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Cover Design by Arthur Thrall, Lawrence University

TRANSACTIONS OF THE

WISCONSIN ACADEMY

OF SCIENCES, ARTS

AND LETTERS

LXI— 1973

Editor

ELIZABETH McCOY

TRANSACTIONS OF THE

WISCONSIN ACADEMY

Established 1870

Volume LXI

THE NEW ERA OF ACADEMY GREATNESS 1

F. Chandler Young

GOVERNOR EDWARD SALOMON, W. YATES SELLECK,

AND THE SOLDIERS’ CEMETERY AT GETTYSBURG 11

Frank L. Element

MAN’S CREEPING JURISDICTION OVER

OCEAN SPACE 29

John B. Ray

CULTURAL DIVERSITY IN CENTRAL WISCONSIN 45

Maurice E. Perret

PRELIMINARY REPORTS ON THE FLORA

OF WISCONSIN. NO. 63. THE GENUS TRIFOLIUM

—THE CLOVERS 59

John M. Gillett and Theodore S. Cochrane

THE MAMMALS OF DANE COUNTY, WISCONSIN 75

A. W. Schorger

GERMS, LUMBERJACKS, AND DOCTORS IN

NINETEENTH CENTURY MARINETTE 87

Carl E. Krog

THE EASTERN SUBTERRANEAN TERMITE, RETICULITERMES

FLAVIPES (KOLLAR), AND THE COMMON THIEF ANT,

SOLENOPS1S MOLESTA (SAY), IN THE LABORATORY,

WITH NOTES ON OTHER ASSOCIATED ANT SPECIES 95

R. V. Smythe and H. C. Coppel

SOCRATES ON CIVIL DISOBEDIENCE: THE APOLOGY

AND THE CR1TO 103

Peter S. Wenz

DISTRIBUTION OF PHOSPHORUS, SILICA, CHLOROPHYLL a,

AND CONDUCTIVITY IN LAKE MICHIGAN AND GREEN BAY 117

D. C. Rousar and A. M. Beeton

PRE- AND POSTSETTLEMENT POLLEN FROM A SHORT

CORE, TROUT LAKE, NORTH-CENTRAL WISCONSIN 141

Thompson Webb III

MARKET STRUCTURE AND BANK PERFORMANCE:

WISCONSIN 1870-1900 149

Richard H. Keehn

LA QUERELLE DU CID: CLASSICAL RULES

OR POLITICAL EXPEDIENCY? 157

Edmund Roney

THE EFFECTS OF HARVESTING AQUATIC

MACROPHYTES ON ALGAE 165

Stanley A. Nichols

ANNOTATED LIST OF TRICHOPTERA (CADDISFLIES)

IN WISCONSIN 173

Jerry L. Longridge and William L. Hilsenhoff

ORGANISMS, ESPECIALLY INSECTS, ASSOCIATED WITH

WOOD ROTTING HIGHER FUNGI (Basidiomycetes)

IN WISCONSIN FORESTS 185

J. L. Ackerman and R. D. Shenefelt

VACATION RESORTS IN ONEIDA COUNTY (WISCONSIN).

A Study of 1950-1968 Trends and Owner-operator

Characteristics 207

L. G. Monthey and Daniel Zielinski

MAJOR CAUSE OF ALGAE AND WEEDS IN

LAKE MENDOTA

229

M. Starr Nichols

ISOLATION AND CHARACTERIZATION OF A POSSIBLE

ALLELOPATHIC FACTOR SUPPORTING THE DOMINANT

ROLE OF HIERACIUM AURANT1ACUM IN THE

BRACKEN-GRASSLANDS OF NORTHERN WISCONSIN 235

Dana S. Dawes and N. C. Maravolo

DETERMINATION OF MERCURY LEVELS IN WISCONSIN

RESIDENTS: A PROJECT INVOLVING SECONDARY

SCHOOL STUDENTS 253

Leon M. Zaborowski

NITRATE CONTENT OF WELL WATER IN

WEST-CENTRAL WISCONSIN 259

Milan W. Wehking, Janies W. Pavlik, Paul Strege and Dawn Gilles

AN ORDINATION OF CORTICOLOUS LICHEN COMMUNITIES

IN THE POPPLE RIVER BASIN OF NORTHERN WISCONSIN 267

James A. Jesberger

ABOUT THE AUTHORS AND THEIR PAPERS 285

EDITORIAL POLICY

The Transactions of the Wisconsin Academy of Sciences, Arts and Letters

is an annual publication devoted to original, scholarly papers, some prefer¬

ence being given to the works of Academy members. Sound manuscripts

dealing with features of the State of Wisconsin and its people are especially

welcome, although papefs on more general topics are occasionally published.

Subject matter experts will review each manuscript submitted.

Contributors are asked to submit two copies of their manuscripts to the

Editor. The manuscripts should be typed on 8V2 x 11" bond paper. The title

of the manuscript should be centered at the top of the first page and should

be typed in capital letters throughout. The author’s name should appear

below the title and toward the right hand margin. It should be typed with

capitals and lower case and underlined for italics. Each page of the manu¬

script beyond the first should bear the page number and author’s name for

identification; e.g. 2-Brown, 3-Brown etc. A note on separate sheet should

identify the author with his institution, if appropriate, or with a brief

personal address. Other facts of importance or interest are requested, to

be used in About the Authors to be printed at the end of the volume; this

personal information will be verified with the authors at the time of proof

reading.

The style of the text should be that of scholarly writing in the field of the

author but to minimize printing costs the Editor suggests that the general

form of the current volume be examined. For the Science papers an AB¬

STRACT is requested. Documentary Footnotes may be useful, especially

for the Arts and Letters manuscripts, and should be typed and numbered

for identification as to place in the text; such footnotes as a group should

be separate from the text pages and may occupy one or more pages, as

needed. For the Science papers, or others at the authors’ choice, the refer¬

ences should be typed together on page or pages at the end of the manu¬

script to be printed under REFERENCES. Citations in the text may be

either by number in parentheses or by author and date as is preferred in

the authors’ field.

The cost, of printing the Transactions is great. Therefore, papers in ex¬

cess of 25 printed pages will not normally be accepted. In the rare instance

of such a paper or of some especially costly printing or illustration, the

contributor may be asked to subsidize publication.

Galley proofs and manuscript copy will be forwarded to the author for

proof reading prior to publication; both should be returned to the Editor

within ten days.

Papers received on or before November 1 will be considered for publica¬

tion of the current volume.

Manuscripts should be sent to :

Elizabeth McCoy

Editor: Transactions of the Wisconsin Academy

W.A.S.A.L. Office

1922 University Avenue

Madison, Wi 53705

REPRINT POLICY

Authors of papers which appear in Transactions will be provided by the

Editor with one complimentary copy of the issue in which their work is

published. Single copies only of the article are available to other persons

at 10 cents per page and single copies of the volume at $5.00 per copy from

the Wisconsin Academy office, 1922 University Avenue, Madison, Wi 53705.

Individuals may copy articles from Transactions in limited quantities for

educational purposes without request.

Large orders for copies of the articles should be placed with: Reprint

Division of the American Printing and Publishing Inc., 2909 Syene Rd., Madi¬

son, Wi 53713. Cost will be based upon the length of papers (units by 4

page intervals) and 100 copies is the minimum order. Prices and order

blanks will be sent with the proof and authors should deal directly with this

printer for routine orders up to 500. Very large orders (more than 500)

should be cleared by the W.A.S.A.L. office. Reprints by this plan will be

available within 2 weeks time and can be made either at the time of issue

of the Transactions volume or any time later, if desired. All orders should

be accompanied by institutional purchase order (or requisition #) or full

payment.

F. CHANDLER YOUNG

50th President

WISCONSIN ACADEMY OF SCIENCES, ARTS

AND LETTERS

THE NEW ERA OF ACADEMY GREATNESS

Presidential Address

F. Chandler Young

President 1971-72

Stevens Point, Wisconsin

May 6, 1972

Let me start boldly and abruptly. I am convinced that the

Wisconsin Academy is on the threshold of a new era of greatness.

We must recognize this and, in doing so, do our part for the

Academy and for the State of Wisconsin.

These opening remarks are so abrupt that I am at once reminded

of a professorial comment on the first draft of the Introduction

to my doctoral dissertation. My professor wrote: “You let the

clutch out too fast.” You may wonder why I have ignored this sage

advice. Let me simply say this. Whether as speaker or audience,

I have learned how easily after dinner speakers can lose their

audience to the comforts of sleep. I regard my time with you this

evening as precious. I shall not abuse the honor and privilege

granted me by this distinguished audience.

The evidence to support my thesis of Academy greatness in

these coming years derives from a number of observations about

the past and the present which have significance for the Academy’s

future.

I shall begin by drawing upon some of the wisdom in our history.

We do need an awareness of our illustrious past for guidance in

charting our future. I shall continue with some observations of

the present. We need to understand the conditions of the society

in which the Academy exists today. They are quite different from

what they were a hundred years ago. The Wisconsin Academy

itself is stronger than ever before. We need to know about this new

strength.

Then, with an awareness of our past and present, we can look

toward a great future.

Early in my studies of the Academy’s past, I found a sentence of

special interest to me, personally. Thomas Chamberlin had written

about our first president, John Hoyt. In that one sentence charac¬

terizing Hoyt, our first president, he in fact characterized me, your

last president. He wrote: “Though not a special worker in any line

1

2 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

of research, his intellectual sympathies were wide, his aspirations

were high, his dream for the Academy was ambitious. ” My dream

for the Academy is at once ambitious and realistic.

Back in 1872, John Hoyt had reported to the Governor of the

State of Wisconsin. This was when the Academy was just two years

into its first century. Now, in 1972, I report to you, the members

of the Academy. This is when the Academy is just two years into

its second century.

John Hoyt had told the Governor about what little progress had

been made in Wisconsin in the Sciences, Arts, and Letters. He

showed how necessary the Academy was to the State of Wisconsin.

A few quotes from his 1872 address give us some of the thrust and

flavor of his remarks.

In commenting about the Fine Arts in our early history, he

wrote: “The practice in Architecture, both in the construction of

private dwellings and buildings for public use, gave here, as else¬

where in our country, painful proof of a prevailing ignorance of

the principles of art.,, In remarks about the intellectual achieve¬

ments of that era, he wrote: “If [the record] shows that in the

Practical Arts — in the rough work of civilization — we had

achieved marvelous results for a State of but twenty-two years,

it reveals, on the other hand, how little has been accomplished in

those higher fields of human activity, the scientific, literary, and

aesthetic . . . [which are] essential to those high intellectual

achievements which exalt man as an individual and make the other¬

wise half-civilized community an enlightened and refined common¬

wealth.”

We know that the Transactions , the Academy Review, and some

special monographs and papers written for our centennial celebra¬

tion attest to our many worthwhile accomplishments. In these

writings we not only get a sense of our purpose, we also can find

some specifics in our history which we must use. I have time for

only one example.

Charles Sumner Slichter was president of the Academy in 1902.

In his presidential address, I found a paragraph worth repeating.

His address was entitled, “Recent Criticism of American Scholar¬

ship,” and he was discussing the then new Carnegie Institution.

He wrote in 1902 :

The Carnegie Institution should take its chief warning from the un¬

fortunate history of the Smithsonian Institution, which at one time

promised so much for American science. This institution instead of becom¬

ing the one place in the United States where the highest science could

always find a home, has become very largely a routine institution. It

spends its money for salaries and administration in true American fashion

and has a minimum to show for its more than fifty years of existence.

At the present time about four-fifths of its income goes for salaries and

1973]

Young — New Era of Academy Greatness

3

expense of administration. The American is a great man for stipends,

and stenographers, and card catalogs. Fortunate would it have been if

a Helmholtz had had charge of this institution. He would have been so

absorbed in his science that he would have forgotten about his clerks

and typewriters, but his suggestions and plans, given to his scientific

workers, would have made at Washington an institution conspicuous

“for the increase and diffusion of knowledge among men.”

We must heed this advice. We must be aware of the ideas re¬

corded in our history. But we must resist the temptation of dwell¬

ing too heavily upon our past. We must understand the present

and then dwell upon our future.

In recent years I have experienced what may be called emotional

attacks. As one who has survived 21 years as a college adminis¬

trator on the Madison campus, I know that these attacks were not

upon my person, but rather upon my position. What bothers me the

most is that these attacks were leveled at an institution devoted

to teaching and research, to intellectual pursuits, to the search for

truth.

My South Hall office was fire bombed in 1968. My Bascom Hall

office window was trashed in 1969. Tear gas has caused me to weep

on more than one occasion.

I have been publicly criticized. In 1968, when I became Vice

Chancellor of Student Aifairs, a local newspaper announced it with

the foreboding headline: ‘‘Young gets UW ‘Hot Spot’ position.”

Some time later, a local paper's front page story about a regent

meeting reported: ‘‘The Regents spent more than three hours

lambasting and questioning UW administrators . . .” In reporting

my answer to a question put to me by the Regents at that meeting,

the article stated: “Young’s answer was drowned out in a chorus

of ‘why not’ from several of the Regents.”

I have been confronted by other angry people. For three and a

half hours a legislative investigating committee questioned me,

not for facts or for information but as an occasion to attack me

verbally, publicly, and politically. One question was, in essence,

why I had permitted the Daily Cardinal to use four letter words.

Another was, in effect, why I did not rid the campus of communists.

On another occasion, eight students entered my office, posted

guards at my door, pointed a camera and a tape recorder at me and

insisted upon an immediate response to thirteen demands.

Another day, during a disturbance, an elderly citizen called me

late at night to demand that I get the students out of the neighbor¬

hood. A little later that same night, a student in the same neigh¬

borhood demanded that I get the police out and ended with the

icy comment, “What is a highly paid administrator like you doing

in bed on a night like this?”

4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

I could, of course, continue with many similar experiences,, but

I shall cease with a brief account of one more incident. This one

contained a bit of humor, a little reason, and even considerable

decency.

This took place in the height of campus turmoil. An attempt was

being made to halt classes. The idea seemed to be that drawing

attention to a particular issue was more important than the intel¬

lectual pursuits of the campus. Some twenty students were block¬

ing one of the main entrances to Bascom Hall. I went up to the

group and said cheerfully: “It doesn’t look like I can get through

here, does it?” The leader asked me pleasantly: “Are you a stu¬

dent?” I said “No.” Although I was tempted to say thanks for the

compliment. Then the leader asked: “Are you a teacher?” “No,” I

said, “I’m just a Vice Chancellor on my way to a meeting.” With

this, the leader said : “Let the Vice Chancellor through.” He waved

his arm and the group made a path through its midst. As I went

through, my host said eagerly: “Please remember how polite we

have been.”

One hundred years ago we had no dean of students, let alone any

vice chancellor. Students caused trouble, of course, and leaders

in educational institutions constantly worked with the political,

economic, and social forces of the time to the benefit of teaching

and research in higher education. I doubt, however, that the anti¬

intellectual forces have ever been more dramatically visible nor

more intense than at present.

It is the irrationality of man that makes it difficult. Many of the

persons in the emotionally charged sessions of my experience seem

to have rejected the intellectual approach. Whether politically radi¬

cal or ultra conservative, whether in fact in positions of authority

or in fact trying to seize power, some persons seem to prefer

ignorance to knowledge. They seem to prefer persuasive rhetoric

based on falsehoods and half truths rather than facts and the

principles of logic and causality. They seem to like the emotional

comfort of over simplification. They often dismiss the realities

of complexity. They seem to be motivated by fears, hates, and

angers, seemingly derived from perceived threats to their own

well being or to their own belief systems.

It is this kind of irrationality, at work in our society today,

which threatens the interests of our Academy and of the citizens

of the State of Wisconsin. I am not alone in my concern about

threats to the forces of intellect.

Kingman Brewster, Jr., in a recent Phi Beta Kappa address,

expressed this concern when commenting on an orthodoxy of pas¬

sion and politics. He said: “If we should slip into a dark age of

irrationality, far better for the Universities to live meagerly, while

1973]

Young — New Era of Academy Greatness

5

sticking to the integrity of reason, than to prosper elegantly in the

corruption of a new orthodoxy/’

Another intellectual expressed this concern in a somewhat differ¬

ent way. Jacques Barzun, in his book, The House of Intellect ,

arraigns our modern democratic culture for its unfortunate atti¬

tude toward intellectuals. He is intensely critical of our educational

system and at one point is quite caustic in his assessment of the

college graduate: “After sixteen long years of an ordinary edu¬

cation, we observe young men and women of unquestionable gifts,

energy, and zest whose fine intelligence is not matched by strength

of intellect. Freshness of taste and openness to novelty, yes; even

refinement of sensibility, sometimes prematurely delicate. Though

here are college graduates, many of them cannot read accurately or

write clearly, cannot do fractions or percentages without travail

and doubt, cannot utter their thoughts with fluency or force, can

rarely show a handwriting that would pass for adult, let alone

legible, cannot trust themselves to use the foreign language they

have studied for eight years, and can no more range conversation¬

ally over a modest gamut of intellectual topics than they can

address their peers consecutively on one of the subjects they have

studied.”

Perhaps college graduates are less equipped and perhaps less

skilled intellectually than is desired. Nevertheless I believe that

college graduates today are much more aware of the problems of

our society than they were a hundred years ago. Even though their

behaviors may have at times appeared irrational, one underlying

rationale seems to have been to stir things up in ways which make

the public more fully aware of existing problems and thus bring

public pressure to bear on persons in authority to make desirable

changes.

Let me just mention a few of the concerns which students ex¬

press about our society. Each set of problems is exceedingly com¬

plex but each exists and is in need of solution.

Persons tend to be discriminated against if they are different

from the majority in regard to age, sex, color, ethnic origin, or

religious beliefs. What could be more irrational than to treat

persons differently simply because they belong to some minority

group ?

The problem with the military industrial complex is essentially

the misplacement of priorities ; that is, a greater concern for mili¬

tary and economic power than for the well-being of persons.

Poverty and hunger have no rational basis for existence in our

affluent society, yet the prevailing attitude seems to be that these

human miseries have always been and thus always will be a part

of our society. What could be more irrational?

6 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

The value put upon material things seems to be higher than the

value put upon persons and upon our natural environment. For

example, what is the rationale for spending millions on automobiles

and highways at the expense of the well-being of persons, not to

mention the resulting irrevocable damage to our environment?

My main point is that fierce irrational forces are at work. I have

felt these personally. I have found modern day thinkers who are

concerned about this same inattention to intellectual matters and

the same difficulty of making intellectual forces effective. I have

identified some of our urgent problems, seemingly created by

irrational forces and certainly in need of solution by knowledge¬

able, reasonable, intelligent men and women.

Society, of course, had its problems a hundred years ago. The

record shows that the Wisconsin Academy played a part in finding

solutions to these problems then. The Academy did this through the

encouragement of research and the spread of knowledge. The prob¬

lems today are quite different and in my view much more serious

and more urgent than ever before. The mission of the Academy is

still essentially that of providing intellectual strength through

the encouragement of original investigation and through the spread

of knowledge.

Academy greatness in the years to come can be expected because

intellectual strength in the minds of men in every walk of life is

needed more today than ever before and the Wisconsin Academy is

better equipped than ever before to provide that strength.

Let me turn to a brief discussion of a new and growing strength

in the Wisconsin Academy.

On the financial front we are far ahead of John Hoyt’s fondest

dream. In 1872, our assets were less than one thousand dollars

when John Hoyt had expressed the hope of increasing our assets

one hundredfold. Now our assets have been increased a thousand¬

fold. My hope is that we will more than double our current assets

of over a million dollars.

Even though there were special ceremonies during our centen¬

nial celebrations to honor our great benefactor, Harry Steen-

bock, I want to say again how grateful and appreciative we are

for his outstanding gift. We have invested this wisely. We have

used only the income from that investment and have spent this

income cautiously and sensibly. It is interesting to note that Harry

Steenbock was one of us. It is true that we have had some financial

help from the legislature and some from business and industry.

The University of Wisconsin-Madison has helped us in important

ways over the years and continues to do so. Up to now, however,

these other contributions have been relatively small. This tells us

something. First, we must intensify our efforts for financial sup-

1973]

Young — Neiv Era of Academy Greatness

7

port from business and industry. Second, we must think seriously

of ways to approach our own members. We are the ones who under¬

stand the Academy and its importance to society. There are those

among* us who wish to add to our financial assets through gifts or

bequests. We must encourage these people to step forward.

Our membership is larger and more active in Academy affairs

than ever before. A century ago we had only fifty-five members.

Now we have over 1500. More important, there is every indication

that our membership will increase significantly; not simply in

sheer numbers, but also in degree of active participation.

Back in 1955, my only task as an Academy Vice President was

to write a paragraph for the back cover of the Academy Revieiv.

Although it seems strange to be quoting myself, I do believe my

comments then support my present thesis that our membership

will become even stronger than it is today. I had written :

“Many Wisconsin people have the kind of inquiring mind, the degree

of fascination with new knowledge, and the depth of appreciation for

scholarly endeavor to enjoy membership in this Academy. We who are

members can understand how the Academy can enrich one’s life and

contribute to the cultural and intellectual life of the State.”

I should like to identify briefly three conditions of society today

which further convince me about our new growth in membership

participation.

One is the condition of boredom. Perhaps it is our affluence.

Perhaps it is the lack of any need for intellectual effort in many

occupations. Perhaps it is because we have more leisure time or

because we are less preoccupied with our own financial security.

Many persons will welcome Academy activities as a worthwhile

way to fill their leisure time with exciting intellectual and creative

pursuits.

A second condition is that a greater variety of people are be¬

coming interested in higher learning. New egalitarian trends in

higher education are already evident. College populations are

drawing persons to campuses from a much wider base. Ethnic

origin, still an irrational barrier, is now becoming somewhat less

of a reason to bar a person from higher education. Whatever one’s

view may be about women’s liberation, the truth is that more and

more women are being welcomed into the academic ranks. Sons

and daughters of parents in non-professional and non-managerial

occupations are coming to campuses in greater numbers. Age is

now less of a barrier to higher learning and more and more persons

over thirty are seeking higher learning.

A third condition is an increasing general attitude of granting

individuals greater freedom “to do one’s own thing.” There seems

to be a new emphasis on nature and the natural, on encouraging

8 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

each person to live his or her own life, unhampered by the pres¬

sures of conformity or the expectations of others. Perhaps it is a

new kind of individualism.

These conditions of a greater need to escape boredom, an in¬

crease in number of persons entering the intellectual arena, and

a greater concern for freedoms for the individual, present us with

unusual opportunities for new memberships and greater member¬

ship participation.

In addition to our fresh economic power and our revitalized

membership, we have new administrative assistance and new

physical facilities. It is not that we are great “for stipends,

stenographers, and card catalogs.” It is rather that we will be great

because our members, our officers, our Council, our committees,

and our editors can come forth with suggestions, can make plans,

can accomplish tasks and thus make our society conspicuous “for

the increase and diffusion of knowledge among men.” Our highly

competent Executive Director and his limited staff are not re¬

garded as substitutes for volunteers. He and his staff make it

possible for us to carry out our work cooperatively, efficiently,

thoughtfully, and above all, effectively.

Those of us who have participated actively in Academy Affairs

this past year have come to appreciate the importance of competent

administrative help. We have been able to get our house in order,

to care properly for our precious investments, to up-date our con¬

stitution and by-laws, to re-awaken dormant committees and create

new ones, and to introduce new qualities of excellence in our

publications, in our fall gathering, and in our annual meeting.

Taken all around, the Wisconsin Academy is better equipped for

its mission than ever before. It has an illustrious past upon which

to build. Its financial position is strong. Its growing membership

is freer than ever before and more highly motivated than ever

before to engage in activities of the kind we all treasure. The

society in which we live is faced with urgent problems which not

only threaten the quality of life but threaten the very existence

of mankind. These are problems that need solution by the knowl¬

edgeable, reasonable, intelligent, concerned persons of the kind

who are members of our Academy.

This is the threshold to greatness upon which we now stand. But

we cannot stand. We must move forward with renewed enthusiasm

and with a full awareness of our capabilities.

In my view, our most important task is to strengthen the intel¬

lectual and creative forces among the citizenry of the State of

Wisconsin. Intelligent, knowledgeable and reasonable men and

women are everywhere, whether they be rich or poor, craftsman

or professional, worker or manager, man or woman, young or

1973]

Young — New Era of Academy Greatness

9

old, white or of color, urban or rural, or whatever. These are the

thinking men and women. These are the people who shape public

opinion, who use their time and talent for the public good, whether

in occupation or at leisure. They now know or can easily come to

know about “those higher fields of human activity, the scientific,

literary, and aesthetic . . . which exalt man as an individual and

make an otherwise half-civilized community an enlightened and

refined commonwealth.”

Our Long Range Planning Committee has prepared a number of

working papers which many of you have read and discussed. These

are the charts for our future courses of action, for carrying out

our primary mission. We must pay close attention to these ideas

and where there is general agreement we must implement them,

not file them away.

I am especially pleased with the ideas for expanding our ways of

spreading knowledge, of reaching the thinking citizens of the

State. Our new publications committee will explore and hopefully

implement new ideas for our publications. We need also to explore

new uses of radio, television, and other modern communication

devices.

We have been pleased with the ways our Junior Academy has

reached our younger citizens. The new age of majority legislation

shows the respect we have for the young men and women of our

State. The Junior Academy of Science has now become the Junior

Academy of Sciences, Arts, and Letters. Whether over or under

the age of 18, the younger citizens of our State need opportunities

which will encourage their intellectual pursuits.

We need to cooperate more fully with our affiliated organizations

and seek out other interest groups engaged in activities germane

to our mission. We must learn from them and, where appropriate,

assist them in their task of informing our citizenry.

As we work together in our efforts to spread knowledge, we can

enjoy a personal sense of identity. I think each of us, whatever

his or her own particular creative or intellectual pursuits may be,

likes to belong, to have an identity with a total effort, to share ideas

and information.

As we read through the history of the Academy, we can note

shifts in emphasis regarding the Sciences, Arts, and Letters. In

my view, there is emerging a new emphasis upon the fine arts and

I, for one, sincerely hope that this trend continues. My guess is

that our new president-elect also shares this view.

Finally, I believe that we should devise new ways of cooperating

with the Governor, the legislature, and others in positions of gov¬

ernmental authority. Far too long we have approached our govern¬

ment with a tin cup. We must now approach it with the persuasive

10 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

power of knowledge and reason. We must have the courage to take

positions on issues when there is common agreement. Even if an

issue is so controversial that we cannot reach any agreement, we

should be willing to present both the pros and cons with intellectual

honesty and scholarly objectivity.

In closing let me simply say this. I believe that we are just

entering a new era of Academy Greatness. I hope that you believe

this too and that we will all do our part for the Academy and for

the State of Wisconsin.

GOVERNOR EDWARD SALOMON, W. YATES SELLECK,

AND THE SOLDIERS’ CEMETERY AT GETTYSBURG

Frank L. Klement

Marquette University

Milwaukee, Wisconsin

Two Wisconsin residents played an important role in the estab¬

lishment of the Gettysburg-based soldiers’ cemetery, a cooperative

project of eighteen states. Governor Edward Salomon endorsed

the project, promised that Wisconsin would pay its fair share of

the costs, and named a state agent to cooperate with Pennsylvania

authorities. W. Yates Selleck, as state agent and Salomon’s rep¬

resentative, spent considerable time in Gettysburg, lent energy

and ideas to the project, played a role in the dedication cere¬

monies of November 19, 1863, and served as secretary of the

cemetery association.

Governor Salomon, who promised that Wisconsin would be a pay¬

ing partner in the enterprise, was an effective administrator, if

not a popular one. In a way, chance and expediency had made

Salomon their prisoner. Born in Prussia, he accompanied an older

brother to Wisconsin, living first in Manitowoc, then moving to

Milwaukee in 1852 to enter the fields of law and politics. He started

out as a Democrat, but, influenced by Carl Schurz, he deserted

that party for Republicanism and the slogan “Free soil and free

men.” At their state convention in 1861 Republicans gave Salomon

the second spot on the state ticket in a bid for votes of the German-

Americans. The strategy helped put Louis P. Harvey in the execu¬

tive mansion and made Salomon the lieutenant governor. Chance

again intervened. When Governor Harvey was in the Shiloh area

distributing supplies and hospital stores, he fell into the Tennessee

River while stepping from one steamboat to another during the

night and was drowned — elevating Salomon to the governorship.1

While Governor Salomon worked in the glare of the public spot¬

light, W. [illiam] Yates Selleck lived in the shadows, on the fringe

1 Edward Salomon was born at Stroebeck, near Halberstadt, Prussia, on August 11,

1828. He was a student, first at the college at Halberstadt, and then at the University

of Berlin. Political repression, part of the story of the revolutions of 1848, prompted

him to leave for the United States. He settled in Manitowoc, serving successively as

school teacher, county surveyor, and deputy clerk of the circuit court. After moving

to Milwaukee, he read law in the offices of Edward G. Ryan, being admitted to the

bar in 1855. He set up a law practice with Winfield Smith and engaged in politics,

supporting Democratic candidates. He bolted the Democracy to support Lincoln in

1860.

11

12 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

of obscurity. A young man who had come to Milwaukee during

the 1850’s, he found employment' with an insurance firm and

turned to politics as an outlet for his energy. He was one of the

founders of the Young Men’s Republican Club in March of 1860,

and became the organization’s corresponding secretary.2 He sup¬

ported Lincoln for the presidency and became acquainted with the

Salomon brothers. Republican victory in the 1860 elections netted

Selleck an appointment in Washington.3

The young, energetic and imaginative fellow was one of the or¬

ganizers of the Wisconsin Soldiers’ Aid Society, formed by the

state’s residents-in- Washington to furnish aid to sick and needy

soldiers — “and more especially to visit the hospitals and see that

our soldiers are well cared for.”4 Appointment as state agent (more

correctly “State Military Agent”) to oversee Wisconsin’s sick and

wounded soldiers became a full-time job after the Union defeat at

First Bull Run and as more of the state’s volunteers were assigned

to the Army of the Potomac. Selleck felt compelled to defend the

reputation of the Third Wisconsin Regiment which a correspondent

of the New York Times had mistakenly accused of cowardice.5 In

a letter to the Milwaukee Sentinel, Selleck wrote “ ... it is a base

slander and falsehood, and there is not a word of truth in it .”6

Union defeats in such battles as Second Bull Run, Fredericksburg,

and Chancellorsville placed a heavy load upon the shoulders of

Selleck, who performed his duties with a dedication and ability

unmatched by any other state agent in the Washington area.

When word of heavy fighting in the Gettysburg area went out

over the telegraphic wires, both Governor Salomon and state agent

Selleck expressed apprehension. Approximately 6,000 Wisconsin

soldiers served in the Army of the Potomac, and each major battle

brought concern to state authorities and sorrow to many homes.

Selleck left Washington for Gettysburg as soon as reports reached

the capital that the battle was over.7 Taking his assistant, Wil¬

liam P. Taylor, with him, Selleck took the train to Baltimore in

the hopes of securing passage to Hanover Junction and Gettysburg.

But they could not get a train out of Baltimore until July 6, and

then the train ran only as far as Westminister, Maryland. Un¬

able to secure a conveyance of any kind in Westminister, Selleck

and Taylor set out on foot, carrying bundles on their backs. They

reached Littlestown, seventeen miles away, at sundown. Next

morning they “procured a conveyance” which took them the final

2 Milwaukee Sentinel, March 22, 1860.

a Ibid., April 27, 1861.

4 Ibid., June 9, 1861. James R. Doolittle, one of the state’s U.S. Senators, served

as president of the organization while Selleck held the office of vice-president.

5 New York Times, Aug1. 12, 1861.

6 Selleck to “Editor,” Aug. 14, 1862, published in Milwaukee Sentinel, Aug. 20, 1862.

7 Selleck to Salomon, July 9, 1863, published in (Madison) Wisconsin State Journal,

July 15, 1863.

1973] Klement — Soldiers' Cemetery at Gettysburg

13

sixteen miles to Gettysburg. The two state agents found that Wis¬

consin troops had suffered heavy casualties, the famous “Iron

Brigade” (composed of three Wisconsin regiments and one each

from Indiana and Michigan) being decimated in the first day’s

fighting during the three-day battle.8 He visited Col. Lucius Fair-

child, whose left arm had been amputated above the elbow after

the first day’s fearful and fateful battle. He visited hospitals and

noticed a pathetic shortage of supplies. Leaving his assistant to

“make out lists of all the killed, wounded and missing” (Wiscon¬

sin soldiers), Selleck rushed back to Washington and Baltimore to

“procure supplies.” To get back to Washington, the spirited and

solicitous agent rode “17 miles on a locomotive and 69 miles on

the back end of a freight car.”9

Selleck arrived back in Gettysburg late during the afternoon of

July 10, bringing with him “5 boxes of under clothing and other

hospital stores” as well as “three bundles of crutches.” After dis¬

tributing most of the “stores,” Selleck took time to write about

his activities to Governor Salomon. He promised to send a list of

“the killed, wounded and missing” as soon as possible.10

Selleck and Taylor did noble work, angels of mercy to those in

need. “The Wisconsin wounded at Gettysburg,” Selleck’s aide wrote

to a friend, “were better attended to and cared for than those of

any other State — -not even excepting Massachusetts.”11

Taylor could have added that the wounded and sick soldiers re¬

ceived far more consideration than the dead. Many of the dead

soldiers had been buried hurriedly and carelessly. In some cases

no graves had been dug, and the bodies had merely been covered

with spadefuls of dirt ; mounds of soil dotted portions of the battle¬

field, especially that section where the first day’s fighting had been

heaviest. Heavy rains which followed washed off some of the soil

shoveled over the maggot-infested bodies, exposing arms, legs, and

heads. “I saw one entire skull above the ground,” wrote an ob¬

server, “and in many instances hands & feet are sticking

through.”12 A blazing sun and the summer heat hastened the bloat¬

ing and decomposition of the corpses, filling the air with a sicken¬

ing stench. Worse than that, hogs rooting in various portions of

8 The “Iron Brigade,” the only all-Western brigade in the Army of the Potomac,

consisted of the 2nd, 6th, and 7th Wisconsin, the 19th Indiana, and the 24th Michigan.

