875 it encloses, as in a parenthesis, the bulk of the travel accounts subsequently col¬ lected as Transatlantic Sketches. It is no way surprising that these accounts reflect also, with the intensity of vivid, immediate experi¬ ence, perceptions and responses to pictures which James had learned elsewhere. The travels reinforce one’s inescapable sense of connection in everything James wrote. He rhapsodized in 1873 on Tintoretto, before whose work he feels old doubts and dilemmas to evaporate and the conflict of idealism and realism to be practically solved. That earlier sense which led him to declare that a scene presented with hard material integrity could leave behind a certain moral deposit now makes 19 Review of Gustave Flaubert’s Temptation of St. Anthony, Nation, 18 (4 June 1874) , p. 365. "Review of Emile Montegut’s Souvenirs de Bourgogne, Nation 19 (23 July 1894), p. 62. 21 Review of William Dean Howells’ A Foregone Conclusion, Nation, 20 (7 January 1875) , p. 12. 22 “Charles Kingsley,” Nation, 20 (28 January 1875), p. 61. "Review of Bayard Taylor’s The Prophet: A Tragedy, North American Review, 120 (January, 1875), p. 193. 24 French Poets and Novelists, p. 114. 1969] Emerson — Relation of James’s Art Criticism 15 him speak of Tintoretto as “the most interesting of painters,” whose indefatigable hand never drew a line that was not “a moral line.” Tintoretto’s great merit, to James’s mind, is his unequalled distinctness of vision : “When once he had conceived the germ of a scene, it defined itself to his imagination with an intensity, an amplitude, an individuality of expression, which makes one’s observation of his picture seem less an operation of the mind than a kind of supplementary experience of life.” Veronese and Titian, by comparison, seem to James to be content with much looser specification, so that to place them against Tintoretto is to measure the difference between observation and imagination. Tintoretto grasped the whole scene in his great dramatic compositions, and his work conveys the impression that “he felt, pictorially, the great, beautiful, terrible spectacle of human life very much as Shakespeare felt it poetically.”25 The justness of characterization which James demands in literary criticism is again, in his observations on painters and painting, satisfied only with an account of the artist’s imagination. He proceeds even by negative example with Domenichino, to him a supreme example of “effort detached from inspiration . . . school- merit divorced from spontaneity” for the production of examples of how the artist must never paint. The intensity of James’s feel¬ ing is apparent in his introduction, into a travel account, of a fic¬ tional character, the head-master of a drawing academy who sadly leads his pupils to the disheartening examples of this painter’s work and explains, “Domenichino had great talent, and here and there he is an excellent model ; he was devoted, conscientious, observant, indus¬ trious ; but . . . his imagination was cold. It loved nothing, it lost itself in nothing, its efforts never gave it the heart-ache. It went about trying this and that, concocting cold pictures after cold receipts, dealing in the second-hand and the ready-made, and putting into its performances a little of everything but itself.”26 The same type of discrimination appears in James’s discussion of Sandro Botticelli, whom he finds, in a certain way, the most interesting of the Florentine painters. Although he acknowledges indebtedness to Walter Pater he resolutely puts aside all that he considers recondite in Pater’s interpretation of Botticelli and pro¬ ceeds in typical fashion to conclude, “A rigidly sufficient account of his genius is that his own imagination was active, that his fancy was audacious and adventurous. Alone among the painters 33 Transatlantic Sketches (Boston, 1875), p. 90 ff. 26 Ibid., p. 177. 16 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 of his time, he seems to me to possess invention .” Where the glow of expanding observation sent Botticelli’s contemporaries to their easels, Botticelli possessed a faculty which loved to play tricks with the actual, to sport, wander, and explore on its own account.27 James’s individual judgments of painters or canvasses, of both which numerous critics can give more just accounts, are less inter¬ esting than the bases of his judgments and their relations to his literary criticism. Early and late examples in both fields show his conviction that the quality of imagination in a work of art is all- important. It is the power, he now feels, to conceive greatly and to feel greatly, to organize irreproachably the work of art of what¬ ever kind, and to make it a kind of supplementary experience of life. Of literature James speaks with an authority that is wanting in his criticism of painting, for in dealing with books he can make the kind of technical analysis he eschews in his reports of painters and their work. He can thus, in 1877, condemn Whistler’s work as unprofitable and uninteresting, and at the same time praise the work of Edward Burne-Jones as having, for all its faults, “an amount of imaginative force the mere overflow of which would set up in trade a thousand of the painters who are more generally accepted by the public.”28 Again, in the same year he can declare that “a picture should have some relation to live as well as to paint¬ ing. Mr. Whistler’s experiments have no relation whatever to life ; they have only a relation to painting,” while he praises the art of Burne-Jones as the art of culture, reflection, intellectual luxury, and aesthetic refinement, the art, in short, “of people who look at the world and at life not directly . . . and in all its accidental reality, but in the reflection and ornamental portrait of it fur¬ nished by art itself in other manifestations; furnished by litera¬ ture, by poetry, by history, by erudition.”29 James much later confessed to Charles Eliot Norton that he had come to find the work of Burne-Jones uninteresting,30 but the changed view is of less significance than that James’s insistence on relations in art leads eventually to his conception of criticism as in part the study of connections. Guided himself by the practice of Sainte-Beuve, in 1880 he praised the Frenchman’s sense of his role: “The critic, in his conception, was not the narrow lawgiver or the rigid censor that he is often assumed to be ; he was the stu¬ dent, the inquirer, the observer, the interpreter, the active, inde¬ fatigable commentator, whose constant aim was to arrive at just- 27 Ibid., p. 300. 28 “The Grosvenor Gallery and the Royal Academy,” Nation, 24 (31 May 1877), p. 320. 29 “The Picture Season in London,” Galaxy, 24 (August, 1877), p. 156f. 30 Letters, ed. Percy Lubbock (London, 1920), I, 341. 1969] Emerson — Relation of James's Art Criticism 17 ness of characterization.”31 Four years later he says, “The meas¬ ure of my enjoyment of a critic is the degree to which he resembles Sainte-Beuve.”32 James’s experience as writer inevitably affected his criticism; he more and more cited his own authority. One such authoritative pronouncement is “The Art of Fiction,” of 1884, which charac¬ terizes the novel as a direct impression of life, the value of which depends upon the intensity of the impression. The writer must work from reality and experience, but reality has myriad forms, and experience is never complete. “It is an immense sensibility . . . it is the very atmosphere of the mind; and when the mind is imaginative ... it converts the very pulses of the air into revela¬ tions.” “Imagination assisting,” the artist can deal with anything, for experience is practically constituted of the gifts which are designated as imagination: “The power to guess the unseen from the seen, to trace the implications of things, to judge the whole piece by the pattern, the condition of feeling life in general so completely that you are well on your way to knowing any par¬ ticular corner of it.”33 This declaration explains why in James’s criticism in all fields the imagination is so emphasized, why it is the ground of so many of his discriminations, and why he insists upon a description of the artist’s imagination as part of the discussion of his work. With his enlarging view of criticism as practiced by Sainte-Beuve, James is shortly to remark that works of art grow more interesting as one studies their connections, this study being a function of intel¬ ligent criticism.34 He goes on to insist that everything depends on the qualifications of the critic. “Curiosity and sympathy” form his equipment. To lend himself, to project himself and steep himself, to feel and feel till he understands and to understand so well that he can say, to have perception at the pitch and passion and expres¬ sion as embracing as the air, to be infinitely curious and incor¬ rigibly patient, and yet plastic and inflammable and determinable . . . these are fine chances for an active mind.35 He characterizes himself when he speaks of the critic who has no a priori rule but that a production shall have genuine life.36 31 “Sainte-Beuve,” North American Review , 130 (January, 1880), p. 56. 32 “Matthew Arnold,” English Illustrated Magazine, 1 (January, 1884), p. 242. 33 Partial Portraits (London, 1888), pp. 387, 389. 34 Essays in London (London, 1893), p. 160. 35 Ibid., p. 276. 36 Views and Reviews (Boston, 1908), p. 227. 18 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 James is well known to have remarked rather testily, “Nothing is ever my last word about anything,” but for a variety of reasons he gave over the criticism of painting, and one of his last words has an almost valedictory note. It is his mature expression of the importance which through his life he attached to art, the em- balmer, the magician whom we can never speak too fair or whose importance overstate. For art “prolongs, it preserves, it conse¬ crates, it raises from the dead. It conciliates, charms, bribes pos¬ terity ; and it murmurs to mortals, as the old French poet sang to his mistress, ‘You will be fair only so far as I have said so.’ ”37 This may belong to the realm of deep appreciation, but it is no longer criticism. Aside from the stresses of James’s desperate pre¬ occupation with the stage between 1890 and 1895, his return from defeat there to the magic of his “own old pen,” and his disgust with all journalistic practices and activities, is there not also a partial explanation of James’s ceasing to write on painting in his inability to conduct the kind of analysis which, with novelists, he increasingly made a part of his criticism, and with no one so much as himself? All the last critical essays bear a family resemblance, and the artistic problem is always the general subject, as it is in the Prefaces for the collective New York edition of James’s own work. A final illuminating statement of the office and the effect of criticism completes the perspective of James’s changing views and implies his neglect, in the late criticism, of painting. Painting had been one of the great resources of his imaginative life, but he could not write of it as he could of literature, the field in which he spoke with the authority of the high title he gave himself as “a man of the craft.” The effect, if not the prime office, of criticism, is to make our absorption and our enjoyment of the things that feed the mind as aware of itself as possible, since that awareness quickens the mental demand, which thus in turn wanders further and fur¬ ther for pasture. This action on the part of the mind practically amounts to a reaching out for the reasons of its interest, as only by its so ascertaining them can the interest grow more various. This is the very education of our imaginative life ... we cease to be instinctive and at the mercy of chance.”38 In its scope and development James’s criticism reveals the growth of an artistic mind of high quality, and the evolution of his standards explains the changing estimates he made of painters and 37 picture and Text (New York, 1893), p. 134 f. 38 Notes on Novelists (New York, 1914), p. 315. 1969] Emerson — Relation of James’s Art Criticism 19 writers. This itself is sufficient ground of interest, his views of the painter’s art forming a long and interesting chapter in the whole volume. The early advocate of science and logic turns from judg¬ ment to justness of characterization and at last to deep apprecia¬ tion, with his final word a demand that criticism promote the edu¬ cation of the imaginative life itself. A good deal of James’s artis¬ tic education came from pictures, and he was deeply responsive; but as critic he was authoritative only in his own productive field. VIOLENCE AND SURVIVAL IN THE NOVELS OF IRIS MURDOCH Donald Emerson Professor of English University of Wisconsin— Milwaukee Iris Murdoch is a contemporary Anglo-Irish novelist and philoso¬ pher who besides writing a monograph on Jean-Paul Sartre (1953) continues to lecture and to produce papers in both her professional fields since her resignation from Oxford. Beginning in 1954 she has published eleven novels, none of which can properly be termed a philosophical novel though each is related to the problems of con¬ temporary philosophy and to Miss Murdoch’s own developing thought. She is an artist who believes that men are social creatures who confront the intractibility of a contingent world in which their concerns are not abstractions but personal relationships and the confusing, unpredictable “stuff” of human life. She writes in full awareness of the state of fictional art, with an inexhaustible in¬ ventiveness, humor, irony, and compassion ; and she has a high re¬ gard for the 19th century novel of character and plot although she has attempted it less frequently than the patterned novel which tends to abstraction and symbolism. The novel is one of the contemporary modes of philosophy, and Sartre has gone so far as to declare it a chief mode of expression for his brand of existentialism. As a form it reaches an audience indifferent to other statements, and whether existentialist or not has a persuasiveness more powerful than discursive exposition. It expresses more sensitively than other forms the motives and as¬ sumptions of men in action or, in some of its guises, the convictions, amounting to assessments of the culture of their times, of its authors. The novel is one of the chief cultural documents of the age, and Miss Murdoch considers it “a picture of, and a comment upon, the human condition.” She regularly directs her attention to the principal strategies of the novel in our time. In her criticism and her practice she distin¬ guishes four chief modes: the journalistic novel of thin characteri¬ zation and abundant detail ; the realistic novel of character which is closely related to the great 19th century examples of, say, George “Violence and Survival in the Novels of Iris Murdoch” was read at the meeting- of the Wisconsin Academy in Eau Claire, May, 1968. 21 22 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Eliot or Tolstoy ; the symbolist, almost allegorical novel of the type of William Golding’s The Spire, which she terms a “crystalline” form; and the fantasy-myth which she herself produced brilliantly in her first novel, Under the Net (1954) . Although much of the best modern work has been done in the symbolist novel, she regards the form with misgivings prompted by her conviction that interest in issues rather than people is inappropriate in the novel. To her thinking the novelist proper is “a sort of phenomenologist ... a describer rather than an explainer,” whose eye should be fixed on what we do rather than on what we ought to do. She thus insists on “the stubborn irreducibility of persons” and the contingency of experience, and finds Sartre’s impatience with the “stuff” of human life crippling to his work. Miss Murdoch is herself victim of the paradox which confronts every novelist. On the one hand there is the bumbling confusion of human personalities and relationships, on the other the demands of the novel for some degree of formal coherence. The form has swung between the wonderful, lifelike record of contingency which made Henry James refer to the 19th century English novel as “the para¬ dise of the loose end,” and the contemporary tight, structured form which results in what Miss Murdoch terms “dryness”. Against such dryness she argues that the novelist must portray individuals who are independent of their author and are not puppets to exteriorize some psychological conflict of his own. As she also says, “A novel must be a house fit for free characters to live in; and to combine form with respect for reality with all its odd contingent ways is the highest art of prose.” She demands respect for contingency, yet she writes novels which have sometimes prompted her critics to draw diagrams. In practice she has resolved the paradox of com¬ position with varying degrees of success in fictions which tend to be well-structured although her characters are believably un¬ predictable and her own sense of the ridiculous conveys a lively sense of their contingent world. Some further preliminaries must be dealt with : Miss Murdoch’s philosophical ballast; the relations between her work and that of others; and her own development as a novelist. As an unreformed academic she naturally draws upon her professional knowledge of philosophy when she works as a novelist. In an excellent study, Degrees of Freedom, Mrs. A. S. Byatt documents Miss Murdoch’s indebtedness, sometimes in opposition, to Sartre. Wittgenstein’s net of concepts furnishes the dramatic metaphor of the hero’s pre¬ dicament as well as the title of the first novel. Simone Weil’s life and work illuminate, for example, the portrayals of suffering in The Flight from the Enchanter (1956) and The Unicorn (1963). 1969] Emerson — Novels of Iris Murdoch 23 There are allusions to Kant and Hegel, and to the religious con¬ cepts possible to a novelist who terms herself a Protestant “Chris¬ tian fellow-traveller.” Yet Miss Murdoch says there is little con¬ nection between her books and her academic thinking despite the fact that their shape and moral bases owe something to philosophy. When she published her earliest novels she was bracketed by critics with such Young Angries of the 50’s as John Wain and Kingsley Amis, and her third novel, The Sandcastle (1957), dis¬ appointed those who expected her to continue in a single vein. A Severed Head (1961) was a further surprise which had even some success of scandal— -How far could she go? — and was likened by critics to the Restoration drama of Congreve. An Unofficial Rose (1962), in which she made an ambitious attempt to regain the advantages of the 19th century novel of character led to her being categorized as a “lady novelist,” a term which was repeated with all its denigrative connotations for The Nice and the Good (1968), whatever it may mean. Iris Murdoch is well read in her profession of novelist as in her profession of philosopher, and awareness of the art affects her work much as does her knowledge of philosophy — there is only partial connection between the work she does and what, as critic, she approves, yet the shape of her novels owes much to her critical understanding of the art and of the developments which have brought it to its present state. She began with two novels best described as fantasy-myths, the first concerned with freedom for the individual caught in the conceptual net, the second with the meanings of power in the modern world. Both are social novels, both owe something to existentialist thinking, and both maintain a brisk pace in which respect for contingency leads at times to por¬ trayal of wild and hilarious improbabilities. Since then Miss Mur¬ doch has chosen to deal more frequently with inter-personal rela¬ tions, an emphasis which recalls that of Henry James. Her most successful novel, The Bell (1958), has the solid life which Miss Murdoch praises in the great 19th century novels, for it best dis¬ plays the “real apprehension of persons other than the author as having a right to exist and to have a separate mode of being which is important and interesting to themselves.” Other novels of this type are The Sandcastle (1957), An Unofficial Rose (1962), The Red and the Green (1965), and The Nice and the Good (1968). A Severed Head (1961) is strikingly different from the other novels in style and plot, and is an apparent attempt of Miss Murdoch’s to make a comic substitution of the “hard idea of truth” for the “facile idea of sincerity” with which, to her mind, both Freud and Sartre are associated; and The Italian Girl (1964) is a feebler example. There is one further division within the work: The Uni - 24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 corn (1963) is a return from the realistic complexity of The Bell type of novel to the patterned, mythic novel, no longer fantastic in the fashion of the earliest novels, but stripped and contrived, as is The Time of the Angels (1966). Mrs. Byatt has found the central concern of Miss Murdoch’s novels in the question of freedom, and it is an illuminating ap¬ proach; but other themes deserve analysis, among them violence and survival. Four exemplary novels will serve: Under the Net, The Bell, An Unofficial Rose, and The Time of the Angels. They happen to have appeared at precise four year intervals since 1954, but they are not strikingly better than others for illustrative pur¬ poses. They might equally be used to define such other recurrent themes as the nature of love ; the relation between love and moral¬ ity; the recognition of reality, especially the reality of other per¬ sons ; or the conflict between reality and illusion. Outward violence in Miss Murdoch’s work ranges from simple theft to murder or suicide on the scale of action, from impulse to premeditation on the scale of intention, from accident to catastro¬ phe within the workings of nature; but the inward violence of aggression, subjugation, and enslavement is frequently far more interesting than any outward event, as is the inward struggle for freedom. The terms of survival encompass the degrees between the unthinking safety of self-chosen ignorance, the achievement of stoic endurance, and the liberation of self-awareness. But for Miss Murdoch’s characters survival is not always possible nor even, sometimes, desired. Under the Net is the earliest of the three novels Miss Murdoch has attempted in the first person, and from a male point of view. It is a lively romp, wildly improbable at times, which conveys an encompassing sense of London and, in a different way, of Paris. Jake Donaghue is the ideal narrator for a piece filled with action and ideas, for the substance of his life is the private conversation with himself of a man who sees too much ever to give a straight answer. Besides, in his half-outsider fashion he is as much a phi¬ losopher as Iris Murdoch. Beneath the tumultuous action there is a seriousness which makes Jake’s final grasp of a direction for his life meaningful and convincing, but there is none of the grimness of final choices apparent in some later novels, none of the sense of harried individuals pushed to intolerable limits. There is even a good-natured quality about the violence, which occurs in great variety. Why shouldn’t a sensible man carry a pick-lock and take over his friends’ apartments when locked out of his room for non¬ payment? Why shouldn’t he, when his manuscript has been stolen, steal a valuable dog which the thief has kidnapped? Or help his 1969] Emerson — Novels of Iris Murdoch 25 friend break out of a hospital? Or escape from a riot in a film studio by blowing a hole in the set? Or use a powerful detonator of the type conveniently at hand in well-furnished apartments, to blow open a wall safe? Jake’s sense of the passing of time, how¬ ever, at last turns him to other courses. All work and all love, the search for wealth and fame, the search for truth, life itself, are made up of moments which pass and become nothing. Yet through this shaft of nothings we drive on¬ wards with that miraculous vitality that creates our precarious habitations in the past and the future. So we live — a spirit that broods and hovers over the continual death of time, the lost meaning, the unrecaptured moment, the unremembered face, until the final chop-chop that ends all our moments and plunges that spirit back into the void from which it came. Jake turns from the hand-to-mouth survival of his first thirty years to the possibility of writing well, from the “ragged, inglori¬ ous, and apparently purposeless” life he has known to the possibil¬ ity of doing better work than in his first book, with a strength and joy which make the moment “the morning of the first day.” His survival is not endurance but a plunge into a life made new by being newly understood. And he achieves the understanding for himself. Another novel filled with the odd contingencies of life followed Under the Net, though more sombre in tone. Miss Murdoch then turned to a less highly charged picture of domestic life and per¬ sonal relations, thence to The Bell, her best piece thus far. The action of The Bell centers about an Anglican lay community and the disruptive events which lead to its closing after a public scan¬ dal. It is far too rich a novel for easy summary, but the violence of much of it indicates the direction which Miss Murdoch is tak¬ ing: From the casual, almost merry violence of the first novel she has turned to more consequential acts, not because suicide is more desperate an act than safe-cracking but because all the violence is seen more meaningfully in relation to the values of religion and philosophy. There is more depth to this novel than to the first, and the study of violence in still later novels will show a progression already apparent here. Yet the two principal characters are far less intelligent than Jake Donaghue. Dora Greenfield is neither talented as an artist nor gifted with common sense even in ordinary affairs ; and Michael Meade is a weak, homosexual ex-schoolteacher who fails in all relationships through fumbling attempts at tender- minded goodness. It is Miss Murdoch who is intelligent in this novel, though with that respect for her characters as having lives of their own which she considers typical of the great novelists. 26 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Besides suicide, attempted suicide, adultery, and self-righteous bullying there is the more subtle violence, increasingly apparent in Miss Murdoch’s work, of the sins against love : indifference, failure of feeling, and calculated betrayal. It is these which are destruc¬ tive; the violence is their product. And the terms of survival have changed. For Dora and Michael there is no sudden life-enhancing vision such as Jake Donaghue experienced; instead, there is a slow progress and adjustment to their changed lives. Michael observes that the events at Imber Court increase Dora’s substance ; there is simply “more of her” after the dreadful events have passed. Michael himself, in anguish over the death of a man he might have saved, defeats thoughts of suicide by perfecting his suffering through responsibility for the dead man’s half-mad sister. At last he, like Dora, turns to the hope of life when he can “experience again, responding with his heart, that indefinitely extended re¬ quirement that one human being makes upon another.” After The Bell Miss Murdoch produced a witty, brilliant study of sexuality in A Severed Head, but with An Unofficial Rose she returned to what she terms “the novel proper,” and here there is far less of the strangeness, amounting at times to the effect of en¬ chantment, elsewhere cast about her characters. Miss Murdoch’s characters are never ordinary but they are sometimes fantastic; the figures of An Unofficial Rose are believable without being com¬ monplace. More of this novel is related to the linked problems of violence and survival than any other, yet the surface is compara¬ tively placid. There is one tumultuous scene and at least two sym¬ bolic murders, but with her shift of interest from social to personal relations Miss Murdoch here subdues violence to those crimes only possible between persons who have loved or, what is worse, have failed of love. There is Hugh Peronett, who is willing to sell a pre¬ cious picture to finance his son Randall in an adultery the like of which Hugh was never bold enough to attempt. The selfishness of Randall includes a self-justifying and coldly rationalized hatred of his wife Ann, at the same time that his acceptance of Hugh’s money is a joyful symbolic murder of his father. The daughter of Ann and Randall destroys her mother’s possibility of happiness by ruthless deception which prevents Ann from ever enjoying the love of a good man who has waited until Randall deserted her. Even Hugh’s mistress of long ago manipulates Hugh, Randall, and Ran¬ dall’s new mistress with devilish skill. Yet they are not monstrous, however they may seem, and they survive in their varied fashions beyond such difficulties as are always, one character remarks, solved by violence. For some there is simple forgetfulness — the young will find other interests. For Hugh and his failing mistress 1969] Emerson— Novels of Iris Murdoch 27 there is the anticipation of an early extinction which leaves Hugh thinking only that a brief interval remains : “Perhaps he had been confused, perhaps he had understood nothing, but he had certainly survived. He was free. 0 spare me a little that I may recover my strength; before I go hence and be no more seen.” Randall is simply left in a besotted state with the mistress whom he may decide, when sober, to leave. For Ann there is endurance in her ignorance of those she has never known. “She had not known them. She did not know herself. It was not possible, it was not necessary, it was perhaps not even proper. . . . Tasks lay ahead, one after one after one, and the gradual return to an old simplicity. She would never know, and that would be her way of surviving.” In her next novel, The Unicorn , Miss Murdoch turned more di¬ rectly to the problems of suffering and endurance, but in a form radically different and equally exemplified later in The Time of the Angels. It has been noted that her thought can be related to that of other philosophers and her fictions to those of other novelists. The Unicorn suggests the strangeness of Sheridan Le Fanu while its subject is related to the anatomy of suffering diagrammed by Simone Weil. Both influences are apparent in The Time of the Angels, set in London but a corner of London isolated amidst bombed-out acres, and fog-shrouded into a remoteness as strange as that of any Irish coast. These two novels are departures from the realism to which Miss Murdoch feels the novel must return to recover its vitality. It is as though the modern fascination with myth-making, abstraction, and the creation of patterned fictions has some sort of irresistible appeal. Or possibly Miss Murdoch finds realism in some ways inadequate and unsatisfying and seeks some¬ how to get more directly to the core of reality by rejecting com¬ monplace actuality, “a world in which people play cricket, cook cakes, make simple decisions, remember their childhood and go to the circus; not the world in which they commit sins, fall in love, say prayers, or join the Communist Party.” She has moved, as has been seen, from Jake Donaghue’s tumultuous social world to the portrayal of personal relationships, thence to ever-tighter sets of relationships in progressively restricted groupings of characters. This has been accompanied by a deepening of meaning as she has neglected the everyday world in which people play cricket and cook cakes for the more intense world in which, if they do not pray, they are concerned with God, man, suffering, and evil. And violence has become ever more closely identified with evil, while suffering has acquired an ultimate redemptive power. It is only fair to note that the latest novel, The Nice and the Good (1968), returns, with some modifications, to the realistic mode. 28 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 The Time of the Angels actually contrasts the commonplace world with the enclosed, fantastic rectory dominated by the mad atheist priest Carel Fisher, for into it intrude Carers younger brother Marcus and the one-time mistress of Carel’s older brother Julian, who committed suicide after Carel had seduced Julian’s wife. The weird household includes Muriel, Carel’s daughter by his deceased wife ; Elizabeth, the daughter of that adultery, with whom Carel commits incest; and Pattie O’Driscoll, Card’s mulatto mis¬ tress and housekeeper. Other servants are a father and son, a refugee pair, of whom the son is the more interesting because he is a pathological liar. All is not the grimness of American Southern Gothic with which in subject matter this novel surely could com¬ pete; there are scenes and encounters as funny as some in Faulk¬ ner, if sometimes equally macabre. Much of the violence is in the past, and if theft, incest, and suicide are in the foreground, the meaning of violence has changed. Carel Fisher is a Dostoevskyan character sunk in debasement and at the same time a religious seeker for whom God is dead. Not even evil is real to him any longer. “There is only power and the marvel of power, there is only chance and the terror of chance.” Carel’s most significant act of violence is a blow to his brother’s face, for he is persuaded now that only infliction of pain can prove the existence of others. When his daughters discover the truth of their relationship and when Pattie cannot remain with Carel, though she loves him, he kills himself. And now it is not a question of survival so much as of suffering. Pattie’s love is to be her own torment only. Muriel, watching her father die, realizes she is “condemned to be divided forever from the world of simple innocent things, thoughtless af¬ fections and free happy laughter and dogs passing in the street.” She is bound to Elizabeth, and they will be each other’s damnation. The ultimate violence and perversity of both The Unicorn and The Time of the Angels belong to the literature of extremity, the very thing which Miss Murdoch had avoided in her earlier work. She has already gone on to another portrayal of a more easily recognizable world in The Nice and the Good, but it is a world in which, as before, violence is a fact to which survival or destruction are alternatives. These themes will surely recur in the further work of this endlessly Inventive and interesting novelist. They are cen¬ trally related to her conception of the human condition, and despite her respect for a contingent world and irreducible persons Miss Murdoch seems increasingly drawn to the symbolic novel in which emphasis on violence and suffering is greater than in portrayals in the realistic mode. LIFE AGAINST DEATH IN ENGLISH POETRY: A METHOD OF STYLISTIC DEFINITION* Karl Kroeber with Alfred L. Kroeber and Theodora K. Kroeber Our purpose in this paper is to illustrate a method of defining configurations of literary style through the study of word-choice patterns in poetry. Refined and extended, this method should make possible more meaningful analyses of poetic movements and counter-movements both within and across the conventional classi¬ fications of stylistic periods— neo-classic, romantic, modern, and the like. We began by counting words sure to have significance in poetry, words such as nature , soul, spirit, and words referring to the emo¬ tions, the seasons, and so forth. We soon found ourselves over¬ whelmed by a wealth of possible directions and significances, so we settled on a pattern of life and death words as a starting point, examining four life words and four death words in thirty-five poets, both British and American, beginning with Chaucer and concluding with James Dickey, for a time span of nearly six hun¬ dred years. Our technique is to determine the frequency of the same set of words in (where feasible) the total work of each poet. This sort of survey of course turns for its data to the standard concordances, whose value has too often been underestimated. A mass of ex¬ tracted and ordered stylistic information lies shallowly buried in every concordance, and this information may be illuminating and significant in many diverse ways. For instance, Housman has much to say of the soul, which he mentions thirty-four times in his seri¬ ous verse as against twelve mentions of flesh. But his concordance also shows that he does not once use either the word spirit or the word body — a fact that might easily escape the most devoted stu¬ dent’s observation. Yet this “negative fact” is essential to our * The work reported on in this article was begun by my father, the late Alfred L. Kroeber. Always interested in literature, he became especially intrigued during his later years with problems of describing literary styles. This essay represents one of the last scholarly investigations he initiated. Recent developments in techniques for using computers to compile concordances make it feasible now to carry out systemati¬ cally the extensive stylistic analyses my father first envisaged nearly a decade ago. Karl Kroeber 29 30 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 understanding of the function of soul and flesh in Housman’s poetry. There are as yet, unfortunately, no concordances for some ma¬ jor poets and still very few for minor poets. We are reluctant to base findings on a sample which will someday be superseded by a concordance : the latter is not only complete but also far more re¬ liable. We did, however, make some counts of life and death words to give broader representation to our list. Except for Swinburne, our samples are probably adequate, covering approximately a moiety of each poet’s work. But these results are strictly provi¬ sional, both because they are not based on total output, and because a running tally is almost surely less accurate than a formal con¬ cordance count. Even these provisional findings, however, are adequate to illus¬ trate the nature of our method and (we hope) to encourage others to undertake analogous studies. In work of this kind results are in large measure additive. Indeed, it is our premise that only through the accumulation of many investigators’ findings will such quanti¬ tative discriminations lead to deeper understanding of the qualities of literary style. We counted the nouns death and life, including of course their plurals and possessives; the verbs die and live, including conjuga- tional forms such as dies, dieth, died; the participles dying and living; and the adjectives dead and alive. These eight have the merits of being simple and unescapable Anglo-Saxon, and the four of each set derive from one root. This last is not always the case- — thus in German, death is Tod, but the verb for die shifts to sterben, the Tod-derived verb tbten being used for kill. Some features of our limitation to these four pairs are admit¬ tedly arbitrary, but we have made the limitation in the interest of avoiding complications. Thus live as an adjective may have been used more frequently by some poets than alive, but some concord¬ ances do not distinguish parts of speech, even mechanically group¬ ing the verb lives with the plural noun lives. Too many re-orderings of concordance listings would have introduced a high degree of error into work where at best mistakes are easy. Quick as a syno¬ nym might also have come into consideration, especially in earlier poets. Deadly would perhaps have been desirable to include, but its counterpart lively has moved out of the range of the correspond¬ ing meaning. Mortal might well have been significant, and one could argue plausibly that the contrast of birth to death would be at least as meaningful as that of life to death. The terms we chose are certainly not exhaustive, but one step at a time seemed wisest as a beginning, with consideration for consistency and for equal 1969] Kroeber — Life Against Death in English Poetry 31 pairing on the life and on the death side. The final goal of this method is the creation of a series of something like “semantic fields” upon which to base stylistic judgments. Our eight words, then, should be regarded as the first segment of such a field. The results of our count of the concordances plus our sample counts are given in Table 1. Fourteen of the poets show an excess of life words, twenty of death words. Collins, whose volume is tiny, splits evenly. The general division, then, is not a half and half one, and it is not random. Before 1815, and especially from Milton on, most poets give preference to life; after 1815 death takes the lead. Although clusterings seem most significant, the inevitable individual exceptions are interesting. The results of our counts have throughout been converted into percentages to make them comparable. The absolute numbers in the last three columns of the table are the totals of our counts, included as indication of the reliability of the percentage figures. We cannot invest with much significance the grand total. About all that can be ventured from our absolute figures is that, for the six centuries, life and death words are in approximate equality, and that the total number of poets favoring one or the other may be expected to be more or less in balance when it shall have become possible to make a complete count of all of them. The significance of variability in our count lies in periods and in individuals, not in grand totals. We have not yet attempted the other obvious “totaling” test — determining how large a part life and death words play in the total vocabulary of each poet. The series begins with Chaucer, our only Middle-English poet. The concordance included but we excluded from his count the Boethius and the Parson’s Tale as being in prose and The Romaunt of the Rose as being an outright translation. Their inclusion, as a matter of fact, would not very materially change his proportions — by two percent only— but it would be inconsistent with the method of our other counts. In Chaucer we discover a reasonable balance between life and death, the death words being weighted over the life ones by the small percentage of fifty-four to forty-six. Such a balance sems proper to Chaucer, robust and sanguine of temperament as he was but nonetheless a poet not yet out of the Middle Ages and their preoccupation with the after life, and writ¬ ing a scant century before the addiction of popular North Euro¬ pean art to the macabre Dance of Death, the carrying off of the damned, and their torments in hell. Chaucer’s life and death word preferences lie near the presumable English poetical mean, to judge by our other counts: small but pleasantly corroboratory evi¬ dence for the view that Chaucer was already within the generic Table 32 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Number of Occurrences Total lsOOO\D^OO>AKKO'-00-''t'l'^0'OC'tONO"+NOOO'ONTfO''tTt'N^N-N^ vDK'ANl\vDtsNTfOO'tN>0'^00'AlNO^NOOOClC'l^rANO'AC'l^NOOV' — ^FAvOOO rS — « — tj* Death 00OC‘f'O'--,'^f^N>0Or^(St^v0l^'-0C\0NO001A00-<)'iAts' Tf r<-\ — ■ — — < *— tJ< — — > Life vD — N’tO't(J'MS\Of^>AOONOl''0'iv'OOts'A-'>^lsOO\0'DNOr,'r^>AN^Ol''00'f^ OO^OO'Nst^.^'K^OO— <^Ot^tN?Nr^>J^O'fS00\O>J^tN.0000f,'vDO''C)^''— 00O'CS(N-^,t\O'sO — 00 VN — O' sO — — 1 C-) W-\ rS CS (O- r<-\ 04 — — — 00 04 0C eo rj- o4 — • — 04 — — 04 Percent Death Total T)10^'-O^Nv0^vOONK^OO^OO''taONOv-0N^^NGNfA^vD-K-N-^a^ Dying OOo\— 'rf — o4— '04o4'T-,To4o4— 'for'-\o4roo)''t',Tw-\.— i/^o4r'>o4^troroo4'T'3"V>cooO— 'ro Die o4roroo4000.roOsO'0'NC>'— (^nDO'1^ — 'ONV'U'NWr^'^'tsn'tTfNOO Dead — N^)- — 'O|A0"tN0'00 00IS't^0'-'C'r^Nl^r"vDiAf^0'0"''l^00 001''O1''Ov0 — Death OOr^f^^O-^VNNO — ro — 04 co — ■T)T^r^OTj'NNOOOOr,'OOTfOQOr^O'vOr^OO'tO'i‘^,'000 — ■ 04 — • — NNN- 'NMN - 04 — 1 — — 04 — — 1 — — ' 04 04 _ — __o>— '04O4O4 — — <04''T — u u. Total vOOvOO'OvOr<>i,t\’^ — p^vOOO^O'-vO'-'- — Ov — — K^,^0'fAOsf/'Ov'0^vO Living NN — 'ilf^r^NMOOOXI'OOOO — NN — H z u o £ Live roororo>ooO''*>oCJ'0'OC'£>toco— < sO to >o 04 io — rfo— ■'t^'^'OKt^O'C'ONsO'O — O' CL | Alive «-VA^NNNN0^0NN00'^N0fANrN-^0^^^-NfAN^-'-,1tNN'AfAiA Life Or^OOWOf^'^KKN'rO' — N-^'tNOr^ — ONr>OOOC',J'f^ — ’-,'fl,^0'N^"AO't'0 r^r 89 >a K O irs o o U-N O' 81-200 A. No. Resorts cO O rS 00 vs s o OlAOO't t^s CD O (N! CD CD 341 28.3 41-80 A. No. Resorts ^ (S CO W-N CD ° ° ^ 66 OOOO-H 29 447 37.1 Under 40 A. No. Resorts ONN W 207 OO'tN o o — ^ ITS CM CD tr\ O CN CD No. OF Resorts sD 00 -h O tN. ITN N N 5 l/-s 00 00 00 — O' sD 189 sO — ' 102 1,206 100.0 No. OF Resort Lakes (S K O 162 CAVA 00 VN CD Ian o O lA ^ O' C<1 00 CD o CD | Type and Size Class of Resort Lakes DRAINAGE LAKES: I. Large. . 1 1 . Medium Large . III. Medium . IV. Small . Subtotals . LANDLOCKED LAKES: I. Large . II. Medium Large. . III. Medium . IV. Small . Subtotals . SPRING LAKES: I. Large . II. Medium Large . III. Medium . IV. Small . Subtotals . Totals . % of All Resorts . 84 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Thus, we see that 207 of these 803 resorts (which have WA/R values under 40) are on 32 drainage-type lakes, of which 30 are either small or medium in size. Another 71 resorts are located on 39 landlocked lakes, and 34 of these lakes are under 100 acres in size. The remaining 25 resorts are on 7 spring-type lakes, of which 6 are less than 100 acres. It would appear that those resorts with limited water surface are predominately on lakes under 500 acres in size, with almost one-third of them on 53 lakes of less than 100 acres each. About three-eighths (37.6%) of the 189 resorts located on land¬ locked lakes had a WA/R value of 40 or less; whereas only 22.6% of the 915 resorts on drainage-type lakes were in this category. For spring-type lakes, the 40-or-under group included 24.5% of the 102 resorts located on them. Well over half (62.2%) of the 1,206 resorts had a WA/R value of 80 or less. Of these 750 resorts— each with 80 acres or less water surface — 74 were on Class I lakes, 174 on Class II lakes, 385 on Class III lakes, and 117 on Class IV (small) lakes. The WA/R factor varied from about 5.0 acres (Dog Lake and Minnow Lake) to well over 1,400 acres (Ike Walton Lake) in Vilas County. These were the extremes, and the values for lakes in Oneida County fell within this range. No attempt was made to separate lakes with high concentrations of private dwellings from those with relatively low concentrations, but the dwelling density (cottage and home numbers) per mile of shoreline was calculated for 135 of the 301 resort lakes. (See Table 5 for summary.) It is noteworthy that some of the lakes with the lowest WA/R values also had rather high concentrations of private dwellings and cot¬ tages on their shores. This denotes a crowded condition which could very well have deleterious effects. This may result in rapid deterioration and a shorter life for the lakes concerned. On a particular lake, the surface water acreage per resort (WA/R value) is becoming more important each year, especially with the rapid increase in private dwellings on the preferred lakes. A WA/R value of 40 is considered minimal for a good-quality lake with a low dwelling density (less than 10 cottages per mile of shoreline). The average for all 301 resort lakes was 97.5 A. The spatial requirements of certain aqua sports — notably speed¬ boating and water skiing — are rather great. These sports are pur¬ sued not only by resort guests, but also by residents and transient visitors on most larger resort lakes. One lake authority has stated that about 20 acres of water surface are needed for one power¬ boat and skier making a complete (360°) turn! Thus, a lake with 80 acres of surface area may serve very well for one cottage 1969] Mont hey— Lake Size and Type 85 resort (averaging- seven rental cottages and 30 guests), but it might be somewhat too small for two such establishments when both are operating at or near capacity. If this same 80-acre lake had three or four resorts on it, as some already have, along with 10 or 12 dwellings per mile of shore, the utilization conflicts on a hot July day can be visualized quite readily. On most resort lakes, the surface-water area must be shared with residents, cottage owners, area fishermen, transient boaters, and other non-resort people. Three small lakes in Vilas county had less than 10 acres of water surface per resort (WA/R values of 5, 6, 9), and all three had private cottages as well, with an average density of 9 dwell¬ ings per lake! In the Oneida-Vilas area, there were 87 resorts on 29 lakes which had WA/R values under 21 acres. Over half, 50 resorts, were on Class IV lakes (under 100 acres) ; the re¬ mainder of 37 were on Class III lakes. Landlocked lakes predomi¬ nated in this group; there were 17 of them with a total of 25 resorts, and all were Class IV lakes. There were nine (9) drainage-type lakes with 47 resorts, and three (3) spring- type lakes with 15 resorts. The lakes in this group had an average of 8.38 private dwellings on their shores, or about 6 per mile of shore¬ line. This figure approximates the average dwelling density for all resort lakes in this region. SL/R Ratio Studied Another index of resort distribution on a given lake is the amount of “available” shoreline per resort, or the SL/R ratio. Since it is another measure of resort density, or crowding, it further highlights the spatial requirements in resort-lake usage. This value is obtained by dividing the total miles of shoreline on a given lake by the total number of resorts thereon. The SL/R values obtained reflect the average length of shoreline per resort on specific lakes, disregarding the amount of lakeshore actually owned by the resorts or other landowners on those lakes. The average values (and resort distribution) for our 12 categories of resort lakes are shown in Table 3. For example, a lake with 12.0 miles of shoreline and 12 resorts thereon would have an SL/R value of 1.00- — or somewhat less than the average figures for the 2-county area. (Oneida county is 1.36, while Vilas lakes have an average value of 1.11 miles per resort.) It is true that those lakes having large numbers of resort estab¬ lishments are also attractive to second-home owners, and thus they often have above-average concentrations of homes and cottages. This tends to complicate the interpretation of SL/R values. How- Table 3. Shoreline per Resort (SL/R Ratio) in the Oneida-Vilas Area of Wisconsin 86 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 % OF Resorts Each Class WITH .5 MI. or Less 13.77% 23.26 26.39 25.00 21.64 OO\0^ O O — ' co O O 00 c-s 9.00 0.00 0.00 17. 19 23.53 14.71 19.07 Shoreline Per Resort (SL/R Ratio) 2.01 mi. + No. Resorts C6 lA C6 (N CD oo ia K OO n CD CD 0 11 11 0 CO co 138 11.44 1.01-2.0 mi. No. Resorts c\ 00 0s "'t' o^co- 274 OOOOr^ © OOOO^ CO C<1 CD tv © 30.43 .51-1.0 mi. No. Resorts (N ia n O O — ■ O © 00 tv. O © "'T O' CD CD 471 39.06 Under 0.5 mi. No. Resorts 00 O O O 198 0000 0 tv O O — ^ vr\ 230 19.07 No. OF Resorts © oo — < o tv N M M 915 U-N 00 00 00 — O' © 189 o — . 'st* tv co © 102 1,206 100% No. OF Resort Lakes NrAKO NnOON 162 Us 00 Us C~\ lr\ o O ic O' CO 00 CD o CD Type and Size Class of Resort Lakes DRAINAGE LAKES: I. Large — 1001A+. . . . 11. Med. Large — 5Q1-1Q00A . III. Medium — 101-5 00 A . IV. Small — 100A . Subtotals . LANDLOCKED LAKES: I. Large — 1001A+ . II. Med. Large — 50 1-1 000 A . III. Medium — 10 1-500 A . IV. Small — 100A . Subtotals . SPRING LAKES: I. Large — 1001A+ . II. Med. Large — 50 1-1 000 A . III. Medium — 101-500 A . IV. Small — 100 A . Subtotals . Totals . (/. U 9 o QL < ‘c ) j J i J 1969] Monthey—Lake Size and Type 87 ever, dwelling density is evaluated in a subsequent section of this paper, and it can be related quite easily to the resort distribution on each lake. The SL/R value, in our opinion, should be at least 0.5 mile per resort under average circumstances— especially on landlocked lakes where there is no easy access or channels to other waters— and particularly where the dwelling density exceeds 10 units per mile of shore. Why is “available” shoreline important? First, much of the recreational activity on most lakes takes place within % mile of the shore. The central area of a large lake — particularly one with¬ out islands— is used very little. Secondly, certain major activities such as fishing and canoeing are high users of shoreline, and the “acres per boat” or “people per acre” figures are less significant. Thirdly, lakes vary greatly in shore length per 100 acres of sur¬ face. For example, Dollar Lake and Watersmeet Lake in Vilas County are about 100 acres each. Dollar (nearly round) has 1.4 miles of shore, whereas Watersmeet (spider shaped) has 10.5 miles. Dollar (3 resorts) has an SL/R value of 0.47 miles; Water¬ smeet (8 resorts) has 1.3 miles. These are extremes, of course, but they illustrate the point. Of the 1,206 resorts in the Oneida-Vilas region, about 19.0% (230 resorts) had SL/R values of less than 0.5 mile. Of this group, 38 were on Class I lakes; 60 on Class II lakes, 109 on Class III lakes, and 23 on Class IV (under 100 acres). The big majority of them (86.1%) were on drainage-type lakes. The remainder was almost evenly divided (7.4% and 6.5%) between landlocked lakes and spring lakes, respectively. Resorts and Lake Fertility A reasonably reliable index of lake quality, or biological produc¬ tivity, is the fertility rating or total alkalinity of the water in a given lake. This is a key factor in the quality of resort lakes, since it is not only associated with fishery yield and composition but also the prevalence or abundance of algae, aquatic plants, plankton, etc. Generally speaking, soft- water (acid) lakes are relatively infer¬ tile. Most landlocked lakes are of this type. On the other hand, most hard-water lakes show a medium or high fertility rating. This index, expressed as total alkalinity in Wisconsin lake inven¬ tories and determined by chemical tests, is a measure of the dis¬ solved solids (carbonates, bicarbonates, hydroxides, etc.) in a sample of water. It is commonly reported as parts per million of 88 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 calcium carbonate (ppm CaC03) and is an indicator of lake pro¬ ductivity. The following classification is used to indicate fertility ratings of the 301 resort lakes in this study: Total Water Fertility Productivity Alkalinity Hardness Rating VL = Very low . 1- 4 p.p.m. Ultra soft Very infertile L = Low . 5-2C p.p.m. Very soft Infertile M = Medium . 21-40 p.p.m. Soft Fairly fertile H = Medium high . & High . 41-90 p.p.m. Med. hard Fertile 91+ p.p.m. Hard Very fertile In this report the last two categories are combined into one, i.e., “high” fertility. Lakes in this class are generally the most produc¬ tive and support a large population of fast-growing fish, other organisms, and aquatic plants. Conversely, lakes rated as “very low” or “low” are relatively unproductive — like a poor soil — with a limited population of slow-growing fish and relatively clear, weed- free water. Fish are sometimes stunted, especially if predators and large fish are inadequate to control the population. Table 4 shows the distribution of our 1,206 resorts by lake fertility, type and size class. Four levels of fertility, or produc¬ tivity, are indicated for each of the 12 lake type-and-size categories. The table reports both the number of resorts and the number of resort lakes under each fertility class. It is noteworthy that 78.8% of all resorts were on lakes of medium or high fertility; further that 825 (86.8%) of these 950 resorts were on drainage-type lakes. Thus, it appears that the great majority of resort establishments have selected, wisely or by chance, resort lakes of good or above- average productivity. However, there were 256 resorts on 115 lakes of low or very low fertility. They comprised over one-fifth (21.2%) of all resorts in the region. Almost two-thirds of these resorts (161 establishments) were on infertile landlocked lakes ; 90 were on drainage-type lakes ; only 5 were on spring lakes. At the “very low” level of water fertility — nr productivity — there were 54 resorts on 42 lakes. Thirty of these lakes were small, 100 acres or less, and only two were over 500 acres in size. All 42 were landlocked lakes, and most of them were questionable or undesirable sites for resorts, because of their small size and limited fishery. Table 4. Resort Distribution by Lake Type and Lake Fertility in the Oneida-Vilas Area 1969] Monthey — Lake Size and Type 89 % of All Resorts on Low- Fer¬ tility Lakes O pa O — ' pa O pa O 7.50 0. 17 1.41 6.58 5.25 13.41 o © © o 0.41 21.22% Resort Distribution by Lake Fertility Class* High No. of Resorts Vn \D 201 o o o — - O O 00 o cs 00 c-} 230 No. of Lakes ns ( 1) ( 1) a © (44) Medium No. of Resorts 0s 0s tv 0s OxONN — < — H CO 624 pa — i O' 27 c-} pa ^ 69 720 No. of Lakes " CN Ws *— -» (106) — 1 1 — 1 (11) W~\ "'T © Low No. of Resorts O sQ O ns w-N 06 — H _ ly-N o ' \D PA 107 © © © i/a 202 No. of Lakes xO (23) (47) NS NS (73) Very Low o t! . o $ 8 o o o o o NS ir\ o o o o o No. of Lakes — OO *—• ns (42) (42) No. OF Resorts so 00 — < o N ^ ^ M (S r'N 915 ia 00 0O 00 — O' vO 189 O — 1 tx. ^0 ■ — ■ 102 1,206 No. OF Resort Lakes pa t\ O (S f°\ OO N 162 C6 VsOOlA o 0>A i'O' C<1 00 NS © NS Type and Size Class of Resort Lakes DRAINAGE LAKES: I. Large . 11. Medium Large. . . . III. Medium . IV. Small . Subtotals . LANDLOCKED LAKES: I. Large . II. Medium Large. . . . III. Medium . IV. Small . Subtotals . SPRING LAKES: I. Large . II. Medium Large. . . . III. Medium . IV. Small . Subtotals . Grand Totals. *NOTE — The figures in parentheses indicate number of lakes in each category. 90 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Dwelling Density on Resort Lakes As mentioned previously, non-resort shoreline developments, in¬ cluding cottages, seasonal homes, year-round residences, camps and marinas, affect lake usage by and water availability for resort enterprises. Heavy usage by resorts plus a high concentration of private dwellings can mean an overcrowded condition on the lake concerned. However, both factors can be measured and evaluated, separately and together, to determine the pressure on a given lake in quantitative terms. Table 5 indicates the cottage-dwelling density on 135 resort lakes in Vilas County, where dwelling counts were taken. The average number was 26.4 dwellings per lake, and the average density was just under 6 dwellings per mile. This rates as a “moderately-low” concentration when measured by the following scale : Dwellings per Mile of Shoreline 1 to 4 5 to 10 11 to 20 21 to 30 31 to 50 51 and Over Concentration or Density Rating Low Moderately Low Medium Moderately High High Very High On a given lake, the total number of dwellings (seasonal and year around) is divided by the total miles of shoreline to determine the dwelling density. On the resort lakes studied, this factor varied from 0 to over 20/mile. For the 12 type-size categories of resort lakes, the average dwelling density ranged from 1.18/mile to 9.55/mile, with the greatest variation among landlocked and spring-type lakes. However, this value averaged approximately 6.0 per mile for each of the three major lake types, with spring lakes (6.53) being slightly higher than the two other categories. Class II landlocked lakes had the greatest dwelling density, 9.55 per mile. Class III spring lakes (with 8.14) and Class IV drainage lakes (with 7.75) had the next highest densities in terms of dwell¬ ings per mile of shoreline. As would be expected, the larger lakes have more dwellings per lake, but somewhat fewer per mile of shoreline. However, the differences in dwelling density were rather small, as the following data indicate: 1969] Monthey — Lake Size and Type 91 Size Group No. OF Resort Lakes Dwellings Per Lake Dwellings Per Mile I. 1000 Acres + . 13 66.2 5.86 II. 501-1000 Acres . 23 45.7 5.97 III. 101-500 Acres . 58 23.2 5.95 IV. 100 Acres or Less . 41 7.5 6.37 In general, lakes of the Oneida-Vilas region are relatively un¬ crowded in terms of dwelling density, especially when compared to the majority of lakes in southern Wisconsin. Densities of 20 or more dwellings per mile of lakeshore were evident in only a few cases, mostly on smaller lakes. However, when a 100-acre lake has a medium concentration of 11 or more cottages per mile of shore, plus two or more cottage resorts of average size, we are beginning to approach conditions that could easily lead to overcrowding in a few years. This study of dwelling density involved 3,564 private cottages and homes on 135 resort lakes. This is almost one-third of all such dwellings, which numbered approximately 12,000 in this two-county area in 1964. Lakes With Large Resorts There were 25 resort establishments in the Oneida-Vilas area which had 30 bedroom units or more. These enterprises were lo¬ cated on 22 resort lakes, which range in size from 110 acres (smallest) to 3,870 acres. The lake type, size, and general charac¬ teristics of these 22 lakes are reported in Table 6. In general, the larger resorts were located on the larger, better- quality lakes. Although eight of the 22 lakes used were Class III lakes (101 to 500 acres), almost two-thirds of them were Class I or Class II lakes, and the average size of all 22 was over 1,000 acres. All but one of these lakes were of the drainage type, and all but three were medium to high in fertility rating. These lakes were also popular with cottagers and owners of pri¬ vate dwellings. Only one of the 22 lakes had no dwellings other than resorts, and the other 21 averaged slightly over 7.0 dwellings per mile of shoreline — or about 20% higher than the average dwelling density for all resort lakes. However, this density was more than doubled on four of the 22 resort lakes; these were lo¬ cated fairly close to a major city or village, which may account for the higher densities of 14.5 to 17.4 per mile. Table 5. Cottage and Home Concentrations on Resort Lakes (Vilas County Only) 92 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 ,-v. ,h-v od tu O. <0 > > > CL U"“r < < < < O jZ Wo Co t\ Wo sO 00 WO fo — < C'i OcfK CO cO z w w MvO'tK OO >— 1 Wo to. O a- CO — 1 — 1 Wo O 3 d g u vO Vv Va f\ Wo > — 1 0s Vrs \Q vo O CO 00 CO nO Wo J'S ? > g2 Q J*5o N x-O 3o W W ' a, * — < 0 0) > < < < 0) > < 0 1 z OOirvN'-' OfC'tN CO Wo — H O sD Z f— 0 o-- co 0 CO — CO 00 sO CO O sO O wo xD jO> t\ ^ N ^ CO T— ' — < CO Col (Ol e s< C/3 0 W u y w 5 Z-V 0 © 0s CO ^ C-3 K Oco-hO , a co co co 0 O cO 0s 'T vrs vrs _ — , co — . «— < co CO • 1— 1 L) Q No w/< AN cci £ £ w.3.2 TS £ <0 322 CO £ £ ^ a ' — OONKKOOO'Ni'N'^O^O^ rorO'— >rAMiAO>A\0,tN'-iOO NOx(y>^‘A(M(^>ANTtKOOKNKON(NiACOOO,tf,N N'AOO00r^00Nv£)N'tf,\00,tiAN't00N>r.^ < nNfS" — — — « L M — .S £ 03 '2x'2xx'2'2'2x'2'2^x'2^'2'2'2'2>i'2'2 + CO QJ 8“ o5< OOOOOOOOOOOOOOOw-N^^tOI'x — O OOroN>Air,O\000N""iA\DO,tNNvD'ti,O KNVMA000',O'^r^t\0'^NO'£)ro00^iAKrt' — 1 OOvOOOfON^ — ' OO^OOw^i^irsi^^^c^c^cO — — < »— < wr^N' — i > — i <— i > — i No. OF Resorts on Lake N,t,troir\O'ro00iAKi<,t'ON00>A(}'i'0'(^. — ' 00 10.6 QQQQQQQQQQQQQQQQQQQQjQ u. O w 0. >- H 1969] Monthey — Lake Size and Type 95 many more in Oneida County. A number of resort lakes are in this group. This study revealed 46 resorts on 21 winterkill lakes, 30 in Oneida County and 16 in Vilas County (almost 4% of all resorts). All in all, there were 27 resorts on 16 lakes with a maximum depth of nine feet or less, and a total of 333 resorts on lakes that were under 20 feet in depth. Most of these shallow resort lakes, however, were not subject to severe winterkill. There are also some resort locations in this region that experi¬ ence a periodic “summerkiH” of fish, particularly in algae-laden bays of fertile lakes. Both winterkill and summerkill conditions tend to reduce recreational opportunities or esthetic values for re¬ sort guests, and lakes subject to them cannot be considered as good sites for resort operation or development. Summary and Discussion There are 951 named lakes in the Oneida-Vilas area of North¬ eastern Wisconsin, of which 301 were used by resorts in 1964. Of the 301 resort lakes, 162 were drainage type, 101 were landlocked, and 38 were spring-type lakes. The distribution of resort enter¬ prises was 915 on “D” lakes, 189 on “L” type, and 102 on “S” lakes. Almost three-fourths of all establishments were on “D” lakes over 100 acres in size. About one-fourth of the resorts were on Class I lakes (over 1,000 acres). Another fourth were on Class II lakes (501 to 1,000 acres), and about two-fifths were on Class III lakes (101 to 500 acres). The remainder (10.37%) were on Class IV lakes (100 acres or less). Well over half of all resorts (62.2%) had a WA/R ratio (water acreage per resort) of 80 acres or less. One-fourth of them had under 40 acres of water. About one-fifth had a SL/R factor of 0.5 mile or less, under i/2 mile of shoreline per resort. Over one-fifth of all resorts were on 115 lakes of low or very low fertility; 78.7% were on 186 lakes of medium or high fertility. Almost 5% of the resorts were on 42 landlocked lakes of very low fertility, and well over 3% were on winterkill lakes. On these resort lakes, dwelling densities averaged almost six buildings per mile of shore. Over 75% of all resorts and almost 75% of all dwellings were on drainage-type lakes. A study of the lake characteristics associated with the 25 largest resort establishments in the region was included. This group of 22 resort lakes averaged 1,024 acres each, and all but one were drainage-type lakes. Only three of these lakes had a low fertility rating, 14 were medium and 5 were highly fertile. Five of these lakes showed signs of being overdeveloped, with low WA/R and 96 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 SL/R values coupled with above-average concentrations of dwell¬ ings. Dwelling densities for these lakes averaged slightly over seven buildings per mile, with a range of 1.61 to 17.44 per mile, compared to an average of 5.92 for all resort lakes. In general, the larger resorts, all of which had 30 bedroom units or more, were associated with the larger, more fertile, and better-quality lakes. As a result of this study and related field observations, the fol¬ lowing comments and suggestions are offered: 1. Resorts should not be located on any lake under 50 acres in size. In fact, it is felt that no motors or powerboats should be operated on these small bodies of water, particularly land¬ locked lakes. Yet 4.6% of all resorts in the Oneida-Vilas area were situated on lakes in this group. 2. It is questionable whether any resorts should be situated on landlocked lakes smaller than 100 acres, as they are usually infertile and easily damaged. Motors rated over 10 h.p. should not be used on them. This region had a total of 68 resorts on such lakes. 3. A minimum of 40 acres of surface water per resort is recom¬ mended on lakes where dwelling densities are low or moder¬ ately low (10 or fewer dwellings per mile). With medium dwelling densities (11 to 20 per mile), the WA/R factor should be at least 50 acres. When the density is greater, 21 or more dwellings per mile, this “space” value could be 60 or higher. One-fourth of all lakes in the Oneida-Vilas area had WA/R values under 40, and 87 resorts on 29 lakes showed WA/R values of 20 or less. 4. The SL/R factor, miles of shoreline per resort, ought to be 0.50 or more, since most of the water-based recreation is related to the use of shoreline. Yet 19% of all resorts in this area had SL/R values of 0.50 or less, and 33 resorts on five lakes had SL/R values under 0.25 mile. 5. Resort lakes should preferably have a fertility rating of “medium” or higher. Those with a low — and especially a very low — fertility rating just cannot withstand the present day fishing pressures and produce game fish of satisfactory quan¬ tity or quality. Yet, 54 resorts (4.5% of all resorts in this region) were located on lakes of very-low fertility. 6. The dwelling density on a given lake should be under 20 per mile, if that lake is to provide the kind of seclusion and recre¬ ational opportunity that most resort guests are seeking when they come to Northern Wisconsin. Only a few resort lakes in the Oneida-Vilas area are now approaching this figure. How¬ ever, with the rapid increase in “second homes” and private 1969] Monthey — Lake Size and Type 97 lake cottages, many lakes will probably become rather heavily developed and thus lose their northwoods atmosphere, even with land-use zoning and other building regulations. 7. Certain kinds of lakes should probably be avoided, by new resort enterprises at least, aside from the basic size, type, and fertility characteristics they possess. These lakes might include : a. Very shallow lakes where the maximum depth is under 10 feet, regardless of lake type. b. Winterkill lakes with a known history of regular or periodic fish die-offs during winter months because of oxygen deficiencies. c. Summerkill lakes with low-oxygen areas at certain periods in the summer season. d. Highly-acid, bog-type lakes with brownish water and low fertility. Such lakes usually have a high percentage of soft, marshy shoreline — and thus may never be crowded — but their fishery is usually too limited and the water color and quality is not desirable for recreationists. e. High-algae lakes that produce greenish or brownish “blooms” and scum quite regularly each summer. Such lakes are usually highly fertile, and hence may be good for fishing, but the odors and esthetic characteristics are objectionable to swimmers, boaters and vacationers in general. f. Lakes seriously affected by pollutants of any type. 8. It is urged that a comprehensive lake use and conservation plan be prepared as soon as practicable for each and every inland lake of 500 acres or more in Wisconsin. Ultimately, perhaps by 1975, a detailed plan could be developed for each of our 1,200 lakes and flowages that are over 100 acres in size. These inventory studies, which include the findings and recommendations of qualified lake specialists, would provide the following information : 1. Geographic description 2. Physical features (of lake and basin) 3. Water quality and chemical data 4. Aquatic resources 5. Shoreline and related resources 6. Present land use and development 7. Current and future problems 8. Protection measures needed 9. Lake use possibilities and future development 98 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 This report, which involved over 25% of Wisconsin’s resorts and inland lakes, suggests that we need to take a more careful look at our lake resources and how they are used. These resources are more valuable with each passing year, but they also are quite vulnerable to misuse and are highly perishable. In some cases, it appears that lakes too small, too infertile, and too shallow are being used. A few of the better lakes seem to be overcrowded already, and water pollution is in evidence. Perhaps the time has arrived for a re- evaluation and careful study of resort patterns and the resort lakes being used or developed. Such an effort would help to insure the protection of these most valuable resources and will benefit Wis¬ consin’s recreation industry as well. References Andrews, L. M. and C. W. Threinen. (1966) Surface Water Resources of Oneida County. Conservation Division, Wis. Dept. Natural Resources, Madison. 284 pp. Birge, E. A. and Chauncey Juday. (1914) The Inland Lakes of Wisconsin. Bulletin No. 27, Series 9, Wis. Geol. and Nat. History Survey, Madison. 137 pp. Black, J. J., L. M. Andrews, and C. W. Threinen. (1963) Surface Water Resources of Vilas County. Conservation Division, Wis. Dept. Natural Resources, Madison. 318 pp. Fine, I. V. (1966) Wisconsin and The Vacationer. Wisconsin Department Re¬ source Development, Madison, 25 pp. Fine, I. V. and R. E. Tuttle. (1962) Wisconsin Vacation-Recreation Papers , Vol. II, No. 3. University of Wisconsin, Madison, 24 pp. Fine, I. V. and E. E. Werner. (1960) The Wisconsin Vacationer, Wisconsin Vacation- Recreation Papers, Vol. I, No. 11. University of Wisconsin, Madison, 15 pp. Humphrys, C. R. and J. 0. Veatch. Classifications for Michigan's Inland Lakes. Water Bulletin No. 8, Dept. Resource Development, Michigan State University, East Lansing. 18 pp. Monthey, L. G. (1963) Classification and Inventory of Tourist-Lodging Busi¬ nesses in Wisconsin. Recreation & Touristry Notes RT-01, University of Wisconsin. 10 pp. Monthey, L. G. (1964) The Resort Industry of Wisconsin. Wis. Academy Science, Arts & Letters Transactions, Vol. 53 — Part A, pp. 79-94. Threinen, C. W. (1962) Some Spatial Aspects of Aquatic Recreation. Fish Management Misc. Report No. 6, Conservation Division, Wis. Dept. Natu¬ ral Resources, Madison, 11 pp. Wisconsin Lakes. (1968) WCD Publication 218-68, Conservation Division, Wis. Dept. Natural Resources, Madison. 36 pp. WCD Lake Use Report No. FX-31. Booth Lake, Walworth County, Wisconsin. (1968) Conservation Division, Wis. Dept. Natural Resources, Madison. 16 pp. GLACIAL GEOLOGY OF NORTHERN KETTLE MORAINE STATE FOREST, WISCONSIN Robert F . Black Abstract More than a century ago typical stagnant-ice features were recognized and correctly interpreted in the world-famous Kettle Interlobate Moraine in eastern Wisconsin. The common border of the Green Bay and Lake Michigan lobes of late-Woodfordian age. (Cary) is a drainageway established on and partly let down from the surface junction of the two lobes. In the Northern Kettle Moraine State Forest that drainageway, now 0.5 to 3.0 miles wide, is a marked depression that is floored largely with stratified elastics. Rising abruptly from the center of the depression are numerous striking moulin kames and from the flanks numerous crevasse fills, eskers, kames, and other stagnant-ice features that constitute “end” moraines of the two lobes. The “end” moraines are 0.5 to 3.0 miles wide and merge abruptly up ice into ground moraine with drumlins and scattered stagnant-ice features. Abrupt bends in the interlobate moraine seem related to bedrock topog¬ raphy and local direction of ice movement of the opposing lobes. Many representative stagnant-ice features are preserved in the Northern Kettle Moraine State Forest, but most of the moulin kames are outside it. These kames are the best examples to be found for hundreds of miles around and are among the best to be found in the world. However, they are being destroyed rapidly because of the demand for construction aggregates. Introduction Since its inception in 1937, the Northern Kettle Moraine State Forest and environs has been one of the more popular public recreational areas in Wisconsin. The center of the Forest is only 45 miles north of Milwaukee (Fig. 1) and serves especially the heavily populated area from Chicago north to Green Bay. The Forest contains many excellently developed and representative glacial features which are internationally famous, but others even more important or striking lie just outside it. The kettles — depres- 99 100 Wisconsin Academy of Sciences, Arts and Letters [VoL 57 ssliicot Ffgrscl^ Hlgii r Men as ha/ ■ — HM) Vannli ’’’Two''”' divers afsmss " Yl WEitei aw' ■ Siaskferislge Ik g llliiiil H LAKE Oshkosh : WINNEBAGO. St, tai mz heveiaftti: "MTcrove / MtlefSViSfe ? ?eeb > > isychs^ag Stenbeiii; tales Shehoygon ^eoar Grov^ 4/Tbefg$|j ®ap||| §1 f* iestufs; f Port [iSiCkSOfi fSoc.Hi« Brown: /-Dee r /Bayside — RtwHilb ^.'1 —Fox Point — Glendale,,-^ Whitefish Bay I Shodi^ood'M i~4¥a«watosa , Elm Grova-* - — 'Milwaukee itlhifel': M-lwaukee on%s;'co?e n wwf Ml! (-!’ 0 ♦I few t V; Figure 1. Portion of the official Wisconsin State Highway Map, centering on the Northern Kettle Moraine State Forest. 1969] Black — Glacial Geology of Northern Kettle Moraine 101 sions from the melting out of buried ice blocks — are the first in the world to have been described adequately and interpreted cor¬ rectly (Whittlesey, 1860 and 1866; Chamberlin, 1877 and 1878; White, 1964). Alden (1918, p. 308) cited the area east and north¬ east of Kewaskum (Fig. 2) as . . one of the finest examples of terminal-moraine topography in the United States.” Many thou¬ sands of tourists and students each year who visit the Forest do so in part because of their interest in the various glacial features, of which many may be seen along the Kettle Moraine Drive (Fig. 2). It seems timely to outline for them the glacial history of the area as we now know it, because the earlier literature is largely out of print and not focused specifically on the Forest area. The Northern Kettle Moraine State Forest includes some 24,000 acres in a very irregular area about 22 miles long and 1 to 4 miles wide (Fig. 1). It extends from the vicinity of Glenbeulah in She¬ boygan County southwesterly and then southerly to the vicinity of County Highway H, about three miles southeast of Kewaskum, in Washington County. The Forest encompasses much of the area along the common boundary between Fond du Lac and Sheboygan Counties. In and adjacent to the Forest a variety of topographic features rise abruptly some tens of feet to 200 or 300 ft. above prominent lowlands. The precipitous wooded slopes interspersed between typical well-kept Wisconsin farmlands (Fig. 3) and abundant lakes (Fig. 4) makes the area especially photogenic. The numerous lakes and the wooded hills and trails make for an ideal vacation land close to major centers of population. The Northern Kettle Moraine State Forest lies athwart the inter¬ nationally famous Kettle Interlobate Moraine. The moraine, as the name implies, was built between two ice lobes — the Green Bay lobe on the west and the Lake Michigan lobe on the east (Alden, 1918, p. 308-309) — during Woodfordian time, between 13,000 and 22,000 radiocarbon years ago (Black and Rubin, 1967-68). This is the “. . . master topographic feature of the whole series of glacial deposits in eastern Wisconsin” (Alden, 1918, p. 235) which first attracted the attention of early explorers. The moraine consists of silt and sand and coarse, angular to well-rounded rock fragments of the local light grey Niagaran dolomite particularly, but also of rocks from northern Wisconsin, Upper Michigan, and Canada. The composition and texture of the drift comprising the moraine varies markedly from place to place within the area. The drift was dumped between the two lobes of ice as they butted against each other and was also deposited under and on top of the dirty ice along that junction during the final stagnation and destruction of the ice. Hence, some debris was deposited directly from the ice, 102 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Figure 2. Part of the Northern Kettle Moraine State Forest and environs showing the Kettle Moraine Drive and its relation to the major glacial fea¬ tures. Local direction of ice movement is shown by arrows. Base from por¬ tions of Campbellsport and Kewaskum topographic quadrangles of the U. S. Geological Survey. Mapping was done largely from air photos, with local field checks. 1969] Black — Glacial Geology of Northern Kettle Moraine 103 Figure 3. Typical farm and wooded hills in the Kettle Interlobate Moraine, 1.5 miles south of Glenbeulah. Note light-colored, rounded cobbles of Niagaran dolomite. but other material was displaced and reworked by gravity move¬ ments and running water. Glacial-fluvial and glacial-lacustrine de¬ posits are especially common. The buried blocks of glacial ice subsequently melted out, pitting the surface with thousands of irregular kettles from a few feet (Fig. 5) to several miles in extent. A privately owned area within the northcentral part of the For¬ est, northeast of Dundee (Fig. 2), contains one of the most strik¬ ing groups of moulin kames (conical hills of drift deposited under the ice) to be found anywhere in the world (Figs. 6 and 7). [Moulin (moo'lan') Fr., is defined by Webster’s dictionary as “A nearly vertical shaft in a glacier into which a stream of water pours.” The debris carried in by the water is piled up at the base of the moulin, building the moulin kame.] The kames should be protected immediately from further exploitation as they are the best examples to be found for hundreds of miles around. Several 104 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Figure 4. Butler Lake, a kettle lake, as seen from the top of the Parnell esker, 2.5 miles northeast of Dundee. less well developed or less “showy’’ moulin kames in another group east and north of Kewaskum have already been destroyed. Relatively few studies have been made of the glacial phenomena in the Kettle Interlobate Moraine or of their detailed history. Much of what we know was learned 50 to more than 100 years ago when Wisconsin’s outstanding glacial geologists of that heyday were active in their reconnaissance studies of the State. Far more in¬ formation yet awaits discovery through detailed systematic inves¬ tigations than we have learned in our various rapid reconnaissance observations. This means that even the interested layman may un¬ earth critical discoveries which can perhaps provide important clues to the geologic history of the area. The serious student will use the following U. S. Geological Survey topographic maps in his 1969] Black — Glacial Geology of Northern Kettle Moraine 105 Figure 5. Knob and swale topography, 1.5 miles southwest of Dundee. Some rounded knobs are kames and others are till. The elongate ridges are cre¬ vasse fills; the ponds occupy kettles. visit to the area : Kiel, Kewaskum, West Bend, Campbellsport, and Sheboygan Falls. The place names used herein refer to those maps, but they cannot be reproduced here. The Kettle Interlobate Moraine was last mapped by Alden (1918) as part of a reconnaissance in southeastern Wisconsin and has hardly been touched since. Much study is needed to modify his findings significantly or to understand fully the history of individual forms or even of many large units. Different interpreta¬ tions are possible within the framework of existing data. However, it seems clear that several local fluctuations of the two lobes were involved during Woodfordian time. The junction thus is a zone of partial mixing or interstratifying of material from each lobe. Out- wash gravel and other glacial deposits were reworked and redepos¬ ited, commonly on pre-existing ice, as the junction shifted back and forth. The area is so large and diverse that it is not feasible nor neces¬ sary for purposes of this report to describe each feature. Rather, 106 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Figure 6. Moulin kames from left to right — McMullen Hill, Conner Hill, and Johnson Hill — northeast of Dundee. part of the area is subdivided largely by air photo interpretation into mappable units or groups of similar geomorphic features (Fig. 2). These are not pure units because of the almost infinite detail available within any relatively small segment. Nonetheless, they serve to emphasize such features as end moraines and stagnate-ice or “dead-ice” moraine with knob and swale topogra¬ phy (Fig. 5), moulin kames (Figs. 6-8), outwash, eskers (Figs. 9 and 10), crevasse fills (Fig. 5), kettles (Figs. 4 and 5), and the like. These and other features are described more fully later. Because of its variety and superb development of “text-book” features, its proximity to centers of population and heavy recrea¬ tional use, and its historical importance in the development of con¬ cepts in glacial geology, this area is one of the most important in 1969] Black — Glacial Geology of Northern Kettle Moraine 107 Figure 7. Cut in Garriety Hill, a moulin kame northeast of Dundee. the State. Further expansion of the Forest, in spite of high land values, is exceedingly desirable and cannot wait long before many features will be irrevocably lost. General Description of the Moraine In 1876 T. C. Chamberlin orally presented a paper to the Wis¬ consin Academy of Sciences, Arts, and Letters on the extent and significance of the Wisconsin Kettle Moraine (Chamberlin, 1878). In those days when great geologists were formulating principles of the concepts of glacial geology, Chamberlin was a true giant among them (Fenton and Fenton, 1952). Although today some of his words and phrases are no longer popular and editors would cut and prune his remarks in order to save space, Chamberlin’s description of the moraine bears the test of time so well that this writer feels compelled to quote him directly. In describing the sur¬ face form of the moraine he wrote : “The superficial aspect of the formation is that of an irregular, intricate series of drift ridges and hills of rapidly, but often very gracefully, undulating contour, 108 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Figure 8. A small moulin kame, 0.3 miles southeast of Dundee. consisting of rounded domes, conical peaks, winding and, occasion¬ ally, geniculated ridges, short, sharp spurs, mounds, knolls and hummocks, promiscuously arranged, accompanied by corresponding depressions, that are even more striking in character. These de¬ pressions, which, to casual observation, constitute the most peculiar and obtrusive feature of the range, and give rise to its descriptive name in Wisconsin, are variously known as 'Potash kettles/ Tot holes/ Tots and kettles/ 'Sinks/ etc. Those that have most arrested popular attention are circular in outline and symmetrical in form, not unlike the homely utensils that have given them names. But it is important to observe that the most of these de¬ pressions are not so symmetrical as to merit the application of these terms. Occasionally, they approach the form of a funnel, or 1969] Black — Glacial Geology of Northern Kettle Moraine 109 Figure 9. Part of the Parnell esker and Butler Lake, as seen from the air, at a geologic marker. of an inverted bell, while the shallow ones are mere saucer-like hollows, and others are rudely oval, oblong, elliptical, or are ex¬ tended into trough-like, or even winding hollows, while irregular departures from all these forms are most common. In depth, these cavities vary from the merest indentation of the surface to bowls sixty feet or more deep, while in the irregular forms the descent is not unfrequently one hundred feet or more. The slope of the sides varies greatly, but in the deeper ones it very often reaches an angle of 30° or 35° with the horizon, or, in other words, is about as steep as the material will lie. In horizontal dimensions, those that are popularly recognized as ‘kettles’ seldom exceed 500 feet in diameter, but, structurally considered, they cannot be lim¬ ited to this dimension, and it may be difficult to assign definite lim- 110 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Figure 10. Part of the Parnell esker, near Butler Lake, as seen on the ground. its to them. One of the pecularities of the range is the large number of small lakes, without inlet or outlet, that dot its course. Some of these are mere ponds of water at the bottom of typical kettles, and from this, they graduate by imperceptible degrees into lakes of two or three miles in diameter. These are simply kettles on a large scale. “Next to the depressions themselves, the most striking feature of this singular formation is their counterpart in the form of rounded hills and hillocks, that may, not inaptly, be styled inverted kettles. These give to the surface an irregularity sometimes fit¬ tingly designated 'knobby drift/ The trough-like, winding hollows have their correlatives in sharp serpentine ridges. The combined 1969] Black — Glacial Geology of Northern Kettle Moraine 111 effect of these elevations and depressions is to give to the surface an entirely distinctive character. “These features may be regarded, however, as subordinate ele¬ ments of the main range, since these hillocks and hollows are vari¬ ously distributed over its surface. They are usually most abundant upon the more abrupt face of the range, but occur, in greater or less degree, on all sides of it, and in various situations. Not un- frequently, they occur distributed over comparatively level areas, adjacent to the range. Sometimes the kettles prevail in the valleys, the adjacent ridges being free from them; and, again, the reverse is the case, or they are promiscuously distributed over both. These facts are important in considering the question of their origin. “The range itself is of composite character, being made up of a series of rudely parallel ridges, that unite, interlock, separate, ap¬ pear and disappear in an eccentric and intricate manner. Several of these subordinate ridges are often clearly discernible. It is usu¬ ally between the component ridges, and occupying depressions, evi¬ dently caused by their divergence, that most of the larger lakes associated with the range are found. Ridges, running across the trend of the range, as well as traverse spurs extending out from it, are not uncommon features. The component ridges are themselves exceedingly irregular in height and breadth, being often much broken and interrupted. The united effect of all the foregoing fea¬ tures is to give to the formation a strikingly irregular and compli¬ cated aspect.” (Chamberlin, 1878, p. 202-204). Chamberlin in actuality was referring to the surficial features of the end moraine of what is now called the late Woodfordian or Cary ice as it was deployed through the entire State of Wisconsin and not just the interlobate moraine in what is now the Northern Kettle Moraine State Forest. Nonetheless, his description can scarcely be improved upon for the area. In speaking of the nature of the material, Chamberlin (1878, p. 205) emphasized that “. . . all the four forms of material com¬ mon to drift, vis. : clay, sand, gravel, and boulders, enter largely into the constitution of the Kettle range, in its typical development. Of these, gravel is the most conspicuous element, exposed to obser¬ vation .” Chamberlin (1878, p. 210) further recognized that most bedrock units in Wisconsin and Upper Michigan were represented in any one section of the drift, including native copper from Ke¬ weenaw Peninsula, but that the bulk of the drift was derived locally. Thus, most gravel is composed of the local white to very light gray Niagara dolomite, well rounded by water work. How¬ ever, we now know that more than one local advance of ice was involved and that reworking of outwash gravel by later advances 112 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 was commonplace. Hence, some constructional forms contain non- stratified gravel instead of till. Deposition of the reworked gravel directly from ice without water working took place. Other details of the moraine in Wisconsin were presented early, and it was compared with its counterpart in other states (Cham¬ berlin, 1877 and 1883). In the latter paper, the term “interlobate moraine” was first introduced (Chamberlin, 1883, p. 276) and properly diagnosed as to origin in contrast to normal medial moraines. A reconstruction of the ice flow directions (Fig. 11) demonstrates conclusively the lobate character of the ice and the opposing movements of the junction of the two lobes. This gross story has changed little in the intervening 90 plus years. Chamberlin’s important role in the development of the concepts of glacial geology would not have been possible were it not for the clear observations and lucid writings of his predecessors. In con¬ nection with the Kettle Interlobate Moraine, Charles Whittlesey is singled out. It was he (White, 1964) who in the mid-1800’s first recognized the “kettle moraine” and correctly interpreted the ori¬ gin of the kettle holes to buried glacial ice rather than to drifting icebergs as was in vogue at the time. This was truly astonishing insight, and is but one of the major accomplishments of that amaz¬ ing man. The Greenbush Kettle, two miles south of Greenbush on the Kettle Moraine Drive, has been favored with a geological marker sign for years. It is one of the most symmetrical deep circular depressions visible from the road. Many others are more irregular (Figs. 4 and 5) but just as typical whether with or without water in them. In brief, the Northern Kettle Interlobate Moraine is conspicuous because of its more abrupt irregularity and sharpness of feature compared with the undulating ground moraine with smoothly con¬ toured drumlins and till-covered bedrock rises on both sides. The light grey gravel of the Interlobate Moraine also contrasts mark¬ edly with the reddish brown and light yellowish brown sandy till of the ground moraine. Neither its maximum elevation (1,311 feet at Parnell tower, 5.8 miles northeast of Dundee) nor its general relief of 100 to 200 feet are significantly different from the till plains and drumlins adjoining. However, it is characterized by major lowlands at 950 to 1,000 feet, such as that occupied by Long Lake and the East Branch of the Milwaukee River. The flatness of such lowlands and the abrupt rise of drift deposits flanking them also emphasize the glacial features. Farming of the lowlands con¬ trasts with the wooded drift hills to spice the view. 1969] Black — -Glacial Geology of Northern Kettle Moraine 113 Drainage The Kettle Interlobate Moraine lacks an integrated drainage network. Many closed depressions drain through the coarse gravel below and do not need surface streams. Others intersect the ground water table and have perennial ponds or lakes. Elkhart Lake, a large kettle north of the Forest, with high land around it, drains westward to Sheboygan Marsh and the Sheboygan River. Crystal Lake, next south of Elkhart Lake, has no outlet. However, Mullet River (see U. S. Geol. Survey topographic map — Kiel) flows by only 1,4 mile to the southwest in its arc around the north end of the Kettle Moraine Forest. It continues southeasterly and eastward in a tortuous route to join the Sheboygan River at Sheboygan Falls. Interestingly, those two rivers have adjoining headwaters, and their uppermost courses are parallel yet flowing in opposite direc¬ tions about one mile apart northwest of Long Lake. Both rivers have very intricate courses to Lake Michigan, probably in part controlled by fracture patterns in the stagnating ice which per¬ mitted the supraglacial streams to superpose themselves on the underlying drift and bedrock. The East Branch of the Milwaukee River, flowing southward into the Milwaukee River southeast of Kewaskum, drains most of the Northern Kettle Moraine Forest proper. Its course follows the trend of the moraine and generally lies almost precisely on the re¬ constructed boundary between the two lobes of ice. (This is some¬ what west of the boundary indicated by Alden, 1918, pi. 3). Prob¬ ably its origin dates back to the initial abutment of the ice of the two lobes where it developed in the axial depression along that junction. It has remained apparently in that position since. In the wastage of the Lake Michigan lobe, however, additional channels were formed on the stagnating ice. Mink Creek lies in a channel that starts about two miles northeast of Parnell and flows generally southerly past Beechwood in a course with abrupt right- angle bends. These seem also to reflect the fracture pattern of the ice as the initial stream was let down on the surface below. Many other examples exist in the area, but no detailed three-dimensional field study of any of them has been attempted. They need to be integrated into the history of the Moraine. Origin of the Glacial Features Figure 2 shows the distribution of some glacial features that characterize certain parts of the area. For convenience in the classi¬ fication, each unit is named for the most abundant or striking fea¬ ture or features it contains. These units are; Ground moraine (and 114 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 drumlins), drift, end moraine (or stagnate-ice or dead-ice mo¬ raine) , and special features such as moulin kames and eskers. Ground moraine with drumlins and till-covered bedrock rises com¬ prises most of the area up ice from the front of both lobes. Small stagnate-ice features in that unit are common. The orientation of drumlins, fluted forms, and striae recorded by earlier workers and summarized by Alden (1918, pi. 4) show clearly the regional move¬ ments of the ice of both lobes. Arrows on Figure 2 show local trends of the ice recorded by drumlinoid or fluted forms. Even though the general deployment of ice shown in Figure 11 and by Alden (1918, pi. 4) is not expected to be changed in gross form, detailed field work is needed to show ice movement in relation to individual segments of the moraine. Bedrock striae formed in early advances during Woodfordian time are not everywhere parallel with the alignment of molded forms — the last to be produced. In the area of Figure 2, stratified drift, including outwash, glacial-lacustrine deposits and other water-formed features, are almost as prevalent as end moraine or stagnate-ice or dead-ice moraine formed more directly by the ice. The washed surfaces and deposits reflect in part the cleaner ice of the two lobes juxtaposed and in part the concentration of runoff along the junction of the two lobes. The normal surface gradient up ice in each lobe would have led water to the junction of the lobes, from which its escape could only have been to the south along that junction. Such water- worked stratified drift varies in size from the coarse, bouldery ma¬ terial of glacial streams to the sand, silt, and clay in ponded water. Drift obviously has formed in places on buried ice blocks to leave pitted outwash; elsewhere it seems that entire portions of stream beds or lake sediments have been dropped down as con¬ tinuous ice below melted out. Most parts of the well-washed drift, however, were formed adjacent to ice, but not on it. Original stratification is preserved. Even during deglaciation the widening and northward migrating gap between the two lobes effectively concentrated glacial-fluvial activity between the lobes. Thus, it was the locus for many striking forms. Eskers (Figs. 9 and 10) and moulin kames (Figs. 6 to 8) formed under the stagnate ice by subglacial streams fed through moulins or openings through the ice sheet. Their subglacial wa¬ ters also flowed toward that same gap. Crevasse fills (Fig. 5), topographically commonly like short eskers, were formed in cre¬ vasses open to the sky in part by supraglacial streams and in part by mass movement of surface debris into the crevasses. Small moulin kames are scattered throughout much of the area, but none is better developed or displayed than those in the group 1969] Black — Glacial Geology of Northern Kettle Moraine 115 Figure 11. Diagram showing glacial movements in eastern Wisconsin by T. C. Chamberlin, 1876. 116 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 northeast of Dundee (Fig. 2). There, in the widest part of the washed drift area, are some of the best moulin kames to be found anywhere in the world. Beautifully conical hills, such as McMullen (Fig. 6), Garriety (Fig. 7), Conner, and Johnson, rise at the angle of repose of the material more than 100 feet above the flat, washed, drift plain surrounding them. Numerous smaller kames, only a few tens of feet high, are commonly less conspicuous among the drift ridges and are too small to show on Figure 2. Many are just as symmetrical as the larger ones in the lowlands. Other more irregular moulin kames, such as Dundee Mountain (which has a geologic marker), are also present and grade into crevasse fills or into ice-walled lake deposits (openings so enlarged that lakes formed within the glacial ice walls). Such forms originated where melt waters on the ice dropped through moulins or crevasses, dumping their detritus at the base. Openings ranged from nearly vertical, circular pipes (moulins) to very elongate fractures and rounded to irregular large openings; commonly, water and debris were fed into the fractures at more than one place along the sides and ends of crevasses, building irregular forms below the ice. Many large fractures were fed not just with running water, but also with mud flows, debris slides, and the like. Ponded water in some also trapped deltas and lacustrine sediments. Thus, the mate¬ rial in such features as moulin kames and crevasse fills ranges from normal till, through the available sizes of water- transported material, to ponded sediments. The cross section of Garriety Hill is typical (Fig. 7). It shows rounded to angular gravel, sand, silt and clay deposited as unsorted till in irregular masses, and as sorted sediment in alluvial flows, pond sediments, and the like. Water that formed the northern group of moulin kames drained westward under the ice to join the drainageway through Long Lake Valley. Their channels are readily discernible on aerial photographs. End moraine and stagnate-ice or dead-ice moraine are not dif¬ ferentiated on Figure 2 because of their general similarity of ori¬ gin. The terms are used loosely here for lack of detailed under¬ standing of their genesis. They might have been subdivided for descriptive purposes into those areas characterized by elongate ridges and valleys and those with circular knobs and swales. In the interlobate area all are believed to result from ice stagnation and the melting out of blocks of ice of the appropriate geometry to fit the surface depressions. Such geometry is predicated on the move¬ ment of the ice at the time the ice and debris were mixed, on its fractures, or on the manner of burial by overriding ice, outwash, debris slides, etc. 1969] Black — Glacial Geology of Northern Kettle Moraine 117 The detailed deployment of the moraines in the Northern Kettle Interlobate Moraine is of considerable interest in the reconstruc¬ tion of events as related to the flow of ice. From the vicinity of Kewaskum north to Dundee and to Long Lake, the trend of the Interlobate Moraine is almost north. From Long Lake the Inter¬ lobate Moraine turns fairly abruptly to the northeast to Elkhart Lake where it again swings to the north. At least part of the ex¬ planation of the bends may lie in the topography of the bedrock which unquestionably has exercised some control on the deploy¬ ment of the ice. The deep pre-glacial valley at Sheboygan Marsh and Elkhart Lake must have provided relatively easy access for the ice of the Green Bay Lobe, leading it more rapidly and farther to the southeast than was possible over the bedrock hills south of that Marsh. The hills restrained the ice of the Green Bay Lobe, allowing the ice of the Lake Michigan lobe to push farther west¬ ward. Such kinks and bends in the terminal area are commonplace along the entire late Woodfordian front in Wisconsin. They are of considerable importance in understanding the development of such features as are found in the Northern Kettle Interlobate Moraine, but space does not permit their reconstruction here. Much field¬ work is called for to unravel the details of their history. Small moulin kames in the stagnate-ice moraines are probably contemporaneous with the related features, immediately preceding kettle formation. However, the precise timing of the formation of the main group of moulin kames versus the main moraines to west and east is conjectural. The writer hypothesizes that shortly after the two lobes butted together, the thickness of ice gradually increased from 100 to 300 feet at the start to a thickness perhaps of several thousand feet when the ice extended southward into the center of Illinois. Abla¬ tion (loss of ice) particularly by melting aided by a surface stream at the junction of the two lobes would be countered by ice move¬ ment from the base of the ice sheet diagonally upward to that junction at the surface. Upward flow at the terminal zones of gla¬ ciers is commonly at angles of 10 to 45 degrees, bringing debris from the base toward the surface to replace ice lost in the ablation zone and to maintain the surface profile of the glacier. When ice was at its maximum thickness at the junction, the basal debris may not have reached the surface. As the ice thinned during the waning of the late Woodfordian glaciation, it would intersect the surface. As thinning continued to perhaps 200 or 300 feet of ice, fractures penetrated in favorable places, aided by meltwaters, to the bottom of the glacier. In them the moulin kames, eskers, and crevasse fills began to grow. However, at that time the thicker ice 118 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 away from the junction was continuing to move forward even though the terminal zone was stagnate. The shear planes and flow layers that brought debris up from the base presumably angled obliquely downward and away from the actual surface junction of the two lobes to the general location of the main moraines on both sides of the drift area. At the locus of the moraines, basal ice and debris were interstratified by flow of ice while the basal ice closer to the junction was stagnated and remained relatively free of debris. Thus, the two main moraines, one for each lobe, are in a sense end moraines even though they do not mark the terminal position of the ice nor were they deposited at the outer edge of the ice. They represent the outer edge of the active ice for each lobe and were separated by a zone of stagnate ice shaped like a very broad, low wedge with its apex upward, at least during the waning of the glaciation. It seems relatively clear that stagnation took place over much of the area because so many small ice-contact, washed-drift features are superposed on all other forms. Conclusions Many details of the reconstruction of the events that led to the surface features in the Northern Kettle Interlobate Moraine are imperfectly known. New topographic maps and aerial photographs unavailable to Alden (1918) and earlier workers now permit an analysis of surface forms to be made in far more detail than was possible for him in his reconnaissance study. Surface analysis, however, is only part of the story. Serious mistakes have been made in the past in the interpretation of glacial forms by morphology alone. Sub-surface exploration must be carried on concurrently before a firm foundation can be laid that would permit us to change significantly the gross picture of the Kettle Interlobate Moraine as commonly accepted. Such detailed study has had little economic incentive, but should be undertaken before gravel pit operations remove or modify evidence that might be the key to part of the story. A beautiful story can be constructed on evidence available, but an even larger part of the story is still unsupported in fact. The prospects in future study are especially intriguing. Thus, in brief, the heavy use of the area for recreation and con¬ sequent loss of land for cottages and commercial development re¬ quire our immediate action to preserve many glacial forms, like kames, eskers, and stagnate-ice features. Demands for gravel are increasing and many glacial forms are being removed en toto. We must protect not only the many striking forms but also the “nor¬ mal” forms now before they are exploited. Many shown in the mapped area of Figure 2 are outside of the Forest proper. It is 1969] Black — Glacial Geology of Northern Kettle Moraine 119 hoped that some of the better ones ultimately will find their place in the public trust. If not, the gravelly deposits will disappear as have some moulin kames and crevasse fills immediately east of Kewaskum, on the north side of Highway 28. References Cited \lden, William C., 1918, The Quaternary geology of southeastern Wisconsin: U. S. GeoL Survey Prof. Paper 105, 356 p. Black, Robert F., and Rubin, Meyer, 1967-68, Radiocarbon dates of Wis¬ consin: Wis. Acad. Sci., Arts, and Letters, v. 56, p. 99-115. Chamberlin, T. C., 1877, Quaternary Formations — the drift: Chap. V, p. 199- 246, in Geology of Wisconsin, v. 2, 768 p. Commissioners of Public Printing. Chamberlin, T. C., 1878, On the extent and significance of the Wisconsin kettle moraine: Wis. Acad. Sci., Arts, and Letters, Trans., v. 4, p. 201- 234. Chamberlin, T. C., 1883, Terminal moraine of the second glacial epoch: U. S. Geol. Survey Third Annual Report, p. 291-402. Fenton, Carroll Lane and Fenton, Mildred Adams, 1952, Giants of geol¬ ogy: Doubleday and Co., 333 p. White, George W., 1964, Early description and explanation of kettle holes: Jour. Glac., v. 5, p. 119-122. Whittlesey, Charles, 1860, On the drift cavities, or “potash kettles” of Wis¬ consin: Amer. Assoc. Advancement Sci., Proc., 13th meeting, 1859, p. 297- 301. Whittlesey, Charles, 1866, On the fresh-water glacial drift of the north¬ western states: Smithsonian Contr. Knowledge, No. 197, 32 p. Acknowledgements Robert F. Black is Professor of Geology, the University of Wis¬ consin, Madison. Field work leading to this report was supported by the National Parks Service, by the National Science Founda¬ tion, by the Research Council of the University of Wisconsin Grad¬ uate School from funds supplied by the Wisconsin Alumni Research Foundation, by the Wisconsin State Highway Commission, and by the Wisconsin Geological and Natural History Survey. AGE AND GROWTH OF THE WALLEYE IN LAKE WINNEBAGO Gordon R. Priegel Bureau of Research Department of Natural Resources Madison, Wisconsin Introduction The walleye, Stizostedion vitreum vitreum (Mitchill), is an abundant sport fish in Lake Winnebago and connecting waters and it attracts more fishermen to the Lake Winnebago area than any other sport fish. This report describes the age and growth of this species in Lake Winnebago and is part of a study on the life his¬ tory of the walleye. Maintenance stocking of walleyes is not practiced in the study area and accordingly this paper refers exclusively to a natural population. Materials and Methods The 1,237 walleyes used in this study consisted of 1,017 yearling or older fish collected during October and November, 1960 and 220 young-of-the-year walleyes collected from June — October, 1961. All young-of-the-year fish were taken with 12-foot otter trawls while Lake Erie type trap nets, 12-foot otter trawls and an A.C. shocker unit were used to collect the older fish. The length measurements of adult walleyes were made on fresh specimens. The total lengths were measured to the nearest tenth of an inch on a standard measuring board. The length measurements of young-of-the-year walleyes were made on preserved specimens (10 percent formalin). The weight of each fish was determined to the nearest hundredth of a pound ; no young-of-the-year fishes were weighed. All fish, for which length and weight were recorded (1,017 fish), were used in the study of the length-weight relation. Key scales from 1,237 fish were taken from above the lateral line on the left side and came from the intersection of the third row above the lateral line and the first scale row before the first dorsal spine. The scales were impressed on cellulose acetate slides, 0.03 inch thick, by a roller press similar to that described by Smith (1954). Butler and Smith (1953) demonstrated that this method 121 122 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 of preparation does not affect the measurements of scales. The ex¬ amination and measurements of scales were made by means of a micro-projector at the magnification 44X. The length of each scale and the distances from the focus to each annulus were measured along the anterior radius most nearly collinear with the focus as described by Hile (1954) and recorded to the nearest tenth of an inch. Ages were determined by counting the annuli and are given in terms of completed years of life. They are expressed by Roman numerals corresponding to the number of annuli so that fish in the second year of life belong to age-group I (Hile, 1948). Sex and state of maturity were determined for all fish except the 220 young-of-the-year fish collected in 1961. Determination of sex in adult walleyes is easy as the testes have a whitish-gray appear¬ ance, and the ovaries are yellowish with readily visible eggs. Size and shape of gonads, blood vessels on gonads, and color of gonads were used to sex immature fish (Eschmeyer, 1950). In fish of com¬ parable size, the gonads are distinctly the wider in the female. The testis tapers towards the apical and over a considerable portion of its total length, while the region of tapering is much shorter in the ovary. Ordinarily, at least one of the ovaries tends to be translu¬ cent. The dorsal blood vessel of the testis lies in a groove; that of the ovary is on the surface. Veins are usually visible passing across the ovary, while this cross-venation is not found on the testis. The fecundity of 130 walleyes over a size range of 16.6-25.2 inches in total length and a range of 1.39-6.00 pounds was deter¬ mined. The ovaries from these walleyes were preserved in 10 per¬ cent formalin. The weight of each ovary was determined just prior to sampling and a transverse section was made through an ovary. The section was weighed, and the number of eggs within was de¬ termined by actual count. The section represented 1.2 to 7.4 per¬ cent of the entire ovary. The total number of eggs per fish was estimated on a proportional basis. Results and Discussion 1. Age and Growth The precise time of annulus formation of Lake Winnebago wall¬ eyes was not established. Annulus formation probably occurs in May or early June. Carlander (1945) reports that walleyes form an annulus in late May or early June in Lake of the Woods, Min¬ nesota and Cleary (1949) reported the same for Clear Lake, Iowa. 1969] Priegel -Age and Growth of the Walleye 123 Use of the scale method to determine the age of the Lake Win¬ nebago walleyes is justified by the following observations : 1. Fish known to be young-of-the-year had no annuli on the scales. 