The story of this “famous fighting unit” is well told in Alan T. Nolan, The Iron

Brigade (New York, 1961). No participant in the Battle of Gettysburg wrote a better

account than a first lieutenant serving cn the staff of John Gibbon, commanding the

Iron Brigade at Gettysburg; see Prank A. Haskell, The Battle of Gettysburg (Madison,

1908).

“Selleck to Salomon, July 9, 1863, published in Wisconsin State Journal, July 15,

1863.

w Ibid.

11 Taylor to Leonard J. Farwell, July 18, 1863, published in part in Wisconsin State

Journal, July 24, 1863.

12 Ellen Mead to “My dear Mrs. Dean,” Aug. 12, 1863, Lucius Fairchild Papers, State

Historical Society of Wisconsin (Madison).

14 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

the onetime battlefield, desecrated the graves and devoured por¬

tions of the rotting human bodies.13 -

In his work as state agent, Selleck became well acquainted with

David Wills, a kindred soul and a Gettysburger whom the governor

of Pennsylvania, after a visit to the battlefield, had named his rep¬

resentative to supervise “the removal of all Pennsylvanians killed

in the late battles, furnishing transportation for the body and one

attendant at the expense of the State.”14 In pursuing their assign¬

ments, Wills and Selleck noticed that many of the markers over

the graves were temporary and inadequate and that some of the

names were faded and quite unreadable. . . three thousand men

lie in and around Gettysburg,” noted one who had tramped over

the battlefield, “in cornfields, in meadows, in gardens, by the way

side, and in the public road, buried hastily where they fell, and

others in long rows, with a piece of box lid or board of any kind,

with the name of the person and the day he died written with lead

pencil, ink, or whatever they had to make a mark with.”15 Time,

aided by sun and rain, made some of the inscriptions quite illegible,

while occasional so-called graves had no headboards at all.

Wills, encouraged by state agents like W. Yates Selleck of Wis¬

consin and Henry Edwards of Massachusetts, envisioned a cooper¬

ative state cemetery as both a possibility and a necessity. Massa¬

chusetts agents had already asked their state authorities to con¬

sider buying a portion of the battlefield — preferably atop Cemetery

Hill — as a burial ground for Massachusetts soldiers whose bodies

had not been returned to the state and whose ill-kempt graves

were scattered here and there, wherever troops had been engaged.

Selleck, like Wills, believed that most of the eighteen states which

had lost sons at Gettysburg would partake in a project to rebury

the dead in a common cemetery created out of a portion of the

battlefield. Encouraged by Selleck and other state agents, Wills

conveyed the idea to his governor, suggesting that Pennsylvania

purchase a plot for the proposed cemetery. He added an impas¬

sioned plea: “Our dead are lying in the fields unburied (that is no

graves being dug) with small portions of earth dug up alongside

of the body and thrown over it. In many instances arms and legs

protrude and my attention has been directed to several places

where hogs were actually rooting out the bodies and devouring

them — and this on Pennsylvania soil. . . . Humanity calls on us

to take measures to remedy this. . . .”16

13 David Wills to Governor Andrew Curtin, July 24, 1863, Curtin Letterbooks, Execu¬

tive Correspondence, 1861-1865, Pennsylvania State Archives (Harrisburg-),

14 (Gettysburg) Adams County Sentinel, July 28, 1863.

15 Andrew B. Cross, head of the Christian Commission of Pennsylvania, in an appeal

(undated) entitled “To the Patriotic of the Land — A Cemetery for those who Fell

at Gettysburg-,” published in Harrisburg Daily Telegraph, July 29, 1863.

10 Wills to Curtin, July 24, 1863, Curtin Letterbooks, Executive Correspondence,

1861-1865.

1973] Klement — Soldiers’ Cemetery at Gettysburg 15

Governor Curtin, busy with affairs of state, authorized Wills

to purchase the necessary acreage for the proposed cemetery, con¬

tact authorities of the other seventeen states, make arrangements

for the reburials, and superintend the entire project. After each

of eight agents (including Selleck) whom he had “consulted,”

endorsed the cemetery proposal “semi-officially,”17 Wills took steps

to purchase seventeen acres atop Cemetery Hill, adjacent to the

local cemetery already established there. He composed a telegram

to each of the governors who should have had an interest in the

cooperative venture. Wills’ telegram of August 1, 1863, to Governor

Salomon read : “By authority of Gov. Curtin I am buying ground

on or near Cemetary [sic] Hill in trust for a cemetery for the

burial of the soldiers who fell here in the defence [sic] of the

Union. Will Wisconsin cooperate in the project for the removal of

her dead from the field? Signify your assent to Gov. Curtin or

myself and details [will] be arranged afterwards.”18

Governor Salomon replied promptly: Yes, Wisconsin would coop¬

erate in the project. The governor, however, was concerned with

the “details,” for he wanted no loose ends which might be used

to hog-tie him politically. Having committed his state, Salomon

instructed W. Yates Selleck, who had returned to Washington, to

go back to Gettysburg and “confer with Mr. D. Wills” in reference

to “the detail of arrangements for removal of the Wisconsin dead

to the Cemetery grounds.”19

Salomon took some influential Republicans, including editor

Horace Rublee of the (Madison) Wisconsin State Journal , into

his confidence. Rublee, in turn, endorsed the cemetery project in

the public press. “Governor Salomon of this State,” wrote Rublee,

“has signified, in response to Governor Curtin’s proposition, the

readiness of Wisconsin to contribute her proportion towards this

laudable and patriotic enterprise.”20

Selleck, meanwhile, returned to Gettysburg to seek out Wills and

ask questions about progress on the cemetery project and about

the so-called “details.” Wills was then negotiating for the purchase

of five plots totalling seventeen acres — he eventually purchased

two plots at $225 per acre, one at $200, another at $150, and the

fifth at $135. 21 Wills also talked to Selleck and other state agents

17 Wills to Curtin, July 30, 1863, ibid.

18 Wills [as agent for Gov. Curtin] to Salomon, Aug. 1, 1863, Telegrams, 1861—1865,

Executive Department (Administration), Archives Section, State Historical Society

of Wisconsin.

19 William H. Watson (Salomon’s military secretary) to Selleck, Aug. 3, 1863,

Letter Books General, Executive Department (Administration), 1861-1865, Archives

Section, State Historical Society of Wisconsin.

20 Wisconsin State Journal, Aug. 24, 1863.

21 David Wills, report of March 21, 1864 (to Gov. Curtin), published in Revised

Report of the Select Committee Relative to the Soldiers’ National Cemetery, Together

with Accompanying Documents as Reported to the House of Representatives of the

Commonwealth of Pennsylvania (Harrisburg, 1865), pp. 7-8.

16 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

about the specifications for the proposed reburials, seeking their

aid and advice in drafting a circular letter to each of the governors

involved in the project.22 Wills and the agents also discussed the

advisability of getting William Saunders, an employee of the De¬

partment of Agriculture and a well-known landscape gardener,

to come to Gettysburg and “lay out” the proposed soldiers’ ceme¬

tery.

After Wills composed his circular letter to the governors, he had

copies printed and put into the mails. Instead of sending Salomon’s

copy he gave it to Selleck to forward to his governor. Selleck

promptly forwarded the circular letter and included a letter of his

own.

In his letter Selleck stated that Wills had purchased “a very

suitable piece of ground, on one of the most prominent parts of

the battlefield,” and urged Salomon to act promptly upon the

cemetery proposal. “It is desirable to have as little delay as pos¬

sible in getting your reply,” he wrote to his governor, “as the

bodies of our soldiers are in many cases so much exposed as to

require prompt attention, and the grounds should be speedily ar¬

ranged for their reception.” Selleck added, “Governor Curtin

authorized me to say to you that if your State desires a conveyance

in fee simple, of your burial ground in this cemetery, Pennsylvania

will make a deed to you for it. Otherwise she will hold the title

in trust for the purposes designated in the circular.”23

Wills’ carefully composed circular letter, dated August 12, stated

that Pennsylvania had purchased a portion of the battlefield “to

be devoted in perpetuity” for a soldiers’ cemetery, that the dead

would be reburied in the new cemetery, that the grounds would

be “tastefully laid out, and adorned with trees and shrubbery,”

and that the “whole expense,” not to exceed $35,000, would be ap¬

portioned among the cooperating states — each “to be assessed

according to its population, as indicated by its number of repre¬

sentatives in Congress.” Wills closed his lengthy letter with the

request that each governor appoint “an agent” to help carry out

the reburial project and serve on the cemetery commission.24

Salomon acted promptly. “You will please state to Mr. Wills,”

he instructed his efficient and capable representative, “that the

32 Wills to Curtin, Aug. 11, 1863, Curtin Letter Books, Executive Correspondence,

1861-1865.

33 Selleck to Salomon, Aug-. 12, 1863, Executive Department (Administration), Civil

War Memorials Correspondence, 1861-1913, Archives Division, State Historical Society

of Wisconsin. The author is indebted to Miss Patricia Hosey, a research assistant in

the History Department and a graduate student [she received her M.A. in May of

1972] at Marquette for locating- the above correspondence — about 20 of these misfiled

letters were pertinent to this article.

24 Wills to Salomon, Aug-. 12, 1863, ibid. The letter, in Wills’ own hand, was evi¬

dently the basis for the printed circular letter which the author found in the David

Tod Papers (Ohio Historical Society) and the Austin Blair Papers (Burton Historical

Collection, Detroit Public Library).

1973] Klement — Soldiers’ Cemetery at Gettysburg 17

State of Wisconsin will bear its portion of the contemplated ex¬

penditure within the limit, and on the basis proposed, and you are

hereby designated as the Agent of this State to act in connection

with Mr. Wills and the agents of other States, in making the

necessary arrangements for completing the work.” The governor

added a timely warning, saying that he would expect a “full re¬

port” which would be laid before the legislature and suggesting

“the preserving of all necessary memoranda for the purpose.”25

Selleck, back in Washington when he received Salomon’s letter

which officially designated him as state’s agent for the Gettysburg

cemetery project, confirmed receipt of his new assignment by re¬

turn mail. “I will follow out your instruction,” he assured his gov¬

ernor, “and preserve all necessary vouchers and memoranda and

report in full in due time.”26 Selleck also promptly informed Wills

of his official appointment as Wisconsin’s agent to work with him

in the transformation of a portion of the battlefield into a national

cemetery,27 and after putting his Washington house in order he

hurried off to Gettysburg.

When William Saunders, charged with “laying out the grounds,”

arrived in Gettysburg, Wills, Selleck, and several other state agents

accompanied him to the site of the proposed cemetery. They walked

over the seventeen-acre tract as Saunders studied the high and

low portions and considered the best spot to locate the central

monument. In time, the eminent landscape gardener recommended

that the monument be placed on the highest reach of ground and

that the parcel to be allotted to each state run toward the common

center, fitting together in a semicircular arrangement. He promised

the services of a surveyor to lay out the 12-foot wide semicircular

parallels, allowing five feet for a walk between the parallels and

seven feet for each grave.28

Selleck spent some of his time during this stay in Gettysburg

calling on the Wisconsin soldiers recuperating in hospitals or in

private homes. He fretted about the intense summer heat, fearing

its effect upon some of the wounded. He was pleased, on the other

hand, that the citizens of the Gettysburg area had been so solicitous

about the welfare of the wounded and sick soldiers. “The people of

Gettysburg,” Selleck reported to Salomon, “are very attentive in

supplying the wants and looking after the comforts of the wounded

soldiers.” Selleck added a sentence about the cemetery project:

“I will write you to-morrow respecting the plans and arrangements

25 Salomon to Selleck, Aug-. 24, 1863, Letter Books General , Executive Department

(Administration), 1861-1865.

20 Selleck to Salomon, Aug-. 30, 1863, Executive Department (Administration), Civil

War Memorials Correspondence, 1861—1913.

27 Selleck to Wills, Aug-. 31, 1863, ibid.

28 Report of William Saunders, published in Revised Report of the Select Committee

. . pp. 37-38.

18 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

proposed for the laying out of the National Cemetery on the late

Battle Field.”29

In the weeks which followed Selleck returned to Washington to

minister to the needs of soldiers in that area while Wills dealt with

the problems arising out of the proposed cemetery. A controversy

developed over the question whether the reburials in the cemetery

would be “by state” or “promiscuous” and for a time Massa¬

chusetts even threatened to withdraw from the venture unless

she could have her way — each state’s soldiers buried together. At

Governor Curtin’s prompting, Wills was tempted to discard his

“promiscuous” plan and surrender to Massachusetts demands.

Then, after Saunders’ surveyor laid out the state sections and

cemetery plots, Wills was ready to publicize the reburial specifica¬

tions and plan the dedication program. Actually, he did not plan

to have the reburying begun until after the grounds were dedi¬

cated in ceremonies he set for October 22, 1863. Nor did he wish

to begin the reburial program until late fall. “I think it would be

showing only the proper respect for the health of this community

not to commence the exhuming of the dead, and removal to the

cemetery,” he wrote to Governor Curtin, “and in the meantime the

grounds should be artistically laid out, and consecrated by appro¬

priate ceremonies.”30

After Wills invited Edward Everett, the renowned Massachusetts

orator, to give the oration for the occasion, he wrote another round

of letters to the governors. He informed them that satisfactory

progress had been made on the cemetery project, that he had set

October 22 as the dedication date, and that Everett had been in¬

vited to give the day’s oration. He asked each governor — including

Salomon — whether his state preferred promiscuous reburials or

grouping by states. And he invited each governor to attend the

ceremonies in person and expressed the hope that each state would

have a good-sized delegation in attendance.31

Salomon again replied promptly. He preferred that Wisconsin’s

dead soldiers, killed at Gettysburg, be buried together, so that im¬

mediate comrades in life could be together in death. “The selection

of Hon. Ed. Everett as orator,” Salomon added, “is eminently

satisfactory.”32

Everett, meanwhile, expressed the desire to give the oration for

the dedication of the cemetery, but commitments in hand would

29 Selleck to Salomon, Aug-. 18, 1863, published in Wisconsin State Journal , Aug-. 19,

1863.

30 Wills to “His Excellency, A. G. Curtin,” Aug-. 17, 1863, published in Report of the

Select Committee . . . (Harrisburg, 1864), pp. 67-68.

31 Wills to Salomon, Sept. 15, 1863, Executive Department (Administration), Civil

War Memorials Correspondence, 1861-1913.

^William H. Watson [for Salomon] to Wills, Sept. 22, 1863, Letter Books General,

Executive Department (Administration), 1861—1865.

1973] Klement — Soldiers' Cemetery at Gettysburg 19

make it impossible to compose and memorize an oration before

November 19. 33 Since Wills had his heart set on Everett as the

day’s orator, he had to change the date of the dedication ceremonies

from October 22 to November 19.

While Wills and Everett were exchanging letters and changing

the date of the proposed ceremonies, Selleck wondered about the

progress of the cemetery project while he performed his regular

duties in the hospitals and army camps in the Washington area.

He fretted about the lack of information from Wills, and since he

was returning to Wisconsin for a hurried trip, he wanted to take

a progress report to his governor. “I intend going to Wisconsin

the last of next week and shall see the Governor,” he wrote on

October 9, “and I desire to be able to give him some information

in reference to the matter.”34 Thus prodded, Wills wrote a brief

note to Selleck, assuring him that he was “pushing the matter”

vigorously and enclosing a W ills-to-Salomon letter. “The enclosed

letter,” Wills wrote to Selleck, “gives you the information desired

in your letter.”35

The “enclosed letter” told Salomon that, at Edward Everett’s

insistence, the proposed date for the dedication ceremonies had

been changed to November 19. The change in dates also made it

imperative that the reburials begin before, rather than after the

official dedication ceremonies. In the first place, heavy frosts and

an early winter might delay the exhumations until spring. In the

second place, there was fear of a pestilence in the Gettysburg area.

Furthermore, Edward Everett had suggested that “it would make

the scene more impressive to have the dead interred in the Ceme¬

tery for the occasion.” Wills added that he had adopted Everett’s

suggestion and planned to make “arrangements” to commence the

“reburials” about the “26th” of the month. The Secretary of War

had promised to furnish “the requisite number of coffins- — supply¬

ing them at the rate of one hundred per day.” Curtin’s competent

agent also expressed the hope that Governor Salomon, with a large

Wisconsin delegation, would be present “to participate with the

consecration exercises” on the 19th of November.36

Consulting Selleck and several other state agents, Wills finalized

the “reburial specifications” and invited interested parties to bid

on the grisly chore. These “specifications” not only set standards

and procedures for exhuming the bodies and reburying them in

the proper plots in the new cemetery, but limited the number of

33 Everett to “My dear Sir” [Wills], Sept. 26, 1863, published in Report of the Select

Committee . . ., pp. 69—70.

34 Selleck to Wills, Oct. 9, 1863, Executive Department (Administration), Civil War

Memorials Correspondence, 1861—1913.

35 Wills to Selleck, Oct. 13, 1863, ibid.

30 Wills to Salomon, Oct. 13, 1863, ibid.

20 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

reburials to one hundred per day. Everything must be done with

care and under close supervision. 37-

After opening the bids on October 22, Wills awarded the con¬

tract to Frederick W. Biesecker, the lowest bidder — the bids had

ranged from a low of $1.59 per body to a high of $8.00. Then

Wills hired Samuel Weaver to superintend the exhumation, identi¬

fication, and reburial of the Union dead.38

While Wills kept a watchful eye on the progress of the reburials

and planned the program for the 19th, Selleck worked long hours

as state agent in and around Washington. He visited sick and

wounded soldiers in the nearby hospitals and performed sundry

miscellaneous chores. He secured furloughs for several of the

“walking wounded,” checked out a report that a Wisconsin soldier

had been “disloyal,” delivered a new regimental banner to the

battle-scarred Seventh Wisconsin, and looked into the case of “2

soldiers at the General Hospital in Baltimore.”39 “Mr. W. Y. Sel¬

leck, the State Agent,” a newspaperman reported, “is again at his

post, and he is busy, as is also his faithful assistant, Wm. P. Tay¬

lor, Esq., attending to the wants of the wounded Wisconsin men

as they come into the [Washington] hospital.”40 Governor Salo¬

mon, on a trip to the national capital, stopped at Selleck’s quarters

to inquire about his work and to say that affairs back home would

prevent him from attending the dedication ceremonies on Novem¬

ber 19.41

Selleck, thus, inherited the unpleasant task of telling Wills that

Wisconsin’s governor would be among the missing at the dedica¬

tion rites. “He also instructs me to inform you,” Selleck wrote,

“that he has authorized me to act for him on behalf of the State

of Wisconsin in the matter pertaining to the Soldiers’ National

Cemetery at Gettysburg.” Selleck added that he was returning with

Governor Salomon to Wisconsin and would not get to Gettysburg

until early in November.42

In late October Wills asked Ward H. Lamon, Lincoln’s confidant,

bodyguard, and U.S. Marshal for the District of Columbia, to serve

as chief marshal for the procession and master-of-ceremonies for

the program of the 19th. Lamon, in turn, wrote to each governor,

asking him to name “two suitable persons” to serve as assistant

37 “Specifications,” dated Oct. 15, 1863, published in Revised Report of the Select

Committee . . pp. 14-15.

38 Report, Wills to Curtin, March 21, 1864, published in ibid.

30 William H. Watson [for Salomon] to Selleck, Aug-. 25, 28, Sept. 10, Oct. 7, 14,

Nov. 7, 1863, Letter Books General, Executive Department (Administration), 1861-

1865.

40 Milwaukee Sentinel, Nov. 17, 1863.

41 Selleck to Wills, Oct. 16, 1863, Executive Department (Administration), Civil

War Memorials Correspondence, 1861—1913.

42 Ibid.

1973] Klement — Soldiers’ Cemetery at Gettysburg 21

marshals on the day of the dedication of the soldiers’ cemetery,

helping to supervise the day’s events.43

Since Selleck had performed his agent’s duties well and since

Washington was much closer to Gettysburg than Madison, Gover¬

nor Salomon asked him to assume the additional responsibility of

serving as assistant marshal in Gettysburg on the day of the dedi¬

cation program. Selleck, therefore, took the train for Gettysburg

to confer again with Wills and to discuss his new responsibilities

with Ward H. Lamon, chief marshal for affairs on the 19th.

Wills, pleased with the cooperation he had received from the

various state agents, let his enthusiasm run away with him. He

expected an “immense concourse of people” to descend upon Gettys¬

burg for the ceremonies and he supposed that most of the gover¬

nors would be present in person with large delegations.44 Caught

up in the enthusiasm emanating from Wills, one newspaper editor

predicted that the consecration ceremonies would be “the most

interesting ever witnessed in the United States” and that the affair

would be “one of the most imposing spectacles of this century.”45

Both Wills and Curtin expected most of the governors to attend

the ceremonies, and they therefore arranged for a special Harris-

burg-to-Gettysburg train, leaving the capital city at one o’clock in

the afternoon. Most of the governors, Curtin assumed, would come

to Harrisburg the day before the ceremonies, then they could pro¬

ceed to Gettysburg together as his guests. “We learn that the Gov¬

ernors of all the loyal States will assemble in this city on the 17th

inst.,” wrote the editor of a Harrisburg newspaper, “where they

will remain until the 18th, and on that day proceed in a body to

Gettysburg” — aboard the governor’s special train. Believing that

no “like assemblage” had ever taken place “in the career of the

country,” the editor hoped that the city’s citizens would give “the

distinguished guests . . . the hospitable welcome for which the

people of the capital of Pennsylvania are celebrated.”46

Evidently Wills had failed to inform Curtin that the governor

of Wisconsin would not be present in person for the ceremonies.

“I will be pleased,” Curtin telegraphed to Salomon on November

12, “to see you at Harrisburg on the seventeenth or eighteenth

on your way to Gettysburg — -arrangements will be made to leave

Harrisburg at one p.m. on the eighteenth for Gettysburg.”47 On

the same day Wills also sent a telegram to Salomon : “Let your

delegation bring your state flag for the nineteenth (19) inst.”48

43 Lamon to Tod, Nov. 5, 1863, Tod Papers.

44 Wills to Curtin, Nov. 7, 1863, published in Adams County Sentinel, Nov. 10, 1863.

45 Harrisburg- Evening Telegraph, Nov. 10, 1863.

46 Ibid., Nov. 13, 1863.

47 Telegrams, 1861—1865, Executive Department, Archives Section, State Historical

Society of Wisconsin.

48 Ibid.

22 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Selleck witnessed the inundation of Gettysburg- by a human flood.

Some had come on foot, having trudged the dusty sideroads and

the turnpikes ; some had come on horseback, riding either bareback

or in saddle ; many had come in carriages and farm- wagons ; most,

however, had taken one of the numerous special or excursion trains

which had come into the once sleepy city. Every house and shed

and stable was turned into a lodging-house.49 Most of those who

arrived on the 18th spent a part of the day touring the battlefield.

Selleck, delegated to “shepherd” Wisconsin citizens who came to

attend the dedication ceremonies, invited every Wisconsin soldier

or civilian he met to march in the procession and follow the state

flag on its trip from downtown Gettysburg to the top of Cemetery

Hill where a platform had been erected for the formal program.

During the evening of the 18th, Selleck attended a briefing ses¬

sion called by Ward H. Lamon, in charge of the next day’s proces¬

sion and program. Lamon explained the duties of the assistant

marshals and specified on which street corner each state’s delega¬

tion would gather and what place each state had in the procession.50

Earlier, Lamon had spelled out the “proper attire” for his corps

of assistant marshals:

1. Plain black suit (preferably a frock coat), black hat, and white gloves.

2. White satin scarf, five inches wide, to be worn over the right shoulder

and carried across the breast and back to the left hip, and there

fastened with a rosette, the ends to be fringed, and to extend to the

knee. At the center on the shoulder the scarf should be gathered and

mounted with a rosette.

3. Rosette, four inches and raised in center to be made of black and

white ribbon, the outer circle only to be white.

4. Rosette of red, white, and blue on left breast. The initials of state

in center for identification. The saddle cloths on their horses, of white

cambric bordered with black.51

Earlier the Chief Marshal had also instructed Selleck and the other

assistant marshals to secure their own horses and he promised each

a place on the program platform after the procession reached the

top of Cemetery Hill.

Dressed in the assistant marshal’s regalia and mounted on a

horse, Selleck spent the early morning hours of the 19th directing

Wisconsin soldiers and civilians to the street corner assigned the

state’s delegation. As the crowd in downtown Gettysburg swelled

to “immense proportions,”52 it became a milling mass of humanity,

resisting efforts of the assistant marshals to transform it into a

procession.

49 Gettysburg Compiler, Nov. 23, 1863.

50 Entry of November 23, 1863, in Diary of John Hay (microfilm), John Hay Papers,

Library of Congress.

51 Adams County Sentinel, Nov. 17, 1863; Washington Daily Morning Chronicle, Nov.

13, 1863.

52 Gettysburg- Compiler, Nov. 23, 1863.

1973] Klement — Soldiers’ Cemetery at Gettysburg 23

Shortly after ten o'clock, the scheduled hour for the procession

to begin its one-half mile southward trek, President Lincoln

emerged from David Wills' house to take his assigned place in the

cortege. It took nearly an hour for Ward H. Lamon and his assist¬

ant marshals to get the show on the road, down the Taneytown

Road and to the summit of Cemetery Hill. “Pennsylvania,'' an

observant newsman reported, “furnished the largest numerical

representation, Ohio next, Wisconsin third and Massachusetts

fourth."53 Selleck, evidently, had done a good job of rounding up

Wisconsin's soldiers and civilians to march in the state's delegation.

It took time to put the military units in their previously assigned

places atop Cemetery Hill, seat the dignitaries on the 20' x 12'

platform and herd the thousands of marchers into a semicircle in

front of it. Selleck, serving both as Lamon's aide and the state's

representative on the proposed cemetery commission, was the lone

Wisconsin resident to occupy a place on the platform.

Selleck, thus, was in a good position to hear every speaker and

watch the crowd's reaction to each and to the musical numbers.

Borgfield's Band of Philadelphia opened the formal program with

a solemn dirge. The Rev. Thomas H. Stockton, chaplain of the

House of Representatives and a popular Washington preacher,,

gave the invocation — “ a prayer which thought it was an oration."54

The Marine Band then played Luther's hymn, “Old Hundred."55

Next Ward H. Lamon, serving as master of ceremonies, introduced

Edward Everett, orator of the day. Although Everett delivered

his two-hour recitation “with his accustomed grace," it was

“smooth and cold," without “one stirring thought, one vivid pic¬

ture, one thrilling appeal."56 A chorus of twelve members of the

Maryland Musical Association then chanted an ode, drew consider¬

able applause, and set the stage for Lincoln's “few appropriate

remarks."

After Lamon introduced “the President of the United States,"

Selleck watched Lincoln arise from his chair and step forward to

keep his rendezvous with destiny. The crowd interrupted Lincoln's

brief address with applause five times and complimented him with

“tremendous applause" when he finished the dedicatory “remarks."

The responsive audience then gave three cheers for the president

and three more for the governors.57

63 Boston Journal , Nov. 23, 1863.

54 Entry of November 23, 1863, in Diary of John Hay, John Hay Papers.

65 Adams County Sentinel, Nov. 24, 1863.

66 Harper’s Weekly, Dec. 5, 1863.

57 The Cincinnati Daily Commercial, Nov. 23, 1863, and the Washington Daily

MorwAng Chronicle, Nov. 20, 1863, carried the most complete accounts of the program.

Martin R. Potter of the Commercial not only witnessed the ceremonies but sent back

a report which filled seven full columns of type. The best secondary account, marred

by occasional errors of fact, is Louis A. Warren, Lincoln’s Gettysburg Declaration:

“ A Neiv Birth of Freedom” (Fort Wayne, Ind., 1964).

24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

After Lincoln’s memorable performance, a mixed chorus of

Gettysburgers sang a doleful dirge and finally the Rev. Henry L.

Baugher, president of Gettysburg (Lutheran) Seminary, closed the

formal program with a brief benediction. An eight-round salute,

fired by the Fifth New York Artillery, formalized the end of the

ceremonies. Marshals reformed the military portion of the proces¬

sion to escort President Lincoln back to Gettysburg and the Wills

house, while the civilian portion “dispersed in all directions.”58

“Many lingered until the shades of evening approached,” noted

one newspaperman, “seemingly loath to leave the ground conse¬

crated to the blood of those heroes who fought, and died, and found

a grave there.”59

Selleck returned to Washington, D. C., to pursue his manifold

duties in the nearby camps and hospitals. Later he submitted a bill

of $30.90 for “expenses in attending the consecration of the Na¬

tional Cemetery at Gettysburg.”60

Perhaps Selleck was disappointed that his governor, Edward

Salomon, had failed to attend the ceremonies. Selleck also could

have complained that some Wisconsin men who came to Gettys¬

burg for the dedication of the cemetery, neither marched in the

procession nor gathered around the platform atop Cemetery Hill

to watch and hear the speakers and the music. 1st Lt. Frank A.

Haskell, who had fought in the three-day battle and later wrote a

famous letter which described the encounter, spent November 19

visiting those parts of the battlefield where Wisconsin boys had

fought and died. “We obtained horses,” Haskell wrote to his

brother, “and during the afternoon of . . . the 19th we rode all over

the field ... we had little interest in the ceremonies. . . .”61

The conclusion of the dedication ceremonies did not sever Sel-

leck’s link to the cemetery project. Early in December, Wills wrote

a final round of letters to the cooperating governors — including

Edward Salomon of Wisconsin. The letter set December 17 as the

meeting date of the “commissioners” of the cooperative state

cemetery, and Wills expressed the hope that each of the eighteen

states which had soldiers buried at Gettysburg would have repre¬

sentatives present at the Jones House in Harrisburg to devise “a

plan for the protection and preservation of the grounds,” complete

the work already begun, arrange for the proper adornment and

care of the grounds, and provide for expenses already incurred.62

£8 Cincinnati Daily Commercial, Nov. 23, 1863.

69 Washington Daily Morning Chronicle, Nov. 20, 1863.

60 William H. Watson to Selleck, Dec. 23, 1863, Letter Books General, Nov. 11,

1863-Feb. 18, 1863, Executive Department (Administration).

B1 Frank A. Haskell to Hiram M. Haskell (brother), Nov. 20, 1863, Haskell Papers,

State Historical Society of Wisconsin.

62 Wills to Selleck, Dec. 3, 1863, Executive Department (Administration), Civil War

Memorials Correspondence, 1861—1913.

1973] Klement — Soldiers' Cemetery at Gettysburg 25

Governor Salomon, of course, instructed Selleck to represent the

state at the December 17 meeting of the commission. On that day

twelve commissioners (including W. Yates Selleck) representing

ten states assembled at the Jones House. Wills, presiding over the

ad hoc assembly, suggested formal organization as a commission

or association. The delegates responded by electing Wills as presi¬

dent and Selleck as secretary. The commissioners then adopted

five resolutions, all concerned with the completion and operation

of the eighteen-state “Soldiers’ National Cemetery.”63 On motion

of Commissioner Levi Scoby of New Jersey, Wills appointed a

five-member committee “to procure designs of a monument to be

erected in the cemetery.”64

After the commission concluded its business, Selleck returned to

Washington to resume his chores as state agent and to write a

report on his role in the establishment and administration of the

Soldiers’ National Cemetery. In the concluding paragraphs of this

report he called Salomon’s attention to the fact that the amount

expended for the cemetery nearly doubled that stated by Wills in

“his circular letter of August last.” Selleck listed the reasons : “1st.

There are seventeen acres to be enclosed instead of fourteen as at

first proposed. 2d. The sum to be expended on the Monument,

$25,000,, instead of $10,000, as at first proposed. 3d. That in the

laying out and ornamenting of the grounds and the finishing and

placing of the head-stones to the graves of soldiers, would, if

properly done, be more expensive than at first calculated.”65

Selleck stated that the sum of $63,500, “designated for the com¬

pletion of the Cemetery” should “more than cover the expendi¬

tures”— “if judiciously handled.” Wisconsin’s share of the $63,500,

Selleck projected, should be $2,523.13, or $420.53 for each of her

six members in Congress.66

In his lengthy report, Selleck also stated that many Wisconsin

soldiers, mostly belonging to the “Iron Brigade,” were buried in

that section of the cemetery reserved for “the unknowns.” “Nearly

all of the remains of the Union Soldiers killed in the battle of

Gettysburg,” Selleck added, “have been removed to the Cemetery;

all of those killed in the first day’s fighting have been removed; a

great many of them were not identified ; such, are placed in the lots

that are marked unknown!”®1

63 The cemetery remained a cooperative state venture until 1895, when it became

a national cemetery within the Gettysburg- National Military Park.

64 Report of the commissioners, dated December 17, 1863, and signed by David Wills

as “President” and W. Yates Selleck as “Secretary,” in Tod Papers. A copy of the

printed proceedings is in the Civil War Memorials Correspondence, 1861-1913.

65 Selleck to “His Excellency Edward Salomon,” Dec. 28, 1863, published in Journal

of the Proceedings of the Assembly of Wisconsin . . . 186 Jt (Madison, 1864), pp.

302-306.

66 Ibid.

Ibid.

26 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

The Wisconsin State Legislature, at the governor’s recommenda¬

tion, passed a bill appropriating $2,523 to cover the state’s “share”

of the costs of the Gettysburg cemetery project, indemnifying, in

effect, both Salomon and Selleck for their roles in the cooperative

venture.68

In the days and months which followed, the cemetery project

moved towards its completion. After spring’s mild weather thawed

the frozen ground, the contractor responsible for exhuming the

bodies and reburying them in the appropriate plots of the “Sol¬

diers’ National Cemetery,” returned to his grisly task. The reburial

crew finished its work on March 18, 1864. “And I here most con¬

scientiously assert,” the superintendent of the project proudly

wrote, “that I firmly believe that there has not been a single mis¬

take made in the removal of the soldiers to the cemetery by taking

the body of a rebel for a Union soldier.”69

Of the 3,512 reported buried by March 18, 1864, seventy-three

were recorded as Wisconsin soldiers, including twenty “unknowns”

who could not be identified by name.70

In late March, 1864, the legislature of Pennsylvania passed an

act to incorporate the “Soldiers’ National Cemetery,” thus final¬

izing David Wills’ dream — transforming a portion of the battle¬

field into “a final resting place for those who gave their lives that

the nation might live.” The articles of incorporation listed “W.

Yates Selleck, Wisconsin” as one of the “corporators.”71

Selleck, subsequently, attended two meetings of the Board of

Commissioners of the new cemetery, one on April 6-7 and the

other on June 10, 1864 — both held in Gettysburg. The committee

named to select the design for the national monument to be located

in the cemetery recommended a 60-foot high memorial, with a

massive pedestal twenty-five feet square at the base and the fancy

column crowned with a colossal statue representing the Genius of

Liberty. The monument, completed as designed, would cost $102,-

000. The commissioners, concerned with the burgeoning costs of

the project, adopted a resolution asking Congress to appropriate

“the sum of fifty thousand dollars” toward the monument.72

In late July, 1864, Selleck returned to Milwaukee for a visit

and then took a trip to Madison to report on his work as a com¬

missioner of the “Soldiers’ National Cemetery” and as state agent

in the Washington area. General U. S. Grant’s heavy losses in the

68 Journal of the Proceedings of the Senate of Wisconsin . . . 1864 (Madison, 1864),

p. 709.

69 Samuel Weaver, report of March 18, 1864, published in Revised Report of the

Select Committee . . ., pp. 16—18.

79 Ibid.

71 Milwaukee Sentinel, April 5, 1864.

72 Report, Selleck to James T. Lewis, Feb. 15, 1865, published in Journal of the

Proceedings of the Assembly . . . 1865 (Madison, 1865), pp. 409—411.

1973] Klement — Soldiers' Cemetery at Gettysburg 27

battles of the Wilderness (May 5-6), Spottsylvania (May 8-12),

and Cold Harbor (June 3), taxed Selleck’s resources and multiplied

his work. The Milwaukee Sentinel reported on Selleck’s return

and praised him as an “efficient” state agent.73

During the early months of 1865, as the war moved into its final

phase, W. Yates Selleck wrote his second official report regarding

his role in the establishment of the soldiers’ cemetery at Gettys¬

burg. He addressed this second report to Governor James T. Lewis

who had replaced Salomon as the state’s chief executive early in

January.74 Selleck reported on the April 6-7 and June 10, 1864,

meetings which he had attended and he justified the proposal for

a $102,000 national monument. “Every portion of the work and

material used in and about the Cemetery for its adornment and

preservation,” Selleck stated, “is of the most substantial kind and

put up in the most durable manner, and will, when completed, be

a most attractive and beautiful place; a credit to the country and

a noble monument to the gallant and loyal dead who rest within

its limits.” Wisconsin’s share of the added expenses for the com¬

pletion of the “Soldiers’ National Cemetery” were $2,623.00. And

he asked the governor to request an appropriation from the state

legislature.75

The legislative Committee on State Affairs quickly and quietly

recommended payment of the $2,623 which Selleck requested.

“Other states,” the committee recommendation read, “are con¬

tributing to this national and patriotic object, and Wisconsin,

never behind in all concerning her own and the nation’s honor,

and the lives and memory of her gallant volunteers, will not be so

in helping to complete a work which largely commemorate the

devotion of worthy sons whose deaths have passed to her credit

on the scroll of honor and heroic fame.”76

The end of the war and the demobilization of Wisconsin troops

signified that Selleck’s work as state agent was about at an end.

He tendered his resignation as “Military Agent for the State of

Wisconsin” on May 29, 1865, asking that “it take effect on the

1st of June, 1865.” In his Iketter of resignation he stated that he

had conscientiously looked after “the interests and welfare of Wis¬

consin soldiers” for three years, “rendering to them aid as far as

the means placed in my hands” would allow. “As the rebellion is

7:5 Milwaukee Sentinel , July 29, 1864.

7t The Republicans did not nominate Salomon for re-election, choosing- Lewis in his

stead, a bitter pill for Salomon who had performed his duties efficiently and with

ability. Salomon resumed law practice in Milwaukee. Unhappy that Wisconsin Re¬

publicans had failed to support him for re-election, Salomon moved to New York City

in 1869. After his retirement from law in 1894, Salomon returned to Germany to live

at Frankfort-on-Main, where he died and was buried.

75 Report, Selleck to Lewis, Feb. 15, 1865, published in Journal of the Proceedings

of the Assembly . . . 1865 , pp. 409-411.

76 Report of the Committee on State Affairs, James Ross, chairman, in ibid., 489.

28 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

over and our victorious troops are now about to return to their

homes/’ he concluded, “I feel that my services will not be too much

longer required, and my private affairs need my attention and

impose upon me the necessity of resigning.”77

Although Selleck resigned his post as “state military agent/’ he

remained Wisconsin’s representative on the Gettysburg cemetery

commission. In that capacity, he attended the July 4, 1865 cere¬

monies held in connection with the laying of the cornerstone of the

National Monument — Wisconsin’s governor, James T. Lewis did

not attend.

In a sense, the dedication of the monument indicated that David

Wills’ dream of a national cemetery at Gettysburg had been

realized. Both Salomon and Selleck had helped Wills transform his

dream into a reality. Salomon supported the project from the very

beginning, always cooperating with both Wills and Selleck. And

W. Yates Selleck, as a competent and conscientious state agent,

worked closely with Wills to develop, promote, and finalize the

project, transforming a portion of the battlefield, consecrated by

the blood of the “Iron Brigade” and other Wisconsin troops, into

“a final resting place” for “the honored dead.”

77 Selleck to “His Excellency, Jas. T. Lewis, Gov. of Wisconsin,” May 29, 1865,

published in Milwaukee Sentinel, June 23, 1865. Six months after resigning- as state

agent, Selleck left Wisconsin for Connecticut. The Milwaukee Sentinel of Jan. 26, 1866,

carried the following story: “W. Y. Selleck, Esq., of this city, leaves to-day for Hart¬

ford, Conn., where he has accepted the responsible position of Special Adjusting Agent

of the Traveler’s Insurance Company. Mr. Selleck is well known to a great number

of Wisconsin soldiers as late the efficient Military State Agent of Wisconsin in

Washington. The duties of this office were performed with great faithfulness and

tact, and many a hundred of the Badger State’s ‘boys in blue’ will hold his name in

grateful remembrance for the valuable service which he has rendered them, and will

unite with his friends in Milwaukee in wishing him success in the new horizon in which

he is now to labor. We congratulate the insurance company in securing so efficient

an agent.”

MAN’S CREEPING JURISDICTION OVER

OCEAN SPACE

John B. Ray

University of Wisconsin —

Whitewater

ABSTRACT

For the past three decades scientists have been identifying the

wealth of natural resources existing in the oceans and have been

developing the technology to extract and to utilize these resources.

Following such developments, coastal states unilaterally have begun

to declare larger segments of ocean space under their jurisdiction

and sovereignty. Recent trends include : the seaward extension of

the territorial sea from a small band of three miles to a maximum

of two hundred miles ; the establishment of exclusive fishery zones ;

the creation of conservation zones designed for pollution and fishery

control; and the extension of sovereignty over the adjacent conti¬

nental shelf beyond the limits of national jurisdiction. Immediate

definitive action toward the creation of a viable international ocean

regime, beneficial to all mankind, is essential to prevent a future

competitive scramble for ocean space.

As population pressure expands and technology advances, enor¬

mous stresses are being placed upon the earth’s natural resources ;

thus the world community is beginning to focus greater attention

upon the chemical, biological, and geological resources of ocean

space. Scientists are recognizing that the rewards to be had from

the oceans are substantial; but to whom do these rewards belong?

Are they the exclusive possession of the coastal states or are they

a common heritage of all mankind? The problem of ownership of

the seas’ resources is becoming an increasing source of world con¬

flict as vested interests expand.

In past centuries, geographers have viewed the oceans in the

dual role as highways for ships and as hunting grounds for fisher¬

men. With the age of discovery, the oceans assumed a new dimen¬

sion as avenues of conquest. Ultimately these activities led to a

period of saltwater imperialism as continents were discovered,

settled, and penetrated by Europeans. At first, sailing ships carry¬

ing goods of commerce linked these continents. Later with the

invention of the steamship these commercial endeavors were

29

30 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

intensified. The steamship, too, along with other technological

advances in shipping made possible easier connections with distant

fishing grounds and greatly enlarged the fish catch.

Yet while the surface of the oceans was being utilized more and

more intensively by man, the ocean depth, including the seabed,

continued to be shrouded in mystery and legend with little thought

given to exploitation.

In time, however, the seas began to lend themselves to tunneling,

the laying of cables, and submarine travel. Today they are a prin¬

cipal area of military deployment and maneuver and harbor mili¬

tary weapons and equipment. And much to the exacerbation of

marine geographers and other thinking people around the world,

the ocean environment has become an open sewer for man with

every conceivable pollutant poured into it, including the recent

atomic waste and mustard gas.

Problems associated with the surface of the oceans are centuries

old. Yet the question of the continental shelf has emerged only

recently, in fact since 1945. In that year President Truman issued

his historic Proclamation No. 2667 in which the United States

laid claim to the subsoil and seabed of the adjacent continental

shelf, beyond the limits of national jurisdiction, but not to the

superjacent waters which lay above. This document, commonly

known as the Truman Declaration, marks the beginning of a new

era in ocean exploitation.

This era is recognized as one of rapid technological advances,

opening up both the biological and mineral resources of the seabed

to possible exploitation. In addition, this era has produced a new

period in international relations in which the continental shelf

and superjacent waters are being subject to an unprecedented

number of exclusive and competing national controls for which

there are no bases in law or custom, yet against which there is

little effective counter in international law enforcement.

Over the last three and one-half centuries, a complex body of

international law has evolved concerning the oceans, much of

which has been incorporated into the four United Nations’ Geneva

Conventions on the Law of the Sea adopted in 1958. 1 Unfortu¬

nately, these conventions have proven to be of limited value in

resolving international conflicts over access to ocean resources.

It appears that self interest and power, rather than any adherence

to principles laid down by the Geneva Conventions, most often

determine the course a state shall take in these matters.

1 These conventions include : The Convention on the Territorial Sea and the Con¬

tiguous Zone, The Convention on the High Seas, The Convention on Fishing and

Conservation of the Living Resources of the High Seas, and the Convention on the

Continental Shelf.

1973] Ray — Creeping Jurisdiction Over Ocean Space 31

“Never have national claims in adjacent seas been so numerous,

so varied, and so inconsistent,” wrote Boggs (1) two decades ago.

This geographer might well have been describing the increasingly

accelerated pace at which coastal states are presently carving out

interest areas within adjacent seas. Table 1 summarizes the current

patterns of claims exercised by coastal states in terms of the

breadth and status of the territorial sea, exclusive fishing zones,

conservation zones, and the continental shelf. The date shown in

parentheses after each country indicates the year in which a

specific claim was declared.

A three mile territorial sea has been commonly recognized in

the past, but there is no prohibition against broader claims as long

as these claims are recognized by the community of nations. This

pattern of expanding claims (Tables 1 and 2) can be most con¬

fusing, but a degree of rationale is evident in the fact that the

maritime powers maintain modest claims for themselves and gen¬

erally object to wider claims by other coastal states that would

restrict movement in any way upon the high seas.

The 1958 Geneva Convention on the Territorial Sea and the

Contiguous Zone is imprecise in not stating a maximum breadth

for the territorial sea. It did, however, recommend that the breadth

of this zone should not exceed 12 miles (4). A discernible upward

trend in territorial sea claims can be noted in Tables 1 and 2 ; and

yet except for a bloc of Latin American with 200 mile claims, the

trend is toward consolidation of a 12 mile limit in keeping with

the recommendations set forth in the Geneva Articles.

According to the 1958 Geneva Convention on the Continental

Shelf, a coastal state may exercise the sovereign right of exploring

and exploiting both the biological and mineral resources of the

continental shelf adjacent to its coast to a depth of 200 meters

(658.2 ft.) or beyond that limit to where the depth of the super¬

jacent waters permits exploitation of such natural resources (5).

This last exploitability clause has precipitated a rash of claims

to exclusive jurisdiction over the shelf beyond the 200 meter mark.

As a result there are in 1973 54 such claims as opposed to 20 in

1960. Basing its claims upon the exploitability clause, a coastal

state could presumably move its boundaries seaward to any dis¬

tance from the coast provided the state has the technology to ex¬

ploit the seabed resources. Unlikely, yet somewhat unsettling, this

clause could ultimately lead to the division of the ocean floor

among coastal states based upon the median line concept.

More specific functional claims, in Table 3, reveal few claims

in excess of 12 miles. However, those claims exceeding 12 miles

include a number of exclusive fishing zones and fishing conservation

zones. By these methods, coastal states hope to accomplish two

TABLE 1. BREADTH OF THE TERRITORIAL SEA AND RELATED ZONES (2)

32 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

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34

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36

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TABLE 1. (CONTINUED)

38

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40 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

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42 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

TABLE 2. TERRITORIAL SEA CLAIMS, 1960 and 1973

* Source of 1960 data was Brown (3).

distinct goals; 1) to prevent foreign nationals from taking fish

and depleting the fisheries near their shores; and 2) to provide

adequate breeding stock to replenish existing fisheries. The coun¬

tries of the Indian subcontinent and of west Africa (Table 1),

for example, are attempting to protect their coastal fisheries from

the covetous assault of the technologically advanced fishing coun¬

tries until such time as they are able to develop the technology to

effectively exploit the waters themselves.

Canada among the world states has shown unusual concern for

the problems of ocean pollution by creating a 100 mile pollution

prevention zone within the Arctic (Tables 1 and 3). The immediate

stimulus for this legislation was the historic 1969 round-trip

voyage of the U.S. Tanker SS Manhattan from a United States

east coast port to Prudhoe Bay, via the Northwest Passage (6).

Its general purpose was to demonstrate the feasibility of utilizing

ice-breaking supertankers to carry petroleum from the Alaskan

North Slope to Atlantic seaboard markets. The success of the ven¬

ture and the subsequent creation of the pollution prevention zone

a year later, underlined the urgency with which the Canadians

view the necessity of protecting the delicate ecosystem of the

Arctic.

TABLE 3. FUNCTIONAL ZONE CLAIMS IN EXCESS

OF THE TERRITORIAL SEA CLAIMS, 1973*

* Compiled from Table 1.

1973] Ray — Creeping Jurisdiction Over Ocean Space 43

CONCLUSION

One of the most urgent problems of our time is the unilateral

action taken by coastal states in extending their political control

over large segments of ocean space that were once free and open

to all peoples of the world community.

It is apparent that present trends cannot be permitted to con¬

tinue, otherwise they may precipitate a collapse in the present

law of the sea. It is also apparent that these trends cannot be

reversed until new regimes of political control are developed to

deal with the new situation. Each year that elapses without the

establishment of some form of international control, the domain of

vested interests expands ; and the area of ocean space to which an

international regime might be applicable shrinks.

Geographers and other social scientists view with anticipation

the Geneva Convention on the Law of the Sea which is scheduled

to convene in the spring of 1974. The ultimate goal of this con¬

vention will be to establish a broad international regime of prudent

ocean management over the last remaining frontier on earth.

Within this regime will be built expanded opportunities for the

small as well as the large states, the underdeveloped as well as the

technologically advanced states, and the landlocked as well as the

coastal states. Based upon international cooperation, the harvest

of chemical, biological, and geological resources of ocean space

will indeed be exploited and utilized as a common heritage, bene¬

ficial to all mankind.

REFERENCES

1. BOGGS, S. WHITTEMORE. 1951. National Claims in Adjacent Seas.

Geographical Review, 41 : 186.

2. Compiled from: Ocean Science News, 1972, 1973. New York Times, 1971,

1972, 1973. Christian Science Monitor, 1971, 1972, 1973. International

Legal Materials, The American Society of International Law, New

York, Vol. X, No. 6, November, 1971. FAO, Legislative Series, No. 8,

Rome, 1969.

3. BROWN, E. D. 1971. The Consequences of Nonagreement, Proceedings of

the Sixth Annual Conference of the Law of the Sea Institute. Univer¬

sity of Rhode Island, Kingston. Lewis M. Alexander, ed., 1971, p. 14.

4. United Nations Conference on the Law of Sea. Convention on the Terri¬

torial Sea and Contiguous Zone, April 29, 1958. (U.N. Doc./Conf.

13/L52, Article 24).

5. United Nations Conference on the Law of the Sea. Convention on the Con¬

tinental Shelf, April 29, 1958. (U.N. Doc. A/Conf. 13/L. 55, Article 1).

6. BILDER, RICHARD B. 1970. The Canadian Arctic Water Pollution Pre¬

vention Act: New Stresses on the Law of the Sea. Michigan Law Re¬

view, 69: November 1970, 3.

CULTURAL DIVERSITY IN CENTRAL WISCONSIN

Maurice E. Perret

University of Wisconsin —

Stevens Point

In rural areas and in small towns, America is not the melting

pot that idealists would like to see. Even though most of the farm

land has been permanently occupied for more than a century, in

many regions one ethnic group is still predominant. Immigrants

from Europe generally came in groups settling at one place. They

formed homogeneous communities of people with the same back¬

ground : language, religion, education, customs, political affinity,

and although mass immigration has long since come to a stand¬

still, many communities that were established by foreign pioneers

have persisted.

Central Wisconsin shows a number of such ethnic groups (see

Map 1). In Portage County, for instance, we find areas of concen¬

tration of Polish, German, Scandinavian, Bohemian people, to

which we may add the Amish who, although they came to this

country more than two hundred years ago, form a definite ethnic

group which still has more pecularities than any other foreign

group. Polish people occupy the central part of the county; their

area of concentration expands north to Marathon County and west

to Wood County. There are also a number of clusters of Polish

farms in all other towns. Scandinavian people are in the eastern

part, towards Waupaca County where this ethnic group is well

represented and where we find, for instance, the village of Scandi¬

navia; there are also a few clusters of Scandinavian farms in the

south. We note three areas of German people ; the largest around

Kellner, a smaller one in the northwest, both related to larger Ger¬

man colonies in Wood County. A third area of Germans is around

Almond. We find small clusters of Amish south of Amherst. Finally,

a number of Bohemian farmers are located near the Wood County

line, west of Junction City.

Each ethnic group presents some distinctive features. For in¬

stance, in the Polish section, we find still a few log constructions,

farm houses, barns, sheds, where the logs are assembled according

to the half-notch system (Fig. 1). Polish people are devout Cath¬

olics : In the region where they live, they have built several large

45

46 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

MAP 1. Areas of concentration of non Anglo-Saxon ethnic groups in Portage

County, in 1972.

(P = Polish, S = Scandinavian, G = German, A = Amish, B .= Bohemian)

MARATHON CO.

Kilometers

brick churches, some of which are dedicated to saints venerated

in Poland : Saint Casimir, Saint Bronislava, Saint Adalbert, Saint

Stanislaus. One may see also, along rural roads, small brick

shrines containing a crucifix or statues of the Virgin Mary and

of saints (Fig. 2) or high wooden crosses, with a crucifix pro¬

tected by a small tin roof, sometimes in a small enclosure full of

flowers (Fig. 3). They are similar to crosses also found along

rural roads in Central Europe. Often, near the farm house are

small statues of the Virgin Mary or of saints. In the cemeteries,

numerous monuments have crosses and many epitaphs are in

Polish. One may also see some “zoraw”, the well with a pole used

1973] Perret — Cultural Diversity in Central Wisconsin

47

FIGURE 1. Log construction near Stevens Point (Polish Farm house)

to pull up the buckets of water (Fig-. 4). In the area where Polish

are settled, we find many taverns, established at road intersec¬

tions. Some of them have large ball-rooms, used especially at

weddings. A few place names are typical : Polonia, Torun named

after a city in Poland, Pulaski School after the Polish hero of the

Indepedence War of the United States, and Lake Glisczinski.

Scandinavian farms present numerous buildings. In their re¬

gion we may see some old fences of the split rail type (Fig. 5)1.

There are also some log constructions, especially tall granaries,

in which the logs have been assembled according to the dove-tail

system (Fig. 6)2. Scandinavians do not favor taverns. The town

of New Hope is completely “dry”. In the cemeteries, many epi¬

taphs are in Norwegian, Danish or Swedish and the monuments

have no crosses. These farmers are Lutheran and in their region,

we find some isolated Lutheran churches. A few place names refer

1 Although typical of Scandinavian farms in Wisconsin, the split rail fences do not

seem to have been brought from the Northern countries of Europe, as no example

of them is found in the open air museums of Oslo, Copenhagen and Stockholm.

Norwegians probably saw them on some farm when they were coming to Wisconsin

and adopted them here.

- It seems that neither in Norway, nor in Sweden, nor in Denmark, the dove-tail

system of assembling logs has been used in rural constructions. In the open air

museums of the Northern countries, the only buildings with this system are from

cities in Sweden. All rural constructions seem to use the double notch assembling

system and have protruding log ends. The only rural building that is with dove-tail

assembled logs is a cabin built in 1871 by a Norwegian in Kindred, South Dakota,

and brought to the open air museum at Oslo. As in Wisconsin, the ends of the logs

have been sawed to form square corners.

]

48 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

FIGURE 2. Shrine on a farm near Stevens Point (Polish)

1973] Ferret — Cultural Diversity in Central Wisconsin

49

FIGURE 3. Cross with a crucifix, near Stevens Point

(Note the small tiny roof above the crucifix.) (Polish)

50

Wisconsin Academy of Sciences, Arts and Letters [Vol.

FIGURE 4. “Zoraw” — Pole or bascule well near Stevens

Point (Polish)

1973] Perret — Cultural Diversity in Central Wisconsin 51

FIGURE 5. Split rail fence near Rosholt, Portage County (Scandinavian)

FIGURE 6. Log granary, near Nelsonville, Portage County (Scandinavian

farm)

52 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

to Scandinavian settlers : The village of Rosholt was named after

one Norwegian who came from Waupaca County where he was

born. Lake Budsberg and Lake Reton bear names of Norwegian

families.

Germans have also some log constructions. They used the half¬

notch assemblying system, in the same manner as the Poles.

German farmers are Protestant, mostly Lutheran, but there are

also members of the Moravian sect and they have a church in

Kellner. In the cemeteries, before the first World War most epi¬

taphs of Germans were in the German language and many had

verses. The village of Kellner was named after one of the early

German settlers.

The Amish are new settlers. They are not pioneers but they

bought old established farms. Since their religion opposes modern

ways of life and modern technique, they are dressed in their tra¬

ditional garb. The women wear plain color dresses and white

aprons and have at all times a white headdress; when thev go

out they are draped in a black cape and wear a black bonnet. Men

wear black suits without buttons, shirts without collars, and black

felt hats. The young men are clean shaven but they grow a beard

when they get married. They speak between themselves their own

German dialect. They have their own one-room school-house and

on Sundays, they gather at one farm or another for the religious

services. They shun electricity, motors, telephone, radio, television

and for that reason, one does not see electrical wiring, automobiles

or tractors on their farms, but they use teams of horses for their

farm work, and for traveling they have horse-drawn buggies.

The present situation is not definitive. It is but one stage in

the evolution of the landscape. The first occupation of the land

by farmers was different and in most places, the present land-

owners are not descendants of the first settlers, nor do they be¬

long to the same ethnic group.3

The region was first inhabited by Indians who never were nu¬

merous. Some mounds are the only trace of their presence here.

Except the Wisconsin River, there is in the county no place name

of Indian origin. The first white settler was a Frenchman, John

Baptist Dubay who established in the early 1830’s a trading post

near the Wisconsin River, north of Stevens Point and he became

the first postmaster of the post office called Eau Pleine, the only

3 Maps 1 and 2 are based on the names of landowners as given in the platbooks.

They give a generalized picture. Most areas, however, do not form compact blocks

but among farms of the majority group are some farms of other background and

sometimes land belonging to corporations or to the government. Some names could

not be attributed with certainty to one ethnic group, there may be therefore some

slight errors. In the areas where no ethnic group is indicated, there may be in some

places a majority of Anglo-Saxon farmers, in other places no clear majority of one

ethnic group, elsewhere the land is owned by the government, by corporations or

institutions, or by owners who are not farmers.

1973] Perret — Cultural Diversity in Central Wisconsin 53

name of French origin in Portage County. A few years later, the

region which had large stands of white pines, attracted lumber¬

men. Sawmills were erected along the Wisconsin River and on

its tributaries : Mill Creek on the west side and the Plover River

on the east side. Most newcomers were Yankees from the East,

with some Canadians and Irishmen (one of these was the first

foreigner to apply for citizenship in Portage County, in 1845).

Following the lumbermen came some farmers who settled on the

land. Although mostly Yankees, there were some groups of French¬

men, Englishmen, Scots, and after the great famine in Ireland,

a number of Irishmen. As early as 1846, one Norwegian came

with the lumbermen and in 1850, a group of Norwegians settled

in the eastern part of the county and in the neighboring part

of Waupaca County. A few Danes joined them. Three distinct

German settlements were founded in the early fifties ; two were

Protestant, in the towns of Almond and Grant, the third one was

Catholic in the town of Sharon. The last important group to ar¬

rive in Portage County were the Poles. A few individuals, coming

about 1860 to the towns of Sharon and Stockton, were followed

by a continuous stream of immigrants of the same background.

At present the farmers of Polish origin constitute the largest

ethnic group. About fifteen years ago, a few Amish families,

coming from Iowa, bought farms in the vicinity of Amherst. Their

number has increased to some twenty five families and may still

increase, since a recent judgment of the Supreme Court gave the

Amish of Wisconsin the right to keep their children out of public

schools.

The oldest known plat map of Portage County shows the situa¬

tion in 1876, and most probably the names of the farm owners are

those of the original settlers or of their direct descendants. (Map

2 is based on it) . We find the large Scandinavian colony in the

eastern part of the county. The village of Rosholt is indicated, but

only for reference, as it did not exist in 1876. The Polish colony

has Polonia for center and extends west beyond the Wisconsin

River. We notice three German settlements, the largest near Al¬

mond, others around Kellner and in the town of Sharon. There

is a small French colony in the town of Buena Vista (which in¬

cidentally was not named by Spanish speaking immigrants but

by Yankees4). Irish farmers are found in three locations, in the

towns of Dewey, Stockton and Lanark. The rest of the county

was partly occupied by Yankee farmers or was still in woods,

swamps, prairies, either in the hands of the government or be¬

longing to corporations or to some landowners who were not

4 At the origin there was a tavern named after another tavern in Southern Wiscon¬

sin which had received its name just after the battle of Buena Vista.

54 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

MAP 2. Foreign colonies in Portage County, in 1876.

(G = German, P = Polish, S = Scandinavian, I = Irish, F = French)

MARATHON CO.

6 2 4 6

4 6

t-HH

6

o

cn

oo

Kilometers

farmers. This explains why most of the place names are of

Anglo-Saxon origin, some named after cities from which immi¬

grants came, such as Belmont in the state of New York, Amherst

in Nova Scotia, Lanark in Scotland; others after names of early

settlers or prominent local citizens, such as Alban, Bancroft, Car-

son, Ellis, Fancher, Heffron, Keene, Nelsonville, Stevens Point,

Stockton, Whiting; some in honor of great men of the time:

Garfield, Custer, Dewey, Grant; others after natural features:

Linwood, Pine Grove, Sunset Lake, Wolf Lake.

The comparison between Maps 1 and 2 shows the changes that

took place in one hundred years. In 1876, most of the clusters of

non-Anglo-Saxon farmers formed foreign colonies. Until the first

1973] Ferret — Cultural Diversity in Central Wisconsin 55

World War there were immigrants from Europe and therefore

one still could speak of foreign colonies. But in 1970, almost all

the farmers were born in this country and are American citizens.

One might expect that after the end of mass immigration the

foreign colonies would become gradually assimilated and by now

would have become a more or less homogeneous whole that might

be called rural America in opposition to the urbanized parts of

the country. This is not true and the map of 1970 shows that if

some colonies have in fact decreased and even completely van¬

ished, others have developed and continue to develop. They can

no longer be designated as foreign colonies, but they are formed

of a number of families of the same ethnic background who stay

together. We shall speak therefore of areas of concentration of

non-Anglo-Saxon ethnic groups.

The changes between 1870 and 1970 are due to several causes.

The Irishmen left their country after the Great Famine and for

them, farming was probably connected with hardship. They did

not teach their children attachment to the land, and so their

sons and daughters left the farm to go to the cities. At present

only a few isolated farmers are of Irish origin. However, traces

of the Irish occupation remain, such as churches; for instance

Saint Patrick in the town of Lanark, Saint Mary in the town of

Stockton, or cemeteries in which many monuments indicate Irish

names and often carry the name, parish, county of birth in Ire¬

land. The Germans in the town of Sharon were not numerous

and their group dissolved. Their church, Saint Martin, built in

1866, was also used by Catholics of other ethnic groups, but in

1971, it ceased to be used regularly. The German colony near

Almond has decreased, probably because the children did not

stay on farms but went to the cities. Around Kellner the number

of German farms increased and there is another area of German

farms in the northwestern part of the county. Both are near the

Wood County line and are in part an expansion of the large German

colonies in that county.

The Scandinavian colonies increased first through immigration,

but as English is a language close to the Scandinavian languages,

many of the children did not remain on farms and moved to the

cities. After a period of development, the area of concentration

of Scandinavian people has decreased.

The French group was small and soon vanished, not leaving

any trace, except the names of a few farmers scattered in the

county.

The Polish people seem to have found in Portage County the

region that was most suitable to them. The climate is similar to

Poland. The land is almost similar to some regions in Poland, and

56 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

although there are sections where conditions are not very good for

agriculture, this county has become what is probably the largest

rural Polish colony in America. The Poles have large families;

ten children are not exceptional. As the English language is very

different from Polish, foreign born Poles have much difficulty

to learn it and continue to speak Polish at home. Therefore chil¬

dren of Polish descent, born here, have often a strong accent and

for that reason do not mix easily with other people. Many of the

children become farmers; others may go to the cities and work

in factories ; a few get a higher education, but most remain in the

region. In Stevens Point, the Polish element is important not

only in industry, but also in businesses, in administration and in

the professions. Until the first World War, immigration of indi¬

viduals or families coming from the old country was important.

After the war Polish immigrants still continued to come, not

directly from Europe, but from American cities, especially from

Chicago. From Poland, they had gone to Chicago where they

started to work in factories, but they left the big citv for

Stevens Point or for farms in Portage County when they had an

opportunity to find an occupation similar to what they were doing

in their home-country.

For the Amish also, farming is the way of life that suits them,

in fact it is the only way of life allowed in their religion. As

they have large families and as each boy will eventually have a

farm of his own, their colonies expand and when there is no more

land available in the vicinity, they migrate to some other region.

Therefore their colony in Portage County may increase in the

future.

In Portage County, the population of the villages and towns

also reflects the ethnic groups that are represented in the sur¬

rounding area and as a consequence there are typical church de¬

nominations. This is particularly noticeable in small towns with

populations around five hundred inhabitants. Rosholt started as

a Scandinavian center and has only one church, which is Lutheran.

Amherst which had Yankees as first inhabitants has one Meth¬

odist and one Episcopal church representing the first group of

settlers; it has besides one Lutheran church (Missouri synod)

for the German group, another Lutheran church for the Scandi¬

navian group and one Catholic church for the Polish element.

Almond which was started by Yankees has one Methodist and

one Baptist for the Anglo-Saxon group, one Lutheran church

for the German element and one Catholic church for the people

of Polish or Irish origin ; it has besides one Seventh Day Adventist

church. Stevens Point, the county seat, has four Catholic churches ;

originally two were for the Poles, one for the Germans, one for

1973] Perret — Cultural Diversity in Central Wisconsin

57

the Irish; it has two Lutheran churches, one having- at the origin

German parishioners and the other Scandinavian ones. There are

besides several other churches (Episcopal, Presbyterian, Meth¬

odist. Baptist, etc.) and one Jewish synagogue, either for the

Yankees, or for immigrants who came directly to the city, or for

people who left the church of their ancestors to adopt a new faith.

At first some of the churches were very exclusive. St. Mary’s

at Custer in the town of Stockton, which was the church of the

Irish community would accept Polish people only as associate

members with limited privileges; until 1930, persons of Polish

origin could not be buried in the churchyard. At present, how¬

ever, there is no longer any discrimination, but in many churches,

especially the rural ones, one ethnic group is still dominant.

How long will Central Wisconsin still show a cultural diversity?

It seems that in the near future the number of Polish farms will

still increase and occupy more space. So will also the Amish

farms. Both ethnic groups have special interest in farming. Scan¬

dinavian and German farms will probably decrease in number,

as in these ethnic groups families are smaller and many children

get a higher education and quit the farm. Perhaps with time the

differences between ethnic groups may become weaker and even¬

tually there may be one unified American group, but this time

seems very remote and may never come.

REFERENCES

1. ROSHOLT, MALCOLM. 1959. Our County, our Story: Portage County,

Wisconsin. Portage County Board of Supervisors, Stevens Point, Wis¬

consin.

2. NASH, G. N. and F. R. MORGAN (compilers and publishers). 1876. Map

of Portage County, Wisconsin. Milwaukee.