2. The number of annuli increased with the size of the fish. The body-scale relationship was determined from the measure¬ ment of 1,237 walleyes which were grouped into half-inch groups from 1.0 to 24.5 inches. The mean body length for each group was plotted against the corresponding mean lengths of the anterior scale radii and the relationship may be expressed as : L = 1.443 + 3.171 (R) where L = total length in inches and R = anterior scale radius X 44 The body-scale relationship was linear. The calculations of length at each annuli were made from meas¬ urements of the anterior radius applied in the formula : Li = C + Si (L — C) S where Li is the length of the fish at the time of each annulus formation, C is the length of the fish at the time of scale forma¬ tion, Si is the length of the anterior radius of the scale at each an¬ nulus, S is the length of the anterior radius at capture and L is the total length of the fish at time of capture. The length of the fish at the time of scale formation is 40 millimeters (1.6 inch) as determined from close examination of 220 specimens (Priegel, 1964). The regression line of the body-scale relationship intercepts the abscissa at 1.443 which is slightly less than the value deter¬ mined for body length at time of scale formation. The average calculated lengths of males and females in different age groups of walleyes gave evidence of sex differences in growth rate so the data for males and females were kept separate (Tables 1 and 2.) Two estimates of general growth are given in the bottom section of Tables 1 and 2. One is based on the grand average calculated total lengths and the second on the summation of the grand average annual increments of length. The present discussion is based on the sums of increments, since this curve should represent the aver¬ age growth that walleyes might have if the population was not subjected to selective destruction of individuals with the more rapid growth. (Figure 1.) 124 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Table 1. Calculated Total Length at End of Each Year of Life of Each Age Group of Lake Winnebago Male Walleyes and Average Growth for the Combined Age Groups Age Group Number of Fish Length (Inches) at End of Year 1 2 3 4 5 6 7 8 I . 58 7.1 II . 27 5.8 11.7 Ill . 62 5.2 10.2 13.6 IV . 60 5.4 9.9 12.7 14.8 V . 89 5.6 9.9 12.2 13.7 15.2 VI . 80 5.4 10. 1 12.5 13.7 14.7 16.0 VII. . . . . 38 5.6 10.2 12.6 13.7 14.6 15.4 16.4 VIII . 11 5.6 10.2 12.6 13.9 14.8 15.4 16.0 16.8 Grand average calculated length . 5.6 10.2 12.7 14.2 15.1 15.6 16.2 16.8 Mean annual increment . 5.6 4.6 2.5 1.4 1.0 0.7 0.6 0.7 Growth based on summa¬ tion of increment . . . 5.6 10.2 12.7 14. 1 15.1 15.8 16.4 17.1 Table 2. Calculated Total Length at End of Each Year of Life of Each Age Group of Lake Winnebago Female Walleyes and Average Growth for the Combined Age Groups Age Group Number of Fish Length (Inches) at End of Year 1 2 3 4 5 6 7 8 I . 111 7.6 II . 84 6.2 12.1 III. . 79 5.3 10.2 14.6 IV . 90 5.4 9.6 13.4 16.8 V . 109 5.7 9.6 12.8 15.2 17.7 VI . 77 5.6 9.8 12.8 14.8 16.5 18.9 VII . 36 5.8 9.7 13.5 14.6 15.9 17.3 19.3 VIII . 6 5.4 9.7 13.3 15.3 17.0 18.4 20.4 21.0 Grand average calculated length . . . . 6.0 10. 1 13.4 15.6 17.3 18.6 19.5 20.5 Mean annual increment ..... 6.0 4. 1 3.2 2.2 1.8 1.6 1.2 1.0 Growth based on summa¬ tion of increment ........ 6.0 10.1 13.3 15.5 17.3 18.9 20. 1 21.1 Total Length (Inches) 1969] Priegel — Age and Growth of the Walleye 125 Year of Life Figure 1. General growth in length and annual increment in length of Lake Winnebago walleyes. Males, solid line; females, broken line. 126 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Comments on general growth and a comparison of the growth of the sexes are best made from Table 3 which was prepared from data of Tables 1 and 2, The total lengths of the sexes in the first year of life showed a 0.4 inch advantage for the females but a 0.1 inch advantage for the males at the end of the second year of life. The advantage of the females increased from 0.6 inches at the end of the third year to 4.0 inches at the end of the eighth year. If the 13-inch size limit was still in effect, this difference in growth rate between male and female fish would have affected the age at which the legal size was reached. The male walleye took 4 years to reach legal size and the female 3 years. The greatest increase in length for both sexes took place during the first year of life (6.0 inches for the females and 5.6 inches for the males). The amount of growth dropped during the second year, and the decrease continued for the females through the eighth year; but, the males made nearly the same amount of growth each year after the fifth year (0.6 inches to 0.7 inches). Many authors have reported on the growth rates of walleyes in various bodies of water (Table 4). The walleye population in each individual body of water differs in growth rate from other bodies of water. The greatest growth occurred in southern reservoirs, Norris Reservoir and Clayton Lake. The Lake Winnebago walleye is one of the slowest growing fish when compared to the walleye populations mentioned in Table 4. Only the growth rate of the walleye in Lake Gogebic, Michigan, is similar to the Lake Winnebago walleye. Lack of forage fishes and competition from other fish species (burbot, sauger and yellow Table 3. Calculated Total Lengths (Inches) and Length Increments of Male and Female Walleyes of Lake Winnebago in Different Years of Life Year of Life Mai LES Fer/O \LES Size Advantage OF Females Calculated Length Increment Calculated Length Increment I. ........... . 5.6 5.6 6.0 6.0 +0.4 II. ........... . 10.2 4.6 10. 1 4. 1 I -. + 0 . 1 III. ............ 12.7 2.5 13.3 3.2 + 0.6 IV. ............ 14.1 1.4 15.5 2.2 + 1.4 V . . 15.1 1.0 17.3 1.8 + 2.2 VI. . . . 15.8 0.7 18.9 1.6 + 3.1 VII. . . . . . . . . 16.4 0.6 20.1 1.2 + 3.7 VIII. ............ 17.1 0.7 21.1 1.0 +4.0 Table 4. Calculated Growth of Walleyes Reported from Various Waters 1969] Priegel—Age and Growth of the Walleye 127 GO DJ X u Z <-—i CO w cC £ H H < Gd ZI> u •) CO -1 D J O < 2 od H > < & D Q 3 W < S u g wd o W 2 PJ > < I I I I I I I I I EX o o 00 00 1 1 1 EX CA 1 1 00 —1 EX 1 d 1 cx CX — 1 cx CX cx vO o — cx EX o 00 o O 1 1 cx EX r^s 1 00 tx o d 1 Ea Ex cx cx < I— 1 — < CA Ex CA o 00 — < CX CA O' — CX —H Vn EA tX 00 O' O' d O' lA sO o’ d tx 00 7—1 7—1 *” * cx -a ' 1 ’ 1 cx --H ^-1 l^s cx £ d >; cd SdT 4P ea co n 4> di C 3 P 4) w 4) „ Id j s w T"? co U > TJ T3 cw gw Z 2 M >> bfi C O 5 0*5 j^W co d 4J O' g « i— I co ;cW d CO w- §8 •M CP cd 0 §2 O'-' z 128 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 perch) in Lake Winnebago are probably the limiting factors for the slow growth rate in the walleye. The long spawning migration (90 miles maximum) may also be a factor related to slow growth since these migrations must result in great energy loss. 2. Age At Maturity Only those females showing eggs forming in the ovary were con¬ sidered mature, and the males were considered mature if the testis showed the characteristic whitish-gray color. Since all fish were collected during the late fall, no difficulties were encountered be¬ tween distinguishing immature and mature fish. The information on the age and degree of maturity of the wall¬ eye included in this sample is presented in Table 5. The average age of maturity was considered as that age at which 50 percent of the fish reach maturity. (Table 5). The male walleye would gener¬ ally be considered as mature at the end of the third year of life. At the end of the sixth year of life, all males were mature. The female walleye would be considered as mature at the end of the sixth year of life. Only at the end of the eighth year of life were all females mature. The males show a definite tendency to mature earlier in life. The average total length at which more than 50 percent of the males are mature is 12.7 inches. The average total length at which more than 50 percent of the females are mature is 18.9 inches. Hile (1954) reported that 50 percent of the Saginaw Bay walleyes had reached sexual maturity at 15.5 inches for the male and 17.0 inches for the females. Eschmeyer (1950) for Gogebic Lake, Michigan, showed that males mature at 12.2 inches in total length and females at 15.4 inches. In Red Lake, Minnesota, males mature at age group Table 5. Sex Composition of Age Groups of Lake Winnebago Walleyes AND (IN PARENTHESES) PERCENTAGE MATURE Age Group Number of Males Number of Females I . 58 111 II . 27 (37) 84 Ill . 62 (73) 79 IV . 60 (93) 90 (8) V. . 89 (96) 109 (32) VI . . . . . 80 (100) 77 (67) VII . . 38 (100) 36 (81) VIII . . . 11 (100) 6 (100) Total . 425 (77) 592 (22) 1969] Priegel- — Age and Growth of the Walleye 129 5 and females at age group 6 (Smith, Krefting, and Butler 1952). Balch (1951) reported that about one-half of the male walleyes are mature by the time they reach 15.5 inches and that one-half of the females in the 17-inch group were mature in Northern Green Bay waters of Lake Michigan. 3. Length- Weight Relation Length-weight relation was calculated from fish grouped by half inch total length intervals from 7.0 to 24.5 inches. There was no significant difference between sexes so all fish were combined. Length-weight relation of Lake Winnebago walleyes is expressed by the regression: Log W = -5.3596 + 3.2162 Log L where W = weight in pounds and L = total length in inches In the graphical representation of the length-weight relation (Figure 2) the smooth curve represents the calculated weights, and the dots the empirical ones. The agreement of the calculated and empirical weights was satisfactory. The discrepancies were more pronounced among the larger fish; but, on the whole, distribution of the disagreements had no particular trend. Discrepancies among the larger fish resulted from the smaller number of fish and actual weights were great enough to make relatively modest disagree¬ ments seem larger. Calculated growth in weight (Table 6) was determined by ap¬ plying calculated lengths (sum of the average increments of length) of Tables 7 and 8 to the length-weight relation. The an¬ nual increments of weight for the males increased irregularly, while the annual increments of weights for the females showed a gradual increase during the first six years. Increments in individ¬ ual years of life for the females were 0.05 pounds in the first year of life to a maximum of 0.56 pounds in the sixth year of life. It took the males five years to reach 1 pound and the females slightly under 4 years to obtain 1 pound. 4. Fecundity A few estimates have been published on the egg production of the walleye, but most of these estimates have been made on a small number of fish and the size range has been limited. Vessel and Eddy (1941) who had the largest sample (62 fish) from Cut-Foot Sioux Lake, Minnesota, estimated the egg production of walleyes from 1.5 to 5.0 pounds at 39,000-128,000 eggs. Eschmeyer (1950) estimated egg production from Lake Gogebic, Michigan, walleyes Weight (Pounds) 130 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Figure 2. Length-weight relation of the Lake Winnebago walleyes. Dots repre¬ sent the empirical data and the smooth curve is the calculated data. 1969] Priegel—Age and Growth of the Walleye 131 Table 6. Calculated Weights in Pounds at the End of Each Year of Life of Lake Winnebago Walleyes Year of Life Male Female Calculated Weight Increment Calculated Weight Increment I . 0.04 0.04 0.05 0.05 II . 0.28 0.24 0.27 0.22 Ill . 0.60 0.32 0.68 0.41 IV . . 0.84 0.24 1.11 0.43 V . 1.05 0.21 1.63 0.52 VI . 1.20 0.15 2.19 0.56 VII . 1.37 0.17 2.68 0.49 VIII . 1.59 0.22 3.08 0.40 Weights are from the general length-weight relation and correspond to lengths at the end of year of life on the general growth curve for scales taken above the lateral line. Table 7. Estimated Egg Production of Lake Winnebago Walleyes Length Groups in Inches (T. L.) 16.5- 16.9 17.0-17.4 17.5- 17.9 18.0-18.4 18.5- 18.9 19.0-19.4 19.5- 19.9 20.0-20.4 20.5- 20.9 21.0-21.4 21.5- 21.9 22.0-22.4 22.5- 22.9 23.0-23.4 24.0-24.4 25.0-25.4 Number Sampled 1 1 6 6 11 15 24 21 12 13 7 6 2 3 1 1 Average Weight of Fish 1.39 1.50 1.95 2.12 2.34 2.61 2.73 2.89 3.18 3.60 3.94 4.04 4.67 4.63 5.20 6.00 Average Weight of Ovaries (grams) 38.1 24.4 64.5 57.8 74.2 89.3 94.7 1C4.4 117.2 141.1 159.9 158.9 243.7 189.0 291.9 155.1 Average Sample Weight of Ovaries (grams) 2.0 1.8 1.9 1.9 2.3 2.3 2.5 2.7 3.5 3.3 3.2 3.2 4.0 3.1 3.5 3.6 Average Percent¬ age of Ovaries Counted 5.2 7.4 3.2 3.3 3.3 2.7 2.8 2.6 3.1 2.4 2.0 2.1 1.6 1.7 1.2 2.3 Average Calcu¬ lated Number of Eggs 62,617 43,255 65,616 77,832 93,532 98,904 105,417 108,052 114,738 138,414 148,913 143,794 89,377 169,250 227,181 127,569 132 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 (34 fish) at 36,871-154,906 eggs for fish from 16.0-22.9 inches in total length. Smith (1941) calculated that three Norris Reservoir walleyes of 25.0-26.5 inches in total length produced from 77,500- 87,400 eggs. The estimates of the egg production for walleyes by half-inch size groups are given in Table 7. The egg production ranged from 43,255 eggs for a 17.4 inch, 1.50 pound walleye to 227,181 eggs for a 24.2 inch, 5.20 pound walleye. The heaviest walleye (6.00 pounds and 25.2 inches) had a count of 127,569 eggs. Summary 1. Age determinations and growth histories were calculated by the scale method from a sample of 1,237 walleyes. 2. Body-scale relation is expressed by the formula: L = 1.443 + 3.171 R where L = total length in inches and R = anterior scale radius X 44 3. Difference in growth rate for the sexes was noted. The advan¬ tage of the females increased from 0.6 inches at the end of the third year to 4.0 inches at the end of the eight year. If the 13-inch size limit was still in effect, it would take the male walleye four years to reach legal size and the female three years. 4. The annual increments of weight for the males increased ir¬ regularly while the annual increments of weight for the females showed a gradual increase during the first six years. It took the males five years to reach one pound and the females slightly under four years to obtain one pound. 5. The average age of maturity was considered as that age at which 50 percent of the fish reach maturity. The male walleyes would be considered as mature at the end of the third year of life. At the end of the sixth year of life, all males were mature. The female walleyes were considered as mature at the end of the sixth year of life. Only female fish at the end of the eighth year of life were all mature. 6. The Lake Winnebago walleye is one of the slowest growing walleye when compared to the walleye populations of other waters. 7. The egg production of the Lake Winnebago walleye ranged from 43,255 eggs for a 17.4-inch, 1.50-pound walleye to 227,181 eggs for a 24.2-inch, 5.20-pound walleye. 1969] Priegel—Age and Growth of the Walleye 13B 8. Length-weight relation is expressed by the formula : Log W = -5.3596 + 3.2162 Log L where W = weight in pounds and L = total length in inches Literature Cited Balch, Robert. 1951. The age and growth of the yellow pike-perch, Stizoste - dion v. vitreum (Mitchill) in the Green Bay waters of Lake Michigan. Invest. Rpt. No. 652, Fish Mgt. Div., Wis. Cons. Dept., mimeo. Butler, Robert L. and Lloyd L. Smith, Jr. 1953. A method for cellulose ace¬ tate impressions of fish scales, with a measurement of its reliability. Prog. Fish. Cult., 15:175-178. Carlander, Kenneth D. 1945. Age, growth, sexual maturity, and population fluctuations of the yellow pike-perch, Stizostedion vitreum vitreum (Mitchill) with reference to the commercial fisheries, Lake of the Woods, Minnesota. Trans. Amer. Fish. Soc., 73:90-103. Cleary, Robert E. 1949. Life history and management of the yellow pike- perch, Stizostedion vitreum vitreum (Mitchill) of Clear Lake, Iowa. Iowa St. Univ. J. Sci., 23(2) :195-208. Eddy, Samuel and Kenneth D. Carlander. 1939. Growth of Minnesota fishes. Minn. Conservationist 69:8-10. Eschmeyer, Paul H. 1950. The life history of the walleye, Stizostedion vitreum vitreum (Mitchill) in Michigan. Bull. Inst. Fish. Res. (Mich.) 3:1-99. Hile, Ralph. 1948. Standardization of methods of expressing lengths and weights of fish. Trans. Amer. Fish. Soc. 75:157-164. - . 1954. Fluctuations in growth and year-class strength of the walleye in Saginaw Bay. U.S. Fish and Wildlife Service, Fish. Bull. 56:7-59. Priegel, Gordon R. 1964. Early scale development in the walleye. Trans. Amer. Fish. Soc. 93 (2) : 199-200. Rose, Earl T. 1951. Notes on the age and growth of Spirit Lake Yellow pike- perch ( Stizostedion v. vitreum). Proc. Iowa Acad. Sci. 58:517-526. Roseberry, Dean A. 1951. Fishery management of Claytor Lake, an impound¬ ment on the New River of Virginia. Trans. Amer. Fish. Soc. 80:194-209. Schloemer, Clarence L. and Ralph Lorch. 1942. The rate of growth of the walleye pike, Stizostedion vitreum vitreum (Mitchill) in Wisconsin’s in¬ land waters, with special reference to the growth characteristics of the Trout Lake population. Copeia. No. 4, pp. 201-211. Schmulbach, James C. 1959. Growth of the walleye in the Des Moines River, Boone County, Iowa. Proc. Iowa Acad. Sci., 66:523-533. Smith, Charles O. 1941. Egg production of walleyed pike and sauger. Norris Reservoir fish differ from some species in other localities. Prog. Fish Cult. 54:32-34. Smith, Lloyd L., Jr., Laurits W. Krefting and Robert L. Butler. 1952. Movements of marked walleyes, Stizostedion vitreum vitreum (Mitchill) in the fishery of the Red Lakes, Minnesota. Trans. Amer. Fish Soc. 81:179-196. Smith, Stanford H. 1954. A method of producing plastic impressions of fish scales without the use of heat. Prog. Fish Cult. 16:75-78. Strong, Richard H. 1949. Growth of Norris Reservoir walleye during the first twelve years of impoundment. Journ. Wildl. Mgt. 13 (2) : 157-177. Vessel, Matt F., and Samuel Eddy. 1941. A preliminary study of the egg production of certain Minnesota fishes. Minn. Bur. Fish Res. Invest. Rept. 26:26 pp. REGULARLY OCCURRING FLUCTUATIONS IN YEAR-CLASS STRENGTH OF TWO BROOK TROUT POPULATIONS Ray J. White and Robert L. Hunt * During a 12-year period (1953-04) year class strength of wild brook trout (Salvelinus fontmalis) fluctuated rhythmically in two central Wisconsin streams, Lawrence Creek (Adams and Mar¬ quette Counties) and Big Roche-a-Cri Creek (Adams and Wau¬ shara Counties). In this paper we discuss the nature of these rhythmic year class fluctuations and examine some possible reasons for their regularity. Population dynamics of trout have been studied since 1953 in Lawrence Creek primarily to test angling regulations (emphasis switched to trout habitat management in 1964), and since 1957 in the Roche-a-Cri to assess effects of trout habitat management. Normal statewide angling regulations for trout applied to the Roche-a-Cri : season from early or mid-May until September 7, a minimum legal size limit of 6 inches and a bag limit of 10 trout per day. Angling pressure was greatest during seasons when trout were most abundant. At Lawrence Creek, length of the fishing sea¬ son was the same, but 6 combinations of experimental restrictions on size, bag and gear were tested in various parts of the stream during 1955-64. Changes in angling regulations caused angler ex¬ ploitation of the trout population to vary greatly (Hunt, Brynild- son and McFadden, 1962 ; Hunt and Brynildson, 1964; Hunt, 1964). Description of the Streams Although the streams lie only 18 miles apart, their physical, hydrological and vegetational characteristics differ. Lawrence Creek usually has somewhat better living conditions for brook trout. The stream arises in a terminal moraine and flows eastward through a rolling landscape of glacial drift. Our study area, extend¬ ing from the headwaters to an artificially impounded lake, com¬ prises 8.4 stream miles. The Roche-a-Cri, directly north of Law¬ rence Creek, flows westward from the same moraine across a * Mr. White, formerly a Biologist 'with the Wisconsin Conservation Division, is now at the Laboratory of Limnology, University of Wisconsin — Madison. Mr. Hunt is leader of the Trout Research Group of the Wisconsin Conservation Division. 135 136 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 glacial outwash plain. The study area covered in this paper includes the upper 6.1 miles of trout water. Each stream passes through patches of forest, marshy meadows and thickets of brush. Many distinct springs feed Lawrence Creek. Its discharge at the down- stream end of the study area is about 25 cfs during periods of base- flow. The streambed falls an average of 11.5 feet per mile. In con¬ trast, the Roche-a-Cri receives less ground water seepage and flows through flatter terrain (its gradient = ca. 7 ft. /mile). At a point 4 miles below the stream’s source, baseflows vary from 5 to 9 or more cfs depending on recent precipitation. Figure 1 contrasts the relatively stable discharge of Lawrence Creek with the greatly fluctuating streamflow of the Roche-a-Cri. Figure 2 shows monthly low-flows for the Roche-a-Cri. These low-flow data give some indi¬ cation of varying limitations on space available to brook trout in that stream. With its greater discharge and steeper gradient Lawrence Creek has larger areas of gravel streambed relatively free of sand and silt. In summer and fall an abundance of watercress (Nasturtium officinale) and veronica (Veronica connata) offer hiding cover for trout and support a rich trout food supply. In the Roche-a-Cri, Figure 1. Monthly mean streamflow discharges at gaging stations on Law¬ rence Creek (2.5 miles below its source) and on Big Roche-a-Cri Creek (3.9 miles below source). 1969] White and Hunt — Fluctuations in Brook Trout 137 Figure 2. Lowest streamflow discharge recorded each month at the Big Roche- a-Cri Creek gage. sand covers most of the bottom. Here, perhaps owing to the greater variability of streamflow, watercress beds often do not develop until autumn. Only in years of ample rainfall or during a general trend of streamflow increase, for instance in 1959-60 (Figures 1 and 4) , does the Roche-a-Cri’s cress flourish in springtime or early summer. Three miles of the Roche-a-Cri that flowed through grazed land were fenced during 1956-58 to keep cattle away from the stream. Fencing allowed streambank vegetation to thrive and provide trout with hiding places among twigs and leaves that dangled into the water. From 1958 through 1962 current-deflectors and overhang¬ ing bank covers were constructed along 4 miles of stream in the study area. These devices added more cover for trout and concen¬ trated the current to clean sand off streambed gravel. The Trout Populations Wild brook trout are the predominant fish in both streams. Dur¬ ing our investigations, trout population densities ranged from less than 20 pounds per acre in some sections of both streams in poor years to nearly 250 pounds per acre in upstream sections of both streams during favorable years. But on the average, Lawrence Creek maintained larger populations. The Roche-a-Cri’s main con¬ centration of brook trout occupies a section of stream approxi¬ mately equal to Lawrence Creek in length, but this section has only half the surface area of Lawrence Creek. Few wild brown trout ( Salmo trutta) or rainbow trout (S. gairdneri ) occur in the Roche-a-Cri, though at times during sev¬ eral decades prior to the study these species were heavily stocked in the study area as well as in the 20 or more miles of water tol- 138 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 erable to trout (but lacking spawning grounds) below the study area. Even within the study area where brook trout reproduce well, brown and rainbow trout spawn with poor success. At vari¬ ous times during the study small numbers of hatchery-reared brown and rainbow trout moved into the lower portion of the study area from stocking sites several miles downstream. During 1953- 60 about 1,200 age-0 and age-1 hatchery-reared brook trout were also stocked annually in the study area as part of the routine fish¬ ery management program. However, in view of the usual high mor¬ tality of stocked trout during the first few weeks after release, this stocking was probably a minor supplement to the total spring¬ time population of trout in the study area. Hatchery brook trout are not included in the data to be discussed since the 1 % to 3% of such stocked trout that survived to maturity did not contribute significantly to total spawning. Other fishes common in the Roche- a-Cri are: mottled sculpin (Cottus bairdi), pearl dace (Margaris- cus margarita), creek chub (Semotilas atromaculatus ) , brook stickleback (Eucalia inconstans) and white sucker (Catostomus commersoni) . Lawrence Creek is free of brown trout and has only a sparse population of wild rainbow trout. The stream has not been stocked with hatchery trout since 1948. Other than trout, Lawrence Creek contains mottled sculpin, white sucker, creek chub, brook stickle¬ back and blacknose dace (Rhinichthys atratulus) . Methods Trout populations were estimated by mark-and-recapture elec¬ trofishing in April, prior to the angling season, and in September soon after angling ceased. In this paper September estimates of age group 0 are used as initial measures of year class strength. Since age-0 trout are too small in April to be efficiently captured by our electrofishing gear, they could not enter into estimates at that time. During electrofishings in September, 60% to 80% of the age-0 trout were marked by removing fins in combinations denoting year class, i.e., the year of birth. Population estimate procedures, pre¬ cision of the estimates (± 2 to 6% for age group 0; ± 1 to 8% for age group I, but generally around ± 2 to 4% for both groups) and methods of calculating egg production annually are discussed by McFadden (1961), Hunt et ah (1962) and Hunt (1966). Results and Discussion In both streams year-to-year fluctuations in abundance of age-0 brook trout followed a rhythm of alternating upward and down- THOUSANDS OF TROUT 1969] White and Hunt — Fluctuations in Brook Trout 139 ward turns, that is, one having 2 years between peaks (Figure 3; Table 1). Among the combined 20 data-years for the 2 streams, encompassing 18 between-year changes in population level, the pat¬ tern was interrupted only once : the 1956-57 interval at Lawrence Creek. In Lawrence Creek numbers of age-0 brook trout present in September varied from 4,166 in 1958 to 22,646 in 1959. The mean number present during 12 successive Septembers was 10,712. The strongest year class was 5.4 times larger than the weakest year class and 2.1 times larger than the average numerical strength. In the Roche-a-Cri over an 8-year period, the number of age-0 brook trout present in September ranged from 2,012 in 1957 to 9,915 in 1960. Mean strength of the 8 year classes was 5,396. The strongest year class was 4.9 times as numerous as the weakest year class and 1.8 times larger than the mean abundance (Table 1). Keith (1962) suggests that most biologists would favor the definition of “population cycle” as defined by Davis (1957) : “In ecological usage the term ‘cycle’ refers to a phenomenon that occurs at intervals. These intervals are variable in length, but it is implied that their variability is less than one would expect by chance and that reasonably accurate predictions can be made.” Figure 3. Numbers of brook trout in the study areas of Lawrence and Big Roche-a-Cri Creeks. (Although no inventory of the Roche-a-Cri population was made in spring, 1960, the number of age group I trout was probably low since the 1959 year class was a weak one.) 140 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Table 1. Age Structure of April and September Populations of Wild Brook Trout in Lawrence and Big Roche-a-cri Creeks Year Lawrence Creek April September Age I Age 11 + Age 0 Age I Age 11 + 1953 . 10,113 2,040 277 1954 . . . . — — 13,523 2,749 296 1955 . 7,782 1,647 5,720 2,754 324 1956 . 2,012 1,207 10,853 816 133 1957 . 7,029 483 13,258 3,370 251 1958 . 8,485 2,069 4,166 4,393 635 1959 . 1,815 2,707 22,646 1,044 654 1960 . 8,510 409 8,507 3,324 51 1961 . 3,602 842 14,313 2,360 246 1962. . 8,567 1,221 7,611 4,523 225 1963 . 4,644 2,540 10,367 2,388 792 1964 . 7,489 1,669 9,680 4,382 569 Average 1955-64 . 5,993 1,479 10,712 2,935 388 Big Roche-a-Cri Creek Year April September Age I Age 11 + Age 0 Age 1 Age 11 + 1953 . 1954. . . . — — — • - 1955. ....... — — — — — 1956 . — — — ■ — . — 1957 . 2,034 333 2,012 1,135 15 1958 . 1,743 634 6,229 474 24 1959 . 3,749 152 2,637 1,817 58 1960 . — — 9,915 1,257 220 1961 . 5,038 * 4,361 2,630 156 1962 . 2,030 * 5,632 1,609 422 1963 . 3,262 * 4,964 1,623 256 1964 . 1,925 * 7,420 1,072 218 1957-64 . 2,826 — 5,396 1,452 171 *Not yet calculated. 1969] White and Hunt — Fluctuations in Brook Trout 141 According to this definition the fluctuations we observed con¬ stitute cycles of offspring abundance. Both cycles followed 2-year intervals, the shortest possible interval for an animal that repro¬ duces once annually. Although the frequency of fluctuation was regular, levels of abundance did not always swing above and below a long-term mean value, as would be necessary to meet the strict mathematical definition of a cycle. Our contention is, however, that there did seem to be a recurring ecological phenomenon worthy of critical examination. The rhythmic fluctuations in the 2 streams, while of equal fre¬ quency, were out of phase. In the years when Lawrence Creek con¬ tained large numbers of age-0 trout, the Roche-a-Cri had low numbers. This phase difference persisted all 8 years of simultane¬ ous study. The cycles persisted in both streams despite somewhat different environmental characteristics and different general levels of trout populations and they persisted despite large changes in density of total stocks and in age composition of those stocks in each stream (Table 1). Great variation in percentage of brook trout stocks removed from Lawrence Creek by anglers also failed to upset the cycle. Under liberal angling regulations — toward the beginning of the study-anglers took 32% to 65% of preseason populations and as much as 129% by weight. Under very restrictive size and bag limits, angler exploitation fell to 1 % by number and 7% by weight (Hunt et al., 1962). These relatively low angler harvests resulted in higher populations of older trout and egg production became less dependent upon age-I trout as more adults survived the fishing season to spawn a second or third time. These changes in survival rates and structure of the spawning stocks may have contributed to the lower amplitude of fluctuations in age group 0 populations in Lawrence Creek during the later years of our study. In the Roche-a-Cri, the cyclic fluctuations also diminished in amplitude toward the end of the study as populations of age-II and older trout increased (Table 1). Progress in the management of trout habitat may have induced these changes, but despite these trends of improvement in the trout habitat and the trout population, the cycle persisted. It is especially interesting that the cycles were out of phase in the 2 streams. Consequently it is unlikely that a climatological factor was governing the cycles. If that component of the climate, whatever it may be, which is most influential on trout would have had an alternate-year fluctuation during the period of study, it should have affected both trout populations similarly. 142 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 This is not to say that climatological changes failed to affect the trout populations, and in particular age group 0. The highest age-0 population in the Roche-a-Cri, that of 1960, coincided with sharp increases in precipitation (Figure 4) and in streamflow (Figure 2) during autumn 1959 through summer 1960. Not only was vol¬ ume of the Roche-a-Cri greater than usual in the spring and sum¬ mer of 1960, but in-stream vegetation, particularly watercress, was comparatively lush during May, much earlier than in most years. Space and concealment for young trout was undoubtedly greater than normal-— and, judging by Tarzwell’s (1937) findings on the food-harboring capacities of underwater vegetation, the trout food supply was probably better too. Such rather sudden en¬ vironmental improvements, coinciding with whatever factors cause the “even-year” (1958, 1960, 1962, etc.) peaking on the Roche-a- in UJ i o z z o < CL O UJ cr Q. Figure 4. Numbers of age-0 brook trout in September in Big Roche-a-Cri Creek and records of precipitation from its vicinity. Plotted above each trout level are precipitation for September through November of the 'preceding year and precipitation during the 12 months (October-September) prior to trout inventory, i.e. the approximate period covered by development of age group 0. Precipitation data are from the Hancock Experimental Farm, 5 miles south of the stream and from the U. S. Weather Bureau compilation for central Wisconsin stations. 1969] White and Hunt — Fluctuations in Brook Trout 143 Cri, probably reinforced the upward amplitude for one year of the cycle. Lawrence Creek, with a more stable hydrological regime, appeared to have no unusual streamflow during 1960 (Figure 1). Similarly but conversely, an environmental event in Lawrence Creek could have accounted for the upset of the cycle in that stream during 1957. According to qualitative observations by one of the authors (White), watercress was unusually abundant in Lawrence Creek that year. While regional rainfall was low in 1956-57 (Figure 4) and it is unlikely that local variation was great enough to have stimulated aquatic vegetation through increased streamflow; nevertheless, throughout most of the study area the stream was walled with cress on each side to an extent not noted since. Perhaps at no other time during our study did age-0 brook trout in Lawrence Creek have such good hiding cover. Thus, there is some reason to suspect that a low phase of the population cycle was counteracted by an environmental change especially favorable to survival of young trout during 1957. In any event, the cycle was interrupted that year and thrown into a new phase, one having highs in odd numbered years. Environmental crises, on the other hand, might be expected to exaggerate cyclic lows and cancel highs, but no phenomenon of this sort was evident. Despite the unlikelihood that climate maintained the cycles, monthly and seasonal streamflow data for the Roche-a-Cri (Fig¬ ures 1 and 2), were examined to see if there were any patterns of fluctuation that coincided with the fluctuations of year class strength in that stream. No similarities were found. However, since our streamflow records did not cover the first year of the study and were not complete with respect to high flows, a search was made through precipitation data from a weather station near the Roche- a-Cri. (There is none near Lawrence Creek) . One set of these data, precipitation during September through November, showed a pat¬ tern of year-to-year fluctuations resembling that of the trout cycle (Figure 4). While rainfall might be interpreted as having influ¬ enced streamflow at spawning time (October into early December) and hence as having affected success of natural reproduction, there is nothing in the streamflow data to support this contention. Neither is September-N ovember precipitation significantly corre¬ lated with the following-year abundance of age group 0. Conse¬ quently, some mechanism intrinsic to both brook trout populations and acting after the egg stage seems a more likely regulator of the cycles. If, as is the case in these two populations, age group I com¬ prises the great majority of the “mature” (age I and older) trout, then such 2-year cycles could, given an initial disparity between 144 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 any two consecutive year classes, be simply and directly maintained through the following processes: (1) domination of egg production by age group I which results in alternate-year variation similar to that of “even-year” and “odd-year” pink salmon (Onchyrhyn- chis gorbuschka) populations (Neave, 1952 and 1958), (2) a sup¬ pressive effect (predation and/or competition) of age group I on simultaneously occurring age-0 populations. Such suppression would be strong and weak in alternate years. Should both proc¬ esses occur within the same population, they would complement each other, that is, act in synchrony. High egg production would push age group 0 upward one year and strong suppression by numerous age I trout from the same year class would push age group 0 downward the next year (Figure 5). For these two brook trout populations, however, the first process seems unlikely. Even though most of the egg production in most years is attributable to age group I (Table 2), numbers of age 0 trout in September are not correlated with the numbers of eggs from which they originated (Figure 6). Since mortality at the egg and sac-fry stage is known to be less than 10% (McFadden, 1961) and since June inventories (Hunt, 1966) show that age-0 mortal¬ ity progresses at a rather slow rate after the 4th or 5th free- swimming month, most age 0 mortality — and, in fact, the greater share of total mortality in the life of a year class— -takes place dur¬ ing the first few months following emergence of fry into the stream. The second regulatory mechanism, the suppression process, seems more plausible. Age-I trout could be preying on age-0 trout or could be limiting in some way the accessibility of a rather fixed environmental resource. Indeed, numbers of age-0 trout in Lawrence Creek are inversely and significantly correlated with numbers of simultaneously occurring age groups I (Figure 7 and Table 8). If age-I trout were determining the level of age-0 abundance, the time of this effect would most logically be during - ->. Figure 5. Curves of numerical trends within year classes of a hypothetical brook trout population in which: (1) reproduction is entirely by age group I, (2) strong year classes, producing high numbers of eggs, spawn strong year classes, (3) weak year classes spawn weak ones, (4) many age-I trout are present during the fry stages of weak year classes and (5) few age-I trout exist during the fry stages of strong year classes. Survivorship after com¬ pletion of spawning is approximated from Lawrence Creek data (Hunt, 1966, Appendix Table 22; and McFadden, 1961) and is kept uniform for all year classes, thus eliminating compensatory complications. Were age group I suppressing age-0 abundance, survivorship curves of weak year classes should be considered steeper than shown here. SPAWNING BY STRONG YEAR CLASS 1969] White and Hunt— Fluctuations in Brook Trout 145 DURING PERIODS OF HIGH FRY MORTALITY Table 2. Estimated Number of Eggs Produced by Brook Trout of Various Age Groups in Law- 146 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 fa < fa £ fa Eh O H fa o fa fa < £ 02 fa m 9 fa fa £ 0 | I* Ph g k 1-1 < % g 2 £ fa g§ fa n . o !z fa c w fa fc r^vfl'-'OONNO'OO'NK NOOO'KOOOOOOf^Kn NvOf^O'ONOOts^N'O ^hVwDvm^O'^vDK'-''^ ONt\Os\£>00c^t\00— '\0'— 1 A OO^N'tvO-'OOOOOOON QOO-"Mr\OO^N^O ICvCOsOCC^'^frCO' - O\£J'tTtNNN00^VA,i< OvKOONO';t^Or'NK'- NO'A'O'Af^1 — 'OOOf'M^ '^fu-s(N\Dt\-^lO,--|rNSOsoo — I I —I I CO CO CO K KKKOOtX'tO'KOOxO •— > 00 1 u-\ ir\ cO vr\ O CO irs O' CiOOOOO'O'tOOO'O^ 1 vrs — i OO'rO'^KO'ONiA OncOOO\DO''D’-hOO'cOCi re > — iaO'c^vnOvDOOia^ i^O^OOics n^icsONCOlJ'C — con m vrs i/e iaUmaiavOvOvO^O O'O'O'O'O'O'O'O'O'O'O' bD 03 s_ t) > < 1969] White and Hunt — Fluctuations in Brook Trout 147 THOUSANDS OF EGGS Figure 6. Numbers of age-0 brook trout in Lawrence Creek in September plotted against numbers of eggs from which they developed. Numbers by the points denote year classes. Solid points are those for year classes coinciding with lower-than-average April abundances of age group I. spring when fry mortality is greatest. Hence, age-0 levels would be more closely correlated with springtime rather than autumn levels of age group I. The coefficient of correlation for age groups 0 against April age-I populations is higher than the correlation for age groups 0 against September age-I populations (Table 3), but with the low number of observations involved, the difference between the 2 coefficients is not statistically significant. 148 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Table 3. Coefficients of Correlation and Values of Student’s t for Regressions of Numbers of Age-0 Brook Trout in September on Numbers of Older Trout in Lawrence Creek During 1953-64 Regression of Number of Age 0 in September on: Number of Years of Observations (n) Coefficient of Correlation (r) (-VALUE (d/=n- 2) April Populations — 3.218* Number of age I . 10 -.753 Number of age I plus older trout. . 10 -.732 -2.977* Number of age 1 1 plus older trout . 10 .205 0.592 n.s. Total weight of all trout . 9 -.174 -0.055 n.s. September Populations Number of age I . 12 -.608 -2.421* Number of age I plus older trout. . 12 - .564 — 2. 161 n.s. April and September Populations Averaged -3.017* Number of age I . 10 -.729 Number of age I plus older trout . . 10 -.702 -2.781* ^Significant at the 0.05 confidence level, n.s. =not significant at the 0.05 level. Numbers of age-0 trout are not correlated with numbers of trout older than age-I. Perhaps body size or behavior of these older trout reduce interactions with post-emergent young. Cannibalism or competition for space and/or food are the sorts of processes that would tend to cause an inverse correlation such as that in Figure 7.1 Cannibalism has been cited in other studies as a factor accounting for changes in abundance of young fish, but conclusive evidence of cannibalism in our streams is lacking.2 For 1 A disease-crowding- process would seem a further but less likely possible mechanism behind the observed relationship. All we know of disease in these streams is that almost every brook trout carries gill copepods ( Salmincola edwardsii), that emaciation of smaller trout due to this burden appears worse than that of larger trout, and that during times of high trout population density the number of copepods on each trout’s gills appears greater than during population lows. It is thought that the passing on of relatively high infestations of copepods by the higher populations of age-I trout to fry would be too erratic to account for the inverse relationship of age-0 and age-I densities (Figure 8). 2 Cannibalism was postulated as the governor of a possible 4-year cycle among brown trout in 2 New Zealand streams (Burnett, 1959 — another out-of-phase cycle!). Survival of stocked fingerling brook trout increased when older brook trout were re¬ moved from a lake (Smith, 1956). When the number of larger trout in a small lake increased, survival of stocked fingerling rainbow trout decreased (Miller and Thomas, 1957). There are logical yet indirect indications that predation on sockeye fry /, Onchyrhynchus nerka) by smolts and residual non-migrants of preceding strong year classes may cause the 4-year cycle of that salmon in the Fraser River (Ricker, '1950). In a mixed population of warm water fishes in an Illinois lake, 4- or 5-year cycles of abundance were attributed to predation by dominant broods of crappies (Pomoxis sp. ) but no data on predation were furnished (Thompson, 1941). 1969] White and Hunt— Fluctuations in Brook Trout 149 THOUSANDS OF AGE I BROOK TROUT IN APRIL Figure 7. Numbers of age-0 brook trout in September plotted against age-I abundance during April of the same year in Lawrence Creek. Numbers by the points denote the year classes of age groups 0. Correlation is significant at the 0.05 confidence level (Table 2). 150 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 example, during 1960-66, we examined 1,400 stomachs of age-I- and-II brook trout collected in all seasons by daytime electrofishing and angling. Not a single case of cannibalism was found (unpubl. data of R. L. Hunt and D. A. White). To account for even 10% of the average mortality of age group 0 between emergence and mid-J une, each fish in the average population of older trout during that time would have to consume approximately 15 fry per day. However, no fry were found in 160 stomachs collected from adult trout during such a 5-month period. Although cannibalism among brook trout has been demonstrated elsewhere,3 there are no indica¬ tion that it takes place on a scale sufficient to account for the hun¬ dreds of thousands of fry that perish in Lawrence Creek and Roche-a-Cri each spring. Competition rather than predation seems a more likely relation¬ ship between age-0 and age-I brook trout, especially in view of the investigations of LeCren (1965) who varied the numerical density of brown trout fry in small experimental waterways and found their survival and growth to be inversely density-dependent. This result was apparently due to ferritorial behavior. Fry not able to secure a territory drifted downstream and died by starvation usually between 20 and 40 days before feeding began. Density- dependent “disappearance” of age-0 trout from Lawrence Creek is indicated in analyses of September age-0 levels as a function of egg production plus age-I abundance (McFadden, in press). A rough representation of such an analysis can be seen in the nega¬ tive slope of the solid black points in Figure 6. Greater dispersal of young from main nursery areas in years of higher age-0 density (Hunt, 1965) implies that the apparent compensatory mortality in Lawrence Creek prior to September may be partially attrib¬ utable to movement downstream out of the study area. If space competition from age-I trout is similarly affecting fry or finger- lings, this could account for the observed numerical relationships between the 2 age groups, and for the 2-year cycle of yearclass abundance. Although the number of age-0 trout that disappear is not corre¬ lated with the number of contemporary age I trout (Figure 8), age 3 Following release of several thousand brook trout fry into a small Ontario stream, collection of 16 age-I-and-II brook trout mainly by means of a flashlight and handnet at night revealed 4 stomachs containing fry, the greatest number in any one stomach being 8 (H. C. White, 1924). After stocking fry in a Prince Edward Island stream, 319 “adult and yearling” brook trout were captured, “those whose stomachs were distended” (number not reported) were examined, and only one fry was found (H. C. White, 1927). In sections of the same stream, screened to permit only certain types of predators to operate on fry stocks of known size, comparison of mortality rates led to the conclusion that “competition and cannibalism” by larger trout were greater menaces to fry than were predation by birds or sticklebacks ( Eucalia inconstans), but no proof of cannibalism was put forth (H. C. White, 1927 and 1930). 1969] White and Hunt— Fluctuations in Brook Trout 151 group I may act as a “level-setter” of year class strength. The “rather fixed environmental resource”, be it space, food or what¬ ever, could determine a general numerical density (carrying capacity) to which age group 0 must diminish each year regard¬ less of the number of age-I trout. With this carrying capacity at a level of (for instance) 40,000 to 80,000 fry, the greatest share of post-emergence mortality has occurred by the time this level is attained. The amount of additional fry mortality that might be dependent upon the density of age I trout would be a relatively small component of total fry mortality but an important component in the ultimate determination of year class strength. While such a mechanism may bring cannibalism back into the realm of possi- CL LU 10 Figure 8. Absolute mortality of brook trout in Lawrence Creek during the 10 to 11 months from the egg stage to the next September plotted against number of age-I trout present during April of this period. Numbers by the points denote the year classes of age groups 0. 152 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 bility, the negative evidence regarding such predation in Lawrence Creek makes competition seem the more likely process needing further investigation. To investigate more closely relationships between stocks of age-I and age-0 trout, experiments similar to those of LeCren (1965) should be conducted utilizing various sizes and densities of age-I trout introduced as factors modifying existing fry-space relation¬ ships. Suppression of age-0 brook trout by older brook trout could also be tested under the more natural conditions in our streams by measuring survival of trout fry at selected high and low spring¬ time densities of older trout. Low densities of older trout could be achieved by electrofishing to remove them in winter after the spawning season. High densities could be attained by stocking wild brook trout from other nearby streams. Our study points up the need for identifying the causes of mor¬ tality among wild trout fry and for direct observation of behavioral relationships between age groups and size groups of brook trout under wild conditions. Acknowledgements For helpful suggestions during preparation of the manuscript, we are grateful to D. R. Thompson and W. S. Churchill of the Wis¬ consin Conservation Division, Madison, and to A. Buckmann, M. Gillbricht and G. Hempel of the University of Hamburg, Germany. References Cited Burnet, A. M. R. 1959. Some observations on natural fluctuations of trout population numbers. N. Z. J. Sci. 2:410-421. Davis, D. E. 1957. The existence of cycles. Ecology 38(1) : 163-164. Hunt, R. L. 1964. Evaluation of fly-fishing-only at Lawrence Creek (a three- year progress report). Misc. Res. Rept. No. 10 (Fisheries). Wis. Conserv. Dept., Madison, Wis. (mimeo). - . 1965. Dispersal of wild brook trout during their first summer of life. Trans. Am. Fish. Soc. 94 (2) :186-188. - . 1966. Production and angler harvest of wild brook trout in Law¬ rence Creek, Wisconsin. Tech. Bull. No. 35. Wis. Conserv. Dept., Madison, Wis. - , and O. M. Brynildson. 1964. A five-year study of a headwaters trout refuge. Trans. Am. Fish. Soc. 93 (2) : 194-197. - , O. M. Brynildson and J. T. McFadden. 1962. Effects of angling regulations on a wild brook trout fishery. Tech. Bull. No. 26. Wis. Conserv. Dept., Madison, Wis. Keith, L. 1962. Wildlife’s ten-year cycle. University of Wisconsin Press. LeCren, E. D. 1965. Some factors regulating the size of populations of fresh¬ water fish. Mitt. Internat. Verein. Limnol. 13:88-105. 1969] White and Hunt — Fluctuations in Brook Trout 153 McFadden, J. T. 1961. A population study of the brook trout, Salvelinus fontinalis . Wildl. Monog. No. 7. - . (In press.) Dynamics and regulation of salmonid populations in streams. A symposium, “Salmon and Trout in Streams”, sponsored by the H. R. MacMillan Lectures in Fisheries at the University of British Columbia, February 22-24, 1968. Miller, R. B., and R. C. Thomas. 1957. Alberta’s “pothole” trout fisheries. Trans. Am. Fish. Soc. 86:216-218. Neave, F. 1952. “Even-year” and “odd-year” pink salmon populations. Trans. Roy. Soc. Canada XLVL: 55-70. - . 1953. Principles affecting the size of pink and chum salmon popula¬ tions in British Columbia. J. Fish. Res. Bd. Can. 9 (9) :450-491. Ricker, W. E. 1950. Cycle dominance among the Fraser sockeye. Ecology 31: 6-26. Smith, M. W. 1956. Further improvement of trout angling at Crecy Lake, New Brunswick, with predator control extended to large trout. Can. Fish Culturist 19:13-16. Tarzwell, C. M. 1937. Experimental evidence on the value of trout stream im¬ provement in Michigan. Trans. Am. Fish. Soc. 66:177-187. Thompson, D. F. 1941. The fish production of inland streams and lakes. In A symposium on hydrobiology. Univ. of Wis. Press: 206-217. White, H. C. 1924. A quantitative determination of the number of survivors from planting 5,000 trout fry in each of two streams. Contr. Can. Biol. Fish. n. s. 2(9) : 137-139. - . 1927. A preliminary report on trout investigation in Forbes Brook P. E. I. in 1925 and 1926. Contr. Can. Biol. Fish. n. s. 3(15) :365-376. - . 1930. Trout fry planting experiments in Forbes Brook P. E. I. in 1928. Contr. Can. Biol. Fish. n. s. 5 (8) : 203-212. DISTRIBUTION, STANDING CROPS, AND DRIFT OF BENTHIC INVERTEBRATES IN A SMALL WISCONSIN STREAM' John J. Peter ka? Department of Natural Sciences Wisconsin State University Platteville , Wisconsin Abstract Bottom samples showed genera of Tricorythodes , Stenonema , Cheumatopsyche and Chimarra occurred more frequently at down¬ stream than upstream stations, while Neophylctx occurred only near the main spring. Principal organisms found in the stream drift were Gammarus pseudolimneaus Bousfield and Baetis vagans McDunnough. Both had higher drift rates at night than during the day. Total stream drift of Gammarus caught upstream apparently did not affect the drift caught 152 m downstream. Introduction The many small spring-fed streams in southwestern Wisconsin appear to be excellent for studies of the ecology of an entire stream and its watershed. They are short (some less than 800 m) and narrow (some 0.3 m wide) ; they receive a constant supply of spring water ; and they appear productive in benthic invertebrates. The purpose of this study, undertaken in one such stream, was (1) to determine the standing crops of its principal organisms and (2) to measure the kinds and quantities of organisms (drift) carried by the stream current. Stream drift, an important source of food to fishes, has been reported by Needham (1938), Bendy (1944), Waters (1962 a), Miller (1963), and others. Waters (1962 b) used drift rates and standing crop measurements to determine the production of inver¬ tebrates appearing in the drift. The feasibility of using his method in this stream was examined. 1 A grant from the Research Board of the State Universities of Wisconsin supported the research. 2 Present address : Division of Natural Sciences, North Dakota State University, Fargo, North Dakota 58102. 155 156 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 Description of the Stream Samples were taken from Bear Branch (local name), a small stream in Grant County, Wis. (T3N, R2W, Sec. 32). The stream flows south and receives a constant supply of water from a spring located 1,675 m from the mouth at the Little Platte river. Stream widths range from .6 to 2.5 m. Average depths in riffle areas are 7 to 10 cm. Following rainstorms, water levels had extreme fluctu¬ ations. A normal discharge rate, during October 1965, was 0.21 m3/sec. Above the spring, the stream is often dry and no samples were taken there. The temperature of the spring water is fairly constant through¬ out the year, ranging from 9.0 to 9.5°C in 1965. During the sum¬ mer, water temperatures increased rapidly downstream from the spring. For example, on 1 August 1965, temperatures were 14.0, 16.5, 19.0, 23.0, 24.0 and 28.0°C at the respective distances below the spring of 180, 240, 300, 485, 850 m and at the mouth. In the winter, relatively warm spring water resulted in higher tempera¬ tures near the spring than downstream. On 24 January 1965, the temperature dropped from 9.0°C at the spring to 7.0 and 4.0°C at 240 and 850 m, respectively, below the spring. Because the stream is shallow, water temperatures fluctuate during the day. At 240 m below the spring, they were 13 to 15 °C October 16, 8 to 10°C October 23, and 8 to 9°C November 19, 1965. Total alkalinity of the stream water during July 1966 ranged from 303 to 308 ppm, as determined by titration with .02N H2S04, and methyl orange as an indicator. The upper 300 m of the stream is relatively straight, with pools, riffles and a growth of watercress, Nasturtium officinale. Below this, the stream meanders and widens. No trees or shrubs overhang it, and its banks are well-cropped by cattle. Materials and Methods Bottom samples were collected in riffle areas of the stream with a sampler (Waters and Knapp, 1961), which encompassed 0.1 m2 of bottom; the mesh size of its net was 256 jn. Three stations were sampled. Stations 1 and 4, located 60 and 240 m, respectively, below the spring, will be referred to collectively as upstream areas ; station 15, 910 m below the spring, as the downstream area. Organisms found in the drift were captured with stationary nets (Waters, 1962 a,b) . Each was made of 256 ^ mesh Nytex3 nylon netting, and fitted to a 0.3 X 0.2 m frame made of brass welding 8 Trademark of Tobler, Ernst and Traber, Inc., New York. 1969] Peterka — Invertebrates in Small Wisconsin Stream 157 rod. The nets were held in place by iron rods driven into the stream bed. To establish the total daily drift passing from riffle areas, the stream was completely blocked by drift nets at the downstream end of the station 4 riffle for 24-hr periods on 7 June, 31 August- 1 September and 22-23 October 1966, and at the downstream end of a riffle at station 2 (91 m below the spring) on 7 June. The nets were lifted, emptied and replaced in intervals usually ranging from 1 to 3 hrs. Wet weights were determined by centrifuging the organisms in wire-mesh cones to remove surface moisture, and then weighing them to the nearest 0.0001 g on an optical analytic balance. Samples were preserved in 5% formalin and no corrections were made for weight loss caused by preservation. Results Qualitative Analysis of Bottom Samples Invertebrates found in bottom samples from one station were sometimes scarce or absent at others. Nymphs of mayflies belong¬ ing to the genera Tricorythodes and Stenonema were common at station 15, infrequent at station 1 and absent at station 4 (Table 1). Those found at station 1 were probably carried from above the spring, where cursory inspection indicated their presence. The species Baetis vagans McDonnough, dominant mayfly nymph of the upstream stations, was less frequent and abundant downstream. Table 1. Percent Frequency Occurrence of Invertebrates Found in 0.1 m2 Bottom Samples Taken from September 1965 Through April 1966 Organism Percent Frequency Occurrence Upstream Stations Downstream Station Station 1 Station 4 Station 15 Gammarus . . . . . 100 100 50 Baetis . . 82 100 75 Tricorythodes . . . 9 0 50 Stenonema . . . 18 0 100 Hydropsyche . 90 90 100 Cheumato psyche . 0 64 75 Chimarra . 0 9 100 Neophylax . 36 0 0 Number of bottom samples . 11 22 4 158 Wisconsin Academy of Sciences, Arts and Letters [VoL 57 Of the Trichoptera larvae, the genera Cheumatopsyche and Chimarra were found more frequently downstream than upstream. Hydropsyche occurred with about equal frequency in both areas. Neophylax was restricted to station 1. Gammarus pseudolimnaeus Bousfield was always present in upstream areas, but less frequent downstream. There was no detectable difference among stations in the fre¬ quency occurrence of the other major groups of organisms in the bottom samples: Turbellaria (Dugesia) ; Annelida (Enchytraei- dae) ; Decapoda (Orconectes) ; Hydracarina; Hemiptera; Megalop- tera (Sialis) ; Coleoptra (Elmidae, Dytiscidae) ; Diptera (Chry- sops, Tipula, Antocha, Limnophora, Atherix) ; Simuliidae; Tendipedidae ; Gastropoda ( Physa, Limnaea, Ferissia); and Pele- cypoda (Sphaerium). Quantitative Analysis of Bottom Samples At station 1, Gammarus comprised 51% of the total weight of 11 combined samples, trichopteran larvae, 38%, Baetis, 2%. The remaining 9% was mostly chironomid larvae. At station 4, Gammarus comprised 22% and trichopteran larvae 63% of the combined weight of 21 samples. Baetis comprised 2%; the remaining 13% was largely chironomid larvae. At station 15, Gammarus comprised only 6%, trichopteran lar¬ vae 62%, mayfly nymphs 1% of the combined weight of 4 samples. The remaining 31% was mostly dipteran larvae of which chirono¬ mid larvae contributed a little more than half. In the upstream areas, the maximum standing crop for any one sample collected from 11 September 1965 to 4 April 1966 was 0.36 g/0.1 m2 at station 4 for Baetis nymphs on 2 April 1966. Two major emergences of Baetis were directly observed: one from late October through November and another from early April to early May. The maximum standing crop for trichopteran larvae, mostly Hydropsyche, was 7.4 g/0.1 m2 at station 4 on 15 January 1966. For Gammarus, the maximum standing crop was 2.5 g/0.1 m2 at station 4 on 14 November 1965 and 6 February 1966. Measurement of Drift Gammarus and Baetis nymphs were the major components of the drift. Other organisms included chironomid larvae and adults and occasionally other dipteran larvae. Fig. 1 shows the hourly drift rates of Gammarus and Baetis on 7 June 1966, indicating greater total drift at night for both species. Drift rates of Gam¬ marus were nearly identical at stations 2 and 4 (Fig. 1, Table 2), 1969] Peterka — Invertebrates in Small Wisconsin Stream 159 Figure 1. Drift rates of Gammarus and Baetis in g/hr at station 2 (solid line) and station 4 (broken line) on 7 June 1966. Stippled area indicates time of daylight. whereas the hourly drift rates of Baetis were generally larger at station 2 than at 4. The average standing crops, determined from two bottom samples taken at each station on 8 June 1966 were 0.10 (range 0.067 to 0.126 g) and 0.03 (range 0.040 to 0.019) g/0.1 m2 for Baetis at stations 2 and 4, respectively; and 1.48 (range 0.12 to 2.85) and 1.21 (range 0.658 to 1.754) g/0.1 m2 for Gammarus at stations 2 and 4, respectively. The higher drift rates of Baetis at station 2 may be the result of higher standing crop at that station, while the nearly uniform drift rates of Gammarus correspond to nearly identical standing crops at both stations. The total stream drift of Gammarus is less variable than that of Baetis (Table 2), ranging from 2.7 to 13.3 g per day, a factor 160 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Table 2. Total Stream Drift (g/day) of Gammarus and Baetis, for Three 24-hr Periods, 1966 Total Stream Drift in G/Day Date, 1966 Station 2 Station 4 Gammarus Baetis Gammarus Baetis J une 7 . 8.3 0.72 9.4 0.34 Aug. 31 -Sept. 1 . _ * _ * 13.3 5.00 Oct. 22-23 . _ * _ * 12.8 1.30 *No samples taken. of approximately 5; whereas the total stream drift of Baetis ranged from 0.34 to 5.00 g per day, a factor of approximately 15. Discussion Tricorythodes, Stenonema, Cheumatopsyche, and Chimarra were more frequent at downstream than at upstream stations, while Neophylax occurred only near the spring. The principal organisms found in the stream drift were Gammarus and Baetis . While Gam¬ marus is abundant in the drift during most of the year, Baetis occurs in significant quantities only during late winter — early spring and late summer— early fall, periods just preceding major emergences. The maximum standing crop of Baetis of 3.6 g/m2 reported in this study is low when compared with the maximum standing crop of 10.0 g/m2 reported in Valley Creek, Minn. (Waters, 1962 b). These low standing crops may limit the use of the drift method (Waters, 1962 b) for estimating production rates in Bear Branch. A diurnal periodicity of drift rates for both Gammarus and Baetis agrees with results reported by several workers and reviewed by Waters (1965). A clearer picture of the diurnal periodicity might have resulted in this study if shorter sampling intervals had been used and if possible disturbances by cattle in the stream had been eliminated. The complete blockage of the stream with drift nets at station 2 on 7 June 1966 did not influence the drift of Gammarus entering the nets which also completely blocked the stream at station 4. This suggests the drift is not accumulative from upstream to down¬ stream areas for distances of at least 152 m. 1969] Peterka — Invertebrates in Small Wisconsin Stream 161 Acknowledgements I am grateful to Robert F. Meyers for his help in collecting and processing the data and Bryon L. Stephens for his permission to conduct the study on his land. Literature Cited Dendy, J. S. 1944. The fate of animals in stream drift when carried into lakes. Ecol. Monogr. 14:333-357. Muller, Karl. 1963. Diurnal rhythm in “organic drift” of Gammarus pulex . Nature. 198:806-807. Needham, P. R. 1938. Trout streams. Comstock Pub. Co., Ithaca, New York. 233 pp. Waters, T. F. 1961. Standing crop and drift of stream bottom organisms. Ecology. 42:532-537. - . 1962 a. Diurnal periodicity in the drift of stream invertebrates. Ecol¬ ogy. 43:316-320. - . 1962 b. A method to estimate the production rate of a stream bottom invertebrate. Trans. Am. Fish. Soc. 91(3) : 243-250. - . 1965. Interpretation of invertebrate drift in streams. Ecology. 46(3) : 327-334. - ., and R. J. Knapp. 1961. An improved stream bottom fauna sampler. Trans. Am. Fish. Soc. 90(2) : 225-226. BIOLOGY OF THE COREIDAE IN WISCONSIN T. R. Yonke 1 and J. T. Medler* From July 1, 1962, through June 15, 1967, numerous observa¬ tions and collections were made of various members of the order Hemiptera. Special interest centered around species in the alydid- coreid-rhopalid complex. Much of the literature in these groups has been of a taxonomic nature with relatively little or no work reported on the biologies and held histories of the respective spe¬ cies. To fill this void in the knowledge of the Coreoidea a study was initiated on the biologies and held histories of members of the Alydidae, Rhopalidae and Coreidae that are found in Wisconsin. Previously information was reported on the biologies of 4 spe¬ cies of Alydidae in Wisconsin, including Megalotomus quinque - spinous (Say) (Yonke and Medler, 1965) ; Alydus conspersus Montandon, A. eurinus (Say) and A. pilosulus Herrich-Schaeffer (Yonke and Medler, 1968). Limited observations have been con¬ ducted on 2 Rhopalidae, Corizus crassicornis (Linnaeus) and Har- mostes reflexulus (Say) (Yonke and Medler, 1967). This is a report on the study of the Coreidae in Wisconsin. Ten species have been recorded for the state. Three species were ex¬ tremely rare — Anasa armigera (Say), Chariesterus antennator (Fabricius), and Leptoglossus oppositus (Say). Coriomeris humilis Uhler and Merocoris distinctus Dallas were uncommon. Five spe¬ cies frequently encountered were Acanthocephala terminalis (Dal¬ las), Anasa tristis (DeGeer), Archimerus alternatus (Say), Catorhintha mendica Stal, and Euthochtha galeator (Fabricius). The hrst 4 species mentioned were not collected by the authors, but were represented in the Department of Entomology Museum, University of Wisconsin. Of the latter 5 species the squash bug, A. tristis , has been thor¬ oughly studied. The squash bug is of economic importance on the cucurbits. The most comprehensive work to date was that of Beard (1940). Also, the biology of C. mendica has been adequately deter¬ mined (Balduf, 1942, 1957). For the other 8 species, however, instructor, Department of Entomology, University of Wisconsin. Presently Assistant Professor, Department of Entomology, University of Missouri, Columbia, Missouri. Information given in this paper was submitted in partial fulfillment of the require¬ ments for the degree Doctor of Philosophy in Entomology at the University of Wisconsin. a Professor, Department of Entomology, University of Wisconsin, Madison. 163 164 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 there is no published information on the biologies except for Blatchley (1926) who presented only a few brief notes. The prin¬ cipal part of this text reports studies conducted on A. terminalis, A. alternatus, and E. galeator. Methods and Materials Collection trips were made throughout southern Wisconsin from April until December in order to obtain field information. The principal collection sites were at Gibraltar Rock, Columbia County ; Devils Lake State Park, Ferry Bluff and Parfreys Glen in Sauk County; and Wyalusing State Park, Grant County. Surveys were made by net-sweeping of prairie, woodland, and disturbed “weedy” habitats to determine if the bugs were present. If encountered, a more intensive search of the area was made to determine the host plants and make observations. The term, host plant, is used here to denote the actual feeding of the bug on a particular plant in the field. A “collection record” signifies only that the specimen was collected from a particular plant which may or may not have been a host plant and for which no feeding was observed. A special attempt was made to collect parasitized specimens and rear out the adult parasites. All adults and nymphs observed in the field with Dipterous eggs on them were brought back to the laboratory, and the specimens placed in rearing cartons where they were held until the parasites emerged. Field collected eggs were also placed in laboratory rearing cartons in an attempt to obtain Hymenopterous parasites. All stages of coreids found in the field were brought alive to the laboratory in Madison and placed in 1 pint rearing containers. This technique was developed by Scheel, Beck, and Medler (1956). The substitute food used in the laboratory cartons consisted of fresh green beans (Fig. 1). The food was changed 2 or 3 times a week or as needed. Nymphs and adults of E. galeator and later instar nymphs and adults of A. alternatus and A. terminalis fed well on this diet. Rearings in the laboratory were used to complement field infor¬ mation and to some degree determine the biology of these bugs. The term “laboratory biology” is employed to designate that the bugs were reared under laboratory conditions and on a substitute non-natural food. The cultures were maintained at room tempera¬ ture (25 ± 4°C), and approximately normal daylight. Daily records were kept on preoviposition period, the number of eggs produced, incubation period, oviposition period, time spent in each instar, copulation frequency and duration, and adult longevity. Notes were made on any peculiar or interesting habits exhibited 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 165 Figure 1. Feeding aggregation of 7 second-instar nymphs of Euthochtlia galeator on the laboratory diet of fresh green bean. 166 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 by these insects. The old cartons and cotton rolls were replaced with fresh ones approximately every 3 weeks or as needed to re¬ duce contamination. Results and Discussion Acanthocephala terminalis (Dallas) The main held study area for this species was at Ferry Bluff (Sauk County) where it was collected from June through Septem¬ ber. Additional observations and collections were made at Wyalus- ing State Park, Devils Lake State Park, and Parfreys Glen. Figure 2 shows the seasonal occurrence of A. terminalis in south¬ ern Wisconsin during 1962-1967. There was 1 generation per year. Adults appeared June 13 and were found throughout the summer until September 24. Members of the overwintered genera¬ tion were collected and brought to the laboratory where gravid females oviposited bright whitish eggs. No eggs or hrst-instar nymphs could be found in the held; therefore, the data for these stages were obtained from eggs laid by females while transporting them to, or in, the laboratory. No eggs were oviposited by females collected from the held after July 15. Of 47 eggs obtained in this manner all but 2 hatched in from 7-14 days (x = 9.68). Data pre¬ sented for Instar 1 were determined by recording the dates of eclosion. Second-instar nymphs were collected in the held from June 30-August 11 ; third-instar nymphs from June 30-August 11 ; fourth-instar nymphs from July 8-August 11; and hfth-instar Adult Insfar 5 Instar 4 a> §* Instar 3 Instar 2 Acanthocephala terminalis (Dallas) overwinter t > m > t* 4 , i-4 M 'Instar 1 'Egg 30 July Sept. 30 Oct. Figure 2. Seasonal occurrence of Acanthocephala terminalis in Wisconsin, 1962-1967, showing inclusive dates and actual collection records (peaks). 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 167 nymphs from July 23-September 24. Eggs and first- and second- instar nymphs would have to occur in the field earlier, and third- and fourth-instar nymphs later than figure 2 shows. Also, it was expected that there was an overlap in late July and August of the occurrence of the overwintered adults and the new summer generation adults. Fifth-instar nymphs collected from the field and brought into the laboratory began molting into the adult stage as early as July 27. On August 10, 1966, 1 second-, 3 third-, 4 fourth-, and 4 fifth-instar nymphs were collected on Vitis riparia Michx. at Ferry Bluff. Three host plants were established for the nymphs and adults of A. terminals (Table 1). Four females were found at Wyalusing State Park feeding on the tender shoots of staghorn sumac, Rhus typhina L., on June 18, 1964; as was a second-instar nymph on July 12, 1965. Second- through fifth-instar nymphs and adults were collected over the entire season at Ferry Bluff from both V. riparia, river grape, and Physocarpus opulifolius (L.) Maxim., nine-bark. Fifth- instar nymphs and adults were also collected from V. riparia at Parfreys Glen on August 18, 1966. Both nymphs and adults were observed feeding on the succulent stems and petioles of these plants. One fourth-instar nymph was found feeding on the upper surface of a leaf of V. riparia. Adults were easily disturbed and were very rapid fliers. Imma- tures were occasionally found resting on the upper surfaces of leaves but more frequently they were hidden under leaves or along the main stems of the plants. This secretive habit along with their dark color made them somewhat difficult to find. At no time were they in great abundance. Immatures and adults were also collected from Fraxinus sp. (ash), Rubus (Eubatus) sp. (blackberry), Tilia americana L. (basswood), Desmodium acuminatum (Muhl.) Wood (tick tree- foil), TJlmus rubra Muhl. (slippery or red elm) (Table 1). No feeding was observed in any of these instances so that no definite host plant association could be made ; however, it was possible that any or all were fed on and could have served as hosts. No parasites were obtained from either nymphs or adults, and no parasite eggs were found on any of the bugs. Field collected specimens were brought to the laboratory and placed on green beans in the cartons. Although individuals from the first-instar nymph to the adult stage fed on this diet they did not do well. Nymphal mortality was very high from the second- instar through the fifth. Only 2 individuals went through their entire development from egg to adult on beans. One female spent 3, 9, 6, 8, and 17 days in each of the 5 respective stadia for a total of 43 days ; the other, a male, spent 4, 9, 7, 13, and 28 days in the Table 1. Host Plant Kecords of Acanthocephala terminalis, Archimerus alternatus •, and Euthochtha galeator 168 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 ><>< II I I I I I I X I I II I I II II X r i h-- n i 101 1 1 1 1 1 1 1 1 1 1 1 1 1 ix 1 1 1 1 11 1 1 1 1 1 1 IX 1 I lx* I I XX II lx I lx 1 lx Ixx I I I I I I I ir\ ltn Lr\ w~\ w-\ ltn a. 4 1 1 1 I I I I I I I I I I I IX I I I I I I I I I tu I I I I I I I I I I I I I I I I lx I I I 1 1 1 1 1 1 1 1 1 1 1 ri 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 a .2 S23 <5,3 CO E •Z v. so8 ^ S'S-2 M i I -O 50 a ^3 d£ ^ * . C fco 3-s‘a-e » §3-1 2 .3 ,9* a w S-q a ^ *0 cs S 2 c b § IS ? d a ■Cl <*> o a - s - • • «0 3 Q C • | o £ g g dj 5 ■S c »s § 1 (Continued) 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 169 w 0 3 HD < 1 1 x 1 1 1 1 x 1 1 1 1 III 0 ^0 0 -£ jC a £ 1 1 r 1 1 1 1 1 v Hi I] ■2 0 & jc " CUD bfl w 1 IX |X | | Hi |i 1 1 IX | 4_. »3 3 0 c 3 T> < 1 1 1 1 1 lx 1 2X22 i l ll jC 0 «0 W-4 £ >> 2 Mill 1 | 1 H INI 2 0 < bfi bB UJ IX | | | | | | | | | | | | || •2 ~o c 1 3 T3 < x 1 l|lx 1 1 1 1 1 lx lx ■22 c* .a 0 4h a £ ^ I I I I r< VMM * irs I ia c A 8 >> 2 1 1 1 1 1 1 co 4 1 1 1 1 1 1 1 OC) 0 -g c bfl 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 X W Ml I II | 1 HI 1 III II i !■ ,0 3 EE a clt ^ - o co VJ '•£ § E 3 I ^ sS|§ a a a aS.Q a-fyc O -3 ^ O- O r* O O g - txo K ^ Q Q 3 o c J- ■§* a ?*>~3 3 ^ ^ ~ ~o -o *^033 fc ~ Q,Q,0(0i^c<^a a 3 3 ' c 2 aw 3-2 ^ ~o 3 a 3 > 3 C cn CO X, a E >> c 170 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 5 stages for a total of 61 days. Out of 66 adults, both field collected and laboratory reared, 34 were males and 32 were females. The time spent in nymphal development was 58.2 days (Table 2) based on accumulated mean values. Values for Table 2 were compiled primarily from data on second- through fifth-instar nymphs col¬ lected from the field and brought to the laboratory. Growth ratios were fairly consistent (Yonke and Medler, In Press, a). The closeness of fit for the values indicated a uniform growth rate, and therefore tended to substantiate the data pre¬ sented in Table 2 on the time required for nymphal development. However in comparing laboratory data from Table 2 with field occurrence records in Figure 2, the time required for nymphal development would appear to be longer in the laboratory than in the field. Anasa tristis (DeGeer) The following observations in Wisconsin were consistent with the biology of this species as determined by Beard (1940) in Con¬ necticut. The squash bug was not especially abundant in the state. On July 23, 1963, 7 eggs wore collected from the upper surface of a leaf of a squash plant. Four hatched on July 27, and 1 on July 28. They molted to the second instar on July 30, but then died. Two fifth-instar nymphs collected from squash plants on September 17, 1963, molted to the adult on September 29. Four adults were also collected and placed on green bean in the laboratory. They were observed copulating frequently, but no eggs were laid. They lived for 42, 183, and 190 days, respectively. Eleven fourth- and fifth-instar nymphs and 4 adults were collected from squash on September 21, 1963. A Dipterous larva emerged from an adult squash bug and pupated on October 8. After 76 days an adult T. pennipes emerged. Another T. pennipes was Table 2. Duration of Nymphal Stadia of Acanthocephala terminalis in the Laboratory Instar . 1 2 3 4 5 Number of Observations. . 67 42 14 7 8 Range (days) . Mean =*= standard deviation . Accumulated mean days . 3-7 3.6±0.81 7-18 12.9=*=3. 13 16.5 6-18 12.8^3.74 29.3 8-13 9.9±2.2 7 39.2 14-28 19.0=*=5.37 58.2 1969] Yonke and Medler— Biology of Coreidae in Wisconsin 171 placed in the carton and the 2 were observed copulating twice; however, no eggs were deposited on the coreids. Archimerus alternatus (Say) The main study areas for this species were at Parfreys Glen, Wyalusing State Park, Ferry Bluff, Gibraltar Rock in Columbia County, and Devils Lake State Park. This was the most abundant species of the Coreidae found in Wisconsin. Figure 3 shows the seasonal life history of A. alternatus in Wis¬ consin during 1962-1967. There was 1 generation per year with overwintering in the adult stage. Adults were found continuously from June 2-October 8. This species was well represented in the museum collection of the Department of Entomology, University of Wisconsin. These records showed its occurrence from May 12- October 9. An overlap of overwintered adults and those of the new summer generation probably occurred in late July and August Eggs were found in the field on June 13, 15, 29, and July 1 and 13. First-instar nymphs were collected in the field from June 28- July 13; second-instar nymphs from June 18-August 11; third- instar nymphs from June 29-August 27 ; fourth-instar nymphs from July 8-September 1; and fifth-instar nymphs from July 15- September 26. Eggs and first-instar nymphs would have to occur in the field earlier and first-and fourth—instar nymphs later than is shown in Figure 3. Table 1 lists the plants on which eggs, nymphs, and adults of A. alternatus were collected in Wisconsin from 1964-1967. Adults Archimerus alternatus (Say) A . .a. . . 1 OVERWINTER Instar 5 Instar 4 q Instar 3 uj Instar 2 ▲ M Instar 1 Egg 10 20 30 10 20 31 10 20 31 10 20 30 10 20 June July Aug. Sept. Oct. Figure 3. Seasonal occurrence of Archimerus alternatus in Wisconsin, 1962- 1967, showing inclusive dates and actual collection records (peaks). 172 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 of this species tended to aggregate on certain species of plants in June. On June 13, numerous adults were found copulating on rag¬ weed, Ambrosia trifida L., and goldenrod, both Solidago gigantea Ait. and S. altissima. Adults were found feeding in the apical region of the stem of S. altissima. These plants grew in open fields or near the edges of woods. On June 14, 1966, in a stand of oak with a maple-basswood understory located 5 miles south of Platteville in Grant County, 4 pair of adults were found feeding on Aster sagittifolius Willd. (Figure 4) and 1 male on bedstraw, Galium concinnum T. & G. However, no eggs were found on any of the plants in the area. In an open area at Parfreys Glen on July 15, more than 40 adults were found on a group of Desmodium acuminatum (Muhl.) Wood plants. From 2 to 7 adults were found feeding a few inches below the apex of each of the developing stems. Their feeding caused wilting and death of the stem. Many pairs of adults were copulating and 7 individually laid eggs were found, of which 2 were on the upper surface and 5 on the under surface of leaves of D. acuminatum. They were collected and brought to the laboratory where they hatched— 2 on July 28 and 30, and July 1, and 1 on July 2. Adults continued to be found predominantly on D. acuminatum throughout the entire season. This same group of plants was examined on July 16, 1966, and over 30 adults, but no eggs, were found. An abundance of adults were also found on them on June 24 of that year. These observa¬ tions would indicate that A. alternatus migrated to D. acuminatum in June. Oviposition occurred primarily on these plants, but eggs could also be found scattered over plants growing nearby. One egg was found on the stem of Agrimonia gryposepala Wallr. growing right next to the clump of D. acuminatum. Females also frequently oviposited on the dorsal surfaces of the head, thorax or wings of other females or males in the vicinity. The eggs would be carried on these adults until they hatched. This would undoubtedly aid in dispersal of the insect. Adults were also found feeding on horse-mint, Monarda fistulosa L., and the sunflower, Helianthus decapetalus L., in June. In addi¬ tion adults were collected from, but not observed feeding on, Aster sp. and Robinia pseudoacacia L. in June. One egg was found on the underside of a leaf of may apple, Podophyllum peltatum L. Nymphs were predominant in the months of July and August. That the second through fifth nymphal stages were especially abundant on D. acuminatum (Table 1) was indicated by the re¬ spective numbers present: July 7, 1965, 13 second- and 1 third- instar nymphs; July 8, 1964, 6 second-, 29 third- and 9 fourth- instar nymphs; July 8, 1966, 44 second- and 3 third-instar nymphs; July 22, 1966, 17 second-, 21 third-, and 1 fourth-instar nymphs; 1969] Yonke and Medler— Biology of Coreidae in Wisconsin 173 Figure 4. Archimerus alternatus adults feeding on Aster sagittifolius on June 13, 1966, Grant County, Wisconsin. 174 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 August 3, 1965, 1 second-, 10 third-, and 16 fourth-instar nymphs; and on August 24, 1966, 8 fourth- and 58 fifth-instar nymphs. Two other legumes also served as host for second- through fifth-instar nymphs. They were readily found on hog-peanut, Amphicarpa bracteata, (L.) Fern, and fourth- and fifth-instar nymphs less fre¬ quently on D. dellenii Dari. On August 18, 1966, 5 fourth- and fifth-instar nymphs were found feeding on the latter species in the apical portion of the stem and on leaf petioles. Nymphs in all stages were generally found in a resting position, inclined anterior to posterior at about a 30° angle, whenever they were not feeding. First- through fourth-instar nymphs appeared to be obligopha- gous. In contrast, fifth-instar nymphs exhibited an interesting feeding habit in that the number of host plant species that they fed on increased, therefore, being similar to the polyphagous habits of the adult stage. It was not until the fifth-instar that nymphs appeared to move about much over the vegetation. No early-instar nymphs were found on any plants except the host plants as previously noted. However, fifth-instar nymphs were found resting and feeding on ragweed, both A. art emisii folia L. and A. trifida; on honewort, Cryptotaenia canadensis (L.) D. C. ; and on D. canadense (L.) D. C. On September 1, 1966, they were observed feeding on white snakeroot, Eupatorium rug o sum Houtt. Also, many fifth-instar nymphs continued to feed on D. acuminatum even after seeds were produced (Fig. 5) . The distribution of A. alternatus may eventually be shown to coincide with the distribution of D. acuminatum. This would be over the entire eastern United States and include the southern two-thirds of Wisconsin (Fassett, 1939). Adults collected in September were found feeding on the daisy, E. annus and on skunk cabbage, Symplo carpus foetidus (L.) Nutt. Adults were also found on Aster sp., A. trifida, and Solidago spp. The reactions of all of the plants were similar where feeding pressure was great, that is, where 2 or more bugs fed for a pro¬ longed period. Adults feeding in June (Figure 4) and second- through fifth-instar nymphs were all found to produce the same effect. The terminal portion of from 2 to 6 inches above the feed¬ ing loci gradually turned black and died. Where feeding pressure was severe early enough in the season no flower or seed production occurred. Adults exhibited a death feign whenever they were disturbed. They would drop from the plants and remain motionless in the litter for as long as 5 minutes. Two new parasite records were obtained from eggs of A. alter¬ natus. Two eggs of this species were collected at Wyalusing State 1969] Yonke and Medler— Biology of Coreidae in Wisconsin 175 Figure 5. Fifth-instar nymph of Archimerus alternate feeding- on the seed petiole of host, Desmodium acuminatum. 176 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Park on June 29, 1964. One was from the underside of a leaf of may apple. From it 7 Hymenopterous parasites, both males and females, of Ooencyrtus clisiocampae (Ashm.) (Encyrtidae) , emerged on July 12. These adults were placed in a rearing carton with 5 eggs of A. alternatus oviposited in the laboratory and an¬ other generation of parasites was obtained from these eggs. Egg parasitism will be discussed in greater detail under Euthochtha galeator. Another Hymenopterous parasite, a female Anastatus pearsalli Ashm., emerged from the other field collected egg of A. alternatus. The date of emergence was not known. On July 13, 1965, while collecting along a roadside 3 miles west of Sauk City 1 egg of A. alternatus was found on a blade of grass. It was brought to the laboratory and on September 8, 1965, a female A. pearsalli emerged. Six adult T. pennipes were reared from A. alternatus in Wiscon¬ sin. The only previous report of this parasite-host relationship was given by Patton (1958) who listed it from Florida. The following specimens were all obtained at Wyalusing State Park. One female collected on June 14, 1963, produced a Dipterous larva on June 22. It spent 11 days in the pupal stage and emerged into an adult on July 3. A male collected on June 18, 1964, pro¬ duced a larva on June 22, which spent 12 days in pupation before it emerged an adult. Another male collected August 5, 1964, pro¬ duced a larva on August 11. It spent 14 days in pupation. In addi¬ tion 3 T. pennipes adults were obtained from field collected fourth- or fifth-instar nymphs that were reared to the adult stage in the laboratory. The parasites emerged on August 17 and 24, and Sep¬ tember 9, and spent 13, 12, and 16 days, respectively, in pupation. Upon examination of the exuviae of these bugs no parasite eggs were found, indicating that they had been parasitized in the field at least 1 instar prior to when they were collected. They then molted and carried the larva with them until they emerged an adult. The presence of the parasite, therefore, does not necessarily result in the death of the nymph. This is consistent with the ob¬ servations of Beard (1940) on the behavior of the larva in nymphs of the squash bugs. Out of 27 second- and third-instar nymphs collected on June 29, 1964, at Wyalusing State Park, 2 second-instar nymphs were each found to have 1 Dipterous parasite egg on the dorsal surface of the abdomen. One molted to the third instar on July 1, and the other on July 2, leaving the parasite eggs on their cast skins. Ex¬ amination under the microscope showed that 1 was in the process of cutting its way out of the egg and the other was still intact 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 177 within the egg. These eggs fit the description of Worthley (1924) for T. pennipes and were probably of that species. Trichopoda pennipes adults mated in the rearing cartons and 1 female deposited 15 eggs on an adult female of A. alternatus. A Dipterous larva emerged after 15 days, pupated, and spent 15 days in the pupal stage before molting into an adult. Desmodium acuminatum plants were collected from the field on July 16, 1965, placed in pots, and brought into the laboratory to facilitate observation of the behavior of the first 3 instars of A. alternatus. Groups of 5 bugs which hatched on the same day and were all of the same instar were placed on each set of plants. A set con¬ sisted of 3 to 5 stems per pot. Three groups of 5 first-instar nymphs each were placed on the upper surfaces of the leaves of 3 sets of plants. All 15 bugs remained for 3 or 4 days on the leaves where they were placed. Three of the 15 were observed with their stylets inserted through the leaf surface, and most nymphs showed a dis¬ tended abdomen indicating that they fed in the first instar. Shortly after molting, the second-instar nymphs began to migrate over the plants. Eight were feeding on the petioles of the flowers or seeds within 1 day after molting. The other 7 nymphs escaped from the plants. Second- and third-instar nymphs after being removed from the rearing cartons and placed on the leaves immediately began moving over the plants, but again they were always found feeding on the stem or petioles within 2 days after release. On July 2, 1966, plants of D. acuminatum were collected from the field, potted, and brought into the laboratory. They were placed in 1 of 2 rearing cages. Again nymphs were observed. Five first- instar nymphs from the laboratory cartons and 2 second-instar nymphs collected the same day from the field were released on the bottom of the cage. The first instars remained there for 2 days and then moved onto the plants and took the “resting position” on the undersides of leaves. However, within 10 minutes 1 of the second-instar nymphs had moved onto the plant and began feeding on the main stem about 4 inches down from the tip of the 18-inch long stem. Two days later the other second-instar nymph assumed a position immediately next to the first. The 2 bugs remained together for 3 more days before moving. Behavior of the third- instar nymphs was similar to that of the second. Nymphs and adults would gather together and feed at about the same position on the stem even though other stems in the same set of plants were free of any bugs. This aggregation behavior was found in the field. It was also found in the laboratory on D. acuminatum in the rearing cages, and on green bean in the 178 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 rearing cartons (Figure 6). The bugs produced the same feeding stress on the plants grown in the laboratory as was found in the field, causing the apical portions of the stems to die above the feeding loci. Some observations were conducted on these bugs in the rearing cartons in the laboratory. Third- through fifth-instar nymphs and adults fed well on green beans; however, the development of the second-instar nymphs on this diet was impaired. Cultures were established each year from eggs obtained from females or nymphs collected in the field in June. Adults were ob¬ served copulating frequently in the rearing cartons. Eggs of this species were oviposited singly as were those of A. terminalis. Heidemann (1911) reported that the eggs of Ar chimeras calcara- tor (Fabricius) were laid in a row, but not joined. Although oc¬ casionally A. alternatus would lay a few eggs in an irregular row there was no definite oviposition pattern. He also reported 14 “chorial processes” for the egg. The number of micropylar proc¬ esses found for A. alternatus were considerably greater, as noted under the description of the egg of this species (Yonke and Med- ler, In press b). A mean of 2.1 eggs per day (n = 17, range = 1.0- 4.5) were oviposited by females in the laboratory, with a mean value of 57 eggs (n = 17, range = 10-128) for the total number of eggs oviposited per female. The maximum number of eggs laid in any 1 day by a single female was 15. The mean oviposition period was 37.2 days (n = 17, range = 5-120). Although it may not be applicable to a univoltine species overwintering in the adult stage, a mean preoviposition period of 33.5 days (n = 17, range = 22-58) was observed in the laboratory. The mean incubation period of 67 eggs was 13.4 days with a range of from 10-20 days. Out of 258 eggs oviposited in the labo¬ ratory 85.3% hatched. Eclosion took place by means of a pseudopercular cap. The bug forced it open by means of a series of pulsating movements. The cap split first at its most ventral point and then proceeded posterio- dorsally. This took only a few minutes. Nymphal development took 48.6 days (Table 3) based on an accumulation of the means. The values, except for those of Instar 2, employed in construction of Table 3 were obtained from observa¬ tions on nymphs feeding on green beans in the rearing cartons. Since this species did not develop from egg through to adult on beans the values for Instar 2 were obtained from observations of second-instar nymphs that developed on D, acuminatum in the laboratory rearing cages. The time spent in nymphal development agreed favorably with the data presented on seasonal history in Figure 3. 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 179 Figure 6. Feeding aggregation of 8 second-instar nymphs of Archimerus alternatus on the laboratory diet of fresh green beans. 180 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Table 3. Duration of Nymphal Stadia of Archimerus alternatus in the Laboratory Instar . 1 2 1 3 4 5 Number of Observations. . . 54 7 26 54 54 Range (days) . . 3-7 5-10 4-16 7-17 10-30 Mean =*= standard deviation . 4.6^0.92 7 . 6 =*= 1 . 89 8. 3 ±2 . 72 1 1 . 2 ± 1 . 86 1 6 . 9 =*= 5.11 Accumulated mean days . 12.2 20.5 31.7 48.6 1Values presented for Instar 2 were taken from observations of the bugs on Desmodium acuminatum in rearing cages. The growth ratios were very constant for this species. (Yonke and Medler, In press b). The closeness of fit for the values indi¬ cated a uniform growth rate and, therefore, tended to substantiate the data presented in Table 3. Out of 484 adults either collected in the field or reared in the laboratory 231 were males and 253 were females. Deformed antennae in the form of a reduction in the size and number of segments were observed on some bugs reared in the laboratory. It was thought that this might have been due to the non-host diet. However, examination of field collected adults also produced specimens with similar abnormalities. Catorhintha mendica Stal Adults, eggs, and nymphs were collected from Mirabilis nycta - ginea growing on railroad embankments in Dane, Jefferson, Rock, and Sauk counties. They were not found on any other plant. Early- instar nymphs were collected from under the involucral bracts which enclosed the flowers and seeds, and later-instar nymphs fed on the petioles. Additional observations of this bug and its host were made at Kankakee, Illinois. On September 3, 1966, many adults and 151 nymphs were collected of which there were — 1 first-, 8 second-, 56 third-, 78 fourth-, and 9 fifth-instar nymphs. Adults and nymphs were collected from the field and brought alive to the laboratory where they were placed on green beans. Adults fed readily and lived from 3-85 days, but the nymphs did not survive. Females laid from 2-7 eggs per batch (n = 20, x — 3.5). They hatched in from 5-8 days (n — 28, x = 6.4). These data are in agreement with the data presented by Balduf (1942) for this species. 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 181 Euthochtha galeator (Fabricius) The main field study area for this species was at Parfrey’s Glen. Additional observations and collections were made at Wyalusing State Park, as well as a few other locations. Figure 7 shows the seasonal occurrence of E . galeator in Wis¬ consin during 1962-1967. There was 1 generation a year with overwintering in the adult stage. Adults were found from May 25- October 8. No adults were found between July 6 and August 11 indicating that the overwintered adults died off at about that time. This species was well represented in the museum collection of the Department of Entomology, University of Wisconsin. These rec¬ ords showed its occurrence from May 5-October 27. There were no records between July 13 and August 6. Field collected nymphs brought back to and reared in the laboratory began molting into the adult stage on July 25. Eggs were found in the field from June 13-July 20. First-instar nymphs were collected in the field from June 13-July 13; second- instar nymphs from June 13-August 3; third-instar nymphs from July 13-August 10; fourth-instar nymphs from July 18-August 10; and fifth-instar nymphs from July 24-September 19. Eggs and first-instar nymphs would have to occur in the field earlier, and third- and fourth-instar nymphs later than Figure 7 shows. On June 13, 1963, while collecting at Wyalusing State Park, 13 eggs and first-instar nymphs of E. galeator were found on the upper surface of a leaf of Aster ericoides L. growing in an open field. The bright red nymphs were still aggregated about the eggs Euthochtha galeator (Fabricius) Adult Instar 5 Instar 4 . ^ 0 Instar 3 Cn Instar 2 Instar 1 4 OVERWINTER Egg 21 31 10 20 30 10 20 31 10 20 31 10 20 30 10 20 May June July Aug Sept Oct. Figure 7. Seasonal occurrence of Euthochtha galeator in Wisconsin from 1962-1967 (peaks). 182 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 when they were collected. On the same date a cluster of 18 more eggs were found on Urtica dioica L., nettle. They were brought back to the laboratory and placed in a rearing carton. Eight first- instar nymphs hatched on June 29 and 1 on June 30. Also on June 28, 12 encyrtid parasites, 0. anasae (Ashm.), emerged from 4 of these eggs. This constituted a new host record for 0. anasae, there being no previous published association of this parasite and host. These adults were placed in a carton with eggs of E. galeator ob¬ tained from females in the laboratory. The adult parasites died by July 20, and were removed from the carton. On July 80, the eggs were examined under the microscope and found to have many small oval masses about the inner surfaces of the chorions. They also were dark in color, in contrast to the reddish or gold color of the normal eggs. More than 70 adult parasites emerged from the eggs on August 1 ; 17 more parasites on August 2 ; 4 on August 4 ; and 1 on August 5. Some parasites escaped from the carton so an exact count could not be made. Both males and females of 0. anasae were obtained. An unsuccessful attempt was made to obtain an¬ other laboratory generation of parasites. Six eggs of E. galeator were found on June 16, 1964, on a blade of grass growing on the sandy bank along the Wisconsin river at the Mazomanie Wildlife Refuge, Dane County. They were posi¬ tioned in a line, end to end. Five hatched on June 22 and 1 on June 23. On July 10, 1965, 6 eggs were found oviposited on the upper surface of a leaf of wild plum, Prunus americana Marsh. Closer observations revealed slightly raised brown spots on the leaf where 7 more eggs had been laid but were no longer attached. One adult Hymenopterous parasite, Anastatus pearsalli, emerged from 1 egg of E. galeator on August 11, and 5 more emerged, each from a single egg, on August 12. Anastatus pearsalli failed to parasitize laboratory reared eggs and they died by August 14. Two additional batches of eggs were found. In 1 there were 11 eggs that had already hatched, and in the other there were 14 eggs. Ten eggs of the latter batch hatched in the laboratory on July 16, and 4 hatched on July 17. All 3 batches of eggs were found in a disturbed weedy habitat at Parfrey’s Glen. In the same area on July 13, 1965, 8 first-instar nymphs were observed in aggregation resting on the upper surface of a leaf of Rhus glabra L. and 9 eggs that had hatched were found on the underside of the leaf. On July 12, 1964, 7 encyrtid parasites, 0. clisiocampae, obtained from an egg of A. alternatus collected in the field on June 29, 1964, were placed in a rearing carton with 20 laboratory eggs of E. galeator. The parasites were observed on the eggs and on July 27, numerous parasites emerged from all 20 E. galeator eggs. A total 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 183 of 124 parasites were obtained from these and 5 additional labora¬ tory eggs of A. alternatus which were also present in the carton. Six host plants were found for nymphs and adults of E. galeator (Table 1). They were determined by observations of the bug feed¬ ing on the respective plants in the field. The host plants were Agrimonia gryposepala Wallr. or roots fibrous, Achillea mille¬ folium L., Aster pilosus Willd., M. fistulosa or horsemint, D. acu¬ minatum, and Quercus ellipsoidalis E. J. Hill. Of these plants, adults were found feeding and copulating most frequently on A. gryposepala. Since adults had been found frequently at Parfrey’s Glen on A. gryposepala in June of 1966, special trips were made in June of 1967 to see if the bugs could be located on these plants again. On June 4, 4 pairs of bugs were found on 4 separate clones of A. gryposepala, 5 were feeding and 2 pair were copulating. On June 15 another pair was found copulating on this plant. While a male E. galeator was being observed feeding on a petiole of A. millefolium, a female flew to the plant and approached the male, but flew away when disturbed. Adults were often found in June resting on the leaves of composites. Both males and females were swept with a net from Solidago altissima Mill, on June 13, 1966, at the Mt. Hope Conservation Area in Grant County. Second-instar nymphs were found feeding on Q. ellipsoidalis and D. acuminatum and fifth-instar nymphs on A. pilosus and A. gryposepala. Nymphs and adults were collected also from A. bracteata, Aureolaria grandiflora var pulchra (Benth.) Pennell, Cary a sp., and Ulmus rubra Muhl. Therefore, it would appear that E. galeator is not monophagous, but rather can and does utilize a number of host plants representing different families. Adults exhibited a death feign. Whenever disturbed they would either drop from the plant and remain motionless, or fly away quickly. Nine adult Dipterous parasites, T. pennipes, were reared in the laboratory from 4 male and 5 female field collected adults. There were no previous published records of T. pennipes from E. galea¬ tor. Four parasites were obtained from adults collected at Parfreys Glen on June 24, 1966. Larvae emerged from the bugs and pupated on June 27, July 2, 10, and 20, 1966; and spent 18, 15, 14, and 20 days respectively, in their pupal stages. Adult parasites did not live longer than 7 days and although they were observed copu¬ lating they did not oviposite on the bugs in the rearing cartons. One female E. galeator that was parasitized laid 3 batches of 19, 10, and 12 viable eggs. A female and a male collected on July 1, 1966, each produced a parasite on July 6 and 12, respectively. 