3. Atlas and Plat Book, Portage County, Wisconsin. 1972. Rockford Map

Publishers, Inc., Rockford, Illinois.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN

No. 63. THE GENUS TRIFOLIUM— THE CLOVERS1

John M. Gillett

Plant Research Institute, Canada Department of Agriculture , Ottawa

and

Theodore S. Cochrane

Herbarium , University of Wisconsin , Madison

The clovers were classified by Bentham and Hooker (1865) in

the tribe TRIFOLIEAE, the third of eleven tribes recognized in

their order Papilionatae. From the other genera included : Ononis

L., Parochetus Buch.-Ham., Trigonella L,, Medicago L. and Meli-

lotus Adans. — Trifolium L. was distinguished by its capitate inflo¬

rescences, marcescent petals with fused claws and membranous

legumes. The chief features of the Trifolieae include the herba¬

ceous, rarely shrubby habit ; the pinnate, rarely palmate 3-foliolate

leaves ; the leaflet veins often extending into the teeth ; the stipules

adnate to the petioles; the axillary, rarely terminal peduncles;

the flowers 1-many in bracteate racemes ; the diadelphous or mona-

delphous 10 stamens with apically dilated filaments ; and the in-

dehiscent legumes.

The tribe is apparently a natural one, for it has remained rela¬

tively stable in content and circumscription. Taubert (1891) main¬

tained the same classification. Hutchinson (1964) described it in

much the same way as his predecessors but emphasized also the

uniformity of the anthers and the estrophiolate seeds. By ex¬

cluding the genus Ononis and establishing a new tribe for it, he

removed the monadelphous member. Hutchinson also added Fac-

torovskya , a monotypic genus described by Eig (1927).

The distribution of the species, habitat information and dates

of flowering and fruiting were compiled from specimens kindly

made available by the directors and curators of the following

herbaria: Iowa State Bniversity-Ames (ISC), University of Iowa-

lowa City (IA), Milwaukee Public Museum (MIL), University

of Minnesota (MIN), University of Wisconsin-Madison (WIS),

University of Wiseonsin-Milwaukee (UWM), University of Wis-

eonsin-Oshkosh (WSO), University of Wisconsin-Stevens Point

(WSP), University of Wisconsin-Superior (WSS) , University of

1 Contribution No. 836 from the Plant Research Institute, Canada Department of

Agriculture, Ottawa, Canada, in part.

59

60 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Wisconsin Center-Rock County (ULJ). Each dot or circle on the

maps represents an exact location where a specimen was collected,

while triangles designate county records without specific locations.

The numbers within the map insets record the amount of flower¬

ing and fruiting material available and indicate the months when

a species may be expected to flower or fruit in Wisconsin. Vege¬

tative specimens and those in bud or immature fruit are not in¬

cluded. The year of earliest collection within each county is re¬

corded for the non-cultivated species. Except for the flowers of

Trifolium arvense, T. aureum, T. campestre and T. dubium the

illustrations are redrawn and rearranged from Fassett 1939, pp.

35, 41.

1. TRIFOLIUM L. Clover

Annual, biennial, or perennial herbs with palmately, rarely

pinnately, 3-foliolate (in Wisconsin) or 5-7-foliolate leaves; leaf¬

lets denticulate or serrulate, the conspicuous stipules adnate to

the petiole. Flowers sessile or pedicellate, naked or bracteate, in

heads or short head-like spikes, racemes, or umbels, rarely in two’s,

three’s, or solitary. Involucre absent, small, or a large dentate or

lobed ring. Calyx campanulate to tubular; lobes subequal or the

abaxial longer, the adaxial pair more or less connate. Petals mar-

cescent, nearly free or united into a tube, the claws usually fused

to the staminal sheath, to each other, or to both. Standard oblong

or obovate; wings narrow, longer than the keel. Stamens diadel-

phous, occasionally nearly monadelphous by partial union of the

free stamen. Filaments all or only 5 of them dilated at the apex;

anthers uniform. Ovary with 8 or fewer ovules, if 8 then several

aborting. Style filiform, curved upward, the stigma capitate or

hooked. Fruit a short legume enclosed by the mareescent calyx

and corolla, thin-walled or membranous, indehiscent or dehiscent

by the ventral or dorsal suture or both. Seeds 1-3, occasionally 4,

estrophiolate (without a caruncle).

Trifolium contains 250-300 species, the majority of which are

native to Asia Minor and the eastern Mediterranean region. Euro¬

pean species extend from Scandinavia to Spain; some extend into

North Africa and eastward to Siberia, possibly to western China.

A small group is endemic to tropical Africa from Ethiopia to South

Africa. The North American species, numbering perhaps 60, occur

chiefly from British Columbia to Mexico along the western moun¬

tains and plains, with but few in the southeastern United States.

About 20 species are found in South America from southern Peru

and southern Brazil to central Chile and Argentina. Twenty species

(Hermann 1953) occur either in cultivation or as weeds in North

1973]

Gillett and Cochrane — Reports on The Clovers

61

62 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

America. Eight of the nine Wisconsin species, all European intro¬

ductions, were treated earlier by Fassett (1939). The native Tri¬

folium reflexum L., found in neighboring Illinois, (Jones 1963),

has not yet been reported from Wisconsin.

Key to Species

A. Standard obovate, straight or curved downward; heads 5-15

mm thick, the flowers yellow; calyx 5-veined; fruit stipitate

in the calyx; petioles usually shorter than the leaflets. (Sect.

CHRONOSEMIUM)

B. Annual; terminal leaflet stalked and petiolulate; stipules

ovate, those of the upper leaves about one-half the

length of the petiole ; legume 3-6 times the length of the

style ; seeds ellipsoid, the testa lustrous. _

_ 1. T. CAMPESTRE (T. procumhens)

BB. Biennial; terminal leaflet subsessile, merely petiolulate;

stipules oblong-lanceolate, equaling or exceeding the

petiole; legume 2 times the length of the style; seeds

ovoid, the testa dull. _ 2. T. AUREUM (T. agrarium).2

AA. Standard oblong, curved upward; heads 15-35 mm thick, the

flowers pink, red, purple, or white (never yellow) ; calyx 5-,

10-, or many-veined; fruit sessile or essentially so; petioles

usually longer than the leaflets.

C. Calyx tube gibbous, greatly inflated and strongly vescicu-

lar in fruit, the upper side much more pubescent than

the lower; corolla resupinate; rare cultivated species.

(Sect. GALEARIA) _ 3. T. RESUPINATUM.

CC. Calyx tube not gibbous, not inflated nor vescicular in

fruit, glabrous or uniformly pubescent; corolla not re¬

supinate ; common cultivated and naturalized species.

(Sect. TRIFOLIUM).

D. Stems pubescent; flowers ebracteate, sessile; calyx tube

pubescent; petal claws united into a tube.

E. Perennial or biennial; well developed leaflets 10 mm

or more wide, 1-2 times as long as broad; heads

globose to broadly ovoid, sessile or subsessile on pe¬

duncles rarely more than 4 mm long, subtended by

a stipular involucre; flowers 12-18 mm long, the

calyx ferruginous-pilose. _ 4. T. PRATENSE.

EE. Annual; leaflets 5 mm or less wide, 3-6 times as

long as broad ; heads ovoid to cylindric, on peduncles

4 mm or more long, not subtended by an involucre;

flowers 4-7 mm long, the calyx densely pink- or gray-

villous. _ 5. T. ARVENSE.

1973] Gillett and Cochrane — Reports on The Clovers 63

DD. Stems glabrous or glabrate; flowers minutely bracteate,

pedicellate; calyx tube glabrous; petal claws free. (Sect.

AMORIA)

G. Stems creeping, rooting at some nodes, 1-1.25 mm

thick {fide Fassett 1939) ; heads borne on scapes

6-30 cm long, the flowers white ; calyx with 10 incon¬

spicuous veins, the lobes shorter than the tube, the

sinuses glabrous; stipules scarious, thin and

wrinkled, pale brown or tinged with green or purple.

_ 6. T. REPENS .

GG. Stems erect or ascending, 2-5 mm thick; heads

borne on axillary peduncles 2-10 cm long, the flow¬

ers becoming pink with age ; calyx with 5 prominent

veins, the lobes equaling or exceeding the tube, the

sinuses pubescent; stipules partly herbaceous, firm

and green. _ 7. T. HYBRIDUM.

1. Trifolium campestre2 Schreber in Sturm Deutschl. FI. Heft.

XVI : 13. 1804.

Low Hop Clover, Large Hop Clover Map 1, Fig. 1.

Trifolium procumbens of Am. authors, not L. Sp. PI. 2: 772.

1753, nom. ambig.

Annual 5-40 cm tall, simple or with ascending slender branches,

the stems thinly appressed-pilose. Petioles of lower leaves 1-1.5

times as long as the leaflets, those of the upper about equal to or

shorter than the leaflets. STIPULES semi-ovate, acute, dentate

or entire, mostly 5-8 mm long, about 1/2 the length of the petioles.

Leaves pinnately 3-foliolate; LEAFLETS obovate, 4-15 mm long,

fine-strigose along the midveins below, denticulate above the mid¬

dle, cuneate; terminal leaflet borne on a rachis segment 1—3 mm

long, petiolulate like the sessile to subsessile laterals. HEADS

axillary, 7-15 mm long, globose to broadly ovoid or becoming hem¬

ispherical by reflexion of the flowers, the peduncles about as long

as or shorter than the leaves. Flowers (10-) 20-40 (-53). CALYX

strongly bilabiate, 1.5 mm long, the adaxial lobes short, deltoid,

0.2 mm long, the laterals and abaxial linear-subulate, each tipped

2 Linnaeus (1753) described 40 species of Trifolium, a number of which have been

transferred subsequently to other genera. Because of obvious practical difficulties

none of these Linnaean species has been properly typified. According to Dandy

( 1958) two of those included here, namely T. agrarium L. and T. jjrocumbens L.,

bear names which have been misapplied consistently and are considered by him to be

uomina ambigua. The names T. aureum Pollich and T. campestre Schreber therefore

were taken up by Dandy and subsequently employed by Coombe (1968) in his treat¬

ment for the Flora Europaea. For the purpose of this treatment we are following

their usage, reserving judgment on the application of these names until their typifi-

cation problems can be satisfactorily resolved.

64 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

by 1 or 2 stiff hairs. COROLLA yellow , 3.5-6 mm long, the stand'

ard obovate when flattened, hooded and enveloping the other petals,

becoming sulcate-striate ; wings divergent, somewhat spoon-shaped

and twisted, 4 mm long. Ovary stipitate, the stipe 1 mm long;

style 1/3 to 1/6 the length of the legume. LEGUME oblongoid,

2-2.5 mm long, usually 1-seeded ; SEED ellipsoid, yellow, lustrous,

1.2-1. 5 mm long, 0.6 mm wide, n — 7 (Larsen 1956) ; 2n = 14

(Karpechenko 1925; Bleier 1925; Kliphuis 1962; Gadella and

Kliphuis 1963).

Native throughout Europe except for the extreme North and

East, North Africa and western Asia, widely naturalized in North

America; infrequent throughout Wisconsin in dry disturbed hab¬

itats, from pastures, fields, open woods and roadsides with such

weeds as Agropyron repens, Medicago lupulina, Oxalis stricta

and Daucus carota to railroad ballast, landfill and city lots. Flow¬

ering 23 June to 15 August; fruiting 6 July to 4 October.

Trifolium campestre is self-compatible and may set seed inde¬

pendent of pollinators. In Europe pollinators include Hymenoptera,

Diptera and Lepidoptera (Knuth 1908).

Trifolium dubium Sibth. (Fig. 4), Little Hop Clover, can be dis¬

tinguished readily from the similar T. campestre by its smaller

heads (5-8 mm in diam), which have fewer flowers (5-20), its

shorter corollas (2. 5-3. 5 mm long), whose standards are scarcely

or not at all sulcate-striate, and its calyx lobes, which are about

equal in length to the tube. Introduced from Europe and natural¬

ized near the coasts, T. dubium occurs only sporadically inland.

There is ample reason to doubt that one collection from Wisconsin

(Sheboygan: Goessl s.n., 1 Sept. 1914, MIL, WIS) was procured

from the locality given or that the plant was a legitimate ad-

ventive. The only other specimen confirming the presence of the

species in the state is also a Goessl collection: Wood Co.: Marsh¬

field, a few plants, roadsides {2987, 21 Sept. 1915, MIL) ,

Trifolium dubium and especially T. campestre are easily con¬

fused with the ubiquitous black medick, Medicago lupulina. The

following key will help distinguish them from the latter species.

A. Stems rounded, the stipules fused to the petioles for one-half

or more of their length; inflorescenses globose to very short- ,

cylindric heads, 8-15 mm long, 3-40-flowered, the peduncles

thinly appressed-pilose ; flowers reflexed after anthesis ; calyx

teeth very unequal; corollas 2.5-6 mm long, the standard

serrulate about the middle, persistent in fruit; legumes ob¬

longoid, brown _

_ TRIFOLIUM CAMPESTRE and T. DUBIUM.

1973] Gillett and Cochrane — Reports on The Clovers

65

AA. Stems angled, the stipules fused to the petioles for less than

one-half of their length ; inflorescences compact spiciform

racemes, 7-10 mm long, 10-50-flowered, the peduncles densely

pilose, often glandular; flowers not reflexed; calyx teeth

nearly equal; corollas 1.5-3 mm long, the standard entire,

deciduous; legumes subreniform, black at maturity. _

_ MEDICAGO LUPULINA.

The flower structure in many species of section Chronosemium,

which includes all the Hop Clovers, differs markedly from that

in red and white clovers. Instead of turning upward at the apex,

the standard remains straight, acting as a hood which surrounds

the other floral parts. In this it is aided by the presence of strong

parallel veins. The wings flare outward, but because the base of

the blade is adnate to the base of the keel, the whole structure

moves downward under the weight of a pollinator, exposing the

stigma and staminal sheath. Another feature of particular in¬

terest is that the base of the wing blade bears a triangular ap¬

pendage rather than an orbicular one. The acuminate apices of the

two blade appendages form a yoke mechanism which straddles the

staminal sheath, when the wing-and-keel mechanism is depressed.

It acts as a mechanical guide to return the structure to its original

position surrounding the staminal sheath and style after the de¬

parture of a pollinator.

2. Trifolium aureum2 Pollich Hist. PI. Palat. 2 : 344. 1777.

Hop Clover, Yellow Clover Map 2, Fig. 2.

Trifolium agrarium of Am. authors, not L. Sp. PI. 2: 772.

1753, nom. amhig.

Annual or biennial 15-40 cm tall ; stems appressed-pilose, striate,

erect with ascending branches. Petioles of lower leaves about 10

mm long, those of the upper 1-2 mm long. STIPULES oblong-

lanceolate, acuminate, 8-20 mm long, equaling the petioles. Leaves

palmately 3-foliolate; LEAFLETS obovate-elliptic, to 2.5 cm long,

retuse with a minute mucro, denticulate above the middle, entire

and cuneate below, the terminal subsessile and merely petiolulate

like the laterals. HEADS ovoid-cylindric, the top flattened in age,

axillary and terminal, 20-80-flowered, the peduncles equalling or

exceeding the leaves. CALYX 2.5 mm long; tube 1 mm long,

the adaxial lobes deltoid, 0.6-0. 9 mm long, about 3/2 the length

of the linear abaxial and laterals. COROLLA gold-yellow, turning

brown in age, marcescent, 5-6.5 mm long; standard obovate,

the parallel veins strongly sulcate, serrulate about the middle;

wings spoon-shaped, flared outward, 1/3 the length of the stand-

66 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

ard. Ovary stipitate, the stipe 1-1.5 mm long, the style 1/2 or

more the length of the legume. LEGUME oblongoid, 3-3.5 mm

long, 1-seeded, the SEED ovoid, pale yellow-green, dull, 1-1.2

mm long, 0.8 mm wide. 2 n — 14 (Wipf 1939; Wulff 1939; Love &

Love 1956) .

Native to Europe, introduced and naturalized in North America;

an infrequent weed in the northwestern two-thirds of Wisconsin

along railroads and roadsides, such as near Windfall Lake, Forest

Co., near a jack pine planting with Oenothera, Apocynum, Achillea

millefolium and Hieracium aurantiacum (Kruschke K— 65— 4-9, 13

July 1965, MIL), in pastures, fields and disturbed woods. Flower¬

ing 6 June to 4 August; fruiting 22 June to 22 October.

Trifolium aureum often behaves as a perennial because cropping

prevents or delays flowering.

The base number of x = 7 is common in section Chronosemium,

but x — 8 has also been recorded.

3. Trifolium resupinatum L. Sp. PI. 2: 771. 1753.

Persian Clover Map 2, Fig. 3.

Glabrous, procumbent, ascending or erect annual 1-4 dm tall,

with many basal branches. Petioles of basal leaves several times

longer than the leaflets, the upper leaves subsessile. STIPULES

membranous, lanceolate to ovate-lanceolate, narrowly attenuate-

aristate, the tip 0.6-1. 8 mm long. Leaves palmately 3-foliolate;

LEAFLETS 5-20 mm long, obovate to oblanceolate, sharply den¬

ticulate or serrulate, the apex round to acute, cuneate, the ex¬

current veins terminating in numerous minute teeth. HEADS

axillary, hemispherical, small, 5-10 mm in diam in flower, larger

and globose in fruit, 6-20-flowered, the peduncles about equalling

the leaves. CALYX 1.7-2; 7 mm long, about 20-veined, strongly

bilabiate, the tube white to pale green with a dark green basal

band, woolly-pilose adaxially, nearly glabrous abaxially, at ma¬

turity inequilaterally inflated to 8 mm, transparent and regularly

reticulately veined; adaxial lobes setaceous, 0.5 mm long, the

laterals and abaxial subequal, narrowly triangular. COROLLA

pink to violet, resupinate ; tube about equalling the calyx before

inflation; standard oblong when flattened, refuse, 4-6 mm long;

wings and keel 2 mm long. LEGUME compressed-ovoid to orbicu¬

lar, 2 mm in diam, 1-seeded; SEED ovoid, 1.2 mm long {fide Isely

1948). 2 n — 14 (Wipf 1939) ; 2n — 16 (Bleier 1925; Karpechenko

1925 ; Evans 1962) .

Native to Europe and North Africa, locally naturalized in the

southeastern United States where sometimes used for forage and

green mulch; in Wisconsin recorded only from lawns that have

1973] Gillett and Cochrane — Reports on The Clovers

67

been seeded earlier in the year, this handsome species an occasional

component of lawn seed mixtures. The following specimens have

been seen from the state : Dane Co. : Madison, Clifford Ct., Randall

Ave. ( Fassett 10297 , 21 Sept. 1939, WIS) ; Milwaukee Co. : Milwau¬

kee, Ivanhoe Place and Terrace Ave. ( Shinners 1463, July 1939,

WIS) ; Rock Co.: Beloit ( Truman , 1935, fide Fassett 1939, p. 38) ;

Walworth Co. : Delavan, 118 S. 4th St. (Wadmond s. n., 12, 27 July

1935, WIS). Flowering July and August.

4. Trifolium pratense L. Sp. PI. 2: 768. 1753.

Red Clover Map 3, Fig. 5.

Trifolium pratense var. sativum Schreb.

Trifolium pratense var. sativum forma flavicans (Vis.) Hayek

Trifolium pratense forma leucochraceum Aschers. & Prantl

Trifolium pratense forma alhiflorum Puskal

Trifolium pratense forma semipurpureum (Strobl) Aschers. &

Graebn.

Cespitose perennial or rarely biennial 2-6 (-10) dm tall. Stems 1-

several, erect to decumbent, glabrate or more commonly pilose to

villous. Lower leaves long-petiolate, the upper subsessile. STIP¬

ULES narrowly ovate to oblong, 1-3 cm long, membranous, with

strong reticulate venation, adnate to the petiole for most of their

length, the free portion triangular, abruptly contracted to a seta¬

ceous tip 4-8 mm long. Leaves palmately 3-foliolate; LEAFLETS

of basal leaves ovate, obovate or even orbicular, those of the upper

leaves oblong-lanceolate or elliptic, 1-3 (-5) cm long, essentially

entire, the apex rounded to retuse, pilose below, glabrate above.

HEADS 1 or 2, terminal, globose to ovoid, sessile or on peduncles

rarely more than 4 mm long, subtended by an involucre of stipules

of the upper pair of leaves. Flowers 40-150, 12-18 mm long.

CALYX narrowly campanulate, 10-nerved, 10 mm long; tube 4 mm

long, ferruginous-pilose, the throat ringed with hairs within;

lobes subulate, the abaxial 5-6 mm long, the equal adaxial and

laterals 3-4 mm long, the tips with a few divergent pilose hairs.

COROLLA tubular, pink to red-violet or rarely yellowish or white ;

standard oblong-oblanceolate, retuse. LEGUME oblong-ovoid,

1-seeded, the SEED ovoid, 1.8 mm long, 1.1 mm wide, yellow to

brown, n — 7, 14 (Povilaitis & Boyes 1956) ; 2 n — 14, 28 (Karpe-

chenko 1925 and many others; cf. Love & Love 1956).

Introduced from Europe and freely naturalized throughout

North America; a ubiquitous weed and commonly cultivated crop

throughout Wisconsin on fertile but well drained soil, in old fields,

pastures, farmyards, edges of woods, disturbed deep-soiled prairies,

68 Wisconsin Academy of Sciences , Arts and Letters

[Vol. 61

1973] Gillett and Cochrane — Reports on The Clovers

69

along roadsides and railroads, growing with Bromus inermis,

Fragaria virginiana, Melilotus alba , Linaria vulgaris , Taraxacum

officinale and Achillea millefolium. Flowering 24 May to 12 Oc¬

tober; fruiting 27 June to 1 November.

Trifolium pratense , the most widely cultivated forage clover, is

exceedingly polymorphic in both natural and cultivated states,

hence the large number of published binomials that apply to it.

However, considerable study is needed to resolve the status and

nomenclature of any infraspecific taxa which might be recognized.

Such study would be complicated by the large numbers of cultivars

selected and developed through its long period of cultivation.

Because of the long tubular corolla, any nectar, which is secreted

at the base of the staminal sheath, is available only to long-tongued

insects. Thus bumblebees are more successful pollinators than

honeybees. Because the stigma protrudes beyond the anthers, pollen

is obtained from visiting insects immediately upon depression of

the keel-and-wing mechanism. Cross fertilization is necessary for

seed production; very few seeds are produced by plants isolated

from pollinators.

Trifolium incarnatum L. (Fig. 7), Crimson Clover, is a villous

annual or winter annual with broadly obovate to suborbicular leaf¬

lets, blunt stipules, dark blood-red flowers in narrowly ovoid to

spike-like cylindric heads and densely villous calices, which in fruit

have the sinal veins thickened and constricted to form a narrow

orifice and the linear-subulate lobes widely divergent. Native to

South and West Europe, now widely naturalized throughout

Europe and parts of the Mediterranean region (Coombe 1968),

cultivated in the southeastern United States as a forage plant and

for soil improvement. The single Wisconsin collection is probably

an escape from cultivation: Dane Co.: Madison ( T release s.n.,

8 Sept. 1883, WIS).

5. Trifolium arvense L. Sp. PL 2 : 769. 1753.

Rabbitfoot Clover, Old Field Clover Map 4, Fig. 6.

Annual 5-30 (-40) cm tall ; stems rather delicate, simple or more

often branched, with appressed or spreading short-villous pubes¬

cence, the hairs white when fresh, turning brown on herbarium

material. Petioles of lower leaves to 1.5 cm long, shorter than the

leaflets , the upper leaves subsessile. STIPULES ovate-oblong with

long setaceous tips. Leaves palmately 3-foliolate; LEAFLETS

5-20 mm long, linear-lanceolate or usually narrowly oblong to

oblanceolate, cuneate, minutely serrulate near the apex, blunt with

70 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

a mucronate tip. HEADS axillary and terminal, subglobose to more

often short-cylindric, 5-30 mm long, 1 cm thick, subsessile or on

peduncles 5-30 mm long. Flowers 10-180. CALYX pink- or gray-

villous, 4-7 mm long ; tube weakly 10-veined, the throat unthick¬

ened and glabrous; lobes setaceous, essentially equal, 2.8-5 mm

long, plumose. COROLLA white to pink, much shorter than the

calyx, 4 mm long, the standard oblong. LEGUME broadly ovoid,

1.3 mm long, indehiscent, the SEED pale yellow, 0.9 mm long.

2 n = 14 (Karpechenko 1925 ; Bocher & Larsen 1958 ; Kliphuis 1962 ;

Gadella and Kliphuis 1963 and many others) ; 2n — 16 (Evans

1962).

Native to Europe, Asia Minor, North Africa and the Canary

Islands; a scattered advent! ve in Wisconsin on dry sandy or grav¬

elly soils, mostly forming colonies along road shoulders with such

species as Matricaria discoidea, Mollugo verticillata, Ambrosia

artemisiif olia and Polygonum arenastrum , occasionally in quarries,

fields, lake shores and open woods, such as in jack pine barrens

near Spring Brook, Washburn Co., with Potentilla, Ceanothus

americanus, Lithospermum caroliniense and Centaur ea maculosa

( Heidel 314, 14 July 1966, WIS). Flowering 30 June to 19 Septem¬

ber; fruiting 14 July to 10 October. Because it is self-pollinated,

seed production takes place throughout the flowering period.

Trifolium arvense is extremely polymorphic, with many de¬

scribed varieties and forms.

6. Trifolium repens L. Sp. PL 2: 767. 1753.

White Clover Map 5, Fig. 8.

Glabrous or glabrate perennial with repent stems rooting at

some nodes, 10-50 cm long. Petioles 5-15 (-20) cm long depending

on local conditions. STIPULES thin-membranous to scarious, pale

brown with dark veins, 8-15 mm long, ovate to ovate-lanceolate,

truncate or obtuse, abruptly tapered to a short subulate tip. Leaves

palmately 3-foliolate ; LEAFLETS obcordate to broadly elliptic,

1-3 cm long, broadly cuneate, the margin sharply denticulate to

serrulate above, the apex rounded to shallowly retuse, often with

characteristic white markings toward the base and light, prominent

veins. HEADS globose, hemispherical in age, 15-25 (-30) mm in

diam; peduncles axillary, 6-25 (-30) cm long , arising from the

repent stems and usually overtopping the leaves. Flowers (17-)

25-80 (-105) white or pink-tinged fading to brown, sweet-scented ;

pedicels slender, curved, 1-1.5 mm long, elongating in fruit to 5

mm, each subtended by a scarious lanceolate bract, the terminal

flowers on the raceme axis often aborted. CALYX tube campanu-

1978] Gillett and Cochrane — Reports on The Clovers

71

late, 2.5 mm long, white with dark green lobes, 10-veined, some¬

times with additional nerves, these often reticulately connected;

lobes narrowly triangular, 2-3 mm long, the adaxials somewhat

united at the base and longer than the others, about equaling the

tube, separated by a narrow, acute sinus, the remaining sinuses

broad. COROLLA 8-12 mm long, the standard oblong when flat¬

tened; wings 7 mm long; keel 6 mm long. LEGUME linear-

oblongoid, 4-5 mm long, 3-4-seeded, constricted between the seeds.

SEEDS yellow to orange, dull, 1 mm long, n — 16 (Tiemann and

Schreiter 1961) ; 2n = 32 (Atwood and Hill 1940 and many

others) ; 2 n = 32, 48 (Moriya and Kondo 1950) ; 2n = 64 (Levan

1942).

Native to Europe, planted in lawns and widely naturalized

throughout cool temperate North America, on dry to moist but

well-drained soil along roadsides and paths through woods, in fields,

pastures and disturbed areas in general; in Wisconsin a common

weed growing with Melilotus spp., Trifolium hybridum, Pastinaca

sativa, Prunella vulgaris, Plantago rugelii, Achillea millefolium

and Cirsium arvense. As one of the most ubiquitous weeds known

to man, white clover quite probably has a broader distribution than

any other species of Leguminosae. Flowering 21 May to 4 (25) Oc¬

tober; fruiting from 5 June to 14 October.

Of the many described segregate taxa, Coombe (1968) recognized

six subspecies in his treatment for the Flora Europaea. Of the

many cultivars selected from this almost entirely cross-pollinated

species, perhaps the best known is cv. ‘Ladino,’ sometimes referred

to as “giant” because of its large size. Both wild and cultivated

types exhibit great variability not only in their morphology but

also in their productivity, persistence and ability to establish them¬

selves in various habitats.

7. Trifolium hybridum L. Sp. PL 2 : 766. 1753.

Alsike Clover Map 6, Fig. 9.

Trifolium hybridum var. pratense Rabenh.

Trifolium hybridum var. elegans (Savi) Boiss.

Subglabrous perennial 15-60 (-80) cm tall; stems someivhat

stout and succulent, erect or decumbent, rarely procumbent,

branched above. Petioles of lower leaves to 10 cm long, progres¬

sively shorter upward but never sessile. STIPULES partly

herbaceous, light green, ovate to oblong-lanceolate, 10-30 mm long,

attenuate to long slender tips. Leaves palmately 3-foliolate ; LEAF¬

LETS to 25 (-35 or more) mm long, ovate to obovate, rhombic, or

elliptic, cuneate, rounded to retuse at the apex, sharply serrulate

72 Wisconsin Academy of Sciences, Arts and Letters

[Vol. 61

9. TRIFOLIUM HYBRIDUM

r0i & r

TRI FOLIUM REPENS

_ A£

1973] Gillett and Cochrane — Reports on The Clovers 73

to denticulate above. HEADS axillary, globose, hemispherical in

age, 2-3.5 cm in diameter, on peduncles 2-10 cm long. Flowers

20-60 (-80) , each subtended by a setaceous bract, borne on slender

pedicels 1-2 mm long, these becoming deflexed and elongating in

fruit to 7 mm. CALYX tube campanulate, 10-veined, the sinuses

obtuse, pubescent, their nerves usually obscure, those of the lobes

prominent; lobes linear-subulate, the adaxials 3 mm long, longer

than the subequal abaxial and laterals and shorter than to usually

almost tivice the length of the tube. COROLLA white, turning pink

and finally brown, 7-11 mm long, the tube 4 mm long; standard

obovate when flattened, 6-8 mm long ; wings 7 mm long ; keel 6 mm

long. LEGUME linear-oblongoid, 3-4 mm long, 2-4-seeded, the

SEEDS dull black when mature, 1.2 mm long, n — 8 (Kawakami

1930) ; 2n — 16 (Turesson 1962; Love and Love 1956; Gadella and

Kliphuis 1966).

Native to Europe from Portugal east to the Caucasus and

Turkey; widespread throughout Wisconsin, undoubtedly in every

county, most frequently in moist to dry fallow or cultivated fields

and pastures, along roadsides and margins of woods where it is

associated with the same species as Trifolium repens. Flowering

26 May to 30 September; fruiting 20 June to 14 October.

Superficially resembling Trifolium repens when prostrate as

the result of grazing, but easily distinguished by its pink flowers

and calyx morphology. In Clover phyllody, suspected to be caused

by Aster Yellow virus in both species, the flower parts are con¬

verted into leaf-like structures, making identification difficult. Posi¬

tion of parts and presence of normal floral parts among the

abnormal heads will aid in identification. Some of the bizarre forms

described for these two species probably represent infections by

this virus.

Bibliography

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cytes of Trifolium repens. Amer. J. Bot. 27 : 730-735.

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Linnaeus, C. 1753. Species Plantarum, ed. 1. Vol. 2. Stockholm.

Love, A. and D. Love. 1956. Cytotaxonomical conspectus of the Icelandic flora.

Acta Horti Gotoburgensis 20: 65-291.

Moriya, A. and A. Kondo. 1950. Cytological studies of forage plants. II.

Legumes. Jap. J. Genet. 25: 131-134.

Povilaitis, B. and J. W. Boyes. 1956. A cytological study of autotetraploid

red clover. Amer. J. Bot. 43: 169-174.

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.

Taubert, P. 1891. Leguminosae, in Engler, A. and K. Prantl, Die Natiirlichen

Pflanzenfam. Vol. II, 2. Wilhelm Engelmann Verlag, Leipzig.

Tiemann, H. and J. Schreiter. 1961. Chromosomen Studien in der Gattung

Trifolium und phylogenetische Betrachtungen zum Weissklee (Trifolium

repens L.). Ziichter 31: 270-273.

Turesson, G. 1962. Results of colchicine doubling in the red, alsike and white

clover. Agric. Hort. Genet. 20: 111-135.

Wipf, L. 1939. Chromosome numbers in root nodules and root-tips of certain

Leguminosae. Bot. Gaz. 101: 51-67.

Wulff, H. D. 1939. Chromosomenstudien an der schleswigholsteinschen Angio-

spermen-Flora. IV. Ber. Deutsch. Bot. Ges. 57 : 424-431.

THE MAMMALS OF DANE COUNTY

A. W. Schorger

University of Wisconsin

Deceased May 26, 1972

Dane county comprises 789,100 acres. The eastern portion of

the county is rolling and level, while the western is very hilly and

has some steep slopes. The extreme western portion is especially

so, since it contains some Driftless Area (unglaciated). Most of

the county originally consisted of prairies and oak openings, with

little forest land. Peat lands covered 52,288 acres.

At the present time, the eastern half of the county is so intensely

cultivated that it contains a lower population of small mammals

than the western. Most of the trapping on which this report is

based was done in the western half of the county. There is a

large element of luck in trapping, since in spite of the large num¬

ber of trap nights only one rare mammal, an Arctic Shrew, was

caught. This study reports approximately 16,500 trap nights be¬

ginning in 1940.

The nomenclature follows Jackson’s Mammals of Wisconsin

(Jackson, 1961).

SPECIES

Opossum (Didelphis marsupialis virginiana) — The opossum was

a comparatively rare mammal in Wisconsin prior to the early

1930’s. It is now common in the county.

Cinerous Shrew (Sorex cinereous cinereous) — The cinerous

shrew is one of the more common shrews and is found in a wide

variety of habitats. Sometimes it approaches being abundant. On

April 30, 1944, I caught four on the south side of Lake Wingra in

thirty-seven traps.

Saddleback Shrew (Sorex arcticus laricorum) — Locally this

mammal is known as the Arctic Shrew. It is about 120mm in

length including the tail of 42mm. In winter a band of black

extending the entire length of the back is bordered with brown.

This shrew is found in moist or wet places, and is not common.

Specimens have been taken in the marshes adjoining Lake Wau-

besa and in the Cherokee marsh.

75

76 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Pigmy Shrew (Microsorex hoyi hoyi) — This species is rare and

is known from a single specimen taken by Clough (1965) in the

Cherokee marsh.

Big Short-tailed Shrew (Blarina brevicauda kirtlandi) — The big

short-tailed shrew is common and is widely distributed. In the fall

of 1969 Larry Flaten and a neighbor trapped thirty-five on their

lots in east Madison. I have taken specimens with white spots.

Little Short-tailed Shrew (Cryptotis yarn a harlani) — This rare

shrew is known only from a skull found in a great horned owl pel¬

let near Klevenville, January 31, 1932, which is now preserved in

the U.S. National Museum (Nelson, 1934).

Prairie Mole (Scalopus aquaticus machrinus) — The prairie mole

is common, especially in the western half of the county. A farmer

at Riley informed me that his cat often brought specimens to the

house but would never eat them. It appears frequently above

ground, and I have found roadkills even in winter. So far, I have

not encountered it east of the Yahara River. In my collection is

an entirely white specimen taken by Jeff Glasbrenner in the town

of Marietta, Crawford County, on June 1, 1969.

Little Brown Bat (Myotis lucifugus lucifugus) — This is prob¬

ably our most common bat. As evidence of its abundance, 124 were

killed at Lake Barney July 2, 1969, by treating the state barn with

ammonia gas. It is sometimes seen in broad daylight.

Eastern Long-eared Bat (Myotis keenii septentrionalis) — The

long-eared bat is rare in the county and is represented by a single

specimen in the Museum of the Zoology Department at the Uni¬

versity of Wisconsin — Madison.

Silver-haired Bat (Lasionycteris noctivagans) — This tree bat is

fairly common. I took a specimen from my yard September 6,

1965, and have examined three others submitted to the State Labo¬

ratory of Hygiene for possible rabies. There is a specimen in the

Museum taken in the Arboretum September 24, 1939.

Big Brown Bat (Eptesicus fuscus fuscus) — The big brown bat is

common and winters frequently in buildings. On October 17, 1963,

I caught one in the attic of my home in a snap trap baited with pea¬

nut butter. The nose of the bat was on the bait.

Red Bat (Lasiurus borealis borealis) — This bat is the most com¬

mon of tree bats. It frequently roosts in bushes as low as five or

six feet.

Hoary Bat (Lasiurus cinereus cinereus) — The hoary bat is un¬

common, and although it occurs in summer there is no breeding

record. The following specimens have been taken : two were

caught in the Arboretum on September 7, 1944, and another was

found on July 28, 1956, on a barb wire fence, also in the Arbore-

1973] Schorger — The Mammals of Dane County 77

turn. On June 22, 1969, I received a crippled specimen found in

the yard of William F. Fey, 5509 Barton Road, Madison.

White-tailed Jack Rabbit (Lepus toivnsendii campanius) — The

jack rabbit is not native, and its presence in Dane County is due

to spreading from plantings. A few years ago it was not uncom¬

mon near Belleville and Pine Bluff, but I have no recent records.

My latest record is a roadkill found in Section 25, Town of Madi¬

son, on October 28, 1951.

Cottontail (Sylvilagus floridanus mearnsii) — The cottontail is

common throughout the county where it is the most popular game

animal. Its population fluctuates. It occurs regularly on University

Heights and various other parts of the city of Madison. On August

12, 1946, I saw a roadkill within two blocks of the capitol. Between

November 10, 1942, and February 28, 1943, 731 rabbits were shot

and trapped in the Arboretum. It was estimated that this was two-

thirds the cottontail population. Shooting was continued each win¬

ter only from 1942-43 until 1964-65, with a high of 415 killed in

1954-55 and a low of 36 killed in 1959-60.

On March 4, 1954, George Knudsen showed me two rabbits in

willow trees east of the bridge at University Bay. One was resting

about 4 feet from the ground on a mat of twigs growing from the

trunk. The other was 6 feet from the ground on the lowest limb

of a large perpendicular willow. It was impossible to reach the limb

except by climbing the trunk. On the fifth day both rabbits were

in the same places but only the one on the twigs remained on the

sixth day.

Woodchuck (Marmota monax monax) — Woodchucks were for¬

merly numerous, but it is now unusual to see one in the county.

For many years farmers have decimated their ranks, and many so-

called sportsmen have made it a business to shoot them. Their food

is almost entirely vegetable though they will not disdain meat, at

least after coming out of hibernation. One spring in northern Ohio

I set out to climb to the nest of a red-tailed hawk that I had found

previously. The Irish setter with me killed a cottontail and, as it

was out of season, I put the rabbit into a hollow log. On returning

the dog showed great interest in the log. With a long pole I ejected

a greatly distended woodchuck. It had consumed nearly the whole

of the cottontail.

Striped Ground Squirrel (Spermophilus tridecemlineatus tri-

decemlineatus) — The 13-lined ground squirrel is an abundant ro¬

dent. Its favorite habitat is land covered with short grasses. Rong-

stad (1965) found that adults had a 29 percent annual survival

rate in the Arboretum, and juveniles 16 percent. On September

25, 1965, I saw one busily engaged in collecting the small acorns

78 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

of the bur oak. Insects are important in the diet, and meat is eaten

readily. It will frequently be seen feeding on roadkills including

its brethren. Van Meter (1881), editor of the New Richmond

Republican, wrote that he saw a ground squirrel eating a mouse

that it dropped on being approached. The victim was fresh and

bleeding. The head and shoulders had been devoured.

Franklin’s Ground Squirrel (Spermophilus franklinii) — This

ground squirrel is of local distribution. Occasionally it is found

on high ground, especially near hazel thickets, but at present is

confined largely to the edge of marshes. There was a small colony

on the southern edge of the former University Bay marsh. For¬

merly there was a large colony near the present site of the sewage

disposal plant, Section 30, town of Blooming Grove.

On August 14, 1970, I visited the home of Mrs. Joan Ohlin,

Section 8, town of Burke, on the edge of Cherokee Marsh. She

had phoned me that she had shot one of many spotted gray animals

living in her yard. They were identified as S. franklinii. Mrs.

Ohlin said that they appeared most frequently in the evening and

that on one occasion she counted 17 of them. There were numerous

burrows in the yard.

Gray Chipmunk (Tamias striatus griseus) — The chipmunk is

common and periodically abundant. It is found regularly on Uni¬

versity Heights and other parts of the city.

Gray Squirrel (Sciurus carolinensis hypophaeus ) — The color

of the gray squirrel in county districts is usually normal, but in

Madison and its suburbs it varies greatly. Some are white on the

belly, others brown. With a few the entire pelage is yellow or

isabelline. Some have a body of normal color and the tail yellow,

while on the grounds of the Mendota State Hospital the tail may

be white. The black phase is unusual. I have seen only four ex¬

amples in the county, three of which were in Madison.

Fox Squirrel (Sciurus niger rufiv enter)— In 1912, the fox squir¬

rel was the only squirrel to be found on University Heights, and

since that time it has been almost entirely replaced by the gray

squirrel. Fox squirrels with a black belly are rather rare. Thure

Kumlien collected one at Busseyville on August 26, 1880, which

he thought would be of interest to the University of Wisconsin.

In the collection of the Department of Wildlife Ecology is a speci¬

men shot near Mt. Horeb on September 7, 1963. I have a female

collected on the campus on September 27, 1962. Though not preg¬

nant, it weighed 866 grams. This is the heaviest squirrel that I

have handled.

The afternoon of August 16, 1963, I met Harry E. Stanz, Jr.,

formerly with the Wisconsin Conservation Department, at the

corner of Princeton and Kendall Avenues, within a block of my

1973] Schorger — The Mammals of Dane County 79

home. He told me that in the forenoon while walking past 1817

Kendall a large fox squirrel killed one of two young gray squirrels

at the base of a tree. When the young one screamed, an old gray

squirrel came running across the street but on arrival did not

interfere with the attack. We went to the tree and found the dead

young. It was a male gray squirrel (Sciurus carolinensis) 307mm

in length and weighing 178.5 grams. The dead squirrel showed

no lacerations except on the nose where a piece had been bitten out.

The fox squirrel was recognizable by having an incomplete tail.

In spite of my daily attempts I was unable to find this squirrel

until an entire month had passed. It was then seen within 100

feet of the tree where the gray squirrel was killed.

Red Squirrel (Tamiasciurus huclsonicus minnesota) — This

squirrel is rare, and no specimen for the county is known. The

sole reason for including it is a statement by Knipping (1950)

that he examined for parasites a specimen taken in Dane County

on December 1, 1948.

Flying Squirrel (Glaucomys volans volans) — The flying squirrel

is common and sometimes becomes a nuisance in houses. During

cold weather several may be found in the same den. On March 3,

1935, in the School Woods, town of Cross Plains, I routed 12

from a crack in an old oak. I have also found as few as three in

the hole of a woodpecker.

Beaver (Castor canadensis michiganensis) — The beaver is pres¬

ent in small numbers. In June, 1948, there was an occupied beaver

dam on Dunlop Creek and in December, 1949, there was a dam on

Black Earth Creek near Cross Plains. On March 21, 1953, I saw

three fresh skins at the farm of the Wisconsin Conservation De¬

partment in the Mazomanie bottoms. I explored Blums Creek,

town of Mazomanie, on March 27, 1954. There was a new dam

about one-fourth of a mile from its mouth. At its entrance into

the Wisconsin River there were many trees which had been felled

long before. One cottonwood was two feet in diameter. On March

18, 1964, there were three abandoned dams and a house on the

Sugar River two miles west of Verona. The beavers must have

arrived by way of the Rock and Pecatonica Rivers. On the same

date there was a new dam on the old site at Blums Creek.

Harvest Mouse (Reithrodontomys megalotis pectoralis ) — This

mouse is not common. It occupies many diverse habitats from

thick crabgrass on high ground to the edge of marshes. I have

specimens taken in the Pheasant Branch marsh, the Mazomanie

marsh, and on high ground near Cross Plains.

Prairie White-footed Mouse (Peromyscus maniculatis hairdii) —

The prairie white-footed mouse is limited to grasslands and usually

shuns cultivated fields. As a result, its habitat now is greatly re-

80 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

stricted. On November 5, 1944, I caught 16 in 50 traps set along the

railway near Middleton where there was a dense growth of sweet

clover, on the seeds of which they were feeding. When the prairies

and oak openings were in primitive conditions, it must have been

the most abundant Peromyscus.

Northern White-footed Mouse (Peromyscus leucopus novebora-

censis — The northern white-footed mouse occurs commonly in

wooded and brushy areas. It is not uncommon in the city, where

I have taken several specimens at University Heights.

Meadow Vole (Microtus pennsylvanicus pennsylvanicus) — The

population of the meadow vole is cyclic and at times it is very

abundant. There is a die-off about every four years. The largest

specimen that I have handled was a nonpregnant female that

weighed 59.5 grams.

Prairie Vole (Microtus ochrog aster ochrog aster) — This vole is

found on high grassy areas and is local in distribution. It is less

numerous than the meadow vole.

Pine Vole (Microtus pinetorum scalopsoides) — This vole is un¬

common. There are specimens in the Museum from the town of

Vermont and the northwest shore of Lake Mendota. In spite of

special attempts to trap this vole, I have never taken but one

specimen, and this was near Cross Plains.

Muskrat (Ondatra zibethicus zibethicus) — The muskrat is com¬

mon in the marshes of Dane County. There are overland move¬

ments in spring, and some are killed on the highways. I have found

roadkills on University Avenue. It is the most valuable fur bearer

in the county. In addition to storing the rhizomes of aquatic

plants, it will also invade cornfields and store the ears.

Norway or Brown Rat (Rattus norvegicus) — The first definite

date for the appearance of the Norway rat in the county was

1854. In summer and fall many rats move into fields, especially

cornfields, but most of them winter in buildings. It is sometimes

found along lakeshores where it survives on dead fish. Errington

(1935), in the early winter of 1930-31, found them living in holes

in the banks of Lake Kegonsa where they subsisted on dead fish,

ducks, and other animal matter.

It is impossible to exterminate this rodent, and the only alterna¬

tive is to keep it under control by poisoning. Dr. Karl Link found

that dicumarol was an effective rat poison and is known as

Warfarin. It was used in the village of Middleton in the fall of

1950, and most of the rats were exterminated. Rats have the ability

to detect poisons and in some cases become more or less immune

to them.

House Mouse (Mus musculus domesticus) — This mouse is com¬

mon and familiar to all. Though usually found in buildings, in

,

1973] Schorger — The Mammals of Dane County 81

summer it will range a considerable distance into the country.

In September, 1943, I caught two in the sedges at the edge of the

former marsh at University Bay.

Jumping Mouse (Zapus hudsonius hudsonius) — The jumping

mouse is found in grassy areas, especially low meadows and

marshes. It is sometimes abundant. In the fall of 1967, in the

marsh on the west shore of Lake Waubesa, I trapped 17 in an

area of about one acre. One of these had a white tip to the tail.

In the fall of 1950 I caught three specimens with white tail tips

at the western end of Six Mile Creek Marsh, town of Springfield,

Dane County. The white tip is supposed to be characteristic of

Napaeozapus (Schorger, 1951).

Coyote ( Canis latrans thamnos) — The coyote was once numerous.

It is doubtful that it has been completely eliminated. In May, 1920,

two students found a den containing eight young on the north

shore of Lake Mendota. J. R. Smith, of the present Department

of Natural Resources wrote to me that he killed one near Mazo-

manie in 1948. Peter Rindisbacher, Swiss artist, came to Lafayette

County in 1826, and painted a small picture of a coyote which

is in the files of the Wisconsin Historical Society.

Timber Wolf (Canis lupus lycaon) — The timber wolf occurred

in small numbers. Black and white phases were known. The last

date of capture was January 18, 1888, when five gray wolves

were seen near Lake Wingra and two of them were killed.

Red Fox (Vulpes fulva fulva) — The red fox is common and is

found usually in open country. The typical red fox is readily

distinguished by the color and the white tip of the tail. Three

variations in the pelage occur occasionally. These are the black

or silver, cross, and Sampson. The cross fox has a dark cross on

the shoulders and back. The Sampson fox, which is rare, lacks

the guard hairs. Only the silver fox has been recorded for the

county. The heaviest red fox that I have examined was a male

weighing 13 lbs.

Gray Fox (Urocyon cinereoargenteus ocythous) — The gray fox

is common and found principally in woodlands. It is the only fox

that has the ability to climb trees. Two females taken at Token

Creek weighed 9 lbs. 4 oz. and 10 lbs. 5 oz.

Black Bear (Euarctos americanus americanus ) — The black bear

was never more than locally common in the county as there was

so little woodland. At intervals there was a migration from the

north apparently encited by a lack of food. The last bear mentioned

for the county was in 1879.

Raccoon (Procyon lotor hirtus) — The raccoon is common and on

occasion is found in almost any part of the city of Madison. At

one time a pair raised its young in a sewer two blocks from my

82 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

house. On October 16, 1967, a roadkill (female) was found that

weighed 17 lbs. 4 oz. Henry Murphy, formerly of Belleville, stated

that the largest raccoon that he ever caught weighed 29 lbs. He

caught one that was light yellow with pink eyes and called a

“sand coon.”

Least Weasel (Mustela rixosa allegheniensis) — The least weasel

is quite common and occurs most frequently at the edge of marshes.

I have specimens that drowned in the trout pools at the Nevin

Fish Hatchery. On July 20, 1949, I caught a half grown specimen

on the Nakoma Hill as it ran across the road. I have a specimen

given me by Cleveland Grant that was captured at Mineral Point

on March 15, 1961. This appears to be the only record for Iowa

County.

Short-tailed Weasel (Mustela erminea bangsi) — This is probably

the most common weasel although there are no skins from the

county. I have seen them alive on several occasions and have found

roadkills. The roadkills were so crushed as to be unpresentable.

Long-tailed Weasel (Mustela frenata noveboracensis) — The long¬

tailed weasel is less common than the short-tailed weasel. There

is a specimen in the collection of the Department of Wildlife Ecol¬

ogy taken in the Arboretum on January 5, 1946. I have found road¬

kills on August 9, 1939, and August 28, 1946.

Mink (Mustela vison letifera) — The mink is decidedly aquatic.

Owing to the abundance of lakes and streams in the county it is a

common mammal. It is active principally at night, although I have

on several occasions encountered it in broad daylight. Due to its

fondness for fish it has been trapped on numerous occasions at the

Nevin Fish Hatchery. It is the chief enemy of the muskrat.

Badger (Taxidea taxus jacksoni) — though not common the

badger is distributed throughout the county. Occasionally it is

found on the edge of Madison. On August 6, 1970, Robert Ellar-

son observed a bloated roadkill near the Kapec Orchards, a short

distance west of Madison.

Striped Skunk (Mephitis mephitis hudsonica) — The striped

skunk is common and well known from its obnoxious odor. It is

unimportant in the fur trade. It has no aversion to dwellings and

on August 28, 1961, I found a roadkill within half a block of Ran¬

dall Stadium.

Otter (Lutra canadensis canadensis) — At present the otter is

found principally along the Wisconsin River. Formerly it occurred

throughout the county, but currently it is rare in the interior.

On February 5, 1948, Dr. Harland W. Mossman photographed

otter tracks found at University Bay.

Puma (Felis concolor schorgeri) — The former status of the

1973] Schorger — The Mammals of Dane County 83

puma is uncertain. There is but one specific record. Locke states

in 1840 that the puma still existed in the Blue Mounds region.

Canada Lynx (Lynx canadensis canadensis) — The lynx appar¬

ently was never common. Several specimens have been killed in

the county. At the second regular meeting of the Wisconsin

Academy of Sciences on July 19, 1870, the following donation to

the Museum of the Academy was presented: “A lynx, killed near

Madison, and presented by Jacob Seiter.” On June 14, 1936, Her¬

bert Stoddard informed me that he assisted Ed. Ochsuer mount

a lynx killed near Middleton in 1907.

Bobcat (Lynx rufus superior ensis) — The bobcat was formerly

common in the county but is now extinct.

Elk (Cervus canadensis canadensis) — There is no record of a

live elk having been seen in the county as it was probably ex¬

terminated prior to 1800. Numerous antlers have been found by

seining in the county, and a nearly complete skeleton was found

in Lake Wingra (Schorger, 1954).

White-tailed Deer (Odocoileus virginianus borealis) — At the

time of settlement, deer were plentiful, but being shot at every

opportunity they became nearly extinct by 1880. At the present

time deer are found throughout the county. In the early 1930’s

wild deer entered the Arboretum and have been there ever since.

It sometimes enters the suburbs and causes destruction by dashing

through windows.

Bison (Bison bison bison) — There is no definite record of the

bison in Dane County, but it is included as it formerly ranged

throughout the entire southern portion of the state. In 1922, Thure

Kumlien stated that: “the early settlers found buffalo horns in

the vicinity of Lake Koshkonong” (Schorger, 1937).

HYPOTHETICAL REPORTS

Snowshoe hare (Lepus americanus phaeonotus) — In December,

1873, a Madisonian hunted near Westport which was then a post

office on the border of sections 21 and 22, town of Westport. The

woods and brush were swarming with huge “Jack Rabbits.”

Though he had hunted in the area for years he had never seen

game of this kind previously. After killing several, he brought

home only two on account of their weight. It would be preferable

that so confusing a note as to species had never been printed. The

date is too early for a jack rabbit. In weight the hare is only a

little heavier than the cottontail, but the long hind legs give the

impression of greater size. If the rabbits were snowshoes, they

should have shown sufficient white to be conspicuous. No color

was mentioned. It is quite probable that the snowshoe occurred

in the tamarack swamps, particularly in the eastern part of the

84 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

county. Formerly there was considerable tamarack along the Ya-

hara in the town of Westport. The snowshoe was found in the sur¬

rounding counties of Jefferson, Dodge, and Sauk. Snyder (1902)

was informed by hunters “of a very large, big-eared rabbit, oc¬

curring in a tamarack swamp near Lost Lake,” town of Calamus,

Dodge County. Jebez Brown (1855) lived near Ironton, Sauk

County. On January 11, 1858, he shot one of two white rabbits,

and on the following November 27, he shot another. Canfield

(1870) dismisses it with “seldom seen.” In view of the slight

amount of suitable habitat in Sauk County, it is surprising that it

occurred at all.

Red-backed Vole (Clethrionomys gapperi gapperi) — This vole

has yet to be taken in the county. The stomach of a snowy owl

collected on Lake Mendota on February 14, 1961, according to

Keith (1963), contained the remains of four red-backed voles.

From the enamel pattern I identified them as pine voles (Microtus

vinetorum) . The identification was confirmed by John L. Paradiso,

U.S. National Museum. The red-backed vole has not been taken

closer to Madison than Beaver Dam.

Porcupine (Erethizon dorsatum dorsatum) — There is no definite

reference to show that the porcupine was indigenous. Several porcu¬

pines have been taken in the county, but in all cases it is highly

probable that they were escapes or liberations after having been

brought into the county.

SUMMARY

Fifty-three species are recognized as having occurred in this

south-central Wisconsin area in recent times. Bison, elk, bobcat,

puma, lynx, black bear, and timber wolf have been extirpated;

jack rabbits have been present for a time as the result of plantings.

White -tailed deer have largely recovered from the overexploitation

of the 19th century. Aquatic mammals, such as the beaver and

otter, are as easily taken as land mammals and still exist in small

numbers.

The only native mammal that shows an increase in numbers

is the opossum, which has moved steadily northward in the present

century.

REFERENCES

BROWN, JEBEZ. 1855-72. Unpublished diary. Wisconsin State Historical

Society.

CANFIELD, W. H. 1870. Outline sketches of Sauk County. Third sketch.

Baraboo. 39 pp.

CLOUGH, G. C. 1965. Physiological effect of botfly parasitism on meadow

voles. Ecology, 46:344-46,

85

1973] Schorger- — The Mammals of Dane County

ERRINGTON, P. L. 1935. Food habits of mid-west foxes. J. Mammol., 16:

192-200.

JACKSON, H. H. T. 1961. Mammals of Wisconsin. Madison. 504 pp.

KEITH, L. B. 1963. A note on snowy owl food habits. Wilson Bull., 75:

276-77.

KNIPPING, P. A., B. B. MORGAN and R. J. DICKE. 1950. Preliminary list

of some fleas from Wisconsin. Trans. Wis. Acad. Sci., Arts and Lett.,

40:199-206.

LOCKE, J. 1840. Earthwork antiquities in Wisconsin territory. In D. D.

Owen, Report of a geological exploration of Iowa, Wisconsin and Illinois.

26th Cong., 1st Ses. House Doc. Vol. 6, No. 239, Ser. No. 368.

NELSON, A. L. 1934. Notes on Wisconsin mammals. J. Mammol., 15:252-53.

RONGSTAD, O. J. 1965. A life history study of thirteen lined ground squir¬

rels in southern Wisconsin. J. Mammol., 46:76-87.

SCHORGER, A. W. 1937. The range of the bison in Wisconsin. Trans. Wis.

Acad. Sci., Arts, Lett., 30:117-30.

SCHORGER, A. W. 1951. Zapus with white tail-tip. J. Mammol., 32(3) :362.

SCHORGER, A. W. 1954. The elk in early Wisconsin. Trans. Wis. Acad. Sci.,

Arts, Lett., 43:5-23.

SNYDER, W. E. 1902. A list with brief notes of the mammals of Dodge Co.,

Wis. Bull. Wis. Nat. Hist. Soc., 2:113-26.

VAN METER, A. C. 1881. New Richmond Republican May 11.

GERMS, LUMBERJACKS, AND DOCTORS IN

NINETEENTH CENTURY MARINETTE

Carl E. Krog

University of Wisconsin Center —

Marinette

Unlike many frontier communities Marinette had the services

of physicians from the very beginning of settlement. Dr. Jonathan

Hall, who came to Marinette from upstate New York, owned and

operated a sawmill during the 1840’s and 1850’s. His mill failed

during the late 1850’s, but he remained, providing medical services

for the growing community during the 1860’s. A second New

Yorker, John Sherman, worked for Hall as a timber cruiser and

bookkeeper until the 1860’s, when with Hall’s encouragement he

went to Chicago to study medicine at Rush Medical School. The

first school teacher, community historian, founder of the first

church, Pioneer Presbyterian, organizer of the first Masonic Lodge

in the region, reform mayor, “Old Doc Sherman” was one of the

most influential members of the settlement in converting what was

once a lumbering camp into a permanent village.

The Chicagoans and Milwaukeeans who owned most of the large

Marinette lumber companies regarded the growing settlement on

the south bank of the Menominee river as a kind of unwanted by¬

product of their sawmills. The community grew from a population

of 400 in 1860 to 4,000 in 1880, and quadrupled in population dur¬

ing the last two decades of the nineteenth century, attracting

merchants and professional people. Both groups suffered from

outside competition ; the merchants from the lumber company

stores and the village doctors from itinerant physicians who like

many of their lumberjack patients were transients, interested in

making a few dollars as they passed through the community.

In July, 1886, citizens learned that a Dr. Edward Fishblatt was

coming to Marinette. Former professor of Atlanta Medical Col¬

lege and present editor of the Neiv York Medical and Surgical Re¬

view, Fishblatt specialized in chronic diseases. Among his credits

were curing patients suffering from ruptures, piles, tumors, and

kidney infection, as well as helping the lame to walk, and young

men troubled by impotency. Physicians like Fishblatt visited the

community about once every six weeks and appear to have at¬

tracted a large number of patients.1

1 Marinette Eagle, July 3, 1886, August 28, 1886, and December 4, 1886.

87

88 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Medical practices and facilities in Marinette gradually improved

during the last decade and a half of the nineteenth century. As

early as April, 1875, the Marinette Eagle announced plans for a

workers’ hospital. The plans called for a program whereby a

worker would pay one dollar the first month and fifty cents each

month thereafter for hospitalization. Nothing came of the plan,

though the need for hospitalization remained.

Both lumberjacks and sawmill workers followed hazardous oc¬

cupations and accidents were frequent. There was a high rate of

turnover in the lumbering industry, and young inexperienced

crews increased the accident potential. Falling trees, drownings in

the river during log drives, and lack of protective screens and

guard rails in the sawmills took their toll. (Drunkenness does not

appear to have been a major contributing factor.) It was reported

that 120 men were injured in the woods of Menominee river pinery

during the winter of 1898-1899 alone. The close, unhygienic con¬

ditions of the lumber camps made epidemics a constant threat.

Lumberjacks, welcomed in small towns for the money they spent

on services, spread their camp-bred diseases to the urban popula¬

tion. Smallpox remained a continuous scourge to lumbering towns

down to the early years of the 20th century.2

The need for medical service in the lumber camps was obvious,

but because of the remote location of many camps and the improvi¬

dent habits of lumberjacks, doctors were reluctant to leave lumber¬

ing towns. A partial solution was found by three Marinette physi¬

cians, Doctors Frank Gregory, J. A. Somerville, and H. E. Mann

during the middle of the 1880’s. The employees of the Peshtigo

Company, living and working at Peshtigo Harbor, a community

of nearly five hundred, had had difficulty securing the services of

physicians. Located at the mouth of the Peshtigo River, Peshtigo

Harbor was fourteen miles from Marinette. Marinette physicians

charged five dollars a call at the harbor, a prohibitive sum for a

laborer earning a dollar or a dollar-and-a-half a day. The three

doctors entered into an agreement with the Peshtigo Company to

furnish medical care to all employees of the company at a rate of

$1.25 per month for a married man and his family, and $0.75 a

month for a single man. The patient had his choice of the doctors.

The Peshtigo Company agreed to deduct the fee from each man’s

pay and remit the sum collected to the doctor designated to re¬

ceive it.

The Menominee River Hospital was organized in 1886, and three

years later a branch was opened in Menominee. The Marinette and

Menominee Hospital Company was incorporated the same year,

2 George Engberg, “Labor in the Lake States Lumber Industry, 1830-1930”. Unpub¬

lished, Ph.D. Dissertation, University of Minnesota, 1949, p. 229. Quoted figures were

taken from the Mississippi Valley Lumberman XXX, March 24, 1899.

1973]

Krog — Germs , Lumberjacks, and Doctors

89

with Dr. H. E. Mann, president, and Dr. J. A. Somerville, secre¬

tary and treasurer. Hospital confinement, except in very serious

cases, was rare in the late nineteenth century, yet by the early

1890’s the two hospitals were treating over five hundred cases an¬

nually. Because of the high number of accidents in the lumber

camps, the Marinette hospital reported that it had an average of

25 patients daily.3

The first hospital quarters were primitive. A boarding house

served as the first hospital building. Injured lumberjacks some¬

times arrived in railroad baggage cars, hand cars, and in one case,

strapped to a log which served as a splint and stretcher for a man

with a broken thigh. Patients were hand-carried by physicians

and attendants to the hospital operating room. Chloroform and

ether were the only anesthetics and lysol the major disinfectant.

In spite of the primitive working conditions and equipment, diffi¬

cult and serious operations were performed from the beginning.

A Caesarean section was done a few months after the hospital

opened. Several months later, a young man entered the hospital

with typhoid fever. He remained for the next half century. To pay

his hospital bill, John E. Boren began working for the hospital.

Shortly, he was made superintendent of the hospital. A number

of his descendants became physicians and have continued to serve

the community. Time brought improvement in hospital equipment

and staff experience. In 1889, Dr. T. J. Redlings, a graduate of the

University of Wisconsin-Madison, came to Marinette following

two years of graduate study in Germany. Redlings brought the

first microscope to the community. Successful in detecting the ever¬

present typhoid germ, he helped lead the community fight to sup¬

press the disease.4

To provide medical service and insure payment from the notori¬

ously improvident lumberjacks, the Menominee River Hospital or¬

ganized a hospital insurance program. Under its jovial salesman,

James J. Stephens, who visited the lumber camps, the hospital pro¬

vided a dual program for medical protection. In exchange for a

five dollar annual ticket, a lumberjack received all the hospital

care he might need, including physician’s service and surgery. For

ten dollars a year the ticket holder received the same hospital care,

plus a dollar a day while hospitalized.

Both the lumbering industry and village prospered and grew

during the 1880’s. The late 1880’s marked the peak production

years and the busy sawmills attracted a large labor force. The

growing population demanded increased municipal services — serv-

3 Marinette Eagle, February 11, 1893; Frederick Burke papers; State Historical

Society of Wisconsin, Madison.

4 Frederick Burke papers, State Historical Society of Wisconsin, Madison, Marinette

Eagle-Star, November 8, 1933 and November 14, 1933.

90 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

ices which could only be provided by a city government. In spite

of the opposition of some of the mill owners, Isaac Stephenson, the

community’s most prominent lumberman, went ahead with plans

to incorporate the settlement and in March, 1887, Marinette joined

the ranks of Wisconsin cities. At incorporation, Marinette already

had a sizeable population and according to the United States Cen¬

sus of 1890, the three year old city ranked eleventh in the state,

well ahead of Wausau, Kenosha and Manitowoc.

A number of serious problems which required the immediate

attention of new municipal government had accumulated during

the decade of 1880’s. The first to be faced by Marinette’s new gov¬

ernment was that of a public water supply. Was water to be taken

from the bay or from the river, which was closer and hence

cheaper? Would a private company or the city provide the water?

The city decided that the bay was less polluted than the river.

The city lacked the revenue to build its own waterworks. Incor¬

poration had not changed the lumber companies’ under-taxation.

The local lumber powers, aware that the city’s chief industrial

days were numbered, were not interested in investing in a water

utility. With the decision that the bay was less polluted than the

river, the city signed a 25 year contract with the American Water

Works and Guarantee Company. The company, owned mainly by

capitalists from Boston and New York, agreed to lay 12 miles of

pipe, build 125 fire hydrants, and provide free water for both pub¬

lic buildings and churches. The contract appeared to be most fa¬

vorable for Marinette. The installation of Marinette’s water system

was to cost 20 per cent less than neighboring Menominee’s. Mari¬

nette was to secure 125 hydrants at $46 per hydrant to Menomi¬

nee’s 94 at $73 apiece.5

The bargain soon revealed itself to be short-lived and short¬

sighted. In 1895, Dr. John Coulter of the city health department

reported that he believed the increased number of sick people in

the city was due to impure water. Marinette-Menominee with a

total of 28,000 people living at the mouth of the Menominee river,

dumped a large amount of offal into the bay. The report of the

water company’s chemist of Western University, Allen, Pennsyl¬

vania, did not say that the water was fit to drink. The report on

the city’s water from a University of Wisconsin chemist was more

explicit — the water was impure. The wealthy people in Marinette

continued to rely on their private wells.6

Nothing was done to remedy the situation by either the city or

the water company until a severe epidemic of typhoid fever struck

in 1899. In March, 33 deaths from typhoid were attributed to the

5 Marinette Eagle, July 13, 1887 and March 13, 1888.

0 Marinette Eagle, March 30, 1895 and April 27, 1895.

1973]

Krog — Germs , Lumberjacks , and Doctors

91

impure water. Finally, in October, 1899, an agreement was reached

between the city and the water company. The water company

agreed to install a water filter at the cost of $25,000. In return,

the city agreed to pay half of the operating cost for seven years.