184 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 The 1 obtained from the female emerged an adult on July 26, but that from the male died in the pupal stage. On August 4, 1964, a fifth-instar nymph of E. galeator was col¬ lected at Parfreys Glen and brought to the laboratory where it molted into an adult male on August 25. On September 3, a Dip¬ terous larva, T. pennipes, emerged from it and pupated. No para¬ site egg was found on the exuvium of the fifth instar indicating that it had been parasitized in an earlier instar and had success¬ fully molted with the larva inside. A female was collected from Mazomanie Wildlife Refuge on August 23, 1963. A larvae emerged from it and pupated on August 24 and emerged to an adult on September 9. One male E. galeator was collected from Kankakee, Illinois, on August 6, 1965, brought back alive to Madison, and placed in a rearing carton. On August 9, a parasite emerged, pupated, and on August 27 emerged as an adult. From 1 to 5 parasite eggs were found randomly oviposited over the bodies of the bugs. Some observations were conducted on the bugs in the rearing cartons to gain information on their biology. Both nymphs and adults fed readily on the laboratory diet of fresh green beans. Cultures were established each year from eggs obtained from females, eggs, or nymphs collected from the field in June. This species oviposited its eggs in batches similar to those of A. armi- gera and A. tristis, with a mean number of 16.0 eggs per batch (number of batches = 62, range = 2-32). In 22 observations indi¬ vidual females oviposited from 3-19 batches of eggs with a mean of 8.2 batches per female. The greatest number of eggs oviposited by a single female in the laboratory was 259. A mean time of 6.0 days (n = 60, range = 1-24) elapsed be¬ tween oviposition of individual batches of eggs. Generally when only 1 or a few days passed between oviposition fewer eggs were oviposited per batch. In 14 observations a mean time of 40.7 days were spent in the oviposition period, with a range of from 11-89 days. A delayed preoviposition period ranging from 26-143 days (n = 5, x = 95.6) was observed for females reared in the labora¬ tory from the nymphal stage. These females would have normally overwintered and not copulated or oviposited until the following spring. This delayed preoviposition time was a reflection of that behavior. In Tact, a preoviposition period, if defined as the time elapsing between the molting of the insect to the adult stage until first oviposition, would not be a pertinent part of the biology of a univoltine species that overwinters as an adult. 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 185 The period of incubation for 566 eggs oviposited in the labora¬ tory ranged from 7-17 days with a mean time of 12,9 days. Out of a total of 720 eggs obtained in the rearing cartons 636 hatched, or 88.3%. Eclosion took place by means of the pseudopercular cap. The bug forced it open by means of a series of pulsating movements. The cap split first at its most ventral point and then proceeded posterio-dorsally. This took only a few minutes. First-instar nymphs were observed in the rearing cartons with their stylets inserted in green beans. It was assumed that they were feeding; however, they would successfully develop in the absence of any food during that stadium as long as moisture through the wick was available. In fact these nymphs were lethargic and frequently aggregated on or near the eggs until they molted into the second instar. Second- through fifth-instar nymphs spent much of the time feeding. Nymphal development took 54 days, with the second, third, and fourth stadia being about equal (Table 4). This was based on an accumulation of the means. These laboratory data agreed fairly well with the field occurrence records given in Figure 7. The growth ratios were fairly consistent (Yonke and Medler, In press c). The closeness of fit for the values indicated a uniform growth rate, and therefore tended to substantiate the data pre¬ sented in Table 4. Frequently nymphs aggregated while feeding. Figure 1 shows an aggregation pattern of 7 second-instar nymphs feeding in a rela¬ tively small area on 1 of the 2 green beans present in the rearing carton. One additional bug was present in the carton. Out of 206 adults either collected in the field or reared in the laboratory 102 were males and 104 were females. Adults generally Table 4. Duration of Nymphal Stadia of Euthochtha galeator in the Laboratory Instar . 1 2 3 4 5 Number of Observations . 136 80 51 54 42 Range (days) .... Mean =*= standard deviation . Accumulated mean days . 2-5 3 . 4±0. 83 6-18 11.1=^3.13 14.5 6-18 10. 2 ±3 .12 24.7 5-18 12 . 2 ±3 . 43 36.9 8-28 17. 1 ±5.49 54.0 186 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 survived for a long time. Two adults collected from the field in June lived 204 and 212 days under laboratory conditions. Merocoris distinctus Dallas This species occurs in Wisconsin, although nowhere common. Adults occurred from June 9-October 26. Also, 1 adult was re¬ corded for “April.” One adult was collected while sweeping Midway Prairie, La Crosse County, on June 15, 1967. Another adult was observed on June 17, 1967, at Crex Meadows, Burnett County, associated with a carrion spot on the pavement of a road. One fifth-instar nymph was collected September 1, 1964, while sweeping a dis¬ turbed weedy area at Parfreys Glen ; however, continued sweeping and investigation of individual plants did not produce more. It was held on green bean in the laboratory, molted into an adult on October 5, and died on October 17. There is apparently 1 genera¬ tion per year in Wisconsin with overwintering in the adult stage. One adult each was collected on September 80, 1963, and October 2, 1965, on goldenrod, from Curtis Prairie, University of Wisconsin, Arboretum, Madison. Summary From July 1, 1962, through June 15, 1967, numerous observa¬ tions and collections were made of coreid species found in Wiscon¬ sin. Ten species were recorded. Anasa armigera (Say), Charies - terns antennator (Fabricius), and Leptoglossus oppositus (Say) were extremely rare. Coriomeris humilis Uhler and Merocoris dis¬ tinctus Dallas were uncommon. Five species frequently encoun¬ tered were Acanthocephala terminalis (Dallas), Anasa tristis (De- Geer), Archimerus alternatus (Say), Catorhintha mendica Stal, and Euthochtha galeator (Fabricius). The field histories and laboratory biologies were determined for Acanthocephala terminalis, Archimerus alternatus and Euthochtha galeator. All 3 species went through 1 generation a year in Wis¬ consin and overwintered in the adult stage. Adults of Acanthocephala terminalis were collected in the field from June 13 to September 4. No parasites were found for this species. Three host plants determined for nymphs and adult were Physocarpus opulifolia (L.) Maxim., Rhus typhina L., and Vitis riparia Michx. Eggs had an incubation time of 7 to 14 days (mean = 9.7). The mean time spent in nymphal development was 58.1 days. Adults of Archimerus alternatus were collected in the field from June 2 to October 8. They were found feeding on Solidago altis - 1969] Yonke and Medler — Biology of Coreidae in Wisconsin 187 sima Mitt., Aster sagittif olius Willd., and Galium concinnum T. & G. in early June. They moved from these plants to Desmodium aciminatuo (Muhl.) Wood, tick trefoil, on which oviposition gen¬ erally occurred. First-through fourth-instar nymphs were restricted to D. acuminatum, D, dilienii Dark, and Amphicarpa bracteata (L.) Fern., hog peanut. Fifth-instar nymphs fed on these 3 plants and also fed on a number of other plants, including Ambrosia artemisiifolia L., Ambrosia trifida L., Cryptotaenia canadensis (L.) D.C., Desmodium canadense (L.) D.C., and Eupatorium rugosum Houtt. Host plants of the summer generation adults in¬ cluded those recorded for the immatures, and in addition Erigeron annus (L.) Pers., dairy; and Symplocarpus foetidus (L.) Nutt., skunk cabbage. Feeding on all plants occurred on the stems and petioles, causing the terminals above to wilt. Two hymenopterous parasites were obtained from field-collected eggs of A. alternatus. They were Ocencrytus clisiocampae (Ashm.) (Encyrtidae) and Anastatus pearsalli Ashm. (Eupelmidae) . These were new records. Adults of the dipterous parasite, Trichopoda pennipes Fabricius (Tachinidae) , were reared from field-collected adults. Feeding aggregations of Archimerus alternatus were observed both in the field and in the laboratory for nymphs and adults. A mean number of 2.1 eggs per day was oviposited by females in the laboratory. Incubation took 13.4 days. The mean time spent in nymphal development was 48.4 days. Euthochtha galeator adults were collected in the field from May 25 to October 8. The 6 host plants found for nymphs and adults were Agrimonia gryposepala Wallr., Achillea millefolium L., Aster pilosus Willd., Monarda fistulosa L., D. acuminatum (Muhl.) Wood, and Quercus ellipsoidalis Hill. Three parasite species were obtained from field-collected eggs and adults of E. galeator. They included 2 Hymenoptera, A. pear¬ salli Ashm. and Ooencyrtus anasae (Ashm)., and 1 Diptera, T. pennipes Fabricius. These were new records. In addition, C. clisio¬ campae parasitized E. galeator eggs in the laboratory. A feeding aggregation was also observed for nymphs of E. galeator. This species oviposited its eggs in batches (mean = 16.0 eggs per batch). A mean time of 6.0 days elapsed between oviposi¬ tion of individual batches of eggs. Incubation of eggs took 7 to 18 days (mean = 12.9). The mean time spent in nymphal develop¬ ment was 54 days. References Cited Balduf, W. V. 1942. Bionomics of Catorhintha mendica Stal. (Coreidae, Hemiptera). Bull. Brooklyn Entomol. Soc. 37:158-166. 188 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 - . 1957. The spread of Catorhintha mendica Stal. (Coreidae, Hemiptera). Proc. Entomol. Soc. Wash. 59:176-185. Beard, R. L. 1940. The biology of Anasa tristis, DeGeer with particular refer¬ ence to the tachinid parasite, Trichopoda pennipes Fabr. Conn (New Haven) Agr. Exp. Sta. Bull. 440:594-679. Blatchley, W. S. 1926. Heteroptera or true bugs of Eastern North America. The Nature Publishing Co., Indianapolis. 1116 p. Fassett, N. C. 1939. The leguminous plants of Wisconsin. Univ. Wis. Press, Madison. 157 p. Heidemann, 0. 1911. Some remarks on the eggs of North American species of Hemiptera-Heteroptera. Proc. Entomol. Soc. Wash. 13:128-140. Patton, C. N. 1958. A catalogue of the larvaevoridae of Florida. Fla. Entomol. 41:29-39, 77-89. Scheel, C. A., S. D. Beck, and J. T. Medler. 1956. Feeding and nutrition of certain Hemiptera. Proc. Tenth Int. Congr. Entomol. 1956: 303-308. Worthley, H. N. 1924. The biology of Trichopoda pennipes Fab. (Diptera, Tachinidae), a parasite of the common squash bug. Psyche 31:7-16, 57-77. Yonke, T. R., and J. T. Medler. 1965. Biology of Megalotomus quinques - pinosus (Say). Ann. Entomol. Soc. Amer. 58:222-224. - . 1967. Observations on some Rhopalidae (Hemiptera). Proc. N. Central States Branch Entomol. Soc. Amer. 22:74-75. - . 1968. The biologies of 3 species of Alydus in Wisconsin (Hemiptera: Alydidae). Ann. Entomol. Soc. Amer. 61:526-531. - . In Press a. Description of the immature stages of Coreidae: 1. Euthochtha galeator. Ann. Entomol. Soc. Amer - . In Press b. Description of the immature stages of Coreidae: 2. Acanthocephala terminalis. Ann. Entomol. Soc. Amer - . In Press c. Description of the immature stages of Coreidae: 3. Archimerus alternatus. Ann. Entomol. Soc. Amer. HOST RECORDS AND PHENOLOGY OF LOUSE-FLIES ON WISCONSIN BIRDS Nancy S. Mueller, Helmut C . Mueller, and Daniel D. Berger Introduction Louse-flies (Diptera: Hippoboscidae) are ectoparasites which feed on the blood of birds and mammals. Most species infest birds and are agile, elusive creatures that spend most of their lives well hidden and protected by the feathers of their hosts. Reproduction depends on at least one mating and involves the production of one young at a time. Embryonic and larval development of the off¬ spring occurs within the body of the female fly. Shortly before pupation the larva is dropped from the fly and falls from the bird to the ground. Some species overwinter as pupae, while others emerge in one to several months. Each new adult must find a suit¬ able host within a few days after emergence, and mating depends on the presence of both sexes on the same host. These interesting flies are relatively rare in collections because few entomologists have access to large numbers of living, wild birds and most ornithol¬ ogists who handle wild birds have neither the time nor the interest to devote to the ectoparasites. A list of records of louse-flies for Wisconsin and surrounding states as well as some information about the habits of the flies is found in MacArthur (1948). Bequaert’s (1952-1956) monograph on the Hippoboscidae of North America provides an exhaustive compilation and analysis of all aspects of the life histories, dis¬ tribution and taxonomy of Hippoboscidae in North America. Our present knowledge of the life history, distribution and host rela¬ tions of the louse-flies which live on passerine birds has benefited from the recent increase in the use of mist-nets and Helgoland traps for capturing wild birds. Bennett (1961) collected louse-flies from passerine hosts in Algonquin Park, Ontario, between mid- May and mid-September in 1957-1960 and kept some of the flies on captive birds to study their longevity and host preferences. Workers in Britain (Corbet, 1956b, 1961; Hill, 1962, 1963) and in Scandinavia (Hill et al., 1964) studied the distribution, life his¬ tories and host preferences of three species of Ornithomyia dur¬ ing the summer, when these louse-flies are most abundant. Much 189 190 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 less is known about the louse-fly infestation of migrating and win¬ tering birds or about the louse-flies of raptorial birds at all seasons of the year. In the present study we have analyzed records of louse-flies col¬ lected in Wisconsin from a wide variety of raptorial and passerine birds. Most of the specimens were taken during spring and autumn migration, but we also collected flies on hawks and owls in winter. This report is based on data from a total of 1,281 individuals of eight species of Hippoboscidae taken on 695 individuals of 60 species of birds. Techniques Raptorial and passerine birds have been trapped regularly dur¬ ing spring and autumn migration at the Cedar Grove Ornitholog¬ ical Station, located on the western shore of Lake Michigan about 64 km north of Milwaukee, Wisconsin. A description of the station area is found in Mueller and Berger (1966). Birds were checked for Hippoboscidae during a total of 11 autumns and 6 spring sea¬ sons, beginning in 1955; during this period 4,898 raptorial birds and 58,979 passerine birds and woodpeckers were handled. A concerted effort was made to check each captured raptorial bird for the presence of Hippoboscidae. Hawks were trapped individually in nets and usually removed from the netting immedi¬ ately after capture. The usual method of checking for louse-flies on hawks and owls was to blow on the feathers or to spread them by hand. The disturbed flies darted among or on the feathers, seeking shelter in another region, or flew to the person handling the bird or, less frequently, to a window of the banding labora¬ tory. A few flies were certainly overlooked or otherwise escaped capture. The flies were captured manually and immediately doused with alcohol or water to immobilize them momentarily until they could be secured in small vials. Passerines and a few smaller raptorial birds, such as the Sharp-shinned Hawk and small owls, were taken in mist-nets set in dense brush. The mist-nets were checked at approximately 40 minute intervals, and the birds removed from the netting as quickly as possible. Passerine birds were transported to the labora¬ tory in opaque containers, and then segregated according to species in sorting cages made of wire screen of 0.2 mm mesh. Some trans¬ fer of flies from one bird to another may have occurred as the birds were being transported to the laboratory; however, we be¬ lieve that the darkness in the containers drastically reduced the activity of the birds and probably also the movements of the flies. Most flies that remained on their hosts until they reached the laboratory became a part of our collection. Undoubtedly more flies 1969] N. Mueller, H. Mueller and Berger — Flies on Birds 191 would have been collected from passerine hosts if the birds had been checked for ectoparasites immediately on capture and prior to their removal from the nets, but the time and facilities available did not permit a more thorough collection of flies from passerine birds. Hawks and owls were also obtained from the State Experimental and Game Farm at Poynette, Wisconsin, during the period of au¬ tumn through early spring in the years of 1958 through 1963 and on several occasions in the autumns of 1956 and 1964. A total of 475 hawks and 137 owls was handled. These birds had been taken in steel traps set on poles, and they were held for one to several days in aviaries before examination, with separate enclosures for owls and hawks. Most of the birds were Great Horned Owls (135) and Red-tailed Hawks (316), although 7 other raptorial species were represented. The birds were transported in individual burlap bags to Madison, Wisconsin, where they were checked for ectopara¬ sites, measured, banded and released. In addition, hawks and owls were captured with the Bal-chatri trap (Berger and Mueller, 1959) in the central and southern part of Wisconsin, principally during the spring migrations. Species from which louse-flies were collected were the Red-tailed Hawk, Broad-winged Hawk, Sparrow Hawk, Great Horned Owl, Barred Owl and Long-eared Owl. They were either checked immediately for flies, banded, etc., or they were first transported to a suitable banding laboratory. A number of persons contributed flies taken from birds caught in a number of localities, and we have no accu¬ rate record of the number of hawks and owls handled. Louse-flies which were collected prior to 1958 were pinned and dried. Since dried debris and badly shrunken abdomens made sexing difficult, subsequent collections were preserved in 70 per cent ethanol, with each vial containing all flies taken from a single host. Except for the few individuals which had been mutilated at the time of capture, those louse-flies preserved in alcohol re¬ mained in excellent condition for determination of species, sex, and occurrence of phoresy and mites. MacArthur’s (1948) key to the Hippoboscidae of the Eastern United States was used for preliminary identification of the flies. The more exhaustive key of Bequaert (1954, 1955, 1956) was used to reexamine all individuals which appeared in any way unusual. Specimens of Ornithomyia were checked against the key of Hill et al. (1964). Woodman’s (1954) key was used for the identification of Mallophaga. Host Records Eight species of Hippoboscidae were collected, but only three of these were at all common: Lynchia americana (896 specimens), 192 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Ornithomyia fringillina (267), Ornithoica vicina (102), Ornithoc- tona erythrocephala (7), Lynchia nigra (5), Lynchia angustifrons (1), Lynchia albipennis (1), and Microlynchia pusilla (1). M. pu- silla is a new record for Wisconsin and neighboring states (cf. MacArthur, 1948). Louse-flies were found on 11 species of Falconi- formes, 6 species of Strigiformes, 2 species of Piciformes and 41 species of Passeriformes. Many of these are new host records for Wisconsin and neighboring states (cf. MacArthur, 1948), indicated by a superscript “a” in Table 1. Host records new to North America (cf. Bequaert, 1956; Bennett, 1961) are indicated by superscript “b”. The number of birds of each species on which flies were found and the total number of birds handled provide a crude minimum estimate of infestation (Table 1). These data are incomplete for the following reasons: (1). A plus ( + ) mark notation in the “No. Handled” column of Table 1 indicates that additional members of this species were trapped by the Bal-chatri and the exact number is not known. (2). Passerine birds were not examined as thoroughly as were raptorial birds. (3). No data are presented for species of birds from which no louse-flies were taken. A com¬ plete list of the passerine birds taken in autumn at Cedar Grove can be found in Mueller and Berger (1968). The sex could be determined for 1,107 of the 1,281 specimens; the condition of the rest of the specimens, most of which were collected prior to 1958, precluded determination of sex. Taking the sample as a whole, 88 per cent of the flies were females. This un¬ balanced ratio in our sample is undoubtedly a result of our collect¬ ing in late summer and autumn; since females live longer than males, they dominate samples taken after the period of emergence of the adults (cf. Bennett, 1961; Hill, 1963). Ornithoica vicina is an extremely small louse-fly that is easily overlooked. We first observed this species in August 1959 in the ears of a Great Homed Owl. A total of 120 Great Horned Owls has subsequently been checked for the presence of louse-flies, and 23 individuals yielded a total of 71 female and 2 male 0. vicina, all from the ears. Seven specimens of this fly were found in the ears of a Barred Owl, and one specimen each was found in the ear of a Screech Owl and a Saw-whet Owl. A variety of other raptorial birds and passerines harbored this fly on the body plumage, but not in the ears (Table 1) . We found no other species of louse-fly in the ears of any bird. Although 0. vicina was reluctant to move from the ear of an owl, this fly was observed to escape from passerines caught in mist-nets, alighting momentarily on the person handling the bird before flying away. Because of their small size, these flies were harder to find on body plumage than in the ears. Despite this 1969] N. Mueller, H. Mueller and Berger — Flies on Birds 193 Table 1. Host Records for Hippoboscidae Collected in Wisconsin, 1955-65 Host0 No. Handled No. In¬ fested Sexes/Flies Total No. Flies* 9 9 & c? A. Ornithoica vicina Red-tailed Hawk . 940 f 1 1 0 1 Sparrow Hawka . 1 70 f 1 1 0 1 Screech Owl a, b . 10 2 2 0 2 Great Horned Owla . 1 38 f 26 71 2 74 Barred Owl a . 10f 1 5 2 7 Long-eared Owl a . 1 7 1 1 1 0 1 Saw- whet Owla . 234 2 1 1 2 Eastern Kingbird a,b . 19 1 1 0 1 Catbird a . 2,650 1 1 0 1 Veery a . 835 1 1 0 1 Swainson’s Thrush a . 9,845 1 0 1 1 Ruby-crowned Kinglet a . 1,721 1 1 0 1 Rusty Blackbird a,b . 2 1 1 0 1 Song Sparrow a . 723 1 1 0 1 Unknown Host . 4 7 0 7 TOTALS — Ornithoica vicina . 45 95 6 102 B. Ornithomyia fringillina Sharp-shinned Hawka . 1,860 10 8 0 12 Red-tailed Hawka,b . 940 f 2 2 0 2 Marsh Hawk a . 308 1 1 0 1 Pigeon Hawk . 241 1 1 0 1 Great Horned Owla . 138f 1 1 0 1 Saw-whet Owl a . 234 2 1 0 2 Yellow-shafted Flicker a . 197 5 5 0 5 Yellow-bellied Sapsuckera . 268 1 1 0 1 Traill’s Flycatcher a . 1,363 1 1 0 1 Blue Jay a . 91 1 0 1 1 Black-capped Chickadee a . 953 3 3 0 3 Red-breasted Nuthatch a,b . 83 1 1 0 1 Catbird a . 2,650 15 10 2 15 Brown Thrasher a . 206 5 5 0 6 Robin . 400 3 2 0 3 Wood Thrush a . 103 1 1 0 1 Hermit Thrush . 2,475 24 24 0 25 Swainson’s Thrush a . 9,845 52 43 4 55 Gray-Cheeked Thrush a . 2,109 9 9 0 9 Ruby-crowned Kinglet a . 1,721 1 1 0 1 Cedar Waxwinga . 613 3 2 0 3 Solitary Vireoa . 140 1 1 0 1 Red-eyed Vireoa . 2,600 5 5 0 5 Philadelphia Vireoa . 624 2 0 1 2 Myrtle Warbler a . 1,015 1 1 0 1 Bay-breasted Warbler a . 77 1 1 0 1 Blackpoll Warbler a . 399 1 1 0 1 Ovenbirda . 1 ,069 3 3 0 3 Northern Waterthrusha . 1,111 3 2 0 3 Mourning Warbler a . 203 2 2 0 2 Redstart a . 1,930 3 3 0 3 Scarlet Tanagera . 87 2 1 1 2 Cardinal . 121 1 1 0 1 Rose-breasted Grosbeak a . 505 2 2 0 2 194 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Table 1. Host Records for IIippoboscidae Collected in Wisconsin, 195 5-6 5 — ( C on tinued ) Host0 No. Handled No. In¬ fested Sexes /Flies Total No. Flies* $ $ cfd1 Evening Grosbeak a . 320 1 1 0 1 Purple Finch a . 422 1 1 0 1 Pine Siskin a,b . 143 1 1 0 1 Rufous-sided Towheea . 125 1 1 0 1 Slate-colored Junco . 1,940 3 3 0 3 Tree Sparrow a . 693 1 1 0 1 White-crowned Sparrow a . 152 1 0 0 1 White-throated Sparrow . 3,615 20 19 1 22 Fox Sparrow a . 1,068 2 2 0 2 Fincoln’s Sparrow . 233 1 1 0 1 Swamp Sparrow . 889 2 2 0 2 Song Sparrow . 723 8 7 0 8 Unknown Passerine . 47 38 3 47 TOTAFS — Ornithomyia fringillina. . . 258 222 13 267 C. Orrxithoctona erythrocephala Sharp-shinned Hawka . 1,860 1 1 0 1 Broad-winged Hawk . 1 4 f 4 1 3 4 Pigeon Hawk a . 241 1 0 0 1 Unknown Hawk . 1 1 0 1 TOTAFS— Orrxithoctona erythroce¬ phala . 7 3 3 7 D. Lynchia americana Goshawk a,b . 136 17 26 4 35 Sharp-shinned Hawka . 1,860 28 20 6 32 Cooper’s Hawk . 229f 9 6 2 10 Red-tailed Hawk . 940 f 159 216 56 355 Red-shouldered Hawka . 40 f 4 2 3 5 Broad-winged Hawka . Swainson’s Hawk a, b . 1 4 1 14 16 0 22 3 1 0 0 1 Golden Eagle a . 2 1 1 0 1 Marsh Hawk a . 308 2 1 0 2 Sparrow Hawk a, b . 1 70 f 1 0 0 1 Barn Owla . 1 1 1 0 1 Screech Owl a . 10 1 1 0 1 Great Horned Owl . 1 38 f 99 315 25 361 Barred Owl . 10f 6 4 1 6 Fong-eared Owl . 1 7 f 1 0 0 1 Evening Grosbeak a, b . 320 1 0 1 1 Unknown Hawk . 47 45 7 61 TOTAFS — Lynchia americana . 392 654 105 896 E. Lynchia angustifrons Broad-winged Hawk . 28 f 1 0 1 1 1969] N. Mueller, H . Mueller and Berger — Flies on Birds 195 Table 1. Host Records for Hippoboscidae Collected in Wisconsin, 1955-65 — ( Continued ) Host0 No. Handled No. In¬ fested Sexes /Flies Total No. Flies* 9 $ cf c? F. Lynchia nigra Sharp-shinned Hawk . 1,860 2 1 1 2 Broad-winged Hawk a, b . 14t 1 0 0 1 Sparrow Hawka . 1 70 f 1 0 1 1 Unknown Hawk . 1 0 1 1 TOTALS — Lynchia nigra . 5 1 3 5 G. Lynchia albipennis Unknown Hawk . 1 0 1 1 H. Microlynchia pusilla a Catbird a,b . 2,650 1 0 1 1 TOTALS — 60 species . 695 x 975 134 1,281 aNew record for Wisconsin and neighboring states (cf. MacArthur, 1948). bNew record for North America (cf. Bequaert, 1952-56; Bennett, 1961). *Total includes flies not sexed because the specimen was mutilated or dessicated. ■(Represents a minimum number of birds of this species handled; we have no record of the number of birds trapped by the Bal-chatri. xSixteen birds harbored more than one species of fly. °Scientific names of birds are given in the appendix. bias in collection probabilities, we feel strongly that owls are im¬ portant hosts for this species. This was suggested, but with reser¬ vations, by Bequaert (1953) . 0. vicina is apparently not very selective as to host; it is quite adaptable to life on a number of species, both passerine and rap¬ torial. On the Great Horned Owl and the Barred Owl infestations may be heavy, and the same bird is likely to harbor Lynchia americana on the body plumage. We found 14 cases of Great Horned Owls infested with both species of fly, and one individual caught at Wild Rose, Wisconsin, in August carried 14 0. vicina in the ears and 7 L. americana on other parts of the body. We col¬ lected no more than one individual 0. vicina from any passerine bird, and Bennett (1961) found that O. vicina was much less com¬ mon on passerine birds in Algonquin Park than Ornithomyia fringillina. Ornithomyia fringillina is resident on a variety of birds; it prefers passerines, but shows little host specificity (cf. Bequaert, 1954) . We have records from 6 species of raptorial birds, 2 wood¬ peckers, and 38 species of passerine birds (Table 1). This louse-fly 196 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 is not common on raptorial birds, even on those such as the Sharp- shinned Hawk which feed principally on passerines; only 10 of 1,860 Sharp-shinned Hawks caught at Cedar Grove were infested with this species. Our collection indicates no pronounced prefer¬ ences among passerine hosts, and infestations of all species were light (cf. Table 1). Our data from passerine birds are in no way comparable with those of Bennett (1961) for Algonquin Park. The relative infrequency of even the most common louse-flies on passerine birds at Cedar Grove (Table 1) can be attributed both to the collecting techniques and to the fact that we handled birds after the peak of infestation (cf. Bequaert, 1954: 137-138; Ben¬ nett, 1961). Bennett (1961) felt that certain passerines were favored over others, particularly those Fringillidae, blackbirds, and thrushes that inhabit the environment near the ground. In Britain and Scandinavia, where there are three species of Ornithomyia, there may be stronger host-preferences than those of 0. fringillina in the New World (cf. Hill, 1962b). According to Hill (1962b) and Hill et al. (1964) the “important hosts” for 0. fringillina in the Old World are exclusively small passerines, mostly hedgerow species; on larger passerines and on raptorial birds 0. fringillina appears to have been supplanted by the larger fly, 0. avicularia, whereas 0. chloropus prefers birds which frequent moorlands. We found Ornithoctona erythrocephala only on hawks. This very large louse-fly was found on the Sharp-shinned Hawk, the Broad-winged Hawk, and the Sparrow Hawk (Table 1). One Sharp-shinned Hawk had both 0. erythrocephala (1 individual) and L. americana (2 individuals), and a Broad-winged Hawk had one 0. erythrocephala and one L. americana. These are our only records of infestation of an individual host by more than one species of fly, excepting those Great Horned Owls that carried 0. vicina and L. americana. Lynchia americana was found on 15 species of raptorial birds (Table 1) . The only specimen from a passerine was on an Evening Grosbeak on 10 November 1963, and this individual (a male) could have been a stray which took up temporary residence on the grosbeak. These flies were generally reluctant to vacate their raptorial hosts even when the bird’s feathers were disturbed ; most specimens were collected as they darted among the feathers, although some flew to the person handling the bird. Infestations were often heavy, e.g. 17 Great Horned Owls, 8 Red-tailed Hawks and 3 Goshawks had more than 6 L. americana: the greatest individual infestation was 23 L. americana on a Great Horned Owl. L. americana prefers large raptorial birds with large-feathered, loose plumage which provides both a good hiding place and an 1969] N. Mueller, H. Mueller and Berger — Flies on Birds 197 environment which does not deter movement of flies as does a densely feathered, compact plumage. Only one L. americana was found on a falcon, on a Sparrow Hawk on 7 September 1956 ; none were found on 241 Pigeon Hawks and 81 Peregrine Falcons, both of which have relatively compact plumage. The Great Horned Owl is clearly favored; nearly 60 per cent of the Great Horned Owls captured at Poynette and Cedar Grove carried on the average four of these flies. Other species on which this fly is common are the Red-tailed Hawk and the Goshawk and probably the Swainson’s Hawk and the Barred Owl. Small hawks and owls are much less likely to harbor L. americana: for example, only 1.5 per cent of 1,860 Sharp-shinned Hawks and one of 234 Saw-whet Owls were infested. Until recently Lynchia fusca (Macquart 1845) was considered a distinct species; MacArthur (1948) called it “The Owl Fly” (also cf. Bequaert, 1953:264). However, after reexamination of the specimens, Bequaert (1955) was convinced that L. fusca is conspecific with L. americana, and Maa (1963:35) supports this synonomy. Using MacArthur’s (1948) key we found that these Lynchia were in fact difficult to distinguish and that the one host on which louse-flies consistently showed the characters of L. fusca was not an owl but the Broad-winged Hawk (specimens of “L. fusca” were found on 9 of the 12 Broad-winged Hawks from which L. americana were taken in the spring) . Other flies with characters of “fusca” were on a Sharp-shinned Hawk (in April), a Red-tailed Hawk (in August) and three Great Horned Owls (September, November and December). MacArthur (1948) reported no “L. fusca” from Wisconsin and only one record for a neighboring state (on a squirrel in Michigan!). We see no reason why L. fusca should not be considered conspecific with L. americana. We have collected 5 Lynchia nigra in Wisconsin; two from Sharp-shinned Hawks (1 May and 9 May 1964), one from an un¬ known hawk on 29 April 1965, one from a Broad-winged Hawk on 4 May 1958 and one from a Sparrow Hawk on 4 April 1958. We also found a L. nigra on a Sparrow Hawk in Zapta County, Texas, on 7 January 1956. According to Maa (1963:115) this species is essentially Neotropical; the Wisconsin specimens were most likely transported some distance on their hosts during spring migration. Two other species of Lynchia were taken from hawks in Wiscon¬ sin : a male of L. albipennis on a hawk on 12 May 1965 and a male of L. angustifrons on a Broad- winged Hawk on 5 May 1961. Since L. albipennis apparently prefers Ciconiiformes (cf. MacArthur, 1948; Maa, 1963), it is unusual to find one on a hawk. Spring and 198 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 summer records of L. angustifrons from the United States and southern Ontario are probably strays (cf. Maa, 1963). A male Microlynchia pusilla was found on a Catbird at Cedar Grove on 4 May 1965. This is the only record of M. pusilla from Wisconsin (cf. MacArthur, 1948) and the northernmost record for North America (cf. Bequaert, 1955; Maa, 1963). Also present in our collection is an individual of this species from a Harris Hawk (Parabuteo unicinctus) in Kennedy County, Texas, on 22 January 1956. This species is fairly common in southwestern United States (Bequaert, 1955), but its presence on a hawk is un¬ usual (cf. Bequaert, 1955; Maa, 1963). Phenology We found only one species of louse-fly, Lynchia americana, to be present in the adult stage all year round in Wisconsin. One species, Ornithoica vicina, was found in every season except spring; Ornithomyia fringillina was present only in late spring, summer and fall. Four species ( Ornithoctona erythrocephala, Lynchia albipennis, Lynchia angustifrons, and Lynchia nigra) were found only in spring and presumably were carried into Wisconsin on migratory hosts that winter in southern areas. The occurrence of Microlynchia pusilla can be considered accidental. A detailed ac¬ count of the seasonal occurrence of the eight species of louse-flies follows, based on our collections and supplmented with information from Bequaert (1952-1956) and Bennett (1961). Ornithoica vicina was absent from birds trapped in Wisconsin during late winter and spring; our earliest record was from an Eastern Kingbird on 14 June 1958, six weeks earlier than Bennett (1961) found it in Algonquin Park. This species was present, but not common, on passerine birds during fall migration. The only heavily infested individuals were owls in August and throughout fall and early winter. The latest date on which a male was taken was 23 October, but females were taken as late as 24 January in the ears of Great Horned Owls. Since these flies were not found in late winter and throughout the spring, our data cannot support Bequaert’s (1953:265) hypothesis that owls represent a temporary reservoir of overwintering flies that are held in reserve to infect passerine migrants in the spring. In Algonquin Park, Bennett (1961) reported the peak of abundance in late August and none later than early October. Bennett (1961 :401) found that at a constant temperature of 75° F the time required for metamorphosis in O. vicina varied consider¬ ably around a mean of 78 days and that developmental time at 75° was prolonged by interspersing an extended period of chilling at 1969] N. Mueller, H. Mueller and Berger — Flies on Birds 199 45° F. Apparently females of this species live for at least 4 months, but males may not live longer than one month (Bennett, 1961:396). Our data and those of Bennett (1961) suggest that the popu¬ lation of 0. vicina gradually builds up during the summer by the emergence and reproduction of flies that overwinter as pupae, not by overwintering adults or by the introduction of any signifi¬ cant number of flies from the south on spring migrants. The persistence of adults into late January argues for the emergence of flies during the fall, presumably from pupae deposited during the summer. The last males die in late fall and no females survive beyond mid-winter. It is possible that there is an alternation of diapause and non-diapause generations in northern North America such that adults emerging from diapause pupae during June and July give rise to a non-diapause generation that emerges in August or September and reproduces until December or January, giving rise to a second generation that overwinters in a pupal diapause. Alternatively, the overwintering pupae may not have a true dia¬ pause, which requires chilling, but they may simply remain dor¬ mant through the cold season and resume development with the return of warm weather. Ornithomyia fringillina is rare in Wisconsin in spring; from nearly 9,500 passerine birds trapped between 1 April and 10 June we collected only three specimens of this fly, two in late May and one in early June. These are more than a month earlier than previ¬ ous records for northern North America (cf. Bequaert, 1954; Bennett, 1961). We caught more passerine birds during September than during any other month, and it is not surprising that most of our collections of 0. fringillina are also from September. We found no males after late September and no females after early November. In Algonquin Park, Bennett (1961) collected no 0. fringillina until mid- July; he reported the peak of abundance in the two-week period around August 1. Assuming an early Au¬ gust peak for 0. fringillina in Wisconsin, then the decline in infestation of passerines was already well underway when we began netting birds in late August at Cedar Grove. The absence of this species in late fall and through the winter and its extreme rarity in spring and early summer argues against the overwintering of adults, even on migrant hosts in their winter home, and against the introduction of any significant number of adults on spring migrants. According to Bequaert (1954) O. fring¬ illina is confined to cool temperate areas, occurring in northeastern United States from 3 July to 6 November. Bennett (1961) and Hill (1963) present evidence that this species has an obligatory diapause with a period of chilling necessary for development to 200 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 resume. We suspect that as metamorphosis occurs through June and July the numbers gradually build up until mid-August and then slowly decline as adult mortality proceeds without further emergence of adults in fall. In this way the emergence of flies appears to coincide with the reproduction of the hosts (cf. Hill, 1963). In Algonquin Park, Bennett (1961) found O. fringillina on 10 per cent of adult passerine birds and 17.5 per cent of immatures; he presents experimental evidence that these flies, when confined with birds in small cages, select immature birds over adults, and he suggests that adults may be more efficient at catching and eating the flies. In nature these flies may not actively select immature birds, but they may be more frequently found on immature birds because these hosts are both more acces¬ sible and more abundant than adults at the time the flies emerge. Ornithoctona erythrocephala was occasionally found on hawks in spring. Bequaert (1954) considers this genus essentially tropical with little evidence that the adults or pupae can withstand cool temperate winters in North America. He suggests (p. 200) “that it is introduced there afresh every spring on some migratory breeding host”. The incidence of infestation of raptorial birds by Lynchia americana shows a peak during the fall, with fairly large num¬ bers of females and a few males present through the winter and spring, particularly on the Red-tailed Hawk and the Great Horned Owl (Table 2). Although we have very few summer records be¬ cause of very little trapping, males appear to be as common as females in late July and during the first half of August, with an increasing preponderance of females in the fall and winter. The persistence of both sexes suggests that breeding may occur year- round, but with the number of adults gradually decreasing during the winter. Some adults are undoubtedly brought up from the southern states on migrant hosts in spring, but the population is not substantially augmented until eclosion of new adults in summer. Bennett (1961:396) found that adult females of L. americana survived in captivity for 4-5 months, whereas most males lived only 15-20 days. The length of the pupal stage is temperature dependent and interrupted without ill effect by prolonged chilling at 45° F (p. 401). If reproduction does occur in Wisconsin in winter, it would seem that the pupae must be resistant to tempera¬ tures far below 45°, even below freezing, for prolonged periods. It would seem that development of this species does not depend on a period of chilling but that cool temperatures are not deleteri¬ ous to development. Table 2. Incidence of Lynchia americana on Hawks and Owls Taken at Cedar Grove and 1969] N. Mueller, H. Mueller and Berger — Flies on Birds 201 E-i z >* o Pk o Z 1/1 > cd 3 5»E < " 1 ~ 1 ~ u 1 Winter X OJ Uh o X! s % Infested fO 1 1 1*0 1 O I 1- I — > O Vr\ d) 8 Q No. Birds oooo^'-'o—^oa'oo t\ Vrs o' Z 8 00r- o 5 CO 3 - % Infested N O' ' — 'OO’ — ' c^N X — ■ •— 1 >— 1 O C<~\ &D 3 < No. Birds -<00000'00-<^00NOt\ c-) co X ^ O^Ow-\ — —' — ' 00 — < — 202 Wisconsin Academy of Sciences , Arts and Letters [Vol. 57 The persistence of adults in winter and spring suggests that pupae deposited during summer and early fall emerge before cold weather sets in ; those deposited in late fall and winter remain dormant through the winter, emerging after a suitable period of warmer weather in spring and early summer. During late winter and spring the population decreases due to gradual mortality of adults, and a lower level of infestation is maintained until meta¬ morphosis of pupae is completed or until sufficient numbers of infested migrants have returned. The coincidence of numbers of “L. fusca” on Broad-winged Hawks in spring might indicate the ‘‘importation’’ by hawks of a population of L. a mericana that lives year-round on birds of a more southern distribution. According to Bequaert (1955), L. americana may not occur north of 48° 10' north latitude, which is approximately the Canadian border in the central and western United States. We suspect that the range of this fly extends as far north as that of its hosts, which in the case of the Goshawk and Red-tailed Hawk is well north of the U.S.-Canadian border. There are few records of this fly from Mexico and farther south, and in western and southern states it is less common than L. nigra (Bequaert, 1955). The birds on which we found L. americana were, in fact, those which winter north of the Mexican border, and the only records we have for L. nigra are from the spring and from hawks that winter farther south. Lynchia albipennis , L. angustifrons and L. nigra are tropical species (cf. Bequaert, 1955) that are occasionally introduced on spring migrants and apparently do not overwinter in a northern climate. Microlynchia pusilla is essentially tropical and rare north of the southwestern United States (Bequaert, 1955). Our specimen, taken from a Catbird in May, undoubtedly “migrated” north from Central America with its host. Phoresy and Infestation by Mites The attachment of chewing bird lice (Mallophaga) to Hippo- boscidae is frequently cited as an example of phoresy, the wingless louse using the winged louse-fly as a means of transportation (references in MacArthur, 1948:385-387; Bequaert, 1952:163- 174; Corbet, 1956a). This phenomenon is difficult to explain be¬ cause each species of Mallophaga is believed to feed exclusively on feathers and epidermis of hosts of a given species. Ornithomyia fringillina, which is the most common carrier of the lice, is found on a variety of host species, and flies which have been experi¬ mentally marked for individual identification have been followed to individuals of several species of birds (Corbet, 1956b; Bennett, 1969] N. Mueller , H. Mueller and Berger — Flies on Birds 203 1961). The lice would presumably not benefit from being trans¬ ported to a host of a different species, and the louse-fly would not seem to be a good host for the lice to parasitize. Bequaert (1952:163) suggests that phoresy of Mallophaga by Hippoboscidae is “on the borderline of true parasitism”. The incidence of phoresy, at least by 0. fringillina, is too high to be accidental. Bequaert (1952) reported phoresy by 6 per cent of some 500 O. fringillina and less frequently by a number of other species of Hippoboscidae, including Ornithoica vicina, Orni- thoctona erythrocephala, Lynchia americana, and Lynchia albi- pennis. MacArthur (1948) found 14 cases, 4 per cent of the total flies examined, and Bennett (1961) reported phoresy by 22.8 per cent of 0. fringillina collected in 1957, with the highest frequency in July and much less in late August and early September. We found 30 cases (11.3 per cent) of phoresy by O. fringillina and one by our only specimen of Microlynchia pusilla (a female Briielia on the fly taken from a Catbird on 4 May, 1965). Avian hosts for flies that carried lice were a Sharp-shinned Hawk, a Yellow-shafted Flicker and 11 species of Passeriformes. In view of the diversity of bird species, it is perhaps surprising that all lice were in the genus Briielia and appeared to be of the same species; adult female lice predominated. All 30 cases of phoresy by O. fringillina in our collection occurred in summer and early autumn (2 in July, 1 in August, 23 in September, and 4 in Oc¬ tober). The lice were firmly imbedded in the integument of the louse-flies by their mouth parts, and the lateral surfaces of the abdomen were preferred places of attachment; in only one case was a louse attached to the prothorax. All were facing in the same direction as the flies. The infestation of Hippoboscidae by epidermoptid mites (Aca- rina) is considered true parasitism (cf. MacArthur, 1948:387; Bequaert, 1952:142-160). We found mite clusters on 52 O. fringil¬ lina (26.6 per cent) and on three L. americana; the latter is a species not previously recorded as parasitized by mites (cf. Bequaert, 1952). The mites were of the genera Microlichus and Myialges. Hosts for the L. americana which carried mites were two Broad-winged Hawks trapped in April and one unidentified hawk in October. The mite-infested 0. fringillina were on 18 species of passerine birds and one Pigeon Hawk, all trapped dur¬ ing autumn migration (2 in August, 31 in September, 18 in October, and one sometime in fall). Mites preferred the underside of the wings in the vicinity of the large veins, and they were rarely found elsewhere on the flies, such as on the abdomen or the upper surface of the wings. Mites usually consisted of a cluster formed by a female and her eggs or young. Clusters were often present on 204 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 the underside of both wings, and only the immature mites tended to stray to other parts of the fly’s body. Summary In the years 1955 through 1965 more than 5,000 hawks and owls and nearly 54,000 passerine birds were trapped alive in Wisconsin, and louse-flies were collected. Most birds were caught during au¬ tumn and spring migrations. A total of 1,281 individuals of eight species of Hippoboscidae was taken from 695 individuals of 60 species of birds. Only three species of louse-fly were common: Ornithoica vicina, Ornihomyia fringillina, and Lynchia americana. O. vicina was most frequently found in the ears of the Great Horned Owl. O. fringillina was found on a wide variety of passer¬ ines and on 6 raptorial species, with no obvious host preferences. Lynchia americana clearly preferred hawks and owls, especially those with large-feathered, loose plumage. Hawks returning from the south occasionally harbored other species of Hippoboscidae: Lynchia nigra, L. albipennis, L. angustifrons and Ornithoctona erythrocephala. One specimen of Microlynchia pusilla was found on a Catbird. The winter climate is probably the most important factor affect¬ ing the occurrence of a given species of fly in Wisconsin. Those species which cannot tolerate cold during the pupal stage (e.g. O. erythrocephala, L. albipennis, L. angustifrons, L. nigra , and M. pusilla) occur in Wisconsin only as vagrants and have essen¬ tially no chance of becoming permanently established. The pupae of O. vicina and L. americana can tolerate cold with a temporary suspension in development, and O. fringillina has a true diapause that depends on a period of chilling in order for development to resume (cf. Bennett, 1961). Adults of L. americana may over¬ winter on resident hosts, even perhaps continuing to reproduce, but there is no supplementation of their numbers until spring mi¬ gration brings in adults that have emerged in the south or until local temperatures become suitable for emergence of new flies. Adults of species infesting migratory birds travel south on their hosts in autumn and die with or without reproducing in the south. Lice of a species of Briielia (Mallophaga) were found attached to 11.3 per cent of the Ornithomyia fringillina. Phoresy was also found in one specimen of Microlynchia pusilla. Parasitic mites of the genera Microlichus and Myialges were found on the wings of three Lynchia americana and 26.6 per cent of the O. fringillina. Acknowledgments Most of the data for this study were obtained while the authors were engaged in a study of bird migration in Wisconsin. The 1969] N. Mueller, H. Mueller and Berger — Flies on Birds 205 following persons helped with the collection of louse-flies: F. and F. Hamerstrom, E. Bishop, C. Sindelar, D. Seal, J. Oar, J. Weaver, E. Schluter, and P. Drake. The Wisconsin State Ex¬ perimental and Game Farm at Poynette gave us raptorial birds trapped on their premises Mites were identified by Professor G. W. Wharton, Ohio State University. Many of the collections were made during the course of a study of bird migration sup¬ ported by a grant (GB-175) to Professor J. T. Emlen from the National Science Foundation. References Cited Bennett, G. F. 1931. On three species of Hippoboscidae (Diptera) on birds in Ontario. Can. J. Zool., 39:379-406. Bequaert, J. C. 1952-53. The Hippoboscidae or louse-flies (Diptera) of mam¬ mals and birds. Part I. Structure, physiology and natural history. Entomol. Amer., 32-33 (N.S.) : 1-442. Bequaert, J. C. 1954-56. The Hippoboscidae or louse-flies (Diptera) of mam¬ mals and birds. Part II. Taxonomy, evolution and revision of American genera and species. Entomol. Amer., 34-36 (N.S.) : 1-611. Corbet, G. B. 1956a. The phoresy of Mallophaga on a population of Omitho- myia fringillina Curtis (Dipt., Hippoboscidae). Ent. Mon. Mag. 92:207- 211. Corbet, G. B. 1956b. The life-history and host-relations of a Hippoboscid fly O mi thorny ia fringillina Curtis. J. Anim. Ecol., 25:403-420. Corbet, G. B. 1961. A comparison of the life-histories of two species of Orni- thomyia (Diptera, Hippoboscidae). Entomol. Gaz., 12:24-31. Hill, D. S. 1962a. Revision of the British species of Ornithomyia Latreille (Diptera: Hippoboscidae). Proc. Royal Ent. Soc. London, (B), 31:11-18. Hill, D. S. 1962b. A study of the distribution and host preferences of three species of Ornithomyia (Diptera: Hippoboscidae) in the British Isles. Proc. Royal Ent. Soc. London, (A), 37:37-48. Hill, D. S. 1963. The life history of the British species of Ornithomyia (Diptera: Hippoboscidae). Trans. Royal Ent. Soc. London, 115:391-407. Hill, D. S., W. Hackman, and L. Lyneborg. 1964. The genus Ornithomyia (Diptera: Hippoboscidae) in Fennoscandia, Denmark and Iceland. Notulae Entomologicae, 44:33-52. Maa, T. C. 1963. Genera and species of Hippoboscidae (Diptera) : types, synonymy, habits and natural groupings. Pacific Insects Monog., 6:1-186. MacArthur, K. 1948. The louse-flies of Wisconsin and adjacent states (Dip¬ tera: Hippoboscidae). Bull. Public Museum City Milwaukee, 8:367-440. Mueller, H. C., and D. D. Berger. 1966. Analyses of weight and fat varia¬ tions in transient Swainson’s Thrushes. Bird-Banding, 37:83-112. Mueller, H. C., and D. D. Berger. 1968. The relative abundance of species caught in mist-nets during fall migration at Cedar Grove. Passenger Pigeon, 29:107-115. Woodman, W. J. 1954. Mallophaga of Wisconsin, (unpub.) PhD Thesis, Univ. of Wisconsin, Madison. 281 pp. 206 Wisconsin Academy of Sciences, Arts and Letters [Vol. 57 Appendix (Scientific names of birds* infested with Hippoboscidae) Goshawk Accipiter gentilis Sharp-shinned Hawk Accipiter striatus Cooper’s Hawk Accipiter cooperii Red-tailed Hawk Buteo jamaicensis Red-shouldered Hawk Buteo lineatus Broad-winged Hawk Buteo platypterus Swainson’s Hawk Buteo swainsoni Golden Eagle Aquila chrysaetos Marsh Hawk Circus cyaneus Sparrow Hawk Falco sparverius Pigeon Hawk Falco columbarius Barn Owl Tyto alba Screech Owl Otus asio Great Horned Owl Bubo virginianus Barred Owl Strix varia Long-eared Owl Asio otus Saw-whet Owl Aegolius acadicus Yellow-shafted Flicker Colaptes auratus Yellow-bellied Sapsucker Sphyrapicus varius Eastern Kingbird Tyrannus try annus Traill’s Flycatcher Empidonax traillii Blue Jay Cyanocitta cristata Black-capped Chickadee Parus atricapillus Red-breasted Nuthatch Sitta canadensis Catbird Dumetella carolinensis Brown Thrasher Toxostoma rufum Robin Turdus migratorius Wood Thrush Hylocichla mustelina Veery Hylocichla fuscescens Hermit Thrush Hylocichla guttata Swainson’s Thrush Hylocichla ustulata Gray-cheeked Thrush Hylocichla minima Ruby-crowned Kinglet Regulus calendula Cedar Waxwing Bomby cilia cedrorum Solitary Vireo Vireo solitarius Red-eyed Vireo Vireo olivaceus Philadelphia Vireo Vireo philadelphicus Myrtle Warbler Dendroica coronata Bay-breasted Warbler Dendroica castanea Blackpoll Warbler Dendroica striata Ovenbird Seiurus aurocapillus Northern Waterthrush Seiurus novaboracensis Mourning Warbler Oporornis Philadelphia Redstart Setophaga ruticilla Rusty Blackbird Euphagus carolinus Scarlet Tanager Piranga olivacea Cardinal Richmondena cardinalis Rose-breasted Grosbeak Pheuticus ludovicianus Evening Grosbeak Hesperiphona vespertina * American Ornithologists’ Union. 1957. Checklist of North American Birds. Ameri¬ can Ornithologists’ Union, Baltimore, Md. 1969] N. Mueller, H . Mueller and Berger — Flies on Birds 207 Purple Finch Pine Siskin Rufous-sided Towhee Slate-colored Junco Tree Sparrow White-crowned Sparrow White-throated Sparrow Fox Sparrow Lincoln’s Sparrow Swamp Sparrow Song Sparrow Carpodacus purpureus Spinus pinus Pipilo erythrophthalmus Junco hyemalis Spizella arborea Zonotrichia leucophrys Zonotrichia albicollis Passerella iliaca Melospiza lincolnii Melospiza georgiana Melospiza melodia Nancy S. Mueller is a native of Ohio. She received her B.A. from the College of Wooster and her M.S. and Ph.D. (Zoology) from the University of Wisconsin. She presently teaches embryology at North Carolina State University at Raleigh. Her husband, Helmut, is a native of Milwaukee and holds B.S., M.S. and Ph.D. degrees from the University of Wisconsin. He currently teaches animal behavior and vertebrate zoology at the University of North Caro lina at Chapel Hill. Daniel D. Berger is also a native of Milwaukee, where he operates a small business and yet manages to spend about half of his time on his avocation, ornithology. He is a veteran of several ornithological expeditions to the arctic. All three have collaborated for some years in studies of bird migration at the Cedar Grove Ornithological Station in Sheboygan County. WISCONSIN ACADEMY OF SCIENCES, ARTS & LETTERS Madison, Wisconsin OFFICERS 1968-69 President Adolph A. Suppan School of Fine Arts The University of Wisconsin — ■ Milwaukee Vice-President (Sciences) John A. Cummings Department of Biology Wisconsin State University — Whitewater Vice-President (Arts) Mary Ellen Pagel 1111 North Astor Street Milwaukee Vice-President (Letters) Charles D. Goff Department of Political Science Wisconsin State University — Oshkosh President-Elect William B. Sarles Department of Bacteriology The University of Wisconsin — Madison Secretary Eunice R. Bonow Department of Pharmacy The University of Wisconsin — Milwaukee Treasurer Jack R. Arndt University Extension The University of Wisconsin — Madison Librarian Jack A. Clarke Department of Library Science The University of Wisconsin — Madison APPOINTED OFFICIALS Editor — Transactions Walter F. Peterson Department of History Lawrence University, Appleton Editor — Wisconsin Academy Review Ruth L. Hine Wisconsin Conservation Division Madison Chairman — Junior Academy of Science Jack R. Arndt University Extension The University of Wisconsin — Madison The Academy Council ing past presidents of the Paul W. Boutwell A. W. Schorger Henry A. Schuette Lowell E. Noland Otto L. Kowalke Katherine G. Nelson ACADEMY COUNCIL includes the above named Academy. Ralph W. Buckstaff Joseph G. Baier Stephen F. Darling Robert J. Dicke Henry A. Meyer Merritt Y. Hughes officers and the follow- Carl Welty J. Martin Klotsche Aaron J. Ihde Walter E. Scott Harry Hayden Clark John W. Thomson ccetLry^O letters 50C.73 W1W&3 Vol. LVIII — 197< * * :* l ,"v | ’» Cover Design by Arthur Thrall, Lawrence University TRANSACTIONS OF THE WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS LVIII— 1970 TRANSACTIONS OF THE WISCONSIN ACADEMY Established 1870 Volume LVIII THE CREATIVE TEMPER IN A COMPUTERIZED SOCIETY 1 Adolph A. Suppan TOWARD DESIGN IN THE VERNACULAR 9 William A. King MUSIC AS VIBRATIONS AND AS FLYSPECKS 15 Donald W. Krummel VICTOR BERGER: SOCIALIST CONGRESSMAN 27 Frederick I. Olson WHEN SEDITION LAWS WERE ENFORCED: WISCONSIN IN WORLD WAR I 39 John D. Stevens POLICE IN A LARGE SOUTHEASTERN WISCONSIN COMMUNITY 61 John C. H. Oh TRENDS IN WISCONSIN’S TOURIST-LODGING INDUSTRY 71 L. G. Mon they TOPOGRAPHIC INFLUENCE ON TORNADO TRACKS AND FREQUENCIES IN WISCONSIN AND ARKANSAS 101 Robert G. Gallimore, Jr., and Heinz H. Lettau THE OTTER IN EARLY WISCONSIN 129 A. W. Schorger VENTIFACTS ASSOCIATED WITH THE CAMBRIAN- PRECAMBRIAN UNCONFORMITY AT NEKOOSA, WISCONSIN 147 Ronald W. Tank PALEO-GEOGRAPHIC IMPLICATIONS OF CLAY BALL DEPOSITS UNDER VALUER AN TILL IN EASTERN WISCONSIN 153 Barbara Zakrzewska NOTES ON THE ECOLOGY OF THE HARVEST MOUSE, RE1THRODONTOMYS MEGALOTIS , IN SOUTHWESTERN WISCONSIN 163 Gerald E. Svendsen AN ANNOTATED CHECK LIST OF THE GEOMETRIDAE (LEPIDOPTERA) OF WISCONSIN 167 Charles V. Coveil, Jr. FOUR NEW SPECIES RECORDS OF SIALIS (MEGALOPTERA: SIALIDAE) FOR WISCONSIN 185 K. J. Tennessen JUNCUS EFFUSUS. I. THE SITUATION IN WISCONSIN 187 Seymour H. Sohmer GROWTH POTENTIAL OF WISCONSIN NATIVE PINES ON WEED-INVADED SOILS 197 S. A. Wilde CORIXIDAE (WATER BOATMEN) OF WISCONSIN 203 William L. Hilsenhoff TROPHIC NATURE OF SELECTED WISCONSIN LAKES 237 Lloyd A. Lueschow, James M. Helm, Donald R. Winter and Gary W. Karl ANNOTATED LIST OF THE FISHES OF WISCONSIN 265 Marlin Johnson and George C. Becker PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 60 TILIACEAE AND MALVACEAE— BASSWOOD AND MALLOW FAMILIES 301 Fred H. Utech PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 61. HYPERICACEAE— ST. JOHN’S-WORT FAMILY 325 Fred H. Utech and Hugh H. litis PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 62. COMPOSITAE VI. COMPOSITE FAMILY VI. THE GENUS AMBROSIA— THE RAGWEEDS 353 Willard W. Payne BIOGRAPHIES 373 EDITORIAL POLICY The Transactions of the Wisconsin Academy of Sciences, Arts and Letters is an annual publication devoted to the original, scholarly investigations of Academy members. Sound manuscripts dealing with the state of Wisconsin or its people are especially welcome, although papers by Academy members on topics of general interest are occasionally published. Subject matter experts will review each manuscript submitted. Contributors are asked to forward two copies of their manuscript to the Editor. 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Documentary footnotes should appear at the end of the paper under the heading “References Cited.” Supplementary or explanatory notes of material too specialized to appear in the text itself should be typed on a separate sheet entitled “Footnotes” and appended to the section entitled “References Cited.” Contributors should avoid unnecessary documentation wherever possible. Other matters of style should be in harmony with current practice in the subject matter area. Galley proofs and manuscript copy will be forwarded to the author for proofreading prior to publication; both should be returned to the Editor within two weeks. Papers received on or before July 15 will be considered for pub¬ lication in the current year. Papers received after that will be considered for publication the following year. Contributors will be given five offprints of their article free of charge. Additional offprints in sets bf 100, 200, etc. may be ordered at the time galleys and copy are returned to the Editor. Price will vary according to quantity desired and the length of the article. Manuscripts should be sent to: Professor Walter F. Peterson Editor, Transactions of the Wisconsin Academy Lawrence University Appleton, Wisconsin 54911 ADOLPH A. SUPPAN U8th President of the WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS THE CREATIVE TEMPER IN A COMPUTERIZED SOCIETY* Adolph A. Suppan I I begin this talk with a short poem. I ask all of you to reserve your aesthetic judgment until I reveal the author. Darling sweetheart You are my avid fellow feeling My affection curiously clings to your passionate wish. My liking yearns for your heart. You are my wistful sympathy, My tender liking. Yours beautifully, M.U.C.1 You will notice that the initials of the author are M.U.C. I am certain everyone realizes that the quality of the writing is some¬ what below the standard of Shakespeare, Keats or T. S. Eliot ; that is, of course, because M.U.C. is a beginner, and it may be some time before he rates a “B” in creative writing. M.U.C. is the Manchester University Computer, and I let him introduce this talk to show that in an age of technology, in a so¬ ciety already dominated by the computer, its tentacles reach every¬ where. We are all personally aware of the prevalence of the computer; it is programmed for everything from manufacturing automobiles to finding ideal mates for single people. Hospitals employ computers to analyze a patient’s ailment, count his blood cells, and compare his symptoms with the size of his bank account. Department stores use them not only in their business offices (for inventories, purchasing, billing) but to regulate escalators and revolving doors. Schools use data-processing systems to enroll their students, grade their ex¬ aminations, and decide how much to sock (hit) them for in the alumni gift campaign.2 An aircraft factory, wanting to know the equation concerning the distance a plane could fly on a given * Address of the retiring- President, delivered at the 99th annual meeting of the Academy, May 3, 1969. 1 Quoted in A. J. Parisi, “The kinetic movement : technology paces the arts,” Product Engineering , Dec. 2, 1968, p. 34. 2 Corey Ford, “A Guide to Thinking,” Think , Jan. 1961, p. 12. 1 2 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 amount of fuel, with a certain type of wing*, had the answer in seven minutes. A man with an old-fashioned desk calculator would have taken seven years; pencil and paper calculation would have taken six generations. In what we call our ordinary lives, computers provide everything from bank balances to ticket reservations to personal horoscopes.3 The computer does have its flaws and its disadvantages. Recently, translating a Russian proverb, “Time flies like an arrow,” a com¬ puter came out with “Time flies enjoys eating arrows.”4 More seriously, some authorities wonder if programmed learning might not affect man’s reasoning process to the extent that he might accept ideas without studying or questioning them ; some scientists also worry that constant graphic presentation might alter the ability to conceptualize.5 Others fear that the computer will enable gov¬ ernments to exert almost continuous surveillance over every citizen.6 II These are some phases of the overall thrust of the computer in our lives; I come now to the main theme of my talk today — the influence of a technological age and the computer upon the creative artist in our society. It is, of course, to be expected that artists react individually and differently to massive developments and events. It is also natural that the reaction can be both benevolent and malevolent. Certainly many painters, sculptors, composers, playwrights, and choreograph¬ ers show by their works that they have been influenced by the tech¬ nological thrust of our society. A reputable music critic, Frederic Grunfeld, has judged Europe’s most successful piece of avant- garde music to be Rolf Liebermann’s Les Echanges, an automated symphony for business machines, which proved to be the major attraction of a trade exhibit at the Swiss National Exposition in Lausanne. Liebermann, known in the United States for his Con¬ certo for Jazz Band and Symphony Orchestra, has scored this percussive composition : . . . for 156 office machines and mechanical devices, including typewriters, adding machines, cash registers, perforators, tape-moisteners, telephones and what-have-you, led by a computer with a mambo beat. The whole thing takes less than three minutes, but it points the way to a solution of all those problems with temperamental prima donnas and dictators of the baton.7 a John Lear, “Can a Mechanical Brain Replace You?” Collier’s , Apr. 4, 1953, p. 62. 4 Dr. Warren S. McCullock, quoted in New York Times , Apr. 24, 1966. 15 “Obsolescence for the Printed Word,” Think, Jan.-Feb., 1969, Vol. 34, No. 7, p. 19. 0 Zbigniew Brzezinski, quoted in Arthur P. Mendel, “Robots and Rebels,” The New Republic, Jan. 11, 1969, p. 16. 7 Frederic Grunfeld, Hi Fi/ Stereo-Review , Dec., 1964. 1970] Suppan — Creative Temper in a Computerized Society 3 Two years ago in New York, more than forty artists and engi¬ neers produced nine evenings of kinetic art which they titled “Theatre and Engineering.”8 In Los Angeles recently, the County Museum of Art announced a project to “mate” art and industry, involving thirty-one companies and twelve artists at work in five plants.9 Leading artists-in-residence will stay at the company plants for twelve weeks. Larry Bell, one of the artist-constructionists in¬ volved, is “group-thinking” with staff members of the Rand Cor¬ poration. He says : . . . we’re discussing light, color perception, architecture, what art really is. We’ve found out that artists and corporations and technologies can co-exist and make each other’s lives productive.30 A recent exhibition in New York’s Whitney Museum of Modern Art was filled with objects that simply would not leave the viewer alone. According to a review in the Milwaukee Journal , all of them were electronic, all of them glowed, and some of them did even more than that: they growled, spun, flashed and hummed; they didn’t just sit there, they performed. The review went on: The theatricality of the exhibition hits the viewer from the start. The show is called “Light: Object and Image,” and it is installed entirely in the dark. The darkness sets the mood, as one steps into those darkened galleries, as the light goes dim and the catalog becomes unreadable, one waits with fascinated expectation for the performance to begin. Mysterious machinery bleeps and hums. Magical surprises have been promised. The intangible aura of show biz is in the air.11 The highly-respected American painter, Robert Rauschenberg, has created a “theatre piece” which begins with an authentic tennis game with rackets wired for transmission of sound. The sounds of balls hitting rackets control the lights. During the game, the sounds turn out the lights one by one. At the game’s end, the hall is totally dark. But the darkness is illusory ; the hall is actually flooded with infrared light invisible to the human eye. A modestly choreographed cast of from 300 to 500 persons enters to be observed and projected by infrared television onto the screens.12 Another artist is produc¬ ing computer drawings based on mathematical equations and using a light source or cathode-ray tube.13 One might suggest that the artists who are “collaborating” with the computer might have some reason to fear its ultimate victory 8 “Single Channeled You Mustn’t Be,” New York Times, Feb. 5, 1967. 9 Grace Glueck, “Coast Art — Industry Project Blossoms,” New York Times , Apr. 17, 1969, p. 54. 10 Larry Bell quoted in Ibid. 11 Paul Richard of Washington Post, in “Light Show Flashes in New York,” Milwau¬ kee Journal, Aug. 25, 1968, Part 5, p. 6. 12 Richard Kostelanetz, “The Artist as Playwright and Engineer,” New York Times Magazine, Oct. 9, 1966, p. 22. 13 A. J. Parisi, op. cit., p. 36. 4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 over them. A. M. Noll tells of an ingenious experiment which, to all intents and purposes, revealed that the computer could possibly out-create the creative artist. He (Noll) gave 100 people an original Mondrian drawing and a drawing made by a computer in Mon¬ drian’s style. He asked them to judge which drawing was artis¬ tically superior, and which was produced by a machine. Of all those asked, only 28 per cent correctly identified the computer pic¬ ture and 59 per cent preferred it to the Mondrian . . . People who said they liked modern art preferred the computer-drawn picture, three to one.14 Noll comments: “I don’t know whether this is overestimating the computer’s artistic ability or underestimating Mondrian’s.”15 Some artists, however, are not too fearful. John R. Pierce says that: ... it isn’t too early for artists and programmers to study man and his arts on the one hand, and the computer and its potentialities on the other, hotly and realistically. We must decide whether men and machines should work gravely or wackily to produce works that are portentous or delicious. The choice is open, and I hope it won’t be made too solemnly.16 The jarring question for the artist, however, remains: Can the computer itself produce art, thus by-passing the artist?17 You will remember both the computer poem I used in the introduction and the episode of the fake Mondrian. Ill As I have already shown, there is a trend of cooperation toward the computer and its possibilities, evidenced by numbers of artists in our society. This is countered by a mood of rebellion which is also evident in other directions taken by the arts. In saying this, I am, of course, fully aware that the artist, being the type of per¬ sonality he is (more of that later), has often been a revolutionary in any age. I need only cite such a giant as Beethoven, whose works are now selected for the conservative portions of our symphony programs. His third symphony, considered by many to mark a tremendous advance in the entire history of music, outraged con¬ vention by its inclusion of a funeral march. His fifth symphony was condemned by a contemporary composer and critic as “an orgy of vulgar noise.”18 ^A. M. Noll in John R. Pierce, “Portrait of a Machine as a Young- Artist,” Science , Art and Communications , C. N. Potter, New York, 1968, p. 151. 15 John R. Pierce, Ibid., p. 151. 10 John R. Pierce, Ibid., p. 158. 17 A. J. Parisi, op. cit., p. 27. ^Wallace Brockway and Herbert Weinstock, Men of Music, New York, 1950, p. 190. 1970] Suppan — Creative Temper in a Computerized Society 5 Some artists of our time, like Jean Tinguely, design machines that make their own commentary on the machine. In a recent work titled Homage to New York, he presented an assemblage of a piano, machine parts, bicycle parts, a weather balloon, and fireworks, which was programmed to be seen by an audience for a number of hours; after which the “machine” destroyed itself.19 Nor is this tendency in the arts limited to sculpture. In theatre there is also a trend toward improvisation in the form of “happenings.” The “aleatory” music of John Cage, dependent entirely upon contingency or chance, is drawing crowds (and some Brooklyn cheers) wher¬ ever he appears. One of his compositions, titled U minutes, 33 seconds, consists of a pianist sitting at the piano without playing a note and then leaving. I understand that when asked “Where is the music?” Cage replied that it is in the sounds you hear in the audi¬ ence while they are just sitting there.20 The distinguished critic of the New York Times, Harold Schonberg, writes: Basically the entire avant-garde manifestation is revolt, unease, a pro¬ found dissatisfaction with current social, religious and cultural standards. At basis is the feeling that nothing means anything, certainly not when the Bomb has taken the place of God in so many minds as the ultimate disposer of the earth. The avant-garde in the arts, deriding the romantic concept of “beauty/’ has deliberately substituted an anti-ethical concept that is intended to demolish the great ethos upon which all art of the past was based.21 Joseph Wood Krutch testifies further to this rebellion. He sug¬ gests that artists have, “in their own way, signed off from their civilization almost as effectively as has the cultivated manufacturer of the shapeless dream.”22 Another observer predicts that: If today’s trend continues in theatre, we may all look forward to an in¬ flux of poorly-constructed plays covering up their inadequacy with a generous hunk of pornography, a liberal sprinkling of four-letter words and a sugar coating of poetry. This pretense at “free expression” is really no more avant-garde than a ready-mix cake.23 The jarring messages given by the rebellious artist to his society correspond with the free admission that he is disgusted with its dehumanization of human life as well as its despoilation of nature. This is a revolt, whether we relate it to our military-industrial com¬ plex or to our mass-society. Even the individual’s name is on the 19 Nathan Knobler, The Visual Dialogue, New York, (n.d.), p. 204. 20 Summarized from Look Magazine, Jan. 9, 1968, p. 45. 21 Harold C. Schonberg-, “Art and Bunk, Matter and Anti-Matter,’’ New York Times, Sept. 24, 1968. 22 Joseph Wood Krutch, “The Creative Dilemma,” Saturday Review, Feb. 8, 1964, P. 17. 23 “To the Editor,” New York Times, July 28, 1968. 6 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 way to becoming meaningless. He is identified merely as a group of numbers by his student admissions office, his insurance man, or his gas station attendant. IV This evidence relating to the artist's confrontation with tech¬ nology and the computer should be followed by the direct question : Why is the creative temper even more necessary to our technological society than to any past society? A recent psychological study of creativity — The Creative Person — made by a group of psychologists on the University of California- Berkeley campus, is very informative here. It concludes (and I sum¬ marize) that the creative person is inclined to be interested and cur¬ ious, more open and receptive than others ; that he is strongly moti¬ vated to achieve in situations where independence of thought and action are called for ;24 that he has an openness to experience, a free¬ dom from crippling restraints and impoverished inhibitions, and a delight in the challenging and unfinished.25 These characteristics marking the creative temper (and I must quickly point out that the creative scientist, as well as the creative artist, was considered) make me ask: In our society — shadowed by urbanization, mechanization, and over-population — where the per¬ son is in danger of becoming a non-person, is not the creative in¬ dividual a last defense? It is by now a truism to state that these qualities of character — independence, originality, open-mindedness — are more needed than ever by our society. These qualities are needed to challenge the forces a technological society has set in motion, forces that obliter¬ ate personality psychologically, not to speak of what can take place when computer-programmed missiles obliterate us physically. Truly, as the arts of a civilization have often served to symbolize a nation's achievement or failure, the treatment of its artists has revealed the degree of freedom or oppression within its borders. One might therefore say that our age, more than any other, will be judged by future historians in relation to how it realizes the debt it owes to these free, independent spirits who might help prevent a society from melting its men into ciphers. This implies, of course, the need for a greater recognition of the creative individual and his contribution to our culture. As Archibald MacLeish has said : What’s wrong is not the great discoveries of science . . . What is wrong is the belief . . . that information alone will change the world. It won’t. ^Donald W. MacKinnon, “What Makes a Person Creative?,” Saturday Review , Feb. 10, 1962, p. 17. 25 Ibid., p. 69. 1970] Suppan — Creative Temper in a Computerized Society 7 Information without human understanding is like an answer without its question — meaningless. And human understanding is only possible through the arts.26 The arts and humanities provide meaning and purpose to our lives. The artist— in many different ways — probes, searches, and reinterprets reality so as to make our lives deeper, wider, and richer because of his efforts. When we need a jolt, he jolts us, with disso¬ nances or happenings ; when we need a shock, he shocks us, often to tell us that we’re taking the wrong road. I am fully aware of Plato’s overt reason for excluding poets from his Republic — -fear of the emotional influence of great art. But I’ve always had a sneaking suspicion that what Plato was really worried about was that the rebellious, independent, poets would upset the applecarts in his neat, controlled, structured, little state. Certainly our nation is in danger of being computerized beyond belief, organized beyond belief, and benumbed beyond belief by the offerings of the mass media — so much so that the high prophet of the electronic revolution, Marshall McLuhan, has changed his gospel from “the medium is the message” to “the medium is the massage.” The creative temper, as I have emphasized in this talk, can re¬ mind, prod, and inspire us to sustain the value of the person in a non-personal world. 26 Archibald MacLeish, “Thoughts on an Age that Gave us Hiroshima,” New York Times, July 9, 1967, Section 2, p. 1. TOWARD DESIGN IN THE VERNACULAR William A. King There exists in this country a discontent, an almost voiceless potential, with little direction and few spokesmen. This discontent is the result of our lack of aesthetically satisfying visual and tactile experiences. It is a voiceless potential because it is the un¬ spoken yearning for harmony and proportion that every man seeks consciously or unconsciously in his surroundings. There is little direction because few in positions of decision-making are concerned with the yearning. Little effort is directed toward giving a unity of expression. Each one of us is part of this underground potential. Its basis is in the biological and psychological needs which should be reflected in the way we live and in the things we use. The way we live is ex¬ pressed in a jumble of diversions. We are surrounded by cacophony, foul air to breathe and offensive visual experiences. Phonographs look like antique chests, plastics imitate marble, kitchens imitate other factories and are merely as efficient. From the design of the development house (boxes within a box) to the form of the latest automobile, there is no effort at appealing to any one aesthetic sense. There is instead only a confusion of many directions. The recent epidemic of ludicrous tail fins on our auto¬ mobiles is symptomatic of our plight. But if there is discontent, it may be asked, why is there no public protest? Perhaps because man, in his infinite capacity to adapt, shuts out what is intolerable. He no longer notices the unacceptable, just as the soldier in battle can ignore sights of death and mutilation. In this paper I wish to trace the development of this phenomenon of life today. In eighteenth-century America, before the industrial revolution had a strong grip, the objects of daily use expressed in a natural way the lives of the people of that time. There was a dialogue be¬ tween the artisan and the user of his product. The consumer knew what he wanted and he got it; the craftsman was qualified by his sensitivity and his apprenticeship. Since there was this natural alliance between the artisan and the consumer, the results were generally satisfying. It was in the design of useful things that the American showed his creative genius. Creative impulses, untram¬ melled by tradition, were released. The character of early American 9 10 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 design was summed up by James Fenimore Cooper, in Notions of the Americans (1828) : “I have seen more beautiful, graceful and convenient ploughs in positive use here, than are probably to be found in the whole of Europe united. In this single fact may be traced the . . . character of the people,, and the germ of their future greatness. Their axe is admirable for form, for neatness, and for precision of weight . . . the actual necessities of society supply an incentive to ingenuity and talent, that are wanted elsewhere . . . the vast multitude of their inventions ought to furnish food for grave reflection to every stranger.” Mass production was possible in the 1830*8, and it gave almost everyone machine-made chairs, carpets and wallpaper. Designers tested the limits of the machine in their competition to come up with the most ornate product. Expediency took the place of art. Walter D. Teague writes of this industrialization : “While the Revolution had none of the grace and charm of childhood, it had the clumsiness, the ineptness, the unintentional cruelty and the pains of a gigantic, lumbering, grimy, immaturity. It had, too, the eagerness and vitality of youth ... It accepted as a matter of course that the new industrial system on which the whole new scheme of life was based should proliferate in sinister black factories that blighted the lives of their workers ... It re¬ ceived with uncritical acceptance the floods of crudely em¬ bryonic wares that poured from these factories to supply our needs ineptly, while they swamped our lives in ugliness.” The end result of the surfeit of badly designed products in this country was that no one was satisfied, as the craftsman and con¬ sumer were satisfied in a less complex age. The disparity between man striving for beauty and the ugliness of the world is not new. In England Josiah Wedge wood endeavored to solve the problem in the mid-1700’s by enhancing commercial pottery with applied decoration, which emphasized already existing beauty. William Morris a century later counseled, through modified medievalism, that man should ignore the machine entirely and re¬ turn to the days of handicraft. “As a condition of life, production by machinery is altogether an evil . . . art must be produced by the people and for the people, as a happiness for the maker and the user.” And he insisted on the importance of aesthetic considera¬ tions in the design of even insignificant objects, an idea which has had far ranging implications up until the present. Perhaps his most important contribution was the establishment of arts and crafts schools throughout England where young designers studied the possibilities for functional yet beautiful design. 1970] King — Toward Design in the Vernacular 11 Morris’ ideas were followed in Europe by the movement called Art Nouveau. Art Nouveau, as its name suggests, attempted to create a new style. International in character, it was known as Art Nouveau in Belgium and France, Sezession in Austria, Jugend- stil in Germany and stile lihertd in Italy. The Belgian Henry van de Velde, one of its leaders, urged . . a logical structure of prod¬ ucts, uncompromising logic in the use of materials, proud and frank exhibition of the working processes.” Significantly for industrial design, the German Hermann Mu- thesius advocated the study of “railway stations, exhibition halls, bridges and steamships . . . whose shapes are completely dictated by the purposes they are meant to serve.” In 1907 Muthesius founded the Deutscher Werkbund, which was a step away from the arts and crafts (Jcunstgewerbe) toward a true industrial art. Its ideal was stated in Muthesius’ inaugural address : “There is no fixed boundary between tool and machine. Work of a high standard can be created with tools or with machines, as soon as man has mastered the machine and made it a tool.” Standardization was the goa(, and it was only through it that reliable taste could be achieved. The arts and crafts movement inspired by William Morris in England made its impact in the United States in the 1880’s. Ex¬ amples of this influence are the glassware of Louis Comfort Tiffany in New York and the Rookwood pottery of Maria Storer in Cincin¬ nati. These were aesthetic protests against the poor quality of factory production. In more modern times serial production, since its standard is that of indefinite repetition of objects, has changed the attitude of the consumer. Uniqueness or skill of craftsmanship is no longer a consideration; only the design is important. Novelty, however, has increasingly occupied the minds of merchandisers. Designers have borrowed criteria from cybernetics and feel that overfamilar- ity produces obsolescence. The greatest amount of pleasure is de¬ rived from newness, because of its ability to surprise us. These ideas have their roots in the writings of the English empiricist Burke, who formulated a functionalist attitude toward art. Burke writes in A. Philosophical Inquiry into the Sublime and Beautiful (1756) : “When we examine the structure of a watch,, when we come to know thoroughly the use of every part of it, satisfied as we are with the fitness of the whole, we are far enough from perceiving anything like beauty in the watch work itself . . . the effect is previous to any knowledge of the use, but to judge of proportion, we must know the end for which any work is designed.” He shows the distinction between beauty and proportion and fitness and knowledge of use. And in the same work he states: “Indeed beauty is so far from belonging to the idea of custom, that in reality 12 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 what affects us is that manner is extremely rare and uncommon. The beautiful strikes us as much by its novelty as the deformed itself . . . For as use at last takes off the painful effect of many things, it reduces the pleasurable effect in others in the same manner.” Concepts such as those expressed by Burke underwent consider¬ able change in the late 19th and early 20th centuries. Functional¬ ism, which was originally understood in its materialistic meaning, now took on psychological implications. Beauty became synonymous with function. Louis Sullivan maintained that “form follows func¬ tion.” In the 1920’s the Bauhaus set out to educate the industrial designer in the premise that beauty and utility meet in the well- designed object. Such men as Klee and Kandinsky, the architects Mies van der Rohe and Marcel Breuer, adapted the new principles of the arts to the unique requirements of serial production. It was hoped that the lowest possible cost would produce the best aesthetic result. Industry would be furnished with functional designs which were clearly made by machines. With the rise of the Nazis in Germany the Bauhaus was closed. Many of its faculty came to America, and the works of Gropius at Harvard, Moholy-Nagy at the Chicago School of Design, Kepes at M.I.T. and Mies van der Rohe in the Chicago area have had their impact. America’s own Frank Lloyd Wright had more influence on Europe than on his own country. Walter D. Teague, Henry Dreyfuss, Raymond Loewy, and Charles Eames have also made an impact with designs ranging from steamships and telephone re¬ ceivers to bent plywood chairs. In America the emphasis is on styling. The change is on the sur¬ face. Objects change form with what is thought to be the latest mood of the consumer. When aerodynamics occupies the primary concern all objects take on free flowing lines. The theory of entropy of communications hopes to produce the maximum amount of sur¬ prise by a deluge of new styled products. It is held that the greatest amount of information is supplied by a form which, because of its newness and unforeseeableness, gives the greatest amount of surprise. The amount of information is in direct proportion to its degree of surprise. When the form is repeated too often, there is a diminishing amount of information (the form is ignored by the consumer.) The idea that novelty produces aesthetic pleasure takes on cynical overtones and gives rise to planned obsolescence. The benefits of the machine are obvious in our time, and there can be no turning back. The Bauhaus offered an answer to the problem of designing intelligently for the machine. It seems that much of the message has fallen on deaf ears. It would seem that the manufacturer has not discovered that additional dimension, and 1970] King — Toward Design in the Vernacular 13 this keeps his product from being well designed. Much of the prob¬ lem centers around the appropriateness of form and the function of decoration. Surely novelty is not the only thing which will appeal to the public. Many business establishments are aware of the necessity of pre¬ senting an image in plant and administrative office appearance. The best architects are often engaged for this purpose. Why then is the consumer product shoddy in so many instances? An indus¬ trial designer, Richard S. Latham, says : “The quality of materials used and the characteristic details become more skimpy and in¬ appropriate, until finally it appears that the worst design, the most inept craftsmanship and the least beautiful workmanship have been relegated to the individual consumer, with higher orders of skill and execution reserved for industrial products, and the high¬ est order of concept and execution reserved for products that hu¬ man beings will hardly ever see.” Industrial design is the only really popular art form. It has the influence to educate the public in the positive values of modern art. Our condition suggests that manufacturers are not seeking pro¬ fessional ideas in the solution of design problems. College and uni¬ versity departments of industrial design have been trying to edu¬ cate students for more than a quarter of a century to solve these problems in an honest way. And yet their efforts are not very apparent. Imitation of handicraft is not an honest solution, and the machine does not do it willingly. Perhaps the market analysts are second guessing the public. Perhaps businessmen are afraid to disturb the situation which has arisen from the misuse of ma¬ chines by misguided men. Susanne K. Langer addressed herself to the problem in Feeling and Form : “The artisan-craftsman has been superseded by the in¬ dustrial designer; and industrial design is next to architecture in shaping the visual scene. So it is in our things — our countless things, multiplied fantastically praeter necessitatem — that we must find some significance : a look of simple honesty in ordinary utensils, of dignity in silverware, and of technological elegance in our machines.” The time is long overdue for industry to bring its products into line with the limitations and the advantages provided by machine production. Forceful leadership by designers and businessmen is called for. Design in the vernacular can be achieved. It can bring a grace to our lives which has been absent. MUSIC AS VIBRATIONS AND AS FLYSPECKS Observations of a Music Bibliographer on the Unifamiliar Effects and Inherent Perniciousness of His Chosen Objects of Research. Donald W. Krummel It is one of the curiosities of our language that “live” music should be that which will not survive. Like fruit, music keeps in cans or when frozen. Like so many other human achievements, music has been pre¬ served in written records. We know the past not through our memory of events but through documentary evidence, most of it preserved on paper. Axiomatically, that music which predates our written records is pre-historic ; so then also is any music which we may hear which has not been notated or recorded. Paper enables the musician to benefit from the past. As we shall see, it also com¬ mits him to the past, developing his art into one of understanding, interpreting, learning from, and building on the basis of the past. This study undertakes to survey the relationship between music and its documents, in terms briefly of (1) basic reasons; (2) his¬ tory; (3) effects; and (4) future prospects. The subject itself in¬ evitably evokes a wide range of responses, from precarious specula¬ tions to the most painful of truisms (“tear up his scores, and where is Beethoven?”). While I shall hope to develop the speculations out of some of the more significant of the truisms, quite clearly my conclusions are contemporary and highly personal rather than the product of any timeless reasoning. The activity of handling musical documents, I believe, could not have found the meaning which I am here proposing without the benefit of rather basic and widespread changes in our general intellectual attitudes during the past few years. To be sure, we have always had misgivings about our musical heritage being preserved on paper. We concede that, until record¬ ings came along, we were completely dependent on notation for saving our great musical masterworks. But we still feel the need to be both skeptical and demeaning of the paper. The notes we laughingly pass off as flyspecks, of which there are two varieties : the dead dots which our tiresome scholars study and analyze, and the silly dots which our mad composers trace in order to make the great idiotic compositions of today. The flyspecks are a mere re¬ flection of the action, the harmonious vibrations in which is em- 15 16 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 bodied music itself. The fact remains that, without notation and its painstaking formulation and study, our music itself would be an achievement far less significant than what we know and enjoy today — considerably less well developed, clearly an anachronism in our age, truly pre-historic. What has led Western Man to go to the trouble of committing music to paper? First and most obvious is our belief that a musical entity is suitable for and deserves re-performance, which can best be accomplished through preservation — that its sounds, or at least their component relationships, ought to be heard again. This belief may be based on two attitudes. One is a moral, even a religious re¬ sponsibility (“we must save this”) , the other a volition which comes from enjoyment (“we want to save this”). We believe — and quite correctly — that oral tradition is fallible, that the passing of a mes¬ sage by word of mouth is not trustworthy, especially when the mes¬ sage is complicated. Apart from preservation, we wish or need to accommodate a middle man. Divisions of labor usually result as our civilization be¬ comes more sophisticated; in this instance the creator becomes separated from the re-creator, that is, the composer from the per¬ former. Sound itself is transmitted by performer to audience ; the notation enables the composer to communicate with the performer. Behind both, apart from but governing both, as something of a Platonic ideal, is the abstract concept of the work of music itself. Third, we seek a wider circulation of a work. The music be¬ comes part of the repertoire, not of one performer exclusively, but of many. Thanks to notation, the performer no longer needs to commit the work to memory. We are thus involved in the act of pub¬ lishing, which requires promotion and publicity. Implicit is the attitude that music should be shared by per¬ formers — an admirable sentiment at any time, and probably the exception in the larger course of music history. Such generosity departs from the practice of the artist’s repertoire being a closely guarded secret. In eras of great virtuosity, to be sure, the notation may become the merest of outlines, in which case the publication is no act of generosity at all. The masterful performer shares the text with his colleagues, and then in comparison to them shows his superiority of skill and taste. Finally, somewhat opposite to altruistic sharing is sharing for profit. Music becomes a commodity, a means of making money, a basis for commercial gain. Subject to copyright — a “literary” or “intellectual property,” of all things — it provides the musician with a means of survival. He can flout the gods who had prescribed his 1970] Krummel — Music as Vibrations and as Fly specks 17 lot as one of starvation, and, with exceptional luck, get rich and lose his musical soul entirely. Out of such considerations, notation on paper — for all intents and purposes a permanent medium — has joined forces with an art form which is essentially fleeting and impermanent, made up of vibrations which are produced, resonate for an instant, and are gone — which live and die in the tragedy of immediacy. (The word “evanescent” was a favorite in describing it in the Romantic era.) In their essence sound waves, and therefore musical compositions, are momentary, and this we should not forget : such is their limita¬ tion, also their virtue, and their significance today. The commitment of music to paper thus results in an alliance between two media, one visual and the other aural, one directed to the ear and the other to the eye. When the occasional and inevitable family conflicts arise between the two, the notation always loses. This is as it should be, since music was originally, and is essentially, sound and not paper. We can see the way notation loses out as we follow the current fashion of pondering our everyday idioms. We adapt an old mili¬ tary expression and speak of a performer “facing the music,” mean¬ ing that he has chosen to do his own thing which is not J. S. Bach’s own thing. The printed page then brings him rudely back to or¬ thodoxy. (Thus, in current colloquialism, have our flyspecks func¬ tioned as the fall guys in the Great Creative Cop-out of Western civilization.) We also use the German term Augenmusik — music of the eyes — in speaking of a composition which is more rewarding in study than in listening. The term is not precisely appropriate: a better term might be Koyfmusik — “head” music, or at least “heady” music. If the fact be known, there has been very little true Augen¬ musik in the sense of music pleasing to the eye. As a graphic art, musical notation through history has fared very badly indeed. There have been very few great masterpieces of music book produc¬ tion. The thrilling prints of Petrucci, the first great music printer, and the handsome early engraving of Domenico Scarlatti’s sonatas come to mind; but beyond this even the most experienced music bibliographer will have trouble finding examples of which he can be proud. The early twentieth century saw several attempts to make music beautiful on the page through specially prepared music type faces, fine paper, elegant design, and tasteful decoration. The re¬ sults were hardly successful. Music which is visually attractive almost inevitably, and most unfortunately, becomes affected in its appearance. The performer wants his instructions stated in as clear and unornamented a version as possible — and in view of the 18 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 speed and exactitude .with which he must grasp his instructions, his needs are indeed critical. In printing, better an ugly legible statement than a beautiful illegible one. Similarly in publication: better an ugly edition of good music than a handsome edition of bad music. Through the course of Western history, at least up to the twen¬ tieth century, music has found permanence by imitating the printed book. Music has enjoyed a free ride in the vehicle of literary texts; and as a result music has had to go where the literary vehicle was willing and able to go. This influence has yet to be extensively studied or appreciated. At this point, then, a survey of the main events in this history is in order. We find the earliest notation of Western music, as it is traced back to the Middle Ages, already involving either numbers or words. Pitch levels are based on mathematical relationships. The names of these levels are assigned with word syllables, as in solmisation, or later with the letters of the alphabet themselves. Musical rhythm is derived either from the natural rhythm of spoken words, or later from mathematical subdivisions of time duration. With the invention of printing in the mid-fifteenth century, the development of music printing a few decades later, and the emer¬ gence of music publishing soon after 1500, music becomes all the more strongly committed to words. One admires and is fascinated by the achievement of the early craftsmen who conceived the first music type faces; one also respects their output, which provided the permanence for most music written during a span of two cen¬ turies. Musical notation by 1500 had already come to resemble what we know today, to the extent that it consisted of symbols arranged in a line, like the letters of a word. To be sure, the staff lines them¬ selves caused the printer some difficulties which he never solved completely ; but it is hard to doubt that movable type, as soon as it was invented in the days of Gutenberg, was destined to be applied to music. Early type could not directly designate instructions for musical color or harmony. These two elements, we might observe, are in themselves less significant in Renaissance and early Baroque music than they were to be later. Someone someday will perhaps defend the hypothesis that print¬ ing contributed to the transition from polyphonic music to that of the continuo period. Would figured bass have been adopted if per¬ formers could have had printed chords to read (and, by having had more of them to read, would have learned to sight-read them) ? Was there also in the late Renaissance, as part of some larger sub¬ conscious arch of civilization, a need and desire for “line,” for simple linear construction in music? For the first time, man dealt 1970] Krummel — Music as Vibrations and as Flyspecks 19 extensively with books, where one thing happens at a time ; and at this time his imagination and interests were first being stimulated by a knowledge of exploration and travel, involving a person going to only one place at a time: perhaps such factors helped to dis¬ courage polyphony, in which several lines are presented at one time. The great sixteenth-century commercial empire of music pub¬ lishing, based on movable type, finally collapsed and was replaced around 1700. As early as 1620, new music type faces were seldom being designed. This is only a detail in the story of this period, to be sure — the Thirty Years War and the various forms of puritanism had ravaged Europe, and in fact new type faces of any kind were rarely to be seen. In music, the old type was used in religious service books, in treatises— again reflecting a tie to the printed word — and in popular song anthologies. Progressive instrumental music suffered in particular. No notes were available for rapid passage- work, and chords could be constructed only by carefully chipping two or more pieces of type and fitting them together. We can thus add the upheaval of 1700 to our list of those musical revolutions which have obligingly happened every century, on the century. In this instance, liberation was not from a tired and cor¬ rupt artistic tradition, but from a book trade best suited to doing other things. Music from this time forward was on its own course in the publishing world, using engraved plates rather than mov¬ able type. With independence came also the loss of the usual chan¬ nels of distribution and registration control : the librarian today can seldom rely on the standard historical bibliographies for evi¬ dence on published music. The circulation of some music even went underground, although partly for reasons of control over per¬ formance : Italian opera, for instance, conquered Europe not through transmission in typeset editions or even engravings, but through a highly developed manuscript copying network. We can also plot a two-hundred-year historical cycle: music printing around 1500, music engraving around 1700, and sound recording around 1900. The implications of the last development are perhaps the most staggering of all. The marvelous Siamese- twin conveniences — permanent storage of the sound itself, and mass-media distribution of sound — are obviously very great tech¬ nological “breakthroughs.” Typically, they have eliminated produc¬ tion workers (i.e., musicians) and require more service workers (i.e., managers and electronics repairmen). At the same time, the surviving production workers are infinitely more effective: they reach a wider audience, and incidentally get paid slightly better. But typically the inventions have also led to many of the ills which beset music today: the virtual elimination of regional non-con¬ formity; audience apathy; the decline of “live performances,” at 20 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 least of the institution of formal concerts ; and in time perhaps the elimination of the performer, the composer in the de-specialized world of tomorrow communicating directly with his audience by creating his own sounds. Even now, the craft of music engraving is dying. Nobody is really sure what music publishers themselves are up to, and while they claim to be happy, neither the composers nor Wall Street seems to care much for them. The era of music on paper may thus now be regarded as the proper domain of the his¬ torian. It has become part of the past from which we are expected to learn something. What then is our heritage of music on paper? What characteristics of our music are the result of paper? We have considered what we want when we commit music to paper: more important is what we actually get. In what ways — pace Mc- Luhan and the ambiguous verb which he has taught us to say to¬ gether — “is” the medium the message? We speculate and surmise, but with no real certainty: there is of course no parallel civiliza¬ tion with music not committed to paper with which we could make a clinical comparison. Even so, we can develop several lines of rea¬ soning which tell us what the marriage has done to one of the part¬ ners. From this we can anticipate the freedom and to a lesser degree the loneliness which will characterize the newly found single life. Let us begin with McLuhan’s concept of the linear-— the idea of progression from one point to another, such as we experience in countless ways : reading a text from one word to the next, travel¬ ing from one place to another, reasoning logically step by step, growing from childhood to adulthood. Before printing we com¬ municated in “auditory” rather than “physical” space. Our com¬ munication, being mostly oral, took place in time rather than through the two-dimensional visual surface of paper or other documents. It is wrong to say that auditory space is not linear, as I some¬ times think (but am really not quite sure) McLuhan would have us believe. In its various forms — primitive, pre-Renaissance, and that since the invention of printing — music is always committed to a temporal “line.” Line as perhaps been emphasized, or more systematically conceived, since the invention of printing. The de¬ vices for notating the elements of music were fixed long before the Renaissance, and then accommodated in movable type. Rameau's formulation of the harmonic progressions in the eighteenth century, the monumental Western codification of its practices, is a complex system of rhetoric and logic rather than a grammar or spelling guide, the appropriate counterpart to the succession of words on a 1970] Krummel — Music as Vibrations and as Fly specks 21 printed page. When confused by new music even today we say “I don’t follow,” as if we were lost in an argument or discussion. Line, the term we use for the sequence of sounds in time, is an essential dimension to all music, the other dimension being the variety of the sounds occurring in