The water company, in turn, received a 25 year charter and the

hydrant rental was raised to $1250 annually. These changes forced

the city to raise its tax rate on $1,000 of property from 3% to

3. 5%. 7

A water filter was vital for the city’s health. Of the water users

in Marinette in 1899, 894 or 84 per cent used city water, but only

13 per cent bothered to boil it. Of 69 cases of typhoid fever re¬

ported that October, 66 used city water. At the end of the year

the death rate, including those from typhoid, was 14.72 per 1,000.

Without the typhoid deaths, the death rate would have been only

8.69. After the installation of the water filter, the scourge of

typhoid fever gradually decreased. During the 12 months after

April 1, 1900, only two out of 199 deaths were attributed to typhoid

fever, giving the city a death rate of 12.23 per 1,000 in 1900.

Nevertheless, as late as 1903, three years after the water filter

installation, the city health commissioner reported 89 cases of

typhoid with eight deaths.8

After 1900, pure milk, like pure water, was important in cutting

the death rate in Marinette. Regulation began in 1894 when the

city passed an ordinance with the specific aim of ending the un¬

sanitary conditions of some dairies. In the late winter of 1897,

both Marinette and Menominee passed ordinances demanding

tuberculin tests for dairy cattle in the two-county area providing

milk for the two cities.

The problem of a high infant mortality rate persisted, nonethe¬

less, during the first years of the 20th century. In 1902, children

under the age of five accounted for 39% of the deaths in Marinette.

The year was a marked improvement over 1900 when children

under five accounted for half of the deaths in the city. Progress

was gradual and uneven. As late as 1908, the city health depart¬

ment could proudly report that “only” seven infants had died of

cholera infantum during the warm days in the middle of Septem¬

ber of that year. The death rate in Marinette continued to go down

after 1900, with the purification of milk and water probably the

most significant factors in the decline. In 1900, the death rate was

14.43; 1901, 12.22; and 1902, 10.76. Many of the other causes of

death were further reduced as the century progressed. Other ma¬

jor causes of death listed by the city health commissioner in 1902

7 Marinette Eagle , October 7, 1899, October 21, 1899, June 18, 1899, and January 6,

1900.

8 Marinette Eagle-Star } January 8, 1904 and October 28, 1902.

92 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

included: respiratory ailments, 21; pneumonia, 16; tuberculosis,

15; external violence, 18; typhoid fever, 7 ; and diphtheria, 7.°

Along with impure milk and water, Marinette’s citizens faced

the hazards of consuming diseased beef. The United States Con¬

gress had passed the Pure Food and Drug Act in 1906, following

the national uproar over Upton Sinclair’s book The Jungle, which

realistically described conditions in the Chicago stockyards. The

federal government, however, inspected interstate meat only.

A few meat peddlers in Marinette bought old and diseased cattle

from nearby farmers and sold the meat at cut-rate prices directly

to customers from their carts. City authorities seized a cow from

such a cart to inspect it in March, 1909, but before the cow could

be checked, it disappeared.

A few months later, a cow which was being led to a butcher col¬

lapsed in the main business district. The peddler claimed that the

animal was merely tired, but he hastily buried the carcass before

investigation could begin. A few months earlier, the city attorney

had recommended that the city council pass an ordinance forbidding

the selling of cows for slaughter during gestation, but nothing

had been done. The tale of the cow’s death was too much for a

number of prominent housewives, who signed a petition asking the

Marinette city council to pass a meat inspection act similar to that

of Milwaukee. They demanded a strict sanitation code providing

for the inspection of meat markets and peddlers’ carts, and pro¬

hibition of selling wild boars as pork and sick and unborn calves

as veal. The city inspection code was passed.10

In passing its code, Marinette was following the lead of large

cities in the United States, which during the decade of the nineties,

were spending large sums on sanitation and health departments

with gratifying results. In Milwaukee the death rate in 1891 was

21 per 1,000; in 1898 it was 11.92. The death rate in Denver de¬

clined from 23.71 to 11.55 during the decade of the 1890’s.

The city of Marinette with 16,000 people spent $472.55 on its

health department in 1900. To match the expenditures of Denver

that year, Marinette would have had to spend $4,503 and to match

Milwaukee, $5,271.

Nonetheless, the death rate in the city compared favorably with

the national rate, which was 17 per 1,000 in the first year of the

20th century. The life expectancy of an American male in 1900 was

only 49 years, the nation’s three deadliest killers being tubercu¬

losis, typhoid fever, and pneumonia. In 1921 the death rate in

9 Marinette Eagle, July 7, 1894, April 3, 1897, July 23, 1901, and Marinette Eagle-

Star, October 28, 1902, and October 7, 1908.

10 Marinette Eagle-Star, April 8, 1909, March 25, 1909, June 29, 1909, and Febru¬

ary 14, 1910.

1973]

Krog — Germs, Lumberjacks, and Doctors

93

Marinette was 11.3. The rate in neighboring- Green Bay was 16.7 ;

Wausau, 11.3; Ashland, 18; Manitowoc, 12.2; and Oshkosh, 13. 3. 11

Humming sawmills and a bustling community left in their wake

a need for better medical care and an effective city government

to provide minimum municipal services. The group insurance plan

of the Peshtigo Company and the Menominee River Hospital both

brought medical care to lumberjacks and sawmill employees where

none had existed. Through the persistent efforts of Dr. John Coul¬

ter, the community was able to secure pure water. These efforts of

the last century paved the way for a longer life expectancy for

Marinette citizens in this century.

11 Marinette Eagle, March 10, 1900, Statistics from Wisconsin State Board of Health,

Bureau of Health Statistics, Madison, Wisconsin.

THE EASTERN SUBTERRANEAN TERMITE,

RETICULITERMES FLAVIPES (KOLLAR), AND THE

COMMON THIEF ANT, SOLENOPSIS MOLEST A (SAY),

IN THE LABORATORY, WITH NOTES ON OTHER

ASSOCIATED ANT SPECIES

R. V. Smythe and H. C. Coppel

University of Wisconsin —

Madison

Lestobiotic ants, those which rob and prey on termite broods

or disabled termites, are generally of the genera Solenopsis, Mono -

monorium, or Carebara (Escherich, 1909; Hegh, 1922). The ants

have minute, pale yellow, blind or myopic workers and nest in the

interstices of the nests of larger ants or termites. The workers

gain access to the nurseries of their prey by excavating tenuous

galleries and devour the brood. Solenopsis molesta and the Euro¬

pean S. fugax Latreille both inhabit termitaria. S. molesta is “not

infrequently” found in the nests of Reticulitermes flavipes (Kollar) ,

R. lucifugus (Rossi), and R. virginicus (Banks) (Wheeler, 1936).

S. texana has similar habits in the southern states and S. caro-

lensis Forel and S. truncorum Forel, occur in termite nests in

North Carolina (Forel, 1928).

A cluster of 2,000 termite eggs was found about 18 in above the

ground under the bark of a dead elm tree at Janesville, Wisconsin

in 1963. One to 2 ft above the egg mass was a colony of S. molesta,

also under the bark. Once disturbed, the ants swiftly removed their

eggs and larvae as well as the termite eggs below them. Within 10

min after the termite eggs were uncovered, the thief ants had

collected and carried off considerably more than the termites had,

and moved them out of sight. On four other occasions that sum¬

mer, colonies of S. molesta were found under the bark of dead elm

trees. On two occasions, R. flavipes was active in runways and

tunnels adjacent to the ants; on one occasion the ants were active

in abandoned termite runways beneath the bark ; and on one occa¬

sion the ants were on a tree where there was no termite activity.

These observations prompted laboratory studies on the interaction

of R . flavipes with S. molesta, particularly, and with other associ¬

ated ant species generally.

95

96 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

MATERIALS AND METHODS

It was considered probable that any interrelations between S.

molesta and R. flavipes would be initiated by the ants. Thus, in

all the experiments, the termites were established first in various

containers and S. molesta individuals were added later. In tests

with other ant species, the ants were sometimes established first

and sometimes the termites. Varying numbers of ants and ter¬

mites were used in the different tests and these details will be in¬

cluded in the discussion of each particular test. Most of the activity

occurred at the base of the test units, consequently, the cultures

were observed by placing the unit on a wooden rack beneath which

was placed a magnifying mirror and a desk lamp to enhance the

illumination. When not under observation, the containers were

maintained at 25 °C.

RESULTS AND DISCUSSION

Two experiments were initiated, in which 200 R. flavipes and 50

termite eggs each were added to a clear plastic unit (4 x 7.5 x 10.5

in) containing damp earth and two small (0.5 x 3 in) pieces of

wood. Twenty-four hours later the eggs in one unit (Exp. 1)

had been moved and were not seen again. After 1 mo, a colony

FIGURE 1. Nest cavity of Solenopsis molesta with ant tunnels and tunnels

of Reticulitermes flavipes. Experiment 1.

1973] S my the and Coppel — Notes on Ant Species 97

of 50 S. molesta adults and 20 larvae were added. The ants quickly

hollowed out an area in the dirt at the bottom of the unit and de¬

posited their larvae in its center. The ant “nest” was surrounded on

all sides by termite tunnels (Fig. 1) . Within the first week the ants

constructed minute passageways from the nest cavity to the termite

tunnels 4 and 5 (Fig. 1, CA and DB). During the second week the

ants moved through tunnels 2, 3, 4, and 5 which they blocked off

for their own use. The ants erected earthen walls in the termite

tunnels and created a circle of diameter about 3 in, which excluded

the termites. The ant nest was approximately at the center of the

circle. Toward the end of the third week, 5 dead termites were

observed in tunnels 1, 6, 7, and 8. On the 25th day 7 dead termites

were seen in a ring around the ants’ nest. We offer no explana¬

tion for this phenomenon and it was not repeated.

The termites dug intermittently at the earthen wall blocking

their entrance to tunnel 4. One soldier was always close behind

the few digging workers. These workers often became excited

and retreated quickly, shaking violently (vibrating their bodies

on stiff legs). A termite worker finally broke through to tunnel 4

but remained in 3 with only his antennae in the ant tunnel. During

this termite activity the ants rarely utilized the portion of tunnel

4 below A but moved their larvae from the center of the nest

and piled them in the cavity as far as possible from the scene of

activity.

The termite which had penetrated the ant tunnel remained in

the same position for about 1 min at which time an ant began

moving toward him. The ant slowly moved to within 1 cm of the

termite, then quickly turned and retreated. The worker termite

withdrew and was replaced by a soldier termite who placed his

head into the confluence of the two tunnels. Within a few minutes

4 ants moved down tunnel 4 toward him. The leading ant moved

to within a few mm of the termite and retreated just as it ap¬

peared the soldier would lunge. The following 2 ants immediately

turned around about 15 mm from the termite and withdrew. The

fourth ant moved directly to the termite’s mandibles and rapidly

turned around. Only after the ant had reversed himself did the

soldier lunge. The ants then began to block tunnel 4 from A up¬

ward for about 15 mm. A similar wall of dirt was established in

tunnel 5 upward from B and a much smaller wall was erected at D.

The latter barrier was only about 2 mm thick. The termites began

to penetrate and widen the ant tunnel DB, apparently having

gained access to it from a tunnel in the soil not visible from below.

One termite reached the thin earth barrier at D and slowly began

to remove some of the dirt. The termite activity greatly activated

the ants which immediately and rapidly began bolstering the

98 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

earthen wall from the other side. For about 2 hr the termites

slowly broke down the wall while the ants rapidly strengthened it.

The end result was a barrier about 1 cm thick. At this point the

termites neglected \\e barrier and retreated. During the frenzied

activity by the ants, some of them seized their larvae and removed

them from the nest cavity and thus out of sight. Within 8 hr after

the cessation of activity by both species, the ants moved most, if not

all, of their larvae to their original position in the nest, and nothing

unusual occurred for the next 6 months.

The ants in the second unit (Exp. 2) immediately established

themselves in a cavity (X) on the floor of the container (Fig. 2).

About 12 ant larvae were placed in the nest. As in the first experi¬

ment, the ants were completely surrounded by termite tunnels.

For the first week no ants moved on the bottom of the container;

however, as never more than 25 ants were visible in the cavity, it

was assumed that they had established or usurped tunnels not visi¬

ble from below. On the tenth day, some ants were observed in

tunnel 1. They established a tunnel (CA) from their nest cavity

to the termite tunnel and blocked off a portion of the tunnel with

earthen walls. Two days later the ants tunneled (DB) to termite

tunnel 2 and moved in it. They erected no visible barrier at B

X - ANT NEST

. « ANT TUNNEL

— « termite tunnel

FIGURE 2. Nest cavity of Solenopsis molesta with ant tunnels and tunnels

of Reticulitermes flavipes. Experiment 2.

1973] Smythe and Copy el — Notes on Ant Species 99

but often seemed to move upward through the soil at this point

in a tunnel not visible from below.

Termites moving along tunnel 3, after the erection of the ant

wall at A, seemed to be aware of the barrier and only rarely did

a termite turn off 3 and enter 1 moving toward A. On all but one

of these sporadic occasions the termite did not proceed to A but

turned around and re-entered 3. Once, a termite moved to the wall

at A, waved his antennae slowly with their tips in contact with

the barrier, then quickly backed away, reversed himself, and re¬

turned to tunnel 3. The termites never tried to break through any

of the ants’ earthen walls nor did the ants attempt to extend their

range. After 9 weeks the experiment was discontinued.

The following tests were initiated to obtain data on the behavior

of the ants toward termite eggs. In test A, an 8 dr shell vial was

ys filled with soil and 25 ants were added. After 48 hr, 15 termite

eggs were placed in a cluster on top of the soil. Two ants almost

immediately found the eggs and picked up several. Within 30 sec

about a dozen ants had converged on the eggs and had placed

them in a pile under a small piece of bark which extended partially

into the soil. Several ants frequently handled the eggs with their

mandibles. For 3 days the ants “cared for” the eggs by handling

them and moving them. On the fourth day the eggs showed signs

of shrinkage and the ants subsequently abandoned them.

For test B, 20 cc of moist soil were placed in a clear plastic

zipper case (2 in diam. x 1.4 in high) and 25 ants were added.

After 48 hr, 2 termite eggs were placed on the soil surface. For 2

days the ants showed no interest in the eggs. An additional 15

ants were introduced and after 6 hr the termite eggs had disap¬

peared. The contents of the test unit were carefully examined but

no eggs were found. Probably they were eaten by the ants.

Test C replicated B but used 15 termite eggs instead of 2. Within

3 hr the ants had located the eggs and moved them underground

where they could not be seen. After 2 days, the test unit was ex¬

amined. The eggs were in good condition in a little pocket near

the bottom of the container. One egg appeared close to hatching.

It was immediately transferred to a new container with moist

earth and ants but the ants ignored it and the termite died with

only its head free of the egg. It is not known whether S. molesta

or other ant species assist young termites in emerging from their

eggs.

Attempts were made to induce the ants to steal termite eggs

from an established termite colony. Twenty cc of moist earth

were placed in a zipper case with 15 termites and 40 termite eggs.

The termites located the eggs almost immediately. A termite would

pick up one or more eggs, carry them a short distance and drop

100 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

them. This was repeated many times. After 20 hr the termites had

gathered about 12 eggs under a small piece of wood at the soil sur¬

face. Fifteen ants were added to the test chamber. Within 2 hr they

had tunneled to the bottom along the wall of the container and

hollowed out a small chamber. In the meantime, some of the ants

and termites were on the soil surface. The two species moved

freely among each other without evidencing any hostile behavior.

Occasionally, when the antennae of the 2 species touched, the ant

would hastily jerk backward, then continue as before. The ter¬

mites rarely backed away. After 24 hr the ants had established

several tunnels through the soil. These were not contiguous with

the termite tunnels. Both species moved freely in their respective

tunnels and seemed indifferent or unaware of the other’s presence.

Careful observation for 6 consecutive days revealed nothing un¬

usual in the behavior of either species. Neither made any attempt

to extend its domain. Occasionally 1 or 2 individuals of both species

would contact each other on the soil surface with the same results

as described previously. After 1 week the termites began moving

more and more on the soil surface which is a preliminary indica¬

tion of an unhealthy colony. An inspection of the eggs showed

early signs of shrinkage. In 10 days the termites appeared un¬

healthy and the experiment was discontinued. Similar experiments

produced no significant results. Although the termites frequently

grouped clusters of their eggs, the ants were never seen to steal

them.

Though S. molesta never was seen to steal guarded termite

eggs, they readily seized unguarded eggs and at times appeared

to care for or tend them. The extent of this practice or its bio¬

logical significance is not known. An interesting behavioral ac¬

tivity of the ant was in walling off and usurping for its own use

the termite tunnels adjacent to its nest. Additionally, there was the

evident lack of hostility between the two species when they encoun¬

tered each other on the soil surface.

INTERACTION OF R . F LAV I PE S WITH

OTHER ANT SPECIES

A cluster of approximately 5,000 termite eggs was found about

5 ft from the ground under the inner bark of a large, partially

dead elm on July 30, 1963. Immediately after exposure of the

eggs, many Aphaenogaster tennessensis (Mayr) ants moved up

under the inner bark at the base of the tree from their nest in the

ground and began moving large numbers of termite eggs. Oc¬

casionally the ants would seize the termites which were also at¬

tempting to remove their eggs, but primarily the ants concen-

1973] Smythe and Coppel — Notes on Ant Species 101

trated on the eggs. When all the eggs were removed, the ants re¬

turned to their nest. Similar behavior records for A. tennessensis

have not been reported in the literature. Limited laboratory studies

were undertaken, but none gave any evidence of similar (egg¬

stealing) behavior. Laboratory studies indicated a strictly preda¬

tory relationship between the ant and termite.

Camponotus pennsylvanicus DeG., Tapinoma sessile (Say), and

Lasius sp. may be considered as inquiline ants. Laboratory experi¬

ments suggested that C. pennsylvanicus and R. flavipes, given suf¬

ficient room and a preponderance of termites, will establish a rela¬

tionship best described as indifferent neighbors. Field observa¬

tions suggest that once this relationship is established, it will exist

even under conditions of great agitation. However, crowding and

a preponderance of ants destroy the inquiline relationship and

the ants kill the termites.

Laboratory experiments with T . sessile indicated that regardless

of whether the ants or termites were established first, after 1

week a condition of indifferent tolerance with no direct contact

between the two species resulted. Although King (1897) reported

a close association between R . flavipes and T. sessile in field obser¬

vations, no such relationship was indicated in the laboratory stud¬

ies. The two species were not observed together in the field.

Lasius sp. and R. flavipes were frequently found closely asso¬

ciated in the field (Smythe and Coppel, 1964). On one occasion,

a weathered piece of wood lying on the ground was overturned

and both species were seen moving together on the ground. Labora-

try tests indicated that Lasius sp. occasionally would usurp por¬

tions of the termites’ tunnels for its own use. After several days,

however, the ants would withdraw and the termites would rein¬

habit their former territory. Mixed cultures of the two species in¬

variably stabilized with the termites inhabiting tunnels in the bot¬

tom half of the test unit and the ants living in the upper half of the

soil and on the soil surface. Frequently, tunnels of the two species

would connect but no intermingling of the species occurred. Occa¬

sionally, the termites would erect barriers in the connecting tunnels

but after several days they would remove them and thereafter re¬

side as indifferent neighbors. Gorging the ants with honey and then

introducing them to a termite inhabited unit in a small wire cage

for 48 hours, so that the odors of the two species could inter¬

mingle without any physical contact, eliminated all ant aggressive¬

ness.

SUMMARY

The eastern subterranean termite, R. flavipes, and the common

thief ant, S. molesta, may coexist harmoniously in uncrowded labo-

102 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

ratory cultures. Encounters on the soil surface between the two

species showed no evidence of hostility. The ants continuously

walled off and usurped portions of termite tunnels close to their

nests. Unguarded termite eggs were removed by the ants ; however,

in the studies undertaken guarded eggs were not removed. A. ten-

nessensis rapidly removed exposed termite eggs in the field. In the

laboratory, the ants C. pennsylvanicus , T. sessile, and Lasius sp.

generally had an inquiline relationship with R. flavipes, which was

often reflected as a neighborly indifference with or without contact.

The relationship with Lasius sp. seemed particularly compatible.

Under either crowded conditions or a preponderance of ants this

was modified in favor of the ants.

REFERENCES

ESCHERICH, K. 1909. Die Termiten oder Weissen Ameisen. Werner Klink-

hardt, Leipzig. 198 pp.

FOREL, A. 1928. The social world of the ants. Vol. II, G. P. Putman’s Sons,

Ltd., New York. 445 pp.

HEGH, E. 1922. Les termites. Louis Desmet-Vexteneuil, Brussels. 756 pp.

KING, G. B. 1897. Termes flavipes Kollar and its association with ants.

Entomol. News 8: 193-196.

SMYTHE, R. V. and H. C. COPPEL. 1964. Preliminary studies on ant-ter-

mite relationships in Wisconsin. Proc. North Centr. Branch Entomol. Soc.

Amer. 19: 133-135.

WHEELER, W. M. 1936. Ecological relations of Ponerine and other ants to

termites. Proc. Amer. Acad. Arts Sci 71 : 159-243.

ACKNOWLEDGEMENTS

Research supported by the College of Agricultural and Life

Sciences, University of Wisconsin, Madison, and by the Research

Committee of the Graduate School from funds supplied by the

Wisconsin Alumni Research Foundation and the Office of Naval

Research, Naval Biology Program, under contract No. N00014-67,

NR306-012. The authors gratefully acknowledge Dr. A. C. Cole,

Department of Entomology, University of Tennessee, Knoxville,

who identified the ant species.

SOCRATES ON CIVIL DISOBEDIENCE:

THE APOLOGY AND THE CRITO

Peter S. Wenz

University of Wisconsin —

Stevens Point

In the Crito (50a-52b)1 Plato represents Socrates as defending

what at least appears to be the view that one ought never disobey

either a law or a lawful command of the state. But in the Apology

Socrates is represented as condoning disobedience. First (29d) he

states that he would “never stop practicing philosophy” even if

so ordered by the jury at his trial. Second (32c-d) , Socrates

proudly tells of his disobedience to the Thirty Commissioners when

they ordered him to go to Salamis to get Leon.2 Thus, there is at

least an apparent inconsistency between the Apology and the Crito

on the question of disobeying the law.

There are two general approaches to this apparent inconsistency.

One is to admit that there really is such an inconsistency and then

to explain why or how it arose. The second is to offer an interpre¬

tation of either or both of the two dialogues in question accord¬

ing to which the inconsistency is merely apparent and never really

arises. In this paper I shall critically review various attempts

along these two lines (sections I and II, respectively),3 and then

present my own analysis, an attempt of the second type, an inter¬

pretation of the Crito according to which it is perfectly consistent

with the Apology .

I

First, it might be maintained that the two dialogues really are

inconsistent and are so due to Plato’s inability to detect the incon¬

sistency. This, however, is incredible, since the inconsistency is,

if anything, quite apparent. It is difficult to believe that anyone

with the ability to write the Apology and the Crito would simul¬

taneously have the inability to detect the apparent inconsistency

between them.

1 All references to the dialogues will be noted in the text. All quotes are drawn

from Plato: Collected Dialogues eds. Hamilton and Cairns, Random House (1963).

2 A third incident, recounted at 32b, is Socrates’ refusal to allow a group of naval

officers to be tried en bloc , but here it is made clear that Socrates was upholding,

not disobeying the law. So this case does not constitute an instance of Socrates’ dis¬

obeying either a law or a lawful command of the state.

3 For sections I and II, I am indebted to A. D. Woozley’s “Socrates on Disobeying

the Law” in The Philosophy of Socrates, ed. G. Vlastos, Doubleday (1971).

103

104 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

A second explanation for the (supposed) inconsistency is that

Plato was conscientiously reporting an inconsistency he had found

in Socrates’ philosophical position. But this is of little help because

it merely trades the unlikely view that Plato was unable to detect

the contradiction, for the equally unlikely view that Socrates lacked

the intellectual ability to detect it.

Then it might be suggested that Socrates did not hold these

inconsistent positions simultaneously; he changed his mind be¬

tween his trial and the appointed time for his execution. But this,

too, seems very unlikely, both because the time involved was quite

short (about a month), and because Socrates insists in the Crito

that the position he is there taking on the question of disobeying

the law is the only one consistent with his earlier philosophical

teaching.

Finally, Grote considered the two dialogues to be really incon¬

sistent but attributed this inconsistency to deliberate falsification

on Plato’s part. He maintained that the view expressed by Socrates

in the Crito (that one ought never disobey the law) was neither

genuinely Socratic nor believed by Plato to be genuinely Socratic.

It was included by Plato to protect the memory of his teacher as

a patriot from those who, according to Xenephon (and justly,

according to Grote), accused Socrates at his trial of “inducing

his associates to disregard the established laws. . . .” Thus, the

inconsistency is due to the fact that the Apology is genuinely

Socratic and the Crito is not.

There are, however, at least three difficulties which make it seem

improbable that this view is correct. First, there is no evidence

(aside from its inconsistency with the Apology) to support the view

that the Crito is not genuinely Socratic. Second, it is difficult to

believe that Plato could have thought he would get away with the

publication of such a falsification, when Socrates had been dead

only a relatively few years. And finally, we have no reason to be¬

lieve that Plato was given to such intellectual dishonesty; quite

the contrary, the Parmenides seems to be evidence of Plato’s ex¬

traordinary philosophical candor.

In sum, whereas none of the above explanations for the incon¬

sistency between the Apology and the Crito can be conclusively

refuted, the accuracy of each seems quite implausible. So let us

see if a more plausible account can be given by those who try to

show that there really is no inconsistency to be explained.

II

It might be suggested first that the doctrine in the Crito is

merely that one ought not to disobey the law, leaving one the op¬

tion to disobey particular commands or judicial decisions which,

1973]

Wenz — Socrates on Civil Disobedience

105

one feels, do not accurately reflect the law. Thus, Socrates can dis¬

obey the command of the Thirty Commissioners to get Leon and

the judicial decision to cease philosophizing, because these are, in

Socrates’ opinion, inaccurate reflections of the law. But this will

not do. Socrates mentions in the Crito no such distinction between

the law and a judicial or executive interpretation of it. Instead,

he makes it quite clear that, since the rule of law would be de¬

stroyed were individuals to substitute their own opinions for those

of the court, the duty to obey the law entails the duty to obey the

judgment of a court (50b and 51b-c) . And this appears incon¬

sistent with the passage in which Socrates declares his unwilling¬

ness to abide by a judicial decision ordering him to cease philoso¬

phizing. So the appearance of inconsistency remains.

One might, of course, adopt a natural law position and say that

according to Socrates, no statute, executive command or judicial

decision which prescribes what is immoral is to be considered a

law (or to have the force of law). Thus, Socrates can disobey the

command of the Thirty Commissioners to get Leon and the judicial

decision to cease philosophizing without thereby disobeying the

law because each of these orders, in Socrates’ opinion, prescribes

an immoral action, the former because it involves the execution

of an innocent man, the latter because it constitutes disobedience

to the gods (and we know from the Euthyphro that Socrates

thought the gods commanded only what was morally obligatory).

This approach contains an important element of truth. For it

certainly does seem to be the case that Socrates’ reason for refus¬

ing both to get Leon from Salamis and to cease philosophizing was

that he considered compliance in these cases to be immoral. But

that does not mean that Socrates was a natural law philosopher

who felt that immoral statutes, executive commands and judicial

decisions lack the force of law. Indeed, in neither the Apology nor

the Crito does Socrates distinguish between moral and immoral

statutes, executive commands and judicial decisions in such a way

as to state or imply that the former have, whereas the latter lack

the force of law. In the Crito , in fact, just the contrary is the case.

Socrates there argues (51d-e) that living in a state constitutes an

implicit agreement to abide by its laws. Thus, when political power

changes hands in the state, the individual should, according to

Socrates, find out how the new regime does things and then either

resolve to obey its laws, or move to a different state. In this way,

Socrates acknowledges at least the legal legitimacy of even an im¬

moral, de facto government, and in so doing acknowledges that so

long as the Thirty are in power, their way of doing things (rule

by decree) is the law and is legally binding, just as the Democ-

106 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

racy’s way of doing things is the law and is legally binding when

it is in power.

Socrates does not, then, question the legality of the order given

to him by the Thirty to get Leon from Salamis, nor would he ques¬

tion the legality of a court order enjoining his further philosophi¬

cal activity. The point is, rather, that because of the immorality

involved in compliance, Socrates refused to obey these orders

despite the fact that he considered them to be legal. This is what

makes him a civil disobedient.

Finally, Professor A. D. Woozley in a recent article, “Socrates

on Disobeying the Law,” claims that the doctrine in the Crito that

“a man must either do whatever his city orders him to do or must

persuade her where the rightness of the matter lies (51c)” enables

one to reconcile Socrates’ injunction in the Crito against disobey¬

ing the law with his declaration in the Apology that he is prepared

to disobey a judgment banning him from further philosophizing in

Athens. Woozley argues that the doctrine (contained in the Crito)

that one must either persuade the state or obey its orders, pre¬

supposes that one always has the right at least to attempt moral

persuasion of the state. Thus, it is perfectly consistent with the

Crito for Socrates, in the Apology, to refuse a court order to cease

philosophizing, since such an order is equivalent to an order to

cease his attempted moral persuasion of the state and Socrates’

position in the Crito (on Woozley’s reading of 51c) is that men

always have the right to such attempts at moral persuasion. As

Professor Woozley puts it,

The disobedience to a lawful command which he (Socrates)

is not prepared to countenance in the Crito is of a kind which

would do violence and injury to the law. . . . And all dis¬

obedience to lawful command is of this kind, with the single

exception of attempting to convince the state that it is wrong

in the law or command concerned.

But this permitted exception to the rule of obedience is precisely

what he had proposed to follow in the Apology when he declared

his willingness to disobey a judicial order to cease philosophizing.

Thus, Socrates’ willingness to disobey such an order is perfectly

consistent with everything in the Crito.

There are, however, some grave difficulties with this approach.

First, consider the passage at 51c in which Socrates says that “a

man must either do whatever his city orders him to do or must

persuade her where the rightness of the matter lies.” It is not at

all clear that this passage implies Socrates’ adherence to the prin¬

ciple that one always has the right to attempt moral persuasion

of the state. For the passage is part of an analogical argument

1973]

Wenz — Socrates on Civil Disobedience

107

(50e-51c) in which the laws are compared to one’s parents.

Socrates, speaking for the laws, argues that just as one ought to

show respect for one’s parents, even more so ought one to show

respect for one’s country and its laws, because “compared with

your mother and father and all the rest of your ancestors, your

country is far more precious, more venerable (and) more sacred”

(51b). The point is that an individual should take no liberty with

his country that he would not take with his own parents. But

earlier (51a) one of the liberties denied the individual relating

to his parents was that of answering back when scolded. This

would seem to mean that an individual can speak only if his par¬

ents give him permission. So if he disagrees with one of his

parents’ commands, it is only if his parents give him permission

to speak and if, upon speaking, he convinces them that he is right,

that he may do as he had originally thought correct.

Continuing the analogy, then, between one’s country and one’s

parents, when the laws say, “If you cannot persuade your country,

you must do whatever it orders,” they are properly understood as

meaning if your country grants you the right to speak, and if,

upon speaking, you persuade your country, then, and only then,

may you do as you had originally thought correct. But if your

country refuses to let you answer back, then it is not your right

to do so, any more than it would be your right to answer back your

parents when scolded. So the passage at 51c will not support Prof.

Woozley’s interpretation; it is not evidence of Socrates’ belief that

an individual always has a right to attempt moral persuasion of

his country on any and all matters.

Indeed, Socrates next quotes the Athenian laws as saying, “We

give (the individual) the choice of either persuading us or doing

what we say.” (52a) The option, then, of attempting moral persua¬

sion of the state is given by the laws and presumably, therefore,

did not exist before it was so given. Thus, it is not, according to

Socrates in the Crito, an “inalienable right.”

Nor is it an inalienable right according to Socrates in the Apol¬

ogy . For he there notes (37b) that some cities require several days

for the completion of a trial for a capital offense, and predicts that

if such were the practice in Athens, his chances of acquittal would

be considerably enhanced. But, he says, he will nevertheless abide

by the Athenian rule which leaves him only one day to persuade

the jury of his innocence, thus, again acknowledging, this time in

the Apology, that the state has a right to limit and control the

extent of his attempts to persuade it.

So one cannot account for Socrates’ refusal to obey a court order

enjoining his philosophical activities by any such appeal as Prof.

Woozley’s to a Socratic belief that an individual always has the

108 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

right to attempt moral persuasion of the state on any and all mat¬

ters. For there appears to be no such Socratic belief in either the

Crito or the Apology.

Another difficulty with Woozley’s approach is that even if one

were to permit his reading of 51c, the Crito and the Apology would

still be unreconciled on the issue of disobeying the law. This is the

case because Socrates' disobedience to the command of the Thirty

Commissioners to get Leon still cannot be justified on the grounds

that “a man must do whatever his city orders him to do or must

persuade her where the rightness of the matter lies." In the first

place,, Socrates states, “When we came out of the Round Chamber,

the other four went off to Salamis and arrested Leon, and I went

home" (32d). There is no indication in the passage that he even

so much as tried to convince the Thirty Commissioners that it was

wrong to execute Leon of Salamis. Instead, he simply listened to

what they had to say and then went home. In the second place, and

this is the really important point, even if he had tried to convince

them where the rightness of the matter lay, the fact remains that

he did not succeed in so convincing them, and should, therefore,

according to the rule stated in the Crito, have obeyed the order.

Thus, unless we are to suppose that Socrates considered the order

to be illegal (which, I have already argued, is quite unlikely), we

must conclude that his disobedience to the command of the Thirty

still appears inconsistent with the Crito.

Ill

How, then, can one reconcile the Apology and the Crito on the

question of disobeying the law? I will begin by explicating Socrates'

views on this matter as they appear in the Apology. Then I will

offer an interpretation of the Crito according to which it is com¬

pletely consistent with the views explicated from the Apology.

We have already seen in the Apology that Socrates is not willing

to do evil. It is for this reason, after all, that he disobeys those

laws, executive commands and judicial decisions which require

him to do what he considers to be immoral; it is for this reason

that he refuses to participate in the unjust execution of Leon and

would refuse to cease philosophizing even if so ordered by the

court at his trial. This is the grain of truth in the attribution to

Socrates of the natural law position considered above, namely,

that the Apology represents Socrates as refusing to do evil under

any circumstances, whether that evil be a requirement of law, a

requirement of a legally sanctioned executive tribunal, or a re¬

quirement of a legally constituted court of law.

So Socrates in the Apology is not willing to do evil under any

circumstances. But he is willing to suffer evil ; he is willing to put

1973] Wenz — Socrates on Civil Disobedience 109

the location and duration of both his body and his material posses¬

sions at the disposal of the state. This is evident in his behavior upon

receiving a guilty verdict at his trial. He refuses to cease philoso¬

phizing (for this he considers immoral). Otherwise, however,

despite the fact that he is innocent of the charges for which he

was convicted, he is willing to suffer a great range of personal

inconveniences. He is willing to pay a fine, and will even put all his

material possessions at the disposal of the state. He is willing to

go to prison (he did so for a month). He is willing to accept the

penalty of exile. But he will also accept, and even prefers to accept

the penalty of death. Thus Socrates demonstrates his willingness

to put the location and duration of both his body and his material

possessions at the disposal of the state.

His encounter with the Thirty Commissioners reveals the same

willingness. Socrates tells us that after receiving the order of the

Thirty to go to Salamis and get Leon, he went home thinking it

not unlikely that he would be punished by death for disobeying.

The clear implication is that Socrates would not have attempted

to evade such a punishment; he would have allowed the state

to dispose of him, even though it was under the immoral rule of

the Thirty. Thus, again, Socrates demonstrates in the Apology his

willingness under any circumstances to put the location and

duration of his body at the disposal of the state (material posses¬

sions not being in question in this case.)

Socrates’ position in the Apology , then, is this : He will do with

his body only what is moral; he will not use it for such immoral

purposes as going to Salamis to get Leon, and he will not cease

to employ it in the practice of philosophy. But the location and

duration of his bodily existence, as well as that of his material

possessions, he leaves at the disposal of the state. He will allow

the state to dispossess him, imprison him, exile him and put him to

death. And he will allow the state to do these things at any time

they wish, for any reason they wish; Socrates imposes no condi¬

tions on the state, least of all that it should so inconvenience him

only when it has just cause.

If this position sounds strange to our modern ears, it is because

it follows from a belief we no longer share: — Socratic dualism,

the belief that the soul is all-important, the body and material

possessions, unimportant. Thus, Socrates says to his accusers at

his trial that they are not harming him because they are only af¬

fecting him bodily and materially and these are unimportant. But,

he says, they are harming themselves because their intentions are

evil, unjust, and this evil harms their souls, which is important.

Other statements of this dualism can be found throughout the

Socratic dialogues, most notably in the Gorgias (477a) and the

110 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Crito (54b). But the important point here is that it appears in

the Apology and is manifested in Socrates’ combined zmwillingness

to do evil, since that would harm is soul, and willingness to suffer

evil, to put the location and duration of his body and his material

possessions at the disposal of the state.

I shall now attempt to show that the text of the Crito is such

as to render quite plausible an interpretation of that dialogue

according to which it is perfectly consistent with the Socratic

position just uncovered in the Apology. Toward this end, the first

thing I would note is that the form of reasoning in the Crito is

very much like that employed in a judicial opinion of a modern

appellate court. In such a judicial opinion one usually finds that

there is: 1) A specific decision to be made, 2) that that decision

is made by appeal to a general principle from which it (the specific

decision) follows deductively, and 3) that the general principle

is itself supported by a) appeals to the consequences of certain

acts, in conjunction with b) some (often inexplicit) ethical max¬

ims which entail that it is immoral that such consequences be so

brought about. Consider, for example, a case (Case I) arising

from a girl’s broken engagement. She broke the engagement due

to the persuasion of her father. Now her former fiance wishes to

sue the father for damages due to the alienation of affection (on

the part of the girl towards her former fiance) resulting from

the father’s persuasion of the girl to break the engagement. The

question before the court is 1) whether or not such a suit is to be

allowed. The decision is made that such a suit will not be allowed,

this specific decision being based on 2) the explicitly stated general

rule that the court should not interfere with the parent-child re¬

lationship. 3) The general rule is itself supported by a) the be¬

lief that such judicial interference would have the consequence

of weakening the parent-child relationship and that the per¬

formance of certain filial duties would thereby be put in general

jeopardy, in conjunction with b) the ethical maxim that such filial

duties ought to be performed.

Suppose now that a new case is before the court. A bank was

robbed and the man performing the robbery did so with the

advice and encouragement of his father. The question before the

court is whether or not this father can be prosecuted as an ac-

cesory for his part in helping to plan the robbery. The defend¬

ant in this case (Case II) claims the decision in our prior case

(Case I) as a precedent; he claims that the general rule in that

case governs the present case as well. That general rule was

explicitly stated to be that the court should not interfere with

the parent-child relationship. The defendant argues that prose¬

cution of the father in the present case would constitute such in-

1973]

Wenz — Socrates on Civil Disobedience

111

terference and is, therefore, precluded by the dicision in Case I.

The prosecution would, of course, argue that the meaning of

the general rule established by the judge in Case I cannot be de¬

termined solely by reference to the literal meaning of the words

used by the judge in stating that rule. To determine what rule the

prior judge himself meant to establish in his opinion, we must

look also to 1) the particular decision made in the case in which

the rule was originally stated, and 3) the justification there

given for the rule. This is so because the regulation of human con¬

duct by general rules is (as Aristotle noted) a very difficult matter.

It is very difficult to state a general rule, whether as parent, mor¬

alist, judge or legislator, which covers all of those cases you wish

to be covered and none of those you wish to leave uncovered. And

this is why it is common practice when attempting to determine

exactly what a given general rule was originally meant by its author

to entail, to consult, besides the literal wording in which the rule is

stated, 1) the context in which that rule is stated and 3) the justi¬

fication given in that context for the rule.

In the case of our legal example, then, the prosecutor would

note that the justification offered by the judge for the general

rule in Case I was that the performance of certain filial duties

that ought to be performed would be jeopardized by judicial inter¬

ference with the parent-child relationship. Thus, it would seem

that the judge in Case I originally meant the rule against such

judicial interference to apply only to those cases which involve

(or at least might reasonably be construed to involve) the jeopardy

of such filial duties. But helping to plan a robbery is not a filial

duty at all. So, the prosecutor could argue (and rightly as I think) ,

the general rule laid down in Case I cannot be used as a precedent

in the present case, despite its literal wording, because the justifi¬

cation for that rule makes it clear that the judge in Case I origi¬

nally meant it to apply only to cases which could at least be reason¬

ably construed to involve filial duties.

Returning now to the Crito , notice that the form of reasoning

is very similar to that employed in Case I. As in a modern legal

case, 1) there is a specific decision to be made, namely, whether or

not Socrates ought to escape. 2) That decision is made by appeal

to a general principle from which it (the particular decision) fol¬

lows deductively, the general principle in the Socratic case being

stated as the rule that one ought never to disobey the law. And

3) the general principle is itself supported by appeals to the conse¬

quences of certain acts (acts of disobedience destroy the rule of

law in a society), in conjunction with some ethical principles

which entail that it is immoral for such consequences to be so

brought about. In the Socratic case the ethical principles are a)

112 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

“Never return evil for evil” which, given the view that destroying

something is doing it evil, entails that one ought not disobey (and

thereby destroy) even unjust laws, and b) “You have a duty and

an obligation to uphold the laws which raised you (and to which

you have since given your tacit consent by remaining in the state) ,

from which it again follows that disobedience, because it destroys

the laws, is morally wrong.

This form of reasoning is, I submit, itself sufficient to warrant

at least the suspicion that the Socratic case is similar to Case 1

also in the respect that a better understanding of 2, the general

rule, is forthcoming upon a consideration of 1, the specific decision

deduced from that rule, and 3, the justification given for the rule.

What is more, however, this suspicion is enhanced by the fact

that in the Crito, just as in Case I, the subject matter is such that

the general rule is a rule for the regulation of human conduct.

One can reasonably suppose that Socrates might, at least, experi¬

ence the same difficulties as others (moralists, parents, judges and

legislators) who attempt to regulate human conduct by general

rules. He, like they, might find it difficult to frame rules which,

when construed according to the literal meanings of the words

used, cover all those cases he wished to cover and none of those he

wished to leave uncovered. It is thus again plausible, at least, that

we can here, as in Case I, obtain a more accurate understanding

of the general rule Socrates meant to endorse in the Crito if we

consider, in addition to the literal meaning of Socrates’ statement

of the rule, those other elements in the total reasoning process

which were so helpful in discovering the meaning of the general

rule in Case I.

I shall, then, offer an interpretation of the general rule in the

Crito . Consider the justification given in the Crito for the general

rule that one ought never disobey the law. The claim is a) that

such disobedience destroys the law (thus doing it evil) and one

ought never do evil even in return for evil, because doing so is

wrong and one should never do what is wrong (49a-50a). It is

then argued b) that the laws are analogous to one’s parents and, like

one’s parents, deserve special respect. They are, therefore, the

last things that one ought to disobey and thereby destroy. So

again, because disobedience destroys the laivs, one ought not to

disobey. Lastly, it is claimed c) that one ought to keep one’s agree¬

ments and that remaining in a state upon attaining maturity

constitutes an implicit agreement to uphold the laws of that state

(51d-52d). So yet again, because disobedience destroys the state’s

laws, one ought not to disobey.

Thus, just as the claim central to the judge’s justification

for the general rule stated in Case I was, “Legal interference with

1973]

Wenz — Socrates on Civil Disobedience

113

the parent-child relationship will jeopardize the performance of

filial duties,” so in the Crito the claim central to Socrates’ justifi¬

cation for his general rule is, “Disobedience destroys the laws.”

Accordingly, it seems at least plausible that just as the judge in

Case I was taken to really mean , “The court ought not to interfere

with the parent-child relationship, when such interference might

reasonably be construed to jeopardize the performance of filial

duties,” so in the Crito Socrates can be taken to really mean, “One

ought never to disobey the law, when such disobedience might

reasonably be construed to destroy the laws of the state.”

But if Socrates really meant in the Crito to disallow only such

disobedience as might reasonably be construed to destroy the laws

of the state, then it is not at all inconsistent with the Apology.

For in the Apology Socrates leaves the location and duration of

both his body and his material possessions at the disposal of the

state. And clearly no individual threatens the laws with destruction

who places the location and duration of his body and his material

possessions at the disposal of the state. A state surely does not re¬

quire, in addition, dominion over the individual’s soul. For order

in the state, even if it is modelled upon order in the soul, as Plato

would have it, is nevertheless order among human activities in the

material world of sense perception. Therefore, a sufficient condition

for a state to be capable of maintaining its order (be that order

good or evil) is that it be able to order the activities of those indi¬

viduals who come in contact with it. And this sufficient condition

can be met in either of two ways. Either the individual can act

in conformity with the state imposed order, or his contact with

the state can be ended. Socrates chooses the latter course. He

will not act in conformity with the state-imposed order, but he will

leave the location and duration of both his body and his material

possessions at the disposal of the state, allowing the state to con¬

trol the extent of his contact with it, and giving the state, thereby,

the ability to withstand whatever disruptive effects his disobedience

might have.

So while Socrates, and others engaging in a similar practice

of civil disobedience, may disturb the state temporarily, they pose

no danger of destruction to that state or its laws. Thus, Socrates’

disobedience in the Apology is not the type disallowed in the Crito ;

it is not such disobedience as might reasonably be construed

as destructive of the laws of the state because it reserves to the

state control over the location and duration of both body and ma¬

terial possessions, and this is all the state needs to preserve its

laws. The Crito and the Apology are, then, perfectly consistent

on the question of disobeying the law.

114 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

CONCLUSION

Socrates’ position was, then, that of a moral man in an immoral

society which required him, occasionally, to participate in its im¬

morality; it required him, for example, to aid in the execution of

Leon, and might have required him to cease philosophizing. There

were, in addition, the following complications: 1) Socrates thought

it immoral to destroy anything, including the immoral laws of

the state in which he lived. 2) His were times of political unrest

when the state, and its laws, were all too easily overturned and

destroyed. And 3) he had a very exaggerated view of the dis¬

ruptive effects of civil disobedience.

What could Socrates do under these circumstances to preserve

his moral integrity? There seem to be only two alternatives, both

of which were recognized by Socrates. One is that he could move

to a different state, a state which is moral or which, if it is im¬

moral, will at least not require of him participation in its immoral¬

ity. Socrates clearly considered this alternative morally acceptable

(51d and 37d). But he rejected it on other grounds, grounds of

patriotism and sentiment.

What, then was the moral alternative available to him if he

did not wish to leave his country? “Political civil disobedience,”

as that term is currently defined,4 will not do because such diso¬

bedience is purposefully disruptive. It constitutes an appeal, through

disobedience, to the conscience of other members of the society

in an attempt to effect the elimination of a given immoral law

or policy. But in Socrates’ case the state, unlike our own but simi¬

lar to some of the so-called “developing nations,” was in a very

precarious political position and could easily have been destroyed.

And Socrates considered it immoral to destroy the state. He could

not, then, risk the purposefully disruptive tactic known as politi¬

cal civil disobedience because under such conditions of instability,

there may be little, if any difference between disruption and sedi¬

tion, each being as likely as the other to destroy the state. So

political civil disobedience was not Socrates’ other moral alterna¬

tive, and he never mentions it.

This is, by the way, one respect in which Socrates’ practice

of civil disobedience differs from that of Martin Luther King.

For whereas both King and Socrates were civil disobedients and

gadflies on the state, King employed civil disobedience as a weapon

in support of his activities as a gadfly, and Socrates did not. In¬

stead, Socrates attempted to effect changes in the state’s moral

behavior solely by verbally confronting its leaders and its citizens

4 See The Morality of Civil Disobedience by Robert T. Hall, Harper Torchbooks

(1971).

1973]

Wenz — Socrates on Civil Disobedience

115

with their own immorality. He did not, like King, use civil dis¬

obedience as a method for promoting such confrontation. And the

reason is clear. Socrates considered disobedience to be extremely

disruptive, as he makes clear in the Crito. He considered it im¬

moral to destroy the state, and he, unlike King, lived in a state

that was on the brink of destruction. Thus, while Socrates was a

political gadfly, which he did not consider to be overly disruptive,

he was not, like Martin Luther King, a political civil disobedient.

Socrates was, then what is currently defined as a “moral civil

disobedient.” To preserve his moral integrity while remaining in

the state, he had to disobey when the state ordered him to partici¬

pate in its immorality. But, realizing the potential for disruption

of any civil disobedience, he did so in the least disruptive manner

possible so as to insure avoiding the immorality of destroying the

state. Thus, so far as we know at least, Socrates did not encourage

imitation on part of the others who, like him, were ordered by

the Thirty to get Leon from Salamis. He did not ask them to

join him in his disobedience of the order because his disobedience

was not political; it was not designed to effect a change in any

law or policy. Socrates’ only political tactic was verbal confronta¬

tion; his civil disobedience was purely moral, designed only to

maintain his own moral integrity.

But Socrates thought that even such disobedience might inspire

imitation and thereby cause sufficient disruption to destroy the

state. To insure that this would not occur, to insure that whatever

imitators he might attract would not cause sufficient disruption

to destroy the state, Socrates, consistent with his belief that the

soul alone is of worth, put the location and duration of both his

body and material possessions at the disposal of the state, insuring

thereby that neither he, nor any others who imitated his prac¬

tice, would jeopardize the state’s existence by their disobedience.

So far as an example for others is concerned, however, it is one

thing to illustrate one’s commitment to putting one’s body and

material possessions at the disposal of the state by paying a small

fine, and quite another to do so by accepting the death penalty.

The latter illustrates the commitment much more vividly and

convincingly. So Socrates, when speaking of his past disobedience

(to the command of the Thirty) made clear his willingness at that

time to accept the death penalty for that disobedience. And then,

to prove the sincerity of his commitment, he decided to sacrifice

whatever philosophizing he might be able to pursue in other coun¬

tries, and goaded the court at his trial into giving him the death

penalty, and then refused the advice of his friends to escape that

penalty.

116 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Thus, his every action seems to indicate that Socrates was a

civil disobedient living in times of political unrest who considered

it immoral to destroy the state in which he lived, who had an

exaggerated view of the disruptive effects of civil disobedience,

and who therefore took every precaution to minimize the disrup¬

tion his disobedience might cause. He never encouraged others to

disobey the law ; he never used disobedience as a tactic for effecting

change in the state’s laws or policies; he had a willingness to

put the location and duration of his body and his material posses¬

sions at the disposal of the state; and he chose the most dramatic

method possible to illustrate that willingness for any who might

want to imitate his disobedience. He was, in short, a perfectly

consistent moral (as opposed to political) civil disobedient.

ACKNOWLEDGMENT

This paper owes much to the helpful advice of my colleagues

in the Department of Philosophy at the University of Wisconsin-

Stevens Point, and particularly of J. Baird Callicott. Its remaining

deficiencies are, of course, my own.

DISTRIBUTION OF PHOSPHORUS, SILICA, CHLOROPHYLL a,

AND CONDUCTIVITY IN LAKE MICHIGAN AND

GREEN BAY1

D. C. Rousar and A. M. Beeton

Center for Great Lakes Studies

University of Wisconsin —

Mihvaukee

ABSTRACT

In July 1971 seventeen stations in central Lake Michigan and

twenty-one stations in southern Green Bay were sampled at vari¬

ous depths to determine the spatial distribution of total phos¬

phorus, silica, chlorophyll a, and conductivity. Water from Lake

Michigan averaged 10.4 /xg P/liter as total phosphorus, 1.0 mg

Si02/liter as soluble reactive silica, 3.0 /xg chlorophyll a/liter as

total chlorophyll, and 256.1 /xmhos/cm as specific conductance. Cor¬

responding values from Green Bay were 87.8 /xg P/litey, 0.9 mg

Si02/liter, 32.9 /xg chlorophyll a/liter, and 257.7 /xmhos/cm. Surface

concentrations of phosphorus, chlorophyll, and conductivity varied

little in the lake but decreased markedly along a 10 mile transect

extending from the mouth of the Fox River to a station in the bay.

Silica was uniformly distributed in the bay but showed some

inshore-offshore differences in the lake. Vertical differences in the

parameters tested were minor at most of the stations in Green Bay.

Conductivity and phosphorus were relatively constant with depth in

Lake Michigan. However, silica increased with depth at all stations

except one, and chlorophyll was highest at the 30 m depth of 13

of 17 stations. Comparisons of the results with published data are

discussed.

INTRODUCTION

Long-term chemical changes have occurred in Lake Michigan.

Beeton (1969) reported substantial increases in total dissolved

solids, sulfate, and chloride during the past 90 years (and pro¬

vided information on the growth of human population in the drain¬

age basin). Published data on increases in nutrients, or eutrophi¬

cation, for a similar time period are generally inadequate. No

trend in phosphorus was found by Beeton due to insufficient and

1 Contribution No. 71, Center for Great Lakes Studies.

117

118 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

conflicting data. Levels of various forms of nitrogen representing

the entire lake are poorly known, but from the period 1928 to

1966 inorganic nitrogen (nitrate) decreased, while organic nitro¬

gen (albuminoid ammonia) increased at a Milwaukee water purifi¬

cation plant intake (Beeton 1969). Schelske and Stoermer (1971),

using data from municipal water intakes near Chicago, cited a

decrease of at least 4 mg Si02/liter from 1926 to 1970. They

have hypothesized that accelerated eutrophication is causing de¬

pletion of silica in surface waters of Lake Michigan during sum¬

mer stagnation due to diatom production.

From the standpoint of spatial distribution of chemicals in Lake

Michigan and Green Bay, surveys by the U. S. Public Health Serv¬

ice in 1962 and 1963 demonstrated major horizontal (inshore-

offshore) differences in total soluble phosphate, silica, and con¬

ductivity (FWPCA 1968). Depletion of silica in epilimnetic waters

during thermal stratification has been reported (Beeton and Mof¬

fett 1964, Schelske and Callender 1970).

The objectives of this study were twofold. One was the spatial

distribution of the parameters tested and their interrelationships.

The other was the comparison of results with previous data to

see if nutrient enrichment had occurred.

MATERIAL AND METHODS

All sampling was done onboard the R/V “NEESKAY” of the

Center for Great Lakes Studies. Station numbers, locations, depths,

and dates of sampling are listed in Table 1. Ports and coastal

geographic features mentioned in the text are depicted on Figs.

1 and 2. All stations were sampled in July 1971, but only the

numbered stations in Lake Michigan were sampled by the U. S.

Bureau of Commercial Fisheries in 1954 (Beeton and Moffet 1964).

Water samples were collected with either a plastic 9-liter Van

Dorn sampler or by Teflon-lined 2-liter Nansen bottles. Tempera¬

tures were recorded by reversing thermometer or manually from

a bucket. Unfiltered water was put into 250 ml pyrex bottles

which were brought back to Milwaukee for conductivity and total

P analysis. Samples for pigment assay were prefiltered through

a 250 pm nylon mesh into polyethylene bottles and stored in the

dark no longer than two hours before being refiltered. Water to

be analyzed for silica was put in polyethylene bottles and filtered

through 0.45 pm HA Millipore filters of 47 mm diameter within

a day after collection. This filtered water was then stored in

polyethylene bottles.

Total phosphorus was determined by the method of Gales et al.

(1966) with the following modifications. A 10 or 35 ml aliquot

1973] Rousar and Beeton — Phosphorus Distribution 119

TABLE 1. STATION LIST WITH LOCATIONS, DEPTHS,

AND DATES SAMPLED

of thoroughly shaken sample was pipetted into a 50 ml pyrex

flask. Then 10 ml of a 0.11 M (0.3g. per sample) aqueous solu¬

tion of potassium peroxydisulfate, K2 S2 08, was added followed

by one ml of sulfuric acid solution. The flasks were then covered

by 30 ml pyrex beakers and autoclaved at 15 PSI for 30 minutes.

After cooling, all samples, standards, and blanks were diluted to

50.0 ml with distilled water. Two ml of molybdate solution was

120 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 1. Lake Michigan stations and geographic features mentioned in

the text. Numbered stations sampled by Beeton and Moffett (1964). All

numbered and x-numbered stations sampled in July 1971.

added, mixed, and followed by two drops of a 4% solution of

SnCl2 in glycerine. The absorbance of the resulting heteropoly

blue was measured by a Hitachi Perkin-Elmer 139 spectropho¬

tometer set at 720 nm and using 10 cm cuvets. This measurement

was made no later than 20 minutes after addition of the color-

1973] Rousar and Beeton — Phosphorus Distribution

121

imetric reagents. Instrument values were then converted to con¬

centrations reading from a standard curve with 5, 10, 20, and 40

fxg P/liter standards. The standard deviation relative to the arith¬

metic mean (coefficient of variation) was 3.6% at 9 jmg P/liter

concentration.

Soluble reactive silica was determined by the heteropoly blue

method (A.P.H.A. 1965) with the following modification. The

122 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

HC1 solution was made by diluting 210 ml concentrated HC1 with

130 ml distilled water. Other reagent solutions were not modified.

To 50 ml of sample, 2.5 ml of molybdate solution, 1.0 ml diluted

HC1, 1.0 ml oxalic acid solution, and 2.0 ml reducing agent were

added sequentially at 5 minute intervals. A Fisher Electrophotom¬

eter II, with a red 650 nm filter and 2.5 cm diameter pyrex

cells, was used to measure absorbance of the reduced molybdosili-

cate complex no sooner than 30 minutes after addition of the

reductant. The coefficient of variation was 1.3% at 1.3 mg Si02/

liter concentration.

Chlorophyll a and phaeo-pigments were measured by the methods

of Strickland and Parsons (1968). A Model III G. K. Turner

fluorometer equipped with a high sensitivity phototube, Corning

CS. 5-60 filter for the excitation light and Corning CS. 2-64 filter

for the emitted light was used. The fluorometric procedure was

calibrated spectro-photometrically using the equations:

fig chlorophyll a/ liter = - - - -

* , . , /r, 26.7 [1.7 (665a) -6650] x v

ug phaeo-pigments/hter — - - - ^ - — -

V x 1

where 6650 is the extinction at 665 nm before acidification, 665a

the extinction at 665 nm after acidification, v the volume of 90%

acetone used for extraction (ml), V the volume of water filtered

(liters), and 1 the path length of the cuvet (cm). The derived

factor for door 3 was then substituted into the equation :

ixg chlorophyll a/liter = door factor x fluorometer reading. This

value was called total chlorophyll a. “Active” chlorophyll a was

calculated from

jug “active” chlorophyll a/liter = F

D'

- 1

(Rb-Ra) where F

D

is the factor for door 3, RB and RA are the relative fluorescences

R

before and after acidification, respectively, and r is the ratio —

i\A

Concentrations of phaeo-pigments were determined by subtracting

“active” chlorophyll a from the total chlorophyll a.

Aliquots (10-250 ml) of prefiltered water were filtered through

two 2.4 cm diameter Reeve Angel glass filters (834 AH grade)

at one-third atmospheric pressure, extracted with 90% acetone in

a tissue grinder, diluted to 10 ml, and centrifuged. Five ml of

supernatant were then transferred to pyrex tubes and measured

fluorometrically. Dilutions with 90% acetone were made as re¬

quired. Two drops of 5% HC1 were next added, the solution mixed

by inverting, and the fluorescence after acidification recorded. The

coefficient of variation was 5.6% at 5.9 jig chlorophyll aj liter.

1973] Rousar and Beeton — Phosphorus Distribution 123

A Leeds and Northrup electrolytic conductivity bridge (Cat. No.

4959) was used to measure specific conductance of unfiltered wa¬

ter stored in glass stoppered pyrex bottles. Samples were brought

to 25 C before testing.

RESULTS AND DISCUSSION

Lake Michigan

Total Phosphorus

Total phosphorus ranged from 4.5 to 23.0 p g P/liter, with an

average of 10.4 (see Center for Great Lakes Studies Special Re¬

port No. 13 for complete tabulation of data). Assuming that a

difference of 10% or more between values from the surface (2m)

and maximum depth sampled (60-90 m except stations 32 and x3)

is significant (i.e., greater than twice the standard deviation), ten

stations showed an increase, four stations no difference, and

three stations a decrease in total phosphorus with depth. All sta¬

tions along the Milwaukee-Grand Haven transect showed an in¬

crease with depth, and the 60 and 70 m depth of station 13 had

the highest concentrations of total P of all Lake Michigan samples.

These relatively high values might reflect nutrient enrichment from

nearby urbanized drainage basins. Only station 32 along the Frank¬

fort to Sturgeon Bay transect showed increase in total P with

depth.

Total phosphorus values at depths of 2 m and 15 m were aver¬

aged to approximate epilimnetic concentrations (Fig. 3), except

for station x3 where the metalimnion was less than 1 m from the

surface. An epilimnetic average for all stations was 9.9 pg P/liter.

The stations on the northern transect had slightly higher average

values than those on the southern transect, but the reason for this

is not clear.

Levels of total phosphorus found by Beeton and Moffett (1964)

show greater variation but little difference from those reported

here when data from the same stations at approximately similar

dates are compared. They analyzed unfiltered water samples for

total phosphorus, using a modification of Harvey (1948) which

included mineralization by adding sulfuric acid and autoclaving

at 25 PSI for seven hours. Their average value for Lake Michigan

exclusive of the extreme southern end was 13 pg P/liter (Beeton

1969). The U. S. Public Health Service collected water from vari¬

ous depths at seven stations located in a 20 mile wide strip from

Sheboygan, Wisconsin to Little Sable Point, Michigan, as part of

a more extensive lake survey in 1962-1963. Risley and Fuller

(1965) and Verber (1966) discussed various aspects of the

USPHS study and reported an average of 0.02 mg PCb/liter (6.5

124 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 3. Approximate epilimnetic concentrations of total

phosphorus in yg P/liter obtained by averaging values from

depths of 2 and 15 m for each station except station x3 where

only the surface value is given.

1973] Rousar and Beeton — Phosphorus Distribution 125

ng P/liter) as total soluble phosphate for the above strip of lake.

However, average values of 0.0 1 and 0.06 mg P04/liter (13.0 and

19.6 fxg P/liter) were found in the southern end of the lake. Since

the water samples were filtered prior to mineralization and color

development, the quantity of phosphorus determined was called

total soluble phosphate. This fraction of total phosphorus does not

include phosphorus associated with particulate matter removed by

filtration. The minimum detectable concentration of their analysis

was 0.01 mg P04/liter (FWPCA 1968), which corresponds to 3.3

fxg P/liter.

Holland (1969) presented data from a transect between Stur¬

geon Bay, Wisconsin and Ludington, Michigan for April to No¬

vember 1965. Total P averaged 7.8 yg P/liter inshore and 6.3 jxg

P/liter offshore for 2, 5, and 10 m depths. Schelske and Stoermer

(1971) state that except in grossly polluted areas the concentra¬

tion of total phosphorus is less than 10 fig P/liter in Lake Michi¬

gan. From May 1970 to October 1971 surface water (4m) samples

from five stations along a Milwaukee, Wisconsin to Ludington,

Michigan transect (two stations are 3 miles from harbor break¬

waters, the other three are one-fourth, one-half and three-fourths

of the distance from the two ports) have been analyzed for macro¬

nutrients (Fee 1971). The average total phosphorus for the in¬

shore station nearest Milwaukee was 15.2 /xg P/liter from 39

samples, and the average for all other stations was 8.4 jxg P/liter

from 156 samples. Unfiltered water was mineralized by boiling

with H2S04 and H202, neutralized, and analyzed colorimetrically

as described by Schmid and Ambiihl (1965). This information is

summarized in Table 2.

Since phosphorus can occur in a wide variety of dissolved and

particulate forms — or fractions — in natural waters, the analysis

for total phosphorus should preclude the difficulty of measuring

and interpreting these variable fractions (Rigler 1964). The use

of filters and their porosities, the completeness of mineralization

(A.P.H.A. 1971), the sensitivity and stability of colorimetric prod¬

ucts, the types of instrumentation, and differences in analytical

skills can produce variances that make comparisons of the above

results problematic. Thus no conclusions can be drawn about gen¬

eral changes in total P from central Lake Michigan during the

past 17 years.

Soluble Reactive Silica

Soluble reactive silica from all depths sampled ranged from 0.4

to 1.9 mg Si02/liter and averaged 1.0 mg Si02,/liter. All Lake Mich¬

igan stations exhibited increasing silica concentrations with depth

probably due to greater assimilation by diatoms in the euphotic

126 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 2. Published total phosphorus values for central Lake Michigan

zone than in deeper water. However, station x3 was homogeneous

throughout most of the water column. The surface value at this

station was markedly higher than those at other stations (Fig. 4)

but of the same order of magnitude as hypolimnetic values. This

observation gives support to the contention that station x3 was in

an area of up welling. The relatively high value of 1.8 mg Si02/liter

at 250 m probably reflects near-bottom hypolimnetic water. The

similarly high values at 60 m and 70 m at station 13 correlate

positively with the total phosphorus concentrations and may indi¬

cate nutrient enrichment producing greater primary production

with subsequent sedimentation and decomposition.

Epilimnetic values at stations nearest shore on the Frankfort-

Sturgeon Bay and Milwaukee-Grand Haven transects were dis¬

tinctly lower than offshore stations on the transects (Fig. 4). This

suggests inshore-offshore differences in depletion of silica due to

diatom production (Holland and Beeton 1972). The epilimnetic

average of all stations was 0.65 mg Si02/liter with a range of

0. 4-1.0 mg Si02/liter.

The silica values given by Beeton and Moffett (1964) were con¬

sistently higher than ours, generally in the 2 to 4 mg Si08/liter

1973] Rousctr and Beeton — Phosphorus Distribution

127

FIGURE 4. Approximate epilimnetic concentrations of soluble

reactive silica in mg Si02/liter obtained by averaging values

from depths of 2 and 15 m for each station except station x3

where only the surface value is given.

128 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

range. However, their determinations were made on unfiltered

samples stored for longer periods of time, and some silica may

have been released from particulate matter. They clearly demon¬

strated an increase in silica with depth during thermal stratifica¬

tion. Ayers et at. (1958) found an average of 2.1 mg Si02l/liter

(range of 1.6 to 3.7 mg Si02/liter) for surface water and an over¬

all average of 2.4 mg Si02/liter (range 1.4 to 4.8 mg Si02/liter)

for 81 samples collected from all depths along three transects:

Montague, Michigan-Milwaukee, Wisconsin; Pentwater, Michigan-

Manitowoc, Wisconsin; and Manistee, Michigan-Sturgeon Bay,

Wisconsin. Samples were collected on 9 and 10 August 1955 by

several vessels. Silica was analyzed as the yellow molybdosilicate

complex according to the ninth edition of Standard Methods

(1946). They reported an analytical reproducibility of 0.2 ppm

silica.

The U. S. Public Health Service survey of 1962-1963 found an

average of 2.9 mg Si02/liter (range of 1.8 to 5.1 mg Si02/liter)

within the 20 mile wide strip mentioned above (Risley and Fuller

1965). They followed the colorimetric heteropoly blue method of

the eleventh edition of Standard Methods to measure silica. The

minimum detectable concentration was 0.02 mg Si02l/liter

(FWPCA 1968).