a single moment — color, and in a static sense, harmony. Line has certainly been conspicuous in the art music which we most highly esteem. We admire and are moved by music which brings out the “long line,” be it an Urlinie in Schenker’s musical analysis or the delicate spinto affectations of a great lyric soprano. Italian and German music, one might speculate, are generally more linear than French. The frequent abandonment of line is regarded as a hallmark of the new music, foreshadowed by the Romantic color made possible by the technol¬ ogy which produced the modern symphony orchestra. Composers are supposed to delight now in bright bursts of sonority— perhaps for purposes of being non-linear, unconsciously or self-consciously, possibly also to wake us up and keep us awake, and perhaps be¬ cause the two are one in the same. In many of the non-Western musics, I am told, the linear element is also less conspicuous. Even in the most advanced music of the future, line is inevitable, since time — like physical space — has dimensions. Music always has a line, although it is possible that because of printing the line is more conspicuous. Second, the printed page of music offers escape — a refuge from the bright glare of musical sound. The metaphor of a “bright glare” of course, is logically inappropriate, and in the same way as musical “color” is. Coming from the world of light rather than sound, it is useful only by way of suggesting the peculiar way in which sound engulfs us. Sound varies in loudness, and usually the hearer can locate the source of the sound. But we can not avoid sound by turning our head as we can avoid looking at a visual object. It is important for a listener to be able to get away from music. This was felt as early as the Renaissance, when the audience came to be placed further away from the performers, especially the large groups of performers. Thanks to opera, the proscenium arch tended to be used for music as soon as it was devised for the theatre. In more recent eras the classic escape at a concert has of course been sleep. Today earphones offer a further element of privacy. Having the music we want when we want it is an unpre¬ cedented and staggering blessing, the only limitation being our ability to absorb very much of it at one time. We will still want and need to get away from it at times. It may prove to be one of the typical ironies of history that, at the very moment when we 22 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 have the totality of the musical repertoire available, we will least care about it or need it. Third, paper makes possible analysis : the printed page helps us comprehend music by allowing us a limited and a different access to it, enabling one set of senses to be reinforced by another. Music on paper lends itself to a varied manner of comprehension, the lay¬ ing out flat on a two-dimensional surface making possible an im¬ pression of the totality at a glance. The score becomes a map of the terrain; and while there is admittedly no way to know the countryside better than through a good walk, we can correct many of our errors if we take along the map. Fourth, and most important in many ways, is the prospect of betterment made possible largely through analysis. The composer can study the past and learn from it. He learns to hear his music “in his mind's ear” as interpreted through his eyes ; and from this he can discover his own errors and correct them, his weaknesses and strengthen them. In an abstract way, his work can evolve in its perfection. He can work as a Beethoven, re-examining his achievements and thereby building an organically conceived type of music — keeping in mind all the time, of course, that there are also Mozarts who are no less great for having comprehended intuitively so many of the relationships which are to him so thoroughly a rational process. Fifth, paper offers tangibility. Sound, being impermanent, is also undependable. We ask the man we deal with to “put it in writing” ; and we argue endlessly after a concert, always about what the per¬ former accomplished, often even about what sounds actually were heard. Control becomes possible with the printed page — the per¬ former's job becomes one of making music in terms of conditions spelled out, the degree of freedom depending on the music. Stravin¬ sky would have the conductor of Le Sacre acting largely as a cuing metronome ; the composer of Neapolitan opera, of a concerto arriv¬ ing at a cadenza, or of a pop tune intended for a jazz combo, draws in only the rough sketch, asking the improvising performer to take off like a liberated bird, making sure only that the flight follows the suggested course or lands at the right airport. In all such situa¬ tions, the written notes, being fixed, are the means of control. Through our copyright laws, they take on the characteristics of real estate and personal property. They get bought and sold, and have resulted in music industries as concerned with self-perpetua¬ tion as our great corporations. The notes engender their own laws and rules ; and they get hauled into court because of those regulations. Finally, they also get us into heaven, if they’re good enough. Permanence, and the prospect for improvement, together lead to 1970] Krummel — Music as Vibrations and as Fly specks 23 immortality — to timeless musical monuments, the concept of the heroic Romantic musical genius leaving footprints on the sands of time. Through paper, music, long assailed by puritanism as sinful and ungodly, achieves revenge, offering its favored practitioners its own brand of salvation apart from the rules and regulations of the church: in effect, “instead of getting to heaven by being good, live it up, write a great symphony, and you’ll make it.” It is thus much in order here to recall a lovely old German canon with the following text : Himmel und Erde mussen vergeh’n, Aber die Musici, bleiben besteh’n. Literally translated, “Heaven and earth must pass away, but the musicians will always remain.” Really quite outrageous. Today the words would probably read instead, Soon the Establishment ceases to swing, Leaving musicians a-doing their thing. Or, as our feelings may become more specific, “When our institu¬ tions collapse of their own cumbersomeness, our cultural centers go bankrupt, our paper turns to pulp— then we’ll be left with music.” The innocuous Sdngervereine who perpetuated this ditty certainly never thought of doing any such thing, but they have indeed brought us face to face with the doom of the flyspecks, the fall of the gods, the movement of the tide which will smooth the sand, erasing the footprints of the 3 B’s. The Armageddon we are talking about is not in itself the great battle going on today for social change, the eradication of poverty, or the rise of the non-white races — although the two are con¬ nected: music is part of society, and there are obvious parallels between our social and our musical establishments. By way of a brief digression, we might observe that even if the parallels did not exist, music would almost surely play a conspicuous role in social conflict. Its well-known emotional appeal is only half of the picture. Existing as it does in time, music is the very essence of change, of creating beauty in a context of impermanence. In days of uncertainty, it is symptomatic that we should so often hear the expression, “Play it by ear.” To the musician the phrase means memorizing the notes and then executing them. The world of com¬ merce flatters him by defining it even more broadly, as going into a difficult situation with no fixed course of action at all in mind. Music’s message is less obvious than that of words and pictures ; thus it becomes the medium for reflecting those pulses and 24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 rhythms, those subconscious feelings and sensations which other forms are unable or unwilling to express or reflect. Music may be harmless and lovely in its purity as an autonomous art form ; but as a means to an end, it can much too easily also be highly potent, intellectually stultifying as only an emotional appeal can be. If musicians are less “involved” today than their music is, perhaps this is because they can see the whole process of reform as merely one more mind-blowing operation, at least at the stage where music gets into the act. They can be sympathetic with the cause of social justice; but they also have strong impressions of how democracy in America has preferred mayhem and inanities on television to live artistry — how popular education has produced technicians rather than humanists, and how the more abundant life resulting from the battle against poverty is conceived largely in terms of Gross National Product. Musicians perhaps have a better pipeline than we give them credit for. Their music has frequently revealed some important things about ourselves which we were not ready to accept. But they have also been all too quickly ridiculed for the attitude “My kingdom is not of this world” or “after all the blue meanies get bumped off, we meek little rascals will inherit the earth — the Bible tells us so.” Thus it is well to return to the little German canon to note that the word is “Musici,” and not “Musica” : what will re¬ main is not the music itself, but the musicians. We really must be allowed to stretch the point here and say that music-making is what will remain. The musicians’ bodies and talents, like their composi¬ tions, must be regarded as part of the Himmel und Erde which will pass away. The musical experience is fixed in the human condition, and beyond this in the vibrations of the stars. Music on paper has obviously played a large role in the process by which music has become increasingly committed to the past. Fifty to a hundred years ago, concerts came to favor the “tried and true” at the expense of the present. Within the past fifty years, our musicologists have sought to fill in the gaps in the panorama of Western musical development. Today the musical experience is largely an archival experience, our values those of the historian. What we make of our musical past may bother our sense of honesty, and quite appropriately. Hitler loved and used his Wagner; and the modern administrator loves and uses his Machiavelli. But to deny that our most cherished musical experiences are important to us and in some way bettering is dangerously close to a denial of that vague but important link between the humanities and humanity. We all piously insist on a need for musical vitality. The price may be expensive indeed. Probably we would need to abandon the institutions which encumber our music, not only the flyspecks and 1970] Krummel — Music as Vibrations and as Fly specks 25 recordings in our libraries but also the stultifying etiquette of our concert life. Also vulnerable are the concert halls themselves — in¬ deed they are probably the very proof of Parkinson’s “law” about institutions deteriorating when they move into an edifice properly suited to their image of themselves. Along with all of these monu¬ ments, alas, must go the Art of Fugue and Messiah , the Mozart con¬ certos and the Beethoven quartets, Otello and even Wozzeck. We will never excel them — such is one of the obvious assumptions to¬ day, and whether inherently true or false, it will be true as long as we believe it. With this in mind, I must take exception to a well-intentioned but wrong-minded defense of the arts in our society today on the basis of their excellence. We do indeed need excellence; and the level of excellence in the arts is indeed high enough to be a model for other activities today. The experience of music, like that of her sister arts, is one of stimulation, accomplishment and pleasure; and such being the case, the already high level of excellence will be further heightened by competition in an inevitably overcrowded profession. In practical terms, the results are likely to be less happy. For economic reasons — supply and demand, together with the tech¬ nological “breakthrough” in sound recording mentioned earlier — the Gradus ad Parnassum is missing some steps near the top. The boy who practices seldom gets to Carnegie Hall. The excellence toward which the vast army of our educators must work must be fitted into a context in which amateurism, rightly and understand¬ ably, equals amateurishness, in which local pride is often an emo¬ tionally charged but valid excuse for quality. The ascent from the great plains, vast if less arid than we imagine, to the Olympian heights, is sudden, steep, and with frightening odds against sur¬ vival. “It’s warm in here— -yes, perhaps for violinists” ; and there¬ fore, “If you can’t stand the heat, get out of the cotton patch.” And as a result, the Global Village Philharmonic will soon be impeccably performing the opera omnia of Western music, giving us with pushbutton convenience all the listening pleasure we want. Rather than justify music on the practical grounds of its excellence, one should perhaps accept its total uselessness as its greatest virtue — it does less harm than politicians or scientists. Far better one should hope and work for the impractical, unpredictable, but now highly possible: an aesthetic right-headedness of some sort, com¬ parable to the recent moral righteousness over Viet Nam in this country, serving to remind us that the musical experience is more rewarding in achievement than in gratification, being in es¬ sence a creative art rather than a consumer commodity. Rather than justifying music in terms of an administrative value in our society today, it would be nice to think that we might seek 26 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 to apply one of the administrator’s favorite laws of positive think¬ ing : when faced with two alternatives, come up with a third which, with instinct and effort going for it, will prove to be brilliantly ap¬ propriate. Can we keep the past without being its slave? The mind boggles at what the answer might involve : but experience leads to the hunch that somebody will be singing about it before the mind stops boggling. A dispassionate and analytical glace at the popular music of today will perhaps help to renew our faith. Its texts usually tend to strengthen our respect for the social values of adolescents ; and similarly, its musical content leads to a sanguine hope that a new creative era is at hand. In comparison with most of the popular (and much of the classical) music of the past, its content is indeed richly varied, imaginative, and frequently quite skillful in its con¬ struction. As for the future of music on paper, this is altogether more pre¬ dictable : the dictator is alive and well and living in central Siberia, available for academic appointment. As long as performers and scholars continue to work with the past, the examination of a composer’s documents will be a necessary experience, not to men¬ tion a moving one. As for the composer today, the notion that he learns from the past appears to be temporarily out of fashion. The Romantic genius saw the past as irrelevant in the light of a divine blessing, and so the composer of today sees it as subverting his originality, no longer relevant. The fact of the matter of course is that composers of any age develop their craft, a skill in the handling of their materials. In the learning experience, musical documents will maintain their importance. They are the giants, in the medieval metaphor of Bernard of Chartres, from whose shoulders the dwarfs of succeeding generations will be able to see more, and more distant things. It is more than a cunning trick of a parsimonious librarian to suggest that our repertoire be shifted, quietly and en bloc , to his watchful custody. The care and feeding, and to a degree even the protection of the giants (sensitive as they are, and susceptible to disease, despite their great strength) should belong to specialists, and not the general public. Rather than become infatuated with the giants, or throw stones at them, our society will be far happier helping our midget composers climb onto the giants’ shoulders-— perhaps watching a few of them become giants in their own right. Musicians learn first to read music, then to recognize the danger of playing the notes and missing the musical experience. As his¬ torians we examine our notation and come to appreciate what it has enabled us to have in our music; we should then look to its larger function, as both a preserver of and a stimulant to music itself. VICTOR BERGER: SOCIALIST CONGRESSMAN Frederick I. Olson Between 1910 and 1929 Victor Berger had not one but three con¬ gressional careers.1 A single term from 1911 to 1913 coincided with, indeed was a major element in, the high tide of the Socialist party movement, both in Milwaukee and in the nation. His second career included no service in the House of Representatives, simply two election victories without seating in 1918 and 1919, and it sig¬ nified his constituents' distaste for the first World War and their defiance of the federal government and the Wilson administration. Berger’s triumphal return to Congress after his election in 1922, and his reelection in 1924 and 1926, were personal victories, devoid of the Socialist party import of 1910 and the anti-Wilson political symbolism of the 1918 and 1919 triumphs.2 His defeat for a fourth consecutive term in 1928 removed the greatest prop for his pride and the major recompense for his party’s decline. Nearing 70 and deprived of the regular income from his congressional salary, he now prepared to sell his majority stock interest in the Milwaukee Socialist daily which he had dominated since its founding in 1911, as a further step in the reorientation of his personal life. His party and his movement shattered, his Congressional seat gone, his news¬ paper about to pass under the control of others, Berger may have lacked that will to live which could overcome the injuries he sus¬ tained in a streetcar accident in the summer of 1929. On August 7, 1929 he died. Nonconformity in politics came early to Berger. Born and edu¬ cated in the decaying Austro-Hungarian empire, he migrated with his family to America in his late teens. He settled down in Mil¬ waukee in the 1880’s, amidst social and political turbulence which exposed him to emerging labor politics and to single tax, anarchist, and socialist solutions to the social problem.3 An omnivorous 1 This paper was read at the annual meeting of the Organization of American His¬ torians in Dallas, April 19, 1968. 2 For a somewhat different analysis of Berger as congressman, see Sally M. Miller, “A Socialist Represents Milwaukee,” Historical Messenger (Milwaukee County His¬ torical Society), 22:132-138 (December 1966). 3 Edward J. Muzik, Victor L. Berger, A Biography, unpublished Ph.D. dissertation, Northwestern University, 1960; and for shorter sketches of Berger: Max and Edna Albers Lerner, “Victor L. Berger,” Dictionary of American Biography , XXI (N.Y., 1944), 72-75, and “Victor Louis Berger” in Dwight L. Agnew et al., eds., Dictionary of Wisconsin Biography (Madison, Wis., 1960), 33-34. 27 28 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 reader, he built a large personal library on social issues. He loved disputation and found in German Milwaukee of the 1880’s and ’90’s, with its Turn societies and its beer saloons, congenial companions to argue with. Such reading and discussion convinced him that one could accept a comprehensive theory for the solution of social ills.4 Through the 1890’s Berger evolved a workable social philosophy to account for the future in terms of the past. This was Marxian socialism with its major components of the class struggle, economic determinism, social revolution, and wage, profit, and capital con¬ cepts. Like so many other Americans of the age who embraced Marxian socialist doctrine, Berger acquired a point of view, a sys¬ tem of analysis, and stereotyped rhetoric which clung to him for life. But he was soon convinced by socialist revisionism, especially of the Bernstein variety, and he was sufficiently American to realize the inapplicability of pure scientific Socialism to conditions in the United States. Moreover, he seems to have been impressed by the openness of American political institutions, their availability to all, even the immigrant, through easy naturalization, universal man¬ hood suffrage, liberal qualifications for public office holders, and, at least locally, a fluid party system. Berger became above all a con¬ servative or reform socialist politician with a burning desire to win public office for himself and for his fellow socialists.5 6 Even while evolving their socialist theories in informal discus¬ sions during the 1890’s, Berger and his Milwaukee colleagues had participated in local politics through the Populist party. But Berger also sought to convert Eugene V. Debs to socialism and his Ameri¬ can Railway Union into a socialist political movement. The found¬ ing of the Social Democracy of America in July 1897, followed by the chartering of the first branch in Milwaukee, was to Berger a first step toward the very political action which the leadership of the new organization had prohibited. Thus, in order to nominate a candidate for mayor in Milwaukee’s spring 1898 elections, Berger’s branch was obliged to seek special permission from the Social Democracy’s national executive committee. The next decade was critical in the evolution of an effective so¬ cialist movement in the United States and of a powerful Socialist party in Milwaukee. Berger’s approach to the propagation of so¬ cialism was strongly political from the start. He persuaded the 4 Part of Berger’s impressive personal library may be identified in the collections of the University of Wisconsin-Milwaukee Library. Among1 his less scholarly Socialist party associates and among Milwaukeeans generally Berger’s library and his reading habits took on a legendary character. 6 In addition to Muzik’s dissertation, see Roderick Nash, “Victor L. Berger : Making Marx Respectable,’’ Wisconsin Magazine of History } XLVII (Summer 1964), 301—308, and two convenient collections of Berger’s own views: Berger’s Broadsides (Milwaukee, 1912) and Voice and Pen of Victor L. B&>~gerT: Congressional Speeches and Editorials (Milwaukee, 1929). 1970] Olson — Victor Berger: Socialist Congressman 29 Milwaukee Socialists after 1898 to endorse candidates for almost all elective offices in metropolitan Milwaukee and many in Wis¬ consin, until they matched the Republican and Democratic slates in city, school board, county, state, and congressional elections. Socialist candidates won seats in the city council, the county board, and the state legislature as early as 1904; for the remaining quarter century of Berger’s life his Milwaukee party always held a sub¬ stantial number of elective public offices. Under Berger’s leadership the Socialists not only ran candidates for office — they ran them to win. Berger adjusted Marxian theory and party doctrine in order to gain votes, particularly when an election victory was in sight. For electoral expediency he accepted regulation of utilities as a practical, short term alternative to public ownership. And on many other occasions he shrewdly calculated the effect of platform planks and candidate selection on Socialist vote tallies, not excluding the tactical advantages to be gained by an occasional abstention from competing with the two old parties for an office. Indispensable to Berger’s party was its alliance with local trade unions and union leaders, without incorporating such unions into the party structure.6 Berger carried his absorption in Socialist politics two steps fur¬ ther. He adapted to Milwaukee and Wisconsin an essentially European concept of party organization and party discipline, con¬ trary to Republican and Democratic traditions. Berger’s party im¬ posed stringent requirements and procedures on access to member¬ ship. It organized ward and foreign language branches as its basic units and coordinated them through a strong county central com¬ mittee. It eschewed the open direct primary in the very state which popularized that device. Instead it determined party nominees through its own machinery. Candidacies for public office were in¬ tended to be, and to a large degree were, party not personal affairs. In theory this meant that a faithful party member dutifully ac¬ cepted any draft and as faithfully swallowed his personal ambition if someone else were preferred to him. In fact it obviously wasn’t that simple. Reputations, brokerage, personal friendship, and per¬ sonal popularity all played a role in party endorsements. Proven vote getters and popular party figures like Berger, Hoan, Seidel, Gaylord, Ameringer, Minkley, and Heath usually received the elec¬ tion opportunities they wanted. They were nonetheless expected to campaign as Socialist party agents. This meant endorsing the party 6 Marvin Wachman, History of the Social-Democratic Party of Milwaukee 1897-1910 (Urbana, Illinois, 1945) ; Frederick I. Olson, The Milwaukee Socialists, 1897-1941, unpublished Ph.D. dissertation, Harvard University, 1952 ; and Olson, “The Socialist Party and the Union in Milwaukee, 1900-1912,” Wisconsin Magazine of History, 44 (Winter 1960-61), 110-116. 30 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 platform and guaranteeing, if elected, to carry it out, on penalty of expulsion from membership or even involuntary removal from office. Campaigning had to be ticketwide, but the party handled all campaign arrangements and expenses, financing the latter through monthly dues, a checkoff from the salaries of officeholders, party benefit functions, and levies on the treasuries of friendly labor bodies. Never in American history, perhaps, has a political organ¬ ization woven so tight a web over so many members as that which enmeshed the Milwaukee Socialists. It is not surprising that some opportunists joined the party to gain the support of so formidable an organization. And there were always some members who re¬ belled at the required pledges of loyalty and of money, at the ex¬ pulsions and the purges, at the irritating limitations on their political independence, and their number grew when party fortunes waned and party decay appeared. As the genius who conceived and organized this tight structure, Berger was charged with being a boss, at first by his anti-Socialist political opponents, but soon by his enemies within the party.7 Berger’s masterplan for pragmatic political action also included influence in national party affairs in order to protect Milwaukee autonomy from national interference. For years his nearly im¬ pregnable spot on the party’s national executive committee provided Berger the necessary oversight. His vigilance extended to the na¬ tional party’s constitution, its election platforms, even the party name. Berger understood the need for a strong national move¬ ment to parallel Milwaukee’s, but especially after the high tide of party ballots in 1912, he preferred to preserve the party’s show- place in Milwaukee at almost any cost.8 Socialist political strength in New York, Chicago, and Los Angeles was always overshadowed by its major party opponents, however large their vote tallies by Socialist standards. Socialists were relatively more important in Berkeley, California, Haverhill, Massachusetts, Reading, Pennsylvania, Bridgeport, Connecticut, and elsewhere. But Berger’s troops were the best organized and the most successful of all. Among the nation’s large cities Milwaukee alone seemed to justify the “socialist” label.9 The first Socialist victories in 1904 foreshadowed the landslide of 1910 when the party won city council and county board majorities and elected its can¬ didate for mayor. Berger knew that as long as party labels pre¬ vailed in local elections, his Socialists could win pluralities in three way races. Beginning in 1912 Republican and Democratic fusion 7 Olson, Milwaukee Socialists, 55-84. 8 David A. Shannon, The Socialist Party of America , A History (N.Y., 1955), 17, 21-25, 62-63, 258-60. 9 Shannon, Socialist Party, 8-4 2, 188-89. 1970] Olson — Victor Berger: Socialist Congressman 31 and non-partisan election laws virtually eliminated such leverage, yet the Socialists continued to win selected elections.10 If Berger was a boss, he declined to play the traditional role entirely behind the scenes. He ran for public office regularly, a visi¬ ble target for his intraparty opponents and his anti-Socialist ene¬ mies. In the 1910 landslide he finally won election as an alderman- at-large and for a year or so played the major role in Mayor Seidel’s administration. But Berger’s ambition was to be Congressman. He ran as early as 1904 in Wisconsin’s Fifth Congressional District, and beginning in 1910 he never missed a campaign for that seat. Berger emerged in the 1904 balloting as a possible threat to the incumbent Repub¬ lican William H. Stafford by polling over 10,600 votes, or 33.8% of the total, for second place. His opponent, a colorless conserva¬ tive, was a native Milwaukeean, 35 years old, a graduate of Harvard law school and a bachelor, who had won the seat for the first time in 1902. Between that date and Stafford’s last victory in 1930, he or Berger won every Fifth District election. Over that period the seat was vacant for 2 years, Stafford served for 20 years, and Berger for eight. Curiously enough Berger and Stafford faced each other only 9 times in 14 Congressional contests between 1904 and 1928.11 What kind of district could alternate between a colorless con¬ servative Republican and the first Socialist ever elected to Con¬ gress? A major clue is found in the changing relationships among three major parties. While Stafford and Berger remained con¬ stants, their Democratic opposition varied. Moreover, old party fusion succeeded against Berger three times. And once Stafford failed to win the Republican nomination. But the Fifth clearly bore the marks of a swing district, even after Berger and Stafford were gone. From Stafford’s defeat in the Roosevelt landslide of 1932 to the election of Democrat Henry S. Reuss in 1954, three Democrats and two Republicans divided 22 years into seven segments, the longest consisting of three consecutive terms. Reuss had already made the seat safely Democratic before reapportionment reshaped it out of all resemblance to its 1904 character. Altogether, fickle Fifth District voters changed Congressman 14 times in 52 years. Only Stafford, at the very beginning of this period, won as many as four consecutive terms. He also shared with Berger and Staf¬ ford’s immediate successor in New Deal days a run of three con- 10 For contrasting1 views, see Myron L. Anderson, “Milwaukee Election Law That Boomeranged,” Milwaukee Journal , December 20, 1959, and Erich C. Stern, “The Non-Partisan Election Law: Reform or Anti-Socialism?”, Historieal Messenger (Mil¬ waukee County Historical Society), 16:8-11 (September 1960). 11 For election statistics, see the biennial Wisconsin Blue Books (Madison, Wiscon¬ sin, odd years), and biennial reports of the Board of Election Commissioners of the City of Milwaukee (Milwaukee, various dates). 82 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 secutive terms. But seven times the incumbent failed to win even a second consecutive term. Redistricting created a district in 1901 which joined the north half of the city of Milwaukee and the north and west suburban and rural-farm areas of Milwaukee County with villages and wealthy farm sections of Waukesha County to the West, the latter account¬ ing for slightly more than a fifth of the District's voters. The com¬ bination of rural-farm- village with big city was a major cause of party alternations. Across the near north side of the city of Mil¬ waukee stretched a band of German settlement which pro- , vided the core of the Socialist strength. The Germans as a whole were frugal, law-abiding members of the working class or lower middle class. Many were Socialist party members; others were sympathizers who often voted Socialist. In the suburbs, as well as in the rural and farm areas of Milwaukee and Waukesha Counties, lived business, professional, and farm folks, typically middle class and traditionally Republican. Scattered in both counties, but heavier in the city of Milwaukee, were traditional Democrats who were predominantly Roman Catholic and Irish or German. To win, the Socialists and the Republicans were obliged to woo not alone the independent voter, but the normally Democratic voter as well. The redistricting of 1901 put two Congressional seats within the grasp of city of Milwaukee voters for the first time. Ten years later the detachment of Waukesha County made the Fifth District even more urban in character; and nearly half a century later, with all of Milwaukee County urban, the central city portion was divorced from the north suburban, and the latter was joined once again to the contiguous suburbanizing portions of Waukesha County. Berger’s election in 1910 gave the Socialist party its first House member, and Milwaukee its first labor-oriented congressman since 1886. The only other Socialist party member ever to win a House seat was Meyer London of New York; his three terms between 1915 and 1923 neatly complement Berger’s subsequent victories to suggest a unique character for all Socialist congressional triumphs. One of Berger’s successors in the Fifth after the second World War was ex-Socialist Andrew J. Biemiller, who had passed through La- Follette Progresslvism into the Democratic party.12 The propaganda value of Berger’s election for the Socialist party all over the nation can hardly be exaggerated. Electing So¬ cialists to common councils and state legislatures was clearly not enough, for only Congress could legislate the major components of socialist doctrine. The Socialists had to demonstrate their capacity to capture Congressional seats if they were to move the nation 12 Olson, Milwaukee Socialists, 197-99, 271-72; Milwaukee Journal, September 30 and October 23, 1955 ; Shannon, Socialist Party, 9, 11-12, 158. 1970] Olson — Victor Berger: Socialist Congressman 33 towards socialism, and up to 1910 their propaganda about the relentless tide of Socialist votes notoriously lacked confirmation on this point. Berger thoroughly enjoyed the personal glory of be¬ coming the first Socialist congressman, but he had to picture him¬ self as a foretaste of growing numbers of Socialists, not as an ef¬ fective instrument itself. Quips about caucusing in a telephone booth merely emphasized his ineffectiveness, for as a minority of one he could be readily tolerated. Patronage and his other per¬ quisites as a Congressman appealed strongly to Berger, but Mil¬ waukee Socialists had gained far more tangible party benefits from their other local victories. In the House Berger was no social revolutionary. He claimed that he represented not alone— or not primarily — the Fifth District, but the working class and all the Socialist voters of the nation. He often acted prudently by doing errands for any of his constituents, even the anti-Socialist or non-Socialist as well as his known sup¬ porters. As the sole Socialist Representative he tried to be true to his apocalyptic role. For their propaganda value he sponsored futile Socialist measures such as old age pensions and national ownership of railroads and communications. His speeches and news handouts were designed not to win votes in Congress but to present Socialist positions and gain Socialist converts outside the halls of Congress. His occasional deviations from the party line to please his Milwaukee constituents brought prompt censure from the party faithful while failing to secure his reelection. His most effective action, in calling for an investigation of the textile workers strike at Lawrence, Massachusetts, depended, as he well knew, on support from and tolerance by colleagues in Congress whom he felt obliged to castigate at every opportunity. A gregarious person who had enjoyed social acceptance by his ideological enemies in Milwaukee, Berger could not now isolate himself from the fellowship of the House of Representatives. Most evident was the gratification of his desire for personal acceptance by fellow Congressmen. That he never recovered from the fascination of his first Congressional ex¬ perience is witnessed by his biennial efforts to regain his seat for the rest of his life. But the crudest dilemma in Berger's congressional service arose not from compromise of his socialism but from diversion of his energies. His central role in the Milwaukee party organization made his absence in Washington and his attention to Congressional duties costly to the Milwaukee movement. The persistent financial strain of ordinary party activities now extended to building a labor temple and founding a daily newspaper, the latter a part of the publishing complex from which Berger drew a salary. The long planned So¬ cialist daily Milwaukee Leader appeared December 7, 1911, coinci- 34 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 dental with the opening of the 62nd Congress. While Berger espoused Socialist doctrine on the floor of the House in the Spring of 1912, the Leader limped along, powerless to prevent a Socialist administration wiped out by a fusion ticket, and the Milwaukee rout in the municipal elections. From afar Berger saw the Seidel Socialists rent by dissension in their leader’s absence.13 Nor could Berger return for the fall campaign in time to develop a winning tactic against Stafford’s coalition with the Democrats. Berger’s plurality in 1910 had been a slim 350 or 1% of 35,224 ballots. The resulting anti-socialist fusion, although incomplete, reduced Berger’s vote but slightly from 38% to 35 or 36%, yet provided Stafford with margins of 1,908 in 1912, 3,946 in 1914, and 3,649 in 1916. After his 1912 defeat Berger adopted the correct party stance, lamenting that the millions of Americans who had voted Socialist were deprived of a voice in Congress. Between Berger’s defeat in 1912 and his khaki election victory of 1918 the Milwaukee Socialists entered a new era. The momentum of local and national election gains and party membership growth was lost between 1910 and 1912. Within Milwaukee and Wisconsin politics the Socialists settled down to a respected but limited role. When Dan Hoan was reelected city attorney in 1914 and recaptured the mayor’s office for the Socialists two years later, no one could foresee how long he would hold it or how remotely Socialist his administration would become before his defeat in 1940. 14 Some Socialists including Berger did foresee the catastrophic potential of the first World War as early as 1914. But America’s Socialists were obliged to reconcile their doctrinaire war beliefs to the votes of their European brethren on the war credits. Initially Berger maintained in his Mihvaukee Leader a peace posture which was distinguishable from pro-Germanism. Stafford rather than Berger embraced an opportunistic pro-German position in 1914 which helped him win easy reelection. Gradually, however, Berger’s reaction to the war took on a more pro-German, anti-French and anti-British tone, not surprising in one who was born and educated in Central Europe. It became more difficult for Berger to apply simple Socialist tests to the events preceding the entry of the United States into the war, and he seems to have convinced himself that the defeat of Germany must be avoided because it would destroy the most promising socialist party in Europe and thus in turn weaken the American movement. Fortuitously, Berger’s cultural preference for Germany coincided with the prejudices of his Mil- w Miller, loc. cit. ; Olson, Milwaukee Socialists, 247-54; Olson, “Milwaukee’s First Socialist Administration, 1910-1912 : A Political Evaluation,’’ Mid- America, 43 (July 1961), 197-207. u Olson, Milwaukee Socialists, 271-72, 310-11. 1970] Olson — Victor Berger: Socialist Congressman 85 waukee constituents and reinforced his concern for the fate of German socialism. Stafford could outbid Berger for the critical German votes in the Fifth District as long as President Wilson did not invoke federal power against Berger. But when Wilson's Postmaster General re¬ voked the Leader's second class mailing privilege on October 3, 1917, and his federal attorney brought indictments against Berger and four alleged Socialist co-conspirators under the Espionage Act on March 9, 1918, Berger was cast as a martyr to an all-powerful government which could not tolerate dissent. Berger, who had con¬ curred in his party's condemnation of the American declaration of war, in his newspaper and by other acts and utterances had crit¬ icized many government policies and practices in prosecution of the war. But he had every reason to believe that he had remained within the letter of the law. That his opposition to the Wilson ad¬ ministration, or rather the latter's prolonged persecution of him for his beliefs, had election appeal was demonstrated in April 1918 when he polled 110,478 votes, or over 25%, in a senatorial election against a pro-war Republican and a Wilson Democrat. Most sig¬ nificant, concentration of Berger's votes gave him a plurality in the Socialist stronghold of Milwaukee. In the November 1918 congressional race Berger won handily despite the pending federal indictment. The Democrats, hoping to convert Wilson’s call for a Democratic Congress into their first victory in the Fifth District, defected from their fusion agreement to make their first serious challenge since 1908. But they merely gained second place while Berger drew enough traditional Republi¬ can and presumably German votes from Stafford to produce the largest plurality of his six victories, 5,470 votes.15 What Berger had regained at the polls, his erstwhile colleagues in the House now withheld. Between his election and the convening of Congress his indictment was converted in federal district court in Chicago into a conviction, with a sentence of 20 years from Judge Kenesaw Mountain Landis. While out on bail Berger fought to be seated, but the House refused him on November 10, 1919, by a 311 to 1 vote. Congressional reasoning was that Berger had given aid and comfort to the nation's enemies and thus invoked the third section of the Fourteenth Amendment, which required denial of his seat. Berger promptly stood for the Socialists in the December 1919 bye-election resulting from the vacancy, and as promptly was re¬ elected by nearly 5,000 votes over his fusion opponent with the attractively Teutonic name of Bodenstab. Again Congress applied the Fourteenth Amendment to keep Berger from serving. 16 Ibid., 339-40, 355-56, 374-84. 36 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 By 1920 Berger could not so readily capitalize on anti- Wilson sentiment. For once fusion was so thorough it not merely weakened the Democratic effort as it had in 1912, 1914, and 1916, but it actually eliminated all non-Socialist candidacies but Stafford’s. Even Berger was not safe from Republican exploitation of anti¬ war and anti- Wilson sentiment, and Stafford won by a 6,773 ma¬ jority, the largest margin of any of their nine contests.10 Yet in defeat Berger was preparing for ultimate vindication. First of all, he appealed his conviction to the United States Supreme Court, which set aside Landis’ sentence on January 31, 1921, and the Harding administration dropped the case without further effort. Next, in the favorable Wisconsin election climate of 1922, wherein a sympathetic Senator LaFollette also won triumphant reelection, Berger defeated Stafford by a 3,771 majority, or 6.7 %, in another two way race, a most dramatic reversal of 1920. Without dissent he was now seated. And in the next two elections he survived the Coolidge landslide and the almost total eclipse of his own party everywhere, including serious setbacks in Milwaukee. But a presi¬ dential contest between Hoover and Smith in 1928 was too formida¬ ble for him to deal with. Hoover’s candidacy capitalized on the fragile prosperity of the era, Smith’s on Milwaukee’s resentment over Prohibition. For the first time in 10 years the Democratic nominee for Congress polled over 20% of the vote, and Berger lost to Stafford by 709. It was his last race.17 As the first Socialist Congressman in 1911, Berger had sym¬ bolized his party’s potential, and both his sense of party responsi¬ bility and his vision of the future had dictated the ideological role he must play. But by 1923 he had survived the terrible buffetings of the war period — persecution of his newspaper, prosecution of himself as a subversive, and double denial of a House seat — while his party, swollen by the transient growth of the war era, split wide open nationally over the Russian revolution and then de¬ clined sharply. What Berger salvaged of Milwaukee Socialism drew no nourishment from a national movement that struggled simply to maintain its name and headquarters. The hopefulness of its 1910 victories had given way to despair in the 1920’s. The Berger brand of gradualist socialism, which had seemed so promising under Mayor Seidel, had deteriorated into a housekeeping version called sewer socialism under Mayor Hoan. It took great faith in a socialist 16 Ibid., 384-88; Edward J. Muzik, “Victor L. Berger and the Red Scare,” Wisconsin Magazine of History, XLVII (Summer 1964), 309—18; Hearings Before the Special Committee Appointed under the Authority of House Resolution No. 6 Concerning the Right of Victor L. Berger to be Sworn in as a Member of the Sixty— Sixth Congress (2 v., Washington, 1919); Zechariah Chafee, Jr., Freedom of Speech (N.Y., 1920), 310-33. 17 Olson, Milwaukee Socialists, 388—90, 441—42, 448-50. 1970] Olson — Victor Berger : Socialist Congressman B7 future even to maintain a party organization from election to elec¬ tion. Ideological disputation, long the stock in trade of Socialists, gave way to factionalism, personality differences, and a scramble for personal preferment. Thus Congressman Berger no longer claimed a socialist, only a Milwaukee, constituency. No one seriously expected a Socialist revival, even in Milwaukee. All one hoped for was to sustain the present officeholders and quiet the dissidents who wondered aloud what had happened to the fiery zeal of 1900. The respect Berger had once sought for his party and his movement he now craved for himself. He was not so much a lone Socialist in Congress ; London had been that too, for three terms. He was rather the vindicated victim of a war time hysteria. The U. S. Supreme Court, Fifth Dis¬ trict voters, and the House itself had in turn confirmed this. While he retained his love for the stereotypes of non-revolutionary so¬ cialist programs and socialist rhetoric, he spoke in Congress most commonly about international affairs and the issues and conse¬ quences of the war. Set apart from most Congressmen by his Central European upbringing and his informed interest in the larger movements of Europe and the world, he addressed himself to the deepening American disillusion with the war and the war settlement and to relations among the great powers of Europe. Here and there an underlying socialist analysis shone through, but generally the viewpoint was more personal and his major concern over civil liberties and Prohibition reflected election needs. Today his remarks reflect the shallowness of contemporary comment ; then they drew attention to his education, his reading, and his essentially foreign cast of mind.18 If Berger continued to stand apart from his House colleagues as in 1911, it was due less to his Socialist label than to his personal independence. Yet he enjoyed far more than in 1911 friendships with other House members. The respect of some who differed from him most like John Nance Garner is reflected in their farewell re¬ marks, while the friendly Fiorello LaGuardia saw Berger as “a pioneer, popularizing ideas of political and social reform long before they are accepted by the many, and while they are still frowned upon by the majority and denounced by political leaders.”19 Berger’s election as the first Socialist in Congress had been a natural by-product of his obsession with local Socialist political activity. He had fashioned in Milwaukee the most thoroughgoing and durable political organization the American Socialists ever saw, and thus won for himself not only a Congressional seat but 18 Ibid., 442-48. Berger, Voice and Pen, passim. 19 Congressional Record, 70 Cong., 2 sess. LXX (Mar. 4, 1929), 5275. 38 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 a place in the Socialist pantheon along with Debs, Hillquit, London, Hoan, and Thomas. But before his death in 1929, the socialist con¬ viction and optimism which had sustained his early party activity had given way to personal advancement and despair. “The Socialist party of Milwaukee/' he had told his closest friends, “will not survive my death by six months." He was wrong. He had already outlived it.20 20 Olson, Milwaukee Socialists, 451-52, 573—74. WHEN SEDITION LAWS WERE ENFORCED: WISCONSIN IN WORLD WAR I John D. Stevens Today, with television and Broadway shows satirizing public officials, with militants calling for armed rebellion, with young men chanting, “Hell no, we won’t go,” it is important to remember that it was not always so. Certainly it was not so during World War I, when men went to prison for chance remarks in bars, rooming houses and on street corners, when the Post Office hounded foreign language papers out of business, when wearing an Industrial Workers of the World pin made you, automatically, a disloyalist.1 Such “crimes” were prose¬ cuted under local ordinances, state sedition laws, and primarily under federal statutes. This paper attempts to examine the en¬ forcement of these federal laws in Wisconsin, which had the na¬ tion’s highest proportion of German descendants and one of the nation’s most active and successful socialist parties.2 Although President Wilson had been urging an internal security law since December, 1915, Congress could not agree on one, so the United States entered the war with only the Conspiracies Act of 18613 and the Treason Act of 1862.4 Neither reached individual utterances. The federal security package included five major laws which cur¬ tailed expression: Threats Against the President Act,,5 Selective Service Act,6 Espionage Act,7 Trading-with-the-Enemy Act,8 and Sabotage Act.9 These were supplemented by many presidential 1 See e.g. Chafee, Zechariah Jr., Free Speech in the United States (1941) ; Johnson, Donald, The < Challenge to American Freedoms (1963) ; Scheiber, Harry N., The Wilson Administration and Civil Liberties (1960). 2 See Petersen and Fite, Opponents of War 1917-1918 (1957) ; Preston, William Jr., Aliens and Dissenters (1963), and Maxwell, Robert S., Emanuel L. Philipp, Wisconsin Stalwart (1959), discuss Wisconsin’s reputation for disloyalty. 3 12 U.S. Statutes 284 4 12 U. S. Statutes 589. This law was used to convict some anarchists; cf., Goldman v. U.S. , 245 U.S. 474 (1918) 6 39 U.S. Statutes 919. 6 40 U.S. Statutes 76. 7 40 U. S. Statutes 217, with amendment, 40 U.S. Statutes 553. The 1918 amendment sometimes is referred to as the Sedition Law. 8 40 U.S. Statutes 425. »40 U.S. Statutes 533. 39 40 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 orders and directives.10 These five laws will be discussed in chrono¬ logical order, with particular emphasis on their enforcement and effects in Wisconsin.11 Threats Act Spurred by the growing menace of war, the House of Representa¬ tives passed the act in June, 1916; the Senate concurred in Febru¬ ary, and the President signed it on Valentine’s Day, 1917. There had been suggestions for such a law since the 1901 assassination of President McKinley. The brief law provided up to five years and $1,000 fine for mail¬ ing any “threat to take the life or to inflict bodily harm upon the President of the United States.” Federal courts interpreted “threats” quite broadly, to mean something akin to the fifteenth- century English high treason law which made it a crime to imagine the death of the sovereign. By June 1918, 85 persons had been con¬ victed under the law, and in Wisconsin nearly twice that many had been indicted.12 Two men were convicted and five others pleaded guilty during the war. Penalties ranged from $100 to 18 months in prison. Without a broad interpretation of “threats,” it seems un¬ likely any of them would have been convicted.13 A Racine policeman was the first person indicted under the law in Wisconsin. Allegedly he told another man that Wilson would be shot within 30 days and that if no one else did it, he would do it himself. During his trial in May, 1918, he denied making the last part of the statement and said the first was based on astrology. He said he was discussing astrology with another man when a third person overheard the conversation and reported it to federal offi¬ cials. The jury was not impressed, and he was sentenced to 18 months in Leavenworth Prison. Both indictments brought in the Eastern District were against Shawano County men, and both were joint actions under the Threats and Espionage Acts. One man had allegedly said, “If I was drafted I would take a straight shot for President Wilson’s house and would do away with him if I got the chance.” His case was dismissed in March, 1919, without coming to trial. The other was indicted and tried for saying in a private home in the presence of 10E.G., Alien Enemies Proclamation (April 6, 1917); Federal Employees Loyalty Order (April 7, 1917) ; Order Creating- Committee on Public Information (April 14, 1917) ; Cable and Telegraph Order (April 28, 1917) ; Order Establishing Board of Censorship (October 12, 1917). 11 Unless otherwise specified, information on Wisconsin cases was taken from federal court records in the Federal Records Depository at Chicago ; in some cases, details were added by contemporary newspapers. 12 Report of the Attorney General (1918), p. 56. 13 Among decisions holding that the threat need not be communicated to the President were U.S. v. Strickrath , 242 F. 151 (1917) ; U.S. v. Jasick, 252 F. 931 (1918) ; U.S. v. Metzdorf, 252 F. 933 (1918) ; U.S. v. Stoho , 251 F. 689 (1918). 1970] Stevens — When Sedition Laws Were Enforced 41 three other persons, “The President is the one that caused this war. He ought to be killed and if I had the chance I would kill him in a minute.” The jury refused to convict him, perhaps because the trial came a month after the Armistice. It should be noted that the only Threats Act indictment which resulted in a conviction in the Eastern District was also the first one brought there. On the other hand, in the Western District the first man indicted entered a plea of guilty and got off with the lightest penalty of all, a $100 fine. In October, 1917, he allegedly said: We ought to clean out the White House. Wilson wants to be shot before they shoot the Kaiser. If Wilson is not shot before he gets out of office, he will be after he gets out of office if he doesn’t get out of the country. Indictments were returned in March, 1918, against three men and one woman. The case against an Ashland man who was also indicted under the Espionage Act for saying “I’d kill the President like a dog” never came to trial. A German-American from Mara¬ thon County was sentenced to six months in the Milwaukee County House of Correction for saying, “I am a socialist. President Wilson is a son of a bitch and I would hang him if I had my way.” A young Eau Claire County farmer was indicted for saying that if he had an airplane he would “get that damned Wilson” and that if he were drafted he “would like to kill that goddamned Wilson.” He pleaded guilty but told the judge he had been drunk. Unmoved, Judge Sanborn sent him to Leavenworth for a year and a day. The wife of a butcher at Milladore in Wood County, after hearing of the sinking of the Tuscania troop ship, which was carrying a large contingent of Wisconsin troops, allegedly threatened to put a bullet in the head of President Wilson. She was convicted in a one-day trial in August, 1918, and sentenced to six months in the Eau Claire County Jail or a fine of $500. She chose the latter. A Dunn County man pleaded guilty in March, 1918, to saying, “The President ought to have been killed long ago, and if some¬ body does not do it, I will.” He served 30 days in the Eau Claire County Jail for his indiscretion. The final case was not related to the war at all and indicates how broadly the statute was being interpreted by federal judges. A Prairie du Chien man, three weeks after the Armistice, allegedly said, “I will shoot Wilson, the son of a bitch, if the country goes dry July 1.” He was indicted May 12, 1919, and two weeks later entered a guilty plea. He was sentenced to 30 days in the Dane County Jail. 42 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 By most standards, none of these 10 persons posed much of a threat to President Wilson. For that reason, it might be instructive to review the only statement recorded about the purpose of the law by the author himself. During the congressional debate, Representa¬ tive Webb said, “The man who makes the threat is not himself very dangerous, but he is liable to put devilment in the mind of some poor fellow who does try to harm him (the President) .”14 Although no one can tell how much “devilment” the quoted re¬ marks put in the minds of hearers — especially since the indict¬ ments did not indicate the context of most of the remarks — it seems unlikely that the hearers in Wisconsin posed much threat to the President. The same could be said about other persons reported in the newspapers as arrested under the Threats Act but who were not actually indicted. From 1921 until World War II, the law was a virtual dead letter. Three cases under the Threats Act were appealed during the war and two have been appealed since. The Threats Act re¬ mains on the statute books, in peace as well as war time, and in 1962 was broadened to protect the Vice-President or other per¬ son next in line for the presidency, as well as the President-elect and the Vice-President-elect. It was 1965, in the aftermath of the Kennedy assassination, before Congress passed a law which pro¬ tected the life of a President.15 Selective Service Act The conscription law of 1917 made it a crime to discourage men from registering or serving in the military services. There were widespread fears of draft riots such as those of the Civil War, but they were not realized. In spite of Governor Philipp’s later recollection that there was no opposition in the state to the draft “save now and then an in¬ dividual,” the dockets of the two federal courts in the state con¬ tain the names of hundreds of men indicted for draft violations, most of which were quietly dropped when the men “volunteered” for immediate army duty. Attorney General Gregory assured a congressman in July, 1918, that this was the usual procedure for men failing to register. He said that through June 8, there had been 8,802 actions under the draft laws through the nation; of these, 4,064 dealt with failure to register.16 From among the many indictments under the draft act, five Wisconsin cases have been singled out which resemble closely in- u Congressional Record , 65 Cong-., 2d Session, 9377. is The Threats Act now is 18 U.S. Code Annotated , Section 871. 16 Philipp, "Wisconsin’s War Activities,” Wisconsin Blue Book, (1919), pp. 301-303; Congressional Record, 65 Cong., 2d Session, 528. 1970] Stevens — When Sedition Laws Were Enforced 43 dictments brought under the Espionage Act for expression. One, in fact, was a joint draft-espionage action. This involved nine members of a “Holy Roller” fundamentalist religious colony in Barron County who were indicted in March, 1918. When they appeared in court June 3, they refused to plead, but instead made anti-war religious speeches. The judge ordered a not guilty plea entered in their behalf and placed them in jail when they refused to post bond. The men were indicted on 13 counts based on 16 separate speeches, personal letters, personal advice, and articles in their paper. After a three-day trial which ended July 20, all were found guilty. The leader was sentenced to 15 months at Leavenworth, and three other members were sent there for a year and a day. The other six were given terms of three to six months in county workhouses, although four of the six were permitted the option of paying fines of $250 to $500. In another case, in June, 1918, three men were indicted jointly under the draft and espionage measure, but the indictment has been lost. A newspaper reported that their arrest at Ashland was for saying Wilson was responsible for the war and for the sinking of the Lusitania. A wealthy Grant County farmer was convicted by a jury in 1918 and sentenced to a year and a day at Leavenworth plus a fine of $1,000. He was indicted on five counts and convicted on two, including the following statement made to draft-age men : This is a rich man’s war and we would not have this war if it had not been for the rich girls in the United States marrying English lords. We had no business to start this war. The issue that Wilson was elected on was not to start war. They loaded supplies for the Allies on boats and hired a few Americans and put them on those boats and they were killed and this started the war. We had no cause to be in this war. The Germans killed no Belgian citizens and there are no orphans in Belgium or France that were caused by the war. The Germans have done no worse than Americans have done to Germans. We have no business in this war. We went into it to protect the money that was loaned to the Allies. The money interests hired a few Ameri¬ cans to ride on those ships so that if they were killed we could get into the war. The jury, according to a newspaper, was out for only a half hour in convicting the 58-year-old father of seven. This was the only case in which the judge’s charge to the jury was preserved in the archives. It was given by special judge Evan A. Evans. In his charge, Evans denied the defense attorney’s con- 44 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 tention that such words had to be spoken to a person already in the service or who was about to be called ; it was enough that they be spoken and there be some reasonable chance that they might some¬ how be conveyed to a soldier or potential soldier. The fourth draft case involving expression was that of a Wood County man whose remarks were made in Polish in the summer of 1918 to some prospective draftees. After a one-day trial he was sentenced to six months in the county workhouse and fined $500 for saying: The kaiser will fare along better than ever because he has made peace terms with Ukrania, but the kaiser will pay no attention to the papers that were signed. It is coming to it that President Wilson will have to crawl on his knees to kiss the kaiser’s boots. The submarines will work havoc with American boats. They have sunk nine boats and of a total of 1,000 persons on those boats, only 40 were saved. It is difficult to imagine where the Wood County man got his “facts,” but the jury decided that his remarks could frighten some rural Wisconsin lads enough that they might not fulfill their mili¬ tary duties. The most celebrated draft case in the state had nothing to do with free expression directly but much to do with it indirectly. This was the action against Congressman John M. Nelson of Madi¬ son and his draft-age son, Byron. To understand the free expression implications of the case, one needs to bear in mind that Nelson had voted against most war measures, including the declaration of war. He had been in Con¬ gress since 1906 and was clearly identified with the La Follette wing of the Republican Party. (In 1924, he managed La Follette’s presidential campaign.) He had alienated many Republican “regu¬ lars” in Congress by his role in the 1908-1910 fight to prune the powers of Speaker Joe Cannon. When, in August, 1917, the federal district attorney announced that he would seek indictment of the congressman’s son for not registering for the draft, Nelson realized that he was the real tar¬ get. Nelson knew his way around Capitol Hill and had many friends. On September 4, he sent a long letter to Attorney General Gregory refuting one by one the charges against his son. Byron was charged with not registering on “Duty Day”; however, those residing in foreign countries were specifically excluded from the registration requirement, and Byron had been managing a family farm in Alberta since May 5. This was 15 days before the draft act was enacted, 28 days before the President’s proclamation estab- 1970] Stevens — When Sedition Laivs W eve Enforced 45 lishing the registration date, and a month and two days before the registration day itself. Nelson said that at the time he had asked his son to go north he had no way of knowing what the terms of the act would be. His son had worked on the farm during sum¬ mers while attending the University of Wisconsin. Nelson even sent Gregory a published “Roll of Honor” which showed that his son ranked fourth among students in donating their time during the spring to work on the university farm. Nelson further assured Gregory that in May he had received assurances from the Provost Marshal General that persons living abroad did not have to register for the draft. He had reaffirmed this in person with the Provost Marshal on September 1 after reading of the district attorney’s intention to prosecute. Nelson cashed in some of his other political debts. For example, he convinced Speaker Champ Clark to write a confidential note to the Postmaster General on his behalf.17 In October, Byron pleaded not guilty to the draft evasion charge in the federal court in Madison. In late November, the Congress¬ man was indicted for conspiracy to avoid the draft and his son for failure to register. Both entered pleas of not guilty and were re¬ leased on bond. One month later, on January 3, the federal judge quashed both indictments. Although the irate district attorney told the press that he would appeal the judge’s decision, he did not. That was the end of the case, at least in the courts. Although the Nelson case was over in the courts, it was “retried” at the polls in the spring primary, and Nelson lost; however, he won back his seat in 1920 and held it until defeated by a gerry¬ mander in 1932. Espionage Act If, as John Roche suggested, World War I was a “black mass celebrated by the elected leaders of the American nation,”18 then surely the Espionage Act must be considered its litany. Only two provisions of the lengthy law affected free expression. Section 3 of Title I made it a crime to interfere with the military or recruiting services, while Title XII made it illegal to mail matter which violated other sections of the law. Strictly construed, these would have had little effect on expression. That they were not so construed was shown by the number of prosecutions. Chafee wrote that 877 persons were convicted under Section 3 and more than 100 publications banned from the mails under Title XII. Chafee based his figure on annual reports of the Attorney General, which for the two federal courts in Wisconsin 17 Nelson to Gregory, May 4, 1917, and Clark to Nelson, December 14, 1917 in John M. Nelson Papers, State Historical Society of Wisconsin, Madison, Box 1. 18 Roche, John P., Quest for the Dream 49 (1963). 46 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 were too low. Other estimates ran higher, but it seems likely that there were at least 2,000 actions and 1,000 convictions under Sec¬ tion 3. Virtually all were for expression.19 Punishments under the law were harsh. At least 35 persons went to prison for the maximum 20 years and another 58 for 10 to 15 years; others received shorter sentences and fines. Still, the man who directed the law’s enforcement admitted that there was not a single proven case of sabotage in the nation after the declaration of war, and Charles and Mary Beard concluded the law did not catch one bonafide spy or saboteur.20 Twelve days after the United States declared war, Senator Lee Overman of North Carolina introduced an omnibus measure which covered such diverse security matters as the embargo, conveying secrets to foreign agents, passports, impounding enemy vessels and counterfeiting the government seal. He described it as substantially the same measure which the Senate passed during the previous session but which died in the House. Both in the press and on the floor, most of the controversy cen¬ tered on a section which provided five years in prison and a $10,000 fine for publishing information declared by the President to be useful or even possibly useful to an enemy. There was much less attention to Section 3 of Title I which, as enacted in 1917, provided 20 years in prison and $10,000 fines for anyone who while the United States was at war should willfully make or convey false re¬ ports or false statements with intent to interfere with the opera¬ tion or success of the military or naval forces of the United States or to promote the success of its enemies: and whoever, when the United States is at war, shall willfully cause or attempt to cause insubordination, disloyalty, mutiny, or refusal of duty in the mili¬ tary or naval forces of the United States, or shall willfully obstruct the recruitment or enlistment service of the United States. Title XII allowed the Postmaster General to refuse to convey any letter or printed matter “advocating or urging treason, insurrec¬ tion, or forcible resistance to any law of the United States.” Maxi¬ mum penalties were set at $5,000 and five years, in prison. In introducing the measure, Senator Overman assured his col¬ leagues the measure in no way limited freedom of the press or of individuals to comment on governmental policies. He assured them that the courts could be trusted to interpret the law in a reasonable manner.21 For three days, the Senate debated the press censorship section, but in the end rejected it. The House accepted the mail section with 10 Chafee, op. cit., supra note 1, at p. 52. 20 O’Brian, John L., National Security and Individual Freedom, 49-50 (1955) ; Beard, Charles and Mary, The Rise of American Civilization, Volume II, 644 (1927) 21 Congressional Record, 65 Cong-., 1st Session, 778-781. 1970] Stevens — When Sedition Laws Were Enforced 47 a few minor wording changes over the warnings of its only socialist member that Title XII would be the death knell for minority opin¬ ion press in this country. The bill passed, 260-106. The Senate then debated the bill for two days in executive session and for three more in public. On May 12, the Senate defeated the censorship section and later at attempt to reinstate it. The upper chamber prohibited postal inspectors from opening sealed letter and adopted the bill, 77-6. Wilson signed it June 15. Once shorn of its press censorship section, the Espionage Act was greeted by newspapers either with praise or with indifference.22 Early in 1918 the Attorney General requested an amendment to Section 3 to protect government bonds from criticism. The Senate Judiciary Committee broadened the amendment to cover disparag¬ ing remarks about the flag, the military forces, the Constitution or form of government. The amendment, sometimes called the Sedition Act of 1918, provided 20-year prison terms for remarks which in¬ terfered, even remotely, with the war effort or aided the enemy. The sponsoring committee admitted the amendment was quite superfluous since most district judges had interpreted the 1917 law to cover such offenses. During the seven days of debate, at least five senators said that such an amendment would help quell the mob spirit in the land. Gregory used the same argument in his 1918 report.23 The Senate strengthened Title XII, permitting the Postmaster General to refuse even to pick up or deliver mail during the war at addresses using the mails in violation of the act, thus crimping socialist and radical defense committee collections. On April 9, the Senate refused to incorporate even the protections of the 1798 sedition law, excluding truthful remarks made with good motives.124 Of the 92 indictments in Wisconsin under the Espionage Act — several in conjunction with the Threats, Draft or Trading acts — all but two involved expression. Since most judges and juries used ill tendency as their standard from the beginning, the 1918 amend¬ ment made little difference. Analysis of the preserved indictments shows that the offensive remarks fell into certain logical categories, shown in Table 1. A few representative Wisconsin cases will be discussed for each category. Thirty-six indictments included remarks praising Germany or the Kaiser or expressing the belief or hope that Germany would 22 Congressional Record, 65 Cong-., 1st Session, 847, 871-887, 1590-1596, 1698-1701, 1717-1720, 1750-1780, 1807-1841, 2055-2072, 2087-2113, 2166-2196, 2241-2270. 23 Congressional Record, 65 Cong-., 1st Session, 3002-3004, 4559, 4562, 4633, 4637, 4645-4646, 4710, 4764-4771, 4824. 24 Congressional Record, 65 Cong’., 1st Session, 4562-4563, 4637, 4784, 4826, 4835- 4839, 4895-4898, 5541. Procedural safeguards did little to protect defendants under the 1798 law, according to Smith, James M., Freedom’s Fetters 421-424 (1952). 48 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Table 1. Types of remarks indicted under Espionate Act 1917-1918 in Wisconsin Praising Germany . 36 Criticizing U. S . 35 “Rich man’s war” . 32 Criticizing Allies . 17 Insulting flag, uniform . 9 Criticizing bonds . 19 Criticizing food laws . 9 Criticizing charities . 15 Obstructing recruiting . 17 Praising ship sinkings . 3 (Total exceeds 90 because some indictments were for multiple remarks) win the war. Such remarks seem more likely to produce a brawl than a breach of national security. While society has an interest in preserving peace and tranquility, punishing a man for remarks on this ground alone is like punishing a rich man for keeping money around to tempt thieves. A similar breach of peace rationale has all but disappeared from American criminal libel law.25 Two Wood County men were convicted for remarks favoring Germany in July, 1917. One, a 44-year-old native of Prussia, was sentenced to two years in prison for saying, “I wish the war would end and the Kaiser would win. Yes, God damn it. We will never have good times until the Kaiser wins.” Although he had lived in the United States for seven years, his record showed that in 1914 he had been denied citizenship papers because he was not a “law abiding and peaceful” man. The jury was sworn in one morning, heard five government witnesses plus the defendant in the after¬ noon, and convicted him before nightfall. The other man was a Pole employed at a Grand Rapids paper mill. Three fellow workers, through an interpreter, told the court the man had said the Kaiser was sure to win the war and to sink all the American troop ships. The defendant told the court that 25 years before he had fled Europe after killing a German officer who disciplined him. His court-appointed attorney told the jury that surely it could not find this man disloyal or pro-German. “He is the only man in this courtroom who bears on his body a scar in¬ flicted by the brutality of Teutonic autocracy.” After hearing the prosecutor tell them that their only duty was to decide if the words had been spoken, not to judge their criminality, the jury proved it certainly could find the Pole guilty. It took them only 90 minutes to convict him, and the judge imposed a six-month term in the county workhouse and a $500 fine. A Madison druggist who long 25 Anon. “Constitutionality of the Law of Criminal Libel,” 52 i Columbia Laiv Rev. 521 (1952). 1970] Stevens — When Sedition Laws Were Enforced 49 had been prominent in German-American societies pleaded guilty to similar remarks and was fined $2,000. Even doubting the worst about the enemy could get a man in trouble, as it did a prosperous 68-year-old Grant County farmer who, among other statements, expressed disbelief about reports of German atrocities. (After the war, investigations showed that nearly all such tales were untrue.) The principal prosecution wit¬ ness was a neighbor who had a long-standing grudge with the de¬ fendant over a land transaction. This hostile witness testified he heard the farmer say, ‘The Germans killed no Belgian citizens and there are no orphans in Belgium or France that were caused by the war. The Germans have done no worse than Americans have done to Germans. . The farmer, the father of seven including a soldier in France, denied making such statements,, but the judge ruled the neighbor’s testimony admissable as he did that of two representatives of the Council of Defense who said there had been rumors about the farmer’s disloyal remarks on other occasions. The jury took 35 minutes to convict. The sentence : a year and a day at Leavenworth prison plus a $1,000 fine. Thirty-three indictments quoted criticisms, often quite pointed, of the American conduct of the war. Some of the offensive remarks said the Administration was too zealous, others that it was lethargic. Such opinions would seem to pose small danger to the success of a war; at least the United States managed to win World War II without prosecuting such criticisms. One of the heaviest punishments — 15 months in prison — was meted out to a 37-year-old Russellite evangelist from Milwaukee who was arrested at Plover in Portage County. He was convicted on four counts in a one-day trial at Eau Claire in July, 1918. His remarks, delivered in Polish, were translated thus in the indict¬ ment : The Constitution of the United States says that the govern¬ ment of the United States cannot compel a person to go to an¬ other country to fight, but Wilson has spoiled the Constitution, and is compelling men to go to other countries to fight. President Wilson started the war and he is now going to run away. A socialist attorney at Milwaukee was indicted in October, 1918, for telling two men (including a judge) that the United States had no business in the war and that its army should not be in Europe. On another date, he allegedly told a woman at Whitefish Bay, “You are upholding the cruelest and most abominable form of govern¬ ment in existence.” 50 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 A traveling salesman from Minneapolis was indicted in the Western District of Wisconsin for a remark he allegedly made in 1918 in a Portage store. He was accused of saying, “This Govern¬ ment was not ready for war. If Mexico were to rise up we could not protect our own country. The President and the Administration are to blame.” He was indicted three weeks after the Armistice but his case was dismissed six weeks later. A Milwaukee machinist was indicted for questioning the constitu¬ tionality of the draft law, and a former postmaster at Fall Creek got into trouble for questioning the constitutionality of the Espio¬ nage Act. Both cases were dropped without coming to trial, but the circumstances surrounding the Eau Claire County case are worth considering. Six men called on the former postmaster to see why he had not taken his assigned “quota” of $500 in Liberty Bonds. Allegedly he told them, “The Government of the United States is dishonest. Freedom of speech has been abridged by the espionage law. Free¬ dom of the press has been denied by the espionage law.” The solici¬ tors told the commissioner who heard the complaint that they had been chased off with a shotgun, but the ex-postmaster said he did not threaten them. He said he was carrying the gun because they were waiting for him when he came in from hunting. As to the remarks, he admitted criticizing the espionage legislation but denied saying the nation was dishonest. Although he never was penalized under the law, he was under indictment until July, 1920. The most frequent words quoted in indictments were “rich man’s war.” The term had been part of the vocabulary of socialists, Wob- blies and other anti-capitalist radicals for many years. In the case of the socialists, at least, they were saying the same things in Ger¬ many about the capitalist Junkers. Some of the remarks included under this heading were close equivalents, but all indicated a belief that it was a money war. Since it is impossible to prove the cause of any war — historians have suggested more than 100 causes for the American Civil War, and a century after the shooting seem no closer to an agreement than they were at the time — a statement about its being a “rich man’s war” can be considered no more and no less than an opinion. Certainly it is not a false report.26 A socialist who ran for the assembly from Sauk County in 1916 was indicated in 1918 for telling two neighbors it was a rich man’s war. He pleaded guilty and was fined $500, the same fine imposed on a Medford physician who ran for sheriff as a socialist and made similar remarks during his campaign. A Wood County man was 211 Anon., “The Espionage Cases,” 32 Harvard Law Rev. 417 (1919), 1970] Stevens — When Sedition Laws Were Enforced 51 sent to the workhouse for three months after pleading guilty to calling the war a “millionaire's graft." Criticizing Great Britain or France was apparently as serious an offense as criticizing the United States' war effort. The first indict¬ ment under the war statutes brought in the Western District was for such remarks. The owner of a Madison hotel allegedly told some draft-eligible Italian- Americans that since the war was being fought for England’s benefit they ought to head back to Italy to avoid the draft. At the time of his arrest in August, 1917, the fed¬ eral district attorney was quoted in the press as saying, “This thing of saying we are fighting for England must stop. I am going to have all such persons tried if I can obtain indictments for them. Any other remarks tending to honeycomb our solid front and to give comfort to the enemy and cause disloyalty among our soldiers will be summarily dealt with if I have my way." The hotel owner pleaded not guilty and was released on bond. Two years later the case was dropped without coming to trial. The 1918 amendment was a new wrinkle in American law, pro¬ tecting symbols from offensive words. Nine Wisconsin indictments referred to verbal attacks on the flag or military uniforms. On August 17, 1918, there was great excitement in the little Racine County town of Corliss. Men and women were scurrying about,, stringing up bunting and flags for a Red Cross parade. The town marshal was just stepping back to admire the decorations when he saw a threshing rig heading straight for the biggest Amer¬ ican flag, draped from a wire across the main street. Apparently the rig operator did not hear the warnings. Suddenly the top of his stream machine caught the flag, yanked it down, and devoured it. Three or four men raced out and waved down the operator, who pulled over to the side of the road. Over the terrible roar of the machine, they carried on an animated conversation about how he had ruined the flag. “Keep the damned flag out of the road," the operator shouted back. “It’s a public highway." Three months later, the operator was indicted for insulting the flag, not as one might expect for tearing it down, but rather for calling it a “damned flag." He pleaded guilty and was fined $200. As was pointed out earlier, the original excuse for requesting the 1918 amendment was to protect government securities from dis¬ paraging remarks. Attacks on Liberty Bonds and Thrift Stamps were quoted in 19 Wisconsin indictments. For telling three other men that the government “makes men buy bonds so that they can get their wages away from them," and that the bonds would be worthless after the war, a Richland Center socialist was sent to Leavenworth for a year and a day. It cost a 52 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 man from near Wausau $500 to say that the bonds would not be worth 40 cents on the dollar after the war. Unfortunately for him, he made the remark in the presence of the vice-president of the local Loyalty Legion. A Washburn man told a Liberty Bond sales¬ man that the bonds were no good for the poor man. Tried and convicted nearly a year after the Armistice, he was fined $200. Truth certainly was no defense, since a Taylor County man was fined $1,250 for saying the money from bonds was being used to help England defeat Germany. A Grant County farmer who said “they can’t make us buy any bonds” was fined $600. Although there was no rationing, a federal Food Commissioner in each state enforced rules against hoarding and profiteering. Eight Wisconsin indictments mentioned criticisms of the food controls; however, of these cases, one stands out because it was one of only two to be reversed on appeal and one of the few to involve a prominent state politician, John M. Becker, circuit court judge at Monroe for 21 years. Becker was a lifelong Democrat who supported La Follette’s stand against the war; in fact, Becker promoted a referendum on whether the nation should enter the war. The election was April 8, 1917, the very day Wilson went before Congress to ask a declara¬ tion of war. The vote was 954 to 95 against entering the war.27 Two days later, Becker addressed a patriotic rally, urging all citizens to be loyal to the nation once war was declared. He threat¬ ened criminal libel actions against those circulating rumors he had been charged with “aiding the enemy.” In August, Becker hinted and in January confirmed that he would run for governor on an anti-war platform. His candidacy, however, was short-lived, be¬ cause the Attorney General announced that Wisconsin law pro¬ hibited anyone from holding a judicial post and running for another office.28 Becker’s troubles were only beginning. In May he was indicted under the Espionage Act for saying on February 5 : The idea of having an administrator of fuel and food is ridic¬ ulous. There is no shortage of food. The idea of a shortage of food is being preached by agents employed by corporations for their own gain, and going about the country on high paid salaries. This is a rich man’s war. We won’t have peace as long as these high-salaried fellows have jobs to protect. There is no labor shortage. There is no seed shortage. Farmers, beware of taxes, war taxes, which must be paid in July. 27 Monroe Evening Times , March 23-April 4, 1917. 28 Monroe Evening Times, April 5—6. May 10 and November 19, 1917, and January 8, 1918. 1970] Stevens — When Sedition Laws Were Enforced 53 (His jury refused to convict on a second count based on testi¬ mony from a German woman bitter at Becker over settlement of her husband’s estate.) The Circuit Court of Appeals detailed the circumstances sur¬ rounding the meeting at which Becker’s food remarks were made. Because Green County had failed to meet its quota in the first two Liberty Loans, the state Council of Defense insisted the county council be reorganized. The county board called a meeting for this purpose, but because of a storm, only six of the 27 elected board members showed up. These six got into an informal discussion with 50 or 60 spectators, and in the ensuing session Becker spoke for 15 to 20 minutes. Two witnesses swore that the thrust of his re¬ marks were patriotic and that the statements quoted in the indict¬ ment were taken out of context. Other witnesses admitted they were talking among themselves and could not hear what Becker was saying. The appellate court reversed the conviction on the grounds that the words were not unpatriotic. The court also objected to testimony introduced during the trial about how many Liberty Bonds the defendant had purchased and to the fact that one wit¬ ness gave a Sunday dinner to the other witnesses.29 The trial was conducted at Eau Claire in August, 1917. One wit¬ ness collapsed from the heat. Almost none of Becker’s questions were about the food remarks but dealt with personal attitudes and his gubernatorial platform. At one point the special deputy district attorney called the platform “bunk.” Two friends helped Becker down from the stand after the long afternoon. The jury consisted of 10 farmers, one miller and one mechanic. After a week of testi¬ mony, it took them six hours to convict Becker. Wolfe told reporters he considered the conviction an important one since it might “deter” lesser individuals who might be inclined toward disloyalty. On August 16, Becker was sentenced to one year in federal peni¬ tentiary on each of three counts, the sentences to run consecutively. He was released on $20,000 bond, pending the appeal which was decided October 5, 1920. Upon his conviction, the state Attorney General ordered Becker to vacate his office. A successor was appointed and sworn in. After the reversal, Becker sued to recover his lost salary, but the Attorney General ruled against him on the ground that state law required resignation only for conviction of an infamous crime and the Espio¬ nage Act did not so qualify. The Attorney General told Becker the earlier order had been only “advisory” and that he complied of his own volition.30 29 Becker v. U.S., 268 F. 195 (1920). 30 Opinions of the Attorney General (1920), pp. 529-534. 54 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 By April, 1917, there were 159 voluntary war relief agencies in the United States, all conducting fund drives. The Red Cross alone collected more than $400 million during the war.31 Fifteen Wis¬ consin indictments quoted criticisms of the war agencies. It was in the conviction of Louis B. Nagler of Madison32 that the legal principle was established that such criticism could be prosecuted under the Espionage Act. Nagler, like Becker, was a prominent supporter of La Follette. While assistant secretary of state, he published a letter in the Madison paper defending the senator’s anti-war stand. His indict¬ ment resulted from remarks in November, 1917, to solicitors for the Red Cross. He told them he was through “contributing to your private grafts.” He named the YMCA, the YWCA and the Red Cross and said “not over 10 to 14 per cent of the money collected goes to the soldiers or is used for the purpose for which it is col¬ lected.” His indictment reasoned that since the Red Cross was chartered by Congress, it was a “government agency” within the meaning of the Espionage Act. Although the YMCA and YWCA were not chartered, they were engaged in similar morale work and logically should be covered, too. Nagler’s trial was scheduled immediately before Becker’s, and the district attorney made it clear they were to be showcase trials. His jury of 10 farmers, a miller and a railroad employee (five of whom had relatives in the army) wasted little time in convicting. The Circuit Court of Appeals upheld the conviction, and to the defense contention that such a decision would encompass even the Jewish Relief Agency, the Knights of Columbus and the Salvation Army, the court wrote, “With this position I agree.” In July, 1921, the United States Supreme Court remanded the case for a new trial, at which point it was quietly dropped. Nagler certainly was not cowed during the appeal. He even wrote a congratulatory letter to Victor Berger on his fine showing in the 1918 senatorial election.33 Among the other 14 Wisconsin indictments which included com¬ ments about voluntary agencies, all but one were based on replies to solicitors. The exception was a Milwaukee woman who said it was foolish to sew for the Red Cross since the rich people took all the garments and they never reached the soldiers. Her case was dropped without coming to trial. The tragedy which brought the war home to many Wisconsin citizens was the sinking of the troop ship Tuscania in the spring of 31 Cutlip, Scott M., Fund Raising in the United States 110-139 (1965). 32 U.S. v. Nagler, 252 F. 217 (1918). See Nagler, “A Fragment of War-Time History,” 121 The Nation 568 (1925). Nagler to Becker, Nov. 8, 1918, in Socialist Party Papers, Milwaukee County His¬ torical Society, Milwaukee, Box 23. 1970] Stevens — When Sedition Laws Were Enforced 1918. Soldiers from many Wisconsin communities went down with the ship. Those who lost friends or relatives were not in the mood for snide remarks about the incident. For example, a Port Huron man went to jail for six months for remarks about the effectiveness of German submarines. One man was fined $300 for saying he hoped the Germans sank every ship that we sent across the sea, while another pleaded guilty to a fed¬ eral charge arising out of his remark that “All those who go across ought to be at the bottom of the ocean.” Section 3 seemed to be aimed at persons who tried to keep others from fulfilling their duties, either by not registering or enlisting or by disobeying superior officers; still, few prosecutions had any¬ thing to do with recruiting.34 Fifteen Wisconsin cases did involve some actual counseling. The most prominent man convicted for such an offense was a former state legislator and president of the Marathon County Telephone Company. During his eight-day trial in September, 1918, the prose¬ cution called 27 witnesses, including the secretary of the Cassel town draft board who testified that Schilling had dissuaded more than 70 men from enlisting. The trial judge permitted testimony about his niggardly contributions to war charities and about his alleged hoarding of food, although neither was mentioned in the indictment. The jury was out only three hours before convicting. The next day the judge sentenced him to 18 months in prison plus a $3,500 fine. A Lithuanian who had lived in this country for 16 years went to prison for a year and a day for saying he would not register and for telling others they were fools for doing so. A Racine man al¬ legedly advised some Armenians not to report for the army if called, and he too was sentenced to a year and a day. Four other indictments cited remarks questioning the legality, not of the draft but of sending drafted men overseas; however, none of these cases came to trial. Trading Act During the summer of 1917, the House devoted three days and the Senate two to enacting this omnibus measure, whose primary purpose was to establish guidelines for seizing and holding prop¬ erty owned by alien enemies, particularly German corporations. Not until the final 15 minutes of debate was there any mention of a provision affecting expression, and then there were two. One was virtually a carbon copy of the new Section 3 later added to the 34 Apparently no prosecutor even attempted to prove injury to military procurement or allow a jury to consider possible effects of utterances. Nelles “In the Wake of the Espionage Act,’' 111 The Nation 684, (1920). 56 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Espionage Act. After its defeat (because the sponsoring Commerce Committee did not want such an unrelated section in the act) , Sena¬ tor William H. King then proposed a requirement that all German- language publications print a parallel English translation. Penal¬ ties were to be $500 or a year in prison. King explained that he earlier had proposed a total ban on foreign-language publications, but that this measure was bottled in committee. The Utah Demo¬ crat asserted that his regulation “certainly works no great hard¬ ships’’ on any publisher while at the same time providing a good check on sedition for the postal officials and for neighbors. A spokes¬ man for the Commerce Committee said he thought the Postmaster General already had sufficient censorship powers under the Espio¬ nage Act and expressed the hope that this irrelevant section would not be tacked on to the Trading Act. It was anyway, and the bill went to conference committee. The committee greatly expanded the mail section to cover publications in any foreign language. Instead of publishing English translations in adjoining columns, publishers now had to file such translations in advance of publication with local postmasters. Only material which dealt in some way with the war was covered by the requirement. The Speaker overruled objections that the conferees had exceeded their authority by add¬ ing new classes of crimes to the mail section, and the House adopted the report by voice vote.35 Most of the newspapers affected by the new regulations were one- or two-man operations, working on a tiny financial margin. Because filing translations cost both time and money, many foreign- language newspapers closed down. Others began publishing in English or tried to make their contents totally bland. That the latter tactic was not always successful for avoiding trouble with the federal officials is illustrated by the Auer case, described below. Auer’s was one of seven actions brought under the Trading Act in Wisconsin. Three of the actions involved possession of explosives and had no expression connotations and are not considered here. Four actions did involve expression under the mail section of the Trading Act. One of these was a joint action with the Espionage Act. Jacob J. Auer, a crotchety old man, was the editor and publisher of a little German-language weekly, the Eau Claire Herold. Most of his readers apparently were about his age. The younger generation had learned English in the schools and read “American” publica¬ tions instead. Even his closest friends and relatives testified that old Jacob was senile and that sometimes he “acted strangely.” He had been in this country more than 30 years and had been bitterly ss i Congressional Record , 65 Cong-., 1st Session, 4840-4879, 4907-4930, 4968-4989, 6949-6958, 7007-7025. 1970] Stevens — When Sedition Laws Were Enforced 57 disappointed a few years before the war when he was not appointed postmaster of Eau Claire. In his edition of December 6„ 1917, Auer published an editorial entitled, “Uncle Sam’s Army Threatened by Slow Destruction.” The menace he was describing was not the German army, but the United States Army’s policy of giving all its recruits smallpox vaccinations. The article was long, rambling and none too coherent. Auer did not file a translation of it with the postmaster for advance clearance, nor did he submit the next week’s issue, which quoted Hindenberg as charging the United States had used the submarine as an issue to enter the war on the side of the Allies. His December 27 edition carried an editorial which suggested Germany’s enemies were on the verge of collapse. In subsequent weeks, he attacked those citizens who were persecuting everything which was German, praised the German-Russion peace treaty and cautioned his readers not to believe everything they read in the press about how many German ships the Allies were sinking. He did not file translations of any of these articles, the local postmaster told a citizens’ protest committee meeting which met on March 15. The postmaster said Auer had filed translations of some other articles and editorials, however. The public meeting, which was sponsored by the county Council of Defense passed a resolution “to personally notify J. J. Auer that his attitude on the war, as expressed by said articles is seditious and disloyal, and warn him against publishing like arti¬ cles in the future.” Auer hardly had the opportunity, since he was arrested by the federal marshal eight days after the meeting. The marshal waited until after Auer had put out that week’s issue before taking him to Madison for arraignment. He was formally indicted in June, 1918, under both the Espionage and the Trading Acts. He first entered a plea of not guilty and then tried to change it to nolo contendere. The special assistant federal district attorney objected because such a plea carried with it only a fine, not im¬ prisonment. The judge refused the plea and Auer pleaded guilty. Before sentencing, a Civil War veteran and long-time friend of Auer spoke on his behalf. He said there had been a noticeable de¬ cline in Auer’s mental capacity during the last two years, a fact confirmed by both Auer’s wife and his son. His relatives testified that the old man had purchased Liberty Bonds and had contributed to the Red Cross and other fund drives. The son told the judge that he did not think that his father really understood the translation regulation. The prosecution charged the judge to show no mercy in assign¬ ing sentence. The judge said he was unmoved by the claims of mental incompetence since the articles seemed clearly written to 58 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 him; however, he would be merciful because of Auer’s advanced age. His version of mercy was a year and a day at Leavenworth. If Auer’s case seemed somewhat pitiful, consider the fate of poor Jacob Mueller, who edited the German-language Dodge County Pioneer at Mayville. In his edition of August 16, 1918, he ran a front-page story about Auer’s conviction, adding at the end : Since October 6, 1917, the German newspapers have been under the knout and have not been permitted to print any war news unless a true translation is filed with the postmaster. Since we have no desires to pester police with translations we print abso¬ lutely no war news. We are not going to be caught. Mueller was wrong about that. The district attorney thought Auer’s conviction was “war news.” With some earlier articles, this one formed the basis for his indictment in March, 1919. Mueller pleaded guilty and got off with a $50 fine. After all, the war had been over for four months. Until March 4, 1918, there had been two German weeklies in Mayville, but on that date the Dodge County Banner had switched to English. Ever since, it had been attacking Mueller for supporting Victor Berger and for having no principles. When Auer was con¬ victed the opposition paper wondered editorially if Mueller would relay that information to his readers. He did, much to his later regret. On October 24, 1918, about two weeks before a special Senate election in Wisconsin in which he was a candidate, Victor Berger and four other officials of the Milwaukee Leader were indicted under both the Trading and Espionage acts. Among other charges, they allegedly had failed to file translations of some letters to the editor which had been written and published in German. The five were released on bond and the case finally was dropped in 1922. Sabotage Act That there were few prosecutions under this broad statute, which punished “malicious destruction or injury to property, no matter how essential the property might be to the conduct of the war,” probably can be attributed to the fact that it was enforced by the Justice rather than the Post Office Department and that it was not signed until less than seven months before the Armistice. It is ironic that the federal government waited so long to prohibit overt acts of sabotage when it was so prompt to prohibit speech which might incite such overt acts. There were three actions under the Sabotage Act in Wisconsin, two of them against employees of the E. I. du Pont Demours Com- 1970] Stevens — When Sedition Laws Were Enforced 59 pany powder plant at Barksdale in Bayfield County. Plant guards were deputized by the sheriff and they kept a wary eye on “agita¬ tors.” They were also stringent in their enforcement of the rule against having matches within the plant gates. One employee was arrested in July, 1918, and charged under the Sabotage Act for carrying four concealed matches in his socks. Two weeks later an¬ other employee was arrested for carrying 12 matches in his socks. In the only Sabotage Act case tried in Wisconsin, a Milwaukee man was convicted by a jury for throwing ice tongs into plant ma¬ chinery; however, the trial judge ordered the case dismissed for reasons not made clear in the preserved record. There was a marked difference in enforcement in the two dis¬ tricts of Wisconsin. Not only were half as many persons indicted under the Espionage Act in the more populous, more socialist, more German Eastern District, but a far higher proportion of cases were dismissed without coming to trial. Table II compares the activity in the two districts. Apparently the major variable was the vigor of the enforcers. Certainly the laws were interpreted quite differently by different federal judges.36 Chafee pointed out that although Massachusetts 38 The Department of Justice issued 204 “Interpretation of War Statutes” bulletins in an effort to increase consistency. Table 2. Actions under Federal War statutes in Wisconsin No. Pled Guilty Convicted Acquitted Dismissed Espionage East . 30 2 1 2 25 West . 60 30 17 3 8 Threats East . 3 0 1 1 1 West . 7 5 1 0 1 Trading East . 3 1 0 0 2 West . 1 0 1 0 0 Sabotage East . 1 0 1 0 0 West . 2 0 0 0 2 Total East . 37 3 3 3 28 West . 70 35 19 3 11 GRAND TOTAL 107 38 22 6 39 60 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 had large military posts, a major port of embarkation, many war plants, and a large alien enemy population (to say nothing of the anti-English population of Irish) , there was not a single Espionage Act indictment in the Bay State. The District Attorney simply re¬ fused to bring any, and Chafee said there was no rash of sabotage there as a result.37 Even more striking was the contrast in punishments in the two districts. In the East, the three defendants who pleaded guilty were fined a total of $900, and none of them went to prison. The three convicted in the Eastern District each went to Leavenworth for a year. By contrast, the 85 defendants who pleaded guilty in the West were fined $600 but sentenced to 28 years and 2 months in prison. The 19 convicted were fined $13,100 and sentenced to prison and workhouses for a total of 26 years and 3 months. The only prison sentences of more than a year and a day came in the Western District, where one man (Becker) was sentenced to three years,, two others to two years, one to 18 months and an¬ other to 15 months. All were for violations of the Espionage Act. Thirteen men (10 of them in the West) received the year-and-a- day sentences. Eleven others were sentenced to terms of 30 days to six months in various county workhouses. The two heaviest fines were both in the Eastern District and both were for violations of the Espionage Act. One man was fined $2,000 and another $1,250. No other fine exceeded $500. In total, the 96 men convicted or entering guilty pleas to war statute violations of the Espionage Act in the two Wisconsin courts were sentenced to 57 years and 5 months in prison or workhouses38 and fined $14,600. During World War II, there were almost no actions against “dis¬ loyal” expression under any of these five laws, although all were on the books in slightly altered form. There were no such actions in Wisconsin. The civil liberty picture of World War II was a much brighter one in spite of the blot of the Japanese evacuation. Perhaps it was because the Administration was more enlightened, or the society was less naive, or there were fewer internal threats. Or perhaps it was because the nation profited from the black mass of World War I. 37 Chafee op. cit., supra note 1, at pp. 59-60. 38 If Berger had not been convicted of Espionage Act violations and sentenced to 20 years in the Northern District of Illinois, Berger v. U.S., 255 U.S. 22 (1921), it is likely he would have been tried on one of the three cases pending in the two Wisconsin federal courts ; in such case, Wisconsin’s penalty total might have been 20 years higher. POLICE IN A LARGE SOUTHEASTERN WISCONSIN COMMUNITY John C. H. Oh Abstract This is a study of the local law enforcement personnel, who not only possess but often exercise a great deal of discretionary power in the course of administering the laws and ordinances in their local communities. Their general attitudes and the manner of con- tacting with the public have very important consequences upon the society. Yet, we know very little about these people. The present study is based on the analysis of the survey ques¬ tionnaires returned by police officers of a large southeastern Wis¬ consin community. It was designed to answer the following three questions : 1. How the police officers felt about the role of the Supreme Court in the criminal justice system in American society, par¬ ticularly in regard to some of the recent Court decisions deal¬ ing with the constitutional rights of the suspect— in the area of criminal law and procedures; 2. How they felt about the law and order— their general atti¬ tudes toward violence, civil rights, and social order; 3. And, how they felt about themselves— their general attitudes toward their work and status in the community. The study shows that police officers generally have a very low regard toward the U. S. Supreme Court, that the majority of police officers tend to develop an inferiority complex in their work (because they believe that the public does not extend due recogni¬ tion to the police officers), and that they are excessively protective of their own work and quite unreceptive to any kind of criticism toward the police officers everywhere. It must be cautioned that these findings are only preliminary and suggestive; however, any concerned reader cannot help but to conclude that it is an urgent national task to intensify our efforts to train the police officers with the ideals of democracy and constitutionalism either through in-service or out-service training, or both. 61 62 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Introduction In recent years social scientists have made some significant con¬ tributions to the understanding of two related questions concern¬ ing the community power structure — “where power lies” and “who makes decisions” in the political communities. Consequently, we probably know more about the “men at the top” at both govern¬ mental and non-governmental levels in various political communi¬ ties than ever before.1 On the other hand, we know very little about the middle and/ or lower level of the governmental personnel, because the same social scientists have generally shunned conduct¬ ing any systematic studies on them. One such area of neglect is the local law enforcement personnel. These people-— whether they are called constables or sheriffs or police officers — not only possess but often exercise a great deal of discretionary power in the course of administering the laws and ordinances in their local communities. Their general attitudes and their manner of contact with the public have very important consequences for the society. The present study is based on the analysis of the survey ques¬ tionnaires returned by 55 of 73 police officers of the Waukesha Police Department during the period of October 14-21, 1968. A brief description of the community being surveyed is in order. Waukesha has experienced one of the most rapid population increases among Wisconsin’s cities and is a city of 36,339 people according to a special census of 1966, as compared to 30,004 in 1960 and 21,233 in 1951. (Waukesha County, which is seated in the city, had an increase of 326.5% during the period 1910-60, which was the fastest population growth rate among the state’s 70 counties.) The city was the fourth largest in the Greater Milwau¬ kee Standard Metropolitan Statistical Area,2 which in 1968 con¬ tained an estimated population of 1,458,100, of which only 83,931 (or 5.8%) were Negro. The city itself had only a dozen or so Negro families, since the bulk of the Negro population in the area resided in the central city (Milwaukee). It must be noted, how¬ ever, that the city is not a suburb of the central city in the strictest sense, because if is a booming industrial community on its own with a number of nationally-known manufacturing firms (i.e. It had some 23 banking and savings and loan associations with total assets of over $150 million at the end of 1966). 3 1 See Floyd Hunter, Community Power Structure ( 1953 ) , Robert Presthus, Men at the Top (1964), Robert A. Dahl, Who Governs ? (1961), and Arnold M, Rose, The Power Structure ( 1967 ) , among- others. 2 The area covers at least 38 separate governmental units which include four counties, 15 cities, 15 villages, and four towns. 3 U.S. Census of Governments, 1962 ; State of Wisconsin, Blue Book, 1968 ; Milwaukee Sentinel, 1969 Wisconsin Almanac (January 7, 1969). 1970] Oh— Police in a Wisconsin Community 63 Politically, Waukesha residents have generally voted for Repub¬ lican candidates. In the latest election they again voted heavily Republican for the presidency, the state-wide ticket (for the gov¬ ernor and four other constitutional officers), the state assembly- man, the state senator, and for the U. S. congressman. The only Democrat who received the plurality of votes in the city was an independent-minded, popular U. S. senator. As far as the incidence of crime is concerned, it was reported that Waukesha had a total crime index of only 295 in 1967, which was not only far below the national average of similar size cities but also lower than the state average of 1,121.1 and that of the Milwaukee SMSA’s 1,613.2. This lower crime incidence may be par¬ tially due to the fact that Waukesha's full-time police employee rate of 1.8 was far in excess of an average number of officers per 1,000 inhabitants for the same size communities in the state and the nation as a whole.4 The present study was designed to answer the following three questions : 1. how the police officers felt about the role of the Supreme Court in the criminal justice system in American society, par¬ ticularly in regard to some of the recent Court decisions deal¬ ing with the constitutional rights of the suspect in the area of criminal law and procedures; 2. how they felt about law and order — their general attitudes toward violence, civil rights, and social order; 3. and, how they felt about themselves — their general attitudes toward their work and status in the community. We felt that October, 1968, was the most opportune time for this type of study because many of the questions used in the survey were the same kind of issues raised by the various candidates in their election campaigns. Our actual subjects included 37 uni¬ formed patrolmen and 18 detectives and police executives. The data in Table 1 show that there were some marked differences between these two groups of officers, in that patrolmen were in general younger (51% under age 30 compared to only one of 18 detectives in that age group), better educated (38% to 17% for some college work), and less experienced in police work (an average of six years service for the former as compared to 13 years for the lat¬ ter) than detectives and executives. 4 Federal Bureau of Investigation, Uniform Crime Reports, 1967 (August 27, 1968). Table 1. Age and Education of Police Officers Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 ill > a s § o < CQ < z o H < U D Q W Q ^ y W 5S >£, H U W H W a b W bo gju o u c8 c f) COl sD ^ Tf sD | in o o r^N tx c-n — ' Q Z £ w 3 5 o □ * 5 - ^ o, gca P| fc < U D Q fxl o mi g3> o U co z D 8“i O 1-rH COl -^OOVSvO vrs r^N O o co ro | ^ 00 kn^n co O' ro vr\ co o£ Pi W W CQ. y s; fcc D u* y 06 o o o r*"\ ^ ir\ AAA o CQfflCQO >_; CO ^ UTS T3 £ D O S-, bfi O cd X! c o w o D Tj <0 < * 1970] Oh — Police in a Wisconsin Community 65 On the Supreme Court and the Criminal Justice Our study shows that police officers generally have a very low regard for the U. S. Supreme Court, to such an extent that the very legitimacy of the highest court is being questioned. Almost unanimously (93%), they felt that the Court in its recent decisions dealing with criminal law and procedures has reduced sharply the effectiveness of the police. However, it was found that such gen¬ eral negative attitudes toward the Court did not prevail consist¬ ently when we asked them a series of questions concerning their particular attitudes toward several selected court decisions which dealt with the constitutional rights of the suspect. They were specifically asked about these five decisions of the Court : Miranda vs. Arizona (1966) ; Mapp vs. Ohio (1961) ; Mallory vs. TJ . S. (1957) ; Gideon vs. Wainwright (1963) ; and Escobedo vs. Illinois (1964). (In asking their reactions to these cases, we described the essence of the decisions and elicited their reactions, instead of by the official legal citations.) First, as to their reactions to the Miranda decision, we asked the police officers whether the Court ruling that requires them to inform a suspect of his constitutional rights before questioning him interfered with their performance of duties. The data5 show that a great majority of detectives and executives (72%) who must question the suspect in the course of their investigatory work felt that it interfered with their work, while only 22% felt it did not. The uniformed patrolmen were almost divided evenly. This change of attitude (from their general negative attitude toward the Court) is very significant in view of the fact that the Miranda decision was the most comprehensive requirement laid down by the Court to protect the constitutional rights of the suspect from being infringed upon by the law enforcement officials. Surprisingly, in the next three cases we find that there were more police officers agreeing with Court opinions than disagree¬ ing. In the Mapp decision, 49% agreed with the Court that evi¬ dence obtained by illegal searches and seizures cannot be intro¬ duced into a state court, while 47% opposed it. In the case of the Mallory decision, only 71% felt that the Court decision (requiring a prompt arraignment of the suspect) is a reasonable one, but 80% stated that it did not interfere with their work in any way. The overwhelming majority (82%) also said that they agreed with the Gideon decision (which requires a state to provide free counsel for defendants who cannot afford a lawyer) . 5 Additional statistical supporting- data are deleted throughout the paper. They are, however, available from the author upon request. 66 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 The police officers in general find the Escobedo decision most objectionable. The Court in 1964 ruled in this case that a defend¬ ant has a right to have his lawyer with him when he is being ques¬ tioned by police officers. Seventy-one per cent of the respondents (88% of the detectives and executives and 65% of the uniformed patrolmen) indicated that this decision interfered with their inves¬ tigative work. Altogether 56% of the officers felt that it was a bad decision. It must be emphasized that what they objected to most was not that the defendants should be given free counsel to defend themselves but the Court ruling that permits the presence of a lawyer at the time of their questioning of the suspects. In this sense, the Escobedo case is the main cause of the Supreme Court’s unpopularity among the police officers. On Social Order, Violence and Civil Rights Our study shows that police officers are generally conservative toward issues such as social order, violence and civil rights. We first asked them the following open-ended question : “There is much discussion over the police action that took place during the 1968 Democratic National Convention in Chicago. Please indicate how you feel about the incident, including whether or not you feel that the Chicago police over-reacted in the situation.” Almost all the respondents (95%) felt not only that the Chicago police did not over-react in the controversial incident but also that what they did was proper because, as one officer indicated, “the demonstra¬ tors were forewarned and if they didn’t want to get hurt they should not have been there.” Not one officer said that the Chi¬ cago police should have handled the situation differently-— in direct contrast to the findings of the Walker Report. Even with Chicago Mayor Daley’s very controversial order “to shoot to kill the loot¬ ers” (as reported by the mass media during the disturbances), only 11% indicate that they did not agree with the mayor. When asked “How do you feel about alleged police brutality?” 83% of the officers claimed that not only does it not exist but such a charge is part of a conspiracy to undermine the police through¬ out the country. In reply to another question on civil disorders in our cities and college campuses, 96% of the officers stated that maximum (rather than minimum) force should be used in quelling such disorders. A substantial majority (69%) also felt that there is too much violence on T.V. and in the movies. It is significant to note that police officers in general felt that in order to stem the rising crime rate the criminal offenders must be given much stiff er penalties than they receive now. Fifty-one per cent of the officers (61% of the detectives and executives and 1970] Oh — Police in a Wisconsin Community 67 46% of the uniformed patrolmen) favor capital punishment for persons convicted of crimes such as first degree murder, kidnap¬ ping, sabotage or treason, in spite of the fact that the state of Wis¬ consin does not allow capital punishment. When asked “whether they felt we needed stiffer penalties in cases involving misdemean¬ ors and/or felonies/’ 64% indicated that there should be stiffer penalties for misdemeanors, and almost all the officers (97%) felt the same way toward felonies (the major crimes) . In a near unani¬ mous opinion, they opposed the parole system as now being prac¬ ticed in the United States, because they strongly feel that when a criminal is given a specific sentence — whether it is life imprison¬ ment or a specified number of years of confinement — he should be forced to serve it without being released on parole. Politically, too, police officers generally leaned toward the con¬ servative side. They have a higher voting turnout than the general population, for 78% of them voted in 1964 (it would have been much higher if 12% were not under the voting age). In that elec¬ tion year, of those who revealed their actual vote (69%), they voted almost two to one in favor of Johnson (45%) to Gold water (24%). However, when asked “Which of the candidates (Nixon, Humphrey, Wallace) have you decided to vote for in 1968?” 35% favored Wallace, 33% Nixon, 5% Humphrey, while 27% were either undecided or gave no response. What surprised us most was that this particular community had not experienced any civil dis¬ order per se in recent years, and yet in view of their general atti¬ tude toward law and order, a substantial majority of officers (68%) were attracted to either Wallace or Nixon. On the Role and Status Perceptions of Police Officers Generally, police officers possessed a strong sense of community service, dedication, and altruism. Sixty per cent of all police people and at the same time serving the community best through law en¬ forcement work. Another 29% felt that the profession gave them a sense of job security. Only 4% felt that police work gave them excitement and adventure not to be found in other lines of work, while 2% chose it because of family tradition. However, they also agreed that the general public does not ex¬ tend due recognition to the police officers. The data show that al¬ though higher ideals motivated them to choose the law enforcement profession, a majority of police officers (58%) tend to develop an inferiority complex in their work, for they believe that the public generally tends to look down on them socially. Eighty per cent of the officers also felt that they are not being rewarded in terms of salaries and fringe benefits as they think they deserve. (This salary 68 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 schedules of this police department ranged from about $580 to $900 per month depending upon rank and seniority of the officers). One surprising finding was that when asked whether they would advise young people to go into police work, 74% answered affirma¬ tively. It suggests that their lower reputational perception is largely due to the general public's apathy toward police work, because most police officers believe that they are contributing something positive to the general well-being of the community. Finally, it must be pointed out that police officers are not only conscious of their own status but quite defensive about their work and the police everywhere. We have already shown how sensitive and defensive they were to the charges of police brutality and the Chicago convention incident. To further check on this feeling we asked them the following question: “What educational require¬ ments do you feel police officers should have?" The data show that 51% of all the respondents felt that education beyond high school is needed, but only 4% felt they should have a college degree. There is, however, marked difference between the relatively younger uni¬ formed patrolmen and the somewhat older detectives and execu¬ tives, since 60% of the former stated that police officers should have more than a high school education as compared to only 34% of the latter. These attitudes tend to reflect their own educational background, for within this department not only is there no single college graduate but the formal education of the detectives and executives is comparatively lower than that of the uniformed patrolmen. (In this connection, it is interesting to not that the governor of Wisconsin early this year proposed the establishment of a state-run police academy to train all police officers within the state.) Conclusion These findings are not entirely unknown, for both the Kerner Report and the National Crime Commission Report made it clear that police officers in general have one of the lowest formal educa¬ tional attainments among all the professional groups in the country. Some of the disturbing findings are that : 1. The majority of police officers tend to develop an inferiority complex in their work, in spite of the higher ideals that mo¬ tivated them to choose the police profession (because they be¬ lieve that the public does not extend due recognition to the police officers). 2. They are excessively protective toward their own work and quite unreceptive to any kind of criticism toward police officers anywhere (e.g. their reactions to the Chicago violence). 1970] Oh— Police in a Wisconsin Community 69 3. They generally believe that social order can best be main¬ tained by the maximum use of force by law enforcement officers and by imposing much stiffer penalties upon the crim¬ inals than they are given today. 4. They apparently have very little faith in the Supreme Court of the United States — the highest court deciding questions of law and the official body for interpreting the U.S. Constitution. It must be cautioned that these findings are only preliminary and suggestive; however, any concerned reader cannot help but con¬ clude that it is an urgent national task to intensify our efforts to train police officers with the ideals of democracy and constitu¬ tionalism either through in-service or out-service training, or both. It may well be that eventually education beyond high school (either college education or relevant police education through police acad¬ emies or special institutes) must become the mandatory require¬ ment for all those who seek a police career. TRENDS IN WISCONSIN’S TOURIST-LODGING INDUSTRY L. G. Monthey Highlights This research study included all of Wisconsin’s lodging establishments that were inspected by the State Board of Health during the years 1961 to 1968, inclusive. It reports significant changes in the number, distribution, seasonality, size, and type of tourist-lodging establishments in that period. Wisconsin lost 1,019 establishments between 1958 and 1968; approximately two-thirds of these were seasonal businesses,, primarily resorts. However, the State’s total capacity in bedroom units (B.U.) has remained near 80,000 for 11 years or longer. Since 1961 the total number of hotels dropped 21%; resort-type businesses declined 14%; motel establishments increased 16%. Small establishments with less than 10 B.U. decreased 16%, while the num¬ ber of large enterprises (30 B.U. or more) increased 17%. Motels were the only type of T-L establishment that increased in all size classes, seasonal and year-round, during the 8 years. Among the seasonal establishments (open 9 months or less) only the motel- type business showed a gain in both firms and B.U. capacity. Likewise, motels were the only type to show a large increase in average size, going from 13.8 to 18.5 B.U. in 8 years. The number of year-round establishments also declined, but their total B.U. capacity went up about 5,000 during the period. The number of year-round resorts dropped 14% between 1961 and 1968. The biggest losses in both number of establishments and total B.U. capacity occurred in the northeast and northwest regions of Wisconsin. Tourist accommodations, and the housing enterprises which provide them, are an important part of Wisconsin’s $900 million travel-recreation industry. However, significant changes have taken place within this business since 1960. This study is an at¬ tempt to determine and measure some of these changes over a period of years, also to identify and quantify the important trends that are taking place in the tourist-lodging industry (here¬ inafter referred to as the “T-L industry”), which is in a state of rapid transition. The methods used are similar to those employed in an earlier study entitled “The Resort Industry of Wisconsin.” (Wis. Acad. Transactions, Vol. 53/Part A/ 79-94, 1964.) In order to obtain comprehensive inventories of Wisconsin’s T-L business, State Board of Health inspection records and mail¬ ing lists for the years 1961 through 1968 were used. The appro¬ priate data were coded for each establishment, transferred to IBM cards, and the results compiled by data-processing techniques. Four 71 72 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 major categories were used in classifying T-L businesses: (1) Hotel type; (2) Motel type; (3) Resort type; (4) Other. The establishment’s name was used to categorize each business, and each category was then studied in detail as to the distribution, number, size, and seasonality of the establishments it contained. An Eleven-Year Look Thanks to an earlier study of the tourist-overnight accommoda¬ tions in Wisconsin,* we can go back to the year 1958 for some limited data on the total number of establishments, their season¬ ality and bedroom unit (B.U.) capacity. This enables us to trace a few of the major trends over an 11-year period (1958-68). The number of T-L establishments has declined markedly, but the total capacity for housing visitors has held its own. In 1958 there were 7,842 firms offering about 79,000 B.U. to the traveling public. By 1968, the number of businesses had dropped to 6,823 — a decline of 1,019 establishments — but the total B.U. capacity re¬ mained near 80,000 units. (Note Tables 1-a and 1-b.) A seasonal establishment is one which operates less than 9 months during a given year. Of the one thousand T-L establish¬ ments that have disappeared since 1958, over two-thirds were sea¬ sonal in nature and predominantly resort-type businesses, as we shall see later. It is noteworthy that the greater part of this de¬ cline, both in total establishments and in seasonal businesses, has occurred since 1965. This suggests that certain trends, at least, have accelerated during the past few years. Tables 1-a and 1-b show not only the big losses in T-L estab¬ lishments — 673 seasonal and 346 year-round — over the 11 years, but also a substantial drop of almost 5,000 in seasonal B.U. * By Fine and Tuttle, School of Business, University of Wisconsin. Table 1-a. Overall Summary: Changes in the total number and SEASONALITY OF WISCONSIN TOURIST-LODGING ESTABLISHMENTS DURING THE PERIOD 1958-1968. Year Studied Seasonal Establish¬ ments Y EAR-ROUND Establish- 'ment$ Total No. Establish¬ ments % Seasonal Businesses 1958 . 5,668 2,174 7,842 72.3 1961 . 5,733 1,983 7,716 74.3 1965 . 5,638 . 1,827 7,465 75.5 1968 . 4,995 1,828 6,823 73.2 Change (1958-68) . (-673) (-346) (-1,019) 1870] Monthey — Wisconsin’s Tourist-Lodging Industry 73 Table 1-b. Overall Summary: Changes in the total number and SEASONALITY OF BEDROOM UNITS (B.U.) AVAILABLE AT WISCONSIN Tourist-Lodging Establishments during the period 1958-1968. Year Studied Seasonal B.U. Year-round B.U. Total No. B.U. Seasonal B.U. (as % of Total) 1958 . 47,577 31,533 79,110 60.1 1961 . 47,608 32,690 80,298 59.3 1965 . 47,085 34,719 81,804 57.6 1968 . 42,611 37,087 79,698 53.5 Change (1958-68) .... i (-4,966) + 5,554 + 588 An Eight-Year Study We now depart from an 11-year comparison and concentrate on a more detailed 8-year analysis of T-L establishments. Except for the above-mentioned tables, all of the remaining statistical mate¬ rial — including figures and discussion — relates to the period 1961- 68 inclusive. Generally speaking, data for the years between 1961 and 1968 are not included herein, since most of the changes and trends observed are quite consistent throughout the period studied. Thus, virtually all of the data used here relate only to the years 1961 and 1968. Despite changes in the number, type and distribution of T-L establishments, the total visitor-housing capacity has remained near 80,000 B.U. and has not varied as much as 3% since 1958. However, the number of year-round B.U. has increased about 5,000 since 1961, reaching a high of 37,087 in 1968. A comparable de¬ cline in seasonal B.U., from 47,600 to 42,600, has tended to off¬ set this gain. In other words, 60% of all B.U. were provided by seasonal establishments in 1961 as compared to only 53% in 1968. It is interesting to note, however, that about 73% of all establish¬ ments, statewide, were classed as seasonal in both years. Table 2 involves the main categories of T-L establishments and includes the total B.U. capacities and number of firms under each category for the years 1961 and 1968. A few definitions may be in order at this point. Some Definitions A hotel is defined as a lodging house, usually more than two stories high, having five or more bedroom units and (usually) a public lounge or lobby plus food service. Most Wisconsin hotels were built prior to World War II. 74 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Table 2. Overall Summary: Changes in the Type and B.U. Capacity of Wisconsin Tourist-Lodging Establishments between 1961 and 1968. Type of Establishment 1961 1968 1961-68 Changes No. B.U. No. B.U. No. B.U. Hotels . 583 19,567 457 15,971 (-126) (-3,596) Motels . 915 12,600 1,056 19,505 + 141 +6,905 Resorts . 4,761 40,465 4,101 36,255 (-660) (-4,210) Other . 1,457 7,666 1,209 7,967 (-248) + 301 Totals . 7,716 80,298 6,823 79,698 (-893) (-600) A motel is a T-L establishment, usually on one level and seldom with over two floors,, having five or more bedroom units and a con¬ venient auto-parking space on the premises at no extra charge to the guests. The distinction between hotels and motels is usually quite clear for the small establishments with fewer than 20 B.U. However, there is almost no definable difference between medium¬ sized and larger hotels and motels (usually called motor hotels), except for structural age. Virtually all of the latter are less than 20 years old, whereas very few hotels of the traditional type have been built since 1950. In any case, our motel category includes the newer motor hotels. A resort is defined as a T-L business situated in or near a scenic and/or recreational environment. In Wisconsin a resort- type estab¬ lishment is usually on or near the water, either a lake or river, but not always. Our fourth category of establishments, designated as “Other”, includes all types of T-L businesses that cannot be identified under the three classifications listed above. As Table 2 shows, there have been some significant changes in the types of T-L establishments between 1961 and 1968. Hotels and resorts have diminished in number. The hotel total dropped from 583 to 457 during the 8 years, a decline of over 20%. Motels, on the other hand, increased by 15% and showed a gain in capacity of almost 7,000 B.U. The totals for resort-type establishments show a loss of 660 firms and a drop of about 4,200 B.U. since 1961. The “Other” (miscellaneous) category of T-L establishments had a significant loss in the number of businesses, but it gained about 300 B.U. during the 8 years. This group is predominately tourist homes and rooming houses, but youth camps, guest farms, dude ranches, and many other kinds of housing facilities are included among its 1,209 establishments. 1970] Monthey - — Wisconsin’s Tourist-Lodging Industry 75 In 1961 resort-type establishments comprised 61.7% of all T-L businesses in Wisconsin; in 1968 they still made up 60.1% of the total. Meanwhile, the motel category increased from 11.8% to 15.5% of the total in just 8 years, and the hotel group declined from 7.6% to only 6.7%. Size of Establishments As Table 3 indicates, there were some noteworthy changes in the size of housing establishments during the 1961-68 period. In 1961, for example, the average size of T-L establishments was about 10.4 B.U. — ranging from 8.8 B.U. for seasonal properties to 16.5 for year-round businesses. By 1968 the over-all average had increased to almost 12 B.U. per establishment, with the seasonal businesses showing only a slight increase to 8.5 B.U. and the year- round operations rising sharply to 20.3 B.U. in average size. The number of small establishments has been dropping rapidly, especially since 1965. Those with less than 10 bedrooms comprised 64.7% of all lodging places in 1961„ but they declined to 61.3% of the total by 1968. In 1961 there were 4,995 such businesses offering a total of 23,046 B.U. to the public. Eight years later there were 4,181 establishments and 19,229 B.U. in this l-to-9 size class, which reflects a loss of 814 establishments and 3,800 B.U. between 1961 and 1968. This drop, the largest in any size group studied, was especially noticeable in the 5-to-9 B.U. range. The intermediate group of establishments with 10 to 19 B.U. also declined in number, dropping from 1,937 to 1,806 properties and showing a loss of almost 1,500 B.U. during the 8 years. This group, however, still makes up a substantial segment of the indus¬ try. It included 25.1% of all Wisconsin T-L businesses in 1961 and 26.5% in 1968, providing about 30% of the State’s 80,000 B.U. in the latter year. Table 3. Overall Summary: Changes in the Size Classification of Wisconsin Tourist-Lodging Establishments between 1961 and 1968. Size Class of Establishment 1961 1968 1961-68 Changes No. B.U. No. B.U. No. B.U. 1-to- 4 B.U . 2,682 7,353 2,325 6,467 (-358) ( -886) 5 -to- 9 B.U . 2,313 15,693 1,857 12,762 (-456) ( — 2,931) 10-to-19 B.U . 1,937 25,366 1 ,806 23,888 (-131) (-1,478) 20-to-29 B.U . 458 10,610 461 10,662 + 3 + 52 30-to-99 B.U . 277 12,643 308 14,352 + 31 + 1,709 100 and over . 49 8,633 66 11,567 + 17 + 2,934 Totals . 7,716 80,298 6,823 79,698 ( — 893) ( -600) 76 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 It is probably significant that all size categories under 20 B.U. per establishment showed substantial declines in capacity, with a total loss of 945 businesses and 5,295 bedroom units between 1961 and 1968. Meanwhile, those properties with 20 or more B.U. increased from 10.2% of all T-L businesses in 1961 to 12.2% of the 1968 total. There were 784 such firms with a total of 31,886 B.U. in 1961 ; by 1968 this group included 835 businesses with 36,581 units. This 20-plus category provided 39.7% of the State's B.U. total in 1961 and 45.9% of it in 1968. There was a most substantial gain of over 4,600 units in the group of larger establishments with 30 or more B.U. per business, which comprised 5.5% of the T-L firms in 1968. However, the one category showing the largest increases of all — both percentage¬ wise and in B.U. capacity — was the 100-plus group which gained nearly 3,000 units from 17 additional establishments. Here we ob¬ serve a 34% increase in the number of businesses and a 35% gain in B.U. in only 8 years, 1961-68. Table 4 shows the 1961-68 comparisons of Wisconsin T-L estab¬ lishments as to type, seasonality, number, and bedroom (B.U.) capacity. It also indicates the average size of establishment under each classification, both seasonal and year-round, and under the combined totals for each year. Fewer but Larger These data clearly illustrate the fact that, in general, Wiscon¬ sin's T-L establishments are getting fewer in number but larger in size. But they also reflect some significant variations among the major categories of establishments. For example, the impact of the newer motels and motor hotels on the traditional hotel-type opera¬ tion is quite apparent from these figures. The general decline in resort-type establishments is also evident in Table 4, where we note a loss of 660 businesses and 4,200 B.U. since 1961. Our data indicate that the greater portion of this resort loss has occurred since 1964, when the rate of decline for these establishments accelerated to about 4% per year. The only group of T-L businesses which has increased steadily in both number of establishments and in B.U. capacity over the 8-year period is the motel category. This group has also shown the largest gain in average size of business (B.U. per establishment). The remainder of this report will consider in more detail the major changes taking place, as well as apparent trends,, within each of the four main categories of T-L businesses. The geographic nature of these changes, especially the regional trends and patterns involved, will also be presented and discussed. Table 4. Type, Seasonality, and Average size of Wisconsin Tourist-Lodging Establishments, Comparing 1961 and 1968. 1970] Monthey — Wisconsin’s Tourist-Lodging Industry Total Estab¬ lishments 583 19,567 (33.6) 915 12,600 (13.8) 4,761 40,465 (8.5) KnOon Von con ^ N D ■ - -VON — < tN^ N_^ 7,716 80,298 (10.4) < "0 co < § $ ’ — < ^--n 'tNo-s Nr\ON nONon CON O O-N r\ O S VO Tf von t\ n r ' — < C<1 )0 O <11 ^ JhCQ CON ' — ' CO — 1 CD CO co co v — > 00 ^ ^Non — -^o-N con 00 ^-n — — < vO t)NvO CON nQ tNOcc CON O CON O £ — 1 ■'f • ro O ■ ^ CnI • O' O ■ t>s nO • - -00 -ION - -00 CO ro 0) '+K'-o V/N CON tN.'-o o cd H < ”0 co r"" H < C o d ^ von c<) — n CON , — s t\ o ^ CON 00 ^-n 00 t\ ^ /“s o $ 00 nO 00 C^s, von Von cO 3 .— I N — < CON ' — ' o ^CQ OT — 1 CD CO co Cd O' O-S riOOoN VON ^ — X vOO'/^ VO —I C w KOvO 00 ^ O' con nD — < — • 00 O' vo O $ ^ • ro nO ■ 00 00 • 00 t\ • O' N O • co C - O' -CNl - -00 -VON - -00 S "ot (ON CON CN> n— ' "'F w CD ' — ^ CD -? ^CQ CON : ; ; ; ; ; ; ; ; ; ; ; c n c/) s^cq 'w £ a) id O N ■§4^ 0 o > B 6 d< OZZ DC V) OT . c-gD Iddd JZ Cw' ot C w id O N & f-j'o'o > uu . . ^ H o o< OZZ 2 c-gD CQ ”ci £ a 00 cc O' wmaOO -'f cO O O' U + 0 ' + No. Motels iaNO'vO O rc —i vn OON O sQ — t\ — O' + Q Z D O oS No. of B.U. 1968 Data 6,216 6,596 3,045 15,857 1961 Data 5,717 3,525 330 9,572 +6,285 cC < w >- No. Motels Motels — 567 187 19 773 Motels — 539 113 2 654 + 119 Seasonal No. of B.U. 2,353 1,295 3,648 2,228 800 3,028 +620 No. Motels 238 45 283 229 32 261 CO CO + Size Class in Bedroom Units Small— 5 to 19 B.U . Medium — 20 to 99 B.U . . . . Large — 100+ B.U . . 1968 Totals . Small— 5 to 19 B.U . Medium — 20 to 99 B.U . Large — 100+ B.U . 1961 Totals . Gains in 8 years . 1970] Monthey — Wisconsin’s Tourist-Lodging Industry 85 Figure 5. The Regional Distribution of Motel-type Establishments (all sizes) in Wisconsin for 1961 and 1968. establishments has been fairly uniform among the seven regions. However,, the increased motel capacities in 1968 (see regional B.U. totals) clearly indicate that the largest gains were in Regions II and III (East Central and Southeast), where over two-thirds of the 6,900 additional B.U. were located. The three southernmost regions (II, III, and IV) contained well over half of the hotel and motel establishments in Wisconsin, and about two-thirds of the hotel-motel B.U. as well, in 1968. These same regions also showed the greatest gains in motels and the largest losses in hotels during the 8-year period. Resort-type Establishments Resorts, despite a substantial decline in numbers, continued to make up 60.1% of all T-L establishments in 1968 compared to 61.7% 8 years earlier. In 1961, 83.0% of all seasonal T-L busi¬ nesses were of the resort type; by 1968 this percentage was still 82.1%. In 1961 resorts provided 50.4% of all B.U. in the State, compared to 45.5% eight years later. However, the great bulk of these were seasonal units— -32,800 out of 36,200 B.U. — 90.6% in 1968 com¬ pared to 92.1% in 1961. Resort-type establishments continue to be highly seasonal in Wis¬ consin, despite many promotional efforts and overtures— -largely by state and regional tourist organizations and agencies — to encourage and expand year-round housing for visitors. In 1961 seasonal re¬ sorts comprised 93.2% of all resort establishments, and the per- 86 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 centage was exactly the same in 1968. Thus, less than 7% of all resort-type businesses are open for 9 months or longer each year, and these are generally located at or near winter-sports facilities. Because resort-type businesses are considerably smaller than hotel-motel establishments, on the average, a different size classi¬ fication is used in this study to show what has taken place. This breakdown (together with the 1961 and 1968 totals) is as follows: Size Class B.U. Range 1961 Totals 1968 Totals Small 1 to 9 B.U. 3,202 Medium 10 to 29 B.U. 1,468 Large 30 or more B.U. 91 2,708 1,301 92 Although these categories show the general trend in resort-type establishments between 1961 and 1968, they may not give an ade¬ quate picture of what is taking place within the so-called “resort industry” in recent years. Thus, a more detailed size breakdown for resorts is given in Tables 7-a and 7-b. Small Resorts Are Declining When we separate the “small” resort group into the l-to-4 B.U. size — which we term non-commercial establishments — and the 5-to-9 B.U. size, some important differences are readily seen. They are depicted in Tables 7-a and 7-b, which classify the Wisconsin resort inventory by size and seasonality for 1961 and 1968, includ¬ ing a breakdown of the B.U. provided by these establishments and the appropriate totals. For the small-resort category as a whole, there was a loss of 2,473 bedrooms — a drop of 16.5% in capacity — during the 8-year period. However, whereas the l-to-4 class lost only 10% or 455 B.U., the 5-to-9 group dropped nearly 20% or 2,017 B.U. Similarly, the non-commercial group declined only 11% in number of businesses, while the 5-to-9 class dropped slightly over 20%. In some areas the l-to-4 class of establishment actually increased in numbers, and it is reasonable to assume that many of these “businesses” are actually recently-built, private cottages or summer houses that are rented out during the prime vacation months of July and August. Of the 4,639 B.U. lost by small- and medium-size resorts-— those with less than 30 B.U. per firm- — 3,886 or 83% were from establish¬ ments in the 5 to 19 B.U. range. Apparently the resorts in this size bracket are closing down more rapidly than those in either the l-to-4 class or the 20-plus group, possibly because they have the least to offer in the way of economic returns. Yet the percentage of Wisconsin resort establishments in the l-to-19 B.U. range has 1970] Monthey — Wisconsin’s Tourist-Lodging Industry m o H CQ ◄ m fa £ H § w m oo TH <£> 3 05 9^ W O H fa H fa fa fa fa fa o m &a & c/) 8 o oo O cc — ' O sD O c^n 00 cc o T> C 0 CO £ £ o oi 03 T5 bD £ o> CO to sO Table 7-b. Classification of Wisconsin Resort-type Establishments as to Size, Seasonality, and B.U. capacities for 1961. Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 8 co oC ,670 ,532 ,232 236 00 cn 00 6 Z D CQ 309 737 vr\ — < O 00 674 265 13 4-J H ir\ Ij'N O' U-s CN O vO O 00 c O ^ C o ft 4-1 o o o C^\ i—i 1970] Monthey — Wisconsin's Tourist-Lodging Industry 89 declined but little — from 93.1% to 92.3%— since 1961. In terms of total B.U. at resorts, this group still provided nearly 74% of our resort capacity in 1968. Unfortunately, there has been no significant increase in either the medium-size or the large-size categories of resort-type busi¬ nesses to offset the decline in small establishments. Thus, the loss in small firms and their B.U. capacities tends to approximate the total loss for the resort industry as a whole. Resort Distribution Figure 6 shows the regional distribution of all Wisconsin resort establishments in 1961 and 1968. In both years, over 60% of the resort enterprises and about two-thirds of the total resort B.U. were located in the two northern regions. In fact, Region VII (the Northeast) alone has long had well over one-third of Wisconsin's resorts and almost 40% of the total resort B.U. capacity in the entire state. Figure 6-a shows the geographic distribution of all resort-type businesses which operate on a year-round basis (open 9 months or longer each year) . The number of establishments in this group has not increased in recent years. In fact, the total of these “all year” resorts has dropped from 322 to 276 since 1961. However, the total B.U. capacity of these enterprises has increased about 6.0% — from 3,203 B.U. to 3,402 B.U. over the 8 years. Figure 6-b depicts the 1961-68 regional distribution of small re¬ sorts with less than 10 B.U., which comprise about two-thirds of Figure 6. The Regional Distribution of Resort-type Establishments (all types and sizes) in Wisconsin for 1961 and 1968. 90 Wisconsin Academy of Sciences, Arts and Letters [Vo-1. 58 Figure 6-a. The Regional Distribution of Year-round, Resort-type Establish¬ ments (all sizes) in Wisconsin for 1961 and 1968. all resort-type establishments in Wisconsin. Both the number of businesses and the total B.U. are shown for each of the seven re¬ gions. It is noteworthy that in both years, over 55% of the small resorts and over 60% of the units they provide are located in Re¬ gions VI and VII of Northern Wisconsin. These are also the regions in which the greatest losses of small resorts have occurred. Figure 6-c details the regional distribution of medium-size (10 to 29 B.U. class) resorts in Wisconsin for the years 1961-68. Over 70% of these establishments were located in the two northern re- Figure 6-b. The Regional Distribution of Small Resort-type Establishments (less than 10 B.U.) in Wisconsin for 1961 and 1968. 1970] Monthey — Wisconsin’s Tourist-Lodging Industry 91 Figure 6-c. The Regional Distribution of Medium-size, Resort-type Establish¬ ments (10 B.U. to 29 B.U.) in Wisconsin for the years 1961 and 1968. gions during both 1961 and 1968. The heaviest losses in medium- size resorts occurred in these two regions and in Region III (South¬ east) during the 8-year period. Figure 6-d reflects the distribution of Wisconsin’s largest resorts (those with 30 or more B.U.) in 1961 and 1968. As can readily be noted, both the number of establishments and the B.U. capacities are relatively small. Except for some changes among the regions, there have been no major gains or losses during the 8 years studied. Perhaps the most significant change has been in the average size of Figure 6-d. The Regional Distribution of Large Resort-type Establishments (30 B.U. and over) in Wisconsin for 1961 and 1968. 92 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 establishment, which has gone up about 4.5 B.U. per firm. Over one-half of these larger resort-type establishments and about 50% of the total B.U. capacity they provide are situated in the two northern regions. However, the major increase in room capacity since 1961 has occurred in Region III (SE). Other T-L Establishments All other types of T-L establishments, particularly those not tallied as hotels, motels or resorts, are categorized and totalled in Table 8 for the years 1961 and 1968. This catch-all group is classi¬ fied as to size, seasonality, and B.U. capacity. Since the average size of establishment is relatively small, the size classification used is identical with that used for resort-type establishments. This “Other” category includes such diverse types as tourist homes, rooming houses, seasonal dormitories (often called “camps”), dude ranches, vacation farms, clubs, taverns, truck stops, and a variety of other lodgings. Over two-thirds of them have less than 5 B.U., and the bulk of these very small ones are private homes with one or more rental bedrooms. Table 8 shows a loss of 249 “other” T-L establishments between 1961 and 1968, and well over 90% of these were in the small (1 to 9 B.U.) category. Taken together, these businesses represented a loss of 888 B.U. However, this twas more than offset by a gain of 1,189 B.U. from additional establishments in the large category. Figure 7 shows the regional distribution of Other T-L estab¬ lishments in Wisconsin for 1961 and 1968, including the number of Figure 7. The Regional Distribution of Other Types of T-L Establishments (all sizes) in Wisconsin for 1961 and 1968. Table 8. Classification of Other Tourist-Lodging Establishments in Wisconsin as to Size, Seasonality and B.U. Capacity in 1961 and 1968. 1970] Monthey — Wisconsin’s Tourist-Lodging Industry 00 co 00 ^ N. t\ sO O' \D sO O' CO 00 lr\ cc cc O r~* 0" t\ 00 o O' co — i co CO rC CO CO 00 O c-n cc — i cc CO OON —i O' co 00 CO ITS sD — 00 ^ O r<» — ■ O' Vn (V-S , vr\ O s O O' co — 1 \0 vr\ O' tr\ *—t O' < — i irs O' t\ O 00 O' sD co o sD CT\ O CO O' CO o 00 tZ 0 © CO >TN O' 7 00 O O O' — sD — ' — ■ 5 © © CN tC + + cO O O' — DO D T5 DD D TS DO CQ CQ' G CO CQ CQ CQ c CO ^ O' O' CO b ^ O' O' CO b 0 0 o 4~> CQ o o O -H CQ *“H tTN o o cc *— * o o cc "3 ■O ~0 2 CO D £ 00 7 >2 Blocky red clay with subangular boulders up to 10 T in diameter Thin layers of buff-colored sandy clay Large red clay balls imbedded in well sorted sand Buff-colored fine sand Large red clay balls in buff-colored sand Well stratified coarse sand and small pebbles Small red clay balls in sand Alternating layers of fine, clayey sand and stratified coarse sand Figure 2. 153). The advancing Valderan ice later eroded these red silts and clays, mixed them with the underlying materials, and deposited them as the red clayey till on the uplands of eastern Wisconsin (6) . In front of the advancing Valderan ice the lake level of the Lake Calumet stage of Glacial Lake Chicago rose to 620 feet (7). Dur- 156 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figure 3. Clay Balls in Stratified Sands and Silts under Valderan Till, Two Rivers, Wisconsin. ing the retreat of the Valderan ice, extensive marginal lakes formed in which the red glacio-lacustrine sediments, also found in this area, were laid down (2, Petersen and others, p. 187). Many of the glacial events of Eastern Wisconsin have been re¬ constructed through the study of the Two Creeks Forest bed site at Two Creeks, Wisconsin, eleven miles north of the area of this study. Wilson suggested (8) and others confirm (9) that the Two Creeks Forest grew in tins area on mid-Wisconsin till and lake deposits during the low stage of Lake Calumet, called Lake Bow¬ man- ville (7), which probably stood at the 580 feet level (1, Thwaites and Bertrand, p. 859). Readvance of the ice in the Val¬ deran glacial substage caused the lake to flood the Two Creeks site and bury the forest in stratified clays, silts, and sands. Subsequent 1970] Zakrzewska — Clay Ball Deposits in Wisconsin 157 Figure 4. Typical Clay Ball in Stratified Sand under Valderan Till, Two Rivers, Wisconsin. advance of the ice broke the trees and covered the site and the surrounding area .with almost 10 feet of red till. According to Th waites and Bertrand (1) and Hough (9, p. 98), well developed Calumet stage level beaches have not been discovered in Eastern Wisconsin. This paper reports on field evidence of what is probably a buried beach of the Calumet II stage of Glacial Lake Chicago. As the clay balls found near Two Rivers occur in several layers in generally undisturbed stratigraphic sections underlying Valderan till (in which we found a large tree log probably derived from the nearby Two Creeks Forest bed), it can be suggested that in this locality the advancing Valderan ice was fringed by a shallow mar¬ ginal lake or a lagoon formed behind offshore sand bars. As a re¬ sult of wave erosion or melting along the advancing Valderan ice, 158 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 small chunks of clayey Valderan till were brought to this marginal lake. The high degree of cohesion and compactness of the till, or its frozen state, could have kept the clay chunks from disintegrat¬ ing. In the shore environment the chunks of till were probably rolled on the beach by successive waves until they acquired a high degree of sphericity. Evidence supporting the suggestion that the clay balls may represent beach environment was found at the nearby Point Beach State Park. Red clay balls are present on the inner part of the beach where red clay (probably from parking lot construction) projects through the sand. These balls must have been formed recently by waves which reached the clay bed, de¬ tached segments of it, and rolled them into round balls. Shallow lakes or lagoons with waves of smaller magnitude could have pro¬ vided a more suitable environment for clay balls to be preserved in successive layers of sands and silts varying in thickness from several inches to a few feet. Xntermittently-dry lagoon bottoms could have also provided thick flakes of red clay curling up between dessication cracks, easy to roll into oval clay balls when washed over by small waves. However, while several elongated clay balls composed of layered rather than blocky clay have been found at the site of study, most balls are composed of blocky red clay containing small pebbles. The present topographic setting of the study site further con¬ tributes to the understanding of the paleogeographic conditions in the area during the formation of the red clay balls. The mile-long north-south trending ridge in which the clay balls are found reaches an elevation of about 640 feet above sea level (see Fig. 1) and is located one and a half miles west from the present shore of Lake Michigan. It rises about 30 feet above a gently sloping plain whose eastern edge is marked by a north-south trending 600 feet contour line interpreted to be an abandoned shoreline of Glacial Lake Nipissing (1, Thwaites and Bertrand, Plate 8). Immediately east of this shoreline lies the Point Beach State Park consisting of a broad overgrown lagoon fringed on the east by crescent-shaped, north-south trending, parallel alternating lagoons and sand bars forming a peninsula-like projection into Lake Michigan. These ridges and lagoons are of post-glacial origin and are not covered by glacial till. The north-south trending ridge in which the buried clay balls are found may be a remnant of a beach ridge analogous to the present beach ridges located to the east of it. On the basis of its altitude, the ridge may be interpreted to be a part of the Glacial Lake Calumet II shoreline, whose elevation is estimated at 620 feet (7). The maximum elevation of the ridge is just over 640 feet, and the elevation of the buried beach ridge, overlain in places by as much as 10 feet of Valderan till, is not much over 630 feet. 1970] Zakrzewska — Clay Ball Deposits in Wisconsin 159 The clay ball deposits are located below that elevation, or around 620 feet. Since the beach ridge is buried by the Valderan till, the lake whose shoreline it represents must have existed just prior to the maximum extent of the Valderan ice. Wayne and Zumberge summarize the chronology of glacial lakes in Lake Michigan Basin as shown in Table 1. Inasmuch as the clay balls are found in the upper strata of water- deposited sands and silts which appear to be of Two Creeks age and are overlain by the red Valderan till, they had to be formed at the end of the Two Creeks period but before the maximum ad¬ vance of the Valderan ice. It is therefore suggested that they rep¬ resent a beach ridge of Glacial Lake Calumet II. A search for other sites containing layers of clay balls which would further help interpret the site described was conducted through a 75 square mile area around Two Rivers (in ten sand and gravel pits), but similar beach-ridge deposits were not found. A distinct layer of red clay balls was found at only one other place : Table 1. Years B. P. Name of Lake and Elevation in Feet 2,000 . 3,000. . 4,000 . Lake Michigan (580) Lake Algoma Lake Nipissing (slow crustal uplift due to glacial unloading) 9,500 . 11,000 . Chippewa (230) Post Algonquin Main Algonquin Kirkfield Valders Maximum . Tolleston (605) Calumet II (620) Two Creeks . Bowmanville 12,000 13,000 . 14,000 . Calumet I (620) Glenwood (640) Source: W. Wayne and J. Zumberge, The Quaternary of the United States, p. 7b. 160 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 a small sand and gravel pit on a gently rolling upland immediately adjacent to and south of West Twin River valley, just west of Shoto (Sec. 29, T19 N, R24 E) . The site is located seven miles from the shore of Lake Michigan at an approximate elevation of 620 feet. The layer of clay balls found here lies below 620 feet and is imbedded predominantly in gravel deposits, overlain by about five feet of well sorted sands, and about two feet of buff-colored till containing ice wedges. A few small clay balls are present in the lowest layers of the sand, but the underlying gravel deposits con¬ tain most of the red clay balls, which are of varying sizes up to 7 inches in diameter and have an occasional distinct pebble armor around them. The clay balls in this site are less consistently round ; many are oval or cigar-shaped with a flat ring suggesting rolling in one direction. Inasmuch as red till is not present in this area, though the area is mapped as Valderan (6, p. 170), the red clay balls were probably brought into this locality with outwash gravels from an area of red Valderan till and, therefore, differ in origin from the clay balls at Two Rivers. Furthermore, the clay balls at Shoto do not occur in a distinct ridge parallel to the shores of Lake Michigan beach. Therefore, there is little to suggest at present that they represent a beach ridge environment such as that at Two Rivers. A few scattered red clay balls were also found near Sheboygan, which is located about 35 miles south of Two Rivers, but the topo¬ graphic and stratigraphic conditions again were different from those at Two Rivers. Horn (2, p. 174) found one clay ball south¬ west of Omro, west of Lake Winnebago, imbedded in sand under 20 feet of stratified material covered by 6 to 12 feet of red Valderan till. However, distinct clay ball layers, such as those at Two Rivers, were not reported. All these sites, therefore, though containing clay balls, are not comparable to the site at Two Rivers and do not con¬ tribute directly to its interpretation. Further search may, however, reveal sites which are comparable to that at Two Rivers. In summary, the field findings presented in this paper suggest that marginal lakes, probably with fluctuating water level, fringed the advancing Valderan ice near Two Rivers. The clay ball deposit is interpreted to be a beach ridge of Glacial Lake Calumet II, which was present in this area just prior to the maximum advance of the Valderan ice. The deposit appears to be a beach deposit because of the presence of distinct layers of highly spherical clay balls im¬ bedded in stratified sands and silts, similar to clay balls found on nearby modern beaches, and because of the topographic position of the clay balls in a north-south trending ridge roughly parallel to the shores of Lake Michigan basin. The deposit is inferred to rep- 1970] Zakrzewska — Clay Ball Deposits in Wisconsin 161 resent a beach ridge of Glacial Lake Calumet 11 on the basis of its geographic position, its specific elevation, and, through its strati¬ graphic position, its place in the chronology of geomorphic events of this area. This conclusion assumes that the sequence of geo¬ morphic events proposed in the literature reviewed (7) is correct. Further search around Lake Michigan for sites containing layers of clay balls imbedded in beach deposits under Valderan till may help determine whether the deposit described in this paper rep¬ resents the shoreline of Glacial Lake Calumet II, and may help map this shoreline. Findings of this nature should help reconstruct the paleogeography of Eastern Wisconsin during the most recent ad¬ vance of the continental ice into this area (10). References Cited 1. Alden, W. C., “Quaternary Geology of Southeastern Wisconsin,” U.S.G.S. Prof. Paper 106 (1918) ; Thwaites, F. T., “Pleistocene of Part of North¬ eastern Wisconsin,” G.S.A. Bull. 54, 84-144 (1943) ; Frey, J. C., and Willman, H. B., “Classification of the Wisconsin Stage in the Lake Mich¬ igan Glacial Lobe,” III. G. S. Circ. 285, 1-16 (1960) ; Wright, H. E., Jr., and Frey, D. C., (ed.), The Quaternary of the United States (Prince¬ ton Univ. Press, 1965) ; Thwaites, F. T., and Bertrand, K., “Pleistocene Geology of the Door Peninnsula, Wisconsin,” G.S.A. Bull. 68, 831-880 (1957) ; Black, R. F. and M. Rubin, “Radiocarbon Dates of Wisconsin,” Trans. Wise. Ac. Sci. LVI, 91-115 (1967-68). 2. Murray, R. C., “The Petrology of the Cary and Valders Tills of North¬ eastern Wisconsin,” Amer. Jour. Sci. 251, 140-155 (1953) ; Horn, M. E., “A Pedological Study of Red Clay Soils and Their Parent Materials in Eastern Wisconsin,” Ph.D. Thesis (Univ. of Wise., 1960) ; Lee, G. B., Janke, W. E., and Beaver, A. J., “Particle Size Analysis of Valders Drift in Eastern Wisconsin,” Science 138, 154-155 (1962) ; Janke, W. E., “Characteristics and Distribution of Soil Parent Material in the Val¬ deran Drift Region of Eastern Wisconsin,” Ph.D. Thesis (Univ. of Wise., 1962) ; Petersen, G. W., Lee, G. B., and Chester, G., “A Comparison of Red Clay Glacio-Lacustrine Sediments in Northern and Eastern Wisconsin,” Trans. Wise. Ac. Sci. LVI, 185-196 (1957-68). 3. Bell, H. S., “Armored Mud Balls — Their Origin, Properties, and Role in Sedimentation,” Jour. Geol. 48, 1-31 (1940). 4. Leney, G. L., and Leney, A. T., “Armored Till Balls in the Pleistocene Outwash of Southeastern Michigan,” Jour. Geol . 65, 105-106 (1957). 5. Kugler, H. G., and Saunders, J. B., “Occurrence of Armored Mud Balls in Trinidad, West Indies,” Jour. Geol. 67, 533-565 (1959). 6. Black, R. G., “Valders Glaciation in Wisconsin and Upper Michigan — A Progress Report,” Univ. of Michigan Great Lakes Research Oivision (1966), pp. 169-175, suggests that these deposits are present only in northeastern Wisconsin, though originally they were also recognized in northwestern Wisconsin and Upper Michigan. 7. Wayne, W. J., and Zumberge, J. H., “Pleistocene Geology of Indiana and Michigan,” in Wright and Frey, op. cit., (footnote 1), Fig. 6, p. 76. It is apparent from Bretz, J. H., “The Double Calumet Stage of Lake Chicago,” Jour. Geol. 67, 675-684 (1959) that he and Hough (footnote 162 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 9) do not agree on the interpretation of Glacial Lake Chicago. For this reason the correlations compiled by Wayne and Zumberge from writings by both Bretz and Hough are used in this paper. 8. Wilson, L. R., “The Two Creeks Forest Bed, Manitowoc County, Wiscon¬ sin,^ ” Trans . Wise. Ac. Sci. 27, 31-46 (1932) ; and “Further Fossil Studies of the Two Creeks Forest Bed, Manitowoc County, Wisconsin,” Torrey Bot. Club Bull. 66, 317-325 (1936). 9. For example: Hough, J. L., “The Prehistoric Great Lakes of North America,” Amer. Scientist 51, No. 1, 100 (1936) ; or Wayne, W., and Zumberge, H., op. cit., pp. 73-80; or Thwaites, F. T., and Bertrand, K., op. cit., pp. 859-860. 10. I thank Professors K. Nelson and N. Lasca of the Department of Geol¬ ogy, University of Wisconsin-Milwaukee, for their comments on the ideas contained in the early version of this paper, and Professor J. Flannery and the UWM Cartographic Laboratory for the preparation of illustrations. Figure 4 was provided by Mr. Charles Collins. NOTES ON THE ECOLOGY OF THE HARVEST MOUSE, REITHRODONTOMYS MEGALOTIS, IN SOUTHWESTERN WISCONSIN Gerald E. Svendsen Introduction The harvest mouse, Reithrodontomys megalotis , occurs through¬ out most of the central and western United States and Mexico. It is the most widespread of all species of harvest mice, preferring thick stands of grass (Hall and Kelson, 1959). Local abundance depends on a variety of vegetation that provides food at all sea¬ sons. Uncultivated fields and areas having grasses bearing large seeds are favored. In Wisconsin, as far as is known, the distribu¬ tion of this species is limited to the driftless region, and it favors more or less open, grassy, neglected fields, grassy borders of cul¬ tivated fields, and grain fields (Jackson, 1961). The first specimens of the harvest mouse collected in Wisconsin were taken in La Crosse county in 1930 by Vernon Bailey, and Francis Hammerstrom procured a specimen from Juneau county in 1936 (Jackson, 1961). Subsequent specimens were collected from Columbia and Sauk counties by Hansen (1944a). Personal records include specimens from La Crosse, Wood, and Vernon coun¬ ties. Hansen (1944a) and Jackson (1961) recognize this harvest mouse to be Reithrodontomys megalotis pectoralis, but Hooper (1952) and Hoffmeister and Warnock (1955) find it indistinguish¬ able from Reithrodontomys megalotis dychei. This paper presents some observations on habitats used by this animal, population densities, and a record of some relationships with associated species in southwestern Wisconsin. Methods and Materials Population and vegetation analysis were determined in a 9 acre field which has not been under cultivation for 8 years. This neg¬ lected field is bordered on the north by a wooded area, on the east by a wet marsh, and on the west and south by a 3 track railroad right-of-way and cultivated fields planted in soybeans and corn. The abandoned field is essentially isolated from any other area of similar vegetational composition. A snap trap grid was arranged 163 164 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 in 1 acre plots with traps stationed every 30 feet. The grids were run for six consecutive nights and the population estimates were made from data based on the removal of the animals (Zippen, 1958). Vegetational analysis was accomplished by randomly selecting five 1 meter plots within the grids and determining species compo¬ sition. Light intensity was measured with a light meter at noon on a clear sunny day, and expressed as percent of overhead light. Habitat Preference Trapping efforts in the wooded area, the marsh, the railroad right-of-way, and in the cultivated fields yielded no harvest mice. The only habitat from which the harvest mouse was collected was the abandoned field. This field is in an early stage of succession, with sapling elm, oak, and sumac (%-! inch d.b.h.) at a density of 3 to 5 per acre. The ground cover is dense, total foliage cover is 90 percent. Light intensity at ground level is 15 to 20 percent of full sunlight. The areas of sparse vegetation are due to places where animals’ digging and denning activities have brought up large quantities of sand. Seventy-two percent of the total species of plants are the grasses Phleum, Agropyron, and Panicum, and the legumes Lespedeza and Trifolium. The forbes Aster, Asclepias, Aplopappas and Solidago, the grasses Andropogon, Elumus and Setaris, the blackberry Eubatus, and the composites Tragopogon and Taraxacum make up 23 percent of the plants. The remaining 5 percent of the species are woody plants and other grasses and forbes. Average height of the vegetation varies from 12 to 20 inches, with no ground litter over most of the area. The soil is a sandy loam with very good drainage. The mice live mainly in burrows. Population Density of Reithrodontomys The population density of Reithrodontomys megalotis was meas¬ ured in the falls of 1967 and 1968 in the abandoned field where previous trapping indicated the presence of a substantial popula¬ tion of this species. The fall population, estimated by the removal method, was 18 animals per acre. This represents 0.046 animals per trap night. The spring trapping yielded 0.012 animals per trap night. The fall population based on animals per trap night is almost four times that of the spring population. Hansen (1945) estimated a maximum density of harvest mice in foxtail-smartweed cover type as 2.4 per acre. Birkenholz (1967) estimated a fall population of 17 animals per acre in central Illinois. 1970] Svendsen — Ecology of the Harvest Mouse 165 Associated Species The harvest mouse is found in the same area with a variety of other small animals. Peromyscus maniculatus is the only species which outnumbers the harvest mouse. The estimated density of Peromyscus maniculatus is 72 animals per acre, representing 56 percent of the total population of animals in the old field study area. The percent of the total population represented by the other animals is Reithrodontomys megalotis 14 percent, Blarina brevi- cauda 10 percent, Microtus pennsylvanicus 7 percent, Mus musculus 4 percent, Peromyscus leucopus 4 percent, Zapus hudsonius 3 per¬ cent, Bor ex cinerus 1 percent, and Spermophillus tridecemlineatus 1 percent. Eleven percent of the total population is composed of two species of predators Blarina brevicauda and Bor ex cinerus , the former be¬ ing the most common. Peromyscus leucopus were all captured not more than 30 feet from the edge of the woods. These individuals were probably wanderers from the woods rather than permanent residents of the field. Microtus were trapped mainly in areas of the field where lespedeza and clover were especially thick, and in¬ frequently over the rest of the field. Mus, Zapus, and Peromyscus were continually trapped with the harvest mouse, especially in areas of the field where the seed-bearing plants were more com¬ mon. These four species appear to compete directly with one an¬ other. Zapus hibernates from late October to May and at this time would be removed from competition, and Catlett and Shellhammer (1962) suggest that Mus and Reithrodontomys form a cospecies social hierarchy and that little competition exists between these species. Peromyscus maniculatus appears, therefore, to compete most strongly with the harvest mouse throughout the year in this study area. Discussion It can be assumed that clearing of woodlands and establishing grasslands and grainfields has aided the distribution of the harvest mouse. Birkenholz (1967) reports an eastward range extension in Illinois during the past century in response to the clearing of wood¬ lands. The harvest mouse populates new habitats suddenly and attains a high local abundance if the habitat alteration favors its establishment. These habitats are usually transitory resulting in a decrease in abundance if the vegetation composition becomes less favorable. The harvest mouse can become established in these local and transitory habitats in two ways. Individuals from widely distri¬ buted but small and relic populations can populate the new areas, 166 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 or a saltatorial colonization can occur from isolated, well-populated pockets. Saltatorial colonization by vertebrates is considered un¬ likely by Smith (1957), although Leopold (1947) reports that out¬ lying individuals are common in species in the process of expand¬ ing their range. Five years of intensive trapping of all types of habitats has yielded only one harvest mouse from an area which could be con¬ sidered nontypical, and Hansen (1944) reported catching one indi¬ vidual on a juniper bluff. The trapping data would, therefore, favor saltatorial colonization of new areas, due to the lack of indi¬ viduals, representing a scattered population, in nontypical habi¬ tats. It is reasonable that saltatorial colonization can take place if the harvest mouse has a tendency to wander, and if it has a low degree of motivation to return home. Fisler (1966) reported that the homing tendencies of the harvest mouse would be best described as nonrandom throughout the terrain and that the mice lack motivation to return home from unknown areas. References Cited Birkenholz, Dale E. 1967. The harvest mouse (Reithrodontomys megalotis) in Central Illinois. Trans. Ill. State Acad. Sci., 60, No. 1. Catlett, R. H., and H. S. Shellhammer. 1962. A comparison of behavorial and biological characteristics of house and harvest mice. J. Mamm. 43: 133-144. Fisler, George F. 1966. Homing in the western harvest mouse, Reithrodon¬ tomys megalotis. J. Mamm. 47 : 53-58. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. The Ronald Press Co., New York. 1083 p. Hansen, H. C. 1944a. A new harvest mouse from Wisconsin. Field Mus. Nat. Hist., Zool. Ser. 29: 205-209. Hansen, H. C. 1945. Small mammal census near Prairie du Sac, Wisconsin. Trans. Wisconsin Acad. Sci., 47: 161-164. Hooper, Emmet T. 1952. A systematic review of the harvest mouse (Reithrod¬ ontomys) of Latin America. Misc. Publ. Mus. Zool., Univ. Mich., no. 77. Jackson, H. H. 1961. Mammals of Wisconsin. Univ.'of Wisconsin Press, Madi¬ son, 504 p. Leopold, A. 1947. The distribution of Wisconsin hares. Trans. Wise. Acad. Sci., Arts and Letters, 37 : 1-14. Smith, P. W. 1957. Analysis of post-Wisconsin biogeography of the prairie peninsula region based on distribution phenomena among terrestial verte¬ brate populations. Ecol. 38 : 205-219. Zippen, C. 1956. An evaluation of the removal method of estimating animal populations. Biometrics, 12 : 163-189. AN ANNOTATED CHECK LIST OF THE GEOMETRIDAE (LEPIDOPTERA) OF WISCONSIN1 Charles V. Coveil, Jr. Upon completing the task of identifying some 1,700 undetermined specimens of Geometridae for the University of Wisconsin Insec- tarium, I felt that a Wisconsin list of this family of moths would be useful to lepidopterists. To this end I examined additional mate¬ rial: 1,300 more specimens from the University of Wisconsin; 1,800 specimens from William E. Sieker; and over 2,000 specimens taken by Harold Bower. Over 7,000 specimens provided the data in this list, and localities in at least 30 counties are represented. Previous check lists including Wisconsin Geometridae are those of Rauterberg (1900), Fernekes (1906), and Muttkowski (1907), all of which are restricted to Milwaukee County. I have not in¬ cluded their records here because of subsequent changes in nomen¬ clature of species, and because some of their identifications are dubious (e.g., Fernekes, p. 54, lists Melcmchroia cephise Cramer, a Florida species). However, most of their records are corroborated here; and other species are recorded from Wisconsin for the first time. Although this list is based upon specimens I have actually ex¬ amined, a few records are included which were taken from recent revisionary literature. These cases are so noted in the text, and appropriate references are cited. The nomenclature and arrangement of taxa follow generally those of McDunnough (1938) , since that list has long been the basis for organization of most collections of North American moths. I have used Forbes (1948) and the revisionary works of McDun¬ nough, Rindge, Capps, Rupert, and others as aids in my determina¬ tions; consequently some digressions from McDunnough (1938) have been made here. Numerous genera, such as Semiothisa, badly need revisionary study. It therefore follows that certain long-stand¬ ing errors in nomenclature are unfortunately continued in this list. These must wait for future correction. Most of the specimens studied were collected by the following, each listed with the localities where he made most of his captures : Harold Bower (Lake Katherine in Oneida County, Milwaukee, and 1 University of Louisville Publications in Biology (New Series) No. 110, 167 168 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Wausau) ; Louis Griewisch (Green Bay and other “Brown County” labels) ; Gary Lachmund (Sauk City) ; Gary Ross (Florence County) ; and William E. Sieker (Bailey’s Harbor in Door County, other “Door County” labels, and Madison). All of the Bower speci¬ mens are in the Los Angeles County Museum, and most of the Sieker and Lachmund material is in my collection; the rest is at the University of Wisconsin. I am deeply indebted to the following for their contributions to this effort: Dr. Roy D. Shenefelt, Curator of the University of Wisconsin Insectarium, and his project assistants, Lutz J. Bayer and Isabel Arevalo, for providing material and important informa¬ tion; William E. Sieker of Madison and Gary Lachmund of Sauk City for numerous specimens; Lloyd M. Martin, Los Angeles County Museum, for making the Bower collection available; Dr. Frederick H. Rindge, American Museum of Natural History, for nomenclatural advice; and Patricia K. Liles, my curatorial assist¬ ant, for compiling data. I am also very grateful to Dr. John A. Dil¬ lon, Jr., Dean of the Graduate School, University of Louisville, for making graduate school funds available to me for the visit to the Los Angeles County Museum. Family Geometridae Subfamily Brephinae 1. Brephos infans Mbschler Apr. 10-May 7 Lake Katherine, Oneida Co. 2. Leucobrephos brephoides Walker Synonym hoyi Grote was described from Wisconsin. Subfamily Oenochrominae 3. Alsophila pometaria Harris Apr. 18; Oct. 25-Nov. 5 Crandon; Madison; Wausau, Marathon Co. Subfamily Geometrinae 4. Nemoria mimosaria Guenee May 14- July 11 Bailey’s Harbor; Florence Co. ; Griffith State Nursery, Wood Co.; Lake Katherine; Marinette Co.; Wausau. 5. Nemoria rubrifrontaria Packard May 20- June 27 Lake Katherine; Sauk City; Vilas Co. 6. Dichorda iridaria Guenee May 31- June 7 Door Co.; Sauk City. 7. Synchlora liquor aria Guenee Lake Katherine. This is, according to Ferguson (1969), subspecies albolineata Packard. 1970] Coveil — Geometridae of Wisconsin 169 8. Synchlora aerata Fabricius June 11-Aug. 27 Columbia Co.; Lake Katherine; Madison; Trout Lake, Vilas Co.; Sayner; Wausau. These records include rubrifrontaria Packard, synonymized by Ferguson (1969). 9. Chlorochlamys chloroleucaria Guenee June 4-Sept. 8 Belleville; Columbia Co.; Door Co.; Green Bay; Harrisville; Madison; Rusk Co.; near Sayner, Vilas Co.; Wausau. 10. Hethemia pistaciaria Guenee June 11-July 10 Florence Co. ; Lake Katherine. 11. Mesothea incertata Walker May 10-30 Cranmoor, Wood Co.; Lake Katherine; Marinette Co. Subfamily Sterrhinae 12. Metasiopsis balistaria Guenee May 14- July 22 Boscobel St. Nursery, Grant Co.; Dane Co.; Griffith St. Nursery, Wood Co.; Milwaukee; Sturgeon Bay, Door Co. 13. Scopula cacuminaria Morrison June 18-Sept. 13 Lake Katherine; N.E. Price Co.; Wausau. 14. Scopula quadrilineata Hulst (probably June-Aug.) “Wis.” on pin label. 15. Scopula ancellata Hulst Aug. Bailey’s Harbor. 16. Scopula junctaria Walker May 7- July 26 Bailey’s Harbor ; Crandon ; Florence Co. ; Lake Katherine. 17. Scopula limboundata Haworth June 10-Aug. 16 Bailey’s Harbor; Crandon; Dousman; Florence Co.; Lake Katherine ; Madison ; Marinette Co. ; Milwaukee ; Sauk City ; Trout Lake, Vilas Co. Known widely as enucleata Guenee. 18. Scopula frigidaria Mdschler July 3 Florence Co. 19. Scopula inductata Guenee May 30-Sept. 14 Columbia Co.; Door Co.; Gordon Nursery, Douglas Co.; Green Bay; Lake Katherine; Madison; Milwaukee; Sauk City ; Waushara Co. 20. Idaea demissaria Hlibner May 25-July 21 Door Co.; Madison. Fletcher (1966, p. 12) synonymizes the familiar St err ha Hlibner to Idaea Treitschke. 21. Haematopis grataria Fabricius May 31-Oct. 4 Badger Ordnance Works, Sauk City ; Boscobel ; Brown Co. ; Columbia; DeForest; Door Co.; Lake Katherine; Madison; Milwaukee; Rusk Co.; Sauk City; Tower Hill State Park, Iowa Co. ; Washburn Co. ; Waushara Co. ; Wood Co. 170 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 22. Calothysanis amaturaria Walker July 14-Sept. 14 Columbia Co.; Lake Katherine; Madison. 23. Pleuroprucha insulsaria Guenee June 9- Oct. 1 Dane Co.; Door Co.; Lake Katherine; Milwaukee. 24. Cyclophora pendulinaria Guenee May 21-Sept. 4 American Legion State Forest, Oneida Co. ; Lake Katherine ; Rusk Co.; Trout Lake; Wausau. 25. Cyclophora packardaria Prout June 16-Aug. 29 Lake Katherine. Subfamily Larentiinae 26. Acasis viridata Packard May 16-31 Lake Katherine. 27. Nyctobia limitaria Walker May 1-June 9 Lake Katherine. 28. Nyctobia anguilineata Grote and Robinson May 28 Lake Katherine. 29. Cladara atroliturata Walker Apr. 15-May 22 ; Aug. 9 Lake Katherine. 30. Lobophora nivigerata Walker May 18- July 12 Brown Co.; Dane Co.; Florence Co.; Green Bay; Lake Katherine; Sauk City; near Sayner, Vilas Co. 31. Heterophleps refusata Walker June 20- July 12 Florence Co.; Milwaukee; Wausau. 32. Heterophleps trignttaria Herrich-Schaffer June 17-July 11 Madison; Milwaukee; Wausau. 33. Dyspteris abortivaria Herrich-Schaffer June 4-Aug. 11 Madison; Selzer Farm near Holy Hill. 34. Trichodezia albovittata Guenee May 26-July 23 Kewaunee Co. ; Lake Katherine ; Madison ; Milwaukee. 35. Oporophtera bruceata Hulst Oct. 15-Nov. 12 Lake Katherine; Wausau. 36. Triphosa affirmaria Walker Apr. 19-Oct. 28 Dane Co.; Florence Co.; Lake Katherine; Wausau. Known widely as haesitata Guenee. 37. Hydria undulata Linnaeus June 11-Aug. 15 Door Co. ; Dousman ; Florence Co. ; Kenosha Co. ; Lake Katherine; Madison; Milwaukee. 38. Coryphista meadi Packard May-Sept. 10 Dane Co.; Green Bay. 1970] Covell — Geometridae of Wisconsin 171 39. Eupithecia miserulata Grote May 30-Oct. 19 Columbia Co.; Dane Co.; Door Co.; Florence Co.; Lake Katherine; Rusk Co.; University of Wisconsin Arboretum, Madison. 40. Eupithecia castigata Hubner June 12-26 Lake Katherine. 41. Eupithecia paipata Packard Florence Co.; Vilas Co. 42. Eupithecia transcanadata MacKay Lake Katherine. 43. Eupithecia columbiata Dyar Apr. 30-May 24 Lake Katherine. Determined by Bower as subspecies erpata Pearsall. 44. Eupithecia herefordaria Cassino and Swett June 9-Sept. 15 Columbia Co.; Florence Co. 45. Eupithecia carolinensis Grossbeck June 10 Lake Katherine. Determined by R. Leuschner. 46. Eupithecia russeliata Swett May 24- July 27 Lake Katherine. 47. Eupithecia indistincta Taylor July 10 Trout Lake, Vilas Co. 48. Eupithecia coagulata Guenee July 21-Aug. 22 Lake Katherine; Milwaukee; Wausau. 49. Eupithecia perfusca Hulst June 18- July 20 Lake Katherine. Determined by Bower as subspecies youngata Taylor. 50. Eupithecia ravocostaliata Packard Apr. 26-30 Lake Katherine. 51. Horisme intestinata Guenee May 30-Aug. 23 Bailey’s Harbor; Brown Co.; Green Bay; Lake Katherine; Milwaukee; Sauk City. 52. Eustroma semiatrata Hulst June 10- Aug. 4 Crandon; Lake Katherine near Hazelhurst; N.E. Price Co. Known widely as E. nubilata Packard. 53. Eulithis diver silineata Hubner June 24-Aug. 30 Brown Co. ; Florence Co. ; Kenosha Co. ; Madison ; Milwau¬ kee; Sauk City; Wausau. Fletcher (1966, p. 15) cites diver - silineata as type of Eulithis Hubner [1821] which ante¬ dates Lygris Hubner [1825]. 54. Eulithis gracilineata Guenee June 24-Sept. 16 Brown Co.; Griffith State Nursery; Madison. June 9 May 19- June 17 172 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 55. Eulithis testata Linnaeus June 14-Sept. 5 Florence Co.; Lake Katherine; Wausau. 56. Eulithis molliculata Walker June 14-Aug. 20 Milwaukee ; Platteville. 57. Eulithis destinata Mdschler July 3-Aug. 22 Florence Co.; Lake Katherine. 58. Eulithis explanata Walker Florence Co. ; Lake Katherine. 59. Eulithis xylina Hulst Lake Katherine. 60. Diactinia silaceata Hiibner Forest Co.; Lake Katherine. 61. Plemyria georgii Hulst Lake Katherine. June 24-Aug. 25 July 6-Aug. 2 June 14-Sept. 16 Aug. 11-18 62. Dysstroma citrata Linnaeus June 4-Sept. 1 Florence Co. ; Lake Katherine ; Trout Lake. 63. Dysstroma hersiliata Guenee June 19-Aug. 28 Bailey’s Harbor; Crandon; Dodgeville; Door Co.; Florence Co.; Lake Katherine; Washburn Co. ; Wausau. 64. Stamnodes gibbicostata Walker Aug. 31-Sept. 4 Lake Katherine. 65. Hydriomena transfigurata Swett May 11 Dane Co. This is subspecies manitoba Barnes and McDun- nough. 66. Hydriomena perfracta Swett May 6-June 24 Lake Katherine; Vilas Co. near Sayner. 67. Hydriomena divisaria Walker May 6-July 16 Lake Katherine; Vilas Co. near Sayner. 68. Hydriomena renunciata Walker May 11-Aug. 5 Bailey’s Harbor ; Dane Co. ; Door Co. ; Florence Co. ; Green Bay; Griffith State Nursery; Lake Katherine; Milwaukee; Summit Lake, Langlade Co.; Trout Lake; Waterloo; Wau¬ sau. 69. Xanthorhoe lacustrata Guenee May 29-Aug. 10 Door Co. ; Florence Co. ; Lake Katherine ; Madison ; Milwau¬ kee; “Peaks Lake” (this is probably Peat’s Lake, Brown Co.) ; Wausau. 70. Xanthorhoe emendata Pearsall May 21- Aug. 1 Lake Katherine; N.E. Price Co. 71. Xanthorhoe ferrugata Linnaeus May 21-Aug. 29 Bailey’s Harbor; Columbia Co.; Door Co.; Florence Co.; Lake Katherine; Madison; Milwaukee; Wausau. 1970] Coveil — Geometridae of Wisconsin 173 72. Xanthorhoe algidata Mbschler July 10-Aug. 3 Florence Co.; Lake Katherine; N.E. Price Co. 73. Xanthorhoe iduata Guenee June 20-July 8 Lake Katherine. 74. Xanthorhoe abrasaria Herrich-Schaffer June 22-Aug. 23 Columbia Co. ; Florence Co. ; Lake Katherine. 75. Xanthorhoe luctuata Denis and Schiffermiiller May 28-Aug. 1 Forest Co. ; Lake Katherine. 76. Xanthorhoe intermediata Guenee May 17-Oct. 13 Dane Co. ; Door Co. ; Lake Katherine ; Milwaukee Co. ; Spen¬ cer; Spooner; Univ. Wise. Arboretum, Madison. 77. Mesoleuca ruficillata Guenee May 27-Aug. 21 Florence Co.; Lake Katherine; Milwaukee; “Peaks Lake” ; Wausau. 78. Epirrhoe alternata Muller June 10-Aug. 6 Florence Co.; Lake Katherine; Wausau. 79. Spargania magnoliata Guenee June 10-Aug. 12 Crandon; Lake Katherine; N.E. Price Co.; Wausau. 80. Percnoptilota obstipata Fabricius May 3-Oct. 13 Bailey's Harbor ; Columbia Co. ; Florence Co. ; Green Bay ; Grant Co.; Lake Katherine; Madison; Milwaukee; Rusk Co. ; Sauk City ; Spooner ; Sturgeon Bay, Door Co. 81. Percnoptilota centrostrigaria Wollaston May 19-Oct. 31 Bailey's Harbor ; Brown Co. ; Door Co. ; Florence Co. ; Green Bay; Lake Katherine ; Madison; Milwaukee; Sauk City; Washburn Co. 82. Rheumaptera hastata Linnaeus June 11-July 6 Green Bay ; Kewaunee Co. ; Lake Katherine ; Madison ; Mil¬ waukee ; “Peaks Lake.” 83. Perizoma basaliata Walker Lake Katherine; N.E. Price Co. 84. Earophila vasiliata Guenee Florence Co. ; Lake Katherine. 85. Earophila multiferata Walker Lake Katherine; Madison. 86. Venusia comptaria Walker Lake Katherine; Milwaukee. 87. Hydrelia condensata Walker Lake Katherine; Milwaukee. 88. Hydrelia inornata Hulst Florence Co.; Lake Katherine. July 4-30 Apr. 26- June 9 May 17-June 10 Apr. 25-May 19 June 5-July 1 June 10- July 20 174 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 89. Hydrelia albifera Walker July 3-25 Florence Co.; Lake Katherine. 90. Eudule mendica Walker June 10-July 24 Bailey’s Harbor; Florence Co.; Green Bay; Griffith State Nursery; Kenosha Co.; Madison; Mather; Milwaukee; Oconto Co.; “Peak’s Lake”; Sauk City; Trout Lake, Vilas Co.; Wausau. Subfamily Ennominae 91. Bapta semiclarata Walker May 15-July 6 Crandon ; Door Co. ; Marinette ; Washburn Co. 92. Bapta vestaliata Guenee May 29-July 11 Bailey’s Harbor; Blue Mounds; Brown Co.; Florence Co.; Green Bay; Lake Katherine; Madison; Milwaukee. 93. Bapta glomeraria Grote May 3-31 Lake Katherine; Waupaca. 94. Cabera quadrifasciaria Packard June 7-24 Green Bay; Sauk City. Fletcher (1966, p. 12) shows Deilinia Hiibner (1825) to be a junior objective synonym of Cabera Treitschke (1825). 95. Cabera variolaria Guenee June 20-Aug. 12 Bailey’s Harbor; Brown Co.; Florence Co.; Forest Co.; Madison; Milwaukee; N.E. Price Co. 96. Cabera erythemaria Guenee May 28-Aug. 26 Bailey’s Harbor; Brown Co. ; Florence Co. ; Green Bay; Mil¬ waukee; N.E. Price Co. 97. Apodrepanulatrix liber aria Walker Rindge (1949, p. 293) states that this species ranges “to Wisconsin.” 98. Syrrhodia cruentaria Hiibner June 25 Milwaukee. 99. Isturgia truncataria Walker May 11-June 29 Crandon; Lake Katherine; Marinette Co. 100. Heliomata cycladata Grote June 2 6- July 2 Brown Co.; Sauk City. 101. Semiothisa aemulataria Walker May 21 -July 30 Bailey’s Harbor ; Florence Co. ; Lake Katherine ; Madison, Dane Co. ; Marinette Co. ; Sauk City ; Wausau. 102. Semiothisa ulsterata Pearsall June 5-22 Lake Katherine; Vilas Co. 103. Semiothisa minorata Packard June 25- July 13 Dane Co.; Lake Katherine; Trout Lake; Wausau. 1970] Coveil — Geometridae of Wisconsin 175 104. Semiothisa bisignata Walker June 23-Sept. 13 Bailey's Harbor ; Florence Co. ; Green Bay ; Lake Katherine ; Madison; Trout Lake; Vilas Co.; Wausau. 105. Semiothisa bicolorata Fabricius May 19- July 26 Lake Katherine. 106. Semiothisa distribuaria Hiibner May 5-July 19 Arena, Iowa Co.; Lake Katherine; Madison; Trout Lake; Wausau. 107. Semiothisa punctolineata Packard Sept. 7 Dane Co. 108. Semiothisa granitata Guenee May 29-Aug. 15 American Legion State Forest, Oneida Co. ; Bailey’s Harbor ; Florence Co.; Lake Katherine; Madison; Trout Lake; Vilas Co.; Wausau. This name represents a complex of several species, needing revisionary study. 109. Semiothisa oweni Swett May 28-July 11 Lake Katherine. 110. Semiothisa sexmaculata Packard Florence Co.; Green Bay; Lake 111. Semiothisa denticulata Grote Milwaukee; Poynette. 112. Semiothisa eremiata Guenee Dane Co. 113. Semiothisa orillata Walker May 24-Aug. 29 Bailey’s Harbor ; Dane Co. ; Door Co. ; Florence Co. ; Green Bay; Lake Katherine; Madison; Sauk City; Wausau. 114. Semiothisa ocellinata Guenee July 8-Oct. 3 Kenosha Co.; Madison; Milwaukee. 115. Semiothisa mellistrigata Grote May 15-Aug. 25 Belleville; Crandon; Door Co.; Florence Co.; Green Bay; Madison; Milwaukee; Oconto Co. 116. Semiothisa snoviata Packard Aug. 30 Columbia Co. 117. Semiothisa gnophosaria Guenee May 13-Aug. 17 Bailey’s Harbor ; Dane Co. ; Florence Co. ; Lake Katherine ; Madison; Wausau; Wood Co. 118. Itame pustularia Guenee June 5-Sept. 8 Bailey’s Harbor; Brown Co.; Florence Co.; Green Bay; Lake Katherine; Madison; Milwaukee; N.E. Price Co.; Sauk City ; Trout Lake ; Wood Co. May 13-Aug. 9 Katherine. June 23-Aug. 1 July 12 176 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 119. I tame ribearia Fitch May 15-July 80 Belleville ; Dodgeville, Iowa Co. ; Madison ; Milwaukee ; Sauk City; Wausau. 120. Itame argillacearia Packard June 9- July 23 Bailey’s Harbor; Lake Katherine. 121. Itame occiduaria Packard June 28- July 20 Lake Katherine. 122. Itame evagaria Hulst June 20-Aug. 2 Crandon; Lake Katherine; Madison; N.E. Price Co.; Wau¬ sau. 123. Itame fulvaria deVillers June 12- July 10 Crandon; Florence Co.; Lake Katherine; Spooner; Summit Lake; Wood Co. 124. Itame subcessaria Walker June 21 -Aug. 20 Crandon; Florence Co. 125. Itame coortaria Hulst June 5-July 30 Door Co. ; Madison ; Milwaukee ; Sauk City. 126. Itame anataria Swett July 7-13 Lake Katherine. 127. Itame bitactata Walker July 3-Aug. 4 Lake Katherine; N.E. Price Co. 128. Itame loricaria Eversmann June 19-July 13 Lake Katherine; Wausau. 129. Eumacaria latiferrugata Walker June 1-Aug. 28 Griffith State Nursery; Lake Katherine; Sauk City; Wau¬ sau. 130. Thysanopyga intractata Walker The type of synonym gausaparia Grote is from Wisconsin. 131. Hesperumia sulphuraria Packard June 18-Aug. 4 Madison; Wausau. 132. Ematurga amitariei Guenee May 30-Aug. 10 Crandon; Marinette Co.; Vilas Co. 183. Eufidonia notataria Walker June 7- July 23 Griffith State Nursery, Wood Co.; Lake Katherine; Mari¬ nette Co.; Northern Highlands; Vilas Co. 134. Eufidonia discospilata Walker June 10-22 Lake Katherine; Vilas Co. 135. Orthofidonia tinctaria Walker May 8-June 17 Lake Katherine; Sauk City. 136. Hypagyrtis pustularia Hiibner July 4-Aug. 21 Lake Katherine; Wausau. 1970] Coveil — Geometridae of Wisconsin 177 137. Hypagyrtis subatomaria Wood June 4-Aug. 23 Dane Co.; Douglas Co.; Florence Co.; Lake Katherine; N.E. Price Co. ; Trout Lake. 138. Hypagyrtis piniata Packard June 19-Aug. 6 Bailey’s Harbor; Florence Co.; Gordon, Douglas Co.; Grif¬ fith State Nursery, Wood Co.; Madison; Summit Lake; Trout Lake, Vilas Co. 139. Tornos scolopacinarius Guenee Rindge (1954, p. 221) includes “southern Wisconsin” in his range notes for this species. 140. Melanolophia canadaria Guenee Apr. 27- July 22 Bailey’s Harbor; Lake Katherine; Madison; Milwaukee; Patton Lake Spur, Florence Co. 141. Melanolophia signataria Walker May 14- July 3 Bailey’s Harbor; Lake Katherine; Patton Lake Spur, Flor¬ ence Co. 142. Protoboarmia porcelaria Guenee June 10- Aug. 12 Bailey’s Harbor ; Florence Co. ; Sayner ; Summit Lake. 143. Cleora manitoba Grossbeck May 8-July 13 Lake Katherine. 144. Pseudoboarmia umbrosaria Hiibner June 2 9- July 28 Crandon; Lake Katherine; N.E. Price Co. 145. Stenoporpia p o lygrammaria Packard June 27 Waushara Co. 146. Anavitrinella pampinaria Guenee May 8-Sept. 15 Bailey’s Harbor ; Bone Rock, Sauk Co. ; Columbia Co. ; Gays Mills; Griffith State Nursery; Lake Katherine; Madison; Patton Lake, Florence Co.; Neshkoro; Platteville; Sauk City ; Waushara Co. 147. Iridopsis larvaria Guenee June 20- July 20 Bailey’s Harbor; Florence Co.; Lake Katherine. 148. Anacamptodes ephyraria Walker June 29-Aug. 12 Crandon; Florence Co.; Forest Co.; Lake Crandon; Lake Katherine; Madison; Milwaukee. 149. Anacamptodes humaria Guenee May 5-Aug. 14 Griffith State Nursery; Lake Katherine, near Hazelhurst; Madison; Neshkoro. 150. Anacamptodes vellivolata Hulst May 8-Aug. 22 Lake Katherine; Wausau. 151. Aethalura anticaria Walker May 30-July 3 Lake Katherine. 178 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 152. Ectropis crepuscularia Denis and Schiffermiiller Apr. 15- Aug. 5 Arbor; Boscobel State Nursery; Door Co.; Florence Co.; Grant Co.; Lake Latherine; Madison; Milkaukee; Patton Lake, Florence Co.; Vilas Co.; Northern Highlands; Wau¬ sau. 158. Phaeoura quernaria Abbott and Smith May 19- July 11 Bailey’s Harbor; Door Co.; Florence Co.; Lake Katherine; Milwaukee; Neshkoro. One reared larva emerged Apr. 4. 154. Phigalia olivacearia Morrison Apr. 17-May 7 Dane Co.; Lake Katherine; Madison; Milwaukee; Wausau. 155. Phigalia titea Cramer Apr. 18-May 12 Griffith State Nursery; Lake Katherine; Madison; Mil¬ waukee. 156. Palaeacrita vernata Peck March 13-Apr. 28 Lake Katherine; Madison; Milwaukee; Wausau. 157. Erannis tiliaria Harris Oct. 6-Nov. 2 Brown Co.; Dane Co.; Kewaunee Co.; Lake Katherine; Madison; Wausau. 158. Biston ursaria Walker Apr. 12-May 23 Dane Co.; Griffith State Nursery; Lake Katherine; Madi¬ son; Milwaukee; Wausau. 159. Biston cognataria Guenee May 15-Sept. 19 Arlington; Bailey’s Harbor; Boscobel State Nursery; Brown Co.; Columbia Co.; DeForest, Dane Co.; Green Bay; Lake Katherine; Madison; Marinette Co.; Milwaukee; Sauk City; Trout Lake, Vilas Co. ; Washburn Co. 160. Eugonobapta nivosaria Guenee June 28-Aug. 12 Bailey’s Harbor ; Door Co. ; Florence Co. ; Madison ; Milwau¬ kee; Sauk City. 161. Lytrosis unitaria Herrich-Schaffer June 24- July 30 Madison; Sauk City; Wausau. 162. Euchlaena serrata Drury “May”; June 28-Aug. 14 Bailey’s Harbor; Brown Co.; Dousman; Door Co.; Green Bay; Griffith State Nursery; Lake Katherine; Madison; “Peaks Lake”; Sauk City; Sturgeon Bay. 163. Euchlaena obtusaria Hubner June 9- July 19 Bailey’s Harbor; Crandon; Dane Co.; Door Co.; Florence Co.; Lake Katherine; Madison; Milwaukee; Summit Lake. 164. Euchlaena effecta Walker June 18-July 30 Florence Co.; Lake Katherine; Madison; Wausau. 1970] Coveil — Geometridae of Wisconsin 179 165. Euchlaena johnsonaria Fitch June 9-Aug. 29 Bailey’s Harbor; Florence Co.; Lake Katherine; Madison; Marinette Co.; Sauk City; Trout Lake; Wausau; Wood Co. 166. Euchlaena amoenaria Guenee June 22 Marinette Co. 167. Euchlaena marginata Minot May 30-July 8 Florence Co.; Lake Katherine; Vilas Co.; Waushara Co. 168. Euchlaena tigrinaria Guenee June 14-July 15 Bailey’s Harbor; Florence Co.; Lake Katherine; Milwau¬ kee ; Summit, Langlade Co. 169. Euchlaena irraria Barnes and McDunnough June 17-July 10 Florence Co. ; Lake Katherine ; Madison ; Marinette Co. ; Sauk City. 170. Euchlaena milnei McDunnough July 1 Sauk City. 171. Xanthotype sospeta Drury June 6-Aug. 25 Bailey’s Harbor; Brown Co.; Crandon; Door Co.; Florence Co.; Green Bay; Lake Katherine; Madison; Milwaukee; “Peaks Lake”; Rusk Co.; Sauk City; Shawano Co.; Star Lake; Trout Lake, Vilas Co. 172. Xanthotype urticaria Swett July 5-Sept. 8 Arpin; Brown Co.; Columbia Co.; Florence Co.; Lake Katherine; Madison; Milwaukee; Wausau. 173. Campaea perlata Guenee June 1-Oct. 2 Bailey’s Harbor; Columbia Co.; Door Co.; Florence Co.; Lake Katherine; Madison; N.E. Price Co.; Oneida Co.; Patton Lake; Sauk City; Summit Lake, Langlade Co.; Trout Lake, Vilas Co.; Verona; Wausau. 174. Gueneria basiaria Walker July 8-11 Florence Co. 175. Homochlodes fritillaria Guenee May 28- July 23 Door Co.; Florence Co.; Lake Katherine; Vilas Co. 176. Tacparia deter sata Guenee May 21- July 8 Florence Co. ; Lake Katherine ; Madison. 177. Lozogramma subaequaria Walker May 1 9-Aug. 10 Brown Co.; Dane Co.; Florence Co. 178. Cepphis armataria Herrich-Schaffer Apr. 21-Aug. 24 Bailey’s Harbor ; Florence Co. ; Lake Katherine ; Madison ; Milwaukee ; Washburn Co. 179. Plagodis ser inaria Herrich-Schaffer May 22- June 27 Florence Co. ; Lake Katherine ; Sauk City. 180 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 180. Plagodis keutzingi Grote July 1-13 Bailey's Harbor ; Door Co. ; Madison. 181. Plagodis fervidaria Herrich-Schaffer May 24- July 5 Bailey's Harbor; Brown Co.; Dane Co.; Door Co.; Sauk City. 182. Plagodis alcoolaria Guenee May 17-July 29 Bailey's Harbor; Dane Co.; Florence Co.; Lake Katherine; Madison; Milwaukee; Patton Lake; Sauk City; Vilas Co. 183. Plagodis pklogosaria Guenee Apr. 30- Aug. 1 Bailey’s Harbor ; Florence Co. ; Lake Katherine ; Milwaukee ; Sauk City; Trout Lake; Wausau. I have seen typical phlo- gosaria, p. keutzingaria Packard, and p. purpuraria Pears¬ all. 184. Anagoga occiduaria Walker May 29- July 12 Bailey’s Harbor; Florence Co.; Lake Katherine; Vilas Co. 185. Hyperetis amicaria Herrich-Schaffer May 29-Aug. 29 Bailey’s Harbor; Florence Co.; Lake Katherine; Madison; Marinette Co.; Milwaukee; Patton Lake; Rush Co.; Vilas Co. ; Wausau ; Waushara Co. 186. Hyperetis alienaria Herrich-Schaffer May 2 2- July 4 Bailey's Harbor; Brown Co.; Florence Co.; Green Bay; Lake Katherine; Milwaukee; “Peaks Lake”; Wausau. 187. Nematocampa filamentaria Guenee June 4-Aug. 26 Bailey's Harbor; Florence Co.; Madison; Milwaukee; Rusk Co. ; Rust Lake ; Sauk City. 188. Metarranthis hypochraria Herrich-Schaffer Apr. 13-Aug. 10 Brown Co. ; Door Co. ; Florence Co. ; Lake Katherine ; Madi¬ son; Marinette Co.; Milwaukee. 189. Metarranthis broweri Rupert May 29-July 8 Bailey's Harbor; Dodgeville; Florence Co.; Madison. 190. Metarranthis apiciaria Packard May 29 Lake Katherine. 191. Metarranthis warneri Harvey June 4-6 Wausau; a paratype $ of M. warneri capp sarin Rupert collected 4 June, 1932, is from Madison. 192. Metarranthis duaria Guenee May 28- June 20 Lake Katherine; Madison; Milwaukee; Vilas Co. ; Wausau; Waushara Co. 193. Metarranthis amyrisaria Walker May 27-July 4 Bailey's Harbor; Florence Co.; Madison. 194. Metarranthis angularia Barnes and McDunnough July 8 Milwaukee. 1970] Coveil — Geometridae of Wisconsin 181 195. Metarranthis obfirmaria Hlibner June 80 Milwaukee. 196. Metanema inatomaria Guenee May 29-Aug. 15 Bailey's Harbor; Dane Co.; Florence Co.; Lake Katherine; Milwaukee ; Rusk Co. ; Sauk City ; Waushara Co. 197. Metanema determinata Walker June 17-Aug. 16 Bailey's Harbor; Florence Co.; Lake Katherine; Milwau¬ kee ; Rusk Co. ; Summit Lake. 198. Selenia alciphearia Walker May 7- June 7 Lake Katherine. 199. Selenia kentaria Grote and Robinson Apr. 30- July 22 Florence Co. ; La Crosse ; Lake Katherine. 200. Ennomos subsignarius Hiibner June 19-Sept. 10 Bailey's Harbor; Brown Co.; Florence Co.; Green Bay; Lake Katherine; Madison; Milwaukee. 201. Pero honestaria Walker May 16-Sept. 1 Bailey’s Harbor; Brown Co.; Dane Co.; Green Bay; La Crosse; Madison; Milwaukee; Platteville; “Shaw” (prob¬ ably Shawano) Co. 202. Pero morrisonaria H. Edwards May 28- July 9 Dane Co.; Door Co.; Florence Co.; Lake Katherine; Wash¬ burn Co. 203. Pero marmorata Grossbeck July 29-Aug. 9 Dane Co. ; Platteville. 204. Caripeta divisata Walker June 23-Aug. 14 Door Co.; Florence Co.; Lake Katherine; N.E. Price Co.; Northern Highlands, Vilas Co.; Trout Lake; Wausau. 205. Caripeta piniata Packard May 2 6- Aug. 5 Lake Katherine ; Northern Highlands ; Trout Lake ; Wausau. 206. Caripeta angustiorata Walker June 4-Aug. 6 American Legion State Forest, Oneida Co.; Florence Co.; Lake Katherine ; Trout Lake, Vilas Co. ; Washburn Co. ; Wausau. 207. Lambdina athasaria Walker May 13- July 8 Florence Co. ; Lake Katherine ; Sauk City. 208. Lambdina fiscellaria Guenee July 10-Oct. 5 Bailey's Harbor; Columbia Co.; Dane Co.; Ephraim; Lake Katherine ; Shanty Bay. 209. Besma endropiaria Grote and Robinson June 1-July 15 Bailey’s Harbor; Boscobel State Nursery; Florence Co.; Lake Katherine; Madison; Milwaukee; Patton Lake; Sauk City. 182 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 210. Besma quercivoraria Guenee May 28-Aug. 19 Arlington; Dane Co.; Florence Co.; Green Bay; Griffith State Nursery; Lake Katherine; Madison; Neshkoro; Platteville; Sauk City. 211. Cingilia catenaria Drury Aug. 9— Oct. 2 Columbia Co.; Eau Claire; Lake Katherine; Madison; “Peaks Lake.” 212. Cingilia canosaria Walker July 11-Oct. 10 Bailey’s Harbor; Dousman; Lake Katherine; Wausau; Wood Co. 213. Cingilia pellucidaria Packard Lake Katherine. Sept. 7-0 ct. 4 214. Sicya macularia Harris June 5-Aug. 18 Bailey’s Harbor; Door Co.; Florence Co.; Griffith State Nursery; Lake Katherine; Madison; Sauk City; Trout Lake. 215. Deuteronomos magnarius Guenee July 30-Nov. 11 Arlington ; Bailey’s Harbor ; Brown Co. ; DeForest ; Green Bay; Kenosha Co.; Lake Katherine; Madison; Middleton; Milwaukee; “Peaks Lake”; Sturgeon Bay; Trout Lake. 216. Apicia confusaria Hiibner June 4- Aug. 29 Bailey’s Harbor ; Brown Co. ; Cedar Grove ; Door Co. ; Dous¬ man; Griffith State Nursery; Madison; Marinette Co.; Mil¬ waukee ; Oconto Co. ; Sauk City. 217. Patalene puber Grote and Robinson Aug. 7 Dane Co. 218. Tetrads crocallata Guenee May 29-July 8 Brown Co. ; Door Co. ; Florence Co. ; Lake Katherine ; Madi¬ son ; Milwaukee ; Sauk City. 219. Tetrads cachexiata Guenee Apr. 21- July 29 Bailey’s Harbor ; Door Co. ; Florence Co. ; Green Bay ; Madi¬ son; Milwaukee; Patton Lake; Sauk City; Spooner. 220. Ahbottana clemataria Abbott and Smith May 14-Sept. 26 Bailey’s Harbor; Boscobel State Nursery; Door Co.; Flor¬ ence Co.; Griffith State Nursery; Lake Katherine; Milwau¬ kee Co.; Patton Lake; Sauk City; Wausau. 221. Sabulodes thiosaria Guenee May 2 9- June 30 Lake Katherine. 222. Prochoerodes transversata Drury June 18-Oct. 5 Bailey’s Harbor; Ferry Bluff; Florence Co. ; Lake Kath¬ erine ; Madison ; Milwaukee ; “Peaks Lake” ; Rusk Co. ; Sauk City; Wausau. 1970] Coveil — Geometridae of Wisconsin 183 References Capps, H. W. 1943. Some geometric! moths of the subfamily Ennominae hereto¬ fore associated with or closely related to Ellopia Treitschke. Proc. U.S. Nat. Mus. No. 93, pp. 115-151, 10 pis. Ferguson, D. C. 1969. A revision of the moths of the subfamily Geometrinae of America north of Mexico (Insecta, Lepidoptera). Peabody Mus. Bull. 29, 251 pp., 49 pis. Fernekes, V. 1906. List of Lepidoptera occurring in Milwaukee County. Bull. Wisconsin Nat. Hist. Soc., vol. 4, nos. 1-2, pp. 39-58. Fletcher, D. S. 1966. Some changes in the nomenclature of British Lepi¬ doptera. Ent. Gazette, vol. 17, pp. 9-18. Forbes, W. T. M. 1948. Lepidoptera of New York and neighboring states. Part II. Cornell Univ. Exp. Sta. Mem. No. 274, 263 pp., 255 figs. McDunnough, J. H. 1938. A check list of the Lepidoptera of Canada and the United States of America, Part 1. Mem. Southern Calif. Acad. Sci., vol. 1, 275 pp. - 1949. Revision of the North American species of the genus Eupithecia (Lepidoptera, Geometridae). Bull. Amer. Mus. Nat. Hist., vol. 93, art. 8, pp. 537-728, 20 figs., 7 pis. - 1954. The species of the genus Hydriomena occurring in America north of Mexico (Geometridae, Larentiinae) . Bull. Amer. Mus. Nat. Hist., vol. 104, art. 3, pp. 241-358, 185 figs., 3 pis. Munroe, E. G. 1959. The phlogosctria complex of the genus Plagodis (Lepi¬ doptera, Geometridae). Can. Ent., vol. 91, pp. 193-208, 63 figs. Muttkowski, R. A. 1907. Additions to the lepidopterous fauna of Milwaukee County. Bull. Wisconsin Nat. Hist. Soc., vol. 5, no. 2, pp. 128-133. Rauterberg, F. 1900. List of Lepidoptera of the County of Milwaukee. Bull. Wisconsin Nat. Hist. Soc., vol. 1, no. 2, pp. 111-126. Rindge, F. H. 1949. A revision of the geometrid moths formerly assigned to Drepanulatrix (Lepidoptera). Bull. Amer. Mus. Nat. Hist., vol. 94, art. 5, pp. 235-298, 14 figs. - 1954. A revision of the genus Tomos Morrison (Lepidoptera, Geo¬ metridae). Bull. Amer. Mus. Nat. Hist., vol. 104, art 2, pp. 181-236, 35 figs. — - 1956. A revision of the American species of Deilinia (Lepidoptera, Geometridae). Amer. Mus. Nov. No. 1810, 23 pp., 30 figs. - 1964 A revision of the genera Melanolophia, Pherotesia, and Mel- anotesia (Lepidoptera, Geometridae). Bull. Amer. Mus. Nat. Hist., vol. 126, art. 3, pp. 243-434, 163 figs., 9 pis. - 1966. A revision of the moth genus Anacamptodes (Lepidoptera, Geometridae). Bull. Amer. Mus. Nat. Hist., vol. 132, art. 3, pp. 177-243, 53 figs. - 1967. The North American moths of the genus Earophila Gumppen- berg (Lepidoptera, Geometridae). Amer. Mus. Nov. No. 2306, 12 pp., 9 figs. - 1968. A revision of the moth genus Stenoporpia (Lepidoptera, Geo¬ metridae). Bull. Amer. Mus. Nat. Hist., vol. 140, art. 2, pp. 67-134, 16 pis. Rupert, L. R. 1943. A specific revision of the genus Metarranthis (Lepidoptera, Geometridae, Ennominae). J. New York Ent. Soc., vol. 51, pp. 133-158, 3 pis. ■ - 1949. A revision of the North American species of the genus Plagodis (Lepidoptera, Geometridae, Ennominae). J. New York Ent. Soc., vol. 57, pp. 19-49, 5 pis. FOUR NEW SPECIES RECORDS OF SIALIS (MEGALOPTERA: SIALIDAE) FOR WISCONSIN K. J. Tennessen There has been very little collecting of adult Megaloptera in Wis¬ consin since Ross’s work on the Sialidae in the 1930’s, when he re¬ ported finding two species in this state, Sialis mohri and S. velata. While recording distribution data on the order Megaloptera during the summer of 1968, I found four additional species of Sialis that had not been previously recorded from the state. The identifications have been confirmed by Dr. Ross. The new records are as follows : Sialis vagans Ross. Twelve males and six females of this species have been taken, as it appears to be quite common, especially in the north-central area. Collection dates range from June 2 in the south to June 20 in the north. County Records — Clark, Dane, Forest, Polk, Price, Taylor, Wash¬ burn, Waukesha, Waushara. Sialis itasca Ross. Three males were taken, and the species ap¬ pears to be as widespread as S. velata. The collection dates, June 10 and 20, indicate that this species emerges comparatively late in the seasonal succession of species. County Records — Bayfield, Florence, Jefferson. Sialis infumata Newman. Only two males of this species have been collected, one each from two central counties, indicating that it may not be as common as the former species. The collection date was June 5. County Records — Clark, Wood. Sialis americana (Rambur) . Three males and seven females were collected on June 12 near the Mississippi River in Grant County. The surrounding area was quite unique, like a lagoon, and there¬ fore this species most likely does not occur much further inland due to a habitat restriction. Adults were present in fair abundance. The collection date suggests a comparatively late emergence period. County Record — Grant. In addition, the following new county records were found for the two previously reported species : Sialis mohri Ross. Kenosha County, Racine County. Sialis velata Ross. Winnebago County. 185 186 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 The specimens are in the University of Wisconsin collection. Future collecting, especially early in May and June, will un¬ doubtedly discover new county records as larvae have been found in a large number of streams and lakes; but as there is yet no key separating the larvae, species determinations must rely on adults. References Cited Ross, H. H. September, 1937, Studies of Nearctic Aquatic Insects. Illinois Natural History Survey Bulletin. 21:57-78. The research was carried out when the author was an undergraduate stu¬ dent at the University of Wisconsin, Madison. JUNCUS EFFUSUS. I. THE SITUATION IN WISCONSIN Seymour H. Sohmer J uncus effusus is a species with world-wide distribution (Wei- mark, 1946). It is a relatively hardy perennial and can withstand considerably more abuse than most of the other species in the genus. Postgalacial conditions in Wisconsin may have been ideal for colo¬ nizing elements of the species. Indeed, such was probably the case wherever land was uncovered by the melting ice. It is believed, in this regard, that colonizing elements from disjunct populations of the species may have entered Wisconsin after the last retreat of the ice (litis, 1968). As would be expected of a species with a wide distribution, the variations of form in Juncus effusus are numerous. Many varia¬ tions have been described for North America, and many more could probably be named. The difficulty and confusion here is the same as in other widely distributed species, in that there are as many opin¬ ions regarding the basis for the delimitation of these varieties as there are taxonomists involved. The hopeful goal in these situations is a compromise between taxonomic conservatives and radicals. Certainly there are more variations in this species that can be easily delimited from one another on a general and practical basis than Gleason (1952) would recognize, but probably less than the number of varietal names presently available. One should strive to avoid carrying the recognition of variations too far. In such a case the logical conclusion is ultimately to describe and name all varia¬ tion present. This is not only cumbersome and impractical, but the same variation may not be present some years hence — even disre¬ garding the destructiveness of man to his environment. A great deal of the individual variation may be due more to environmental differences than genetic ones. (Snogerup, 1963. Sohmer, unpub¬ lished). A method of dealing with such a situation is shown by Davidson and Dunn (1967). The varietal names listed below are presently available to de¬ scribe the variations of Juncus effusus in North America. It is pointed out that though some of these varietal names may not be valid, in the author’s opinion, they will be used to describe the species in Wisconsin where applicable. The question of validity may 187 188 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 be determined at a future date by systematic work now underway in the author’s laboratory. Var. compactus Lejeune et Courtois FI. Belg. ii. 28 (1831), also see Fernald and Wiegand (1910). Var. conglomeratus Engelm. in Gray, Manual, ed. 5, (537, 1867), also see Fernald and Wiegand (1910). Var. decipiens Buchenau in Engler’s Bot. Jahrb. Xii. 229 (1890), also see Fernald and Wiegand (1910). Var. exiguus Fernald and Wiegand (1910). Var. gracilis Hooker, FI. Bor. Am. ii 190 (1840). Var. brunneus Engelm., Trans. St. Louis Acad. ii. 491 (1868). According to Fernald and Wiegand (1910) Juncus effusus var. hesperius Piper is synonomous with this. Var. pacificus Fernald and Wiegand (1910). Var. caeruleomontanus St. John (1931). Var. solutus Fernald and Wiegand (1910). Var. pylaei (Laharpe) Fernald and Wiegand (1910). Var. costulatus Fernald (1922). In Wisconsin the species demonstrates a considerable amount of individual variation, principally within the variety pylaei. This variation, however, does not appear spatially related to given areas. The information concerning the species in Wisconsin that will be presented here has come through study of the specimens available from the herbaria of the Universities of Wisconsin and Minnesota as well as the Milwaukee Public Museum. The variable characters measured were: culm length; number of flowers per inflorescence; length of bract subtending the inflorescence (but in the species ap¬ pearing as a continuation of the culm) ; width of culm above the sheath; length of perianth parts; length of capsule; and length of seed. Figure 1 illustrates the distribution of the species in Wiscon¬ sin as revealed by the specimens mentioned above. There are re¬ ports from 45 counties. Among the specimens available the sepals were found to exceed the petals in most cases. The length of the former ranged from 1.9 mm (N. C. Fassett #22676, Chippewa County) to 3.9 mm (H. Brawn #69228, Forest County). The cor¬ relation between sepal and petal length is illustrated in figure 4. Figures 2 and 3 illustrate the occurrence of petal and sepal vari¬ ability by county within the state. There were no significant differences observed in the length or the shape of the seeds. They were found to vary between .5 and .6 mm in length and were generally fusiform. The bracts varied between 6 cm (D. F. Grether #6855, Jackson County),, and 38 cm (R. Melville #397). The latter was two-thirds the length of the culm itself. 1970] Sohmer — Juncus Effusus 189 Figure 1. Distribution of Juncus effusus in Wisconsin. Each symbol represents an individual. Information from herbarium specimens of the University of Wisconsin, University of Minnesota, and the Milwaukee Public Museum. Map provided by H. litis. The most striking variability is expressed in the length of these culms and the number of flowers per inflorescence. Only mature plants with intact culms and inflorescences were utilized. The length of these culms in mature specimens was found to vary from 25 cm (L. S. Cheney, #1068, Oneida County) to 118 cm (N. C. Fasset, #16190, Oneida County) . It is interesting to not that the minimum and maximum lengths examined occurred in populations within 190 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figures 2 and 3. Spatial relationships of artificial groupings of petal and sepal lengths by county within Wisconsin. Each symbol represents one or more individuals within each county whose specific character attributes fall into the artificial groups shown. 2-petal lengths; 3-sepal lengths. Figure 4. Scatter diagram illustrating the correlation between petal and sepal lengths. Each symbol represents an individual. 1970] Sohmer — J uncus Effusus 191 Table 1. Variability in Culm Length and Flower Numbers in Wisconsin Representatives of Juncus Effusus Culm Length (185 Specimens) Flower Number (183 Specimens) Length (in cm) . % of Plants No. of Flowers % of Plants 31 . 6 21 . 6 31 to 50 . 27 21 to 60 . 28 51 to 70 . 40 61 to 100 . 39 71 to 90 . 17 101 to 200 . 17 90..... . 10 200 . 10 the same county. The number of flowers was found to vary from 11 (H. H. litis, #6882, Juneau County) to 870 (C. Gossel #1398, Clark County) per inflorescence. Table 1 illustrates the proportions of the individuals studied that fall into artificial groups based on culm length and flower number. There does not appear to be a relationship between the number of flowers per inflorescence and location within the state. This is shown in Figure 6. Nor is there an apparent relationship between culm length and location. The shortest and tallest plants, on the basis of the specimens available, seem to have equal probability of occurring anywhere in the state. This is illustrated in Figure 5. The correlation between culm length and the number of flowers per inflorescence is illustrated in Figure 7. The varieties pylaei and solutus are found within the state, with the former much more frequent. There is also a dubius report for var. costulatus from Lincoln county. Most members of variety solutus appear in the northeastern part of the state, with scattered reports elsewhere. Some individual examples of Juncus effusus are shown in figures 8-12. Figure 13 represents the distribution of these varieties within the state by county. Summary Specimens of Juncus effusus from the herbaria of the Univer¬ sities of Wisconsin and Minnesota, and the Milwaukee Public Museum were studied with regard to the kind of variation present. It appears that two varieties are established in the state (pylaei and solutus ) and possibly a third ( costulatus) . There is a consider¬ able amount of individual variation within the delimited varieties, which is not, however, apparently correlated with space within the state of Wisconsin. 192 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figures 5 and 6. Spatial relationships of the variability present in the length of culm and number of flowers per inflorescence respectively. Each symbol represents at least one individual within each county whose specific character attribute falls into an artificial group as shown. Figure 7. Scatter diagram illustrating the relationship between culm length and number of flowers per inflorescence. Each symbol represents an individual. Figures 8-12. Examples of variation in Juncus effusus in Wisconsin. 8-Var. pylaei, L. S. Cheney #1068, Oneida Co.; 9-Var. solutus, N. C. Fassett #16190, Oneida Co.; 10-Var. solutus, N. C. Fassett #5699, Ashland Co.; 11-close-up of inuorescence of plant in figure 10; 12-close-up of inflorescence of plant in figure 9. 194 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 13 Varieties of ef/usus in Wisconsin Figure 13. Distribution of the varieties recognized in Wisconsin. Each symbol represents at least one report for the variety within a county. Acknowledgements The aid of Dr. H. H. litis, curator of the Herbarium, University of Wisconsin, Madison, in obtaining specimens for study and his advice are gratefully acknowledged. The monetary aid, in part, of the institutional Studies and Grants Committee of Wisconsin State University, La Crosse, is also gratefully acknowledged here. This aid has permitted the use of student assistance and helped defray the cost of an extensive field trip. 1970] Sohmer — -J uncus Effusus 195 References Cited Davidson, Robert A. and Rosalie A. Dunn, 1967. A correlation approach to certain problems of population-environment relations. American Journal of Botany 54(5) : 529-538. Fernald, M. L. and K. M. W|57 (&4, 1910. The North American vacations of Juncus effusus. Rhodora 12(137) : 81-93. Fernald, M. L., 1922. Expedition to Nova Scotia. Rhodora 23: 239. Gleason, H. A., 1952. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. Volume I, page 388 New York Botanical Garden, New York. Hartley, Thomas G., 1966. The Flora of the “Driftless” Area. The University of Iowa Studies in Natural History 21(1) : 1-174. Iltis, Hugh H., 1968. Personal communication regarding the possible move¬ ments of certain plants after the last glaciation. Snogerup, Sven, 1963. Studies in the genus Juncus III. Observations on the diversity of chromosome numbers. Botanisk Notiser 116(2): 142-145. St. John, Harold, 1931. New and noteworthy Northwestern plants, part 5. Research Studies, State College of Washington. Weimarck, H. 1946. Studies in Juncaceae. With special reference to the species in Ethiopia and the Cape, Svensk Botanisk Tidskrift 40(2) : 141-178. GROWTH POTENTIAL OF WISCONSIN NATIVE PINES ON WEED-INVADED SOILS' S. A. Wilde “No knowledge can he rated as science unless it is anchored in mathematics. ” — Rene Descartes A reasonably reliable prediction of the outcome of tree planting- can seldom be made without an appraisal of the growth-depressing effect of competing ground vegetation. In most cases, the amount of water and nutrients taken away from trees by ground cover plants is closely related to their biomass, the oven-dry weight of tops and roots. Knowledge of this factor is essential for determina¬ tion of the expected yield of trees to be planted, for establishment of suitable tree spacing, and for estimate of the cost of either chemical or mechanical eradication of weeds. This paper reports a method for determination of the biomass of weed cover and the expected yield of plantations of Wisconsin na¬ tive pines, Pinus banksiana, P. resinosa, and P. strobus. Determination of Expected Site Index The expected site index of the proposed plantation is established on the basis of determination of four soil constituents (Wilde et al, 1964) : fine earth (F), organic matter (H), available phosphorus (P2O5), and available potassium (K20). The site index of the stand or the average height of a stand at 50 years (I) is obtained by solu¬ tion of the following simplified regression equation, adjusted to meet requirements of the three native pines of Wisconsin. I = (8.1 + 0.2 F% + 2.8 H°/o + 0.03 P0O5 lbs/a + 0.01 K20 lbs/a) X 8.4 An application of this equation may be illustrated by an ap¬ praisal of Plainfield sand, a widely distributed non-podzolic soil of Wisconsin glacial outwash. Let us assume that this soil of the proposed planting site has the following composition: fine earth (silt and clay particles)— 9%, organic matter — 2%, available P205 1 Contribution from Soil Science Department, University of Wisconsin, in cooperation with and supported in part by the Wis. Dept, of Natural Resources. Publication ap¬ proved by the Director of the Wis. Agr. Exp. Sta., Madison, Wis. 197 198 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 — 40 pounds per acre, and available potassium— 100 pounds per acre. Consequently: I = (8.1 + 0.2 X 9 + 2.8 X 2 + 0.03 X 40 + 0.01 X 100) X 3.4 = 56.8 feet The site index corresponds to definite yields of timber produced at various ages by different tree species. Thus, for the calculated site index of about 57, the volumes expected at the end of a short, 40-year rotation would be 30 cords for jack pine, 34 cords for red pine, and at best 20 cords per acre for white pine. Because 0.7 cords per acre is the minimum average annual increment, promising un¬ der Wisconsin conditions a reasonable financial return, the calcula¬ tion indicates that the analysed soil has a too low productive po¬ tential for planting white pine. The equation under discussion was derived from analyses of non- phreatic sandy soils of Wisconsin supporting about 300 plantations of jack pine, red pine, and white pine of different site indices and of age range from 15 to 37 years (Wilde et al., 1965). The dis¬ closure of an intimate correlation between growth of forest stands and physico-chemical properties of soils, rendered by this equation, presents a remarkable by-product of recent progress in soil analysis, forest mensuration, and statistics. The degree of the equation’s reliability can be inferred from Tables 1 and 2, incorporating the average results of soil and timber analyses published in 1965 (Lit. cit.) . A substitution of these re¬ sults into the equation and subsequent comparison of calculated site indices with those reported in yield tables (Wackerman et al., 1929, for jack pine; Wilde et al., 1964 for red pine; Gevorkiantz Table 1. Average State of Soil Fertility Factors of Soils Supporting Jack Pine, Red Pine, and White Pine of Different Site Indices (After Wilde et al., 1965). Tree Species and Site Index Reaction pH Silt + Clay Percent Org. Matter Percent Avail. P2O5 Lbs. /Acre Avail. k2o Lbs. /Acre Jack pine, low . 4.9 6.9 0.86 21.0 54.8 Jack pine, medium . 5.1 9.4 1.80 62.8 80.0 Jack pine, high . 5.0 10.0 2.10 72.0 91.4 Red pine, low . 5.0 7.3 1.22 47.2 77.8 Red pine, medium . 5.1 8.5 1.37 59.0 87.5 Red pine, high . 5.1 12.5 2.11 85.2 134.2 White pine, medium . 5.1 13.7 2.90 51.0 70.0 White pine, high . 4.8 26.7 3.20 83.0 90.0 1970] Wilde — Growth of Wisconsin Native Pines 199 Table 2. Average Growth of Jack Pine, Red Pine, and White Pine Plantations on Soil of Different Productivity Ratings; as Given in Table 1 (After Wilde et al., 1965). Tree Species and Site Quality Age Yrs. Ave. Ht. Ft. Ht.: Age Ins. Ave. DBH Ins. Stems No. /A. Basal Area Sq. Ft./A. Volume Cu. Ft./A. Jack pine, low . 22 22.0 12 3.4 1 ,241 74 473 Jack pine, medium . . . 24 30.5 15 4. 1 1,177 107 1,106 Jack pine, high . 23 38.6 20 4.6 1,029 117 1,589 Red pine, low . 23 21.3 11 4.0 1,282 101 688 Red pine, medium. . . . 21 27.5 15 4.8 978 117 1,300 Red pine, high . 20 32.9 19 5.1 1,054 136 1,960 White pine, medium. . 24 28.6 14 5.7 648 118 1,420 White pine, high . 24 39.3 19 6.6 627 136 2,257 and Zon, 1930, for white pine) give a picture of a rather astonish¬ ing accuracy (Table 3) . Nearly all deviations of calculated values from those given in the yield tables are well within experimental error. The maximum deviation of the calculated site index for jack pine of low site quality (3 feet) is due to the deficiency of organic matter in soils of this group and the presence of a large fraction of nutrients in the form of silicate minerals which are decomposable by mycorrhizal rootlets,, but not by weak extracting solutions (Wilde and Iyer, 1962; Spyridakis et al., 1967). Determination of the Biomass of Weed Cover The average weight of tops and roots of weeds is determined by excavating entire plants on several 1/10,000 acre plots (2.1 by 2.1 foot squares). Sampling is done at random, and the number of samples required for obtaining an acceptable standard deviation depends on type of soil and nature of ground cover. In many in¬ stances a half dozen sampled quadrats are sufficient for an area as large as 40 acres. The obtained information facilitates in reaching a decision on tree spacing most advantageous with regard to potential site index, biomass of weeds, species and age of trees to be planted, and other ^ conditions (Wilde et al, 1968). Next, the tree planter must establish the approximate age of the plantation (n) at which the biomass of weeds (b) will be reduced to the harmless level of less than 2 tons per acre. The total of the 200 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Table 3. Relation Between the Predicted and Actual Site Indices. Tree Species Site Indices Deviation in 50 Year Height Growth Feet Calculated from the Equation Determined from the Yield Tables Jack pine . 43 46 -3 57 55 +2 61 62 -1 Red pine . 49 48 + 1 53 55 -2 65 65 0 White pine* . 67 69 -2 82 82 0 *A large number of white pine of low site quality were underplantings suppressed in growth by overhead canopy of aspen and were excluded from statistical analysis. As should be expected, both the calculated and actually determined height growth of white pine plantations on fertile soils by far exceeded the maximum of the yield tables obtained largely on the basis of indigenous stand of this tree (75 feet at the age of 50 years), which undoubtedly lost a fraction of their height increment in the struggle with weeds and volunteer trees. biomass (G), present in the plantation through the years, is then determined from the formula : G = 0.7 bn For trees with heavy crowns, such as red pine, growing* on fertile soils at a spacing of 4 by 4 feet, the n period is about 15 years. How¬ ever, at 6 by 6 foot spacing this period is extended to approximately 25 years. With weed cover of blueberries,, sweet fern, and other heath plants weighing 10 metric tons per acre, total biomass of a 4 by 4 foot red pine plantation would be : G =. 10,000 X 0.7 X 15 = 105,000 kg/a A similar calculation for a red pine plantation, established at a spacing of 6 by 6 feet, gives the active weed biomass of 175,00 kg/a. Determination of the Loss of Timber Volume Due to Competition of Weed Cover Evapotranspiration of weeds consumes under Wisconsin condi¬ tions approximately 85 kg of water per kilogram of oven-dry bio¬ mass (Wilde et al., 1968) . On the other hand, production of one kilogram of merchantable wood, having specific gravity of 0.32- 0.35, requires close to 1,200 kg of transpiration water (Wilde, 1967; 1970] Wilde — Growth of Wisconsin Native Pines 201 Shaw et al, 1968). Therefore, the loss of timber (L) is expressed by the formula : 85 G “ 1200 0.07 G Assuming active weed biomass of a 6 by 6 foot red pine planta¬ tion to be equal to 175 m.t. per acre, the loss of timber would be : L = 175,000 X 0.07 = 12,250 kg/a This weight of oven-dry wood of specific gravity 0.35 approaches 14 cords. In the event calculated site index of the proposed plantation is 60, the expected maximum yield of fully stocked stand at the end of a 40-year rotation should be 40 cords per acre. Actually in our case the volume is likely to be 40 — 14 or 26 cords. The given calculation, featuring 35% loss of merchantable tim¬ ber, is based on observations in Wisconsin. In this state tree plant¬ ing on weed-invaded soils was usually accomplished by plowing deep and wide furrows which greatly reduce adverse effects of weed competition. On the other hand, losses of timber volume ex¬ ceeding 50% were recorded in our study in plantations established without adequate ground preparation. Competing vegetation deprives trees not only of water but also of nutrients. However, this loss is of a temporary nature ; suppres¬ sion of weeds by tree crowns in time returns nutrients to the planta¬ tions cycle via mineralization of plant remains. Summary The productive potential of planting sites for Finns hanksiana, P. resinosa, and P. strobus is predicted on the basis of soil analysis, simplified regression equation, and biomass of weeds. Under local conditions pine plantations require about 1,200 kg of water to pro¬ duce 1 kg of merchantable wood of approximate specific gravity of 0.35,, whereas evapotranspiration of ground vegetation consumes circa 85 kg of water per kg of oven-dry tissues. In turn, each kg of weed biomass present in the plantation through the years reduces production of merchantable wood by 0.07 kg. At this rate, planta¬ tions established on soils with a heavy cover of heath plants may suffer within a 40-year rotation a loss of timber exceeding 15 cords per acre. References Gevorkiantz, S. R., and R. Zon. 1930. Second-Growth White Pine in Wiscon¬ sin. Wis. Agr. Exp. Sta., Res. Bull. 98. Madison, Wisconsin. Shaw, B. H., R. R. Maeglin, and S. A. Wilde. 1968. Soil water supply — its consumption by forest stands and weed cover. Adv. Frontiers of Plant Sci. 21:141-155. 202 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Spyridakis, D. E., G. Chesters, and S. A. Wilde. 1967. Kaolinization of biotite as a result of coniferous and deciduous seedling growth. Soil Sci. Soc. Proc. Amer. 31:203-210. Wackerman, A. E., R. Zon, and F. G. Wilson. 1929. Yield of Jack Pine in the Lake States. Wis. Agr. Exp. Sta., Res. Bull. 90. Madison, Wisconsin. Wilde, S. A. 1967. Production of energy material by forest stands as related to supply of soil water. Acta for. fenn. 1,2:31-44. Wilde, S. A., A. Giordano, B. H. Shaw, A. W. Fedkenheuer, and W. T. Seip. 1968. Tree Spacing in Forest Plantations as Related to Soils and Revenue. Coll, of Agr. and Fife Sciences, Univ. of Wis., Bull. 589. Madison, Wis¬ consin. Wilde, S. A., and J. G. Iyer. 1962. Growth of red pine (Pinus resinosa) on scalped soils. Ecology 43:771-774. Wilde, S. A., J. G. Iyer, Ch. Tanzer, W. L. Trautmann, and K. G. Wat- terston. 1964. The growth of red pine (Pinas resinosa,, Ait.) plantations in relation to fertility of non-phreatic sandy soils. For. Science 10:463-470. Wilde, S. A., J. G. Iyer, Ch. Tanzer, W. L. Trautmann, and K. G. Wat- terston. 1965. Growth of Wisconsin Coniferous Plantations in Relation to Soils. Univ. of Wisconsin, Res. Bull. 262. Madison, Wisconsin. Wilde, S. A., B. H. Shaw, and A. W. Fedkenheuer. 1968. Weeds as a factor depressing forest growth. Weed Res., 8:196-204. Wilde, S. A., G. K. Voigt, and J. G. Iyer. 1964. Analysis of Soils and Plants for Tree Culture. Oxford Books Co., New Delhi, India. CORIXIDAE (WATER BOATMEN) OF WISCONSIN' William L. Hilsenhoff1 2 In 1948 H. B. Hungerford published his monumental monograph on the Gorixidae of the Western Hemisphere, which included a monograph on the Trichocorixa by R. I. Sailer. In it are keys, de¬ scriptions and the recorded distribution for all the known species. Numerous collection records were listed from the four states that border Wisconsin, especially from Michigan and Minnesota, but records from Wisconsin were meager, with only 23 species re¬ ported from this state. In November and December of 1965, large aggregates of corixids were collected from the Wisconsin River and its tributaries. These were mixtures of several species, many of which had not previously been reported from Wisconsin. Subsequent sampling showed that these aggregates occurred only in the late fall, when corixids ap¬ parently congregate in the larger streams to spend the winter. These aggregates often contained 15 or more species, with one collection from the St. Croix River in Douglas County on November 20, 1968, containing 23 different species. Since it was obvious that Wisconsin’s Corixidae were largely un¬ known, an effort was made to collect Corixidae from all areas of the state in 1968, with a special emphasis on collecting from larger streams in October and November to take advantage of the large aggregations of wintering individuals. Collections were made in all counties, but some counties were sampled more thoroughly than others, or were sampled at times more advantageous for collecting corixids. From 1962 through 1968, nearly 22,000 corixids were col¬ lected and identified,, and of the 47 species that were collected, 25 are new records for the state. All specimens were preserved in 70 % ethanol and have been deposited in the University of Wisconsin Insect Collection, along with detailed collection data. Keys to the Wisconsin species, maps of their distribution, com¬ ments on their distribution, abundance, and identification, and a 1 Approved for publication by the Director of the Wisconsin Agricultural Experiment Station. Research supported in part by a grant from the Wisconsin Department of Natural Resources. 2 Department of Entomology, University of Wisconsin, Madison 53706. 203 204 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Figure 1. Hesperocorixa obliqua (modified from Hungerford 1948). A. Dorsal view of male. B. Ventral view of male. C. Lateral view of female. D. Foreleg of male. 1970] Hilsenhoff — Corixidae of Wisconsin 205 summary of the collection records are reported below. The keys are adapted from those of Hungerford (1948), Sailer (1948), and Brooks and Kelton (1967). Figure 1 illustrates the morphological terms used in these keys. Records of previous collections from Wis¬ consin and from its neighboring states are summarized from Hungerford (1948) and Sailer (1948), since there have been no records published for these states since 1948. Descriptions and illustrations of the various species are not included, because Hun¬ gerford and Sailer have thoroughly described and illustrated all of the species. Figure 2. Collections of Wisconsin Corixidae, 1962-1968, of the genera Calli- corixa, Cenocorixa , Corisella, Palmacorixa, Ramphocorixa, and Trichocorixa. 206 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figure 3. Collections of Wisconsin Corixidae, 1962-1968, of the genus Hespero- corixa. Key to Wisconsin genera 1. Rostrum without transverse grooves; pronotum without trans¬ verse dark bands _ Cymatia Rostrum with transverse grooves; pronotum with transverse bands although they may be indistinct _ 2 2. Entire hemelytral pattern usually effaced ; upper surface of male pala deeply incised; vertex of male acuminate; both sexes with palar claw serrate at base ; less than 5.5 mm long _ Ramphocorixa Hemelytral pattern distinct, although limited areas may be ef¬ faced in some species _ 3 1970] Hilsenhoff — Corixidae of Wisconsin 207 • S. bicoloripennis ° S. defecto ©S. mulleWensis Figure 4. Collections of Wisconsin Corixidae, 1962-1968, of the genus Sigara. 0 • S. johnutoni OS. knighti ©S. Irantfifloroto 3. Small shining corixids, the males with sinistral asymmetry ; apex of clavus not, or scarcely, exceeding a line drawn through costal margins at nodal furrows _ Trichocorixa Male asymmetry dextral ; apex of clavus plainly exceeding a line drawn through costal margins at nodal furrows _ 4 4. Pruinose area at base of claval suture short and broadly rounded at apex,, usually about % as long as postnodal pruinose area; prothoracic lobe truncate _ Hesperocorixa Pruinose area at base of claval suture narrowly rounded or pointed at apex and almost as long as postnodal pruinose area; prothoracic lobe rounded _ 5 208 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 p • • • 1 ' 1 y~i. • i ~~p. Figure 5. Collections of Wisconsin Corixidae, 1962-1968, of the genus Sigam . 5. Markings on clavis transverse, those on corium transverse, longitudinal, or reticulate _ 6 Markings on clavus and corium narrow and broken, usually open reticulate with many interconnections _ 7 6. Corial pattern transverse and with little contrast; male strigil absent; male pala with two rows of pegs _ Callicorixa Corium usually with contrasting pattern, either transverse, longitudinal, or reticulate; male strigil present; male pala with one row of pegs (2 exceptions) _ Sigara 1970] Hilsenhoff — Corixidae of Wisconsin 209 7. Rear margin of head sharply curved, embracing a very short pronotum; interocular space much narrower than the width of an eye _ Palmacorixa Rear margin of head gently curved; interocular space about equal to the width of an eye _ 8 8. Smooth, shining insects ; male pala triangular ; prothoracic lobe tapering to a narrowly rounded apex _ Corisella Rastrate, hairy species _ 9 9. Eyes protuberant with inner anterior angles broadly rounded; postocular space broad _ Dasycorixa Eyes normal; postocular space narrow _ Cenocorixa Callicorixa White 1873 Two species of Callicorixa probably occur in Wisconsin, but only C. audeni was collected. The second species, C. alaskensis Hunger- ford 1926, has been recorded from Michigan, Pennsylvania, New York;, and New Hampshire to the east of Wisconsin, and from Utah, Montana, and Wyoming to the west. It has also been found in most of the Canadian Provinces. Key to Wisconsin species 1. First tarsal segment of hind leg unicolor ous _ C. audeni First tarsal segment of hind leg infuscated on distal third _ _ C. alaskensis Callicorixa audeni Hungerford 1928 Distribution and abundance : Common in the northern half of the state (Fig. 2B), with three being collected as far south as Adams County. Identification : This species can be separated from other Calli¬ corixa that may occur in Wisconsin by the lack of a dark spot on the first tarsal segment of the hind leg. The black prothoracic lobe of almost all Wisconsin specimens serves to separate this species from all other Wisconsin corixids. The females may be most easily confused with female Sigara alternata (which may have a smoky prothoracic lobe), but can be distinguished by their longer, more acutely pointed metaxyphus, by a slightly wider mesoepimeron, and by light markings that often extend all the way across the corium. Collection Records: Adams Co. 1. 3, 2 $ ; Ashland Co. 19 3, 12 $ ; Barron Co. 1 $ ; Burnett Co. 1 $ ; Chippewa Co. 1 $ ; Clark Co. 1 3 , 1 $ ; Douglas Co. 8 3,7 2 ; Eau Claire Co. 13,29; Florence Co. 54 3 , 65 $ ; Forest Co. 17 3,20 $ ; Lincoln Co. 14 3,30 $ ; 210 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Marathon Co. 15 3 , 22 9 ; Marinette Co. 2 $ ; Oconto Co. 1 3 ; Oneida Co. 5 3„ 8 $ ; Price Co. 10 3 , 14 2 ; Rusk Co. 6 3 , 10 2 ; Sawyer Co. 4 3,4 2 ; Taylor Co. 9 3,9 2 ; Vilas Co. 12 3 , 9 2 ; Washburn Co. 1 2 . Totals: 398 individuals, 71 collections, 21 counties. Previous Records: None. Recorded from Michigan and Minne¬ sota. Cenocorixa Hungerford 1948 Only two species, C. dakotensis and C. utahensis, have been col¬ lected in Wisconsin, but both are rare. A third species, C. bifida (Hungerford) 1926, has been collected nearby in Minnesota and could occur in Wisconsin. Both Wisconsin records were from large lakes in the extreme northwest, and perhaps intensive collecting of such habitats would yield additional specimens and specimens of C. bifida as well. Key to Wisconsin species 1. Last tarsal segment of hind leg black or dark brown ; hind femur pubescent for about one-third its length _ C. dakotensis Last tarsal segment pale; hind femur pubescent for at least 40% of its length _ 2 2. Shining costal area just anterior to nodal furrow longer than middle tarsus; male peg row entire _ C. bifida Shining costal area just anterior to nodal furrow equal to middle tarsus in length; peg row of male pala divided _ C. utahensis Cenocorixa dakotensis (Hungerford) 1928 Distribution and Abundance: Apparently rare in the western part of the state (Fig. 2A). Collection Records: Douglas Co. 1 2 . Totals: 1 individual, 1 collection, 1 county. Previous Records : None. Reported from Illinois and Minnesota. Cenocorixa utahensis (Hungerford) 1925 Distribution and Abundance: Apparently rare in the western part of the state (Fig. 2A) . Collection Records: Douglas Co. 1 3. Totals: 1 individual, 1 col¬ lection, 1 county. Previous Records: None. Reported from Iowa. 1970] 211 Hilsenhoff — Corixidae of Wisconsin Corisella Lundblad Two species have been collected in Wisconsin, and it is unlikely that any others occur in this state. The members of this genus are rare in Wisconsin. Key to Wisconsin species 1. Less than 6.5 mm long; hind tarsus embrowned _ C. tarsalis More than 6.5 mm long; hind tarsus pale; pattern of clavis effaced at inner, basal angle _ C. edulis Corisella edulis (Champion) 1901 Distribution and Abundance: Rare in Wisconsin, it has been col¬ lected only in the southwestern part of the state (Fig. 2A). Collection Records: Dane Co. 1 $ ; Lafayette Co. 2 $, 6 $. Totals: 9 individuals, 2 collections, 2 counties. Previous Records: None. Reported from Iowa and Minnesota. Corisella tarsalis (Fieber) 1851 Distribution and Abundance: Rare in Wisconsin, this species ap¬ pears to be confined to the southern two-thirds of the state (Fig. 2A) . Collection Records: Columbia Co. 1 $, 2 $ ; Door Co. 1 S ; Wau¬ kesha Co. 2 $ . Totals: 6 individuals, 4 collections, 3 counties. Previous Records: “Parco” (probably Portage Co.) 1 $ . No records from adjoining states. Cymatia Flor 1860 The only described North American species, C. americana Hus¬ sey 1920, has not been found in Wisconsin but should occur in the northern half of the state. It has been reported in Michigan and Minnesota. Dasycorixa Hungerford 1948 One species, D. hybrida (Hungerford) 1926, has been collected in Minnesota and could occur in the northwestern part of Wisconsin. Hesperocorixa Kirkaldy 1908 Twelve species of Hesperocorixa have been collected in Wiscon¬ sin, eleven in this present study. Four of these species are very common. One additional species, H. nitida (Fieber) 1851, probably 212 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 occurs in Wisconsin, since it has been found in all four of the neighboring states. Its distribution is generally southern,, and it would most likely be found in the southern counties of the state. Key to Wisconsin species 1. Mesoepimeron at level of scent gland osteole as broad or broader than the lateral lobe of the pro thorax _ 2 Mesoepimeron plainly narrower than the lateral lobe of the prothorax _ 6 2. Mesoepimeron at level of scent gland osteole about equal in width to the lateral lobe of the prothorax; a conspicuous V- shaped yellow band bordering the apex of the corium _ _ H. atopodonta Mesoepimeron plainly broader than the prothoracic lobe _ 3 3. 8 mm long or longer ; tip of metaxyphus blunt or truncated _ _ H. kennicottii 7.5 mm long or less ; tip of metaxyphus pointed _ 4 4. Dorsal surface of hind femur with two stout spines _ _ H. minor ella Dorsal surface of hind femur armed with many spines _ 5 5. Corial pattern crossbanded _ H. michiganensis Corial pattern in longitudinal series _ H. semilucida 6. Pattern of hemelytra reticulate; pronotum nonrastrate _ - H. laevigata Pattern of hemelytra not reticulate; pronotum rastrate _ 7 7. Pronotal disc short, less than half as long as wide _ _ H. scahricula Pronotal disc more than half as long as wide _ 8 8. Color pattern of corium effaced laterally _ H. lucida Color pattern normal _ 9 9. Pale bands of corium beyond hemelytral suture forming slender transverse series. Membrane not plainly separated from corium - II. vulgaris Pala bands of corium beyond hemelytral suture in an inter¬ rupted transverse series. Membrane may be distinctly sepa¬ rated from corium _ 10 10. Hind femur with a row of about 10 spines ventrally on distal portion of rear margin _ H. nitida Hind femur with only about 6 spines on rear margin _ 11 11. Corium and membrane not separated by a coalescing of the pale figures; upper distal angle of male pala acutely, obliquely produced _ H. obliqua Corium and membrane separated by a coalescing of the pale figures _ 12 213 1970] Hilsenhoff — Corixidae of Wisconsin 12. Interocular space almost equal to the .width of an eye ; metaxy- phus as broad as long; male pala rounded at tip; male strigil oval _ _ _ H. lobata Interocular space much narrower than width of an eye; met- axyphus longer than broad; male pala truncated at tip; male strigil very long _ H. interrupta Hesperocorixa atopodonta (Hungerford) 1927 Distribution and Abundance : This species is found throughout Wisconsin (Fig. 3B) and is very common in the northern two- thirds of the state. Identification: The mesoepimeron that is the same width as the prothoracic lobe and the conspicuous V-shaped yellow band border¬ ing the apex of the corium readily distinguish this species. The male pala with the last peg out of line is also distinctive. Collection Records: Adams Co. 14 <2, 11 2 ; Ashland Co. 26 £, 30 2 ; Barron Co. 1 £, 1 2 ; Burnett Co. 9 £ , 2 2 ; Chippewa Co. 16 £ , 15 2 ; Clark Co. 5 £ , 4 2 ; Columbia Co. 1 £ , 2 2 ; Crawford Co. 2 £ ; Dane Co. 12 £ , 9 2 ; Douglas Co. 32 £ , 33 $ ; Dunn Co. 7 £ , 8 2 ; Eau Claire Co. 29 £ , 33 2 ; Florence Co. 89 £ , 92 2 ; Fond du Lac Co. 2 £ , 4 2 ; Forest Co. 20 £„ 22 $ ; Grant Co. 1 $ ; Green Co. 1 £ , 1 $ ; Green Lake Co. 1 $ ; Iowa Co. 5 £ , 3 $ ; Juneau Co. 65 £, 93 $ ; Kenosha Co. 1 £ ; Langlade Co. 1 £ ; Lin¬ coln Co. 77 £,58 2 ; Manitowoc Co. 1 2 ; Marathon Co. 45 £, 36 2 ; Marinette Co. 1 £ , 2 2 ; Marquette Co. 1 2 ; Milwaukee Co. 5 <2,5 2; Monroe Co. 1 <2, 4 2 ; Oconto Co. 7 £,12 2 ; Oneida Co. 9 £ , 10 2 ; Ozaukee Co. 1 <2 ; Polk Co. 5 <2 , 8 2 ; Portage Co. 9 £, 12 2 ; Price Co. 25 $ , 39 2 ; Racine Co. 1 £ ; Richland Co. 3 <2 , 5 2 ; Rusk Co. 40 $ , 45 2 ; Sauk Co. 2 $ , 4 2 ; Sawyer Co. 8 $ ; Taylor Co. 9 $ , 9 2 ; Trempealeau Co. 42 <2, 36 2 ; Vilas Co. 15 £ , 16 2 ; Washburn Co. 7 £, 6 2 ; Waupaca Co. 17 £,92; Winne¬ bago Co. 22 £ , 28 2 ; Wood Co. 5 £, 4 2 . Totals: 1U09 individuals, 11*5 collections, U7 counties. Previous Records: Bayfield Co. 2 2 ; Dane Co. 6 £, 7 2 ; Douglas Co. 1 £, 1 2 ; Fond du Lac Co. 1 £ , 1 2 ; Polk Co. 1 2 ; Rusk Co. 1 2 ; Sauk Co. 2 2 ; St. Croix R. 3 2 ; Wisconsin 2 £ , 2 2 . Also reported from Michigan and Minnesota. Hesperocorixa interrupta (Say) 1825 Distribution and Abundance: The only Wisconsin record is one male collected at Beaver Dam (Dodge Co.) by W. E. Snyder in 1909 (Hungerford 1948). Most of the records for this species are from states south of Wisconsin, indicating that it would most likely occur in the southern part of the state. 214 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Identification: A series of specimens from the Snow Collection was examined. This species is similar to H. lobata, but can be sepa¬ rated from that species by its narrow interocular space, which is only % the width of an eye. In H. lobata the interocular space is about 12/13 the width of an eye. The metaxyphus is longer and more pointed than that of H. lobata. Males have a pala that is truncate at the tip as compared to the rounded pala of H. lobata, and they have a very elongate and large strigil. H. interrupta can be separated from the smaller H. nitida by the characters given in the key. Previous Records: Dodge Co. 1 8. Also reported from Illinois and Michigan. Hesperocorixa kennicottii (Uhler) 1897 Distribution and Abundance: This species is common in all but the extreme southeastern corner of the state, where it was not collected but probably occurs (Fig. 3C). Identification: This species is very distinctive. It has a very wide, pale mesoepimeron, a metaxyphus that is truncate or broadly rounded at the tip, and a golden-brown membrane that is almost devoid of markings and is separated from the corium by the wide, yellow, V-shaped border of the corium. Collection Records: Adams Co. 1 8,2 2 ; Ashland Co. 1 2 ; Barron Co. 1 8 ; Columbia Co. 2 8 ; Crawford Co. 4 8,2 $ ; Doug¬ las Co. 23 8, 34 2 ; Dunn Co. 1 8 , 3 2 ; Florence Co. 2 8 , 5 $ ; Forest Co. 3 2 ; Iowa Co. 1 8 ; Juneau Co. 1 $ ; Langlade Co. 1 8 ; Lincoln Co. 20 <2,11 $ ; Marathon Co. 18 8 , 28 $ ; Marquette Co. 1 8 ; Oconto Co. 1 8,1 $ ; Oneida Co. 2 8,1 2 ; Polk Co. 1 2 ; Portage Co. 1 8,2 2 ; Price Co. 1 2 ; Rusk Co. 2 8,1 2 ; Sauk Co. 2 2 ; Sawyer Co. 12,2 2 ; Taylor Co. 1 8,2 2 ; Trempealeau Co. 3 8 ; Washburn Co. 8 8 , 3 2 ; Waupaca Co. 1 8 , 8 2 ; Winne¬ bago Co. 47 £,50 2 ; Wood Co. 1 8,2 2 . Totals: 309 individuals, 51 collections, 29 counties. Previous Records: Dane Co. 44 8 , 75 2 ; Douglas Co. 1 8 . Also reported from Illinois, Michigan and Minnesota. Hesperocorixa laevigata (Uhler) 1893 Distribution and Abundance: Although apparently rare in Wis¬ consin, having been collected from only two sites (Fig. 3C), at one of these sites it was abundant and thousands could have been collected. This is a common western corixid, and Wisconsin is east of its principal range. 1970] Hilsenhoff — Corixidae of Wisconsin 215 Identification: The reticulate pattern and non-rastrate pronotum set this species apart from all other Hesperocorixa. Collection Records: Adams Co. 1 $ ; Iowa Co. 24 S, 36 2. Totals: 61 individuals, 2 collections, 2 counties. Previous Records : St. Croix R. 1 $ (Hungerford 1948) . Also re¬ ported from Illinois and Minnesota. Hesperocorixa lobata (Hungerford) 1925 Distribution and Abundance: Although collected in only live northern counties in this study (Fig. 3C) , its distribution in the United States (Hungerford 1948) and a previous collection from Dane County indicate that it probably occurs throughout the state. Identification: The separation of this species from H. interrupta was discussed under H. interrupta. The male pala is rounded at the tip, and is not as illustrated by Hungerford (1948). A series of specimens from the Snow Collection was studied, as well as those collected in Wisconsin. Collection Records: Barron Co. 3 $ ; Chippewa Co. 1 & ; Florence Co. 4 3,3 2 ; Polk Co. 1 $ , 1 $ ; Washburn Co. 2 $ . Totals: 15 individuals, 5 collections, 5 counties. Previous Records: Dane Co. 1 S, 1 2 . Also reported from Mich¬ igan and Minnesota. Hesperocorixa lucida (Abbott) 1916 Distribution and Abundance: A distinct rarity in Wisconsin, H. lucida was collected from only two counties in northern Wiscon¬ sin (Fig. 3D). Its distribution to the south of Wisconsin indicates that it may be found in all parts of the state. Identification: The lack of markings on the clavis and the effaced markings of the corium separate this species from other Hespero¬ corixa. Collection Records: Douglas Co. 1 3, 1 2 ; Lincoln Co. 1 $ . Totals: 3 individuals, 2 collections, 2 counties. Previous Records: None. Reported from Illinois and Michigan. Hesperocorixa michiganensis (Hungerford) 1926 Distribution and Abundance: Although it was also collected in the extreme south, the distribution is mostly throughout the north¬ ern two-thirds of the state (Fig. 3A). In the north this is a very common species. Identification : The mesoepimeron is pale,, much lighter than either H. minorella or H. semilucida , the other two small Hespero- 216 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 corixa with a wide mesoepimeron. The pale markings on the mem¬ brane are also more distinct than in either of these two species. Collection Records: Adams Co. 6 3,5 2 ; Ashland Co. 45 3, 20 2 ; Barron Co. 4 3, 10 2 ; Burnett Co. 31 3 , 15 $ ; Clark Co. 7 3,5 $ ; Dodge Co. 1 3 ; Douglas Co. 10 3,8.$; Eau Claire Co. 2 3, 9 $ ; Florence Co. 203 3, 197 2 ; Forest Co. 82 3 , 99 2 ; Green Co. 1 3,2 2 ; Juneau Co. 1 2 ; La Crosse Co. 1 3 ; Langlade Co. 3 2 ; Lincoln Co. 72 3, 84 2 ; Manitowoc Co. 2 2 ; Marathon Co. 42 3,29 2 ; Marinette Co. 2 3,1 2 ; Marquette Co. 33,1 2 ; Monroe Co. 1 2 ; Oconto Co. 30 3 , 12 2 ; Oneida Co. 22 3 , 17 2 ; Outagamie Co. 1 2 ; Polk Co. 3 3,6 2 ; Portage Co. 4 3,2 2 ; Price Co. 53 3 , 60 2 ; Rusk Co. 33 3 , 5 2 ; Sauk Co. 1 3 ; Sawyer Co. 7 3 , 12 2 ; Taylor Co. 14 3, 7 2 ; Trempealeau Co. 10 3, 22 2 ; Vilas Co. 26 3, 44 2 ; Waupaca Co. 4 3,2 2 ; Winnebago Co. 3 3, 4 2 ; Wood Co. 7 3,7 2 . Totals: 1U22 individuals, 120 collections, 35 counties. Previous Records: Dane Co. 1 3,3 2 ; Douglas Co. 3 3, 7 2 ; Wisconsin 2 2 . Also reported from Michigan and Minnesota. Hesperocorixa minor ella (Hungerford) 1926 Distribution and Abundance: A very common species of Hespero¬ corixa that is widely distributed throughout the northern two- thirds of the state (Fig. 3A) . It is most abundant in the far north. Identification: This very dark species can be distinguished by the two spines on the dorsal surface of the hind femur. Collection Records: Adams Co. 1 3„ 10 2 ; Ashland Co. 18 3, 18 2 ; Barron Co. 1 3 ; Burnett Co. 3 3,7 2 ; Chippewa Co. 2 3, 2 2 ; Clark Co. 2 3,5 2 ; Douglas Co. 17 3, 39 2 ; Dunn Co. 2 3, 3 2 ; Eau Claire Co. 36 3 , 54 2 ; Florence Co. 434 3, 458 2 ; Forest Co. 195 3, 260 2 ; Iron Co. 8 3 , 20 2 ; Jackson Co. 3 3, 4 2 ; Juneau Co. 18 3 , 14 2 ; Langlade Co. 1 2 ; Lincoln Co. 44 3 , 78 2 ; Manitowoc Co. 1 2 ; Marathon Co. 14 3,30 2 ; Marinette Co. 4 3,3 2 ; Marquette Co. 1 3 ; Monroe Co. 1 3,1 2 ; Oconto Co. 38 3 , 66 2 ; Oneida Co. 62 3„ 119 2 ; Outagamie Co. 1 2 ; Polk Co. 1 3 ; Portage Co. 7 3,6 2 ; Price Co. 29 3 , 52 2 ; Rusk Co. 62 3 , 58 2 ; Sauk Co. 1 2 ; Sawyer Co. 25 3 , 34 2 ; Shawano Co. 1 3,1 2 ; Taylor Co. 12 3, 12 2 ; Trepealeau Co. 1 2 ; Vilas Co. 198 3 , 231 2 ; Washburn Co. 11 3 , 21 2 ; Waupaca Co. 3 2 ; Waushara Co. 2 3, 1 2 ; Winnebago Co. 5 3, 6 2 ; Wood Co. 4 3, 7 2 . Totals: 2889 individuals, 125 collections, 39 counties. Previous Records: Douglas Co. 1 3 ; Wisconsin (no sex or number) . Also reported from Michigan and Minnesota. 1970] Hilsenhoff — Corixidae of Wisconsin 217 Hesperocorixa obliqua (Hungerford) 1925 Distribution and Abundance: This species was collected only in the extreme southern part of the state, where it is uncommon (Fig. BD) . Identification: The acute and obliquely produced upper distal angle of the male pala is distinctive, but the females closely re¬ semble those of the much more abundant H. vulgaris. The pale transverse lines of the corium are shorter in H. obliqua , and seldom traverse the entire coriurq, while the longer lines of H. vulgaris extend entirely across the corium in the basal third. Collection Records: Dane Co. 5 S, 4 $ ; Kenosha Co. 5 $ , 9 $ ; Racine Co. 1 $ ; Rock Co. 1 S, 1 $ . Totals: 26 individuals, 6 col¬ lections, A counties. Previous Records: None. Reported from Illinois, Iowa, Michigan and Minnesota. Hesperocorixa scabricula (Walley) 1936 Distribution and Abundance: Apparently rare in Wisconsin, this species was collected only from one site in Florence County (Fig. 3D). Its occurrence in Illinois, Michigan, and Minnesota suggests that it may occur throughout Wisconsin. Identification: Readily distinguished by its large size and the very short pronotal disc. Collection Records: Florence Co. 5 $ . Totals: 5 individuals, 1 col¬ lection, 1 county. Previous Records: None. Reported from Illinois, Michigan, and Minnesota. Hesperocorixa semilucida (Walley) 1930 Distribution and Abundance: This uncommon species is appar¬ ently distributed throughout the southern half of the state (Fig. 3D), although it has not been collected in the extreme southeastern counties. Identification: The small size and wide mesoepimeron distinguish this species,, H. minorella, and H. michiganensis. The mesoepimeron is dusky, as in H. minorella , but the arrangement of the corial pat¬ tern into a definite longitudinal series separates it from both H. minorella and H. michiganensis. Collection Records: Crawford Co. 3 $ , 7 $ ; Green Co. 1 $ ; Richland Co. 6 4 ? ; Trempealeau Co. 1 $ , 1 $ ; Waupaca Co. 3 $ ; Winnebago Co. 1^,1 $ . Totals: 28 individuals, 9 collections, 6 counties. Previous Records: None. Reported from Illinois and Michigan. 218 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Hesperocorixa vulgaris (Hungerford) 1925 Distribution and Abundance: This species is very common throughout the state (Fig. 3B) . Identification: The narrow metaxyphus, the lack of any pale figures separating the corium from the membrane, and the long, narrow, transverse pale markings of the corium distinguish this species. The truncated tip of the male pala is slightly indented at the middle. Collection Records: Adams Co. 10 £,10 2 ; Ashland Co. 1 £, 4 $ ; Barron Co. 21 £,29 2 ; Brown Co. 1 £ ; Buffalo Co. 1 £ ; Burnett Co. 5 £„ 5 $ ; Chippewa Co. 3 £,11 $ ; Clark Co. 1 £, 6 2 ; Columbia Co. 10 £, 10 $ ; Crawford Co. 1 $ ; Dane Co. 39 £, 22 $ ; Dodge Co. 7 £ , 11 $ ; Door Co. 1 £ ; Douglas Co. 16 £,17 $ ; Eau Claire Co. 34 £, 42 2 ; Dunn Co. 3 £ ; Florence Co. 12 £, 112; Forest Co. 5 £ , 5 2 ; Fond du Lac Co. 1 2 ; Grant Co. 2 £, 2 2 ; Green Co. 16 £,13 2 ; Green Lake Co. 3 £, 42 ; Iowa Co. 8 £ , 12 2 ; Jackson Co. 4 £ , 1 2 ; Juneau Co. 5 £, 11 2 ; Kenosha Co. 24 £ , 13 2 ; La Crosse Co. 1 £ , 1 2 ; Langlade Co. 2 £ „ 3 2 ; Lincoln Co. 6 £ , 7 2 ; Manitowoc Co. 1 £ , 2 2 ; Marathon Co. 12 £ , 6 2 ; Marquette Co. 5 £, 1 2 ; Milwaukee Co. 30 £,39 2 ; Monroe Co. 3 £, 2 2 ; Oconto Co. 2 £ , 9 2 ; Oneida Co. 1 £, 1 2 ; Outagamie Co. 6 2 ; Ozaukee Co. 1 £,2 2 ; Polk Co. 6 £, 3 2 ; Portage Co. 2 £ ; Price Co. 6 £,15 2 ; Racine Co. 22 £,12 2 ; Richland Co. 3 £ , 5 2 ; Rock Co. 2 £, 6 2 ; Rusk Co. 6 £„ 17 2 ; Sauk Co. 24 £ , 33 2 ; Sawyer Co. 6 £ , 7 2 ; Taylor Co. 3 £ , 5 2 ; Trempealeau Co. 25 £,22 2 ; Vernon Co. 2 £, 1 2 ; Vilas Co. 2 £, 2 2 ; Walworth Co. 28 £ , 45 2 ; Washburn Co. 1 £, 1 2 ; Wash¬ ington Co. 1 £ , 1 2 ; Waukesha Co. 4 £ , 8 2 ; Waupaca Co. 2 £, 2 2 ; Waushara Co. 3 £, 2 2 ; Winnebago Co. 52 £, 78 2 ; Wood Co. 2 2 . Totals: 1083 individuals, 160 collections, 59 counties. Previous Records: Dane Co. 1 £,19 2 ; Dodge Co. (no num¬ bers) ; Douglas Co. 1 £, 2 2 ; St. Croix R. 8 £ , 6 2 . Also reported from Illinois, Iowa, Michigan and Minnesota. Palmacorixa Abbott 1912 Three species of Palmacorixa occur throughout Wisconsin, but none of them is very common. The males of the three species are easily distinguished, but the females are very difficult to separate, especially those of P. nana and P. buenoi. The identification of the females collected in this study is based on the following key, and on other criteria listed under each species, but the separation of P. nana from P. buenoi remains uncertain. 1970] Hilsenhoff — Corixidae of Wisconsin 219 Key to Wisconsin species 1. The pronotal disc with well marked anterolateral depressions; male pala very broad, almost disc-like, with poorly defined pegs _ P . gillettei Anterolateral depression on pronotum weak or absent; male pala elongate _ _ 2 2. Middle femur of male with a longitudinal row or pegs on its ventral surface; female less than 5.2 mm long _ P. nana Middle femur of male without a row of pegs ; female 5.4 mm or longer _ P. buenoi Palmacorixa buenoi Abbott 1913 Distribution and Abundance : This is the most common species of Palmacorixa in Wisconsin. It has been collected throughout most of the state, being fairly common in the southeastern half and less common in the northwest (Fig. 2C). Identification: The females of this species are difficult to sepa¬ rate from Palmacorixa nana. Size is perhaps the most important criteria, with P. nana females being less than 5.2 mm long and P. buenoi females being 5.4 to 6.5 mm. Also, the dark posterior border of the pronotum is usually wider than in P. nana. The pronotum lacks the strong antero-lateral depressions that are found in females of P. gillettei. Collection Records: Adams Co. 2 $ ; Brown Co. 2 $ ; Columbia Co. 12 $ , 19 $ ; Dane Co. 2 $ ; Douglas Co. 1 ? ; Eau Claire Co. 2 $ ; Green Lake Co. 1 $91 $ ; Juneau Co. 5 S , 7 $ ; Kenosha Co. 4 $ ; Kewaunee Co. 12 £ , 7 $ ; La Crosse Co. 1 $ ; Lafayette Co. 2 $ ; Oconto Co. 1 $ ; Oneida Co. 1 $ ; Pepin Co. 1 $ ; Pierce Co. 1 $ ; Polk Co. 1 $ ; Price Co. 2 $ ; Racine Co. 1 S ; Rock Co. 3 $ ; Sauk Co. 2 $ , 2 $ ; Sawyer Co. 1 ? ; Washington Co. 2^,3 2 ; Waupaca Co. 1 $ ; Waushara Co. 1 $ , 3 $ . Totals: 106 individuals, SO collections, 25 counties. Previous Records: None. Reported from Iowa, Michigan and Minnesota. Palmacorixa gillettei Abbott 1912 Distribution and Abundance: Scattered records from throughout the state indicate a statewide distribution, with the possible excep¬ tion the extreme southeastern counties (Fig. 2C). This species is fairly common, but less common than P. buenoi. Identification: The males are easily distinguished by their dilated and flattened palae. The females can be identified by the strong 220 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 anterolateral depressions of the pronotum and the wide, dark, posterior border of the pronotum. Collection Records: Ashland Co. 1 $ ; Barron Co. 1 2 ; Dane Co. 2 2,8 $ ; Douglas Co. 1 2 ; Florence Co. 2 $ ; Iowa Co. 8 2,3 $ ; Lafayette Co. 2 $ ; Oconto Co. 2 2,2 $ ; Outagamie Co. 1 $ ; Rich¬ land Co. 1 $ ; Taylor Co. 2 2,6 $ ; Vilas Co. 1 2 ; Waupaca Co. 1 2,2 $ ; Waushara Co. 1 $ . Totals: U2 individuals, 19 collections, 11> counties. Previous Records: None. Reported from Michigan, Iowa and Minnesota. Palmacorixa nana Walley 1930 Distribution and Abundance: The scattered records indicate that this uncommon species probably occurs throughout the state (Fig. 2C) . Identification: The row of pegs on the middle femur separates the males from those of P. buenoi. The females can be distinguished by their small size (less than 5.2 mm) and the very narrow, often interrupted, dark posterior border of the pronotum. Collection Records: Dane Co. 1 $ ; Florence Co. 1 2 ; Forest Co. 12,6$; Manitowoc Co. 12,1 $ ; Outagamie Co. 1 $ ; Taylor Co. 7 2,9 $ . Totals: 28 individuals, 6 collections, 6 counties. Previous Records: None. Reported from Michigan and Minnesota. Ramphocorixa Abbott 1912 Only one species occurs in Wisconsin. Ramphocorixa acuminata (Uhler) 1897 Distribution and Abundance: This species is rare in Wisconsin, with but a single specimen having been collected. This individual was collected with a black-light trap at Madison (Fig. 2B). Identification: The effaced pattern of the clavis and corium of this species serves to separate it from all other small Wisconsin corixids. The male is also recognized by its accuminate vertex. Collection Records: Dane Co. 1 2. Totals: 1 individual, 1 collec¬ tion, 1 county. Previous Records: None. Reported from Illinois and Minnesota. Sigara Fabricius 1775 This is the most common corixid genus in Wisconsin, 23 species having been collected in the state. The only species likely to occur in Wisconsin that have not been collected are S. hubbelli (Hungerford) 1928 and S. modesta (Abbott) 1916. Both are distributed through- 1970] Hilsenhoff — Corixidae of Wisconsin 221 out the southeastern United States and have been collected in the neighboring states of Illinois, Iowa and Minnesota. S. hubbelli is closely related to S. dejecta while S. modesta resembles S. grosso- lineata. Both can be separated by the characters given in the key. Key to Wisconsin species 1. Pronotal disk with median longitudinal pale line _ 2 Pronotal disk without a median pale longitudinal line _ 7 2. Tip of mesoepimeron as close or closer to scent gland osteole than to mesosternum _ 8 Scent gland osteole remote from tip of mesoepimeron, farther than distance from tip to mesosternum _ 5 3. Hemelytra nearly black; pale markings wide but obscure, transverse on clavis and somewhat longitudinal on corium ; male pala thickened, with a prominent keel on the outside __ _ _ _ S. variabilis Markings on hemelytra narrower and more distinct _ 4 4. Male pala with a single row of pegs ; female abdomen with anal lobes not notched on inner ventral margin _ S. johnstoni Male pala with two rows of pegs; female with anal lobes notched on inner ventral margin _ S. knighti 5. Pronotum and hemelytra boldly cross-barred; vertex produced beyond the eye curve in both sexes _ S. transfigurata Pattern less striking; vertex not noticeably produced _ 6 6. Pattern of membrane effaced or indistinct; pale figures on corium and clavis transverse _ S. compressoidea Pattern of membrane usually distinct; pale figures on corium and distal half of clavis arranged more or less longitudinally _ S. mackinacensis 7. Large species, greater than 7.0 mm long _ 8 Small species, less than 7.0 mm long _ 11 8. Metaxyphus broad, truncated or notched at the tip; claw of pala serrate at base in both sexes _ S. decorata Metaxyphus pointed; palar claw normal _ 9 9. Palae of both sexes with only 14 to 16 lower palmar hairs __ - S. decoratella Palae with from 18 to 22 lower palmar hairs _ 10 10. Interocular space plainly narrower than width of an eye; hemelytra dark, with the pale markings of the corium and distal half of clavis arranged in definite longitudinal series _ - S. penniensis Interocular space equal to width of an eye; pale markings of corium and clavis bold and transverse; vertex of male pro¬ duced; palae long and slender _ _ _ S. conocephala 222 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 11. Corial pattern in definite longitudinal series _ 12 Corial pattern not in definite longitudinal series _ 16 12. Clavus and corium with pale lines in wavy longitudinal series ; hypoocular suture ending laterad of middle line of eye _ I _ S. douglasensis Claval lineations not in wavy longitudinal series _ 18 13. Small, less than 4.3 mm long; antennae usually 3-segmented _ _ S. lineata Larger, greater than 4.5 mm long; antennae 4-segmented __14 14. Scent gland osteole remote from tip of mesoepimeron _ _ S. trilineata Scent gland osteole near tip of mesoepimeron _ 15 15. Corial lineations distinct; male pala with two rows of pegs; female pala short; anal lobes of female only very slightly notched on inner ventral margin _ S. mullettensis Corial lineations fairly distinct; male pala with one row of pegs _ 23 16. Metaxyphus longer than broad _ 17 Metaxyphus not longer than broad _ 19 17. Mesoepimeron at level of scent gland osteole about equal in width to lateral lobe of prothorax _ S. dolabra Mesoepimeron at level of scent gland osteole much broader than width of lateral lobe of prothorax _ 18 18. Pale bands on base of clavus entire, bands on corium plainly transverse; dorsal surface of hind femur with only 3 or 4 pegs _ S. solensis Pale bands on base of clavus more or less broken and confused ; dorsal surface of hind femur with two or three rows of pegs _ S. signata 19. Scent gland osteole nearer lateral bend of mesoepimeron than tip _ 20 Scent gland osteole near tip of mesoepimeron _ 22 20. Head with median longitudinal brown line ; mesoepimeron with a deep incision at or near the lateral bend _ S. mathesoni Head without line; mesoepimeron without incision _ 21 21. Osteole almost in lateral bend of mesoepimeron, at least 4/5 from tip ; metaxyphus with a truncated point grossolineata Osteole not so far laterad, 1/2-3/5 from tip ; metaxyphus with a rounded point _ _ S. modesta 22. Corial pattern in a more or less definite longitudinal series __23 Corial pattern not in a longitudinal series _ 24 23. Pronotal disc laterally reduced; 4 to 6 pegs on dorsal surface of hind femur; pattern of membrane obscure _ _ S. hubbelli Pronotal disc not reduced laterally; row of at least 12 pegs on 1970] Hilsenhoff — Corixidae of Wisconsin 223 dorsal surface of hind femur; pattern of membrane distinct; distal pegs of male pala becoming widely separated ; anal lobes of female notched on inner ventral margin ; pala of female long _ _ _ S. dejecta 24. Metaxyphus small and rounded at tip ; male pala with peg row close to palm; anal lobes of female not notched _ _ S. bicoloripennis Metaxyphus pointed at an angle of less than 90 degrees _ 25 25. Pronotum crossed by 5 or 6 dark bands _ S. macropala Pronotum crossed by 8 or 9 dark bands _ S. alternata Sigara alternata (Say) 1825 Distribution and Abundance : The most common Wisconsin corixid, this species is very common in the northern half of the state and abundant in the southern half. It has been collected in almost every county (Fig. 4A) . Identification: The short, pointed metaxyphus forms an angle at the tip of about 70°. This, the alternate dark and pale transverse markings on the clavis, and the vermiform markings of the corium separate this species from others with a narrow mesoepimeron. Collection Records: Adams Co. 78 3, 114 9 ; Ashland Co. 4 3, 6 $ ; Barron Co. 12 3, 16 $ ; Bayfield Co. 1 9 ; Brown Co. 1 3„ 9 9 ; Buffalo Co. 10 3,6 9 ; Burnett Co. 14 3 , 34 9 ; Calumet Co. 2 3,4 $ ; Chippewa Co. 2 3,4 9 ; Clark Co. 2 3,3 $ ; Columbia Co. 104 3, 128 9 ; Crawford Co. 46 3 , 62 $ ; Dane Co. 155 3, 203 9 ; Dodge Co. 4 $ ; Door Co. 11 3,3 $ ; Douglas Co. 66 3, 93 $ ; Dunn Co. 3 3,6 $ ; Eau Claire Co. 24 3 , 30 9 ; Florence Co. 68 3, 77 9 ; Forest Co. 1 3,5 9 ; Fond du Lac Co. 1 3,6 9 ; Grant Co. 11 3, 13 9 ; Green Co. 40 3, 49 9 ; Green Lake Co. 7 3 , 23 9 ; Iowa Co. 31 3 , 22 9 ; Iron Co. 1 3 ; Jackson Co. 7 3, 10 9 ; Jefferson Co. 1 9 ; Juneau Co. 50 3 , 61 9 ; Kenosha Co. 70 3 , 93 9 ; Kewaunee Co. 1 3,8 9 ; La Crosse Co. 7 3 , 14 9 ; Lafayette Co. 3 3,7 9 ; Langlade Co. 2 3,1 9 ; Lincoln Co. 29 3„ 20 9 ; Manitowoc Co. 3 3,8 9 ; Marathon Co. 483, 54 9 ; Marquette Co. 9 3 , 17 9 ; Menominee Co. 1 3 ; Milwaukee Co. 61 3, 61 9 ; Monroe Co. 3 3 , 32 9 ; Oconto Co. 16 3, 28 9 ; Oneida Co. 9 3, 11 9 ; Outagamie Co. 6 3, 7 9 ; Ozaukee Co. 1 3,1 9 ; Pepin Co. 1 3 ; Polk Co. 1 9 ; Portage Co. 28 3 , 31 9 ; Price Co. 15 3, 14 9 ; Racine Co. 30 3 , 34 9 ; Richland Co. 13 3,9 9 ; Rock Co. 8 3,19 9; Rusk Co. 1 3 „ 1 9 ; St. Croix Co. 3 9 ; Sauk Co. 45 3 , 39 9 ; Sawyer Co. 4 3,9 9 ; Shawano Co. 2 3 ; Taylor Co. 12 3 , 10 9 ; Trempealeau Co. 78 3 , 92 9 ; Vernon Co. 11 3,6 9 ; Vilas Co. 3 3,8 9 ; Walworth Co. 59 3, 89 9 ; Washburn Co. 2 3 ; Washington Co. 1 3 ; Waukesha Co. 19 3 , 16 9 ; Waupaca Co. 6 224 Wisconsin Academy of Sciences, Arts and Letters [VoL 58 3 , 9 2 ; Waushara Co. 1 3 ; Winnebago Co. 17 3, 28 2 ; Wood Co. 6 3 , 6 $. Totals: 3168 individuals, 2U8 collections, 69 counties. Previous Records: Burnett Co. 1 $ ; Dane Co. 2 $ , 5 $ ; Douglas Co. 1 3,1 5 ; Fond du Lac Co. 1 $ ; Jackson Co. 1 2 ; Lafayette Co. 1 2 ; Sauk Co. 1 3„ 6 2 . Also reported from Illinois, Iowa, Michigan and Minnesota. Sigara bicoloripennis (Walley) 1936 Distribution and Abundance: This species is fairly common throughout the northern two-thirds of the state (Fig. 4B). One specimen was collected in Kenosha County in the extreme south¬ east. Identification: The small, rounded metaxyphus separates this species from S. alternata. The females might be confused with S. defecta, but the pala is shorter in S. bicoloripennis and the anal lobes are not notched on the mesal margin. Collection Records: Adams Co. 8 <2 , 20 2 ; Ashland Co. 1 3 ; Barron Co. 1 3,2 2 ; Burnett Co. 3 3,7 2 ; Douglas Co. 6 8,6 2 ; Florence Co. 39 $ , 44 2 ; Forest Co. 2 2 ; Juneau Co. 1 $ ; Kenosha Co. 1 $ ; Langlade Co. 1 2 ; Lincoln Co. 1 $ , 1 2 ; Marathon Co. 1 8 , 2 2 ; Oconto Co. 2 8 ; Oneida Co. 1 8,1 2 ; Outagamie Co. 6 8, 5 2 ; Polk Co. 6 8 , 3 2 ; Price Co. 1 8 , 3 2 ; Sawyer Co. 1 8 ; Taylor Co. 11 8, 11 2 ; Trempealeau Co. 9 3 , 10 2 ; Washburn Co. 1 8 ; Waupaca Co. 1 2 ; Winnebago Co. 1 2 . Totals: 220 in¬ dividuals, 3U collections, 23 counties. Previous Records: Dane Co. 3 3,12 2 . Also reported from Mich¬ igan and Minnesota. Sigara compressoidea (Hungerford) 1928 Distribution and Abundance: While common in the northern two- thirds of the state, it also has been collected in two extreme south¬ eastern counties (Fig. 5C) . Identification : The pale longitudinal line on the prothorax and the effaced membrane distinguish this species. It might be con¬ fused only with S. mackinacensis , but the lines on the basal third of the clavis are mostly entire and not zig-zag, and the corial pat¬ tern is transverse while that in S. mackinacensis is arranged in a longitudinal series. Collection Records: Adams Co. 2 3,1 2 ; Ashland Co. 33,1 2 ; Barron Co. 1 3 ; Bayfield Co. 1 2 ; Burnett Co. 1 3,2 2 ; Chippewa Co. 13,3 2 ; Clark Co. 9 3,6 2 ; Douglas Co. 2 3 ; Eau Claire Co. 3 3,1 2 ; Florence Co. 69 3 , 93 2 ; Forest Co. 5 2 ; Green Lake Co. 1 2 ; Juneau Co. 1 3 ; Kenosha Co. 1 2 ; Langlade Co. 3 3, 3 2 ; Lincoln Co. 41 3 , 37 2 ; Marathon Co. 82 3 , 109 2 ; Mar- 1970] HilsenhofS — Corixidae of Wisconsin 225 quette Co. 2 3,1 2 ; Oconto Co. 9 3 , 15 2 ; Oneida Co. 3 3 , 7 $ ; Polk Co. 6 $ ; Portage Co. 1 2 ; Price Co. 4 3, 18 $ ; Rusk Co. 6 £ , 5 2 ; Sauk Co. 3 3 , 1 $ ; Sawyer Co. 5 $ ; Taylor Co. 33 3 , 50 2 ; Vilas Co. 9 3, 17 2 ; Walworth Co. 1 3,1 2 ; Washburn Co. 1 3,1 2 ; Waupaca Co. 1 3 ; Waushara Co. 1 2 ; Wood Co. 8 3,1 2 . Totals: 692 individuals, 96 collections, 33 counties. Previous Records: Dane Co. 7 2 . Also reported from Michigan and Minnesota. Sigara conocephala (Hungerford) 1926 Distribution and Abundance: Fairly common in the northeast,, this species is apparently confined to the northern two-thirds of the state (Fig. 5A). Identification : The strongly produced vertex identifies the male, while the female can be recognized by her large size and unusually long palae. Collection Records: Adams Co. 1 3,1 2 ; Ashland Co. 1 3 ; Douglas Co. 1 3 ; Florence Co. 34 3 , 66 2 ; Forest Co. 2 3,5 2 ; Oneida Co. 1 $ ; Trempealeau Co. 1 3,2 2 ; Vilas Co. 1 3 . Totals: 116 individuals, 23 collections, 8 counties. Previous Records: None. Reported from Michigan and Minnesota. Sigara decorata (Abbott) 1916 Distribution and Abundance: This species is fairly common throughout the state (Fig. 4C) . Identification: S. decorata is easily distinguished by its large size and broadly rounded or truncated metaxyphus. Collection Records: Adams Co. 14 3 , 35 2 ; Barron Co. 2 2 ; Burnett Co. 1 2 ; Columbia Co. 1 3,2 2 ; Dane Co. 5 3,2 2 ; Door Co. 2 3,1 2 ; Douglas Co. 1 2 ; Eau Claire Co. 23,1 2 ; Florence Co. 1 3 ; Green Co. 1 2 ; Juneau Co. 3 3,3 2 ; Lincoln Co. 1 2 ; Marathon Co. 3 3,2 2 ; Marquette Co. 2 3,1 2 ; Outagamie Co. 1 2 ; Polk Co. 1 2 ; Price Co. 2 2 ; Racine Co. 2 3 ; St. Croix Co. 1 3 ; Sauk Co. 2 3„ 6 2 ; Taylor Co. 4 3,2 2 ; Trempealeau Co. 14 3,9 2 ; Vilas Co. 1 3 ; Winnebago Co. 5 3,6 2 . Totals: 11+2 individuals, 39 collections, 21+ counties. Previous Records: None. Reported from Illinois and Minnesota. Sigara decor atella (Hungerford) 1926 Distribution and Abundance: This is a common species that is distributed throughout the state (Fig. 5A). Identification: The reduced number of lower palmar hairs is dis¬ tinctive for this large species. 226 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Collection Records: Adams Co. 27 <2, 47 2 ; Ashland Co. 4 2 ; Barron Co. 1 <2 ; Burnett Co. 2 2 , 5 2 ; Chippewa Co. 1 <2 ; Colum¬ bia Co. 1 2,1 $ ; Dane Co. 1 2,2 $ ; Douglas Co. 4 2,2 $ ; Florence Co. 21 2, 21 $ ; Forest Co. 9 2,4 $ ; Green Co. 1 2 ; Juneau Co. 1 2,1 2 ; Kenosha Co. 1 2 ; Lincoln Co. 3 2 , 8 2 ; Marinette Co. 1 2 ; Marquette Co. 1 2 ; Oconto Co. 1 2 ; Oneida Co. 2 2 ; Outagamie Co. 2 2,2 2 ; Pierce Co. 1 2 ; Polk Co. 4 2 ; Price Co. 7 2, 8 2 ; Sauk Co. 2 2,14 2 ; Shawano Co. 1 2 ; Taylor Co. 1 2,4 2 ; Vilas Co. 1 2 ; Waupaca Co. 1 2 , 1 2 ; Winnebago Co. 18 2 ,11 2 . Totals: 251 individuals, 60 collections, 28 counties. Previous Records: Dane Co. 1 2, 4 2 ; St. Croix R. 8 2, 11 2. Also reported from Iowa, Michigan and Minnesota. Sigara defecta Hungerford 1948 Distribution and Abundance: Although not collected in the ex¬ treme southwestern counties, it probably occurs throughout the state (Fig. 4B) . It is fairly common in many parts of the state. Identification: The male pala is distinctive, but the female can be easily confused with S. bicoloripennis or S. mullettensis. The pala is longer than that of S. bicoloripennis, and much longer than that of S. mullettensis. The anal lobes are distinctly notched mesally, while those of S. bicoloripennis are unnotched and those of S. mullettensis are only very slightly notched. Collection Records: Adams Co. 12 3, 25 2 ; Ashland Co. 1 2 ; Burnett Co. 3 2,3 2 ; Columbia Co. 1 2 ; Douglas Co. 42 2 , 40 2 ; Dunn Co. 1 2 ; Florence Co. 7 2,4 2 ; Forest Co. 1 2 „ 1 2 ; Juneau Co. 1 2,1 2 ; Kenosha Co. 1 2 ; Lincoln Co. 1 <2,1 2 ; Milwaukee Co. 1 2 ; Oconto Co. 4 2,4 2 ; Polk Co. 5 2 ; Portage Co. 1 2 ; Price Co. 3 2,4 2 ; Racine Co. 3 2, 1 2 ; Sauk Co. 1 2 ; Taylor Co. 1 2,1 2 ; Washburn Co. 1 2 ; Winnebago Co. 2 2 . Totals: 178 individuals, 30 collections, 21 counties. Previous Records: Dane Co. 1 <2,2 2 ; Sauk Co. 1 2. Also re¬ ported from Illinois, Michigan and Minnesota. Sigara dolabra Hungerford and Sailer 1942 Distribution and Abundance: S. dolabra has been collected only in the northern half of the state and it is uncommon (Fig. 5D). Identification: The very long, pointed metaxyphus and the meso- epimeron equal in width to the prothoracic lobe separate this species. Collection Records: Burnett Co. 1 2 ; Florence Co. 3 2, 12 2 ; Forest Co. 1 2 ; Lincoln Co. 1 2 ; Marathon Co. 1 2 ; Marinette Co. 1 <2 ; Onedia Co. 1 2,1 2 ; Vilas Co. 2 2 . Totals: 29 individuals, 12 collections, 8 counties. Previous Records: None. Reported from Michigan and Minnesota. 1970] Hilsenhoff — Corixidae of Wisconsin 227 Sigara douglasensis (Hungerford) 1926 Distribution and Abundance : The distribution of this fairly com¬ mon species is confined to the northern third of the state (Fig. 5B) . Identification: The wavy longitudinal lines on the clavis and corium separate this species, and the location of the hypo-ocular suture is distinctive. Collection Records: Douglas Co. 2 2,1 2 ; Florence Co. 2 3,4 $ ; Forest Co. 3 3 ; Lincoln Co. 1 2 , 3 $ ; Oneida Co. 1 2 ; Polk Co. 3 3,2 2 ; Rusk Co. 1 3 ; Sawyer Co. 13,1 2 ; Vilas Co. 5 2,6 9 ; Washburn Co. 1 9. Totals: 37 individuals, 17 collections, 10 counties. Previous Records: None. Reported from Michigan and Minnesota. Sigara grossolineata Hungerford 1948 Distribution and Abundance: An abundant species, it occurs throughout the state (Fig. 4A). Identification: Closely related to S. modesta, which has not been collected in Wisconsin but might occur in the south. In S. modesta the scent gland osteole is only 1/2 to 3/5 from the tip of the mesoepimeron to the lateral bend, and not close to the lateral bend as in S. grossolineata. Also in S. modesta the dark pattern of the clavis is effaced along the margin bordering the pronotum, while in S. grossolineata the lines may become narrow, but they remain distinct. Several specimens of S. modesta from the Snow Collection were examined. Collection Records: Adams Co. 2 2„ 7 2 ; Ashland Co. 3 3,6 2 ; Brown Co. 12 3, 28 2 ; Calumet Co. 7 3 , 16 2 ; Chippewa Co. 5 2,9 2 ; Clark Co. 13,1 2 ; Columbia Co. 3 3,8 2 ; Crawford Co. 28 3, 60 2 ; Dane Co. 26 2 , 39 2 ; Dodge Co. 11 3 , 15 2 ; Door Co. 10 3,9 2 ; Douglas Co. 1 2 ; Dunn Co. 2 3,8 2 ; Eau Claire Co. 1 3,2 2 ; Florence Co. 11 3, 21 2 ; Fond du Lac Co. 5 2,5 2 ; Forest Co. 13 2, 19 2 ; Grant Co. 14 2 , 37 2 ; Green Co. 13 3„ 17 2 ; Green Lake Co. 5 2,8 2 ; Iowa Co. 77 2 , 81 2 ; Jefferson Co. 11 2,6 2 ; Juneau Co. 6 2, 13 2 ; Kenosha Co. 4 2 , 13 2 ; Kewaunee Co. 8 2,7 2 ; La Crosse Co. 3 2 ; Lafayette Co. 4 2, 14 2 ; Langlade Co. 2 2,5 2 ; Lincoln Co. 10 2 , 14 2 ; Mani¬ towoc Co. 9 2,21 2 ; Marathon Co. 19 2 , 27 2 ; Marinette Co. 12,3 2 ; Marquette Co. 2 2,3 2 ; Menominee Co. 1 2 ; Milwaukee Co. 2 2,4 2 ; Oconto Co. 15 2, 20 2 ; Oneida Co. 2 2,5 2 ; Outa¬ gamie Co. 8 2,12 2 ; Ozaukee Co. 3 2 ; Polk Co. 2 2 ; Portage Co. 3 2 „ 6 2 ; Price Co. 8 2, 10 2 ; Racine Co. 3 2 , 10 2 ; Richland Co. 4 2 ; Rock Co. 2 2,2 2 ; Rusk Co. 2 2,1 2 ; St. Croix Co. 3 2 ; Sauk Co. 18 2, 21 2 ; Sawyer Co. 2 2, 3 2 ; Shawano Co. 228 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 1 3,6 2 ; Sheboygan Co. 3 3,8 2 ; Taylor Co. 29 3 , 59 2 ; Trem¬ pealeau Co. 3 3,45; Vernon Co. 1 3,6 2 ; Vilas Co. 19 3 , 33 2 ; Walworth Co. 19 3 , 29 2 ; Washburn Co. 1 3 ; Washington Co. 4 3,6 2 ; Waukesha Co. 3 3,3 2 ; Waupaca Co. 5 3,8 2 ; Winne¬ bago Co. 11 3, 17 2 ; Wood Co. 6 3,2 2 . Totals: 1309 individuals, 212 collections, 62 counties. Previous Records: Dane Co. 15 3 , 22 2 ; Douglas Co. 3 3,4 2 . Also reported from Illinois, Iowa, Michigan and Minnesota. Sigara johnstoni Hungerford 1948 Distribution and Abundance: This species is fairly common in the northwest half of the state (Fig. 4D). The Illinois record suggests that it might occur in the southern part of the state as well. Identification: The females may be confused with those of S. variabilis or S. knighti. The anal lobes are not notched mesally as they are in S. knighti; those of S. variabilis are slightly notched. Most distinctive is the metaxyphus, which is very slightly notched at the tip in S. johnstoni. Collection Records: Ashland Co. 1 2 ; Burnett Co. 1 2 ; Clark Co. 1 3,2 2 ; Douglas Co. 33; Dunn Co. 1 3 ; Eau Claire Co. 1 3 ; Florence Co. 1 3 ; Forest Co. 1 3,3 2 ; Jackson Co. 1 2 ; Juneau Co. 2 3, 2 2 ; Langlade Co. 6 3,6 2; Lincoln Co. 4 3, 5 2 ; Marathon Co. 11 3,6 2 ; Oneida Co. 1 3 ; Portage Co. 5 2 ; Price Co. 2 2 ; Rusk Co. 2 3 ; Sawyer Co. 4 3,3 2 ; Taylor Co. 3 3,7 2 ; Vilas Co. 1 3 ; Wood Co. 2 3,, 2 2 . Totals: 90 individuals, 3U collections, 21 counties. Previous Records: None. Reported from Illinois and Minnesota. Sigara knighti Hungerford 1948 Distribution and Abundance: This uncommon species occurs only in the northern third of the state (Fig. 4D). Identification: The females may be separated from S. johnstoni and S. variabilis by their mesally notched anal lobes. Collection Records: Ashland Co. 1 2 ; Bayfield Co. 2 3,1 2 ; Douglas Co. 1 3 ; Florence Co. 5 3,6 2 ; Forest Co. 8 3 , 12 2 ; Langlade Co. 4 2 ; Price Co. 4 2 ; Vilas Co. 2 3,3 2 . Totals: Jj9 individuals, 12 collections, 8 counties. Previous Records: None. Reported from Michigan and Minnesota. Sigara lineata (Forster) 1771 Distribution and Abundance: Where it occurs, this species may be found in tremendous numbers. It is common throughout much of the state, and has been collected mostly in areas where there is sandy soil (Fig. 5B). 1970] Hilsenhoff — Corixidae of Wisconsin 229 Identification: The very small size and striped hemelytra make this species easy to recognize. Collection Records: Adams Co. 76 2 , 101 2 ; Ashland Co. 93 2 , 164 $ ; Columbia Co. 335 2 , 329 5 ; Crawford Co. 7 2 , 14 $ ; Dane Co. 7 2 , 13 $ ; Dunn Co. 7 2 , 3 $ ; Eau Claire Co. 1 $ ; Florence Co. 491 2„ 515 $ ; Juneau Co. 24 $ , 19 5 ; Lincoln Co. 2 2,1 $ ; Marathon Co. 91 3, 157 2 ; Marquette Co. 1 2 ; Oneida Co. 1 2, 2 2 ; Outagamie Co. 3 2 ; Portage Co. 8 2, 17 2 ; Price Co. 18 $ , 20 2 ; Rusk Co. 4 2,4 2 ; Sauk Co. 62 2,59 2 ; Taylor Co. 5 2 , 4 2 ; Trempealeau Co. 41 $ , 122 2 ; Vilas Co. 84 2 , 76 2 . Totals : 2981 individuals, 78 collections, 21 counties. Previous Records: None. Reported from Illinois and Minnesota. Sigara mackinacensis (Hungerford) 1928 Distribution and Abundance: The distribution of this species is restricted to the northern half of the state where it is fairly com¬ mon (Fig. 5C). Identification: The distinctly marked membrane separates it from S. compressoidea, the only species with which it might be confused. Collection Records: Barron Co. 1 2 ; Chippewa Co. 1 2 ; Clark Co. 2 2,1 2 ; Douglas Co. 1 2 ; Eau Claire Co. 2 2 ; Florence Co. 5 2,8 2 ; Forest Co. 4 $ , 5 2 ; Lincoln Co. 4 2,2 2 ; Oconto Co. 1 2 ; Oneida Co. 2 2„ 3 2 ; Polk Co. 4 2 ; Sawyer Co. 3 2 ; Vilas Co. 4 2, 7 2 ; Washburn Co. 25 2 , 36 2 ; Wood Co. 4 s, 4 2. Totals: 129 individuals, 27 collections, 15 counties. Previous Records: None. Reported from Michigan and Minnesota. Sigara macropala (Hungerford) 1926 Distribution and Abundance: This species is uncommon in Wis¬ consin, and has been collected only in the extreme north (Fig. 5D) . Identification : The large dorsal extension of the male pala is distinctive. The sharply pointed metaxyphus separates this species from S. bicoloripennis and S. defecta, and it can be easily separated from the larger S . alternata by the pattern on the clavis and the fewer number of black bars on the prothorax. Collection Records: Burnett Co. 1 2 ; Douglas Co. 1 2 ; Florence Co. 21 2 , 20 2 ; Washburn Co. 11 2 , 11 2 . Totals: 65 individuals, 5 collections, J counties. Previous Records: None. Reported from Michigan and Minnesota. Sigara mathesoni Hungerford 1948 Distribution and Abundance: A very common species throughout the entire state, it seems to have an affinity for spring ponds (Fig. 4C) . 230 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Identification : The median brown stripe on the head and the deep incision at the lateral bend of the mesoepimeron are distinc¬ tive. Collection Records: Adams Co. 4 3,9 9 ; Ashland Co. 1 3, 2 $ ; Barron Co. 1 $ ; Calumet Co. 1 3 ; Chippewa Co. 1 3 , 2 $ ; Colum¬ bia Co. 12 28 $ ; Crawford Co. 12 3„ 16 9 ; Dane Co. 36 3, 56 $ ; Dodge Co. 13 3, 16 $ ; Door Co. 16 3 , 51 9 ; Florence Co. 30 3, 86 9 ; Forest Co. 1 3,3 9 ; Grant Co. 1 3 ; Green Co. 1 3 ; Iowa Co. 39 3 , 29 9 ; Jefferson Co. 5 3, 13 9 ; Juneau Co. 2 3, 4 9 ; Langlade Co. 2 9 ; Lincoln Co. 16 3 , 22 9 ; Marinette Co. 1 9 ; Marquette Co. 4 9 ; Monroe Co. 29 3 , 33 9 ; Price Co. 83 3 , 124 9 ; St. Croix Co. 3 9 ; Sauk Co. 1 3 ; Vernon Co. 1 3 ; Vilas Co. 1 3, 3 9 ; Walworth Co. 2 3, 5 9 ; Washburn Co. 1 3,2 9 ; Washington Co. 1 9 ; Waukesha Co. 67 3 , 106 9 ; Waushara Co. 5 3,4 9 . Totals : 1007 individuals, 69 collections, 33 counties. Previous Records: Dane Co. 4 3,5 9 ; Douglas Co. 14 3 , 13 9 ; Sauk Co. 1 9 . Also reported from Michigan and Minnesota. Sigara mullettensis (Hungerford) 1928 Distribution and Abundance: This species is fairly common throughout the northern two-thirds of the state, with only two scattered records from the southern counties (Fig. 4B). Identification: The females could be confused with those of S. defecta, which also tend to have the corial pattern in a longitudinal series. In S. mullettensis the female pala is much shorter, and the anal lobes are only slightly notched. Collection Records: Ashland Co. 7 3, 6 9 ; Burnett Co. 4 3, 4 9 ; Chippewa Co. 1 3 ; Clark Co. 1 3 ; Door Co. 1 9 ; Douglas Co. 1 9 ; Eau Claire Co. 1 3 ; Forest Co. 1 9 ; Juneau Co. 1 3 ; Langlade Co. 2 3,5 9 ; Lincoln Co. 2 3,5 9 ; Marathon Co. 1 3 , 3 9 ; Oconto Co. 3 3,1 9 ; Outagamie Co. 2 3,3 9 ; Ozaukee Co. 1 3 ; Polk Co. 1 9 ; Portage Co. 13,1 9 ; Price Co. 1 9 ; Sawyer Co. 2 3 ; Taylor Co. 16 3, 14 9 ; Vilas Co. 3 9 ; Winnebago Co. 11 3, 27 9 . Totals: 133 individuals,, 35 collections, 23 counties. Previous Records: Dane Co. 1 9. Also reported from Michigan and Minnesota. Sigara penniensis (Hungerford) 1928 Distribution and Abundance: This species is fairly common in the northern two-thirds of the state (Fig. 5A). Identification: This large, dark colored Sigara can be recognized by the narrow interocular space and the pale markings of the corium being arranged in a definite longitudinal series. 1970] Hilsenhoff — Corixidae of Wisconsin 231 Collection Records : Adams Co. 1 2 ; Burnett Co. 2 2,3 2 ; Clark Co. 1 2 ; Door Co. 1 $ ; Douglas Co. 4 2, 10 2 ; Forest Co. 1 2 ; Florence Co. 3 2,7 2 ; Jackson Co. 2 2 ; Lincoln Co. 7 2 , 9 2 ; Oconto Co. 3 2 ; Oneida Co. 1 2,3 2 ; Polk Co. 1 2 ; Vilas Co. 6 2, 2 $ ; Washington Co. 3 2,1 2 . Totals: 71 individuals, 26 collections, 74 counties. Previous Records: None. Reported from Michigan and Minnesota. Sigara signata (Fieber) 1851 Distribution and Abundance: This species is common in the northern two-thirds of the state but has not been collected in the southern counties or the counties bordering Lake Michigan or the Mississippi River (Fig. 5D). Identification: This dark little species looks like a miniature H. minorella. It is much darker than S. solensis, and can be easily separated by the characters given in the key. Collection Records: Adams Co. 1 2,7 2 ; Ashland Co. 3 2,3 2 ; Barron Co. 1 2 ; Burnett Co. 1 2,6 2 ; Chippewa Co. 1 2 ; Clark Co. 6 2 ; Douglas Co. 3 $ ; Eau Claire Co. 3 2,9 2 ; Florence Co. 34 2„ 67 2 ; Forest Co. 3 2,8 2 ; Iron Co. 2 2 ; Juneau Co. 5 2, 5 2 ; Langlade Co. 2 2,1 2 ; Lincoln Co. 26 2 , 29 2 ; Marathon Co. 9 2, 13 2 ; Marinette Co. 2 2 ; Marquette Co. 1 2 ; Monroe Co. 1 2,1 2 ; Oconto Co. 5 2,7 2 ; Oneida Co. 13 2, 17 2 ; Polk Co. 1 2, 5 2 ; Portage Co. 2 2 ; Price Co. 3 2,4 2 ; Rusk Co. 5 2,7 2 ; Sawyer Co. 4 2,2 2 ; Shawano Co. 1 2 ; Taylor Co. 4 2 , 4 2 ; Vilas Co. 19 2, 45 2 ; Washburn Co. 2 2 ; Waupaca Co. 1 2 ; Waushara Co. 1 2,1 2 ; Wood Co. 1 2,4 2 . Totals: Ifll individ¬ uals, 96 collections, 31 counties. Previous Records: Douglas Co. 1 2 . Also reported from Illinois, Michigan and Minnesota. Sigara solensis (Hungerford) 1926 Distribution and Abundance: Except for the extreme southern and eastern counties,, this species has been collected throughout the state (Fig. 4C) . It is fairly common in many areas. Identification : Readily distinguished by the characters in the key. Collection Records: Adams Co. 5 2, 6 2 ; Ashland Co. 1 2 ; Dane Co. 2 2,2 2 ; Douglas Co. 1 2 ; Eau Claire Co. 22,1 2 ; Florence Co. 11 2 , 23 2 ; Forest Co. 1 2 ; Green Lake Co. 2 2 ; Lincoln Co. 5 2,3 2 ; Marathon Co. 2 $ ; Marquette Co. 12 2 , 10 2 ; Oconto Co. 1 2 ; Oneida Co. 1 $ ; Polk Co. 1 2 ; Rusk Co. 1 2 ; Sauk Co. 1 2 ; Taylor Co. 3 2,2 2 ; Trempealeau Co. 1 2 ; 232 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Vilas Co. 1 2 ; Winnebago Co. 34 8 , 37 2 . Totals: 172 individuals, UO collections, 20 counties. Previous Records : Dane Co. 24 8 , 23 2 . Also reported from Michigan and Minnesota. Sigara transfigurata (Walley) 1930 Distribution and Abundance: This rare species was collected at only two sites, one in Vilas Co. in the extreme north and one in Marathon Co. in the central part of the state (Fig. 4D). Identification: The bold cross-bars on the corium separate this species from others with a pale longitudinal line on the pronotum. Collection Records: Marathon Co. 1 8 ; Vilas Co. 2 8. Totals: 3 individuals, 2 collections, 2 counties. Previous Records: None. Reported from Michigan. Sigara trilineata (Provancher) 1872 Distribution and Abundance: Collected throughout the state,, except in the extreme eastern and western counties (Fig. 5B) . Like S. lineata, with which it often occurs, it seems most prevalent in sandy areas. Identification: The bold longitudinal stripes on the hemelytra and the wide mesoepimeron separate it from all other species. Collection Records: Adams Co. 129 8 , 199 2 ; Barron Co. 1 8, 1 $ ; Chippewa Co. 1 8, 2 2 ; Columbia Co. 3 8, 3 2 ; Dane Co. 7 8,1 2 ; Douglas Co. 2 8 ; Eau Claire Co. 1 $ ,, 1 2 ; Florence Co. 101 8, 159 2 ; Forest Co. 2 8 ; Iowa Co. 50 8 , 41 2 ; Juneau Co. 56 8 , 82 2 ; Lincoln Co. 6 8 , 9 2; Marathon Co. 44 .8 , 50 2 ; Marinette Co. 6 2 ; Marquette Co. 2 8 ; Monroe Co. 2 2 ; Oconto Co. 6 8 , 8 2 ; Portage Co. 1 8,2 2 ; Price Co. 1 8 ; Rusk Co. 13 8 , 6 2 ; Sauk Co. 22 8 , 29 2 ; Taylor Co. 2 8,2 2 ; Vilas Co. 78 8, 162 2 ; Washburn Co. 1 8,4 2 ; Waupaca Co. 2 8, 5 2 ; Wood Co. 1 8 . Totals: 1312 individuals, 83 collections, 26 counties. Previous Records: Burnett Co. 1 8, 4 2. Also reported from Michigan and Minnesota. Sigara variabilis (Hungerford) 1926 Distribution and Abundance: Although it was collected only in the northwestern half of the state (Fig. 5C), collections in Illinois and Michigan suggest that this uncommon species should be found in the rest of the state as well. Identification: The pala of the male is distinctive, but the females resemble S. johnstoni. They can be separated from that species by the lack of an incision in the metaxyphus, the wider and more widely 1970] Hilsenhoff—Corixidae of Wisconsin 233 spaced pale figures on the corium, and the slight mesal indentation of the anal lobes. Collection Records: Crawford Co. 2 $ , 1 $ ; Douglas Co. 3 $ ; Dunn Co. 1 $ , 1 $ ; Juneau Co. 1 $ ; Marathon Co. 1 $ ; Oneida Co. 3^,5 $ ; Portage Co. 1 $ ; Trempealeau Co. 1 $ , 3 $ . Totals: 23 individuals, 10 collections, 8 counties. Previous Records: None. Reported from Illinois and Michigan. Trichocorixa Kirkaldy Four species of Trichocorixa have been collected in Wisconsin, with all species being most common in the southern third of the state (Fig. 2D). A fifth species, T. macro ceps (Kirkaldy) 1908, has been found in Illinois and Michigan and may occur in Wisconsin. Key to Wisconsin species Males 1. Strigil small and round _ 2 Strigil very elongate _ 3 2. Nodal furrow appearing absent or at apex of embolar groove; length of pronotal disc about one-fourth its width _ T. macroceps Nodal furrow dividing embolar groove; length of pronotal disc one-third or more its width _ T. naias 3. Strigil extremely narrow, little more than a heavy dark line and usually curved abruptly upward at the mesal end _ T. calva Strigil about 7 times as long as wide _ 4 4. Strigil noticeably widened in region of the bend _ T. borealis Strigil about the same width along its entire length, any slight widening occurring near the lateral end _ T. kanza Females 1. Nodal furrow appearing absent or at apex of embolar groove __2 Nodal furrow dividing embolium _ 3 2. Length of pronotal disc about one-fourth its width -T. macroceps Length of pronotal Disc one-third or more its width _ T. naias 3. Length of apical area of embolar groove exceeding length of middle tarsus _ T. borealis Length of apical area less than that of middle tarsus _ 4 4. At least 2 patches of bristle-like setae on right side of seventh abdominal sternite _ T. kanza Only fine pubescence on right side of seventh abdominal sternite _ _ T. calva 234 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Trichocorixa borealis Sailer 1948 Distribution and Abundance : This species is common throughout the southern third of the state and has been collected as far north as Barron and Langlade Counties (Fig. 2D). Identification: The strigil of the male, which is widened at the bend,, and the long postnodal pruinose area in the female separate this species. In most females there is a distinct outward projection at the anterior end of the polished prenodal area. Collection Records: Adams Co. 13 3 , 15 2 ; Barron Co. 2 2 ; Brown Co. 12 3 , 31 2 ; Columbia Co. 16 3 , 49 2 ; Crawford Co. 1 3,2 2 ; Dane Co. 64 3 , 100 2 ; Dodge Co. 12 3 , 25 2 ; Dunn Co. 3 $,2 2 ; Green Co. 1 2 ; Green Lake Co. 84 3 , 141 2 ; Iowa Co. 5 3,6 $ ; Jefferson Co. 2 3,5 2 ; Juneau Co. 3 $ ; Kenosha Co. 18 3, 26 $ ; Kewaunee Co. 2 $ ; La Crosse Co. 1 2 ; Lincoln Co. 2 3,1 2 ; Manitowoc Co. 1 2 ; Marathon Co. 13,2 2 ; Marquette Co. 4 3,8 2 ; Outagamie Co. 63,7 2 ; Racine Co. 2 3 ; Rock Co. 2 2 ; Sauk Co. 24 3 , 25 2 ; Sheboygan Co. 1 3„ 1 2 ; Trempealeau Co. 1 2 ; Walworth Co. 65 3, 69 2 ; Washington Co. 1 3,2 2 ; Winnebago Co. 1 3. Totals: 867 individuals, 65 collections, 29 counties. Previous Records: None. Reported from Iowa and Minnesota. Trichocorixa calva (Say) 1832 Distribution and Abundance: This species is fairly common in the southern half of the state, and apparently does not occur in the north (Fig. 2D). Identification: The males can be identified by their extremely narrow strigil. The females can be separated from T. kanza by the lack of patches of setae on the right side of the seventh abdominal sternite. Collection Records: Adams Co. 1 2 ; Columbia Co. 1 2 ; Craw¬ ford Co. 30 3 , 35 2 ; Dane Co. 2 2 ; Dodge Co. 1 3,1 2 ; Fond du Lac Co. 1 3 ; Grant Co. 6 3 , 10 2 ; Green Co. 1 3,3 2 ; Green Lake Co. 1 2 ; Iowa Co. 1 2 ; Jefferson Co. 1 2 ; Juneau Co. 2 3 , 1 $ ; Kenosha Co. 80 3 , 90 $ ; Kewaunee Co. 1 $ ; Lafayette Co. 1 3,3 2 ; Manitowoc Co. 1 3 ; Marquette Co. 14 3, 15 2 ; Milwau¬ kee Co. 2 3,1 2 ; Monroe Co. 1 3 ; Outagamie Co. 1 2 ; Racine Co. 7 3, 11 2 ; Rock Co. 1 2 ; Sauk Co. 1 2 ; Trempealeau Co. 2 3, 1 2 ; Walworth Co. 1 3, 8 $ ; Washington Co. 3 3,2 2 ; Wau¬ kesha Co. 1 $ . Totals: 356 individuals, 37 collections, 26 counties. Previous Records: Dane Co. (no numbers). Also reported from Illinois, Iowa, Michigan and Minnesota. 1970] Hilsenhoff — Corixidae of Wisconsin 235 Trichocorixa kanza Sailer 1948 Distribution and Abundance: Found only in the extreme south¬ west corner of the state, this species is uncommon (Fig. 2D). Identification : The male strigil is much wider than that of T. calva, and slightly narrower than T. borealis. The female has three small and distinct patches of setae on the right side of the seventh abdominal sternite. Collection Records: Dane Co. 1 £, 2 9 ; Grant Co. 1 £ , 1 $ ; Lafayette Co. 8 3, 4 9. Totals: 17 individuals, 7 collections, 3 counties. Previous Records: Dane Co. (no numbers). Also reported from Illinois and Iowa. Trichocorixa naias (Kirkaldy) 1908 Distribution and Abundance: This species is fairly common in the southern fourth of the state, and has been collected as far north as Door and Lincoln Counties (Fig. 2D). Identification: The absence of a postnodal pruinose area in the female and the small rounded strigil of the male separate this species from other Wisconsin Trichocorixa, except T. macroceps, which has a very short pronotal disc. Collection Records: Columbia Co. 16 £,22 9 ; Dane Co. 39 £, 41 9 ; Dodge Co. 3 9 ; Door Co. 4 £ , 4 9 ; Iowa Co. 1 9 ; Jefferson Co. 1 9 ; Lincoln Co. 3 £ , 3 9 ; Manitowoc Co. 4 £ , 6 9 ; Marathon Co. 1 £ ; Outagamie Co. 2 £, 1 9 ; Ozaukee Co. 3 £, 1 9 ; Racine Co. 1 £ ; Walworth Co. 2 9 ; Waukesha Co. 1 9 ; Wood Co. 6 £ , 3 9 . Totals: 168 individuals, 30 collections, 15 counties. Previous Records: Dane Co. (no numbers). Also reported from Illinois, Iowa, Michigan and Minnesota. Literature Cited Brooks, A. R. and L. A. Kelton. 1967. Aquatic and semiaquatic Heteroptera of Alberta, Saskatchewan, and Manitoba (Hemiptera). Mem. Entomol. Soc. Canada No. 51. 92 pp. Hungerford, H. B. 1948. The Corixidae of the Western Hemisphere (Hem¬ iptera) Univ. Kansas Sci. Bull. 32: 1-288, 408-827. Sailer, R. I. 1948. The genus Trichocorixa (Corixidae, Hemiptera). Univ. Kansas Sci. Bull. 32: 289-407. TROPHIC NATURE OF SELECTED WISCONSIN LAKES Lloyd A. Lueschow, James M. Helm , Donald R. Winter and Gary W. Karl Appreciable differences in the biota of Wisconsin lakes are ap¬ parent to even the most insensitive eye. Some lakes, such as Geneva (Walworth County) and Crystal (Vilas County), are essentially clear; others, such as Winnebago (Winnebago County) and Dela- van (Walworth County), develop nuisance growths of algae, weeds, or both. Limnologists have long recognized these differences in biological composition and productivity to be related to the accumu¬ lation of nutrients and have referred to this aging process as “eutrophication.” Most lakes when formed are nutrient poor or oligotrophic. They subsequently proceed to a nutrient rich, eutro- phic condition. Although the distinction between oligotrophic and eutrophic lakes is, by definition, based on nutrient status, many parameters are often used to reflect the trophic status of a lake. The object of this study was to evaluate several water quality parameters from well-known lakes where the recreational potential is recognized by the public and where long records of observation and aquatic nuisance control are available. Each of these “trophic parameters” is evaluated in terms of how well it relates to both the trophic status and the observed recreational values of each lake. A better understanding of these relationships should lead to more meaningful interpretation of water quality data, more efficient sampling programs, and better diagnostic and interpretive tech¬ niques for evaluating lake problems. Lakes Included in the Study i The 12 lakes shown in Table 1 were selected because they are all well known and because they represent a broad range of con¬ ditions. Of the lakes selected, three (Pewaukee, Delavan, and Win¬ nebago) have nuisance algae “blooms” during most of the summer months which render the waters less desirable for recreational purposes. A fourth (Mendota) has occasional nuisance algae popu¬ lations. Rooted weed growths and Char a sp. growths also impair recreational potential, and Lakes Middle, Oconomowoc, Pine, and Pewaukee all have had extensive weed control activity. Lakes Men¬ dota and Winnebago have received less weed control effort, but weeds produce local nuisances on both during the summer months. 237 Table 1. General Lake Information 238 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 o TD 03 O c c 03 -cm X •OO^t £ ^UO'oo woo — —' c § g C £ o bfl ,q °0 Op C .a >> rx > W xO sO cd C — ^ • ITS CJ O cm 03 oo — — X C ju o 2 73 ^ 0) I O — t^— c • — 00 ■? OOM- C 8^ 8 E 0) = n Q — S > V C3 O cm O O CO C > -xD X 'OO « O ?NVn 0) 00 CM — _C C 3 $r£! o _2 oo o c _ f— 1 > — £ o o O c c c Q O ■? oj 3 w-x 03 C >> S- -oo S 8 s ;> -ics x • oo 03 •? o> 3> cm -t- o) oo cm — J3 C£ OnxO CxO^cM 03 03 03 a 3 03 xQ >CK I> — xQ 03 •S ° nfo07 C c • ^ ^ c £ ^ — c 03 03 o o c c a 0 = £ £ Jpa6 £?o G'O'y M S 03 x ^ Sailed 03 03 1970] L. Lueschow, J. Helm, D. Winter and G. Karl — Lakes 239 Lakes Oconomowoc and Pine have had extensive Chara sp. control programs, and Big Green Lake has had an occasional Chara sp. nuisance but no control activity. Trophic Parameters and Methods Parameters selected for this study included the nutrients nitro¬ gen and phosphorus, temperature, dissolved oxygen, water trans¬ parency and plankton analysis. Nitrogen was analyzed as organic nitrogen, Ammonia-N, Nitrate-N, Nitrite-N. The NH3-N, N03-N, and NO2-N concentrations were combined as inorganic nitrogen and considered available for utilization. Phosphorus was analyzed as soluble phosphorus (orthophosphates) and total phosphorus. All nutrient analyses were performed by the Wisconsin Laboratory of Hygiene in accordance with Standard Methods, 12th edition, 1965, and results are expressed in mg/1 of nitrogen and phosphorus. Temperature profiles were determined with an electronic re¬ sistance thermometer. Dissolved oxygen concentrations were deter¬ mined by the modified Winkler technique, Standard Methods, 1965. Water transparency was measured with a Secchi disk and also with a Whitney light meter. Light meter values are expressed in per cent of surface incident light at each depth. Plankton evaluations were based on net collections using a Clarke-Bumpus plankton sampler. The Clarke-Bumpus apparatus measures a known volume of water passed through the net, and the volume of plankton captured can then be used to evaluate the con¬ centration of plankton in the water. The net used was a No. 20 standard mesh (.007 inch bar). Most plankton samples required a 1-5 minute tow depending on the quantity of plankton in the water. Samples were transferred from the plankton cup to 180 ml. storage jars, and 10 ml. of commercial formaldehyde were added for preser¬ vation. The samples were then returned to the laboratory for evaluation at a later date. The volume of plankton constituents was estimated by visual observation under a dissecting or compound microscope. The sam¬ ples were then dried and ashed to determine total and volatile solids. Plankton quantity was then reported as total solids and expressed as micrograms of solids per liter of water filtered. Results and Discussion Dissolved Oxygen and Temperature The dissolved oxygen concentration in the hypolimnion may be used as a trophic parameter. Organic material produced in the trophogenic zone eventually settles through the thermocline into the tropholytic zone where it consumes oxygen during chemical and Table 2. Temperature and Dissolved Oxygen by Month — 1966 240 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Table 2. Temperature and Dissolved Oxygen by Month — 1966 — Continued 1970] L. Lueschow, J. Helm, D. Winter and G. Karl — Lakes < H O Q DO 8.5 0.8 0.0 8.2 1 .9 0.0 o o o 00 o o 8.5 8.8 8.8 9.6 9.0 10.2 Z w 2 77 59 57 O fe I tN* *T\ j X Tt- I X ^ 1 o o o 44 44 44 ' ; u Q DO 8.1 11.0 0.0 7.6 1 .6 0.0 8.8 1 . 1 0.0 8.0 7.7 5.2 2 H O' O' CO t\ its irv 70 58 47 X ^ — 1 X X X-N ^ ^ ^ 41 41 41 • H DO 8.1 7.0 6.2 8.7 6.4 2.1 8.65 5.4 3.0 O' — 1 O' 00 X — £01 S- 01 9 01 a h 72 50 00 O 1 X W-N 1 64 52 52 52 52 40 40 40 z UJ W DO 8.4 ‘1° ha 1 8.7 9.9 9.6 9.1 oo oo I g H ^ 1 r\ itn | CO o z t\ hs I\ O O vC SS | 42 42 < w O Q ITV 0O CO sD ^ 7.6 5.5 3.8 0^0 O ^ N 8.3 8.0 1.0 urs ^ O' O' O' z © 0 h IX O' I tX Vv I O O' I tx i/-v | O' 1 1 X 1 1 SS | 46 46 46 < H -1— O Q ■ O GO O 00 O 00 8.2 6.8 5.0 x 32 1C OO >A 8.8 8.6 8.0 o o o >- a ; u H « 00 t\ ^ O' — o X V-v X (X X '> p X y x£ f-S ° o ° S ) • U o 0 o c ) U U X! =3 * 2 - ^ 5 < it u u u X X o X % c .w x 2 3 ^ ^ D. x 2; — ■§ 3 * vO 1 vD Delavan DO rj< t ■£) ia 0 £ 0 2 0 z 0 o O DO 12.5 13.1 12.8 11.2 11.2 11.3 11.2 12.2 11.9 11.6 11.0 10.7 10.2 8.5 8.0 6.8 h a N © © tT pa pa pa pa | pa pa | pa ^ ^ ts >a ia J anuary Surface . Middle of Hypolimnion . 1 M above bottom . February Surface . Middle of Hypolimnion . 1 M above bottom . March Surface . Middle of Hypolimnion . 1 M above bottom . April Surface . Middle of Hypolimnion . 1 M above bottom . May Surface . Middle of Hypolimnion . 1 M above bottom . June Surface . Middle of Hypolimnion . 1 M above bottom . Table 2. Temperature and Dissolved Oxygen by Month — 1966 — Continued 1970] L. Lueschow, J. Helm, D. Winter and G. Karl — Lakes 243 0 < CQ DO 8.0 7.9 imple 8.1 8.3 12.8 Z z £ H CO g 1 g z S 1 1 £ 1 1 S ll / z < > DO 7.6 0.0 0.0 7.4 0.0 0.0 6.7 0.0 0.0 7.9 8.2 8.0 11.0 11.0 11.0 E) Id G H vO pa O ^ 00 N 00 nD sO nD \Q \Q N Van ian \Q \Q i an ian van van Pewaukee DO 8.3 0.0 0.0 7.2 . 0.0 5.9 0.0 10.1 10.0 9.7 11.2 11.2 11.2 H tv ©I'D ia >a N O' O' O' — h — — ■ t^vOvD n. 1 >/■' sOnO1^ Pine DO 8.9 2.6 0.1 8.3 2.7 1 .6 9.0 0.35 0.0 9.1 8.4 0.0 9.6 9.6 9.6 tx ^ o o | o n | 0s oo oo ^ K lA 1a N 1a | \0 Tf 1 ^ ^ ^ ^ ^ OCONOMOWOC DO 8.6 2.2 0.4 7.7 0.7 0.3 8.7 1 .0 0.0 10.2 9.4 9.2 8.2 8.4 8.3 H — < t\ t\ O'CT'C^ ^ O 00 Van l/N sQ Its Van \Q Van Ian lA\ Van Van xH Round DO 8.7 6.9 1 .0 8.2 6.0 2.5 9.2 0.4 2.35 9.0 5.1 0.15 9.3 9.3 9.2 H lA (S N Tf N \D 00 GO \D rA PA CA PA t\ 1a Tr ^ sQ ^ ^ VAN IAN ^ ^ ^ July Surface . Middle of Hypolimnion . 1 M above bottom . August Surface . Middle of Hypolimnion . 1 M above bottom . September Surface . Middle of Hypolimnion . 1 M above bottom . October Surface . Middle of Hypolimnion . 1 M above bottom . November Surface . Middle of Hypolimnion . 1 M above bottom . *Temperature in degrees Fahrenheit, f Dissolved oxygen in mg/1. 244 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 biological stabilization. One may therefore assume that eutrophic lakes with a high productivity or standing crop would produce the most significant oxygen sag in the hypolimnion. These stabilization effects would of course be tempered by the volume of water in the hypolimnion, but nevertheless the quantity of dissolved oxygen in the hypolimnion is indicative of the trophic nature of a lake. Fur¬ thermore, a low dissolved oxygen concentration in the hypolimnion would affect the salmonoid and whitefish populations that might be present in the lakes. It should be noted that even the most oligotrophic lake sampled (Crystal) revealed some reduction in the dissolved oxygen con¬ centration of the hypolimnion. During August, the 5 mg/1 ob¬ served at 17 meters was only 45 per cent of saturation. Trout Lake and Lake Geneva also revealed substantial dissolved oxygen re¬ ductions in the hypolimnion although zero values were not observed. All of the more productive lakes revealed zero dissolved oxygen values during most of the summer months. These dissolved oxygen observations suggest that Lakes Delavan, Pewaukee, Pine, Ocono- mowoc, Middle, and Mendota produce more organic matter in the trophogenic zone than can be aerobically assimilated by the hypolimnion. Lakes Geneva, Trout, Crystal, and perhaps Pine have not reached that level of production and consequently retain sufficient dissolved oxygen in the hypolimnion to sustain fish and fish food organism populations intolerant of low dissolved oxygen values. Lake Winnebago is too shallow to stratify, so although it is a productive lake, a deoxygenated hypolimnion does not develop. Observations by Birge on Mendota in 1906 suggest that oxygen depletion in the hypolimnion occurred at that time as well as now. Transparency and Light Penetration The transparency of a lake as measured by the Secchi disk may also be used as a measure of the trophic condition. The basic assumption is that the magnitude of organic production affects the color and turbidity of the water. Beeton, 1965, used the Secchi disk readings in the interpretation of the trophic nature of the Great Lakes and reported Lake Michigan as oligotrophic with a reading of 19*4 feet. Values of fifteen to eighteen feet reported for Lakes Erie and Ontario were considered by Beeton to be com¬ paratively eutrophic. Only Crystal Lake had a mean Secchi disk reading that would be considered oligotrophic when compared with the Great Lakes. However, in all lakes where the mean Secchi disk values were greater than twelve feet, there has been essentially no deteriora¬ tion of recreational potential due to plankton growths. Lakes Table 3. Transparency of 12 Wisconsin Lakes by Month — 1966 (Meters) 1970] L. Lueschow, J. Helm , D. Winter and G. Karl — Lakes 245 C ^ - o O — TJ • • C ro ^ 42£ I X O' sD X I O' "3- — oo — t". oo I X co 4 S - 4 4 oo 5 4°°4 X X 4 £ _ 4^2 4^2 4~ C-l — O X I oo -4* o — ■ — 7^0-’ x 44 4-^7 .i 7 — 5 "§ # 2 4 >> O kO >- (u U to | ~ Q X co k_l bo ^ J r o Q co ^ 54 54 54 2 0 Green Seech i 5% Li Trout Secchi 5% Li Middl x 2 o w • co ^ •§ x C o w o 2 c£ 246 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 1970] L. Lueschow , J. Helm, D. Winter and G. Karl — Lakes 247 Mendota, Pine, Pewaukee, and Delavan have all had an extensive history of planktonic nuisance and all have mean seasonal Secchi disk values of ten feet or less. Those lakes that have a history of plankton nuisance conditions also reveal a 5 per cent incident light sesonal mean of twenty-five feet or less. In this respect there is good agreement between Secchi disk and per cent of incident light. However, the absolute correlation between Secchi disk and per cent of incident light on a particular lake is not readily apparent. Plankton Population The plankton populations on the twelve lakes studied are repre¬ sented in Figure I for the months when no ice cover was present. The evaluations were based on Clarke-Bumpus net tows (No. 20 mesh net) and the results are expressed in jxgJ 1 total solids. In the twelve lakes under consideration, Crystal, Trout, Geneva, Big Green and Round Lake revealed less than 200 jig/ 1 of solids consistently, and nuisance-producing blue-green varieties were de¬ tectable in trace quantities only. The plankton populations in these lakes were most typically composed of zooplankton and diatoms. Middle and Oconomowoc Lakes did not reveal plankton concen¬ trations that would be considered a nuisance, although the solids analyses revealed between 700 and 1,500 /xg/1 on five occasions. The plankton were principally diatoms or zooplankton, which do not have the nuisance-producing capability characteristic of the blue-green algae. Past aquatic nuisance control records of the De¬ partment of Natural Resources suggest that Delavan Lake has nuisance algae conditions during the entire summer. The plankton catches from June through October revealed 2,500 jig/l on four of five occasions. The plankton populations were dominated by blue-green varieties with Anacystis sp., Anabaena sp., and Oscilla- toria sp. predominating. The aquatic nuisance control records further reveal that shoreline accumulations are a general problem on Delavan Lake and are treated with copper sulphate weekly to control odors and increase the aesthetic and recreational value of the shoreline areas. Lake Winnebago had 1,500 /xg/1 of solids in the April through October samples. There is no question that plankton populations produce periodic nuisance conditions on Lake Winnebago, but the species composition estimates reveal the principal constituents of the population are diatoms and zooplankton. On only one occasion (June 15, 1966) did blue-green algae varieties (Anabaena sp., Aphanizomenon sp., and Anacystis sp.) thoroughly dominate the plankton populations. This observation was substantiated by the •o 248 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 I I I Figure 1. Plankton Volumes and Secchi Disks (April through November, 1966). 1970] L. Lueschow , J. Helm, D. Winter and G. Karl — Lakes 249 increased percentage of volatile solids. During the five months when varieties other than blue-green algae dominated the plankton population, the volatile solids averaged approximately 40 per cent. In June, when blue-green algae dominated the plankton volume, the volatile solids represented 90 per cent of the total solids. Pewaukee Lake has had a long history of algae nuisance control activity, and the blue-green varieties represented a significant por¬ tion of the plankton in six of the seven summer-fall collections. A general observation for Pewaukee Lake on June ninth revealed approximately 400 jig/1 total solids dominated by zooplankton with no indication of nuisance plankton. On August fourth, 700 fig/\ total solids were recorded dominated by Anabaena flosaquae. The visual observation of the lake suggested a distinct “bloom” con¬ dition. On October twenty-eighth, nearly 2,900 fig/ 1 of solids were recorded as a seasonal high in Pewaukee Lake. Lake Mendota is a lake that is sporadically plagued by nuisance algae blooms. On June eighth, a visual observation suggested a high algae population, and the Clarke-Bumpus sample recorded over 3,100 /mg/ 1 total solids dominated by Aphanizomenon sp. Seven other monthly visual observations on Lake Mendota failed to suggest bloom conditions. Total solids were less than 500 fig/ 1 in all other collections except September when a Ceratium sp. dominated 1,400 fig/ 1 total solids. This population was not visually noticeable at the time of collection and would not affect recreational potential. Oconomowoc and Middle Lakes revealed 500 fig/ 1 total solids on five of sixteen occasions, but the populations were dominated by diatoms or zooplankton. At no time were nuisance plankton con¬ ditions observed in the lake water. Round Lake revealed the lowest plankton values of the twelve lakes studied, even though organic nitrogen and Secchi disk obser¬ vations suggested that there should have been more plankton present. This apparent inconsistency may be partially explained by the fact that small algae cells will not be captured in the No. 20 mesh net. In the case of Round Lake, Chlorella sp. was observed in the water samples in sufficient quantities to color the lake water. In this case the plankton analysis techniques were not adequate to represent the actual condition of the lake. Figure I, which illustrates the plankton population, also shows Secchi disk readings at the time of plankton collection. In general, Delavan, Pewaukee, and Winnebago, which have relatively high plankton populations, have low Secchi disk readings. Lake Mendota, with variable plankton populations, had Secchi disk readings of twelve to sixteen feet in spring and fall but only six to nine feet in midsummer, with no apparent correlation to algae populations. 250 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 The lakes with relatively low plankton populations had higher Secchi disk readings, but the cause of variability was not deter¬ mined. Crystal Lake clarity was obvious both by visual observation and by Secchi disk readings. Nutrients Lake eutrophication, by definition, is an accumulation of nutri¬ ents. Many elements and compounds act as nutrients for the de¬ velopment of weeds and algae. However, nitrogen and phosphorus are usually considered the limiting nutrients and as such have received the most emphasis. Sawyer (1947) indicated that lakes containing 0.3 mg/1 inorganic nitrogen (N03-N, N02-N, and NH3N) and 0.015 mg/1 soluble phosphorus at time of spring turnover are capable of producing nuisance algae growths. Gerloff and Skoog (1957) suggested that the nitrogen-phosphorus ratio as it occurred in water was an indication of the ability of that water to produce algae, and their laboratory studies indicated that a ratio of 60 to 1 (nitrogen to phosphorus) was an appropriate ratio for optimum growth. Gerloff and Skoog further suggested that nitrogen was generally the limiting factor in algae production, while other studies (Federal Water Pollution Control Administra¬ tion, Lake Sebasticook) suggest phosphorus may be the limiting factor since ample nitrogen is available from the atmosphere through nitrification by bacteria and blue-green algae. Whatever the limiting factor in primary production may be, it appears that the concentration of nitrogen and phosphorus is useful in evaluat¬ ing the trophic condition of a particular lake. Table IV is a sum¬ mary of monthly nitrogen and phosphorus concentrations from the epilimnion and hypolimnion of the twelve lakes studied. Inorganic Nitrogen The inorganic nitrogen is available for utilization in the produc¬ tion of organic matter. The most obvious source of available nitrogen is nitrate nitrogen (N03). Nitrite nitrogen (N02) is an unstable state, and even eutrophic lakes typically have less than 0.01 mg/1. Ammonia nitrogen (NH3) is readily oxidized by nitri¬ fying bacteria in the presence of oxygen to nitrate nitrogen and is therefore essentially available for organic production in the epilimnion. Five of the twelve lakes studied in this investigation (Round, Mendota, Delavan, Pewaukee, and Winnebago) revealed an eleven- month mean total inorganic nitrogen concentration greater than the 0.3 mg/1 regarded as critical by Sawyer. Of these five lakes, Table 4. Monthly Nitrogen-Phosphorus Concentrations (milligrams per liter) for Twelve Wisconsin Lakes — 1966 1970] L. Lueschow, J. Helm, D. Winter and G. Karl — Lakes 251 Table 4. Monthly Nitrogen-Phosphorus Concentrations (milligrams per liter) for Twelve Wisconsin Lakes — 1966 — Continued 252 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 — — tv vO K - OO vO 00 Wo CO O — »0 O ffl >0 N i1 V/-s — i o ^ O O © © o © © wo tv O 00 00 of G N N O ^ O O 00 00 to co O tV (V G © tv © co tv — (V tV co of — oo oo tv of — of © G of © © oo oo of 00 O G of G G of — • O a ^ O' tv wa o tv wa — — © —i pa At tv © © IA (A N - 'A O' 00 © O O PA PA 00 — — ©©*-<©© -H -H -H -H -H -H -H At O >a K O (A M O' ^ \D OO tv At © tv — < © At wa pa © tv © © ■— ■ tv tV tv. 00 -A 00 00 © Wa © PA © tV (V) — tV _ 00 V\ OO "■ © O' o (As pa © pa tv — — « O O tV tv tv © © tv © — At PA © — i © — i — 1 oo oo O tv PA PA © © © — I WA ^ O - VA - - PA PA 00 © O tv © — i © — © o — © © © ’ V WA WA © oo © oo oo pa © tv © © © © wa tV {V At O t O' © oo © © O O >A N t' N N Y PA PA tV O WA O At © — - © tv «N © © © © o tV tV O PA © © © 00 Tf „ O „ in t >A V fV fV © oo © o oo oo — © (V — © -A tv © © ■ V <::::::: § I £ 6 6 § a cl UJhZZZhtih 2 pA> ~ tC © At © PA © © © -H -H -H -H -H -H -H lA lA t N fA © Af "t PA PA PA © © WA © © IV (V 00 tv tv © OO O oo At tv .— © © © tV PA At © — ©©©©© -H -H -H -H -H -H -H © © tV © © © \0 tv 05 O' *“* tv At a N N fA - N ® N O pa tx o pa oa — < tx o o o o o o o -H -H -H -H -H -H -H pa tx cx O' O O' OA CX O At PA — — 0 0 0 0 — 00 -H -H -H -H -H M -H - N IA vO N K N IA o O 't fA >C pa O O pa rf O O O' — O O' 00 00 Oa K - O K - A -t o o o o - - o -H -H -H — o o — 00 00 sO o o o N 00 O fA fA !A O tx — O co tx — tx O O O O O o o •H -H -H -H -H IX ■t — N -h N OO K IX Oa O lx O — A N o o o o o — -H -H oo oo tx tAi N PA O 00 - PA — O O — PA O O O sD — 00 PA IA o 00 A N — OOO — OO (X PA o At O O PA PA © O > o § o o 'C ■ § 32 £ a u h 6 S, 6 S oiHdo'axoono xsojai - — oiHdo^xng isojaj *Middle of hypolimnion. 1970] L. Lueschoiv, J. Helm, D. Winter and G, Karl — Lakes 263 3. The plankton sample collected with a No. 20 mesh net does not fully measure the standing crop of organic production since it ignores rooted or attached growths, and small algal cells may pass through the net. However, when 1,000 fig/ 1 of blue-green algae are recorded, there are sufficient algae in the water to be distinctly noticeable. Often a recording of 500 fig/ 1 in open water will be associated with wind-blown nuisance accumulation on spe¬ cific shorelines. Diatoms and zooplankton did not develop nuisance conditions on any of the lakes considered, so that absolute nuisance concentrations cannot be established. 4. The monthly analysis for organic nitrogen produced a ranking of the lakes that was reasonably consistent with transparency and plankton recreational potential. Furthermore, it offered an opportunity to translate plankton nuisance conditions into abso¬ lute values. The lakes that had a total organic nitrogen annual mean of less than 0.2 mg/1 had no algae or plankton nuisance. Lakes that had an organic nitrogen mean between 0.2 and 0.4 mg/1 have had rare algae nuisances, and lakes between 0.4 and 0.6 mg/1 had periodic algae blooms or substantial weed growths or both. Lakes with an annual total organic nitrogen mean of greater than 0.8 mg/1 had nuisance algae during most of the growing season. 5. The monthly analyses for soluble phosphorus had a coefficient of variation which approached 100 per cent, indicating that the eleven monthly samples were insufficient to develop any reliable confidence interval about a mean. 6. The monthly analysis for total phosphorus suggested that lakes with an annual mean of less than .03 mg/1 would be free of aquatic nuisances. Lakes with a total phosphorus annual mean between .03 and .05 mg/1 would be essentially free of aquatic nuisances. Lakes with an annual mean total phosphorus more than 0.1 mg/1 would experience nuisance weed growths or algae blooms during most of the growing season. 7. Eutrophication is not a simple process. It involves complex interrelationships between a variety of water quality parameters and an even greater variety of organisms. Although this study pro¬ vides some insight into these relationships, a great deal more will need to be learned in order to cope successfully with the demands currently being made for lake management. References Cited Birge, Edward A., Chancy Juday, 1911, The Inland Lakes of Wisconsin, Wis¬ consin Geological and Natural History Survey, Vol. No. 22. Beeton, Alfred M., 1965, Eutrophication of the St. Lawrence Great Lakes, Journal of Limnology and Oceanography, Vol. 10, No. 2, 264 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Sawyer, C. N., 1947, Fertilization of Lakes by Agricultural and Urban Drain¬ age, Journal New England Water Works Association, Vol. 61. Gerloff, G., and F. Skoog, 1957, Nitrogen as a Limiting Factor for the Growth of Microcystis Aeruginosa in Southern Wisconsin Lakes, Ecology 38, 4 (556). Federal Water Pollution Control Administration 1966, Fertilization and Algae in Lake Sebasticook, Maine. Technical Services Program, Federal Water Pollution Control Administration, Robert A. Taft, Sanitary Engineer Center, Cincinnati, Ohio. The authors are all public health biologists employed by the Department of Natural Resources, Division of Environmental Protection. ANNOTATED LIST OF THE FISHES OF WISCONSIN Marlin Johnson and George C. Becker In light of the increasing amount of work being done by state and federal agencies and by various state institutions on systematic and ecological as well as recreational and economic aspects of our fish fauna, the need has arisen for an up-to-date list, briefly noting the status of each species within the State of Wisconsin. Recent local studies on the distribution of fishes in various parts of the state have made possible the preparation of this list. Moreover, in recent years nomenclature for many species has been modified or clarified by national and international agencies ; these changes have been incorporated. Many significant changes in the distribution of certain Wiscon¬ sin fish species have occurred in recent years. These range changes have been brought about through fish rescue operations, crossover areas, canals, plantings by federal and state agencies, and the omnipresent fisherman’s minnow bucket. Wholesale modification of the landscape, through forestry and agricultural practices as well as dam and industrial construction, has left its mark on streams and lakes and is reflected in widespread alterations in species’ com¬ position and numbers. These changes are pointed out in the annota¬ tions. The list includes all the native fishes for Wisconsin and those exotic species planted with the intent that they become a char¬ acteristic part of the fish fauna. At the end of the paper, a separate list of problematical species includes those not known to be re¬ producing in the state and those found in nearby waters but not yet reported from the state. The status of each species is indicated by its general distribution and relative abundance within the state and by occasional reference to specific habitat. The following scale was used for indicating distribution and abundance : Rare — species which are taken at highly infrequent intervals with one or two specimens per collection. Uncommon — species which are taken infrequently and in very small numbers. 265 266 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Common — species which are taken frequently and in moderate numbers. Abundant — species taken frequently and in large numbers. The information has been compiled from the following sources : literature on Wisconsin fishes, personal communication with workers now actively engaged in fish research, fish collection rec¬ ords in the University of Wisconsin Zoological Museum and Biology Museum at Wisconsin State University-Stevens Point, and personal observations made by the authors. During the period from 1964- 1966 the senior author, in conjunction with Field Zoology classes, has made extensive collections (850 stations) throughout southern Wisconsin. The junior author has made detailed surveys of several large watersheds in the state. Specimens of most species lie in the Zoological Museum, University of Wisconsin, Madison, and in the Museum of Biology, Wisconsin State University, Stevens Point. The arrangement of orders and families follows the classification of Greenwood et al , 1966. Except where Dr. Bailey (see acknowl¬ edgements) has advised recent changes, nomenclature is accord¬ ing to Bailey et al, 1960. Certain synonomies have been included to facilitate cross reference to other literature dealing with fishes of our area (particularly Greene 1935, and Forbes and Richardson 1920) . Pertinent literature on distribution, ecology, and taxonomy is cited. Key rivers and lakes mentioned in the text are found on Map 1. The “lower Wisconsin River” refers to that section from the Prairie du Sac dam down to its juncture with the Mississippi River. The “lower Wolf River and its lakes” starts at the Shawano dam and includes Lake Winnebago. Acknowledgements We are grateful to W. Dryer and Stanford Smith of the U. S. Bureau of Fisheries, to Don Mraz, Arthur A. Oehmcke, Clarence Wistrom, John Truog, Cliff Brynildson, Lyle Christenson, John Brasch, Ruth Hine, and C. W. Threinen of the Department of Nat¬ ural Resources and to Dr. W. G. Reeder of the U. W. Zoology De¬ partment for their critical reading of parts of the manuscript and helpful suggestions. The help of E. J. Trimberger, formerly of the Wisconsin Department of Natural Resources, in obtaining distribu¬ tion data from the Mississippi River area, and of Philip Smith, Illinois Natural History Survey, in checking synonomies is also greatly appreciated. We thank the following commercial fishermen who during 1968 provided specimens from Lakes Michigan and 1970] Johnson and Becker — -Fishes of Wisconsin 267 Superior: Evald Heinonen, Harvey Hadland, Wm. G. Bodin, Howard Weborg, Everett Marks, John Reynen, Bernard Moes, Nor- bert Swaer, Frederick Topel, and Walter Voight. Special thanks are due Reeve Bailey, Curator of Fishes, Museum of Zoology, Univer¬ sity of Michigan, for verification of certain identifications and for help with nomenclatural changes, and Miss Peggy Brogan for draft¬ ing the map. Support for publication costs came from University of Wisconsin Center System. Map 1. Map of Wisconsin showing major drainage units, counties, key rivers, and lakes and cities. 268 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 ORDER PETROMYZONTIFORMES PETROMYZONTIDAE — LAMPREYS Ichthyomyzon castaneus Girard — Chestnut lamprey Uncommon to common in the lower Wisconsin, Yellow (Wood Co.), Mississippi, St. Croix and Nemakagon Rivers. In Great Lakes drainage present in the upper Fox R. at Eureka (Winnebago Co.) and reported from the lower Wolf R. and its lakes. 16, 65 Ichthyomyzon fossor Reighard & Cummins — Northern brook lam¬ prey Uncommon in Wisconsin R. and its tributaries. Common in small to medium-size streams of central Wisconsin flowing into L. Michi¬ gan. Distribution widespread in streams flowing into L. Superior. 7, 16, 25 Ichthyomyzon unicuspis Hubbs & Trautman — Silver lamprey Ichthyomyzon concolor (Kirtland) F&R* Uncommon in Mississippi and lower Wisconsin Rivers, Green Bay, and the large lakes of the lower Wolf R. basin. Common in streams flowing into L. Superior. 9, 13, 15, 16 Lampetra lamottei (Lesueur) — American brook lamprey Entosphenus appendix (DeKay) G.f Lampetra wilderi Gage F&R Common in creeks of the Mississippi basin throughout southern and central Wisconsin. So far not reported from streams of Great Lakes drainage in the state, although reported as common in upper Michigan streams of eastern L. Superior. 7, 13, 16, 58 Petromyzon marinus Linnaeus — Sea lamprey Recently abundant in L. Michigan. Common in L. Superior. Spawning and ammocete development occurring in streams in their watersheds. Exotic, invading L. Michigan in mid-1930’s and L. Superior in the late 1940’s. Currently subject to control with 1am- pricides and other methods, but in last two years increasing in numbers (Moore, James D. 1969. Lake trout lamprey scarring in North Green Bay. Wis. Dept. Nat. Resources. Oshkosh. 5p. mimeo). 16, 69. * Forbes & Richardson, 1920. f Greene, 1935. 1970] Johnson and Becker — Fishes of Wisconsin 269 ORDER ACIPENSERIFORMES ACIPENSERIDAE — STURGEONS Acipenser fulvescens Rafinesque — Lake sturgeon Acipenser ruhicundus Lesueur F&R Common in the Menominee R. (Wis.-Mich. border) and in the lower Wolf R. and its lakes, particularly Poygan and Winnebago. Common in L. Wisconsin (Sauk & Columbia Cos.). Common in St. Croix R. to Gordon Dam and in Namekagon R. below Trego dam. Common in both the Chippewa and Flambeau Rivers. Present in Benson Lake, the widespread of Manitowish R. (Vilas Co.) and in the Clam R. (Burnett Co.). Verified report from Big Cedar L. (Washington Co.) in 1961, resulting from 1936 planting. Rare in Lakes Michigan and Superior. 9, 13, 65, 91. Scaphirhynchus platorynchus (Rafinesque) — Shovelnose sturgeon Uncommon to common in the main channels of the Mississippi and lower Wisconsin Rivers and in the lower Chippewa and lower Red Cedar Rivers. Presence reported up to St. Croix Falls dam on the St. Croix R. 13 POLYODONTIDAE— PADDLEFISHES Polyodon spathula (Walbaum) — Paddlefish Formerly abundant on the Mississippi R., now uncommon in the Mississippi and lower Wisconsin Rivers. 13, 23, 108 ORDER SEMIONOTIFORMES LEPISOSTEIDAE — GARS Lepisosteus osseus (Linnaeus)— Longnose gar Common in most large lakes and quiet waters of larger rivers over lower two-thirds of Wisconsin. In northwestern Wisconsin common in Big Sissabagama, Big Court Oreilles, Grindstone, and Big Sand Lakes (Sawyer Co.). Common in the St. Croix R. below dam at St. Croix Falls; abundant in the Island Lake Chain (Rusk Co.) and the Long Lake Chain (Chippewa Co.). Uncommon in northeastern Wisconsin. 9, 10, 13, 65 Lepisosteus platostomus Rafinesque— Shortnose gar Uncommon to common in lower Wisconsin and Mississippi Rivers and lower portions of their tributaries; in the St. Croix up to the St. Croix Falls Dam. Formerly reported from Lake Mendota (Dane Co.). Recently appearing in Lake Winnebago (Great Lakes drain- 270 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 age), possibly entering via the Fox-Wisconsin canal at Portage (Columbia Co.) . Several specimens taken summer 1968 in southern third of Green Bay (L. Michigan). 9, 13, 67, 87 ORDER AMIIFORMES AMIIDAE — BOWFINS Amia calva Linnaeus — Bowfin Uncommon to common in quiet waters of large rivers and large lakes. Wider dispersal and less common northward. 9, 10, 13, 65, 69 ORDER ANGUILLIFORMES ANGUILLIDAE — FRESHWATER EELS Anguilla rostrata (Lesueur) — American eel Anguilla bostoniensis (Lesueur) G. Anguilla chrysypa Rafinesque F&R Rare in the Mississippi R. and its tributaries. Report from L. Neshonic (La Crosse Co.) about 1954 (letter from Lyle Christen- son, April 25, 1969). Reports for 1966 from mouth of St. Croix (Burnett Co.) and Chippewa (Buffalo Co.) Rivers, and Lake Ne- bagamon (Douglas Co.). Taken regularly on Red Cedar R. up¬ stream to L. Menomin (Dunn Co.). Single specimen collected from L. Superior (Beaver Lake Cr., Alger Co., Mich.). Single specimen collected at Red Banks in Green Bay of L. Michigan, summer 1968. Also collected in Lake La Belle, 1959, and Lake Nemahbin (Wau¬ kesha Co.), 1951. 46, 67, 69 ORDER CLUPEIFORMES CLUPEIDAE — HERRINGS Alosa chrysochloris (Rafinesque) — Skipjack herring Pomolobus chrysochloris Rafinesque F&R G. Probably extinct. Formerly found throughout the Wisconsin por¬ tion of the Mississippi R. and in the St. Croix up to St. Croix Falls. 23, 46 Alosa pseudoharengus (Wilson) — Alewife Abundant recent (1952) addition to fauna of L. Michigan and has become a nuisance. Appeared in L. Superior in 1954, becoming common. 68, 79, 99 Dorosoma cepedianum (Lesueur) — Gizzard shad Abundant in Mississippi and lower Wisconsin Rivers. Uncommon to common in lower portions of their larger tributaries. Common in 1970] Johnson and Becker — Fishes of Wisconsin 271 St. Croix R. upstream to St. Croix Falls Dam. Rare, southern L. Michigan. Uncommon in lower third of Green Bay of L. Michigan where collected by commercial fishermen summer 1968. 18, 22, 65, 68 HlODONTIDAE — MOONEYES Hiodon alosoides (Rafinesque) — Goldeye Amphiodon alosoides Rafinesque G. Rare in Mississippi R. as far north as Lake Pepin. 23, 46 Hiodon tergisus Lesueur— -Mooneye Common in Mississippi, St. Croix R. upstream to St. Croix Dam, lower Wisconsin R., and uncommon in lower portions of their larger tributaries. Common in lower Wolf R. and its lakes (Great Lakes drainage) . Rare in Green Bay and L. Michigan. 9, 13, 22, 65 ORDER SALMONIFORMES SALMONIDAE — TROUTS, WHITEFISHES Coregonus clupeaformis (Mitchill) — Lake whitefish Common in L. Superior (up to 35 fathoms) ; common in L. Mich¬ igan but reduced in recent years by the sea lamprey. Reported from L. Lucerne (Forest Co.), Keyes L. (Florence Co.) , and Trout L. (Vilas Co.). 29, 31, 34, 75, 91, 107 Leucichthys alpenae (Koelz) — Longjaw cisco Formerly common in L. Michigan (20 to 60 fathoms) , but becom¬ ing very rare in recent years. 96 Leucichthys artedii Lesueur— Cisco or lake herring Argyrosomus artedi (Lesueur) F&R Common but declining in Lakes Superior and Michigan. Com¬ mon in many deeper inland lakes in northern tier of counties ; rare to common in lakes of Waukesha Co. and in L. Geneva (Walworth Co.). Nearing extinction in Lake Mendota (Dane Co.). 20, 24, 31, 42, 53, 63, 96 Leucichthys hoyi (Gill)— Bloater Abundant and dominant Leucichthys at 20 to 70 fathoms, but some found to greatest depths in both Lakes Superior and Michi¬ gan. Increased rapidly in numbers and extended range in L. Michi¬ gan during 1950’s and early 1960’s but declining in recent years as alewives increased. 31, 61, 96, 110, 111 272 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Leucichthys johannae ( Wagner) — Deepwater cisco Perhaps now extinct, but formerly common in deeper waters of L. Michigan (30 to 90 fathoms). Last seen and taken in the early 1950’s. 96 Leucichthys kiyi (Koelz) — Kiyi Common in deep areas of L. Superior. Have become very rare in recent years in L. Michigan (60 to 100 fathoms). 31, 96 Leucichthys nigripinnis (Gill) — Blackfin cisco Once common but now rare in L. Superior (15 to 100 fathoms). Formerly common in L. Michigan but now probably extinct — last records from the mid-1950’s. 31, 96 Leucichthys reighardi (Koelz) — Shortnose cisco Rare in western L. Superior (up to 50 and possibly 65 fathoms). Once common in L. Michigan (20 to 60 fathoms) but very rare in recent years. 31, 96 Leucichthys zenithicus (Jordan & Evermann) — Shortjaw cisco Common in L. Superior (10 to 90 fathoms but most common at 30 to 70 fathoms) . Once common in L. Michigan (20 to 70 fathoms) but decreasing to very rare in recent years. 31, 96 Prosopium coulteri (Eigenmann and Eigenmann) — Pygmy white- fish Common in L. Superior (10 to 59 fathoms, but most common at 20 to 50 fathoms) . 31, 37 Prosopium cylindraceum (Pallas) — Round white-fish Prosopium quadr Hater ale quadrilaterale (Richardson) G. Common in L. Superior (shallows to 19 fathoms; rarer up to 40 fathoms) . Uncommon to common in shallower areas of L. Michi¬ gan north of Sheboygan. Rare southwards. 31, 73 Salmo gairdneri Richardson — Rainbow trout Introduced in late 1800’s. Common locally in streams and lakes over the state. Continuously stocked in L. Superior and its tribu¬ taries; spawning successfully in some larger tributaries. Continu¬ ously stocked in recent years along Wisconsin shore of L. Michigan ; natural reproduction insignificant. 19, 27b 1970] Johnson and Becker — Fishes of Wisconsin 273 Salmo trutta Linnaeus — Brown trout Salmo fario Linnaeus G. Introduced in late 1800’s. Common in cold-water streams of southern and central Wisconsin, and in recent years playing a larger role in stream fishing of northern Wisconsin. Large brown trout are taken frequently in shore areas of Lakes Superior and Michi¬ gan, where they are maintained by extensive stocking. 18, 27c, 69 Salvelinus fontinalis (Mitchill) — Brook trout Common in streams of central and northern Wisconsin; rare to uncommon in southern Wisconsin, except Richland, Columbia, Dane and Sauk Cos. where common in some streams. Coasters present along Wisconsin shores of Lakes Superior and Michigan (especially Door and Kewaunee Cos.) ; these populations sustained by stocking. 15, 17 Salvelinus namaycush namaycush (Walbaum) — Common lake trout Cristivomer namaycush (Walbaum) G. & F&R Common in L. Superior (10 to 39 fathoms). Uncommon in L. Michigan but returning in numbers. Inland waters with spawning trout: Trout and Black Oak L. (Vilas Co.), Big Green L. (Green Lake Co.). Recently introduced in Lac Court Oreilles (Sawyer Co.) 27a, 31, 35, 36, 48 Salvelinus namaycush siscowet (Agassiz) — Siscowet Common in deeper waters (40 to 125 fathoms) of L. Superior. 33 OSMERIDAE — SMELTS Osmerus mordax (Mitchill) — American smelt Common in L. Superior and Michigan and occasionally taken in large tributary streams. First taken off Wisconsin shores in L. Michigan in 1928 from 1912 stocking of Crystal R., Benzie Co., Michigan. Reached Wisconsin shores of L. Superior in late 1930’s. Populations reproducing in L. Lucerne (Forest Co.) ; also reported from Sand Bar, Tomahawk and Big Diamond Lakes (Bayfield Co.). 26, 31, 106 Esocidae— pikes Esox americanus vermiculatus Lesueur — Grass pickerel Esox vermiculatus Lesueur G. & F&R Common in scattered localities in lakes and sluggish waters of southern one-third of state. Also found in Fishtrap and High Lakes 274 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 and headwaters of Manitowish R. (Vilas Co.) and Minocqua Thoroughfare (Oneida Co.), where it was probably accidentally in¬ troduced during fish transfer operations in the early 1930’s. 10, 64 Esox Indus Linnaeus- — Northern pike Common to abundant in lakes throughout state and in slow waters of large streams and rivers. Absent from the Chippewa R. and its lakes above Radisson (Sawyer Co.). 7„ 9, 10, 12, 13, 69, 102, 103 Esox masquinongy Mitchill — Muskellunge Common in the lakes and rivers in the headwater regions of the Chippewa, Flambeau, St. Croix (upstream to Trego Dam), and Wisconsin Rivers. Uncommon to rare in the middle one-third of the state. Stocked annually as far south as L. Wisconsin (Columbia Co.) . Populations developed from stocking in Little Green L. (Green Lake Co.), Pewaukee L. (Waukesha Co.), and Lenwood L. (Wash¬ ington Co.). Occasionally caught in L. Superior and in Green Bay (L. Michigan). 9, 10, 43, 64, 69, 81, 83 Esox masquinongy female X Esox lucius male — Tiger muskellunge Natural crosses are reported in Lac Vieux Desert (Vilas Co.), Star, Big St. Germaine and Plum Lakes (Vilas Co.), and Toma¬ hawk and Minocqua Lakes (Oneida Co.). Hybrids have been pro¬ duced in Wisconsin hatcheries in 1940, 1946, 1947, 1963, 1965 and stocked frequently in landlocked bass lakes. This hybrid has a growth rate more rapid than either parent species. Experiments indicate that it will backcross with Esox lucius . 14, 65, 69 UMBRIDAE — MUDMINNOWS Umbra limi ( Kir t land) — Central mudminnow Common to abundant in small streams and marshes throughout state except in southwestern quarter, where uncommon. 7, 9, 12, 13, 65, 69 ORDER CYPRINIFORMES CYPRINIDAE— MINNOWS AND CARP Campostoma anomalum pullum (Agassiz)— Central stoneroller Abundant in small swift-flowing streams of southern Wisconsin. Occasionally taken in quiet pools. 13, 57, 80 1970] Johnson and Becker — Fishes of Wisconsin 275 C amp o stoma anomalum oligolepis Hubbs & Greene — Largescale stoneroller Abundant in medium-size, swift-flowing streams in central and northern Wisconsin. 13, 57, 80 Carassius auratus (Linnaeus)— Goldfish Common in some Milwaukee Co. lagoons. Occasionally found in southeastern Wisconsin streams and in Peters Lake (Walworth Co.). Introduced in part through a fish exchange program with the Nebraska Fish Commission in 1903, 1904, and 1908. 67, 78, 113 Cyprinus carpio Linnaeus— Carp Abundant in large shallow waters of southern and central Wis¬ consin. Becoming common in some northern Wisconsin waters in recent years. Introduced through plantings by Wis. Commissioners of Fisheries, 1881-1895. 30, 41, 69, 74, 113 Chrosomus eos Cope — Northern red-belly dace Abundant in small streams and in bog lakes of central and north¬ ern Wisconsin. 7, 12, 69 Chrosomus erythrogaster (Rafinesque) — Southern red-belly dace Abundant in small to medium-size streams in southern Wisconsin. Rare to uncommon in central Wisconsin, apparently moving into a number of new localities in recent years. 13 Chrosomus neogaeus (Cope) — Finescale dace Pfrille neogaea (Cope) G. Uncommon to common in small streams and ponds in north¬ eastern Wisconsin and in the streams of the L. Superior drainage. Rare in headwater streams of central Wisconsin. 12, 69 Clinostomus elongatus (Kirtland) — Redside dace Uncommon in small to medium-size streams in widely scattered basins of southern, central and eastcentral Wisconsin. 7, 13 Dionda nubila (Forbes) — Ozark minnow Hybognathus nubila (Forbes) F&R Rare in medium-size streams of gentle current Platte R. basin (Grant Co.). Reported from streams in Barron, Lafayette, Iowa, Walworth, Rock, and Waukesha Cos. in late 1920’s. 13, 46 276 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Hybognathus hankinsoni Hubbs — Brassy minnow Common in slow-flowing streams over state except northwestern portion where uncommon. Rare in large rivers. 7, 13, 46 Hybognathus nuchalis Agassiz — Silver minnow Uncommon in the lower Wisconsin R. and the Mississippi and in the lower portions of their tributaries. 13, 46 Hybopsis aestivalis (Girard) — Speckled chub Hybopsis hyostomus (Gilbert) F&R Extrarius aestivalis (Gilbert) G. Generally uncommon over shallow sand flats in lower Wisconsin and Mississippi R. and lower portions of their larger tributaries. 13, 46 Hybopsis biguttata (Kirtland) — Hornyhead chub Hybopsis kentuckiensis (Rafinesque) F&R Nocomis biguttatus (Kirtland) G. Common in dear-water, medium-size streams of northern and central Wisconsin. Uncommon in the southwestern quarter of state except in the southern tier of counties where common. 7, 12, 13, 46 Hybopsis plumb ea (Agassiz) — Lake chub Couesius plumb eus (Agassiz) G. Uncommon in shoal waters in the vicinity of stream mouths in Green Bay and L. Michigan. Common near mouths of streams in Bayfield and Douglas Cos. (L. Superior drainage). 45, 69 Hybopsis storeriana (Kirtland) — Silver chub Uncommon in flowing sections of the lower Wisconsin R., in the Mississippi R., and in the lower portions of their tributaries. 13, 46 Hybopsis x-punctata Hubbs & Crowe — Gravel chub Hybopsis dissimilia (Kirtland) F&R Erimystax dissimilis (Kirtland) G. Probably extinct in state. Taken only once from the Sugar R. (Green Co.) in the late 1920’s. 46 Notemigonus crysoleucas (Mitchill) — Golden shiner Abramis crysoleucas (Mitchill) F&R Common to abundant in lakes, slow-flowing streams and rivers over the entire state. 7, 9, 10, 12, 13, 22, 69 1970] Johnson and Becker — Fishes of Wisconsin 277 Notropis amnis Hubbs & Greene — Pallid shiner Pare. In recent years this minnow has been collected from the lower Wisconsin and from the Mississippi Rivers in water of moderate flow. 13, 46 Notropis ano genus Forbes — Pugnose shiner Rare. Earlier reports from Burnett, Waupaca, Kewaunee, Mar¬ quette, Columbia and Dane Cos. In recent years this minnow has been collected only from Pewaukee Lake (Waukesha Co.) and L. Poygan (Winnebago Co.) . 2, 9, 10, 46 Notropis atherinoides Rafinesque — Emerald shiner Common to abundant in L. Michigan, Superior, Winnebago and other large inland lakes of central and southern Wisconsin. Present in Yellow Birch L. (Vilas Co.). A common minnow in the lower Wisconsin and Mississippi R. and lower portions of some of their tributaries. 9, 13, 22, 69 Notropis blennius (Girard) — River shiner Notropis jejunus (Forbes) F&R Common in the lower Wisconsin and the Mississippi R. and lower portions of some tributary streams. An isolated population in L. Winnebago of the Great Lakes Basin. 9, 13„ 46 Notropis buchanani Meek — Ghost shiner Rare or nearly extinct in the state. Last collected in 1944 (UW Museum of Zoology — Madison) from the Mississippi R. opposite Crawford Co. 46 Notropis chalybaeus (Cope) — Ironcolor shiner Camm Swift, Dept, of Biological Sciences, Florida State Univer¬ sity, Tallahassee, writes that two collections which Greene (46) catalogued originally as Notropis texanus richardsoni are Notropis chalybaeus (letter November 8, 1968). The two series follow: From UMMZ 66537 (7 of 41) Wise., Collumbia Co., Fox R. opposite Lock 25. VIII :26 :1925 Green and Jones. From UMMZ 74054 (2 of 75) Wise., Waupaca Co., Blake Cr., 5 ml. W. Symco. VII :9 : 1926 Greene and Lo Criccho. Notropis cornutus (Mitchill) — Common shiner Abundant and one of the commonest of stream and river min¬ nows found under a wide variety of conditions. Occasional in dear- water lakes over clean bottom. 7, 9, 10, 13, 44, 65, 69 278 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Notropis chrysocephalus (Refinesque) — Stripped ishiner Common in the Saukville-Grafton sector of the Milwaukee R. (Ozaukee Co.). Single specimen (Museum WSU-Stevens Point) collected from Green Bay near the city of Sturgeon Bay. Older records indicate presence from Kenosha to Dane Cos. 44 , 46 Notropis dorsalis (Agassiz)- — Bigmouth shiner Notropis gilherti Jordan and Meek F&R Common over sand-bottom, medium-sized streams in the Missis¬ sippi drainage. Recently established in the east-central streams within the L. Michigan drainage of Wisconsin. 7, 13 Notropis heterodon (Cope) — Blackchin shiner Uncommon in central and southern Wisconsin. Absent from southwestern quarter. Locally common northward in lakes and bog ponds. 9, 10, 12, 46, 65 Notropis heterolepis Eigenmann & Eigenmann — Blacknose shiner Notropis cayuga Meek F&R Rare in southwestern quarter of state. Elsewhere common in widely separated lakes and slow-moving streams, often in heavily- silted habitats. 7, 9, 10, 12, 46, 65 Notropis hudsonius (Clinton) — Spottail shiner Common locally in very large inland lakes, in Lakes Michigan and Superior, and in large slow-moving rivers as the lower Wis¬ consin and Mississippi, and in the St. Croix R. upstream to St. Croix Falls Dam. 9, 10, 12, 13, 22, 46, 65, 69 Notropis rubellus (Agassiz)- — Rosyface shiner Notropis rubrifrons (Cope) F&R Common in medium-sized, swift-flowing streams of central and southern Wisconsin. Uncommon northward — apparently absent from the extreme northwest corner of the state. 7, 12,, 13, 46, 69 Notropis spilopterus (Cope) — Spotfin shiner N otropis whipplii (Girard) F&R G. Abundant in medium to large-sized streams and rivers. A com¬ mon minnow in large lakes. Absent from the northern tier of counties. 7, 9, 10, 12, 13, 65, 69 1970] Johnson and Becker — Fishes of Wisconsin 279 Notropis stramineus (Cope)— Sand shiner Notropis blennius (Girard) F&R Notropis phenacobius Forbes F&R Notropis deliciosus (Cope) G. Abundant in medium to large-sized streams and rivers of central and southwestern Wisconsin. Elsewhere uncommon to common. 7, 9, 12, 13, 69 Notropis texanus (Girard)— Weed shiner Notropis nux Hubbs & Greene G. Uncommon in the lower Wisconsin and in the Mississippi Rivers and in the lower portions of their tributaries. Older records indi¬ cate presence from east-central Wisconsin (Great Lakes drainage). 13, 22, 46 Notropis umhratilis (Girard) — Redhn shiner Locally common in slow-moving, turbid waters of southeastern Wisconsin. Older records show presence in widely isolated streams throughout the southern half of the state. 46, 54 Notropis volucellus volucellus (Cope)— Northern mimic shiner Notropis blennius (Girard) F&R Locally common in medium-sized streams and in lakes over the state except in the southwestern quarter, where rare. Distribution sites widely isolated. 7„ 22, 65, 69 Notropis volucellus wickliffi Trautman — Channel mimic shiner Rare in the Mississippi R. Seldom taken in recent years. 13, 46 Opsopoeodus emiliae Hay— Pugnose minnow Uncommon in slow-water and sloughs of the Wisconsin R. up¬ stream to Du Bay (Marathon Co.) and in the Mississippi R. In the Great Lakes drainage taken only from the west end of L. Poy- gan (Waushara Co.) . 9, 13, 22, 46 Phenacobius mirabilis (Girard) — Suckermouth minnow Uncommon to common in small and medium-sized tributaries to the lower Wisconsin and Mississippi Rivers. In the southwestern quarter of the state, from Vernon Co. southward. Uncommon in Rock R. drainage in southeastern quarter. 13, 46 280 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Pimephales notatus (Rafinesque) — Bluntnose minnow Hyborhynchus notatus (Rafinesque) G. Abundant in streams and lakes over the entire state. Uncommon in the larger rivers. This species, from the standpoint of distribu¬ tion and numbers, is perhaps the most successful Wisconsin fish. 6, 7, 9, 10, 12, 13, 22, 65, 69 Pimephales promelas Rafinesque — Fathead minnow Locally common over the entire state. Frequently associated with turbid water. 7, 12, 13, 22, 46, 69 Pimephales vigilax (Baird & Girard) — Bullhead minnow Cliola vigilax (Baird and Girard) F&R Hy par gyrus velox (Girard) G. Common to abundant in the lower Wisconsin and the Mississippi. Rarely associated with other large streams in the southwestern quarter of state. A recent collection from the Fox R. (Marquette Co.) near the Portage Canal indicates a late crossover into the Great Lakes drainage basin. 7, 22 Rhinichthys atratulus (Hermann) — Blacknose dace Rhinichthys atronasus (Mitchill) F&R. G. Common to abundant in small cool headwater streams. Uncom¬ mon to common in medium-sized streams supporting trout. Dis¬ tributed throughout the state. 6, 8, 12, 13, 69 Rhinichthys cataractae (Valenciennes) — Longnose dace Common to abundant in fast water of medium-sized streams of the northern half of Wisconsin. Common in small fast-water streams of southwestern Wisconsin. Common in wave-swept shal¬ lows of Lakes Michigan and Superior. 6, 8, 12 ,13, 69 Semotilus atromaculatus (Mitchill) — Creek chub Abundant in small and medium-sized streams and rivers over the entire state. Rare in large rivers and in lakes. One of our common¬ est fishes. 7, 12, 13, 65, 69 Semotilus margarita (Cope) — Pearl dace Margariscus margarita (Cox) G. Common in very small streams of central and northern Wis¬ consin except in streams of L. Superior drainage where rare. Un¬ common in larger streams. 7, 12, 65, 69 1970] Johnson and Becker — Fishes of Wisconsin 281 CATOSTOMIDAE— SUCKERS Carpiodes carpio (Rafinesque) — River carpsucker Common in the lower Wisconsin R. and the Mississippi R.,, and their larger tributaries. 13, 22, 46 Carpiodes cyprinus (Lesueur) — Quillback Carpiodes thompsoni Agassiz F&R Carpiodes velifer (Rafinesque) F&R Abundant in the lower Wisconsin R. and the Mississippi R. and their larger tributaries. Common Lakes Poygan and Winnebago. 9, 13, 65 Carpiodes velifer (Rafinesque) — Highfin carpsucker Carpiodes difformis Cope F&R Common in the lower Wisconsin R. and the Mississippi R. and in their larger tributaries. 13, 46 Catostomus catostomus (Forster) — Longnose sucker Common in L. Superior and its tributaries during spawning. Formerly common, now rare in L. Michigan. 31, 46, 69 Catostomus commersoni (Lacepede) — White sucker Abundant and generally distributed in lakes and streams over the state. One of the most widely distributed and abundant fish species in the state. 6, 7, 9, 10, 12, 65, 69 Cycleptus elongatus (Lesueur)-— Blue sucker Rare. Found only in the lower Wisconsin R., the Mississippi R., and the St. Croix upstream to St. Croix Falls Dam. 13,, 46, 65 Erimyzon oblong us ( Mitchill ) — Creek chubsucker Erimyzon sucetta oblong us (Mitchill) F&R Rare. Taken only twice in the southeastern corner of Wisconsin during the late 1920’s from the Des Plaines R. (Kenosha Co.) and a tributary. 46 Erimyzon sucetta (Lacepede) — -Lake chubsucker Erimyzon sucetta oblongus (Mitchill) F&R Rare to uncommon locally in the larger rivers and the lower por¬ tions of tributaries to them in the southern half of Wisconsin. Report from White Clay L. (Shawano Co.) needs substantiation. Occasionally taken in larger lakes, especially in southeastern Wis¬ consin. 9, 10, 13, 46 282 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Hypentelium nigricans (Lesueur) — Northern hogsucker Catostomus nigricans Lesueur F&R Common locally in riffle areas of medium to large streams and rivers. Generally distributed throughout the state. 7, 12, 13, 65 Ictiobus bubalus (Rafinesque) — Smallmouth buffalo Uncommon in the lower Wisconsin R. and in the Mississippi. Recent record from Island L. (Vilas Co.), needs verification. 13, 22, 46 Ictiobus cyprinellus (Valenciennes) — Bigmouth buffalo Megastomatobus cyprinella (Cuvier & Valenciennes) G. Uncommon to common in medium to large rivers in southern Wisconsin. Reported from L. Delavan (Waukesha Co.), L. Kosh- konong (Jefferson Co.), Beaver Dam L. (Dodge Co.), the Madison Lakes (Dane Co.) and Long Lake (Waupaca Co.). The last is the first Wisconsin record from the L. Michigan drainage. Present in the Mississippi R. and in the St. Croix up to St. Croix Falls Dam. Reports needing substantiation from Manitowish Chain and Big Lake (Vilas Co.) . 13, 22, 46, 65, 67 Ictiobus niger ( Rafinesque ) — Black buffalo Ictiobus urus (Agassiz) F&R Rare on the lower Wisconsin R. Uncommon on the Mississippi R. 13, 45 Minytrema melanops (Rafinesque) — Spotted sucker Common locally in the lower Wisconsin and in the Mississippi Rivers and their larger tributaries. Common in Lake Poygan; occasional in L. Winnebago. It has been reported from the upper Fox R. (Columbia to Winnebago Cos.) and from the lower Wolf downstream from the Shawano Dam (Shawano Co.). Recently collected in Des Plaines R. (Kenosha Co.). 9, 13, 22 Moxostoma anisurum (Rafinesque) — Silver redhorse Common locally in large streams in the western half of Wis¬ consin and in streams in the L. Superior drainage. Uncommon in east-central Wisconsin (L. Michigan drainage). 7, 65, 68, 69 Moxostoma carinatum (Cope)— River redhorse This species has not been collected by Wisconsin workers but is reported in boundary waters in Lake St. Croix on the St. Croix R. between Minnesota and Wisconsin. 82b 1970] Johnson and Becker — Fishes of Wisconsin 283 Moxostoma duquesnei (Lesueur) — Black redhorse Probably extinct. Taken only once from Black Earth Creek (Dane Co.) in the late 1920’s. 46 Moxostoma erythrurum (Rafinesque) — Golden redhorse Moxostoma aureolum (Lesueur) F&R Uncommon in northern half of state (absent from northern tier of counties). Common in medium to large rivers in the southern half of the state. 7, 9, 12, 13 Moxostoma macrolepidotum (Lesueur) — Northern redhorse Moxostoma breviceps (Cope) F&R Moxostoma aureolum (Lesueur) G. Common statewide in medium to large rivers. A common species in large lakes of central and northern Wisconsin. 7, 9, 12, 13, 22, 65, 69. Moxostoma valenciennesi Jordan — Greater redhorse Moxostoma rubreques Hubbs G. Rare and probably nearing extinction. Old records indicate gen¬ eral distribution in the state. A purported specimen recently re¬ ported from the lower Wisconsin R. misidentified as this species, actually M. macrolepidotum. 13, 46 ORDER SILURIFORMES ICTALURIDAE — FRESHWATER CATFISHES Ictalurus furcatus (Lesueur) — Blue catfish Rare, probably extinct. Only two old records from the Mississippi R. (Crawford and Pepin Cos.). 46 Ictalurus melas (Rafinesque) — Black bullhead Ameiurus melas (Rafinesque) F&R G. Abundant throughout the state in lakes and warm-water streams of all sizes. 7, 9, 10, 12, 13, 22, 65, 68, 69 Ictalurus natalis (Lesueur) — Yellow bullhead Ameiurus nebulosus (Lesueur) F&R. G. Common throughout the state, generally in clear medium-sized streams and occasionally in clear lakes. 7, 9, 10, 12, 13, 22, 65, 69, 89 284 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Ictalurus nebulosus (Lesueur) — Brown bullhead Ameriurus nebulosus (Lesueur) F&R. G. Uncommon in sloughs of rivers and in lakes. Discontinuous dis¬ tribution throughout state. 7, 9, 12, 13, 22, 69, 89 Ictalurus punctatus (Rafinesque) — Channel catfish Villarius lacustris (Walbaum) G. Uncommon to common in the Mississippi R. and the lower Wis¬ consin R. (upstream to Castle Rock Dam) and their larger tribu¬ taries, and in the St. Croix R. to Gordon Dam and in the Name- kagon to Trego Dam. Uncommon to common in the Wolf and Fox River system of the L. Michigan basin. Rare in L. Michigan and Green Bay. In the L. Superior drainage a single recent record from the St. Louis R. (Douglas Co.). 9, 13, 22, 65, 69 Noturus exilis Nelson — Slender madtom Schilbeodes exilis (Nelson) F&R. G. Rare in Bark R. system (Waukesha Co. ). 20, 46 Noturus flavus Rafinesque — Stonecat Uncommon to common in medium-sized streams of moderate cur¬ rent in southern two-thirds of state. Rare northward except in L. Superior tributaries (Bayfield and Douglas Cos.), where common. 7, 12, 13, 65, 69 Noturus gyrinus (Mitchill) — Tadpole madtom Schilbeodes gyrinus (Mitchill) F&R. G. Common statewide in medium to large rivers. Frequently found in lakes over debris-covered bottom. 7, 9, 12, 13, 22, 69 Pylodictis olivaris (Rafinesque) — Flathead catfish Leptops olivaris (Rafinesque) F&R Uncommon to common in Mississippi, lower Wisconsin and Peca- tonica Rivers. In recent years reported occasionally from lower Wolf and upper Fox Rivers and their lakes (L. Michigan drainage) . 9, 13, 22, 65 Aphredoderidae— pirate perch Aphredoderus say anus (Gilliams)- — Pirate perch Uncommon to rare in sloughs of the Mississippi R. and in the Wisconsin R. up to central Wisconsin. Occasionally found in lower portions of tributaries to these waters. Uncommon in Des Plaines R. (Kenosha Co.). 13, 22 1970] Johnson and Becker — Fishes of Wisconsin 285 ORDER PERCOPSIFORMES PERCOPSIDAE — TROUT-PERCH Percopsis omiscomaycus (Walbaum) — Trout perch Percopsis guttatus Agassiz F&R Uncommon in the Mississippi and Wisconsin Rivers. More com¬ mon northward, in the Chippewa R. and connecting lakes of Sawyer Co., and in L. Superior and tributary streams. Reported from Trout L. (Vilas Co.). Rare to uncommon in L. Michigan drainage except in Lakes Winnebago and Poygan (Winnebago Co.), where abun¬ dant. 9, 18, 31, 86 ORDER GADIFORMES GADIDAE — CODFISHES AND HAKES Lota lota (Linnaeus) — Burbot Lota maculosa (Lesueur) F&R. G. Rare to uncommon in widely separated large rivers and lakes statewide. Common in dark water streams of the Flambeau R. watershed and tributary streams to L. Superior; common in Lakes Poygan and Winnebago (L. Michigan drainage). Young occa¬ sionally taken in small streams opening into large bodies of water. Decreasing in L. Michigan in recent years. 9, 12, 13, 31, 65, 69 ORDER ATHERINIFORMES CYPRINODONTIDAE — KILLIFISHES Fundulus diaphanus (Lesueur) — Banded killifish Common in lakes of southeastern Wisconsin. Uncommon in widely isolated sites in northern half of state and absent from southwestern quarter. 9, 12, 46. Fundulus notatus (Rafinesque) — Blackstripe topminnow Uncommon in sloughs and lakes of the Mississippi drainage in the southeastern quarter of state. A single recent report from the Wisconsin R. at Woodman (Grant Co.). In L. Michigan drainage, present in the upper Fox R. (Columbia and Marquette Cos.). Re¬ cently collected from lower Wisconsin R. 13, 46 Fundulus notti (Agassiz) — Starhead topminnow Fundulus dispar (Agassiz) F&R. G. Rare. Recently collected from quiet water in Coon Creek (Rock Co.) and sloughs of the Wisconsin R. (Iowa Co.). The later is the first for the Wisconsin R. basin. Older records from Walworth and Waukesha Cos. (lower Fox R. system). 46 286 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 ATHERINIDAE — SILVERSIDES Labidesthes sicculus (Cope) — Brook silverside Common in lakes and quiet waters of rivers and large streams in southern half of the state. Distribution discontinuous northward where uncommon. Absent in L. Superior drainage. 10, 13, 20, 22 ORDER GASTEROSTEIFORMES GASTEROSTEIDAE — STICKLEBACKS Culaea inconstans ( Kirtland) —Brook stickleback Eucalia inconstans (Kirtland) F&R. G. Abundant in dense vegetation of small to medium-size streams throughout state. Although taken most frequently in clear water, this species may be found in highly turbid waters. 4, 7, 12, 13, 69 Pungitius pungitius (Linnaeus) — Ninespine stickleback Pygosteus pungitius (Linnaeus) F&R Common in shoal areas of L. Superior and uncommon in streams of its drainage basin. Rare in shoal areas of L. Michigan. 31, 46, 69 ORDER SCORPAENIFORMES COTTIDAE — SCULPINS Cottus bairdi Girard — Mottled sculpin Coitus ictalops (Rafinesque) F&R Uranidea kumlieni Hay F&R Common in cold headwater streams throughout the state. Occa¬ sionally in large lakes: L. Metonga (Forest Co.), L. Winnebago (Winnebago Co.), shoal areas of L. Michigan. 7, 9, 12, 13, 28, 69 Cottus cognatus Richardson — Slimy sculpin Uncommon to common in streams tributary to and in L. Superior. Uncommon in L. Michigan. Recently taken from Citron Creek (Crawford Co.), Camp Creek (Richland Co.) and Big Green L. (Green Lake Co.). 13, 28, 31 Cottus ricei (Nelson)— Spoonhead sculpin Uncommon in shallow to deep waters of Lakes Michigan (2 to 73 fathoms) and Superior (20 to 60 fathoms). 28, 31, 46 Myoxocephalus quadricornis (Linnaeus)— Fourhorn sculpin Triglopsis thompsonii Girard G. Common in deep water of Lakes Superior (40 to 200 fathoms) and Michigan (25 to 100 fathoms). 28, 31, 46 1970] Johnson and Becker — Fishes of Wisconsin 287 ORDER PERCIFORMES SERRANXDAE — SEA BASSES Roccus chrysops (Rafinesque) —White bass Lepibema chrysops (Rafinesque) G. Common in large lakes and rivers of southern half of the state and in the St. Croix upstream to St. Croix Falls Dam. Abundant in L. Winnebago. 9, 10, 13, 22, 117 Roccus mississippiensis (Jordan & Eigenmann) — Yellow bass Morone interrupta Gill F&R G. Uncommon to common in the Mississippi and lower Wisconsin Rivers; common and increasing in Lakes Poygan and Winnebago. Introduced into the Madison lakes (Dane Co.) and into lakes of the lower Wolf R. (L. Michigan basin) during fish transfer operations of the 1930's and 1940's. Abundant in L. Mason, Adams Co. Re¬ cently stocked in the Manitowoc, Sheboygan, and Milwaukee R. basins in eastern Wisconsin. 9, 13, 50, 51, 67, 99 CENTRARCHIDAE — SUNFISHES Ambloplites rupestris (Rafinesque)-— -Rock bass Common in clear, medium to large streams and in lakes through¬ out the state except in southwestern quarter where rare. 7, 9, 10, 22, 53, 65, 69 Chaenobryttus gulosus (Cuvier) — Warmouth Rare to uncommon in southern third of state; in sloughs of rivers and impoundments. Common in the Long Lake Chain (Chip¬ pewa Co.). Reported in L. Nebagamon (Douglas Co.). 13, 46 Lepomis cyanellus Rafinesque-— Green sunfish Apomotis cyanellus (Rafinesque) G. Common in lakes and medium-sized streams of southern one- third of state. Discontinuous distribution northward. Not reported from the L. Superior drainage. 10, 12, 13, 22, 60 Lepomis gibbosus (Linnaeus) — Pumpkinseed Eupomotis gibbosus (Linnaeus) F&R. G. Common in clear medium-sized rivers and in lakes throughout the state except in the southwestern quarter and in L. Superior drainage where uncommon. 7, 9, 12, 13, 22, 69 288 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Lepomis humilis (Girard) — Orangespotted sunfish Allotis humilis (Girard) G. Uncommon to common in Mississippi R. (Vernon Co. southward) and lower Wisconsin R. Rare to uncommon in sloughs and back¬ waters of larger streams in Richland, Iowa, Green, and Lafayette Cos. Recently collected from Sugar River (Dane and Green Cos.) and lower Fox R. (Kenosha Co.) . Lepomis macrochirus Rafinesque — Bluegill Lepomis pallidus (Mitchill) F&R Helioperca macrochirus (Rafinesque) G. Most abundant centrarchid in Wisconsin. Found in medium-sized streams to large rivers and in nearly all lakes throughout the state. 7, 9, 10, 12, 13, 22, 65, 69, 97 Lepomis megalotis (Rafinesque) — Longear sunfish Xenotis megalotis (Cope) G. Uncommon in southeastern quarter of state. Recently collected in the Milwaukee R. (Ozaukee Co.) and in the Mukwonago R. (Wau¬ kesha Co.). Reported rare from L. Geneva (Walworth Co.). 46 Micropterus dolomieui Lacepede — Smallmouth bass Common in medium to large streams and large clear-water lakes throughout the state. Common in upper Green Bay area of L. Michigan and Chequamegon Bay of L. Superior. 7, 9, 10, 12, 13, 65, 69, 70, 71, 114 Micropterus salmoides Lacepede — Largemouth bass Aplites salmoides (Lacepede) G. Abundant in medium to large rivers and in lakes throughout the state. 7, 9, 10, 12, 13, 22, 65, 69, 71, 75, 76 Pomoxis annularis Rafinesque — White crappie Common in the Mississippi and lower Wisconsin Rivers and their larger tributaries. Reported from L. Mendota (Dane Co.). Recently taken from the upper Fox and lower Wolf River basin in east-central Wisconsin (L. Michigan drainage). 9, 13, 22 Pomoxis nigromaculatus (Lesueur) — Black crappie Pomoxis sparoides (Lacepede) F&R. G. Common in lakes and larger rivers throughout the state. Intro¬ duced into many lakes in northern Wisconsin where it has become abundant. 7, 9, 10, 12, 13, 22, 65, 69 1970] Johnson and Becker — Fishes of Wisconsin 289 PERCIDAE — PERCHES Ammocrypta asprella (Jordan) — Crystal darter Cryst allaria asprella (Jordan) F&R. G. Rare in the Mississippi River. Collected recently on the lower Wisconsin R. near Orion (Richland Co.), on the lower Chippewa R. between Durand (Pepin Co.) and Meridian Ferry Landing (Dunn Co.). 18, 46 Ammocrypta clara Jordan & Meek — Western sand darter Ammocrypta pellucida (Baird) F&R. G. Common locally in shallow riffles over sand flats in the lower Wis¬ consin R., in the Mississippi R. and in the St. Croix upstream to the St. Croix Falls Dam. Recently collected in the Waupaca R. (Great Lakes drainage near its junction with the Wolf R. in Waupaca Co.) . 11, 13, 65 Etheostoma asprigene (Forbes) — Mud darter Etheostoma jessiae (Jordan & Brayton) F&R Poecilichthys jessiae Jordan & Brayton G. Rare to uncommon in sloughs of the lower Wisconsin and Missis¬ sippi Rivers and in the lower portions of tributaries to them. 13, 22 Etheostoma caeruleum Storer — Rainbow darter Poecilichthys coeruleus (Storer) G. Common locally in central and southeastern Wisconsin. Uncommon in southwestern Wisconsin. 6, 12, 13, 46 Etheostoma chlorosomum (Hay) — Bluntnose darter Boleosoma camurum Forbes F&R Rare. This southern darter has been collected as far north as the Root R., Houston Co., Minnesota. Records (Zoological Museum UW, Madison; University Museums UM, Ann Arbor, Michigan) from the Mississippi River come from small isolated ponds between New Albin and Minnesota slough on the Iowa-Minnesota border just across from Victory, Vernon Co., Wis. 33, 49 Etheostoma exile (Girard) — Iowa darter Poecilichthys exilis (Girard) G. Uncommon to common locally over the state but found primarily in glaciated areas, where it is taken in small streams and bog lakes. 7, 9, 12, 13, 22, 69 290 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Etheostoma flabellare Rafinesque — Fantail darter Catonotus flabellaris Agassiz G. Abundant over rocks and gravel in the smaller streams of the state. Occasionally taken in medium to large rivers. 7, 12, 13 Etheostoma microperca Jordan & Gilbert — Least darter Microperca punctulata Putnam F&R. G. Uncommon. Although found in widely separated areas over the state, it has been collected most frequently from the southeastern counties. 7, 46 Etheostoma nigrum Rafinesque — Johnny darter Boleosoma nigrum (Rafinesque) F&R. G. Abundant in all waters from the smallest stream and pond to the largest river and lake over a wide variety of bottom types. It is the most successful member of its family. 7, 9, 10, 12, 13, 22, 65, 69, 105 Etheostoma zonale (Cope) — Banded darter Poecilichthys zonalia Cope G. Common locally but of spotty distribution over the lower two- thirds of the state. Rare to uncommon in northeastern Wisconsin. In clear-water streams of medium to large size. 7, 12, 13 Perea fiavescens (Mitchill) — Yellow perch Abundant in lakes, ponds, impoundments and large rivers. Com¬ mon in Chequamegon Bay (L. Superior) and in L. Michigan. 5, 7, 9, 10, 12, 13, 22, 63, 65, 69, 98 Percina caprodes (Rafinesque) — Logperch Common in medium to large streams and rivers and in large lakes. Distribution statewide. 7, 9, 12, 13, 22, 65, 69 Percina evides (Jordan & Copeland)— -Gilt darter Hadropterus evides (Jordan & Copeland) F&R. G. Uncommon in the Black R. and in the St. Croix R. in the vicinity of St. Croix Falls. 46, 65 Percina maculata (Girard) — Blackside darter Hadropterus aspro (Cope & Jordan) F&R Hadropterus maculatus (Girard) G. Common in streams and rivers of all sizes in clear to turbid water. Distribution statewide. 7, 9, 12, 13, 69 1970] Johnson and Becker- — Fishes of Wisconsin 291 Percina phoxocephala (Nelson)— Slenderhead darter Hadropterus phoxocephalus (Nelson) F&R. G. Uncommon in the lower Wisconsin R., the Mississippi, and in the larger streams tributary to them. Also in the Pecatonica R. (La¬ fayette Co.) and Sugar R. (Green Co.). In Great Lakes drainage found in L. Winnebago and in waters of the Upper Fox R. 9, 13, 46 Percina shumardi ( Girard ) —River darter C otto g aster shumardi (Girard) F&R Imostoma shumardi (Girard) G. Uncommon in the lower Wisconsin and in the Mississippi Rivers. Recently collected in Lakes Winnebago,, Poygan, and the lower Waupaca R. (L. Michigan basin). 9, 13, 46 Stizostedion canadense (Smith)— Sauger Common in L. Winnebago and in the lower Wisconsin and Mis¬ sissippi Rivers. Uncommon to common in St. Croix R. up to St. Croix Dam. Elsewhere in state uncommon, rare or absent. 9, 13, 22, 65, 88 Stizostedion vitreum vitreum (Mitchill) — Walleye Common locally in large rivers and almost all of the large lakes in the state. Introduced in many large lakes, especially in the north. Common in Chequamegon Bay area of L. Superior. 7, 9, 10, 13, 22, 65, 69, 77, 88, 90 SCIAENIDAE — DRUMS Aplodinotus grunniens Rafinesque — Freshwater drum Common in the lower Wisconsin R., the Mississippi, the St. Croix upstream to St. Croix Falls Dam. Sporadic in some large lakes of southern one-half of state. Abundant in L. Winnebago. 9, 10, 13, 22, 65 Problematical Fishes The following list includes those fishes currently not found in Wisconsin waters but which may be expected in the future. Also listed are those species and hybrids planted by various agencies but which are not known to reproduce naturally. Salmonidae Oncorhynchus gorhuscha (Walbaum) — Pink salmon Accidentally introduced in 1955 into L. Superior from the Port Arthur, Ontario fish hatchery. Several successive generations have 292 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 been reported. Six adults observed in the Cross R. near L. Superior, Cook Co., Minnesota, on Sept. 20, 1963. Although reported to this date only from Ontario and Minnesota waters, on the basis of its past movements we may expect to find it in Wisconsin waters. 14, 69, 93, 95 Oncorhynchus kisutch (Walbaum) — Coho salmon Unsuccessfully introduced into L. Michigan and Riley L. (Chip¬ pewa Co.) in 1951. Planted in May, 1966, in State of Michigan tributaries of Lakes Michigan and Superior and apparently spread¬ ing throughout these lakes. Large numbers netted by commercial fishermen from L. Michigan off Michigan City, Indiana, during April and May, 1967. In 1968, 25,000 stocked in Ahnapee R. at Algoma. Stockings planned for 1969 include Algoma, Kewaunee, Sheboygan, and Manitowoc. 15, 27c, 104 Oncorhynchus tshawytscha (Walbaum) — Chinook salmon Introduced. Between 1876 and 1879 a number of plantings were made, among them L. Geneva (Walworth Co.), Devils L. (Sauk Co.), Wautoma L. (Waushara Co.), L. Mendota (Dane Co.), Wis¬ consin R. at Portage (Columbia Co.), tributaries to the Mississippi (Grant Co.) . In 1881 a mature female was taken from L. Michigan off Cedar Grove (Sheboygan Co.). Unsuccessfully stocked in Sun¬ set L. (Portage Co.) in 1951. The State of Michigan stocked this species in 1967 in streams tributary to Lakes Superior and Michi¬ gan. Merryll Bailey reports capture of a 5-lb. Chinook off Ashland (L. Superior) in May, 1969. A 1969 Wisconsin release of 60,000 is. anticipated for the Sturgeon Bay ship canal. 15, 33, 113 Salmo clarki Richardson — Cutthroat trout Introduced. In the Report of the Commissioners of Fisheries of Wisconsin for 1895-96 an entry is made of “450 black-spotted trout (full grown) distributed in 1896.” A later entry in the same report mentions 500 black-spotted trout having been planted in “Pike’s Creek,” with no further data. Stocked in Black Earth Cr. (Dane Co.) in 1942 and in Gould Stream (Walworth Co.) in 1943. In 1959 the Twenty-Five Sportsmen’s Club of Hubertus planted 500 legal cutthroat trout in Friess L. (Washington Co.). 113 Salmo salar Linnaeus — Landlocked salmon Introduced. In 1875 landlocked salmon were stocked in L. Men¬ dota (Dane Co.), Oconomowoc L. (Waukesha Co.), and Devils L. (Sauk Co.). In 1876, 10,000 were planted in L. Geneva (Walworth Co.). In 1879, plantings were made in “Clear L., Silver L., Geneva 1970] Johnson and Becker — Fishes of Wisconsin 293 L., Nagawicka L., and Green R.” In 1907„ 10,000 were planted in Trout L. (Vilas Co.) . Two purported specimens of this species were taken Nov. 12, 1937, from a stream entering L. Geneva. Conserva¬ tion personnel who examined the fish in 1939 believe them to be misidentified brown trout (letter Jan. 6, 1939, from L. A. Wood¬ bury to F. C. Hewitt on file with the Wis. Cons. Dept.). 84, 113 Salvelinus namaycush female X Salvelinus fontinalis male — Splake This hybrid was produced in Wisconsin as early as 1884. It has been stocked experimentally in lakes of northern Wisconsin in recent years. According to Canadian workers some backcrossing with the lake trout occurs. 15, 113 Thymallus arcticus (Pallas) — Arctic grayling Introduced. Reports of the Commissioners of Fisheries of Wis¬ consin from 1878 to 1881 refer to grayling held in the Madison Hatchery, but no stocking occurred. In 1902, 180,000 fry were dis¬ tributed from the Bayfield Hatchery with no locales indicated. In 1906, 30,000 eggs or fry were distributed at Lake Nebagamon to N. Clay Pierce, who had an estate on the Brule R. (Douglas Co.) . These may have been released in the Brule. In 1908, 50,000 fry were planted in the Namekagon R. at Cable (Bayfield Co.). Unsuc¬ cessfully introduced in Mosquito Brook (Sawyer Co.) in 1937 and Pine R. (Waushara Co.) in 1938. 113 Cyprinidae S cardinius erythrophthalmus ( Linnaeus ) — Rudd Introduced into Oconomowoc L. (Waukesha Co.) in 1917 by Wisconsin Conservation Department. Reported to have spawned successfully in the lake in 1918, but has not been seen since. 20 Notropis lutrensis (Baird and Girard) — Red shiner Recent records from Menominee Cr., 3 mi. ENE of East Dubuque (Jo Daviess Co.), and from Winnebago Co. in northern Illinois place this minnow almost on the Wisconsin state line (letter from Philip Smith, June 6, 1967). Semotilus corporalis (Mitchill) — Fallfish Appearance of the fallfish in the Cedar Cr., Thunder Bay District of Ontario, Canada is explained through the use of northern Lake Superior as a migration route. Not yet reported from Wisconsin waters. 1 294 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 ICTALURIDAE No turns insignis (Richardson) — Margined madtom Introduced population in Sylvania Tract, Upper Peninsula, Mich¬ igan. Since this population is only a few miles from Wisconsin, it may possibly spread into our waters (letter from Reeve Bailey, April 4, 1967) . Percidae Etheostoma spectabile (Agassiz) — Orangethroat darter The Illinois Natural History Survey has records of this species from Lake and McHenry Cos. which are adjacent to the Wisconsin state line (letter from Philip Smith, June 6, 1967). Etheostoma blennoides Rafinesque — Greenside darter A recent record needing substantiation from Lake Co., Illinois (L. Michigan drainage), places this species next to the Wisconsin state line. 95 References 1. Allin, A. E. 1951. The Fallfish, Semotilus corporalis, from the Lake Superior drainage of Western Ontario. Copeia 1951(4) :300. 2. Bailey, Reeve. 1959. Distribution of the American cyprinid fish Notropis anogenus. Copeia 2:119-123. 3. Bailey, Reeve et al. 1960. A list of common and scientific names of fishes from the United States and Canada. Amer. Fish. Soc. Spec. Publ. No. 2. 102 pp. 4. Bailey, Reeve and M. O. Allum. 1962. Fishes of South Dakota. Univ. of Mich. Misc. Publ. Mus. of Zoology. No. 119. 131 pp. 5. Bardach, John E. 1949. Contribution to the ecology of the yellow perch (Perea flavescens Mitchill) in Lake Mendota, Wisconsin. Ph.D. Thesis, Dept. Zoology, Univ. of Wis., Madison, Wis. 6. Becker, George C. 1952. Preliminary survey of Rocky Run Creek, Colum¬ bia Co., Wisconsin. Unpub. Manuscript. Wisconsin Cons. Dept., Madison, Wisconsin. 59 pp. 7. - . 1959. Distributio'n of central Wisconsin fishes. Trans. Wis. Acad. Sci. Arts & Lett. 48:65-102. 8. - . 1962. Intraspecific variation in Rliinichthys c. cataractae (Val¬ enciennes) and Rhinichthys atratulus meleagris (Agassiz) and ana¬ tomical and ecological studies of Rhinichthys c. cataractae. Ph.D. Thesis. University of Wisconsin, Madison, Wisconsin. 279 pp. 9. - . 1964. The fishes of Lakes Poygan and Winnebago. Trans. Wis. Acad. Sci. Arts & Lett. 53:29-52. 10. - . 1964. The fishes of Pewaukee Lake. Trans. Wis. Acad. Sci. Arts & Lett. 53:19-27. IT; - — ^ — . 1965. The western sand darter. Ammocrypta clara, in the Great Lakes Basin. Copeia 1965 (2) :241. 12. - -. 1966. Fishes of northeastern Wisconsin. Unpubl. data. Dept, of Biology, Wis. State Univ., Stevens Point. 1970] Johnson and Becker — Fishes of Wisconsin 295 13. - . 1966. Fishes of southwestern Wisconsin. Trans. Wis. Acad. Sci. Arts & Lett. 55:87-117. 14. Black, John and Lyman O. Williamson. 1947. Artificial hybrids between muskellunge and northern pike. Trans. Wis. Acad. Sci. Arts & Lett. 38:299-314. 15. Borgeson, David P. and Wayne H. Tody. 1967. Status report on Great Lakes fisheries. Fish Div., Mich. Dept, of Cons., Lansing. Fish Man. Rep. #2:35. 16. Brasch, John. 1950. The lampreys of Wisconsin. Wis. Cons. Bull. 15(9) : 13-16. 17. Brasch, John, James McFadden and Stanley Kmiotek. 1958. The eastern brook trout, its life history, ecology and management. Wis. Cons. Dept. Publ. 226. 11 pp. 18. Brynildson, Oscar M., Vernon A. Hacker and Thomas A. Klick. 1963. Brown trout, its life history, ecology and management. Wis. Cons. Dept. Publ. 254. 15 pp. 19. Burdick, Milton E. and E. L. Cooper. 1956. Growth rate, survival, and harvest of fingerling rainbow trout planted in Weber Lake, Wis. Jour. Wild. Man. 20:233-38. 20. Cahn, Alvin R. 1927. An ecological study of southern Wisconsin fishes. The brook silverside (Labidesthes sicculus ) and the cisco ( Leucichthys artedi) in their relations to the region. Ill. Biol. Monogr., 11(1) :151 pp. 21. Carlander, Harriet Bell. 1954. A history of fish and fishing in the upper Mississippi River. Upper Miss. River Conservation Committee. 96 pp. 22. Christenson, Lyle M. and Lloyd L. Smith. 1965. Characteristics of fish populations in upper Mississippi River backwater areas. U. S. Dept. Interior. Fish & Wild. Serv. Circ. 212. 53 pp. 23. Coker, Robert E. 1930. Studies of common fishes of the Mississippi River at Keokuk. U. S. Govt. Print. Off. Bur. of Fish. Docu. No. 1072:141-225. 24. Cooper, Edwin L. 1956. What’s Happened to the Cisco? Wis. Cons. Bull. 21(3). 4 pp. 25. Churchill, Warren S. 1945. The brook lamprey in the Brule River. Trans. Wis. Acad. Sci, 37:337-46. 26. Creaser, C. W. 1925. The establishment of the Atlantic smelt in the upper waters of the Great Lakes. Mich. Acad. Sci. Arts & Lett. 5:405-424. 27a. Daly, Russell, Vernon A. Hacker and Lawrence Wiegert. 1962. The lake trout, its life history, ecology and management. Wis. Cons, Dept. Publ. No. 233. 15 pp. 27b. Daly, Russell. 1968. Preliminary notes on management of the rainbow trout in western Lake Michigan. Wis. Dept. Nat. Resources. Oshkosh. 4 p. mimeo. 27c. Daly, Russell. 1968, Progress report of fish management on Lake Michigan. Wis. Dept. Nat. Resources. Oshkosh. 18 p. mimeo. 28. Deason, Hilary J. 1939. The distribution of cottid fishes in Lake Michi¬ gan. Pap. Mich. Acad. Sci. Arts & Lett. 24:105-115. 29. Druschba, Leonard. 1959. Are the carp moving north? Wis. Cons. Bull. 24(11) :4 pp. 30. Dryer, William. 1963. Age and growth of the whitefish in Lake Superior. U.S. Fish and Wildlife Service, Fishery Bulletin, 63(1) : 77-95. 31. Dryer, William R. 1966. Bathymetric distribution of fish in the Apostle Islands region. Lake Superior. Trans. Amer. Fish. Soc. 95 (3) : 248-259. 296 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 82. Duck, Harvey. 1967. Salmon spawn, new hope for Great Lakes fishing. National wildlife. 5(3): 18-19. 33. Eddy, Samuel and Thaddeus Surber. 1947. Northern fishes. Univ. of Minn. Press. 276 pp. 34. Edsall, Thomas. 1960. Age and growth of the whitefish, Coregonus clupeaformis, of Munising Bay, Lake Superior. Trans. Amer. Fish Soc. 89(4) : 323-332. 35. Eschmeyer, Paul H. 1955. The reproduction of lake trout in southern Lake Superior. Trans. Amer. Fish. Soc, 84:47-74. 36. Eschmeyer, Paul H. 1956. The early life history of the lake trout in Lake Superior. Mich. Dept. Cons. Inst, for Fish. Pec. Misc. Pub. 10, 21 pp. 37. Eschmeyer, P. H. and R. M. Bailey. 1954. The pygmy whitefish, Core¬ gonus coulteri, in Lake Superior. Trans. Amer. Fish. Soc. 84:161-199. 38. Fernholz, Willis. 1966. Mississippi River commercial fishing statistics — 1965. Wis. Cons. Dept. Fish. Man. Div. Stat. Rep. 31 pp. (Mimeo.) 39. Fukano, K. G., H. Gowing, M. J. Hansen, and L. N. Allison. Introduc¬ tion of exotic fish into Michigan. Institute for Fisheries Research, Univ. of Mich. An'n Arbor, Rep. No. 1689. 50 pp. 40. Forbes, Stephen A. and Robert E. Richardson. 1920. The fishes of Illinois. Ill. Nat. Surv. Bull. 3:357 pp. 41. Frey, David G. 1940. Growth and ecology of the carp Cyprinus carpio Linnaeus in four lakes of the Madison region. Wisconsin. Ph.D. Thesis, Dept. Zoology, Univ. of Wis., Madison, Wis. 249 pp. 42. Frankenberger, Ludwig. 1957. Whitefish and Ciscos — Forgotten Fish. Wis. Cons. Bull. 22(12) 3 pp. 43. Gammon, James R. 1961. Contributions to the biology of the muskellunge. Ph.D. Thesis, Dept. Zoology, Univ. of Wis., Madison, Wis. 144 pp. 44. Gilbert, Carter R. 1961. Hybridization versus integradation : an inquiry into the relationship of two cyprinid fishes. Copeia 1961(2) : 181-192. 45. Greene, C. W. 1927. An ichthyological survey of Wisconsin. Mich. Acad. Sci. Arts & Lett. 7:299-310. 46. Greene, C. Willard. 1935. Distribution of Wisconsin Fishes. Wis. Cons. Comm., Madison, Wis. 235 pp. 47. Greenwood, R. H., D. E. Rosen, S. H. Weitzman, and G. C. Meyers. 1966. Phyletic studies of teleostean fishes, with a provisional classifica¬ tion of living forms. Bull. Amer. Mus. Nat. Hist. 131(4) : 344-451. 48. Hacker, Vernon A. 1956. Biology and management of lake trout in Green Lake, Wisconsin. Trans. Amer. Fish. Soc. 86:13 pp. 49. Harlan, James R. and Everett B. Speaker. 1956. Iowa fish and fishing. State Cons. Comm. Des Moines, Iowa. 377 pp. 50. Helm, William T. 1958. A “new” fish in Wisconsin. Wis. Cons. Bull. 23(7) : 10-12. 51. Helm, William T. 1964. Yellow bass in Wisconsin. Trans. Wis. Acad. Sci. Arts & Lett. 53:109-125. 52. Herman, Elmer et al. 1959. The yellow perch, its life history, ecology and management. Wis. Cons. Dept. Publ. No. 228. 14 pp. 53. Hile, Ralph. 1936. Age and growth of the cisco Leucichthys artedi (Le Sueur), in the lakes of the northeastern highlands, Wisconsin. Bull, of the U. S. Bureau of Fisheries. 48(19) :211-317. 54. Hile, Ralph. 1941. Age and growth of the rock bass, Amhloplites rupestris (Rafinesque) , in Nebish Lake, Wisconsin. Trans. Wis. Acad. Sci., Arts & Lett. 33:189-337. 1970] Johnson and Becker — Fishes of Wisconsin 297 55. Horrall, R. M. 1961. A comparative study of two spawning populations of the white bass Roccus chrysops ( Raf . ) , in Lake Mendota, Wisconsin, with special reference to homing behavior. Ph.D. Thesis, Dept. Zoology, U'niv. of Wis., Madison, Wisconsin. 56. Hoy, P. R. 1883. Catalogue of the cold-blooded vertebrates of Wisconsin. Rep. Geol. Surv. Wis. 1:427-436. 57. Hubbs, Carl L. and C, Willard Greene. 1935. Two new subspecies of fishes from Wisconsin. Trans. Wis. Acad. Sci. Arts & Lett. 20:89-101. 58. Hubbs, Carl L. and Karl F. Lagler. 1964. Fishes of the Great Lakes region (Revised edit.) Cranbrook Instit. of Sci. Bull. 26:213 pp. 59. Hunt, Robert L. 1966. Production and angler harvest of wild brook trout in Lawrence Creek, Wisconsin. Wis. Cons. Dept., Madison. Tech. Bull. 35, 51 pp. 60. Hunter, John R. and Arthur D. Hasler. 1965. Spawning association of the redfin shiner, Notropis umbratilis, and the green sunfish, Lepomis cyanellus . Copeia 1965 (3) : 265-281. 61. Huver, Charles W. 1961. Variation and speciation in coregonid fishes. M. S. Thesis, Dept. Zoology, U'niv. of Wis., Madison, Wis. 27 pp. 62. Jobes, F. W. 1949. The age, growth and bathymetric distribution of the bloater Leucichthys hoyi (Gill) in Lake Michigan. Pap. Mich. Acad. Sci. Arts & Lett. 33(1947) : 135-172. 63. John, Kenneth. 1954. An ecological study of the cisco Leucichthys artedi (Lesueur) in Lake Mendota, Wis. Ph.D. Thesis. Univ. of Wis., Madison. 121 pp. 64. Kleinert, Stanton J. and Donald Mraz. 1966. Life history of the grass pickerel (Esox americanus vermiculatus) in southeastern Wiscon¬ sin. Wis. Cons. Dept. Tech. Bull. 37, 40 pp. 65. Kuehn, Jerome H., Wallace Niemuth and Arthur R. Petersen. 1961. A biological reconnaissance of the Upper St. Croix River. Joint Survey Minnesota Conservation Dept. (St. Paul, Minn.) and Wisconsin Con¬ servation Dept. (Madison, Wis.). 25 pp & 21 appendices.. 66. McCutchin, Thayre N. 1946. A biological survey of Pewaukee Lake, Wau¬ kesha Co., Wisconsin. Wis. Cons. Dept. Madison. Sect, of Fish. Biol. Invest. Rep. No. 573. 28 pp. (Mimeo.) 67. McNaught, Donald C. 1963. The fishes of Lake Mendota. Trans. Wis. Acad. Sci. Arts & Lett. 52:37-55. 68. Miller, Robert R. 1957. Origin and dispersal of the alewife, Alosa pseudoharengus and D or o soma cepedianum in the Great Lakes. Trans. Amer. Fish. Soc. 86:97-111. 69. Moore, Harry H. and Robert A. Bream. 1965. Distribution of fishes in U.S. streams tributary to Lake Superior. U. S. Fish & Wildl. Serv., Wash., D. C. Spec. Sci. Rep. Fisheries No. 516. 61 pp.. 70. Mraz, Donald. 1960. Prelim, report on the Lake Geneva smallmouth bass studies (1958-1959). Wis. Cons. Dept., Fish. Man. Div., Res. Report No, 1, 22 pp. 71. Mraz, Donald. 1964. Observations on large and smallmouth bass nesting and early life history. Wis. Cons. Dept., Res. Report No. 11, 13 pp. 72. Mraz, Donald. 1964. Age, growth, sex ratio and maturity of the whitefish in Central Green Bay and adjacent waters of Lake Michigan. U. S. Fish & Wildlife Serv. Fish. Bull. 63(3) : 619-634. 73. Mraz, Donald. 1964. Age and growth of the round whitefish in Lake Michigan. Tra'ns. Amer. Fish. Soc. 93(1) : 46-52. 74. Mraz, Donald and Edwin L. Cooper. 1957. Natural reproduction and survival of carp in small ponds. Jour. Wild. Man. 21:66-69. 298 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 75. Mraz, Donald, Stanley Kmiotek and Ludwig Frankenberger. 1961. The largemouth bass, its life history, ecology and management. Wis. Cons. Dept. Publ. 232. 13 pp. 76. Mraz, Donald and C. W. Threinen. 1955. Angler’s harvest, growth rate and population estimate of the largemouth bass of Brown’s Lake, Wis¬ consin. Trans. Amer. Fish. Soc. 85:241-256. 77. Niemuth, Wallace, Warren Churchill and Thomas Wirth. 1959. The walleye, its life history, ecology and management. Wis. Cons. Dept. Publ. 227. 14 pp. 78. Noland, W. E. 1951. The hydrography, fish, and turtle population of Lake Wingra. Trans. Wis. Acad. Sci. Arts & Lett. 40:5-58. 79. Norden, Carroll R. 1967. Age, growth and fecundity of the alewife, Alosa pseudoharengus (Wilson), in Lake Michigan. Trans. Amer. Fish. Soc. 96(4) : 387-393. 80. Nybakken, James W. 1961. Analysis of the sympatric occurrence of two subspecies of the cyprinid fish Campostoma anomalum (Rafinesque) in Wisconsin. M. S. Thesis. Univ. of Wis., Madison, Wis. 35 pp. 81. Oehmcke, Arthur A., Leon Johnson, John Klingbeil and Clarence Wistrom. 1965. The Wisconsin Muskellunge, its life history, ecology and management. Wis. Cons. Dept. Publ. 225. 12 pp. 82a. O’Donnell, D. J. and W. S. Churchill. 1954. Certain physical, chem¬ ical, and biological aspects of the Brule River, Douglas County, Wis¬ consin. Trans. Wis. Acad. Sci. Arts & Lett. 43:201-255. 82b. Phillips, Gary L. and James C. Underhill. 1967. Revised distribution records of some Minnesota fishes, with addition of two species to the faunal list. Jour. Minn. Acad. Sci. 34(2) : 177-180. 83. Poff, Ronald J. and C. W. Threinen. 1965. Surface water resources of Columbia County, Wis. Cons. Dept. Lake & Stream Classification Project. 55 pp. 84. Pope, T. E. B. 1938. Landlocked salmon in Wisconsin. Trans. Wis. Acad. Sci. Arts & Lett. 31:559-565. 85. Priegel, Gordon R. 1962. Winnebago winter menu. Wis. Cons. Bull. 27(1) : 20-21. 86. Priegel, Gordon R. 1962. Plentiful but unknown. Wis. Cons. Bull. 27(3) : 13. 87. Priegel, Gordon R. 1963. Dispersal of the shortnose gar, Lepisosteus platostomus, i'nto the Great Lakes drainage. Trans. Amer. Fish. Soc. 92(2) :178. 88. Priegel, Gordon R. 1963. Food of walleye and sauger in Lake Winnebago, Wisconsin. Trans. Amer. Fish. Soc. 92 (3) :312-313. 89. Priegel, Gordon R. 1966. Age-length and length-weight relationship of bullheads from Little Lake Butte des Morts, 1959. Wis. Cons. Dept. Res. & Planning Div. Res. Rep. 17:6 pp. 90. Pycha, Richard L. 1961. Recent changes in the walleye fishery of north¬ ern Green Bay and history of the 1943 year class. Trans. Amer. Fish. Soc. 90(4) : 475-88. 91. Roeloffs, Eugene W. 1958. Age and growth of whitefish, Coregonus clupeaformis (Mitchill), in Big Bay de Noc and northern Lake Michi¬ gan. Trans. Amer. Fish. Soc. 87:190-199. 92. Schneberger, Edward and Lowell A. Woodbury. 1944. The lake sturgeon, Acipensor fulvescens Rafinesque, in Lake Winnebago. Wis. Trans. Wis. Acad. Sci. 36:131-140. 1970] Johnson and Becker — Fishes of Wisconsin 299 93. Schumacher, E. E. and S. Eddy. 1960. The appearance of pink salmon, Oncorhynchus gorbusclia (Walbaum), in Lake Superior. Trans. Amer. Fish. Soc. 8:371-373. 94. Schumacher, Robert E. and John G. Hale. 1962. Third generation pink salmon, Oncorhynchus gorbusclia (Walbaum), in Lake Superior. Trans. Amer. Fish. Soc. 91(4) :421-422. 95. Smith, Philip W. 1965. A preliminary annotated list of the lampreys and fishes of Illinois. Ill. Nat. Hist. Surv. Biol. Notes. No. 54. 96. Smith, Stanford H. 1964. Status of the deepwater cisco population of Lake Michigan. Trans. Amer. Fish. Soc. 92 (2) : 155-163. 97. Snow, Howard, Arthur Ensign and John Klingbeil. 1960. The blue- gill, its life history, ecology and management. Wis. Cons. Dept. Publ. 230. 14 pp. 98. Stone, Roderick C. 1961. Preliminary investigations of winter activity and movements of the yellow perch Perea flavescens (Mitchill) in Lake Mendota, Wisconsin. M.S. Thesis. Dept, of Zoology, Univ. of Wis., Madison, Wisconsin. 78 pp. 99. Tibbles, J. J. G. 1956. A study of movements and depth distribution of the pelagic fishes in Lake Mendota. Ph.D. Thesis, U'niv. of Wis., Madi¬ son, Wisconsin. 193 pp. 100. T'hreinen, C. W. 195-. Life history, ecology, and management of the alewife. Wis. Cons. Dept. Publ. 223. 8 pp. 101. Threinen, C. W. (Co-editor). 1961 to present. Surface water resources publications (a continuing series of each Wis. Co.). Wis. Cons. Dept. Lake & Stream Classification Project. 102. Threinen, C. W. and Arthur Oehmcke. 1950. The northern invades the musky’s domain. Wis. Cons. Bull. 15(9) : 10-12. 103. Threinen, C. W., C. Wistrom, B. Apelgren, and H. Snow. 1966. The northern pike, its life history, ecology, and management. Wis. Cons. Dept. Publ. 235. 16 pp. 104. Tody, Wayne H. and Howard A. Tanner. 1966. Coho Salmon for the Great Lakes. Mich. Dept, of Cons. Fish Div. Fish Management. Report No. 1, 38 pp. 105. Underhill, James C. 1963. Distribution in Minnesota of the subspecies of the percid fish Etheostoma nigrum , and of their intergrades. Amer. Wildl. Nat. 70(2) : 470-478. 106. Van Oosten, John. 1936. The dispersal of smelt Osmerus mordax (Mitch¬ ill) in the Great Lakes region. Trans. Amer. Fish. Soc. 66:160-171. 107. Van Oosten, J., Ralph Hile and Frank Jobes. 1946. The whitefish fishery of Lakes Huron and Michigan with special reference to the deep-trap-net fishery. U.S. Fish & Wildlife Serv. Fishery Bull. 40. 50:297-394. 108. Wagner, G. 1908. Notes on the fish fauna of Lake Pepin. Trans. Wis. Sci. Arts & Lett. 16:230-237. 109. Weed, A. C. 1921. Fishes from southern Wisconsin. Copeia (1921) (99) :69. 110. Wells, LaRue and Alfred M. Beeton. 1963. Food of the bloater, Coregonus hoyi in Lake Michigan. Trans. Amer. Fish. Soc. 92(3) :245- 255. 111. Wells, LaRue. 1966. Seasonal and depth distribution of larval bloaters (Coregonus hoyi) in southeastern Lake Michigan. Trans. Amer. Fish. Soc. 95(4) : 388-396. 112. Whitley, G. P. 1950. New fish names. Proc. Royal Zool. Soc. New S. Wales, 1948-49:44. 300 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 113. Wisconsin Commissioners of Fisheries. 1875-1912. Annual and Biennial Reports of the Commissioners of Fisheries of Wisconsin. Madison, Wisconsin. 114. Wis. Cons. Dept. 1958. Smallmouth bass streams in Wisconsin. 115. Wis. Cons. Dept. 1964. Wisconsin Lakes. Wis. Cons. Dept. Publ. 218-64. 67 pp. 116. Wis. Cons. Dept. 1966. Wisconsin trout streams. Wis. Cons. Dept. Publ. 213-66. 75 pp. 117. Wright, T. D. 1964. Aspects of the early life-history of the white bass (Roccus chrysops). M. S. Thesis, Univ. of Wis., Madison, Wisconsin. PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 60 TILIACEAE AND MALVACEAE— BASSWOOD AND MALLOW FAMILIES Fred H. Utech The woody, mostly tropical Tiliaceae has only one native and two cultivated species in Wisconsin, while the Malvaceae, characterized by monadelphous stamens, is represented not only by several noxi¬ ous Eurasian weeds as velvet-leaf (Ahutilon theophrasti) , flower- of-an hour (Hibiscus trionum) and cheeses (Malva neglecta and M. rotundifolia) and showy exotics as hollyhock (Alcea rosea) and high mallow (Malva sylvestris) , but also by several rare, handsome natives : poppy mallow (Callirhoe triangulata) , rose mallow (Hibiscus militaris ) and glade mallow (Napaea dioica). The present treatment is a revision of an earlier preliminary report on Tiliaceae and Malvaceae by Hagen (1932) and is based on specimens in the herbaria of the University of Wisconsin (WIS), University of Wisconsin-Milwaukee (UWM), Milwaukee Public Museum (MIL), University of Minnesota (MIN), Uni¬ versity of Minnesota-Duluth (DUL), State University of Iowa (IA), Oshkosh State University, La Crosse State University, Northland College (Ashland, Wis.), Beloit College and the private herbarium of Katherine Rill (Clintonville, Wis. — RILL). Grateful acknowledgement is due the curators of these herbaria for the loans of specimens. Map dots represent exact locations, triangles, county records. Some locations have been added from Thomas Hartley’s unpub¬ lished “Flora of the Driftless Area” (1962), Paul Sorensen’s un¬ published range maps from his Glacial Lake Wisconsin studies (1966), Jones and Fuller’s Flora of Illinois (1955) and stand records for Tilia americana from the UW Plant Ecology Laboratory The map inset numbers record Wisconsin flowering and fruiting dates. Plants with vegetative growth only, in bud or with dispersed fruit were not included. For introduced species the year of earliest collection within a county is also recorded. Nomenclature and order of genera and species follows that of Gleason and Cronquist (1963) and Fernald (1950). Field work and preparation of manuscript was supported in part by the Research Committee of the University of Wisconsin, on funds from the Wisconsin Alumni Research Foundation. 301 302 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Special thanks are due the author's major professor, Dr. Hugh H. litis, for his uncanny ability to inspire and stimulate botanical awareness and observation. Numerous people deserve special acknowledgement for their assistance in manuscript preparation: Mrs. Katherine Snell, for encouragement and timely aids; Mr. Brian G. Marcks and Mr. Michael H. Nee, for their advice, com¬ ment and criticism; Mr. Eugene G. Coffman, for photographic reproductions; Miss Bethia Brehmer, for artistic work; and Drs. H. H. litis and John W. Thomson for critical manuscript reading. TILIACEAE A. L. De Jussieu Basswood Family 1. TILIA [Tourn.] L. Basswood, Linden, Lime, Bee-tree. Large deciduous trees with soft white wood, fibrous inner bark, and numerous deep lateral roots. Winter buds large, obtuse, with few scales, the terminal hud lacking. Leaves alternate (2-ranked) , cordate, palmately veined, serrate, oblique to truncate, acute to acuminate; stipules deciduous. Flowers fragrant, entomophilous, yellowish-white, in cyme-like clusters each on an axillary peduncle adnate to the middle of a short-petioled, thin, wing-like bract. Flowers perfect, 5-merous, hypogynous. Stamens many, free or united into 5 bundles and then opposite petals ; filaments often forked distally; anthers 2-celled, opening longitudinally. Stami- nodes 5, petaloid , opposite petals, always present in native species (and in some Eurasian cultivars). Drupes globular, dry and woody, indehiscent, 1-locular, 1- to 3-seeded ; seeds with endosperm ; cotyledons broad, 3- to 5-lobed. About 25 tree species, native to north temperate deciduous forests, from Japan and Siberia to Europe, eastern N. America and Mexico. The chromosome number of Tilia (x = 41) is the highest odd basic number known in higher plants (Derman 1932). Tilia as a generic name is of special botanical interest, for it was from these noble trees that Carolus Linnaeus (Linne) ac¬ quired his surname. His ancestors were peasants. Several of his relatives, who had quitted the plough for the Muses, changed their family name with their profession and borrowed the name of Lindelius or Tiliander (Linden-tree-man) . . . , a custom not unfrequent in Sweden, to take fresh appellations from natural objects. The father of Linnaeus, as the first learned man of his family, could not withstand following the example which his kindred had set before him. He likewise bor¬ rowed of the same tree a name which his son rendered afterwards famous and immortal in 'every quarter of the guobe. (Stoever 1794) In North America, Tilia is called basswood in forestry, linden in horticulture, while in Britain, lime, apparently an altered form of linden or lind. The Anglo-Saxon lind means shield, i.e., of 1970] Utech — Flora of Wisconsin No. 60 303 linden wood. The white, light, diffuse-porous wood is used for do¬ mestic utensils. Leaves, both fresh and dried, were fed to cattle by the Romans, a contemporary custom in northern Europe. The flowers produce excellent honey, hence the local American “bee- tree”, while the dried steeped flowers serve as linden tea. Ropes, mats, shoes and baskets are made from the tough, fibrous inner bark, morphologically phloem bast fibers or bass, hence basswood. Leaf pubescence on flowering branchlets usually differs from that on sprout leaves. Most Tilia lower leaf surfaces have small tufts of simple trichomes in the principal vein axils ; these barbulae, domatia or acaradomatia are supposed to be symbiotic adaptations for mites (Jones 1968). Key to Species A. Leaves of flowering branchlets glabrous beneath, except for tufts of simple trichomes in lateral vein axils; fruit ribs ob¬ scure or lacking. B. Staminodes 5 ; stamens 45-60 ; axillary tufts whitish-gray, lacking at petiolar leaf insert; cymes 6- to 15-flowered ; common native forest tree _ 1. T. AMERICANA. BB. Staminodes lacking; stamens 25-30; axillary tufts rusty brown, well developed at petiolar leaf insert; cymes 5- to 7- (11-) flowered; commonly cultivated European tree __ _ 2. T. CORDATA. A A. Leaves of flowering branchlets pubescent beneath and often above; axillary tufts white; fruit strongly 3- to 5-ribbed; stamens 30-40; staminodes lacking; rarely cultivated Eur¬ opean tree _ 3. T. PLATYPHYLLOS. 1. Tilia Americana L. American Basswood, Linden. Map 1. Tilia glabra Vent. Trees to 35 m tall, 1-1.5 m DBH, often clustered with two or more trunks, often with several to many basal sprouts. Bark on old trunks firm, dark gray with longitudinal furrows, young bark smooth, light gray. Winter buds shiny, dark red, ellipsoid-ovate, 2-ranked; terminal bud absent. Leaves broadly ovate-cordate, the blades of fertile shoots 8-20 cm long, 6-16 cm wide, cordate to truncate at base, abruptly acuminate, coursely serrate with gland- tipped teeth, glabrous beneath, with tufts of whitish-gray hairs in vein axils, these lacking at petiolar leaf insert. Petioles, pedicels and bracts of inflorescence glabrous. Cymes 6- to 15-flowered. Petals 5, yellowish-white, oblong to oblanceolate, 6-9 mm long. Staminodes 5, oblanceolate, 5-7 mm long, opposite petals. Stamens 304 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 1970] Utech — Flora of Wisconsin No. 60 305 45-60, free. Floral bracts 7-10 cm long, short stalked or sessile. Drupes globose, 6-12 mm wide, thick-shelled, without ribs. 2N = 82 (Derman 1932). Native to deciduous forests of eastern North America (New Brunswick to southern Manitoba, southward to Penna. and Okla¬ homa), in its southeastern range partially sympatric along the glacial maxima with the Appalachian centered Tilia heterophylla Vent. (cf. maps Fowells 1965, Jones 1968), whence, Braun (1960) suggests, on vegetative characters, the putative “hybrid swarm” origin of Tilia neglecta Spach. However, according to Jones (1968) most specimens labelled T. neglecta consist of sprout leaves. The modern distributions suggest survival of T. americana in the Ozarks, T. heterophylla in the Appalachians, and post-glacial mi¬ gration, overlap, and hybridization, a clinal pattern similar to that found between Acer saccharum ssp. nigrum and A. s. ssp. saccha- rum (Desmarais 1948, 1952), with Tilia the main deciduous forest co-dominant. Flowering from mid-May to late July, fruiting from early June to mid- August. 2. Tilia cordata Mill. Small-leaved Linden. Cultivated trees to 25 m tall. Leaves orbicular-cordate, 2-7 (-10) cm wide, cordate, abruptly acuminate, the margin sharply serrate, dark green and glaucous above, blue-green and glabrous beneath, the axillary hair tufts rusty brown. Petioles glabrous, 1.4-3 cm long. Cymes pendulent or upright, 5- to 7- (11-) flowered. Stamens equaling petals, ca. 25-30, the filaments connate basally into 5 bundles alternating with petals ; staminodes usually lacking. Floral bracts 3-8 cm long, glabrous. Drupes 1-seeded, globose- apiculate, thin- walled, slightly or not ribbed. 2N = 82 (Derman 1932). Native from Siberia to England, Spain, Italy and the Balkans, commonly planted as lawn, street or park trees in many southern Wisconsin cities. Flowering in late June to early July. 3. Tilia platyphyllos Scop. Large-leaved Linden. Cultivated trees to 30 m tall. Leaves orbicular-ovate, 6-12 cm wide, obliquely cordate, abruptly acuminate, serrate, pubescent beneath and often above; axillary tufts white. Petioles densely pubescent, 2-5 cm long. Cymes pendulent, S- (rarely J- to 6-) flowered. Floral bracts 6-12 cm long, pubescent on lower midrib. Drupes 1-seeded, subglobose to pyriform, apiculate, densely pubes¬ cent, strongly 3- to 5-ribbed. 2N = 82 (Derman 1932). Native in central and southeastern Europe (England to Spain, Asia Minor and Caucasus), rarely cultivated in Wisconsin as a 306 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 shade tree : Milwaukee Co. : Milwaukee, 20 Sept. 1935, Goessl s.n. (WIS). Milwaukee, cultivated, 14 May 1938, Skinners 12899 \ (UWM) . MALVACEAE A. L. De Jussieu Mallow Family Herbs (shrubs and trees in tropical regions), often stellate- pubescent, with simple alternate, mostly palmately -veined and -lobed leaves. Flowers solitary or cymose, 5-merous, regular, hypogynous and bisexual (unisexual in Napaea). Calyx gamose - palous, persistent, often subtended by calyx-like bracts (paracalyx, epicalyx, involucre) . Petals separate, but slightly coalescent or frequently adnate to the base of the filament tube. Stamens numer¬ ous, monadelphous (filaments coalescent into a tube about ovary and style) ; anthers 1 -celled. Ovary superior, bearing as many terminal style branches as there are carpels. Carpels 5-many, either loosely coherent in a ring around the base of the single style and then separating at maturity or completely united into a com¬ pound ovary. Fruit a loculicidal capsule (Hibiscus) or carpels splitting ventrally and releasing seeds (Abutilon) or carpels sepa¬ rate, indehiscent, 1 -seeded schizocarps (Malva). A large, predominately tropical family, only one-third of the Wisconsin species native, the majority naturalized from Eurasia, including obnoxious weeds (Cheeses, Velvet-leaf, Flower-of-an- Hour) and showy Eurasian garden cultivars (Hollyhocks, Mal¬ lows), of which some persist after cultivation, becoming locally established (Kearney 1951) . KEY TO GENERA A. Carpels 5-20 or more, loosely united in a ring around a central axis, separating at maturity (schizocarps or mericarps) ; stamen-column antheriferous at summit. (Tribe MALVEAE) B. Carpels reniform, indehiscent, but not beaked, 1 -ovulate ; style-branches stigmatic along the inner face, slender- tipped. C. Flowers perfect; involucre usually present. D. Involucel of 6-9 distinct bractlets, united at base. E. Plants 7-12 dm tall; petals 1.8-3. 5 cm long; staminal tube terete, hairy; schizocarps 18-20, rounded on back, not keeled; rare escaped cul- tivar _ 1. ALTHAEA. EE. Plants 12-25 dm (or more) tall; petals 3. 6-4. 8 cm long ; staminal tub e 5-angled, glabrous ; schizocarps 25-35 (or more), keeled on back; common showy garden escape _ 2. ALCEA. 1970] Utech — Flora of Wisconsin No . 60 307 DD. Involucel of 3 bractlets or none. F. Petals obcordate, white or tinted with blue, pur¬ ple or red ; bractlets 3 ; common weeds _ _ 3. MALVA. FF. Petals truncate, rose red to purple ; bractlets 3 or none; rare, dry prairie species _4. CALLIRHOE CC. Flowers white, small, dioecious; involucre none; large- leaved robust native prairie herbs _ 5. NAPAEA. BB. Carpels with long divergent beaks (3-3.5 mm), dehiscent,, 3-8 seeded ; stigmas terminal and capitate ; flowers yellow ; very common weed _ 6. ABUTILON. AA. Carpels 5, united, the fruit a loculicidal capsule without a cen¬ tral column; stamen-column antheriferous along much of its length; flowers 3 cm or more in diam. (Tribe HIBISCEAE) _ _ _ _ _ _ 7. HIBISCUS. 1. ALTHAEA L. Marsh-mallow. 1. Althaea officinal|. L. Marsh-mallow. Map 2. Tomentose erect perennial 8-12 (-20) dm tall from an enlarged, knotty rootstock. Leaves triangular-ovate to cordate, crenate- serrate, gray to grayish-green, velvet-pubescent, the petioles 1.4-2. 8 cm long. Involucre united at base, the 6-9 lanceolate bract¬ lets densely stellate and hirsute, 4-5 mm long. Petals obcordate, 18-28 (-35) mm long, white to pale pink. Stamina! tube terete, hairy, commonly 12 mm long, bright violet; anthers purplish-red. Carpels 18-20, densely short stellate, convex and not keeled on back, the lateral face not radially veined. Seeds purplish black, 3. 6-3. 9 mm long. 2N = 42 (Skovsted 1935). Facultative salt-marsh plant, native to the drainage systems of the Caspian, Black and Baltic Seas, in North America as an escape along borders of saline or fresh water marshes. Formerly culti¬ vated for the perennial root which yielded the original non-synthetic mucilaginous marshmallow paste; collected twice in Wisconsin: Crawford Co.: Lynxville, 11 Sept. 1915, Davis s.n. (WIS). She¬ boygan Co.: Sheboygan, Aug. 1919, Goessl s.n. (WIS). Flowering in August to October. 2. ALCEA [Tourn.] L. Hollyhock. Linnaeus (Sp. PL 1753) followed TourneforPs distinction be¬ tween Althaea (marsh-mallow) and Alcea (hollyhock), though Willdenow, De Candolle, Bentham and Hooker and others have fused the two genera into Althaea. Recent studies by Zohary (1963a, 1963b) in SW Asia have demonstrated the validity of readoption 308 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 of this generic segregation : the staminal tubes in Alcea are 5-angled and glabrous, in Althaea, terete and hairy; the keeled schizocarps of Alcea are divided by an internal septum into an upper, empty cell and a lower one with a single seed, in Althaea, the schizocarps are rounded (on back), unilocular, one-seeded. Alcea, an Old World genus with 60 species, is an Irano-Turanian element (mainly East Mediterranean, SW & central Asia). 1. Alcea rosea L. Hollyhock. Map 3. Althaea rosea (L.) Cav. Robust biennial or perennial with spire-like stems to 2.5 m tall. Leaves cordate, shallowly 5- to 7- (9-)lobed, rarely dissected, densely stellate-pubescent, crenate; mid-stem leaves 5-11 cm long (blade petiole junction to lobe tip), 6-13 cm wide, the petioles (-3) 5-14 cm long. Flowers showy, 7-10 cm wide, solitary or 2-1}./ leaf axil or in a terminal raceme. Involucral of 6 (-9) lanceolate bractlets, united at base, densely stellate, the lobes 10-13 mm long. Calyx lobes 5, united, densely long-stellate, 11-16 mm long, the involucre and calyx accrescent at maturity. Petals 5 or double in horticultural forms, broadly obcordate, 3.6-5 (-6) cm long, white, yellow, pink, carmine-red to purplish-black or brownish- black, the claws bearded. Staminal tube 5-angled, glabrous. Pe¬ duncle stout, 1.8-2. 3 cm long at maturity. Carpels 26-30, pale brown, ribbed, densely stellate-hirsute on tips, the lateral face radially veined. Seeds light gray, 3-3A mm long. 2N — 42 (Skov- sted 1941). Native to the north-eastern Mediterranean region (Aegean Islands, and adjacent Balkan Peninsula), and not, as customarily ascribed, to China (Linnaeus, Sp. PI. 1753), which is beyond the natural range of the genus (Zohary 1963b) ; exact European intro¬ duction unclear, though long naturalized in SE Europe, Italy and S France; early herbalists Caspar Bauhin and Albertus Magnus noted that, like the tulip, introduction probably came in the 16th century from Turkey (Hegi 1925) . In Wisconsin frequently cultivated and persisting in the vicinity of gardens, dumps and waste places. Flowering from June to August or till frost. The cultivated hollyhock is undoubtedly polymorphic (Zohary 1963b), consisting at least in part of hybrids of A. pallida (Willd.) Waldst. & Kit., A. rosea L. and A. lav at erae flora (D.C.) Boiss. In recent years its cultivated popularity has decreased due to a wide-spread Chilean fungal leaf infection (Puccinia malvacearum Bert.). 1970] TJtech — Flora of Wisconsin No. 60 309 A horticultural variety, ALCEA ROSEA L. VAR. SIBTHROPII iBoiss., differs from A. rosea in having large fig leaf-like leaves with 7-elongate lobes; carpels 35 or more, densely stellate at tips, deeply keeled; seeds reddish brown, minitely papillose, 3. 8-4. 2 mm long; in Wisconsin a rare and sporadic escape in dumps and rubbish piles, occasionally cultivated singly or mixed with A. rosea. Mil¬ waukee Co. : Milwaukee, C. & N. R.R., on Locust St., one clump near patch of A. rosea, all probably sprouting from garden refuse, 3 July 1939, Skinners 518 (WIS). Winnebago Co.: Oshkosh, dump at end of Oak St., probably escaped from soil scraped off a garden, 2 July 1966, Harriman 925 (Oshkosh State Univ.) . Flowering in July. 3. MALVA [Tourn.] L. Mallow, Cheeses. Annual, biennial or perennial herbs with orbicular-renif orm, palmately dissected or lobed, crenate leaves. Flowers solitary or fascicled in leaf axils. Involucral bractlets (2-) 3, free, linear to obovate. Calyx 5-lobed. Petals 5, truncate, notched or obcordate. Anthers terminal on the staminal column. Styles as many as car¬ pels, stigmatic on the inner surface. Fruit of 8-20 radial carpels surrounding a central depression, these separating at maturity into as many one-seeded, indehiscent, round-reniform, laterally flattened schizo carps. Old World genus, native to Eurasia and North Africa, with 30 species, including several world-wide weeds and cultigens, such as the medical herbs Malva sylvestris and M. neglecta, which provide a leaf concoction (i.e. Malvae folia) that serves as an emollient and demulcent. Malva is an Old Latin name from the Greek malache or moloche, referring to the emollient leaves. The fruits are called cheeses since they resemble cheese wheels, and are eaten by chil¬ dren and poultry, and had been served on medieval tables. Reported hybrids (in Flora Europaea, Tutin et al. 1968) include M. alcea X M. moschata, M. sylvestris X M. neglecta, and M. neglecta X M. rotundifolia. Key to Species A. Upper leaves deeply 5- to 7-parted to below the middle or nearly to the base ; flowers chiefly solitary in the upper axils, showy; petals 2-3 cm long; escaped garden plants (Section BISMALVAE). B. Pubescence of spreading simple hairs; carpels densely hirsute on back; bractlets linear-lanceolate _ _ 1. M. MOSCHATA. 310 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 BB. Pubescence of short stellate hairs; carpels glabrous; bractlets oblong-ovate _ M. ALCEA. AA. Upper leaves reniform-cordate, with scalloped margins, only rarely lobed to the middle ; flowers fascicled in the axils ; petals 0.4-2. 3 cm long (Section MALVA). C. Bractlets oblong-ovate; petals rose-violet, 1. 4-2.3 cm long ; erect, escaped garden plant _ 2. M. SYLVESTRIS . CC. Bractlets linear or narrowly lanceolate; petals white with pale blue or purple tinged tips, 0.3-1. 2 cm long. D. Carpels 12-15, when mature puberulent and not rugose on back; lateral face of carels not radially veined ; petals 6-12 mm long; common prostrate weed _ _ _ _ 3. M. NEGLECTA. DD. Carpels 8-11, ivhen mature glabrous and rugose-reticu¬ late on back (Figs. 1 & 2) ; lateral face of carpels radially veined ; petals 3.2-6 mm long. E. Stems commonly prostrate to ascending; fruiting pedicels 10-45 mm long; bractlets 2. 9-3. 8 mm long; petals 3.2-4. 5 mm long ; common weed _ _ 4. M. ROTUNDIFOLIA. EE. Stems erect; fruiting pedicels 8-15 mm long; bractlets 3. 7-5.4 mm long; petals 4.8-6 mm long; rare garden escape _ M. VERTICILLATA. 1. Malva moschata L. Musk-malow. Map 4. Erect biennial to perennial, 3-8 dm tall, with knotty root-stocks, pubescent throughout, chiefly of simple divergent hairs. Leaves cordate, deeply 3- to 5- (7-) parted, the upper linear bipinnatifid, the lowest broadly 5-lobed, faintly musk scented (hence common name). Flowers solitary to 3 in upper axils, the fruiting pedicels 1.6-4 cm long. Bractlets 3, linear to oblanceolate, ciliate, glabrous on back, 5-6 mm long. Calyx 5-lobed, with simple trichomes, in¬ flated in fruit. Petals 5, triangular-ob cordate , pale rose-violet, 2- 2.4 cm long. Fruiting carpels 11-15, rounded not keeled and densely hirsute on back. Seeds reniform, gray brown, glabrous, 1.8-2 mm long. 2N = 42 (Skovsted 1935). Native of Europe to North Africa, (rail-transported) adventive over most of Europe, often cultivated as a garden plant for its showy flowers, in Wisconsin infrequent, escaping and persisting locally on sandy roadsides, dumps, waste places and beaches. Flow¬ ering and fruiting from mid June to mid September. 1970] Utech — Flora of Wisconsin No. 60 311 Malva alcea L. Vervain mallow. Erect robust perennial, 4-8 (-12) dm high, short stellate pubes¬ cent. Similar to M. moschata, but upper leaves more broadly di¬ vided. Flowers solitary, on short pubescent pedicels, these 1.6-2. 6 cm long in fruit. Bractlets 3, oblong to ovate, 4.5-6 mm long, densely stellate on back. Calyx 5-lobed, stellate pubescent. Petals 5, obcordate, notched, pale purple to white, 2-2.8 cm long. Carpels 18-20, when mature keeled and glabrous or sparsely pubescent on back. Seeds reniform, dark gray brown, glabrous, 2-2.3 mm long. 2N = 82 (Skovsted 1935). Native of Europe (Sweden to Spain,, the Balkans and S Russia), a pontic-Mediterranean element of dry and calcareous sites, in Wis¬ consin escaping sporadically from gardens but not persisting : Calu¬ met Co. : Roadside n of Stockbridge, 7 Aug. 1907, Goessl s.n. (WIS). Milwaukee Co.: Milwaukee, vacant lot on Hopkins St., % block s of Villard Ave., flowers light pink, garden escape, 15 July 1942, Fuller F-U2-86 (MIL). Flowering in late summer. 2. Malva sylvestris L. High-mallow. Map 5. Erect biennial to perennial, 4-8 dm tall from a shallow branched taproot, the stems glabrous or very sparsely hirsute. Leaf blades round-cordate or reniform, broadly 5-lobed, the terminal lobe ob¬ tuse to rounded, 2-9 cm long (petiolar leaf insert to lobe tip), 3-11 cm wide. Petioles 6. 5-9. 5 cm long, pubescent in a single line on the upper surface. Flowers fascicled in upper leaf axils; fruit¬ ing pedicels 2-3.5 (-5) cm long. Bractlets 3, oblong to ovate or obovate, 3.8-U.7 mm long, subglabrous, ciliate. Calyx 5-lobed, stellate-pubescent. Petals obcordate, notched, rose-violet, 1. 4-2.3 cm long. Mature carpels commonly 10, rugose-reticulate on the back, glabrous. Seeds round-reniform, 1. 9-2.1 mm long, blackish brown, subglabrous. Native Euro-siberian element (Europe, N. Africa, Asia Minor to Siberia), world-wide adventive, in Wisconsin grown in old gardens, locally persisting as an escape : Green Co. : Monroe, 7 Aug. 1894, Stuntz s.n. (WIS). Fond du Lac Co.: Pea field near old homestead, 1 mi. w from hwy 175 on Cemetery Rd., 1 Oct. 1967, Jeffers s.n. (WIS). Ozaukee Co.: Port Washington, growing wild, 17 Aug. 1887, Runge 137 (MIL). Sheboygan Co. : Sheboygan, July 1914, Goessl s.n. (WIS). St. Croix Co.: Baldwin, Anderson s.n. (WIS). Milwaukee Co. : Milwaukee, 1873, Sherman 673 (IA). The Wiscon¬ sin specimens have glabrous or sparsely hirsute stems, obtuse-lobed leaves and dark flower color, thus belong to var. mauritiana (L.) 312 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Boiss. (Fernald and Wiegand 1910). This is the common type grown in European farm gardens (Hegi 1925). Flowering from mid- July to early October. 3. Malva neglecta Wallr. Common mallow, Cheeses. Fig. 1 & 2. Malva rotundifolia of some American authors, not L. Map 6. Biennial or perennial with prostrate to ascending stems, 1-7 dm tall, from a deep branching taproot. Leaves round-cordate, shallowly 5-9 lobed, crenate, 2-4 (5.5) cm long (petiolar leaf insert to lobe tip), 3-6 (-9) cm wide, long petiolate. Flowers 1-5 in leaf axils, pedicels slender, 1-5 cm long. Calyx 5-lobed, stellate-pubes¬ cent at anthesis. Bractlets 3, linear, ciliate, 3. 6-4. 6 mm long. Petals 5, triangular-obcordate, 6-12 mm long, white with pale blue or purplish tips, the claws bearded. Staminal tube ca. 6 mm long. Styles and stigmas purple. Outline of carpel ring scalloped, the depressed center ca. 1/3 the diameter of the head (Figs. 1 & 2). Mature carpels 12-15, commonly 11>, densely puherulent, not reticu¬ late on back , the lateral faces smooth. Seeds round-reniform, 1 .7-1.9 mm long, laterally depressed, blackish brown, glabrous. 2N ±= 42 (Mulligan 1961)/ Native to temperate Eurasia (from England to L. Baikal) and N. Africa, widely adventive in Wisconsin mostly about dwellings, fields, lawns, roadsides, gardens and especially in highly nitrogenous waste places, becoming a noxious pest, most common south of the northern highlands, being limited by the colder climate, e.g. earlier frost, shorter growing season — cf. map of Galium aparine (Urban & litis 1957). Flowering continually from April to No¬ vember, fruiting May to November. 4. Malva rotundifolia L. Cheeses. Map 7, Figs. 1 & 2. Malva borealis Wallm. Malva pusilla Smith ex Withering Procumbent to ascending, branched biennial to perennial, 1-5 dm tall, from a slender unbranched taproot. Leaves round-cordate with 5-9 shallow lobes, crenate, 1-5 (-6) cm long (petiolar leaf in¬ sert to lobe tip), 3-9 cm wide, long petiolate. Flowers 2-6 in leaf axils, the fruiting pedicels 1-3.5 (-4.5) cm long. Calyx 5-lobed, with simple pubescence at anthesis, accrescent at maturity. Bractlets 3, linear, ciliate, 2. 9-3. 8 mm long. Petals 5, oblanceolate-obcordate, 3. 2-4.5 mm long, white with pale blue or purplish tips, the claws bearded. Staminal column ca. 3 mm long. Styles and stigmas purple. Marginal outline of carpel ring circular, the depressed center ca. 1/5 the diameter of the head (Figs. 1 & 2). Mature carpels 9-11, 314 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 • • «w mm m mm •••• M al va neg lecfa ■ M. ro+undifolia • RATIO : Diameter of Central Depression (mm.) / Diameter of Fruit (mm.) Figure 1. Scatter-diagram comparing the number of carpels per fruit and the width ratio of the central depression diameter divided by the fruit diameter of Malva neglecta and M. rotundifolia. Length (mm) Malva rotund i f o 1 i a Ma I va neg 1 ecta Bract lets 2.9 - 3.8 3.6 - 4.6 Petals 3.2 - 4.5 6.0 - 12.0 Stam i na 1 ca . 3 ca . 6 co 1 umn Figure 2. Comparative carpel illustrations and floral measurements for Malva neglecta and M. rotundifolia. 1970] Utech — Flora of Wisconsin No. 60 315 commonly 10, when young tomentulose, glabrous with maturity , conspicuously rugose-reticulate on back, the lateral face radially veined. Seeds reniform, 1.8-2 mm long, laterally depressed, dark brown, glabrous. Native of eastern Europe and western Asia (central Europe to Siberia and India), a world-wide adventive like Malva neglecta though not as frequent, a weed in southern Wisconsin in sandy, gravelly soils, about dwellings, waste and disturbed places, road¬ sides and railroad tracks. Similar in appearance to M. neglecta, but with fewer and glabrous carpels (Fig. 1) and shorter bractlets, staminal tube and petals (Morton 1937). Some European floras use M. pusilla Smith (ex Withering), because M. rotundi folia is considered a nomen ambiguum. Flowering and fruiting from April to November. Malva verticillata L. var. crispa L. Curled-mallow. Erect perennial to 1 m (or more) tall, with large reniform leaves crenately 5- to 7-lobed, the margins crisped. Petioles pubescent along a single line on the upper surface, 5-11.7 (-17) cm long. Flowers crowded in axillary fascicles, nearly sessile, the fruiting pedicels 8-15 mm long. Bractlets 3, linear-lanceolate, 4.4-5.4 mm long, with simple trichomes. Calyx 5-lobed, acuminate, with short stellate pubescence. Petals 5, ovoid-obcordate, notched, 5-8-6 mm long, white with pale blue tips, Mature carpels 8—11, glabrous, obsurely reticulate near carpel margins. Seeds ca. 2 mm wide, gray-brown, glabrous. Native of China, naturalized from S Asia to SE Europe, intro¬ duced to European gardens with Alcea rosea in the 16th century, sporadically adventive in North America, now seldom grown as a salad green or garden plant (Hegi 1925), in Wisconsin rarely persisting after cultivation: Racine Co.: Racine, (ca. 1860), Hale s.n. (WIS). Sheboygan Co.: Sheboygan, waste land, Goessl s.n. (WIS). Walworth Co.: Darien, brought in from Pflaum garden, 31 July 1935, Wadmond s.n. (WIS). Flowering from July to August. 4. CALLIRHOE L. Poppy mallow. Perennial herbs with erect stems fascicled from a thick fusiform taproot. Leaves triangular-cordate, crenate to deeply divided. Calyx 5-lobed, either naked or with a 3-leaved involucre at its base. Petals 5, triangular, broadly truncate, erose or fimbriate at summit, pur¬ ple, red-purple, occasionally pink or white. Staminal column antherif erous along more than half its length. Styles slender, stig- 316 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 matic along- inner surface, greatly elongated at anthesis. Carpels 10-15, these separating at maturity into as many one-seeded, indehiscent, reniform schizocarps. New World genus with 7 species of the Great Plains and northern Mexico. Key to Species A. Leaves triangular, crenate; involucre of 3 spatulate bractlets ; carpels keeled, strigose, the lateral faces not radially veined; native, dry sand prairies _ 1. C. TRIANGTJLATA. AA. Leaves deeply 5- to 7 -parted, the segments incised; involucral bractlets lacking; carpels strongly rugose, the lateral faces radially veined; rare, waif introduction _2 .C. ALCAEOIDES. 1. Callirhoe triangulata (Leavenw.) Gray Poppy mallow. Map 8. Erect perennial herbs with several stems to (2.5-) 6-8 dm tall, from a stout fusiform taproot, to 3 dm long. Stems, leaves and calyx harshly stellate. Leaves triangular-hastate, 6.5-13 cm long, 3. 6- 9. 5 cm wide, coarsely crenate, truncate to cordate at base, with variable petiole lengths 3-17 cm. Stipules oblong, persistent, 6-11 cm long. Floivers showy, panicled from axillary peduncels, 8.6- 20 cm long, the pedicels 1.6-2. 2 cm long at maturity. Bractlets 3, spatulate, ciliate, 6-9.5 mm long. Calyx deeply 5-parted, the lobes 2. 1-4.7 mm long at maturity. Petals magenta-purple, broadly truncate at summit, 1.9-2. 6 cm long. Staminal column 10-13 mm long, antherif erous laterally, the anthers yellowish brown. Carpels 10-12, one seeded, indehiscent, short beaked, keeled, strigose, not rugose on back, the lateral faces not radially veined. Seeds reni¬ form, laterally depressed, dark brown, 2. 3-2. 9 mm long. Rare showy sand prairie species of the Ozarks and Great Plains, confined in Wisconsin to the dry and sandy valleys of the Wisconsin and Mississippi Rivers : in Prairie du Chien cemetery on Mississippi River sand terrace, ( Jones U16, WIS) ; in Grant Co., Boscobel, prairie remnant along railroad, ( Musselman 2036, WIS) ; sandy upland, Midway Prairie, La Crosse Co., (Hartley 1U0U, WIS) ; sandy Pinus banksiana woods E of Gotham, Richland Co., (Nee 1068, WIS). The preservation of this handsome species is insured by the inclusion of the Midway Prairie as one of the Scientific Areas of Wisconsin. Flowering and fruiting in July and early August. 1970] Utech — Flora of Wisconsin No. 60 317 Callirhoe alcaeoides (Michx.) Gray Perennial with numerous erect slender stems to 4 dm tall from a shallow, swollen taproot. Lower leaves triangular-cordate, shal¬ lowly 5-7 lobed, the upper deeply 5-7 parted, these incised into numerous linear segments . Involucral bractlets lacking. Calyx 5-lobed, the lobes acuminate, elongating to 1 cm at maturity. Flowers 2.5-4 cm wide, the petals 1.4— 1.8 cm long, pale rose to white, fimbriate or erose at summit. Staminal column commonly 6 mm long, antheriferous along sides, the anthers dark red. Styles exceeding staminal column by 1.5 cm at anthesis. Carpels commonly 12, glabrous, strongly rugose on back, the lateral face radially veined. Seeds reniform, reddish brown, glabrous, 2. 1-2.3 mm long. Native of dry sand barrens, plains and prairies of south-central United States, where locally restricted. In Illinois limited to dry, gravelly or sandy, exposed soils, frequent along railroads (Jones and Fuller 1955), the only Wisconsin collection probably a waif introduction : Milwaukee Co. : Bay View, summer 1888, Range HO (MIL). 5. NAPAEA [Clayt.] L. Glade mallow. 1. Napaea dio|ca L. Glade mallow. Map. 9. Napaea dioica L. forma stellata Fassett, Torreya 42:179-180. 1943. [Type : Dane Co. : Along a railroad 3.8 miles west of Cross Plains, Aug. 16, 1942, Fassett 22057, (WIS, Isotypes in US, NY, GH, and several other herbaria.)] Robust, white-Yowered, dioecious perennial, 1-2 m tall from a stout taproot. Leaves 1-3 dm wide (basal leaves larger, to 6 dm), deeply 5-9 (-11) lobed or parted, the lobes coarsely toothed or incised, the lower surfaces strigose-pilose with or without ad¬ mixture of short stellate hairs. Flowers numerous, dioecious, with¬ out involucel, in large terminal panicles. Calyx lobes 5, united, ovate to triangular, 5-8 mm long. Petals white, notched, those of the staminate flowers 5-9 mm long, those of pistillate shorter. Staminate flowers with 16-20 anthers, pistils lacking, the pistillate with a short column of (usually) antherless filaments, the styles stigmatic along the inner surface. Carpels 8-10, when mature irregularly dehiscent into as many reniform, one-seeded schizo- carps, the back with stellate trichomes, rugose and ribbed. Seeds reniform, reddish brown, 3. 5-3. 8 mm long, glabrous. 2N = 30, N = 15 (litis & Kawano 1964) . The phytogeography, ecology and nomenclatural history of Napaea dioica have been reviewed by litis (1963) and Mickelson & litis (1966). The heliophytic habitat preference strongly sug- 318 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 ge sts that it is a wet prairie, not a forest species, which was asso¬ ciated with the formation and extension of the prairie peninsula. Flowering late June to mid- August, fruiting July to early September. Napaea is the only endemic gemus confined almost completely to the glaciated north-central United States, and with its monotype N. dioica being the only strictly dioecious Malvaceae in the West¬ ern Hemisphere. The Californian Sidalcea malachroides with 2N = 20 appears to be the closest morphologically, but only as a putative relative. Based on karyotype analysis, litis and Kawano (1964) suggest that Napaea is an ancient allopolyploid hybrid between a 2N = 20 species of Sidalcea and another species ( Sidalcea ?), now extinct, in which 2N = 10. The basic number would then be 5, not 7 as in all other members of the subtribe Malvinae. This would suggest segregation of Napaea and Sidalcea into their own subtribe. Some plants have simple, straight appressed hairs and few stel¬ late trichomes on the lower leaf surface. In others, these simple hairs are lacking, except on the larger veins, and are replaced by the short branched stellate trichomes (forma stellata Fassett, 1943). Frequently, the same plant will have upper leaves with mostly stellate trichomes, while the large basal leaves have mostly simple unbranched trichomes. Under the microscope these extremes are striking, yet there are numerous collections with both types mixed in various proportions (Utis 1963). Though it might be useful to name these extremes, it is impossible to know whether the Linnean type specimen has stellate, simple or mixed pubescence. Fassett assumed, on the flimsy basis of Sprengel’s (Syst. Veget. 3:122) statement that it is a “herba hirsuta”, that the type had simple hairs. This conclusion is not warranted, since any plant may be hirsute with simple hairs. It seems best to ignore forma stellata until the Linnean type has been examined. 6. ABUTILON [Tourn.] Miller Velvet-leaf. 1. Abutilon theophrasti Medic. Velvet-leaf, Butter-print. Map 10. Stout branched annual herb (2-) 5-18 dm tall, from shallow thick taproot 8-17 cm long, softly velvet-pubescent throughout. Leaves cordate, 4-12 (-19) cm long, 3.5-11 (-17) cm wide, acumi¬ nate, the margin entire or slightly toothed; petiole 4-11 cm long. Peduncles 13-26 mm long , jointed above the middle. Bractlets lacking; calyx 5, united at base, ovate-elliptic, persistent, 7-12 mm long. Flowers yellow, 14-28 mm across. Carpels 12-15 (-17) with conspicuous, horizontally spreading beaks, 3. 1-3.5 mm long, the 1970] U tech— Flora of Wisconsin No. 60 319 u schizo carps” ventrally dehiscent and not separating readily from the central axis. Seeds 3-6 (-8),/carpel, 3. 1-3.4 mm long, dark brown with short scattered stellate pubescence. 2N = 42 (Skovsted 1935, Smith 1965). Native of India and central Asia, a world-wide adventive, com¬ mon below the tension zone in southern Wisconsin, climate-limited (Lindsay 1935), as a noxious weed in cultivated and fallow fields, especially corn, but occasionally in gardens, along fence rows and waste places. The seeds retain their viability for over 50 years and are not killed by siloing (Steyermark 1963). The arrangement of the carpels resembles the print-blocks used on farms for stamp¬ ing rolls of butter. Flowering and fruiting continuously from July to October or till frost. 7. HIBISCUS [Tourn.] L. Rose-mallow. Annual or perennial herbs or shrubs with entire to palmately lobed or dissected leaves. Flowers large, showy, in terminal racemes or solitary in upper axils. Involucral bracts linear, 7-15. Sepals 5, broadly triangular, enlarged in fruit. Stamen-column long, with numerous lateral anthers; stigmas 5, capitate. Fruit a 5-locular, subglobose or prismatic loculicidal capsule, subtended or enclosed by the persistent, accrescent calyx. World-wide, subtropical to tropical genus, with ca. 150 species ; Hibiscus esculentus L., okra or gumbo, a garden vegetable grown for its soft immature edible pods and H. syriacus L., Rose-of- Sharon, a showy ornamental shrub, are common temperate eco¬ nomic species. Key to Species A. Tall perennial herbs, 1-2 m ; petals pink, 5-9 cm long ; fruiting calyx not inflated about the capsule. B. Leaves obovate-lanceolate, canescent beneath; involucral bracts 10-15; seeds not pubescent--!. H. MOSCHEUTOS. BB. Leaves hastate, green on both surfaces, glabrous; in¬ volucral bracts 9-10; seeds with reddish broivn hairs; Wisconsin and Mississippi River bottoms _ _ 2. H. MILITARIS. AA. Low annuals, 3-6 dm; petals pale yellow with purple center, 1. 5-3.0 cm long; fruiting calyx inflated; common southern Wisconsin weed _ 3. H. TRIONUM. 320 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 7. HIBISCUS [Tourn.] L. Rose-mallow. 1. Hibiscus moscheutos L. subsp. palustris (L) Clausen Hibiscus palustris L. Robust perennial to 2 m; stems densely to remotely stellate- pubescent. Leaves ovate to elliptic lanceolate, 8-20 cm long-, 9-15 cm wide, serrate to crenate, rarely lobed, green above, with whitish to grayish pubescence beneath. Petioles adnate to peduncle. In- volural bracts 9-14, linear, 1.4-2. 5 cm long. Petals pink , rarely white, lacking red band at base, 6.5-9 cm long. Styles with spread¬ ing pubescence. Capsules abruptly contracted to a beak, glabrous or essentially so. Seeds dark brown, papillose, 8-3.2 mm long. Saline, brackish or fresh water marshes of the northern coastal plain with inland extension to the Great Lakes region of southern Ontario, southern Michigan, northern Indiana and northeastern Illinois, in Wisconsin known from only one collection and now probably extinct: Rock Co.: Janesville, flowers pink, 25 Aug. 1889, Gertil Skavlem s.n. (WIS). Radford et al. (1968) treat Hibiscus palustris L. as a sub¬ species of the sympatric and southern H. moscheutos L., as previ¬ ously suggested by Fernald (1942, 1950). 2. Hibiscus militaris Cav. Rose-mallow. Map 11. Robust, essentially glabrous perennial 1-2 m tall, with several erect stems arising from a crown root. Leaves triangular to hastate, 8-12 cm wide, 8-15 cm long, acute at apex, basal lobes diverging, green on both sides; petioles 7-13 cm long. Flowers showy, solitary in upper axils, the peduncles 2. 3-5. 8 cm long. Bractlets 9-12, glabrous and linear, 1.7-2. 3 cm long. Petals 5, pink with red-purple base, 6-8.5 cm long. Carpels 5, dehiscent and beaked with 6-8 seeds carpel. Seeds with stiff, brownish-red trichomes, 3-3.3 mm long. 2N = 38 (Nakajima 1936 ex Darlington 1955). Native of marshes and muddy shores of pond and streams of the coastal plain, extending north of the Mississippi Embayment to Iowa, Minnesota, Indiana, Illinois and Wisconsin. The few Wis¬ consin stations are limited to the Mississippi and Wisconsin River bottoms where deposition and silting occur — levees, boat landings and alluvial forests. Leersia lenticularia Michx., Sagittaria montevidensis Cham. & Schlect. ssp. calycina (Engelm.) Bogin, and Rorippa sessiliflora (Nutt.) Hitchcock have similar distributions. Dispersability and habitat preference suggest a recent northward migration. Dispersal is no problem, for the seeds are eaten by ducks and bobwhites (Steyermark 1963). Recent migration is noted for Indiana by Deam (1940) : “I have known well the shores of the 1970] Utech — Flora of Wisconsin No. 60 321 Wabash River near Bluffton for a distance of 5 miles since 1880. The first colony of this species was noted in 1897 and is now com¬ mon all along the muddy shores and on the muddy bars in the river. In the early history of the state our streams were clear and when the forests were removed the streams became muddy and sediment was deposited on the shores and on the gravelly and rocky bars which made a suitable habitat for this species.’' Wisconsin’s earliest collection dates from 1914 : Crawford Co. : Bridgeport, 6 Aug. 1914, Denniston s.n. (WIS). Later collections indicate sites where deposition and silting would occur and create suitable habi¬ tats. Apparently native, though its recent northward extension has probably been due to man’s activity. Flowering late July to early August, fruiting August to September. 3. Hibiscus trionum L. Flower-of-an-hour. Map 12. Low hairy annual, much branched at base, 3-6 dm tall; taproot slender. Leaves deeply 3-parted, 1.5-6 cm long, the segments ob- long-obovate, irregularly incised; petioles 2-5 cm long. Bractlets 10-12, linear, ciliate with white trichomes, 7-12 mm long. Calyx 5-lohed, membranaceous , ribbed. Flower sulfur yellow with a redr- purple center, showy, but ephemeral (hence common name) ; petals 1.5-3 cm long. Mature capsules 5 -locular, black with long yellow trichomes, 5-7 seeds/carpel; the calyx surrounding it highly in¬ flated, translucent, with many vertical dark green ribs, papillose and long hirsute. Seeds brown-black, papillose, 2-2.3 mm long. 2N = 28 (Medvedeva 1936, ex Darlington 1955) and 2N = 56 (Skovsted 1935). Native of SE Eurasian agricultural center, found in the neolithic Aggtelek kitchen middens of Croatia (Yugoslavia), a Kulturbeglei- ter (“culture follower”) now adventive throughout Europe, eastern Asia to China, N. Africa and N. America (Hegi 1925) , its numerous seeds with unusual viability contributing to its spread (Deam 1940), a common weed in southern Wisconsin, in sandy soils of cultivated fields, gardens, roadsides, and disturbed or waste places. Flowering and fruiting from early July to late October or till frost. Bibliography Bailey, L. H. 1906. The Standard Cyclopedia of American Horticulture. New York: Doubleday, Page and Co. Braun, E. L. 1960. The genus Tilia in Ohio. Ohio J. Sci. 60(5) :257-261 Curtis, J. T. 1959. The Vegetation of Wisconsin. Univ. of Wisconsin Press, Madison. 657 pp. Darlington, C. D. and A. P. Wylie. 1955. Chromosome Atlas of Flowering Plants. George Allen & Unwin, Ltd., London. 322 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Deam, C. C. 1940. Flora of Indiana . Dept, of Conservation, Indianapolis. 1236 pp. Derman, H. 1932. Chromosome numbers in the genus Tilia. Journ. Arnold Arboretum 13 : 49-50. DesmaRaxs, Y. 1948. Dynamics of leaf variations in the sugar maples. Ph.D. Dissertation, Univ. of Wisconsin, Madison. - . 1952. Dynamics of leaf variations in the sugar maples. Brittonia 7(5): 347-388. Fassett, N. C. 1943. Some new forms from the Middle West. Torreya 42: 179-181. Fernald, M. L. 1942. Hibiscus moscheutos and H. palustris. Rhodora 44: 266-278. - . 1950. Gray’s Manual of Botany, ed. 8. New York: American Book Co. - and K. M. Wiegand. 1910. A summer’s botanizing in eastern Maine and western New Brunswick. Rhodora 12: 140-141. Fowells, H. A. 1965. Silvics of Forest Trees of the United States. U. S. Dept. Agr., Forest Service, Agr. Hand. No. 271, Washington. Gleason, H. A. and A. Cronquist. 1963. Manual of Vascular Plants of North¬ eastern United States and Adjacent Canada. New York: van Nostrand Co. Hagen, A. 1932. Preliminary Reports on the Flora of Wisconsin. No. 20. Malvales. Trans. Wis. Acad. Sci. Arts and Letters , Vol. 27: 247-249. Hartley, T. G. 1962. The Flora of the Driftless Area. Ph.D. Dissertation, State Univ. of Iowa, Iowa City. Hegi, G. 1925. Illustrierte Flora von Mittel-Europa. Miinchen: J. F. Lehmann Verlag, Vol. 5, I. Iltis, H. H. 1963. Napaea dioica (Malvaceae) : Whence came the type? Am. Midi. Nat. 70(1) : 90-109. - and S. Kawano. 1964. Cytotaxonomy of Napaea dioica (Malvaceae). Am. Midi. Nat. 72(1) : 76-81. Jones, G. N. 1968. Taxonomy of American species of linden (Tilia). III. Biol. Mono. 39, Univ. of Illinois Press, Urbana. - and G. D. Fuller. 1955. Vascular Plants of Illinois. Univ. of Illinois Press, Urban. 593 pp. 1375 maps. Kearney, T. H. 1951. The American Genera of Malvaceae. Am. Midi. Nat. 46(1): 93-131. Lindsay, D. R. 1953. Climate as a factor influencing the mass ranges of weeds. Ecology 34: 308-321. Mickelson, C. J. and H. H. Iltis. 1966. Preliminary Reports on the Flora of Wisconsin. No. 55. Compositae IV — Composite Family IV (Tribe Helenieae and Anthemideae) . Trans. Wis. Acad. Sci. Arts and Letters, Vol. 55: 197-222. Morton, C. V. 1937. The correct names of the small-flowered mallows. Rhodora 39: 98-99. Mulligan, G. A. 1961. Chromosome numbers of Canadian weeds. III. Can. J. Bot. 39: 1057-1066. Radford, A. E., H. E. Ahles and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. 1183 pp. Seymour, F. C. 1960. Flora of Lincoln County Wisconsin. P. F. Nolan, Taun¬ ton, Mass. Skovsted, A. 1935. Chromosome numbers in the Malvaceae. I. J. Genet. 31 : 263-296. 1970] U tech— Flora of Wisconsin No. 60 323 Smith, E. B. 1965. Chromosome numbers of Kansas flowering plants. II. Kansas Acad. Sci. 68: 463-464. Sorensen, P. D. 1966. The Flora of the old bed of Glacial Lake Wisconsin and the adjacent terminal moraine. MS. thesis, State Univ. of Iowa, Iowa City. Steyermark, J. A. 1963. Flora of Missouri. Iowa State University Press, Ames. 1725 pp. Stoever, D. H. 1794. The Life of Sir Charles Linnaeus. London. Sugiura, T. 1936. Studies on the chromosome numbers in higher plants, with special reference to cytokinesis. I. Cytologia (Tokyo) 7: 544-595. T'utin, T. G., et al. 1968. Flora Europaea. Vol. 2. Rosaceae to Umbelliferae. London: Cambridge Univ. Press. 455 pp. Urban, E. and H. H. Iltis. 1957. Preliminary Reports on the Flora of Wis¬ consin. No. 38. Rubiaceae-Madder Family. Trans . Wis. Acad. Sci. Arts and Letters , Yol. 46: 91-104. Zohary, M. 1963a. Taxonomical Studies in Alcea of Southwestern Asia. Part I. Bull. Res. Council Israel , Vol. II D4: 210-229. - . 1963b. Taxonomical Studies in Alcea of Southwestern Asia. Part II. Israel J. Bot. 12 : 1-26. PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 61 HYPERICACEAE— ST. JOHN’S-WORT FAMILY Fred H. Utecli and Hugh H. litis The Hypericaceae, a natural group often segregated from the more polymorphic, woody, tropical Guttiferae, has 2 genera and 14 species in Wisconsin. All but the ubiquitous Common St. John’s- wort (Hypericum perforatum) are native and occur either in dry, exposed sands or wet marly marshes or bogs, sandy swales and lake, river or stream-sides. A southern origin for the Great Lakes- Central Wisconsin endemic Hypericum kalmianum is suggested. Three species are reported here as new for the state: Hypericum prolificum , x H. dissimulatum and Triadenum virginicum. The present treatment revises McLaughlin’s (1931) preliminary report on Hypericaceae. Material from the following herbaria was intensively studied: University of Wisconsin (WIS), University of Wisconsin-Milwaukee (UWM), Milwaukee Public Museum (MIL), University of Minnesota (MIN), University of Minnesota- Duluth (DUL), State University of Iowa (IA) , Oshkosh State University, La Crosse State University, Northland College (Ash¬ land, Wis.), Beloit College and the private herbarium of Katherine Rill (Clintonville, Wis.-RILL). We are grateful to the curators of these herbaria for the loans of specimens. Dots on the maps represent exact locations, triangles, county records. Some locations have been added from Thomas Hartley’s unpublished “Flora of the Driftless Area” (1962), Paul Sorensen’s unpublished range maps from his Glacial Lake Wisconsin studies (1966), Olga Lakela’s Flora of Northeastern Minnesota (1965), Jones and Fuller’s Flora of Illinois (1955) and Frank Seymour’s Lincoln County sight record index (WIS). The map inset numbers record Wisconsin flowering and fruiting dates; plants with vegetative growth only, in bud or with dis¬ persed fruit were not included. For introduced species the year of earliest collection within a county is also recorded. Nomenclature and order of genera and species follows Gleason and Cronquist (1963) and Fernald (1950). 1 Field work and preparation of manuscript supported in part by the Research Com¬ mittee of the University of Wisconsin, on funds from the Wisconsin Alumni Research Foundation. 325 326 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Numerous people deserve special recognition for their help in manuscript preparation: Mrs. Katherine Snell, for encouragement and timely aids; Mr. Brian G. Marcks and Mr. Michael H. Nee, for comment and criticism; Mr. Eugene G. Coffman for photographic reproductions; Miss Bethia Brehmer and Miss Cynthia Loughran for artistic work; and Dr. John W. Thomson and Dr. Preston Adams for critical manuscript reading. HYPERICACEAE Lindley St. John’s-Wort Family Herbs or shrubs with opposite, simple, entire, often pellucid- or black-punctate leaves; stipules lacking. Flowers perfect, regular, hypogynous, solitary, axilliary or in cymes. Stamens 5 to over 100 ; filaments elongate, free or basally connate in 3 or 5 bundles, these opposite petals; anthers 2-celled, longitudinally dehiscent. Ovary superior; carpels 3-6; placentation parietal, pseudo-axile or axile (Fig. 1) ; styles distinct or united. Capsules septicidal, 1, 3 or 5 loculate. Seeds many, small, without endosperm. A small family with 12 genera and 600 species, usually segre¬ gated from the tropical Guttiferae (Clusiaceae) . KEY TO GENERA A. Petals yellow to orange, convolute in bud; stamens numerous to few, distinct or united at base into 3 to 5 clusters ; hypogy¬ nous glands lacking _ 1. Hypericum. AA. Petals pink to mauve-purple, imbricate in bud; stamens 9, strongly triadelphous ; hypogynous glands 3, orange, alternate with the stamen bundles _ 2. Triadenum. 1. HYPERICUM [Tourn.] L. St. John’s-Wort. Annual or perennial herbs or shrubs; leaves simple, frequently pellucid or black-dotted, opposite, entire, frequently with axillary decussant branchlets ; stipules lacking. Inflorescence cymose. Sepals 5, often unequal, persistent. Petals 5, yellow to orange, convolute in bud, often black-dotted. Stamens numerous or few (small fid. spp.) ; filaments free or basally connate. Ovary superior; styles united or separate and divergent; stigmas minute or capitate. Placentation parietal (1-celled), pseudo-axile by intrusion of pla¬ centae (partially 3- or 5-celled) or axile (completely 3- or 5-celled) (Fig. 1). Capsules septicidal, 1- to 5-carpellate. Seeds small (ours 0.5-3. 0 mm), short-cylindric, aerolate. The largest genus of the Hypericaceae, world-wide, throughout temperate and tropical montane regions, with ca. 300 species of annuals, perennials with persistent rhizomes, and woody shrubs. 1970] Utech and litis — Flora of Wisconsin No. 61 327 Key to Species A. Styles united at base into a single straight beak at anthesis, splitting at maturity; stigmas minute, never capitate; sta¬ mens many, distinct. (Sect. MYRIANDRA) B. Small woody shrubs; leaves and sepals articulate at base; withered stamens deciduous soon after anthesis. (Subsect. Centrosperma) . C. Midstem leaves 2. 6-4. 5 cm long, sessile; styles and carpels 5; cymes chiefly terminal, 3- to 7-floivered. _ _ _ 1. H. KALMIANUM. CC. Midstem leaves 3.5-7 cm long, short-petiolate ; styles and carpels 3; cymes terminal and axillary, 11- to 19-flowered; rare. _ 2. H. PROLIFICUM. BB. Perennial herbs slightly suffrutescent at base ; leaves and sepals not articulate at base; withered stamens persistent long after anthesis. (Subsect. Pseudo- brathydium) . D. Plants robust, 30-60 cm tall, rhizomatose, the rootstock often woody; leaves linear-elliptic, 30-58 mm long; seeds 2. 0-2.7 mm long; rare, moist prairies, Green Co. and Rock Co. _ _ 3. H. SPHAEROCARPUM. DD. Plants slender, 15-35 cm tall from horizontal rhizome, the bases herbaceous; leaves elliptic to ovate, 16-33 mm long; seeds 0.5-0. 8 mm long; central and northern Wisconsin lakes and river margins _ 4. H. ELLIPTICUM. AA. Styles free to base, the capsules not beaked ; stigmas capitate ; stamens many to few, connate basally into 3 or 5 bundles (phalanges) . E. Stigmas and styles 5; capsules 5-celled, 8-15 mm wide; flowers 50-60 mm across ; stamens 5-delphous, numerous (over 150) ; larger leaves 5-8 cm long; robust perennial of wet habitats. (Sect. ROSCYNA) _ _ 5. H. PYRAMIDATUM. EE. Stigmas and styles 3; capsules 1- or 3-celled, 1-5 mm wide; flowers 5-30 mm across; stamens 60-5; larger leaves less than 5 cm long. F. Capsules 3-celled, with azle placentae; flowers 6-32 mm across; corolla black-dotted; stamens 60-27, weakly 3-delphous. G. Capsules oblong-conic, 4. 5-6. 5 mm long; styles 4-5 mm long; flowers 15-30 mm across; petals 328 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 black-dotted on margin only; stamens (45) -50- (60) ; common Eurasian weed. (Sect. HYPER¬ ICUM _ 6. H. PERFORATUM . GG. Capsules subglobose-ovate, 3. 8-4. 6 mm long; styles 1.5-2. 5 mm long; flowers 6-10 mm across; petals and sepals marked with black dots and lines; stamens (27) -35- (40) ; native. (Sect. ELINEATA) _ 7. H. PUNCTATUM . FF. Capsule 1-celled, with parietal placentae; flowers 4-7 (-10) mm across; corolla yellow, lacking black dots; stamens 20-5, weakly 5-delphous: (Sect. BRATHYS) H. Leaves linear to elliptic-ovate, 8-44 mm long, 1-15 mm wide, 3- to 7-nerved; inflorescence cymose. I. Capsules broadly ellipsoid or oblongoid, the apex rounded to obtuse; inflorescence diffuse and leafy-bracted, not well defined, the angle between a pair of lateral pe¬ duncles 70° or more (Fig. 5) ; sepals ob¬ long to elliptic, widest near middle. J. Uppermost bracts highly reduced, se¬ taceous; cauline leaves often over 20 mm long, deltoid to ovate, usually cor¬ date-clasping ; sepals acute, equaling capsule ; plants often 3-5 dm tall ; mostly Driftless Area. _ _ 8. H. MUTILUM? JJ. Uppermost bracts foliaceous; cauline leaves 8-15 mm long (rarely longer), ovate to elliptic, sessile but not strongly clasping; sepals obtuse, shorter than capsule ; plants usually 1-3 dm tall ; widespread. _ _ 9. H BOREALE? II. Capsules ovoid to conic, the apex nar¬ rowed; inflorescence rather compact and clearly defined, setaceous-bracted not leafy, the angle between a pair of lateral pe¬ duncles 70° or less (Fig. 5) ; sepals lanceo¬ late to linear, acute or acuminate. K. Leaves elliptic-lanceolate to ob¬ long, the larger 5-12 mm wide, rarely narrower, the bases sub- cordate-clasping; sepals 4-6.5 mm 1970] Utech and litis — Flora of Wisconsin No. 61 329 long ; capsules elliptic-ovate, 5- 7.8 mm long; plants often 3-5 dm tall ; common throughout state. — _ 10. H. MAJUS2 KK. Leaves linear to linear-elliptic or -oblanceolate 1-3 mm wide, 1- (3-) nerved, the bases sessile- attenuate; sepals 2. 5-4.4 mm long; capsules conic, 3-5.2 mm long; plants usually 1-3 dm tall; mostly northern Driftless Area. _ _ 11. H. CANADENSE2 HH. Leaves minute, scale-like, 1.5-3 mm long, 0.5 mm wide, strictly 1-nerved; inflorescence racemose ; branches wiry ; dry sands, Driftless Area. _ 13. H. GENTIANOIDES. 1. Hypericum kalmianum L. Kahn’s St. John’s-Wort. Map 1, Figs. 1-4. Slender shrubs 2-6 (-10) dm high, with branches 4-angled, the branchlets 2-angled. Leaves linear-elliptic to oblanceolate , revolute, coriaceous, sessile, obtuse, mostly 26-45 mm long, 4-8 mm wide (Fig. 4). Cymes 3- to 7-flowered (rarely more), restricted to termi¬ nal node (Fig. 2). Flowers 20-35 mm across. Sepals oblong-elliptic, foliaceous, 6-8 mm long. Petals 5-14 mm long. Stamens numerous, distinct. Ovary 1 -locular, usually with 5 pseudo-axile intruding placentae (Fig. 1) ; styles (3) -5- (6) ; stigmas never capitate. Capsules ovoid, 5-carpellate (rarely 3, 4 or 6), 7-10 mm long, 4-7 mm wide. Seeds light brown, 0.7-1. 1 mm long. N = 9 (Hoar & Haertl 1932; Robson & Adams 1968). Central Wisconsin sand plains and sphagnum-sedge meadows, in rocky shores, sandy swales, behind dunes, and calcareous low prairies about Lakes Michigan, Huron and Erie, to the Ottawa River, Quebec (Fig. 3: cf. maps McLaughlin 1930, Guire & Voss 1963, Adams 1959b), its Wisconsin distribution closely associated with the desiccated beds and outwash plains of Glacial Lakes Wis¬ consin, Oshkosh and Chicago. Flowering late June to early Octo¬ ber; fruiting early July through October. The history of Hypericum kalmianum is of particular interest, since all its stations are in glaciated territory (Adams 1959b, Guire & Voss 1963, McLaughlin 1931) ; this restriction suggests either a a Hybrids between the species 8-11 are not uncommon (cf. Hybrids of Sect. Brathys, 12a, 12b, and 12c, pp. 29-32). Depauperate plants are common but can be keyed with help of full grown ones nearby. 330 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 J^n rid HYPERICUM hypericum' % \ 1 PROLIFICUM KALMIANUM Jl’ J HYPERICUM^ v [_' ellipticum HYPERICUM^ H SPHAEROCARPUM ! - 1 r1- LL I NO S NOI S j". r* HYPERICUM % \ " “"I I PYRAMIDATUM HYPERICUM % H l PERFORATUM Y~ LL mOl S tuug: * ■ j ■ 7 1 ( 1 « - \s£Ml r 1970] Utech and litis — Flora of Wisconsin No. 61 331 (s) H perforatum /Sect. Hypericum! 0 | .2 ' : 3 ( 6 ) H pyramidatum /Sect. Roscynu J Figure 1. Diagrams illustrating the different types of placentation found in Wisconsin Hypericum: 1. H. boreale (Sect. Brathys) — parietal. 2. H. ellipticum (Sect. Myriandra) — pseudo-axile. 3. H. prolificum (Sect. Myriandra) — pseudo- axile. 4. H. kalmianum (Sect. Myriandra) —^pseudo-axile. 5. H. perforatum (Sect. Hypericum) — axile. 6. H. pyramidatum (Sect. Roscyna) — axile. 332 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Hypericum 1 kalmianum Hype ricum prolificum Hypericum loJoocarpum Wa -y A ^ >e / • c \ / c ^ A B C H. kalmianum 0 M/M/ T ota Is/ Node A (terminal) B (axil 1 ary 1 C ( ' ' > . C ( " ) 37 (7.8%) 168 (35.2%) 19 (3.9%) 3 (0.6%) 245 (5 1 .4%) * 19 (3.9%) 23 (4.8%) 4 (0.8%) 477 ( 100 %) 38 7.8 3 0.6 0 0 H. prolificum Totals / Node B (axillary) C ( " ) D ( . ' ' ) 32 (35.2%) 28 (30.3%) 28 (30.3%) * 48 (52.7%) * 52 (57.1%) * 24 (26.3%) 6 (6.6%) II (12.1%) 10 ( 10.9% ) 8 (8.8%) 91 ( 100 %) 90 98.3 60 65.4 6 6.6 H. lobocarpum Totals / Node A (terminal) B (axillary) C ( " ) 0 < " ) 1 (1.6%) 3 (5.0%) 1 (1.6%) 18 (29.5%) 19 (31.1%) 3 (5.0%) 3 (5.0%) 38 (62.3%) * 28 (45.9%) * 22 (36.0%) * 4 (6.6%) 61 ( 100 %) 50 82.0 26 42.6 3 5.0 . Figure 2. Statistical model and data comparing the inflorescences and degree of dichasial branching of Hypericum kalmianum, H. prolificum and H. lobocarpum. 1970] Utech and litis — Flora of Wisconsin No. 61 Glacial maxima H. densiflorum Figure 3. Distributional maps of Hypericum kalmianum L. H. lobocarpum Gatt., H. densiflorum Push and H. proliflcum L., based in part on the unpub¬ lished map of Adams (1959b). 334 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 Figure 4. Scatter-diagram comparing leaf width and length and indicating number of carpels and degree of inflorescence branching of Wisconsin Hypericum kalmianum and Wisconsin and non-Wisconsin H. prolificum. pre-glacial origin with subsequent survival either in unglaciated or in once-glaciated territory between differentially advancing gla¬ cial lobes, or a recent, post-glacial origin from a more wide-spread southern species. The last hypothesis seems to us the most reason¬ able. Adams (1959b, 1962) thinks this ancestor to be the Gulf Coastal and Mississippian Embayment H. lohocarpum Gatt. ( = H. oklahomense Palmer), since this species and H. kalmianum both have mostly 5-carpellate ovaries, differing from the 3-carpellate, but otherwise similar and related H. prolificum L. The eastern coastal and inter-montane H. densiflorum Pursh is also commonly 3-carpellate : many authors de-emphasize carpel number and follow Svenson (1940) in considering H. lohocarpum as a variety of H. densiflorum. Carpel number is extremely variable even in well- defined species of section Myriandra. In any case, the fruits of both 5-carpellate taxa ( H . kalmianum and H . lohocarpum) are very similar, except that those of H. lohocarpum tend to be smaller on the average and more deeply sulcate. 1970] Utech and litis — Flora of Wisconsin No. 61 335 Comparison of degree of dichasial branching and flowers/in¬ florescence (Fig. 2) of H. kalmianum, H. prolificum and H. lobo- carpum reveals a north to south increase, with the northern H. kal¬ mianum usually fertile only in the uppermost node, the southern H. lobocarpum highly floriferous in many (3 to 5 or more) nodes. H. prolificum has the largest number of inflorescence combinations. Both H. prolificum and H. kalmianum have broad foliaceous sepals (4-8 mm long) and flowers to 30 mm wide; H. lobocarpum and H. densiflorum have usually shorter, narrower sepals (2-4 mm long) and often smaller flowers (to 20 mm across). Thus, while H. kalmianum was probably derived from H. lobocarpum (or H. densiflorum) , the reduced inflorescence-branching, foliaceous sepals and larger flowers suggest some H. prolificum introgression into H. kalmianum , which would not be unlikely, considering the geo¬ graphic proximity of the two populations. 2. Hypericum prolificum L. Map 2, Figs. 1-4. Hypericum spathulatum (Spach) Steud. of ed. 8, Gray’s Manual. Erect bushy shrub 3-9 dm tall, diffusely branched ; bark shreddy, gray, the branchlets sharply 2-angled. Leaves oblanceolate-linear , obtuse and often mucronate, the margins strongly revolute , punc¬ tate, the midstem leaves 3.5-7 cm long, 7-15 mm wide (Fig. 4). Petioles 1-4.6 mm long. Cymes 11- to 19-flowered, terminal and axillary (Fig. 2). Flowers 15-27 mm across. Sepals ovate, mucro¬ nate, 4.5-6 mm long. Petals obovate, bright yellow, 7-10 mm long. Stamens numerous, distinct. Ovary 1 -locular usually with 3 pseudo- axile intruding placentae (Fig. 1) ; styles 3 (4) ; stigmas not capitate. Capsules ellipsoid-ovate, 3-carpellate (rarely 4 or 5), 10- 13 mm long, 3.5-6 mm wide. Seeds black, 1.2-1. 8 mm long. N = 9 (Nielsen 1924, Robson and Adams 1968). A variable species in eastern and central United States (Fig. 3) on dry creek beds, sandy or rocky slopes, roadsides and old fields, occasionally cultivated, reported here for Wisconsin for the first time, Swezey’s (1883) use of this name being based on collections of H. kalmianum (Lapham s.n. and Hale s.n., WIS, MIL) . All collections are recent, perhaps escapes from cultivation. Crawford Co.: town of Clayton, S. 11 SE 9 July I960, Densmore s.n. (WIS) ; sandy hillside E of Soldiers Grove on Co. E, with Silphium perfoliatum, Corylus americana, Rhus radicans, 4 Aug. 1960, Schlising & Musolf 17 h 9 (WIS). Dane Co.: Edgerton, Camp Hickory Hill, open, light sandy soil, 5 Aug. 1947, Dorney s.n. (RILL). Flowering late July and early August. 336 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 3. Hypericum sphaerocarpum Michx. Map 3. Hypericum cistifolium of authors, not Lamark Erect perennials from woody branched rootstocks, the deep rhizomes with adventitious shoots. Stems herbaceous, 30-58 cm tall, 4-lined. Leaves linear-oblong to narrowly elliptic, acute to obtuse, sessile, 3-7 cm long, 4-15 mm wide. Cymes compact, com¬ pound, many-flowered; bracts lanceolate. Sepals ovate-lanceolate, 2. 8-4.8 mm long. Petals yellow, 5. 3-8. 6 mm long. Stamens numer¬ ous. Ovary 1 -locular, with 3 intruding parietal placentae. Capsules globose to ovoid, firm, few-seeded, 4.5-6. 7 mm long; styles 3, united in sharp beak. Seeds blackish-brown, coarsely reticulate, pitted, 2. 0-2. 7 mm long, the raphe developed into a keel. Species of low or mesic prairies, limestone outcrops and cedar glades, Ala. to SW Ark., north to Iowa, Ill. and N Ind., rare in Wisconsin, confined to low wet prairies along the Sugar and Rock Rivers, as in low rich prairie (near Monticello, Green Co.), with scattered willows, dogwood, bur oak, Potentilla arguta, Eryngium yuccifolium, and Ratibida pinnata. The plants are almost always pulled up without the very deep slender horizontal rhizomes. Flow¬ ering late June to mid-August, fruiting mid-July through August. 4. Hypericum ellipticum Hook. Creeping St. John’s-Wort. Map 4, Fig. 1. Erect unbranched perennial from a reddish, spongy, slender, creeping rhizome. Stems 15-35 cm tall, obscurely 4-angled. Leaves elliptic to elliptic-lanceolate, the larger 16-33 mm long, 4-13 mm wide, pellucid-punctate, not revolute, the midvein prominent. Cymes terminal, few-flowered; bracts linear to lanceolate. Sepals narrowly obovate to oblanceolate, 4-6 mm long. Petals oblanceolate, 4. 6-8. 6 mm long, often reddish in bud. Stamens numerous. Ovary 1-locular with 3 intruding parietal placentae (Fig. 1). Capsules subglobose to ovoid, 4-7 mm long, many-seeded; styles 3, united at base. Seeds dark reddish brown, striated, pitted, 0.5-0. 8 mm long. N = 9 (Hoar & Haertl 1932). A “northern hardwoods” species, from NE Tenn. to Newfound¬ land, west to Lake Superior, in central and northern Wisconsin along stream banks, pond and lake shores, river flats and sand bars, as in cold streamside Alnus thickets in sunny Car ex meadows, Brule River, Florence Co. The northern-most member of the 30 species of section Myriandra (Adams 1962) and the only Wiscon¬ sin Hypericum with prominent (oft-collected) rhizomes. Flowering late June to early August, fruiting July to latest October. Submerged aquatic plants with simple sterile stems and round to ovate “feather-veined” leaves, resembling Callitriche, have been 1970] Utech and litis— -Flora of Wisconsin No. 61 337 designated as forma SUBMERSUM Fassett (1939, 1960), while terrestrial plants with axillary branches overtopping the mature infructescence, as forma FOLIOSUM Marie-Victorin (Le Nat. Canadien 71 : 201. 1944.) 5. Hypericum pyramidatum Ait. Giant St. John’s-Wort. Hypericum ascyron of Am. authors, not L. Map 5, Fig. 1. Robust erect perennial herbs, 6-16 (-20) dm tall; branches 4- angled ; root-crown woody. Leaves ovate-oblong to lanceolate, the larger 5. 5-9. 6 (-11) cm long, 2.5-4 (-5) cm wide, acute or obtuse, with sessile clasping bases. Stamens numerous (over 150), 5- delphous ; anthers versatile. Sepals ovate-triangular, 6.5-8 mm long. Petals broadly ohovate, persistent, 1 .8-2.8 cm long. Styles 5, 6- 10 mm long, halfway united, divergent above ; stigmas capitate. Capsules conic-ovoid, completely 5-celled, septicidally dehiscent, 15-25 mm long, 8-15 mm wide (Fig. 1). Seeds brown, lustrous, reticulate, 1.1-1. 4 mm long, the raphe keeled. N = 9 (Nielsen 1924). Quebec to Minn., south to Penn., Ind., Mo. and Kansas, in Wis¬ consin in wet and open habitats as gravelly river banks, sphagnous sedge meadows, mesic forest edges and drainage ditches, as in low wet muck meadow near Mauston with Carex, Polygonum, Physo- stegia virginiana, or in a weedy floodplain prairie along Pine River, Richland Co., with Napaea dioica, Artemisia s errata, Silphium perfoliatum, Urtica sp., ( Nee 11^53, WIS). Flowering late June to mid-September, fruiting mid- July to early October. It is closely related to the true Hypericum ascyron L. of eastern Asia (E Siberia, Japan & China). 6. Hypericum perforatum L. Common St. John’s-Wort, Klamath wed. • Map 6, Fig. 1. Erect branching perennials from a subligneous crown with short shallow rootstocks and deep branching taproot. Stems 2-angled, 4-6.5 (-8) dm tall, with numerous sterile basal shoots and leaf axillary decussate branchlets. Leaves linear-oblong to elliptic, pel¬ lucid-dotted, obtuse, sessile, 5-nerved, commonly 12-36 mm long, 3-9 mm wide, reduced on axillary branchlets. Cymes paniculate, flat-topped; flowers numerous, 15-30 mm across. Sepals linear- lanceolate, acuminate, 4-7 mm long. Petals orange-yellow, black- spotted near margins, 9-1 U mm long. Petals and stamens persist¬ ent. Stamens (45)-50-(60) . Ovary 3-loculate ; styles 3, divergent, 3-4.5 mm long; stigmas capitate. Capsules completely 3-celled (Fig. 1) oblong-elliptic to conic, veiny, 4-7 mm long, 3-5 mm wide. Seeds blackish brown, lustrous, reticulate, 0.8-1. 2 mm long. N = 338 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 16, 2N = 32 (Hoar & Haertl 1932, Mulligan 1957, Robson & Adams 1968). A noxious world-wide weed, native to N Africa, W Asia and Europe (Hegi 1925), naturalized in E and W North America, in Wisconsin a common weed in open, sandy, poor or worn soils, chiefly on roadsides, railroads, neglected fields, beaches, sand plains, blowouts and barrens, occurring with such disturbance in¬ dicators as Corny tonia yeregrina, Hieracium aurantiacum, Am¬ brosia art emisii folia, Daucus carota, Ascleyias syriaca and Euyhorbia esula. Flowering from early June to early September, fruiting from late June to early October. This adventive is especially troublesome in the Klamath River Basin (N Calif. & S Ore.). The stem’s numerous resin canals con¬ tain hypericin, which is poisonous to livestock, but probably not fatal (Marsh & Clawson 1930, Kingsbury 1964). Eradication is difficult due to deep perennial roots, vigorous leafy basal offshoots and numerous, highly viable, genetically similar seeds, megasporo- genesis being 97% apomitic (pseudogamous) (Tutin et al. 1968). The name “St. John’s Wort” is derived from the belief that the plant’s dew precented sore eyes on St. John’s eve, June 24, when huge ceremonial bonfires of this plant blazed throughout Europe. Bouquets then gathered were hung in windows as talismans against thunder, witches and other misfortune, while in Switzerland, young women put them under their pillows believing they would marry the men of their dreams. The dark-red pellucid leaf dots supposedly appeared on August 29, the day John the Baptist was beheaded (Hegi 1925, Clohisy 1930). 7. Hypericum punctatum Lam. Map 7. Sparingly branched perennial with terete stems 4-10 (-12) dm tall. Leaves oblong-elliptic to lanceolate, the larger 3-7 cm long, 1-2 (-3) cm wide, dark punctate, blunt or retuse, 5- to 7-nerved; base clasping to attenuate. Corymbs compact; flowers 6-10 mm wide, short-pedicellate. Seyals ovate-oblong , broadly acute, black - syotted and - lined , 3. 8-6.8 mm long. Stamens (27) -35- (40) , weakly 3-delphous ; anthers black-dotted. Ovary 3-locular ; styles 3, free, 1.5-2. 5 mm long; stigmas capitate. Caysules 3-celled, sub- globose to ovoid, S.8-^.6 mm long, 3.6-U.2 mm wide, with elongate oil vesicles. Seeds yellowish-brown, 0.6-0. 7 mm long. N = 8, 2N — ring of 16 at anaphase (Hoar 1931, Robson & Adams 1968). Eastern North America, from Maine to Minn., south to Fla. & Texas, in Wisconsin along forest edges, open wooded slopes, flood- plain thickets, wet prairies, abandoned fields and roadsides : in low, mesic woods (near Tomah, Monroe Co.), with Trillium cernuum, 1970] Utech and litis — Flora of Wisconsin No. 61 339 Uvularia sessilii folia, Mitchella repens, Aster macro phyllus and in bottomland woods on floodplain terraces (Trempealeau Co.), with Quercus bicolor, Fraxinus spp., Betula nigra, Carex spp. Though widely distributed, it is never abundant, this possibly related to the “Oenothera-like” ring chromosome segregation pattern, pro¬ ducing various isolated and inbred populations. Flowering from early July to early September, fruiting August to mid-September. 8. Hypericum mutilum L. Map 8, Fig. 5. H. mutilum var. long [folium Bob. Keller, in Bull. Herb. Boiss. (ser. 2) 8: 184. 1908, ex char. [Type: Visconsin (sic!), Kumelien (sic!), 113.] The variety longifolium is apparently nothing but a long-leaved plant, not taxonomically recognized here. There are no Hypericum specimens in the very fragmntary Kumlien Herbarium deposited at WIS. The type is presumably in G. Erect slender annuals or perennials (1-) 2-6 dm tall, branched diffusely above, the leafy-bracted bases decumbent. Leaves deltoid- ovate to oblong-lanceolate, obtuse, cordate-clasping, 3- to 5-nerved, minutely punctate, the larger 1-2 (-3) cm long, (3-) 8-13 mm wide. Inflorescences lax and diffuse, poorly defined, leafy-bracted, the pedicels very slender, unequal, the angle between a pair of lateral peduncles ca. 70-105°. Uppermost bracts setaceous, 1.5-3. 8 mm long (Fig. 5). Flowers 2. 6-4. 5 mm wide. Sepals linear-oblong, acute, 2-4 mm long, equaling capsule. Petals 2-3 mm long. Stamens 5-10. Ovary 1-locular, the styles 0.5-0. 9 mm long; stigmas capitate. Capsules ovoid to ellipsoid, greenish at maturity, 2-4 (-5) mm long, 1. 6-2.4 mm wide. Seeds yellow, striate, minutely rugose, 0.45-0.55 mm long. N = 8 (Hoar & Haertl 1932). Eastern N. America, from Minn, to Newfoundland, south to Fla. and Texas, in Wisconsin mostly in the lower Wisconsin River valley and Driftless Area, on sandstone cliff ledges, sandy creek margins and river flats, moist sandy or black muck lowland meadows, swales and desiccated temporary pools, rarely in moist woods or abandoned fields. Flowering July to September; fruiting early August to late September. A highly variable species with populations in Brazil, Hawaii and Europe. 9. Hypericum boreale (Britton) Bicknell Map 9, Figs. 1, 5, 6. Northern St. John’s-Wort. Slender, rhizomatous and decumbent, branched perennial herbs with terete to obscurely 4-angled stems 1-3 (-4 in tall grass) dm tall, the bases often with leafy short-shoots. Leaves ovate to ellip- ■^i , [• HYPERICUM HYPERICUM '"'x mutilum" PUNCTATUM I IL I t\|op S y i HYPERICUM HYPERICUM MAJUS L L I N OjV S i n y L HYPERICUM HYPERICUM^\ 1 i— GENTIANOIDES CANADENSE 340 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 1970] Utech and litis — Flora of Wisconsin No. 61 341 HYPERICUM Figure 5. Line-drawi'ngs of several Wisconsin small-flowered Hypericum (Sect. Brathys) , (From N. C. Fassett, 1960, p. 246, with permission). 342 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 HYPERICUM CANADENSE X HYPERICUM DISSIMULATUM A A V V V V V ~t~ HYPERICUM BOREALE State of upper-most bract: Foliaceous Setaceous Nerve number of mid-stem leaves: 1-nerved 3-nerved 5-nerved Primary Dichasial Angle ^ 0-30° 30 j- 60° 60 - 90° (or more) Collection: Jackson Co. , City Point. Utech 68-200. (WIS). T I T— I — T 1 I™”1 1 I I 0.1 0.2 0.3 0.4 0.5 0.6 0.7 ' 0.8 LEAF WIDTH / LENGTH RATIO (mid-stem) Figure 6. Scatter diagram of mass collection (Utech 68-200) clearly showing intermediates (x Hypericum dissimulatum Bicknell) between H. boreale and H. canadense. tic, if aquatic suborbicular, obtuse to rounded, obscurely punctate, sessile, 3- to 5-nerved, the larger 8-15 (-25) mm long, 4-8 mm wide. Inflorescence lax and diffuse, leafy -hr acted, the angle between a pair of lateral peduncles ca. 85—110°. Uppermost bracts foliace¬ ous, resembling small leaves, 2.5— U mm long (Fig. 5). Flowers 3.4-5 mm wide. Sepals oblong -elliptic, 2. 7-3. 8 mm long, equaling petals but shorter than capsule. Stamens 5-10. Ovary 1 -locular ; styles 3; stigmas capitate. Capsules ovoid to ellipsoid, 3-5 mm long, 1.5-3 mm wide. Seeds light brown, 0.6-0.75 mm long. N = 8 (Hoar and Haertl 1932). Northeastern N. America, from Minn, to Newfoundland, south to N.Y., W Penna., N Ind. and E Iowa (disjunct to Tenn. ? — Sharp et. al. 1960), apparently limited to glaciated areas, in Wisconsin common in moist acid habitats as rocky or sandy shores, mud flats, acid tamarack bogs, alluvial marches, floating sedge mats, damp swales and sandy fields, as Sphagnum-— Cyperaceae wet meadows (Black River Falls, Jackson Co.) with scattered Larix, Drosera intermedia, Eriophorum virginicum, Muhlenbergia uniflora, Spiraea tomentosa and J uncus spp. Flowering mid- July to Septem¬ ber, fruiting late July to early October. 1970] Utech and litis — Flora of Wisconsin No. 61 343 Hybridizes in Wisconsin with H. canadense (pp. 26, 29-30; Fig. 6), these hybrids, called x. H. dissimulatum Bicknell (1913) and reported here for the first time for Wisconsin, represent the western-most station of this supposedly “unusually constant and recurring hybrid” (Fernald 1950). 9a. Hypericum boreale forma callitrichoides Fassett A sterile, submerged aquatic form with simple, flexible stems and small, 3-nerved leaves lacking pellucid-punctate dots, occa¬ sional in northern and central Wisconsin lakes (Adams, Bayfield, Jackson, Juneau, Langlade, Monroe and Oconto Counties) and grading shoreward into normal plants. 10. Hypericum majus (Gray) Britton Map 10, Fig. 5. Common St. John’s-Wort. Erect stout perennial with solitary or tufted stems, 1-4 (-6) dm tall, often with small leafy basal offshoots. Leaves commonly ascending, lanceolate or elliptic to oblanceolate or broadly acute to narrowly oblong, acute to obtuse, sub cordate-clasping, 5- to 7 -nerved, the larger 1.5-J+.U cm long, 5-12 mm ivide. Inflorescences well-defined, often compact, the angle between a pair of lateral peduncles only 25-50° ; bracts setaceous-subulate, 1-nerved, 1.8- 3.8 mm long (Fig. 5). Sepals lance-acuminate, 4.2-6. 5 mm long. Petals equaling sepals; stamens 15-20, weakly 5-delphous. Ovary 1-locular ; styles 3, 1-1.5 mm long; stigmas capitate. Capsules nar¬ rowly ovoid to ellipsoid, obtuse, reddish-purple at maturity, 5-7.8 mm long, 2-4 mm wide. Seeds pale brown, lustrous, reticulate, 0.6- 0.7 mm long. N = 8 (Hoar & Haertl 1932). Western and northeastern N. America, from British Columbia, E Wash, and Colorado to Quebec and Penna. (2 disjunct stations in Tenn.— Gillespie 1959; Sharp et. al. 1960), frequent throughout Wisconsin, chiefly in open, moist, gravelly, sandy or sometimes muddy habitats, as shores and beaches, low wet prairies, shrub- carrs, black spruce and tammarack bogs, Carex swales, moist talus and cracks of sandstone cliffs, spring and marly marsh margins, and weedy in pastures, sandy fields, roadsides and cranberry bogs. Flowering late June through September; fruiting earliest July to mid-October. Extremely variable in Wisconsin, especially as to size, hybridiz¬ ing not infrequently with H. boreale and H. canadense (cf. page 32). 344 Wisconsin Academy of Sciences , Arts and Letters [Vol. 58 11. Hypericum canadense L. Canadian St. John’s-Wort. Map 11, Figs. 5, 6. Slender erect annual or perennial herbs, 1-3 (-4.6) dm tall; stems unbranched except above, sharply 4-angled. Leaves linear- oblanceolate to linear, obtuse, sessile-attenuate, 1- or weakly 3 -nerved, 6-25 (-50) mm long, 1-3 mm wide. Inflorescences well- defined but open, the angle between a pair of lateral peduncles ca. 30-65° ; bracts subulate, 2-2.7 mm long (Fig. 5). Flowers 4-7 mm wide. Sepals linear-lanceolate, acuminate, 2. 5-4.4 mm long. Petals 2-5 mm long; stamens 5-10. Ovary 1-locular; styles 3, 0.7-1. 0 mm long; stigmas capitate. Capsules ovoid to conic, acute, reddish- purple at maturity, 3-5.2 mm long, 1.3-2. 6 mm wide. Seeds light yellow, reticulate, 0.5-0. 6 mm long. N 8 (Hoar & Haertl 1932). Eastern N. America, from Ga. to Ala. to Newfoundland, west to Iowa, the Black Hills and SE Manitoba, in Wisconsin mostly in the northern Driftless Area, in sandy-peaty roadsides, along railroads, wet sandy meadows, swales and marshes, moist sandstone ledges, in Sphagnum of Ericaceae-Cyperaceae bog (near City Point, Jack- son Co.) with Ledum groenlandicum, Chamaedaphne calyculata, Larix laricina, Picea mariana, Rhynchospora alba, Carex oligo- sperma, Solidago uliginosa, Eriophorum virginicum, Betula pumila and Aronia melanocarpa. Flowering early June to late August, fruiting mid- July to early October. Of the 30 collections from Wisconsin, only 6 date from before 1930 and these from only 2 stations (McLaughlin 1931) . The recent building of roads and flowages may have made the region’s glacial lake beds more receptive to botanizing and to the establishment of pioneers such as H. canadense and its hybrids with#, boreale and H. majus (cf. below for citations). HYBRIDIZATION IN HYPERICUM SECT. BRATHYS IN WISCONSIN: Four Wisconsin members of section Brathys (mutilum, boreale, canadense, majus) are morphologically and ecologically similar, often two or three growing together in the same station in Wiscon¬ sin’s Driftless Area, especially in the beds of glacial lakes (Maps 8-11), where all but the uncommon H. mutilum tend to hybridize. Hybrids are especially common in sandy, moist, flat, acid habitats. This region, at least post-glacially, has been a very suitable “open habitat” for the establishment of many Coastal Plain species (Mc¬ Laughlin 1932, Peattie 1922), such as Xyris spp., Bartonia vir- ginica, Gratiola lutea, Drosera spp., Rhynchospora spp., Helenium flexuosum (— H. nudiflorum) (Mickelson & litis 1966), and the microevolution of others (Johnson & litis 1963, pp. 267-8). 1970] Utech and litis — Flora of Wisconsin No. 61 345 Potential for long range dispersal in this small-seeded group is probably very great: all four species occur in Europe, probably introduced by birds or in fodder (Heine 1962, Tutin et al. 1968). Hypericum mutilum L. also occurs in Brazil (Keller 1908) and Hawaii (Doty & Mueller-Dombois 1966). Hybrids between these taxa are very common, both putative Fx’s and backcrosses, which is one reason for the great taxonomic diffi¬ culties in this group. In addition, dwarf forms of each species are common and especially difficult to distinguish. Hybrids, being inter¬ mediate morphologically, are only briefly described below. 12. Hypericum roreale (Britton) Bicknell x H. canadense L. Hypericum x dissimulatum Bicknell Fig. 6. Similar to H. canadense, but more lax and branched, with smaller, lanceolate-elliptic leaves and shorter, reddish-green cap¬ sules (Bicknell 1913). H. canadense is evidently a southern species, which post-glacially overlapped the northern H. boreale. The latter species, confined as it is to glaciated territory, has a most anoma¬ lous distribution and its Pleistocene survival or origins is not clear. It may represent a “stabilized hybrid” between H. mutilum, a wide- ranging eastern species, and H. majus, a western element, which is now limited to glaciated territory of northeastern North America. Jackson Co. : Indian Creek, sand flats ca. 1 mi W of City Point, 22 Sept. 1968, TJtech 68-200 (WIS), a mass collection, represented by the hybrid analysis scatter diagram of Fig. 6. 12b. Hypericum boreale ABrittonq Bicknell x H. majus (Gray) Britton Hybrids of these dissimilar species are erect but shorter than H. majus and with diffuse-branched inflorescences and short, ellip¬ tic-ovoid capsules like H. boreale. H. majus is clearly a cordilleran (Pacific Northwest ?) element which, like so many other taxa, in¬ vaded NE N. America post-glacially to hybridized with an east- tern vicarious element (Mason & litis 1965), in this case H. boreale (see above). Barron Co.: mud flat, edge of small lake, ca. 2 mi N of Turtle Lake, 21 Aug. 1956, litis et al. 7280 B (WIS). Juneau Co.: Sprague Flowage (T. 19 N., R. 2 E., Sec. 1), dry sandy-peaty, sedge-grass marsh, 23 Sept. 1967, litis et al. 25,851 pro parte, (WIS — mass collection). 12c. Hypericum majus (Gray) Britton x H. canadense L. Two very similar species form hybrids of shorter stature than H. majus and with leaves of intermediate shape, vein number and width; sepals and capsules also intermediary. The parental species 346 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 represent a vicarious pair (W. and E. North America) which post- glacially became sympatric in glaciated northastern North Amer¬ ica, where they hybridize freely. Clark Co. : Trow, moist ground, 15 July 1915, Goessl 1303 (MIL). Jackson Co.: temporary sandy margined pool, (T. 20 N., R. 1 W., Sec. 22), 22 Sept. 1968, Utech 68-300 (WIS — mass collection). Lincoln Co.: Wilson twp. (T. 35 N., R. 5 E., Sec. 4), ditch, 11 Sept. 1949, Seymour 10,968 (WIS). Monroe Co. : NW of Warrens, railroad ditch, 15 Sept. 1935, Fassett 17.686 (WIS). Oneida Co.: Rhinelander, Silver Bass Lake, on shore stump, 29 Aug. 1945, Hein 3U (WIS). Wood Co. : swale and desiccated pond, % mi W of Dexterville, 22 Sept. 1968, Utech 68- 100 (WIS — mass collection) ; Birch Bluff, S of City Point. (T 21 N. , R. 2 E., Sec. 19), 31 May 1958, litis & Koeppen 12,271a (WIS) . 13. Hypericum gentianoides (L.) B.S.P. Orange-grass, Pine Weed Map 13. Erect, strict, very slender annual 1-2.5 dm tall; stems, 4-angled, punctate, with numerous ascending filiform branches. Leaves subu¬ late, appressed, acute, sessile, 1.5-2. 8 mm long, 0.5 mm wide. In¬ florescence racemose ; flowers minute, nearly sessile. Sepals linear- lanceolate, 1.6-2. 2 mm long. Petals pale yellow, 2. 8-3. 6 mm long. Stamens. 5-10. Ovary 1-locular; styles 3, separate, 0. 6-0.9 mm long; stig¬ mas capitate. Capsules slenderly conic to lance-subulate, 4-4.8 mm long, 0.8-1. 5 mm wide. Seeds yellowish brown, obscurely areolate, O. 3-0.4 mm long. N = 12 (Hoar & Haertl 1932). Eastern United States (Maine to SW Ontario & Minn., south to Fla. and Texas), in open, dry, rock or acid outcrops, sand barrens and sand prairies in southern Wisconsin. At maturity, the entire plant turns copper or brick-color, hence called “orange grass.” Flowering mid-June to mid-September; fruiting late June to late September. 2. TRIADTNUM Raf. Marsh St. John’s-wort. Erect glabrous perennial, stoleniferous herbs with simple, oppo¬ site, entire, often pellucid-punctuate leaves. Flowers 5-merous, regular, perfect, hypogynous, small, in axillary and terminal cymules. Petals oblong, mauve or pinkish to greenish. Stamens 9, the filaments connate into 3 fascicles (3-delphous) and alternating with 3 conspicuous hypogynous glands. Ovary superior, completely 3-loculate; styles 3, separate, divergent; stigmas capitate. Capsules septicidal, 3-carpellate. Seeds small, short-cylindric, reticulate. Triadenum has 4 species in eastern North America and 1 in Asia (Japan, Korea, Manchuria, Ussuri & Amur, Triadenum japonicum 1970] Utech and litis — Flora of Wisconsin No. 61 347 (Bl.) Makino; Ohwi (1965)), which are often considered as sec¬ tion Elodea (Juss.) Choisy [non section Elodes (Adans.) Koch] of Hypericum. Segregated by Rafinesque (FI. Tell. 3: 78. 1836) on its pink petals and 3 hypogynous glands, it differs in addition (Holm 1906) by petals imbricate not convolute in bud, 9 stamens strongly 3-delphous into 3 fascicles alternating with 3 hypogynous glands, prominent veins repeatedly branched laterally to the blade mar¬ gins, and tuberous subterranean stolons with paired scale-like leaves usually with one, rather than many adventitous roots above each bud. Chromosome counts of N = 19 (Hoar & Haertl 1932) do not suggest a relationship to Hypericum , but rather to Crato- xylon, a pan-tropical tree genus. Key to Species A. Sepals elliptic to spatulate, summit obtuse to rounded, 2.8- 4.8 mm long; fruiting styles 0.6-1. 5 mm long; common Wis¬ consin marsh and bog plant. _ 1. T. FRASERI. AA. Sepals oblong to lanceolate, summit acuminate to acute, 4.3- 8 mm long; fruiting styles 2. 1-3.6 mm long; rare, central Wisconsin. _ 2. T. VIRGINICUM 1. Triadenum FRASERI (Spach) Gl. Marsh St. John’s-Wort. Hypericum Fraseri Spach May 13, Fig. 7. Hypericum virginicum L. var. Fraseri (Spach) Fernald Erect, glabrous, stoloniferous perennial herbs 2-6 dm tall, mostly reddish-purple in age; internodes terete, without decurrent lines. Leaves ovate-cordate to elliptic, 2. 3-6. 5 cm long, 1-3 cm wide, emarginate to obtuse, sessile and cordate-clasping. Cymules num¬ erous, terminal and axillary, few-flowered. Sepals 3 -Ip, 8 (-5.2) mm long, elliptic-oblong to spatulate, obtuse to rounded. Petals mauve or pink, oblong, 5.4-8 mm long. Stamens 9, 3-delphous, persistent; hypogynous glands 3, oval, orange. Ovary 3-celled ; styles 3, free, 0.6 -1.5 mm long at maturity; stigmas 3, capitate. Capsules conic- ovoid, 7-10 mm long, 3-5 mm wide, abruptly narrowed to styles. Seeds cylindric, dark brown, reticulate, 0.9-1. 1 mm long. Native of northeast North America (NE Nebraska & SW Mani¬ toba to Newfoundland & Labrador, south to Conn., Penna., N. Y., the mountains of W. Va., N. Ind. and Iowa), abundant in Wiscon¬ sin’s wet acid habitats, as tamarack-black spruce- leather-leaf- sphagnum bogs, sedge meadows, shrub carrs, sloughs and peaty marshes : in Comstock Marsh, Marquette Co., an extensive quaking bog with abundant Drosera rotundifolia, Sarracenia purpurea and patches of Phragmites communis, (Nee 1355, WIS) ; along Turtle Lake, Marquette Co., with Carex spp., Potentilla fruticosa and P. 348 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figure 7. Scatter diagram comparing' sepal and style length of Wisconsin Triadenum fraseri and Wisconsin and non- Wisconsin T. virginicum. 1970] Utech and litis — Flora of Wisconsin No. 61 349 palustris , Chamaedaphne calyculata, sedge meadow and adjoining thickets; in Vilas Co., along Lac Vieux Desert, in shallow acid waters of slow stream draining sphagnum bog with Larix laricina, Picea mariana, Ledum groenlandicum, Sarracenia purpurea, Cal- opogon pulchellus, Nuphar rubrodiscum, Utricularia, Hippurus, etc., (litis 18,149, WIS). Flowering from early July to latest August, fruiting mid- July till late September. 2. Triadenum virginicum (L.) Raf. Map 14, Fig. 7. Hypericum virginicum L. Similar to above, but sepals oblong-lanceolate, acuminate to acute, 4.8-8 mm long. Petals obovate, 6. 3-9. 8 mm long. Ovary 3- celled; styles 3, divergent, 2. 1-3. 6 mm long; stigmas capitate. Cap¬ sules ovoid-cylindric, 7.6-13.4 mm long, 3-5 mm wide, gradually tapering to styles. Seeds cylindric, dark brown, reticulate, 0. 9-1.1 mm long. N — 19 (Hoar & Haertl 1932). An Atlantic coastal plain element, Triadenum virginicum ex¬ tends inland to S. N. Y., S. Ontario, and disjunct to N. Ind. and central Wisconsin : Lincoln Co. : Merrill, dry sandy field along Prairie River with Comptonia peregrina, Hieracium aurantiacum and Robinia pseudoacacia, 19 Aug. 1956, liven 419 (WIS). Bagga Marsh, among cranberry beds along Copper River, 25 Aug. 1957, Schlising & PeroutJcy 660 (WIS). Wood Co.: burned over sphag¬ num bog, 5 mi NW of Babcock, 20 Aug. 1937, Catenhusen s.n. (WIS). Biron township, Huffman farm, 16 July 1953, Dana s.n. (WIS). It is noteworthy that all collections are recent, since 1937. In Wisconsin, T. Fraseri is quite common and easily separatable from both Wisconsin and non- Wisconsin T. virginicum (Fig. 7). Taxonomically and morphologically, Gleason (1947) distinguished them on sepal and style length. Fernald (1936), using var. Fraseri to indicate this difference, notes that, where sympatric, they usually show clear segregation into a southern or lowland and a northern or upland series. Bibliography Adams, P. 1959a. The status of Hypericum proliflcum. Rhodora 61:250-251. - . 1959b. The taxonomy of Hypericum Section Myriandra ( Hyper - icaceae). Ph.D. Dissertation, Harvard Univ., Cambridge, Mass. - . 1962. Studies in the Guttiferae. I. A. synopsis of Hypericum Section Myriandra. Gray Herb. No. CLXXXIX, 1-50. Harvard Univ., Cambridge, Mass. Bartholomew, E. A. 1949. The Family Hypericaceae in West Virginia. Castanea 14:102-109. Bicknell, E. P. 1895. Hypericum boreale (Britton) and related species. Bull. Torr. Bot. Chib 22(5) :211-215. 350 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 — - . 1913. The ferns and flowering plants of Nantucket — XI. Bull. Terr. Bot. Club 40(11) : 610-611. Clohisy, M. 1930. Wisconsin Wild Flowers. Milwaukee. 122 pp. Curtis, J. T. 1959. The Vegetation of Wisconsin. Univ. of Wisconsin Press, Madison. Darlington, C. D. and A. P. Wylie. 1955. Chromosome Atlas of Flowering Plants . George Allen & Unwin, Ltd., London. Doty, M. S. and D. Mueller-Dombois. 1966. Atlas for Bioecology in Hawaii Volcanoes National Park. Hawaii Bot. Science Paper No. 2, Univ. of Hawaii, 507 pp. Fassett, N. C. 1939. Notes from the Herbarium of the University of Wiscon¬ sin — XVII. Elatine and other aquatics. Rhodora 41:376. - . 1960. A Manual of Aquatic Plants , 2nd ed. Univ, of Wisconsin Press, Madison. Fernald, M. L. 1936. Plants from the Outer Coastal Plain of Virginia. Rhodora 44:434-435. - . 1950. Gray’s Manual of Botany, ed. 8. New York: American Book Co. Gillespie, J. 1959. The Hyperiaceae of Tennessee. Castanea 24:24-32. Gleason, H. A. 1947. Notes on some American plants. Phytologia 2:288-291. - and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. New York: von Nostrand Co. Guire, K. E. and E. Voss. 1963. Distribution of distinctive shoreline plants of the Great Lakes region. Mich. Bot. 2:99-114. Hartley, T. G. 1962. The Flora of the Driftless Area. Ph.D. Dissertation, Univ. of Iowa, Iowa City. Hegi, G. 1925. Illus trier te Flora von Mittel-Europa. Munchen: J. F. Lehmann Verlag, Vol. 5, I. Heine, H. 1962. Les Millepertuis americains da'ns la flore d’Europe. Bauhinia 2(1) : 71-78. Hoar, C. S. 1931. Meiosis in Hypericum punctatum. Bot . Gaz. 92:197-204. - - and Haertl. 1932. Meiosis in the genus Hypericum. Bot. Gaz. 93: 197-204. Holm, T. 1906. Art. XXXVI. Triadenum virginicum (L.) Rafin. A morpho¬ logical and anatomical study. Amer. J. Sci. IV, Vol. XVI, No. 95:369-376. Johnson, M. F., and H. H. Iltis. 1963. Preliminary Reports on the Flora of Wisconsin. No. 48. Compositae I — Composite Family I (Tribes Eupa- torieae, Verno'nieae, Cynarieae and Cichorieae) . Trans . Wis. Acad. Sci. Arts and Letters, Vol. 52:255-342. Jones, G. N. and G. D. Fuller. 1955. Vascular Plants of Illinois. Univ. of Illinois Press, Urbana. Keller, R. 1908. Zur Kenntnis der Sectio Brathys des Genus Hypericum. Bull, de Vherbier Boissier ser. 2, 8:175-191. Kingsbury, J. M. 1964. Poisonous plants of the United States and Canada. Englewood Cliffs: Prentice-Hall, Inc. 626 pp. Lakela, O. 1965. A Flora of Northeastern Minnesota . Univ. of Minnesota Press, Minneapolis. Marsh, C. D. and A. B. Clawson. 1930. Toxic effect of St. Johnswort (Hy¬ pericum perforatum) in cattle and sheep. U.S.D.A. Tech. Bull . No. 202: 1-23. Mason, C. T. and H. H. Iltis. 1965. Preliminary Reports on the Flora of Wisconsin. No. 53. Ge'ntianaceae and Menyanthaceae — Gentian and Buck- bean Families. Trans. Wis . Acad . Sci. Arts and Letters, Vol. 54:295-329. 1970] Utech and litis — Flora of Wisconsin No. 61 351 McLaughlin, W. T. 1931. Preliminary Reports on the Flora of Wisconsin. No. 14. Hypericaceae. Trans. Wis. Acad. Sri. Arts and Letters , Vol. 26:281- 288. - . 1932. Atlantic Coastal Plain Plants in the Sand Barrens of North¬ western Wisconsin. Ecol. Mono. 2:335-383. Mickelson, C. J. and H. H. Iltis. 1966. Preliminary Reports on the Flora of Wisconsin. No. 55. Compositae IV — Composite Family IV (Tribe Helenieae and Anthemideae) . Trans. Wis. Acad. Sci. Arts and Letters, Vol. 55: 197-222. Mulligan, G. A. 1957. Chromosome numbers of Canadian weeds. I. Can. J. Bot. 35:779-789. Nielsen, N. 1924. Chromosome numbers in the genus Hypericum. Hereditas 5:378-382.. Ohwi, J. 1965. Flora of Japan. Smithsonian Institution, Washington. 1067 pp. Peattie, D. C. 1922. The Atlantic coastal plain element in the flora of the Great Lakes. Rhodora 24:57-88. Robson, N. K. B. and P. Adams. 1968. Chromosome numbers in Hypericum and related genera. Brittonia 20:95-106. Seymour, F. C. 1960. Flora of Lincoln County Wisconsin. P. F. Nolan, Taun¬ ton, Mass. Sharp, A. J., R. E. Shanks, H. L. Sherman and D. H. Norris. 1960. A Pre¬ liminary Checklist of Dicots of Tennessee. Univ. of Tennessee, Knoxville. 114 pp. (ms). Sorensen, P. D. 1966. The Flora of the old bed of Glacial Lake Wisconsin and the adjacent terminal moraine. MS thesis, Univ. of Iowa, Iowa City. (ms). Svenson, H K. 1940. Plants of Southern United States. III. Woody species of Hypericum. Rhodora 42:8-19. Swezey, G. D. 1883. Catalogue of the phaenogamous and vascular cryptogam- ous plants of Wisconsin. In Chamberlain’s: Geology of Wisconsin, Vol. 1: 376-395. Tutin, T. G., et al. 1968. Flora Europaea. Vol. 2. Rosaceae to Umbelliferae. London: Cambridge Univ. Press. 455 pp. PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 62. COMPOSITAE VI. COMPOSITE FAMILY VI. THE GENUS AMBROSIA — THE RAGWEEDS'-1 2 Willard W. Payne Ambrosia (including Franseria) is one of several genera of the Compositae which are distinguished by exceptional modification for wind pollination or anemophily. All of these genera (Iva, Euphro- syne, Dicorea, Hymenoclea and Xanthium, in addition to Ambrosia) are characterized by similarities in vegetative morphology, chem¬ istry, cytology, and pollen and inflorescence structure that suggest evolutionary affinity. They have been variously classified as a dis¬ tinct family, the Ambrosiaceae (Small, 1913; Rydberg, 1922), as a distinct tribe, the Ambrosieae (Cassini, 1834; Delpino, 1871; Ben¬ son, 1957; Payne, Raven and Kyhos, 1964), and as a subtribe, Ambrosiineae, of the tribe Heliantheae (Bentham, 1873; Fernald, 1950; Cronquist, 1952; Keck, 1959; Solbrig, 1963). There is grow¬ ing evidence to support the derivation of this evolutionary group from helianthoid progenitors, of particular interest in this respect being the pollen wall ultrastructural studies of Larson and Skvarla (1966) . At the same time, however, these plants have certain chem¬ ical and structural attributes more characteristic of the Anthe- mideae (Payne, 1963; Miller, 1967), and it is possible that they have been derived from a progenitor group intermediate between the Anthemideae and Heliantheae. Because of this, and taking into ac¬ count the array of morphological features common to all of the genera, I believe they are best treated, for the present at least, as members of the tribe Ambrosieae. Ambrosia is distinguished from other closely related genera of the Ambrosieae by capitulescences of wholly staminate and wholly pistillate capitula, the staminate with saucer-shaped, lobed in¬ volucres of connate phyllaries, the pistillate with hard, spiny, bur- like fruiting involucres. The usual arrangement of head types is that in which staminate capitula are produced in racemose or 1 This study has been supported in part by a grant (GB 5558) from the National Science Foundation. 2 1 wish to gratefully acknowledge the assistance and advice of Dr. Hugh litis, Michael Nee and Roger Thibault of the Herbarium of the University of Wisconsin, Madison. Specimens which were examined for this study were kindly made available by the curators of the following herbaria : G, IDL, ISC, KY, MICH, MIL, MSC, NY, SMU, TENN, UWM, WIS. 353 354 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 spicate groups at the tips of the main stems and branches, while pistillate capitula are clustered in the axils of leaves and bracts below the staminate racemes (see Figs. 1, 2, 3). Close similarity and extreme floral reduction in Ambrosia, Xanthium and Hymeno- clea suggest that these genera constitute the terminal evolutionary branch of the Ambrosieae, with Xanthium and Hymenoclea inde¬ pendently derived from primitive forms of Ambrosia. The genus Ambrosia is predominantly American ; approximately 31 species are native to North America, eight to South America; two species, closely allied to and probably derived from American progenitors, are found in the Old World. All available evidence sug¬ gests that the genus originated in and diversified from arid and semi-arid regions in southwestern North America, where primitive species are still abundant (Payne, 1964). All of the ragweeds are found in open habitats. Primitive, shrubby species occupy natural sites in deserts and semi-desert areas, being particularly adapted for stream banks, exposed arroyos, and the like. Such species produce fruiting involucres with many straight or hooked spines (often identical to those of the cockleburs) and are adapted for animal dispersal. In addition, involucres of the majority of the species commonly fall into streams or are carried to streams by runoff water, and dissemination by flowing water is probably important in their local distribution. Advanced species of ragweeds occur most abundantly today as weeds in association with man. The physiological characteristics that have allowed prim¬ itive members to survive in open and primary sites in arid regions are undoubtedly those which have been refined to permit derived species to successfully exploit pioneer habitats created by the agri¬ cultural and urban practices of present American cultures. Fruit¬ ing involucres of most advanced, weedy species have few or no vestigial spines, and the reduced spines appear to play no signifi¬ cant role in fruit dispersal (Gebben, 1965). When individuals grow along streams or river banks, or when they occur on slopes, water flow is probably important for local down-stream or down-slope transport of fruits and seeds. Studies of introduced populations of the short ragweed (A. artemisiifolia) in Oregon (G. H. Moose, personal communication) indicate that seeds of this species may be carried short distances by water flow, and temporary colonies of the species often flourish on sand bars and flood plains. I have ob¬ tained evidence from experiments with a captive English sparrow (Payne, 1962) that a small proportion of ingested seeds of A. artemisiifolia can pass unharmed through birds’ alimentary canals and still germinate. In addition to these factors man is a potent, long-distance transport agent for species that occur as weeds of cultivation. Regardless, most seeds of derived species fall and re- 1970] Payne — Flora of Wisconsin No. 62 355 Figure 1. Ambrosia psilostachya DC. A. Capitulescence. B. Habit silhouette showing several young, adventitious shoots developed from runner-like root. C. Representative staminate heads. D. Pale. E. Representative fruiting in¬ volucres with vestigial spines or without spines. F. Leaf silhouettes from rep¬ resentative specimens; each silhouette from different specimen, all from median, cauline nodes. 356 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figure 2. Ambrosia artemisii folia L. A. Capitulescence. B. Representative staminate heads. C. Pale. D. Representative fruiting involucres. E. & F. Heteroblastic leaf series representing leaves of two frequently encountered forms; lowest nodes toward left. G. Population sample, each leaf from a different specimen; all leaves from node five above cotyledonary node. 1970] Payne — Flora of Wisconsin No. 62 357 Figure 3. Ambrosia trifida L. A. Silhouette of seedling. B. Capitulescence. C. Staminate head; note distal striations. D. Pale. E. Representative fruiting in¬ volucres. F. Representative leaf silhouettes. 10 mm 858 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 main in proximity to the parent plants, where they augment seed reserves present in soils that have previously supported the species. All of these factors are important in understanding the develop¬ ment and nature of the population structure of advanced species. The three ragweeds found in Wisconsin are among the most specialized in the genus. They are widespread in the eastern and central United States (Maps 2, 4, 6) , and are well known as sources of allergenic, air-borne pollen that constitutes the most serious natural air pollutant in North America, being the major cause of “hay fever” (cf. Wagner, 1959). An outstanding feature of all Wisconsin ragweed species is their morphological variability. Al¬ though this is particularly striking when the species are examined throughout their ranges, variability is also a pronounced attribute of the members of genetically restricted populations, such as the progeny grown from seeds developed by a single, self-pollinated plant (Jones, 1936). 1 have suggested (Payne, 1962, 1965) that for the short ragweed, heterogeneity may have been originally de¬ veloped or strongly reinforced by fusion (“subspecies amalgama¬ tion”) of once more distinct, subspecific groups. This hypothesis is compatible with present species structure and with the probable Pleistocene and post-Pleistocene history of the species. It may also provide at least a partial explanation for the similar attributes of Wisconsin perennial ragweed (A. psilostachya) and giant ragweed (A. trifida). Although this paper deals specifically with only the ragweeds of Wisconsin, there is presently no treatment covering other genera of the Ambrosia tribe (Ambrosieae) of the Compositae with which the ragweeds are often confused. Therefore, the keys below include all of the members of the Ambrosieae found in the state. Treatment of the genus Iva follows that of Jackson (1960), and of the genus Xanthium that of Love and Dansereau (1959). AMBROSIEAE — Wind pollinated shrubs, perennial herbs and annuals distributed principally in desert and disturbed habitats in the Americas. Leaves alternate to opposite, usually petiolate, often lobed. Pubescence various but always including simple, uniseriate trichomes which are dead at maturity, and biseriate, glandular colleters. Capitulescences paniculate to racemose or spicate, typ¬ ically maturing acropetally ; capitula paleaceous, perfect or unisex¬ ual, often nodding. Perfect heads with few, free or connate phyl- laries, sterile disc florets, and fertile ray florets. Unisexual heads usually with connate phyllaries (phyllaries lacking in staminate heads of Xanthium), the phyllaries of pistillate heads united to form winged or spiny burs with prominent beaks through which the stigmatic lobes project. All florets reduced and specialized for 360 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 anemophily; pappus vestigial or lacking; pistillate florets with reduced corollas or the corollas wanting, without androecia, the stigma lobes elongate and minutely papillose ; staminate florets with anthers weakly connate and often separating during anthesis, lack¬ ing ovaries, and with capitate, penicillate pistillodia which elongate after anthesis to push pollen from the anther cylinders. Pollen oblate, tricolporate, the colpae mostly vestigial, echinulate, cavate, the cavae enlarging to form bladder-like chambers by pronounced shrinkage of the protoplast and invagination of the inner wall layers of the grains after shedding. Base chromosome number .r =18. Key To Wisconsin Ambrosieae a. Staminate and pistillate florets in common heads; ray florets fertile, disc florets sterile (IV A) _ _ b. b. Plants annual; phyllaries free _ c. c. Leaves ovate, coarsely serrate; heads subtended by prom¬ inent bracts ; phyllaries 3-4 _ _ /. annua L. c. Leaves subcordate to ovate, usually coarsely lobed and toothed; heads ebracteate; phyllaries 5 _ _ _ I. xanthifolia Nutt. b. Plants perennial; phyllaries basally connate _ _ _ _ _ _ I. axillaris Pursh a. Staminate and pistillate florets borne in separate heads _ d. d. Staminate heads lacking phyllaries, pistillate heads 2-flow- ered and with many, hooked spines (XANTHIUM) - e. e. Leaves pinnately lobed ; stems bearing long, golden, three- rayed, axillary spines _ —X. spinosum L. e. Leaves coarsely palmately lobed; stems unarmed _ _ X. strumarium L. d. Staminate heads with involucres of connate phyllaries, pistil¬ late heads 1-flowered and with few vestigial spines or none _ _ (AMBROSIA) See following key. Genus 15. AMBROSIA L. Ragweed. In Wisconsin, perennial or annual herbs with petiolate, lobed leaves opposite below or throughout. Staminate heads in terminal, racemose clusters, nodding on short stalks, many-flowered, pale¬ aceous, with few-lobed involucres of connate phyllaries. Pistillate heads clustered in axils of upper leaves, sessile, 1-flowered, tur¬ binate, with few sharp or blunt spines localized near the beaks or without spines. 1970] Payne — Flora of Wisconsin No. 62 361 Key To Species A. Leaves pinnately or bipinnately lobed or parted; staminate in¬ volucres lacking dorsal striations; upper cauline leaves usually alternate _ B. B. Plants perennial with horizontal runner-like underground roots ; involucral spines blunt or absent ; leaves usually coarsely lobed _ 1. A. psilostachya. B. Plants annual with taproots; involucral spines usually sharply pointed; leaves usually delicately lobed and parted _ 2. A. artemisiifolia. A. Leaves palmately lobed or unlobed; staminate involucres marked with dorsal striations; all cauline leaves usually oppo¬ site; plants annual _ 3. A. trifida. 1. Ambrosia psilostachya DC. Prod. 5: 536. 1836. (non A. psilo¬ stachya Grisebach. 1861.) Perennial Ragweed, Western Ragweed (Maps 1, 2; Figs. 1, 4P.) Type: Berlandier 2280 G; Isotype NY. A . hispida Torr. Ann. Lyc. Nat. Hist. N.Y. 2: 216. 1828. (non A. hispida Pursh. 1814). A. coronopifolia T. & G. FI. N. Am. 2: 291. 1842. A. glandulosa Scheele, Linnaea 22: 157. 1849. (non A. glandulosa Rydb. 1922). A. lindheimeriana Scheele, lx. 22: 157. 1849. A. coronopifolia var. asperula Gray, Bost. Jour. Nat. Hist. 6: 226. 1857. A. coronopifolia var. gracilis Gray, lx. 6: 227. 1857. A. psilostachya var. lindheimeriana Blank. Rep. Mo. Bot. Gard. 18: 173. 1907. A. calif ornica Rydb. N. Am. FI. 33: 20. 1922. A. psilostachya asperula (Gray) Blank, ex Rydb. lx. 33: 19. 1922. A. psilostachya calif ornica (Rydb.) Blake, in I. Tidestrom. FI. Utah & Nev. 580. 1925. A. psilostachya var. coronopifolia (T. & G.) Farwell ex Fern. Gray’s Man. Bot., ed. 8, 1470. 1950. Erect, perennial herb, 0.5-10 dm high; proliferating from run¬ ner-like roots. Stems unbranched or branched; pubescent, hirsute to pilose to hispidulous, minutely glandular; light green to yellow¬ ish, occasionally blotched or suffused with red. Leaves opposite below, alternate above, occasionally opposite nearly to staminate portion of capitulescence. Median cauline leaves short-petiolate to subsessile, the petiole usually broadened with decurrent blade tissue. Blade ovate to ovate-lanceolate in outline; pinnately to bi¬ pinnately lobed (rarely nearly unlobed), lobes with entire margins or sparsely serrate; lamina somewhat coriaceous, often densely 362 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 gray-green pubescent ; veins prominent on under surface ; pilose to scabrous. Capitulescence usually little branched or unbranched, gradually blending with vegetative portion of axis. Staminate heads paleaceous ; 10-40-flowered ; stalked to subsessile ; ebracteate. Staminate involucre campanulate, often prominently eccentric ; shallowly toothed, the distal teeth usually larger; persistent after anthesis; pubescent, often minutely glandular-punctate. Staminate florets narrowly campanulate; corolla hyaline, hve-lobed. Pistil¬ late capitula sessile ; often borne singly in axils of bracts and leaves subtending staminate raceme, usually clustered. Fruiting involucre obovate; spines 0-7, terete, usually bunt, often lacking; body to 6 mm long and 3.5 mm broad, reticulate-rugose, rarely striated ; beak short, blunt to vestigial. Haploid chromosome number, n = (18), 36, 54, 72. Ambrosia psilostachya is the least abundant of the Wisconsin ragweeds. It is a plant of sandy soils, commonly found on sandy prairies, often along ancient lake shores, sandy, glacial outwashes, and near the Great Lakes. Typical collection data include refer¬ ences to sandy beaches, sandblows, sandy prairie openings, and dry, sandy, open, upland forests. It is distributed fairly generally through Wisconsin (Map 1) but because of its habitat preferences is seldom abundant in comparison with short and giant ragweeds, and it is of minor economic importance as a weed of cultivated land. Reproduction is principally vegetative, and individuals well adapted for a particular site commonly produce large, clonal popu¬ lations. The tendency for vegetative reproduction is correlated in this species with production of smaller, fewer-headed capitules- cences. Comparative seed production by individual, well-grown shoots, raised in experimental garden plots at the University of Michigan Botanical Gardens in 1960, gave the following results: A. artemisiij olia (annual)- — 38,800 fruits; A. trifida (annual) — 4,700 fruits; A. psilostachya (perennial) — 80 fruits. In contrast, 24 perennial ragweed seedlings planted in similar plots in 1960 produced 3,175 vegetative shoots the succeeding spring (Payne, 1962). This difference in capitulescence development among the species indicates that, even where relatively abundant, A. psilo¬ stachya is a comparatively minor contributor to the atmospheric ragweed pollen load. Like the other Wisconsin ragweeds, A. psilostachya is quite vari¬ able. This morphological variability is probably related, at least in part, to the presence in this species of a polyploid series, although no close correlation between particular ploidal levels and particular morphological expressions has yet been demonstrated. While Wis¬ consin perennial ragweed has not been intensively investigated 1970] Payne — Flora of Wisconsin No . 62 363 cytologically, the most common ploidal level is probably n — 36 (tetraploid, based upon x ~ 18). Octoploid in — 72) and diploid (n = 18, this count not yet verified in my laboratory) plants have been reported only from California and eastern Texas, respec¬ tively. Hexaploid plants (n = 54) are common west and south of Wisconsin, and it is possible that this ploidal level is. represented in the state. Studies of the comparative sesquiterpene lactone chemistry of A . psilostachya clones and populations, currently in progress, indicate that the morphological variability of this species is paralleled by chemical variability (Miller, 1967 ; Miller, et al , 1968). 2. Ambrosia artemisiif olia L. Sp. PL 2: 988. 1753. Short Rag¬ weed, Common Ragweed. (Maps 3, 4; Figs. 2, 4A.) Lectotype : Linnaeus 1HU-U. LINN. (I select this specimen from the Linnaean Herbarium as lectotype because it is representative for the species, is clearly labeled aartemisifolian (sic) by Linnaeus, and is readily available as a photograph in the microfilm edition of the Linnaean Herbarium by the International Documentation Center AB, Tumba, Sweden.) A. elatior L. Sp. PL 2: 987. 1753. Iva monophylla Walt. FL Carol. 232. 1788. A. elata Salisb. Prodr. 175. 1796. A. simplicifolia Raeusch. Nomen. Bot. 274. 1797. A. absynthifolia Michx. FL Bor. Am. 2: 183. 1863. A. paniculata Michx. lx. 2 : 183. 1803. A. heterophylla Muhl. ex Willd. Sp. PL 4 : 287. 1805. A. artemisifolia elatior (L.) Desc. FL Ant. 1: 239. 1821. A. longistylis Nutt. Trans. Am. Phil. Soc. N.S. 17 : 344. 1841. A. artemisiif olia vars. a, p, 8, y. T. & G. FL N. Am. 2: 291. 1842. A. artemisiif olia L. a quadricomis Ktze. Rev. Gen. PL I. 305. 1891. A. artemisiif olia L. p octocornis Ktze. lx. 305. 1891. A. artemisiae folia ssp. diversifolia Piper, Contr. U. S. Nat. Herb. 11: 551. 1906. A. artemisiae folia L. var. paniculata (Michx.) Blank. Rep. Mo. Bot. Gard. 18: 173. 1907. A. media Rydb. Bull. Tor. Bot. Club, 37 : 127. 1910. A. elatior L. var. artemisiif olia (L.) Farw. Rep. Mich. Acad. 15: 190. 1913. A. elatior L. var. heterophylla (Muhl.) Farw. lx. 190. 1913. A. diversifolia (Piper) Rydb. N. Am. FL 33: 18. 1922. A. monophylla (Walt.) Rydb. lx. 17. 1922 A. artemisiif olia var. elatior f. villosa Fern. & Grisc. Rhodora 37: 185. 1935. 364 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Figure 4. Leaf silhouettes representative for the Wisconsin ragweeds A. artemisii folia (A), A. psilostachya (P), A. trifida (T), and their hybrid progeny. Erect, annual herb, 1-7 (-20) dm high. Stems unbranched to much branched ; glabrous to villous, often streaked with reddish to black longitudinal markings. Leaves petiolate, opposite below, alternate above. Lamina of median cauline leaves pinnatifid to tri- pinnatifid, frequently less lobed above and below, blades of upper¬ most cauline leaves occasionally unlobed; margin sparsely serrate, serrations blending with lobes; ovate in outline. Capitulescence much branched, occasionally wholly or predominantly staminate or pistillate, more or less abruptly differentiated from vegetative stem. Staminate heads paleaceous; 10-100 (-200) flowered; stalked, stalks to 4.5 (-15) mm long, seldom bearing more than a 1970] Payne — Flora of Wisconsin No. 62 865 single, terminal head; ebracteate. Staminate involucre campanu- late to cupulate to flattened; lobed, with 5-10 short-deltoid, mar¬ ginal lobes, the sinuses normally extending less than % the distance to the stalk, terminal lobes usually largest; unmarked above; sparsely pubescent, sometimes obviously glandular; remaining at¬ tached after anthesis is complete. Pales linear, usually with mar¬ ginal or terminal, simple or glandular trichomes. Staminate florets narrowly campanulate; corolla hyaline, 5 lobed; stamens 5, the tips of the claws usually long-attenuate; anthers connate. Pistil¬ late heads sessile, 1 flowered; clustered in axils of bracteal and foliage leaves below the staminate racemes; all but the terminal one bracteate. Fruiting involucres obovate, the body to 2.5 mm broad and 3.5 mm long, with one beak to 2 mm long; supplied with 4-12 straight, terete spines, to 1 mm long (rarely longer, or com¬ pletely vestigial and lacking), arranged in one or a few close whorls above center; glabrous or with few to abundant, short to long hairs, frequently glandular punctate, particularly below the beak; somewhat rugose; commonly with reddish-brown to black mottling or striations, sometimes suffused with pigment. Haploid chromosome number, n — 18. Short ragweed is by far the most abundant and economically important of the ragweed species of Wisconsin. It is distributed generally throughout the state (Map 3), being uncollected from only a few counties in the northwestern and west-central portions of Wisconsin, where the species begins to reach the northern limits of its range at these longitudes. It is most frequently a weed of cultivated and ruderal habitats, roadsides, railroad embankments, and similar sites, and may also become abundant in overgrazed and sterile soils. Because urbanites most often encounter A. arte- misiifolia as a roadside weed, it is popularly believed that road¬ side habitats support a major proportion of the plants in an area, and are, thus, the sources for most ragweed pollen production; this belief is defended by manufacturers of chemicals used to spray roadsides in weed control programs. However, three studies car¬ ried out in southeastern Michigan in 1958, 1959 and 1960 (Har¬ rington, et ah, 1960; Gebben, et al., 1962; Gebben, 1965) clearly demonstrate that, in areas encompassing both rural and urban land use, the great majority of plants are found in cereal grain fields, of which wheat fields lead the list. An outstanding attribute of this species is its variability, usually with considerable distance between extremes in expression of virtu¬ ally all characters (see, for example, Fig. 2, D and G). It is prob¬ able that genomes and populations capable of producing variable progeny are selected and maintained in this species. Such a mech- 366 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 anism enhances the ability of the species to survive with man by providing a continually varying supply of genotypes, some of which are capable of exploiting nearly any available primary site. The selective advantage for such a mechanism is obvious when one considers the variable nature of man-created primary habitats, viz., those associated with crop rotation, frequent interruption of dif¬ ferent successional stages in different ways, and so on. A possibly significant factor in this mechanism has to do with seed longevity. The longevity experiments initiated by Beals (Darlington, 1922) and others demonstrate that seeds of A. art emisii folia may remain viable in the soil for periods of 40 years or more. Thus, it is possi¬ ble for plants developed from seeds produced the preceding season to interbreed with plants developed from seeds produced many ragweed generations in the past, bringing together and intermix¬ ing genomes from populations which may have been selected under quite different site conditions. Similar factors influencing popula¬ tion dynamics are probably also important in the biology of A. psilostachya and A. trifida . 3. Ambrosia trifida L. Sp. PI. 2: 988. 1753. Giant Ragweed, Great Ragweed, Horse Cane. (Maps 5, 6; Figs. 3, 4T.) Lectotype: Linnaeus 1114—1. LINN. (As for the previous species treated, this specimen is appropriately labeled and preserved, and a photograph is available from the International Documentation Center.) A. simplicif olia Walt. FI. Carol. 231. 1788. A. integrifolia Muhl. ex Willd. Sp. PI. 4: 375. 1805. A. aptera DC. Prod. 5: 527. 1836. A. trifida var. integrifolia (Muhl. ex Willd.) T. & G. FI. N. Am. 2: 290. 1842. A. trifida L. var. /3 texana Scheele, Linnaea 22: 156. 1849. A. trifida L. a normalis Ktze. Rev. Gen. PL I. 305. 1891. A. trifida L. a normalis var. aptera Ktze. lx. 305. 1891. A. trifida L. a normalis var. heterophylla Ktze. lx. 305. 1891. A. striata Rydb. Brittonia 1 : 96. 1931. A. variabilis Rydb. lx. 97. 1931. A. trifida L. f. integrifolia (Muhl.) Fern. Rhodora 40: 347. 1938. A. trifida var. polyploidea Rousseau, Nat. Canad. 71 : 215. 1944. A. trifida var. trifida Cronquist, Rhodora 47 : 396. 1945. Erect, annual herb, 2-50 dm high. Stems unbranched to much branched ; hispid-hispidulous to scabrous or nearly glabrous, some¬ times tuberculate; somewhat angular and ridged; often with fine, black longitudinal striations, frequently suffused or blotched with red. Leaves opposite throughout or becoming alternate in the ca- 1970] Payne — Flora of Wisconsin No. 62 367 pitulescence ; petiolate, the petiole often more or less winged with decurrent blade tissue. Blade ovate-lanceolate to broadly ovate or ovate-deltoid in outline; upper, bracteate blades often becoming narrowly lanceolate; unlobed or palmately lobed, the 3-7 primary lobes occasionally bearing pinnately arranged secondary lobes ; mar¬ gin serrate, the abaxial surface of each tooth often with a single, black striation; adaxial surface hispidulo-scabrous. Capitulescence little or much branched. Staminate heads minutely paleaceous ; 10- 125 flowered; stalked, stalks 2-8 mm long, rarely subsessile; ebracteate. Staminate involucre shallowly campanulate to saucer¬ shaped; crenulate or toothed at the margin, the three distal teeth usually more pronounced and marked on the upper surface with prominent, black striations; abcissing after anthesis is complete. Staminate florets narrowly campanulate; corolla hyaline, five- lobed, marked with longitudinal striations; stamens with short- attentuate claws. Pistillate capitula sessile, 1 flowered ; clustered in axils of bracteal leaves below the staminate racemes; all but the terminal bracteate. Fruiting involucres obovate; spines terete to radially flattened, 3-11 in one or a few close-set whorls below the beak; body to 17 mm long and 10 mm broad usually ridged and somewhat rugose, frequently marked with black or red, somewhat pubescent. Haploid chromosome number n 12. Giant ragweed is the second most abundant species in Wisconsin, being distributed principally in the southern two-thirds of the state (Map 5). Although ordinarily less abundant regionally than short ragweed in terms of absolute numbers of specimens the greater stature and larger capitulescences make it a heavy pollen producer, and in areas of abundance it may contribute as much or more pollen to the local atmospheric pollen load. It is essentially a floodplain species and is most abundant in moist soils of drainage ditches, low fields, open stream banks, and the like. Like short ragweed, A. trifida is quite variable, outstanding vari¬ ability being associated with fruit size and shape, leaf shape, and pubescence. In the southern and southwestern United States the fruiting involucres are often scarcely larger than those of A. arte- misiifolia, while in the Appalachian region fruits 10-20 times this size may be found (Payne and Jones, 1962). Similar fruiting in¬ volucre variation is common in local populations, and may also apply to different fruits taken from the same plant. Forms with unlobed leaves, or with both lobed and unlobed leaves are common, this aspect of leaf morphology being partially related to ecological conditions. Such plants have played a prominent role in the tax¬ onomic history of the species, generating such epithets as simplici- folia, integrifolia, and variahilis. It is probable that, as with short ragweed, greatly increased population size, mixing of geographic 368 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 races once more distinct, and variable selection pressures associated with modern agricultural practices and urbanization, have con¬ tributed to local variability. The black striations distributed on all parts of the plant are re¬ lated to sub-epidermal canals (resin canals?) which contain a deep red pigment. When cut, the plant bleeds red “blood,” a fact that figured prominently in the “doctrine of signatures” medical prac¬ tices of some American Indians, and which actually led to reverence and fear of the plant by certain tribes (Payne and Jones, 1962) . The most unusual characteristic of A. trifida is its chromosome compliment of n — 12. With the usual base of n — 18 in the genus, this presents the possibility of a progenitor genome of n — 6. Pair¬ ing behavior studies by K. L. Jones (1943) of the hybrid A. arte - misiifolia X trifida have demonstrated, however, that the 12 chromosomes of giant ragweed are homologous with the 18 chromo¬ somes of short ragweed, and the chromosome complement of A. trifida is interpreted to be the result of aneuploid reduction (Payne, Raven and Kyhos, 1964). Furthermore, no similar compliment is known elsewhere in the Ambrosieae, although aneuploid reduction to n — 16 or 17 is common in the genus Iva, and in A. bidentata n — 17. Polyploidy in A. trifida was suggested by Rousseau (1944), but was refuted by the studies of Payne and Jones (1962). To my knowledge, no bona fide report of polyploidy exists for the species. Ecological Relationships and Specialization of Wisconsin Ambrosia The ragweed species described above appear to be specialized within the genus, and are probably more or less recently evolved. The majority of specific character expressions are interpreted as derived expressions within character gradients established by com¬ parison of ragweeds with general conditions in the family Compos- itae. These specialized characteristics, common to the three species treated, include : shallowly lobed staminate involucres, eccentrically borne on relatively short stalks; vestigial pales; single flowered pistillate heads, with few, vestigial spines, localized near the beaks ; prominently lobed or dissected leaves, opposite at the lower nodes or throughout ; herbaceous perennial or annual habits ; and distribu¬ tion in non-arid eastern and northern North America. Although they share these specializations, the three species do not appear to comprise a natural, evolutionary group. Ambrosia psilostachya and A. art emisii folia are very similar and probably are the more spe¬ cialized members of a subgeneric group that includes the Mexican A. cumanensis, the West Indian and South American A. peruviana, the Caribbean A. hispida , and the South American A. tenuifolia and 1970] Payne — • Flora of Wisconsin No. 62 369 A. micro cephala, The entire group bears considerable resemblance to the A. conf ertiflora assemblage (including also A. cmiescens and A. pumila) and is probably derived from shrubby progenitors char¬ acterized by regularly lobed and dissected leaves, hooked involucral spines, and usually non-striated stems, involucres and leaves. Am¬ brosia trifida, however, bears greatest resemblance to species of a distinct, derivative line, characterized by irregularly lobed leaves, straight, flattened involucral spines, and prominent vegetative striations. It can be traced to shrubby progenitors of the least spe¬ cialized sort in the genus (such as A. deltoidea) along an evolu¬ tionary line represented by A. chamissonis , A. nivea, A. cheiranthi- folia, A. grayi , A. tomentosa, and A. acanthicarpa. Ambrosia Hybrids The three species are capable of hybridization, although hybrid individuals are usually uncommon and highly sterile. Hybrid plants are most easily recognized by their intermediate leaf character¬ istics (Fig. 4). Wagner and Beals (1958) have found that the perennial hybrid A. artemisiifolia X psilostachya (A. X inter- gradiens Wagner & Beals) (Fig. 4ap) is persistent and fairly “com¬ mon” in northern and eastern Michigan, where it often forms clonal populations that persist for many years. It is probable that similar populations are frequent in adjoining Wisconsin in areas of sympatry. Palynological evidence (Bassett and Terasmae, 1962) indicates that ragweeds occupied sites in the northeastern United States and adjacent Canada during and since the Pleistocene, and it is prob¬ able that all of the species found today in Wisconsin were here long before the invasion of North America by European cultures. On the other hand, European man, by providing variable primary sites in much greater abundance than were ever before available for ragweed occupation, has greatly influenced the natures of the species and their population dynamics. Literature Cited Bassett, I. J. and J. Terasmae. 1962. Ragweeds, Ambrosia species, in Canada and their history in post glacial time. Canad. Jour. Bot. 40: 141-150. Benson, L. 1957. Plant Classification. D. C. Heath & Co., Boston. Bentham, G. 1873. Notes on the classification, history, and geographical dis¬ tribution of Compositae. Jour. Linn. Soc. Bot. 13: 335-557. Cassini, H. 1826-1834. Opuscules Phytologiques. Tomes I, II, & III. Paris. Cronquist, A. 1952. Compositae, pp. 323-545. In H. A. Gleason (ed.), The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. Yol. 3. Lancaster Press, Inc., Lancaster. Darlington, H. T. 1922. Dr. W. J. Beal’s seed-vitality experiment. Am. Jour. Bot. 9: 266-269. 370 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 Delpino, F. 1871. Studi Sopra un Lignaggio Anemophilo delle Composte. Cellini, Firenz. Fernald, M. L. 1950. Gray’s Manual of Botany. 8th ed. American Book Co., New York. Gebben, A. I. 1965. The ecology of common ragweed, Ambrosia artemisii folia L ., in southeastern Michigan. Unpbl. Ph.D. thesis. Univ. Michigan, Ann Arbor. Gebben, A. I., W. W. Payne, and W. H. Wagner, Jr. 1962. Botanical Phase, pp. 1-16. In J. M. Sheldon and E. W. Hewson (eds.) Atmospheric pollu¬ tion by aeroallergens; prog. rept. 5. Univ. Michigan, Ann Arbor. Harrington, J. B., Jr., E. S. Epstein, D. L. Jones, A. N. Dingle, G. C. Gill, F. C. Elder, E. W. Bierly and E. W. Hewson. 1960. Meteorological Phase, pp. 75-177. In J. M. Sheldon and E. W. Hewson (eds.) Atmospheric pollu¬ tion by aeroallergens; prog. rept. 4. Univ. Michigan, Ann Arbor. Hartley, T. G. 1962. The flora of the “driftless area”. Unpbl. Ph.D. Thesis, Iowa State University. Jackson, R. C. 1960. A revision of the genus Iva L. Univ. Kansas Sci. Bull. 41: 793-876. Jones, K. L. 1936. Studies on Ambrosia: I. The inheritance of floral types in the ragweed, Ambrosia elatior L. Am. Midi. Nat. 17: 673-699. - . 1943. Studies on Ambrosia: III. Pistillate Ambrosia elatior X A. trifida and its bearings on matroclinic sex inheritance. Bot. Gaz. 105: 227-232. Keck, D. D. 1959. Compositae, pp. 1073-1310. In P. A. Munz. A California Flora. Univ. Calif. Press, Berkeley. Love, D. and P. Dansereau. 1959. Biosystematic studies on Xanthium; Tax¬ onomic appraisal and ecological status. Canad. J. Bot. 37 : 173-208. Miller, H. E. 1967. The chemistry and infraspecific variation of sesquiterpene lactones in Ambrosia psilostachya DC. (Compositae). Unpbl. Ph.D. thesis. Univ. Texas, Austin. Miller, H. E., T. J. Mabry, B. L. Turner, and W. W. Payne. 1968. Infra¬ specific variation of sesquiterpene lactones in Ambrosia psilostachya (Compositae). Am. J. Bot. 55: 316-324. Payne, W. W. 1962. Biosystematic studies of four widespread weedy species of ragweeds (Ambrosia: Compositae). Unpbl. Ph.D. thesis. Univ. Mich¬ igan, Ann Arbor. - . 1963. The morphology of the inflorescence of ragweeds (Ambrosia- Franseria: Compositae). Am. Jour. Bot. 50: 872-880. - . 1964. A re-evaluation of the genus Ambrosia (Compositae). Jour. Arnold Arb. 45: 401-438. - . 1965. “Subspecies amalgamation” in North American short ragweed, Ambrosia artemisiif olia (Compositae). Am. Jour. Bot. 52: 649 (abstract). Payne, W. W. and V. H. Jones. 1962. The taxonomic status and archaeological significance of a giant ragweed from prehistoric bluff shelters in the Ozark Plateau region. Pap. Mich. Acad. Sci. 47 : 147-163. Payne, W. W., P. H. Raven and D. W. Kyhos. 1964. Chromosome numbers in Compositae. IV. Ambrosieae. Am. Jour. Bot. 51:419-424. Rousseau, J. 1944. Reconstitution de 1’ Ambrosia prehistorique des Ozark. Nat. Canad. 71: 211-216. Rydberg, P. A. 1922. Carduales (Ambrosiaceae, Carduaceae). In North Amer¬ ican Flora. Vol. 33. N. Y. Bot. Gard., New York. Skvarla, J. J. and D. A. Larson. 1965. An electron microscopic study of pollen morphology in the Compositae with special reference to the Ambrosiinae. Grana Palynolog. 6: 210-269. 1970] Payne — Flora of Wisconsin No. 62 371 Small, J. K. 1913. Flora of the southeastern United States. 2nd. ed. Publ. by author. New York. Solbrig, 0. T. 1963. The tribes of Compositae in the southeastern United States. Jour. Arnold Arb. 44: 436-461. Wagner, W. H., Jr. 1958. The hybrid ragweed, Ambrosia artemisiifolia X trifida. Rhodora 60: 309-316. * - . 1959. An annotated bibliography of ragweed (Ambrosia) . Review of Allergy and Applied Immunology 13: 353-403. - . and T. F. Beals. 1958. Perennial ragweeds (Ambrosia) in Michigan, with the description of a new, intermediate taxon. Rhodora 60: 177-204. Wylie, R. B. 1915. A hybrid ragweed. Proc. Iowa Acad. Sci. 22: 127-128, pi. 21. BIOGRAPHIES CHARLES V. COVELL, JR., is assistant professor of biology at the University of Louisville and is curator of its insect collection. He holds a Ph.D. from Virginia Polytechnic Institute, is an officer in the Kentucky Society of Natural History, and is a member of entomological societies in the United States, Canada, and England. This year he will become editor of the Lepidopterists’ Society News. WILLIAM HILSENHOFF, associate professor of entomology at the University of Wisconsin, holds a Ph.D. from that institution. A specialist in the ecology of water insects, he is a member of several professional societies and has been president of the Midwest Benthological Society. Before teaching at Wisconsin he did classified research in medical entomology. HUGH H. ILTIS is professor of botany and director of the herbarium at the University of Wisconsin. He holds advanced degrees from Washington University and the Missouri Botanical Garden in St. Louis. A specialist in plant evolution and conservation, he has done field work in several locations throughout the Western hemisphere. FREDERICK U. UTECH, who holds an M.S. from the University of Wisconsin, is presently a graduate teaching assistant there. A botany major, he has held an NSF fellowship for tropical ecology studies in Costa Rica. MARLIN JOHNSON, who received his Masters degree in zoology from the University of Wisconsin, is an instructor in biology at the university’s Waukesha County Campus. GEORGE C. BECKER is professor of biology at Wisconsin State University — Stevens Point. A Wisconsin Ph.D., he has been president of the Wisconsin Society for Ornithology and vice-preside'nt of sciences in the Wisconsin Academy of Sciences, Arts, and Letters. WILLIAM A. KING is curator of art at the University of Wisconsin — Green Bay. He has studied at Oklahoma State University, the University of Tulsa, and ITstituto Statale d’Arte in Florence. He has had several one-man shows in both Europe and the United States, and his paintings hang in collections both here and abroad. Associate librarian of the Newberry Library in Chicago, DONALD W. KRUMMEL holds a Ph.D. from the University of Michigan. He is a member of several library and musicology societies and has written many articles and book reviews on music, music bibliography, and library history. He has pre¬ viously been reference librarian of the music division of the Library of Congress. HEINZ H. LETTAU, professor of meteorology and civil engineering at the University of Wisconsin, received his Ph.D. from the University of Leipzig. He is the author of many articles, primarily in the field of meteor¬ ology, and has worked on several committees of the National Academy of Sciences. ROBERT G. GALLIMORE, JR., holds the M.S. degree from the University of Wisconsin and is presently a research assistant in meteorology there. 373 374 Wisconsin Academy of Sciences, Arts and Letters [Vol. 58 LLOYD A. LUESCHOW, who holds an M.S. from the University of Wisconsin, is acting chief of the Laboratory Services Section of the Division of Environmental Protection in the state’s Department of Natural Resources. JAMES M. HELM (M.S. Wisconsin), GARY W. KARL (B.S. Wisconsin), and DONALD R. WINTER (B.S. Wisconsin) are biologists with the Depart¬ ment. All four are primarily concerned with the problem of water pollution. L. G. MONTHEY is travel-recreation specialist with University Exten¬ sion, the University of Wisconsin, and he has written many articles dealing with economic recreation and travel-industry trends. He holds an M.S. from Wisconsin and has held national offices in various professional agricultural societies. Associate professor of political science at Wisconsin State University — Whitewater, JOHN C. H. OH received his Ph.D. from New York University. He has taught previously at the University of Wisconsin and Marquette Uni¬ versity, and he has published articles and reviews in leading political and sociological journals. FREDERICK I. OLSON is professor of history and chairman of the department at the University of Wisconsin — Milwaukee. A Harvard Ph.D., he has also been associate dean of the University Extension, University of Wisconsin. He has published articles and reviews in many journals and is a member of various professional and public historical societies. WILLARD W. PAYNE, who holds a Ph.D. from the University of Michigan, is associate professor of botany and associate curator of the herbarium at the University of Illinois. He is presently on leave from the campus to serve as associate program director in the Ecology and Systematic Biology Section of the National Science Foundation. A. W. SCHORGER is emeritus professor of wildlife management at the University of Wisconsin, from which he received both his Ph.D. and an honorary D.Sc. The author of many articles and books on both chemistry and ornithology, he is a former director of the National Audubon Society and president of the Wisconsin Academy of Sciences. S. H. SOHMER, a member of the biology department at the University of Wisconsin — La Crosse, is also curator of the university’s herbarium. He holds degrees from City College of New York and the University of Tennessee, is the author of several articles, and has participated in programs in Costa Rica sponsored by the Organization for Tropical Studies. Assistant professor of journalism at the University of Michigan, JOHN D. STEVENS received his Ph.D. from the University of Wisconsin. He is an officer in the Association for Education in Journalism and has written a number of articles for leading journalistic periodicals. The paper presented here is based on his doctoral dissertation. ADOLPH A. SUPPAN is dean of the School of Fine Arts of the University of Wisconsin — Milwaukee. A Wisconsin Ph.D., he has held offices in many state organizations, including the chairmanship of the Wisconsin World Affairs Council and the presidency of the Wisconsin Academy of Sciences, Arts and Letters. At UWM he is also professor of English and philosophy. 1970] Biographies 375 GERALD E. SVENDSEN received his B.S. from Wisconsin State Uni¬ versity — River Falls and his M.A. in zoology from the University of Kansas. He is assistant professor of biology at Viterbo College, is a member of several professional societies, and has been a research biologist for the Bureau of Sport Fisheries and Wildlife. RONALD W. TANK, associate professor of geology at Lawrence Uni¬ versity, holds a Ph.D. from Indiana University and a graduate school diploma from the University of Copenhagen. He is the author of articles on regional geology and clay mineral genesis, and he has been a research geologist for Standard Oil of California. KENNETH TENNESSEN received a B.S. degree from the University of Wisconsin, majoring in entomology, and has done graduate work in this area at the University of Florida. He is presently in the armed services. Emeritus professor of soil science and forestry at the University of Wisconsin, SERGIUS A. WILDE received his professional training at the Prague Polytechnical Institute before emigrating to the United States in 1929. He is the author of several books about soil bacteriology which have been translated into many languages. BARBARA ZAKRZEWSKA, who holds a doctorate from the University of Wisconsin, is professor of geography at the University of Wisconsin — Milwaukee. A specialist in land form geography, she has published several articles on this subject in geographical journals. WISCONSIN ACADEMY OF SCIENCES, ARTS & LETTERS Madison, Wisconsin OFFICERS 1969-70 President William B. Sarles Department of Bacteriology The University of Wisconsin — Madison Vice-President (Sciences) Laurence R. Jahn Wildlife Management Institute Horicon Vice-President (Arts) Richard W. E. Perrin Department of City Development Milwaukee Vice-President (Letters) Edgar W. Lacy Department of English The University of Wisconsin — Madison President-Elect Norman C. Olson NML Insurance Company Milwaukee Secretary Corinna del Greco Lobner Department of English Dominican College Racine Treasurer Jack R. Arndt University Extension The University of Wisconsin — Madison Librarian Jack A. Clarke Department of Library Science The University of Wisconsin — • Madison APPOINTED OFFICIALS Editor — Transactions Walter F. Peterson University Librarian Lawrence University, Appleton Editor — Wisconsin Academy Review Ruth Louise Hine Wisconsin Department of Natural Resources Madison Chairman — Junior Academy of Science LeRoy Lee James Madison Memorial High School Madison ACADEMY COUNCIL The Academy Council includes the above named the following past presidents: officers and officials and Paul W. Boutwell A. W. Schorger Henry A. Schuette Lowell E. Noland Otto L. Kowalke Katherine G. Nelson Ralph N. Buckstaff Joseph G. Baier Stephen F. Darling Robert J. Dicke Henry A. Meyer Merritt Y. Hughes Carl Welty J. Martin Klotsche Aaron J. Ihde Walter E. 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