Although most of their stations were either north or south of

ours, Schelske and Callender (1970) found average surface (2 m)

values of 0.15 (±0.07) mg Si02/liter during July 1969 at 16 sta¬

tions in the southern basin and 0.26 (±0.07) mg Si02/liter during

August 1969 at 18 stations in the northern basin. Samples taken

1 m from the bottom had higher values than those from the sur¬

face of all Lake Michigan stations except three. The average bot¬

tom value was 1.25 mg Si02l/liter for 28 samples. Their HA Milli-

pore filtered samples were frozen until analyzed, and freezing can

reduce levels of soluble reactive silica (Kobayashi 1967; Burton

et al, 1970). As mentioned in the introduction, Stoermer (1971)

referred to dramatic decreases in silica from inshore water around

Chicago over a period of 44 years.

An average of 0.8 mg Si02/liter from 158 surface water samples

collected along a Milwaukee-Ludington transect from May 1970

to October 1971 has been reported (Fee 1971). Twenty of these

samples taken at different dates in July 1970 and July 1971 aver¬

aged 0.5 mg Si02/liter. (See Table 3 for a listing of published

values.)

Considering variations in sample pretreatment, storage, and

analytical methods the evidence for a long-term decrease in silica

is suggestive but not conclusive.

1973] Rousar and Beeton — Phosphorus Distribution 129

TABLE 3. PUBLISHED SILICA VALUES FOR CENTRAL

LAKE MICHIGAN

Chlorophyll a and Phaeo-pigments

Total chlorophyll a for all Lake Michigan samples ranged from

0.4 to 6.9 and averaged 3.0 /xg chlorophyll a/liter, and phaeopig-

ments were usually detectable. Comparing surface samples with

those taken from the greatest depth for each station and making

the same assumption as with total phosphorus, 14 stations had

chlorophyll a concentrations increase with depth, two stations no

difference, and one station decrease. Coupled with the low surface

values relative to bottom values was the concentration of total

chlorophyll a at the 30 m depth. The maximum station value oc¬

curred at the 30 m depth for 13 of 14 stations excluding 12, 32,

and x3; and this value ranged from 3.7 to 6.9 with an average of

5.3 /xg chlorophyll a/liter. This may be a result of settling of

phytoplankton from the epilimnion and metalimnion into the up¬

per part of the hypolimnion where increased density retards sedi¬

mentation.

Epilimnetic total chlorophyll a in Lake Michigan averaged 2.2

/xg chlorophyll a/liter and ranged from 0.8 to 5.3 (Fig. 5). Values

130 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

were somewhat higher along the northern transect than along

the southern.

Stoermer (1967) found that during thermal stratification there

is an evident concentration of phytoplankton at, or just below, the

level of the thermocline; but the vertical distribution of total cell

numbers was strikingly uniform before stratification developed.

From April to November 1965 inshore Lake Michigan surface

waters (2 m) averaged 2.4 yg chlorophyll a/liter and offshore wa¬

ters 1.5 (Holland 1969). These values agree fairly well with ours,

but limited published information on chlorophyll a in Lake Michi¬

gan prevents meaningful comparison.

Specific Conductance

This parameter ranged from 215.9 to 273.3 /mihos/cm at 25 C

and averaged 256.1 for all Lake Michigan samples. Fig. 6 also

shows no apparent spatial distribution patterns. Beeton and Chand¬

ler (1963) reported an average specific conductance of 225.8

/xmhos/cm at 18 C (approximately 262 at 25 C using a conversion

factor of 1.16 based on a .01M KC1 solution) for Lake Michigan.

Risley and Fuller (1965) found an average of 220 /unhos/cm for

water in the previously described 20 mile wide strip and higher

average values south of this area.

Green Bay

Total Phosphorus

Values for all samples ranged from 30.5 to 430 fig P/liter and

averaged 87.8. Excluding stations 1, 3 and 7 the average was

47.7 fig P/liter. Thermal stratification was observed only at station

26, where epilimnetic total phosphorus was greater than that found

in the hypolimnion. Except for station 14, all other stations showed

little relative variation in total phosphorus with depth.

The most striking aspect of Fig. 7 is the steep concentration

gradient from the mouth of the Fox River to station 10. This

gradient probably results from the dilution of grossly polluted

river discharge by water from Green Bay. No discrete pattern of

mixing of bay and river water was seen north of station 10,

perhaps due to strong southerly winds during sampling. The aver¬

age surface total phosphorus from Green Bay was several times

higher than the comparable average value in Lake Michigan.

The U. S. Public Health Service (FWPCA 1968) sampled Green

Bay between 26 June and 17 July 1963 and determined total soluble

phosphate levels to average 0.20 mg P04/liter from the mouth of

the Fox River to about ten miles lakeward and 0.06 mg P04/liter

in an area 10 to 30 miles from the south end of the bay. These

concentrations correspond to 65 and 19.6 fig P/liter, respectively.

1973] Rousar and Beeton — Phosphorus Distribution

131

FIGURE 5. Approximate epilimnetic concentrations of total

chlorophyll a in Mg/ liter obtained by averaging values from

depths of 2 and 15 m for each station except station x3 where

only the surface value is given.

132 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 6. Approximate epilimnetic concentrations of specific

conductance in /mihos/cm at 25 C obtained by averaging values

from depths of 2 and 15 m for each station except station x3

where only the surface value is given.

1973] Rousar and Beeton — Phosphorus Distribution 133

FIGURE 7. Surface concentrations of total phosphorus

in j u-g P/liter from 2 m depth.

In August and October 1966 the Wisconsin Department of Natu¬

ral Resources (Wis. Div. Res. Dev., 1968) analyzed surface and

bottom samples for total phosphorus. Five samples taken 1, 4 and

10 miles from the mouth of the Fox River averaged 90 /xg P/liter

and six samples from 20 to 30 miles lakeward averaged 64 /xg

P/liter. There was no apparent difference between samples taken

134 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

1 m from the bottom and 1 m from the surface, and the

range was 32 to 150 jxg P/liter. Their analysis included evaporat¬

ing unfiltered water to dryness with Mg (N03)2, igniting, dissolv¬

ing, and reacting with ammonium molybdate and stannous chloride.

Between 1965 and 1968 the same laboratory (State of Wisconsin

1968) measured total P from 15 Fox River samples taken from

FIGURE 8. Surface concentrations of soluble reactive silica in

mg’ SiOa/liter from 2 m depth.

1973] Rousar and Beeton — Phosphorus Distribution 135

the State Highway 54 bridge in Green Bay and the De Pere dam

and obtained an average of 230 fig P/liter with a range of 80 to

400 fig P/liter.

Holland (1969) found an average total P of 37.6 fig P/liter for

water taken from 2m at three stations about 20 to 30 miles from

the south end of the bay. Sager et al. (1971) have reported con¬

centration ranges of 0.321-1.08 and 0.044-1.065 mg total phos-

phate-P, /liter for the mouth of the Fox River and lower Green

Bay, respectively.

Thus, the waters of southern Green Bay contain high and vari¬

able amounts of total phosphorus, and they are clearly higher

than those occurring in central Lake Michigan. No change in con¬

centration levels since 1963 can be perceived for similar reasons

given regarding Lake Michigan.

Soluble Reactive Silica

Except for low values at station 1, the distribution of silica in

Green Bay did not suggest any gradients or spatial differences

(Fig. 8). Values ranged from 0.4 to 1.5 and averaged 0.9 mg

Si02/liter. As noted above, wind-driven currents may have tem¬

porarily increased mixing of tributary and lake water.

The U. S. Public Health Service (FWPCA 1968) reported aver¬

ages of 1.7 mg Si02/liter from the mouth of the Fox River to 30

miles from the lower end of the bay in June and July 1963.

Schelske and Callender (1970) in August 1969 detected 0.25 and

1.84 mg Si02,/liter at the surface and bottom of their station 5,

which was located in the area of the bay sampled by us. The

paucity of data disallows any statement on the change of silica

concentration over a period of years.

Chlorophyll a and Phaeo-pigments

Total chlorophyll a ranged from 7.0 to 144 fig/ liter for all sta¬

tions ; and, excluding stations 1, 3 and 7, averaged 19.6 fig chloro¬

phyll a/liter. Fig. 9 shows a pronounced increase in pigment con¬

centrations from station 10 in the bay to station 1 near the mouth

of the Fox River. The data of Fig. 9 indicate much higher levels

of chlorophyll a in Green Bay than in Lake Michigan as well as

higher relative amounts of phaeo-pigments in the former.

Holland (1969) measured chlorophyll a from three stations

situated about 20 to 30 miles from the south end of the bay at a

depth of 2 m from April to November 1965 and obtained an aver¬

age of 10.4 fig/ liter. This compares with an average value of 18.6

fig chlorophyll a/liter from 2 m samples for our stations 15-27.

Sager (personal communication) obtained a range of 1.2-57.4 and

an average of 21.9 fig chlorophyll a/liter from seven stations situ-

136 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 9. Surface concentrations of total chlorophyll a in

jug/liter from 2 m depth.

ated near the mouth of the Fox River and extending to about 40

miles north of the river. Samples were collected monthly during

June, July, and August 1971, from 1 m depth. He also noted a

gradient in the lower bay with high values near the mouth of the

Fox River. Monthly samples at the confluence of the Fox River

with Green Bay from June 1970 to October 1971 have yielded a

1973] Rousar and Reeton — Phosphorus Distribution

137

range of 0.2 to 80 and an average of 24 /mg chlorophyll a/liter. He

prefiltered samples through a 10 /un nylon bolted cloth, then fil¬

tered with a 0.8 Millipore filter, extracted with 90% acetone

in a tissue grinder, and measured spectrophotometrically. His pre¬

filtering may have produced noticeably lower results than ours.

FIGURE 10. Surface concentrations of specific conductance in

/unhos/cm from 2 m depth.

138 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Specific Conductance

This measurement ranged from 205.9 to 376.0 /xmhos,/cm and

averaged 257.7 for all Green Bay samples. Again, there is a clear-

cut increase in values from station 7 to station 1 (Fig. 10). All

other stations have similar concentrations which vary little with

depth. Average specific conductance values from Lake Michigan

and Green Bay excluding bay stations 1, 3, and 7 differ by only

5%. Generally, this physical property appears to be more ecologi¬

cally conservative or biologically independent than the other pa¬

rameters studied.

Sager et al. (1971) found specific conductance values ranging

from 348-525 and 240-555 /unhos/cm at 25 C, from the mouth of

the Fox River and lower Green Bay, respectively.

SUMMARY

A comprehensive characterization of the trophic state of a body

of water should include analysis of the interrelationships between

basin morphometry, seasonal environmental influences, and key

physical, chemical, and biotic properties of the water itself. Sam¬

pling should be conducted the year-round from a sufficient number

of sites that together represent the body of water as a whole. Our

samples covered much offshore and some inshore water of the

central region of Lake Michigan and the southern quarter of

Green Bay, but they missed littoral and near-bottom areas. Sea¬

sonal or shorter period changes in trophic conditions cannot be

adequately described by sampling an area once. Moreover, the

minimal number of chemical and physical tests conducted here

limits discussion. Long-term changes in the macronutrients studied

by us can be obscured by analytical differences, small numbers of

samples combined with unrepresentative coverage, and conflicting

results. Thus, published nutrient data have not adequately quanti¬

fied eutrophication rates in Lake Michigan and there is conse¬

quently a great need for a suitable water quality monitoring pro¬

gram to be established on a lakewide basis. Additionally, nutrient

inputs from the drainage basin, groundwater, atmosphere, and

other sources should be measured — as well as various factors af¬

fecting nutrient output — in order to cope with nutrient budgets

and cycles.

Nevertheless, the instantaneous productivity can be estimated

from a single sampling, and this study demonstrated a higher

standing crop of phytoplankton in southern Green Bay — especially

near the Fox River: — than in the central region of Lake Michigan

in July of 1971.

1973] Rousar and Beeton — Phosphorus Distribution

139

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amination of water and wastewater. 12th ed., APHA, 769 pp.

AYERS, J. C., D. C. CHANDLER, G. H. LAUFF, C. F. POWERS, and

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GALES, M. E., E. C. JULIAN and R. C. KRONER. 1966. Method for quanti¬

tative determination of total phosphorus in water. Amer. Water Works

Assoc., 58: 1363-1368.

HARVEY, H. W. 1948. The estimation of phosphate and of total phosphorus

in sea waters. J. Mar. Biol. Assoc. U.K., 27: 357-359.

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HOLLAND, R. E. and A. M. BEETON. 1972. Significance to eutrophica¬

tion of spatial differences in nutrients and diatoms in Lake Michigan.

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R/V NEESKAY in central Lake Michigan and Green Bay, 8-14 July

1971. Center for Great Lakes Studies Special Report No. 13. Univ. Wis¬

consin — Milwaukee, Milwaukee, Wis.

KOBAYASHI, J. 1967. Silica in fresh water and estuaries, p. 41-55. In:

H. C. Golterman and R. S. Clymo [eds.], Chemical environment in the

aquatic habitat. N. V. Noord-Hollandsche Uitgevers Maatschappij, Am¬

sterdam. 322 pp.

RIGLER, F. H. 1964. The phosphorus fractions and the turnover time of

inorganic phosphorus in different types of lakes. Limnol. Oceanogr. 9:

511-518.

RISLEY, C. and F. FULLER. 1965. Chemical characteristics of Lake Michi¬

gan, p. 168-174. In: Proceedings 8th Conf. Great Lakes Res., Great Lakes

Res. Div., Univ. Michigan, 512 pp.

SAGER, P. and J. WIERSMA. 1971. Nutrient inputs in Green Bay, p. 6-8.

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Rept. No. 8. The Univ. Wisconsin Sea Grant Program. 219 pp.

SCHELSKE, C. L. and E. CALLENDER. 1970. Survey of phytoplankton

productivity and nutrients in Lake Michigan and Lake Superior, p. 93-105.

In: Proceedings 13th Conf. Great Lakes Res., Int. Assoc. Great Lakes

Res. 1063 pp.

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SCHELSKE, C. L. and E. F. STOERMER. 1971. Eutrophication, silica de¬

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von Gesamtphosphor im Wasser von Binnenseen. Schweiz. Z. Hydrol. 27:

184-192.

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Wis. Dept. Natural Resources, Div. Environ. Protection, 97 pp.

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VERBER, J. L. 1966. A study of Lake Michigan: chemical, biological and

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1966 and 1967. Wis. Dept. Nat. Resources.

PRE- AND POSTSETTLEMENT POLLEN FROM A SHORT

CORE, TROUT LAKE, NORTH-CENTRAL WISCONSIN

Thompson Webb III

Brown University —

Providence, R. I.

ABSTRACT

Pollen extracted from a short core collected in Trout Lake shows

the drastic changes in the composition of the vegetation around

the lake since logging and settlement of this area. The proportion

of pine pollen decreases by one-half. The proportion of hemlock

pollen also decreases. At the same time, the values for birch, oak,

alder, ragweed, and pigweed all increase. These changes indicate

the removal of white and red pine from the surrounding forests

and the clearance of some of the cut-over land. Secondary succes¬

sion by birch and the growth of weeds are favored by these

activities.

INTRODUCTION

Recent work (Davis, 1967; McAndrews, 1966; Ritchie, 1967;

Lichti-Federovitch and Ritchie, 1968; Webb and Bryson, 1972;

and Wright, 1968) indicates the importance of records of modern

pollen for the interpretation of assemblages of fossil pollen. Analy¬

sis of the surface sediments of lakes and peat bogs provides records

of the contemporary pollen rain. These samples of the modern

pollen can be correlated with modern vegetation and with spectra

of fossil pollen, thus providing a link between the fossil data and

the vegetation they reflect. A major problem in the use of modern

samples is that, unlike the fossil spectra, the modern spectra indi¬

cate not only the effects of climate and soils on the vegetation but

also the effects of man’s activities, e.g., logging and settlement.

In particular, pollen of herbs, especially ragweed (Ambrosia) ,

appears in greater proportion in contemporary sediments than

in sediments deposited prior to the massive disruption of the

natural landscape (Davis et ah, 1972; McAndrews, 1966). It is

important, therefore, to investigate the changes in pollen records

that are related to the land’s settlement by white men and to iden¬

tify, as well as possible, the effects of logging and land clearance

practices on these records.

141

142 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Short cores of bottom sediments from lakes provide a convenient

means for studying these changes. In general, a short column of

sediments will contain pollen from both settlement and presettle¬

ment times. A comparison of the pollen in the presettlement record

with the pollen deposited after settlement will reveal the elements

in the modern records related to man’s activities.

For this purpose, pollen samples were extracted and analyzed

from a short core taken from Trout Lake (46° 03' N, 89° 40' W) in

Vilas County, Wisconsin. According to a map of the early vegeta¬

tion of Wisconsin (Wisconsin Geological and Natural History Sur¬

vey, 1965), Trout Lake lies in a region of pine forests (Fig. 1),

which dominate on the sandy outwash deposits of this region

(Curtis, 1959; Hole and Lee, 1955). According to the vegetation

map of this area, white and red pines (Finns strobus and P. resin -

osa , respectively) were dominant in forests that included red maple

(Acer rubrum), red oak (Quercus borealis), white birch (Betula

papyrifera) , sugar maple (Acer saccharum) , and hemlock (Tsuga

canadensis) (Curtis, 1959). In addition to these species, Potzger

and Richards (1942) found balsam fir (Abies balsamea) growing

fairly abundantly in two tracts of primeval forest near Trout Lake.

METHODS

In 1966, a field crew led by G. Fred Lee of the Water Chemistry

Department of the University of Wisconsin-Madison used a 4-in.

90° W

FIGURE 1. Section of northern Wisconsin showing the location of Trout

Lake (T), Lake Mary (M), and Gillen Nature Reserve (G) within Vilas

County. The indicated vegetation data are taken from a map by the Wis¬

consin Geological and Natural History Survey (1965). Clear areas represent

northern hardwoods; cross-hatched, boreal forest; vertical lines, conifer

swamps; horizontal lines, pine forests; and stippled, pine barrens.

1973]

Webb — Pollen From Trout Lake

143

piston corer to obtain a 105-cm core (WC-59) from the south bay

of Trout Lake. The sediments consisted of greenish-gray to black

gyttja (Bortleson, 1971). The core was divided into 5-cm segments

and portions of each segment were separated for pollen analysis.

During 1967, subsamples taken from eight of the top 13 5-cm

segments were processed by standard palynological procedures

(Faegri and Iversen, 1964), which remove most of the unwanted

sediments and leave a concentrate quite rich in pollen. This residue

was placed in glycerine and mounted on microscope slides. The

pollen grains were identified at between 250 and lOOOx magnifica¬

tion on a Zeiss® microscope, and counts of over 300 grains were

made for each sample.

DESCRIPTION AND COMPARISON OF THE RESULTS

Table 1 contains the numbers of grains counted in each sample;

Fig. 2 is a diagram of the percentage values. A pollen sum includ¬

ing all but aquatic pollen types was used to calculate the per¬

centages plotted on the diagram.

Two zones are readily recognizable on the diagram, with the

zonal boundary lying between 35 and 40 cm depth. In the lower

zone, pine dominates the record, with values of 65-70%; while

hemlock values are 5-10% and birch about 15%. Above 35 cm,

the proportion of pine drops first to 55% and later to 35% ; hem¬

lock values are unchanged at first but decrease in the upper three

samples, ultimately to less than 2%. The values for birch rise to

30%; oak, to 10%; and ragweed, to 20%. The values of alder,

chenopods, and fir also increase. These changes are all related to

the logging and settlement of this area by white men.

The pollen spectra for the core are quite similar to others avail¬

able for this area. In the lower zone, the pollen values resemble

those found by Potzger and Richards (1942) within the upper

strata of four sites they investigated near Trout Lake. These sites

were all within the pine forest and their values of pine pollen were

all higher, while those of birch and hemlock were lower than the

values of these pollen types at sites within the northern hardwoods

(e.g., Lake Mary [Webb, in preparation] and Gillen Nature Re¬

serve [Potzger, 1942 ; see Table 2] ) . A clear separation of these

two forest types is present in these presettlement spectra, and

the pollen spectra at Trout Lake reflect the pine forests that grew

around Trout Lake prior to logging in the late 1800s.

The changes in the pollen spectra that occur above 35 cm are

quite similar to those found in pollen data extracted from short

cores within the pine region in Michigan’s lower peninsula (Webb,

in preparation). Some of these changes are also evident in the

TROUT LAKE, VILAS COUNTY, WISCONSIN

144 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

\

V*

%

^ ‘

<9 %

<s>.

r,

'Vo

V

’S'

* I

w

9k°&\

X~3

I I

m

r~~i

UJ

— !

H

UJ

CO

UJ

cr

o.

I:-.

□

B

o

<\fi

o.

2-,

FIGURE 2. Diagram of the pollen percentages for the levels counted in Trout Lake, Wisconsin.

1973]

Webb — Pollen From Trout Lake

145

TABLE 1. NUMBER OF POLLEN GRAINS COUNTED

TABLE 2. POLLEN PERCENTAGES IN PRESETTLEMENT SAMPLES

FROM PINE AND NORTHERN HARDWOOD FORESTS

Potzger and Richards (1942).

2Webb (1971).

3Potzger (1942).

upper one or two samples of some of the sites studied by Potzger.

At Bog C (Potzger and Richards, 1942), Gilbert Bog, and Birge

Bog (Potzger, 1942), in particular, the pine values decrease and

the oak and birch values increase. Potzger did not comment on

these changes in either of his two papers describing these sites;

his discussion concerned only the zones of longer duration in his

diagrams. Had Potzger used a finer sampling-interval, counted the

herbaceous pollen, and been looking for man-induced changes, he

might well have added an extra zone to each of his diagrams.

146 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

The pollen spectra of the upper zone at Trout Lake were com¬

pared with modern spectra collected in northern Wisconsin (Davis

et ah, 1969; Webb, 1971). These spectra were from either pine or

northern hardwood forests. The Trout Lake spectra most resembled

the modern spectra from regions of pine forests. The distinction

between the Trout Lake spectra and the spectra from the northern

hardwoods, however, is not as great as it was in presettlement

times.

The changes in the pollen record reflect the cutting of the pine

forests and their replacement by open land, with increased numbers

of herbs, and the secondary growth of forests, with increased

numbers of birch and oak. According to Curtis (1959), white birch

is among the first trees to invade bare soils in the successional

sequence within the region of pine forests. Increased growth of

oaks in this region may have been favored by an increased inci¬

dence of fire (Braun, 1950) shortly after the cutting of the pines.

The decline in hemlock at 15 cm, paralleling a rise in both rag¬

weed and pigweed, may be related to land clearance for settlement.

The continued depression of hemlock values may also be related

to the increase in the deer population in this area. A study by

Deboer (1947) shows that deer may be able to eliminate the repro¬

duction of hemlock and, thereby, decrease the numbers of hemlock

relative to other trees in stands.

However, not all environmental changes in the region were

caused by man. After the 1800s, the climate of this area changed

somewhat. Maps by Wahl and Lawson (1970), showing the re¬

corded temperatures and precipitation in the 1850s, indicate that,

compared to the period from 1931-60, northern Wisconsin annually

was about 2°F cooler and received an additional 3 in. of precipi¬

tation. The climatic change to warmer and drier conditions might

partially account for the rise in herbs and oak pollen. In this region,

however, the effects of these climatic changes are probably small

compared to the effects wrought by man, although there is no way

of completely separating the effects of these two factors.

Independent of this study, Bortleson (1971) completed a chemi¬

cal analysis of the sediments in core WC-59. Although both organic

carbon and organic nitrogen decrease sharply in the three samples

above 40 cm, no evidence for increased erosion of inorganic mate¬

rial from the uplands was found. Some peculiarities in the chemical

profiles led Bortleson (1971) to suggest the mixing of the sedi¬

ments by bottom convection currents. The regularity and con¬

sistency in the pollen profiles, however, dictate against excessive

mixing by the currents.

1973]

Webb — Pollen From Trout Lake

147

CONCLUSIONS

Man’s activities have had a severe effect on the vegetation around

Trout Lake. The pollen record in the uppermost sample is still

dominated by pine and birch but the ratio between these two pollen

types has changed drastically. Whereas pine pollen was over four

times more abundant than birch in presettlement time, pine is now

only slightly more frequent than birch. The values for oak have

doubled and the proportions for herbs and shrubs have increased.

Although the pollen values still place the modern spectrum in the

pine-forest region, the modern spectrum is not as clearly distin¬

guishable from samples collected in the northern hardwood region,

as were the presettlement spectra. These differences between the

modern spectrum and the presettlement spectra must be considered,

when modern data are used to aid interpretation of profiles of fossil

pollen.

ACKNOWLEDGMENTS

Contribution No. 162 of the Great Lakes Research Division, The

University of Michigan.

Major support for this work came from NSF grant GA-10651X,

to R. A. Bryson and J. E. Kutzbach at the Center for Climatic

Research, University of Wisconsin-Madison. Some support during

the writing of this paper was provided by NSF grant GB-24836, to

Margaret B. Davis, Great Lakes Research Division, University of

Michigan. I thank G. F. Lee and G. C. Bortleson for making the

core available to me, R. Willard for laboratory assistance, and

L. Brubaker for a critical reading of the manuscript.

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WRIGHT, H. E., JR. 1968. The roles of pine and spruce in the forest history

of Minnesota and adjacent areas. Ecology 49:937-955.

Wisconsin Geological and Natural History Survey. 1965. The early vegetation

of Wisconsin (Map). Wis. Geological and Natural History Survey,

Madison.

MARKET STRUCTURE AND BANK PERFORMANCE:

WISCONSIN 1870-1900

Richard H. Keehn

University of Wisconsin —

Parkside

INTRODUCTION

This paper is a summary of the author’s study of nineteenth

century Wisconsin banking.1 The study is not intended as a com¬

plete history of Wisconsin banking but explores certain questions

of historical interest with respect to Wisconsin economic develop¬

ment which also have applicability to current problems of bank

regulation and control.2 The primary purpose of the study is to

outline the changing statewide and local market structure of the

commercial banking industry in Wisconsin from 1853 through 1914

and to use this information to explore the major determinants of

individual bank performance with special emphasis on the impact

of local market structure on individual bank lending performance.

Economic historians have emphasized the importance of in¬

creased and improved financial intermediation in the process of

economic growth and development, and the existence of under¬

developed capital markets are seen as a barrier to the mobility

of funds which tends to retard development. Increased and im¬

proved intermediation and institutional innovation should improve

the efficiency with which financial intermediaries provide financial

services, increase the mobility of funds between areas and indus¬

tries, and possibly raise aggregate savings rates. In the United

States where geographical distances were substantial and trans¬

portation and communication relatively undeveloped, improved

and increased intermediation should have been of considerable im¬

portance.3 It has also been suggested that many bankers in the

1 Richard H. Keehn, Market Structure and Bank Performance: Wisconsin, 1870-1900

(Madison, unpublished Ph.D. dissertation, University of Wisconsin, 1972). The findings

reported in this paper are from this source.

2 The major studies of the history of Wisconsin banking are: Leonard B. Krueger,

History of Commercial Banking in Wisconsin, University of Wisconsin Studies in the

Social Sciences and History, No. 18 (Madison, the University of Wisconsin, 1933) ;

Theodore Andersen, A Century of Banking in Wisconsin (Madison, State Historical

Society of Wisconsin, 1954).

3 Lance Davis, “The Investment Market, 1870-1914: The Evolution of a National

Market”, The Journal of Economic History, Vol. XXV, (September, 1965), pp. 355-

356.

149

150 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

United States in the nineteenth century were in relatively monopo¬

listic positions which enabled them to exploit their customers to

varying degrees, but very little is known about the influence of

bank market structure on the actual performance of individual

banks.4 Bank regulatory authorities are interested in the impact

of bank market structure on the performance of individual banks

and the issues examined are of interest to these agencies.

METHODOLOGY

A model of individual bank behavior and performance is devel¬

oped that conforms to conditions in nineteenth century Wisconsin.

The model generates predictions about bank performance with

respect to loan rates and loan output, and it is used to explore the

major potential influences on bank behavior and performance.

Banks operate in two markets which it is profitable to separate

because of differing elasticities of demand. The loan market tends

to be geographically limited because high transfer costs (defined

to include information, transportation, convenience and communi¬

cation costs) protect each local market to a considerable degree

from “outside” competition. Loan customers usually do not have

access to financial institutions outside their local area. An indi¬

vidual bank also operates in the investment market which tends

to be regional or national in scope and the bank has little or no

influence on prices charged and the elasticity of demand approaches

infinity. Because of the limited scope of the loan market, the firm

can exert some control over price and bank behavior and per¬

formance will be influenced by the competitive conditions in that

market — the number of rivals and the concentration of assets.

Bank output is defined as earning assets. To maximize profits a

bank will invest in earning assets to the point where marginal

cost equals marginal revenue in the market where the elasticity

of demand is greatest (usually the investment market). It will

invest in loans to the point where the marginal revenue from loans

equals the marginal revenue from investments. The major hypothe¬

sis tested is that the structure of the local loan market will exert

a measurable influence on the loan performance of individual banks.

The model indicates that the more concentrated the local banking

market the more the monopoly power that can be exerted by the

firm or firms in that market. Profit maximization entails lower loan

ratios and higher loan rates in less competitive markets, ceteris

paribus. Other factors exert an influence on the loan demand curve

facing an individual bank and these factors must be considered in

4 Richard Sylla, “Federal Policy, Banking Market Structure and Capital Mobiliza¬

tion in the United States, 1863-1913,” The Journal of Economic History _, Vol. XXIX

(December, 1969), pp. 657-686.

1973] Keehn — Market Structure and Banking 151

estimating the impact of structure on performance. These variables

include demand, organizational form, and internal factors. Specific

hypotheses about the potential impact of these variables on bank

performance are tested along with the primary hypothesis.

Market structure, a measure of competition in local loan mar¬

kets, is measured by the number of banks in the relevant market,

by the share of total bank market assets controlled by the largest

bank in the market, and by variations of the Herfindahl Index —

the sum of the squares of each bank’s market share.5 Non-bank

intermediaries were examined but were quantitatively unimpor¬

tant throughout the period and were not specifically included in

the tests. Individual bank performance is measured by the ratio

of loans to total assets. Higher loan ratios imply larger credit

extensions to the local area and this is generally regarded as

favorable for that area.

Local bank loan markets were defined in two ways. The first

treated each county as the local market and used structural meas¬

ures based on this geographical area. Within each county banks

faced similar conditions and were protected to some extent from

outside competition. In many cases this market may be too broad ;

banks probably operated in more restricted markets in many areas

and time periods. An alternative approach defined the relevant

market as the city or town. This measure is considerably narrower

but may be more realistic. As expected, concentration was much

higher in the latter case.

DATA

The hypotheses were tested with data on each bank in Wiscon¬

sin for census years 1870-1900, obtained from the records of the

various state and national agencies charged with bank supervision.

Balance sheet data on individual banks provided most of the neces¬

sary information. Detailed information on county and local markets

was developed from these and other sources. Multiple regression

analysis was used to estimate the relative importance of the fac¬

tors suggested by the model as potentially influencing bank lend¬

ing with special emphasis on the structural variables. Succeeding

cross-sections were then compared in order to isolate changes in

the determinants over time. Throughout the period commercial

banks were the major financial intermediary in Wisconsin and the

major changes within the financial sector was the rapid increase

in the number of banks; for this reason attention is centered on

these institutions.

5 For a discussion of these measures see: William Paul Smith, “Measures of Bank¬

ing Structure and Competition,” Federal Reserve Bulletin , (September, 1965), pp.

1212-1222.

152 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Why History ?

The issues studied are of both historical and current interest,

and examining them with historical data has several advantages.

The relatively long period involved (1870-1900) encompasses a

time of rapid change and growth in the state’s financial sector

and economy. The diversity within the state increased greatly over

the period in terms of local banking markets and the economic

development of various regions within the state. A viable manu¬

facturing sector developed to complement the agricultural base;

Milwaukee developed as a major urban center and other cities

grew rapidly as centers of population and industry, yet many areas

remained essentially undeveloped throughout the period.

Another advantage is that three distinct bank types operated

during the period and the differences between these were substan¬

tially greater than those existing between member and non-member

or state and national banks today. Private (unincorporated) banks

faced virtually no regulation or control except that faced by the

average business firm. State banks operated under supervision and

control that limited their activities in some ways at least. National

banks operated under a well defined set of regulations and were

closely supervised in an attempt to influence their behavior, espe¬

cially with respect to portfolio policy. Despite the controls imposed

on state and national banks between 1870 and 1914, banking was

less restricted than in earlier and later periods. The existence of

two bank types regulated to differing degrees and an essentially

unregulated segment presents an opportunity to study the effec¬

tiveness of these regulations in altering bank behavior and per¬

formance.

While entry into nineteenth century banking was more restricted

than into other industries, it was relatively easy compared with

more recent experience. A potential private banker faced virtually

no restraints; state and national charters imposed certain condi¬

tions and standards on potential entrants but beyond assuring

compliance with entrance requirements, regulatory authorities,

state or national, exercised almost no administrative control over

entry, and potential entrants were freer to respond to market

forces than in more recent periods.6 In one respect nineteenth

century Wisconsin banks faced limitations similar to those in re¬

cent periods; banks were generally not permitted to branch and

the prohibition was made explicit for state banks in 1909. Because

branch banks were virtually non-existent, it was impossible to

compare the performance of branch and unit banks.

0 The Wisconsin Commissioner of Banking- attempted to deny a new state charter

in 1914 on the grounds that another bank was not needed in the area but he was

overruled by the Board of Appeal which claimed that the Commissioner did not have

the power to pass on charter applications. See Wisconsin State Journal (Madison),

July 9, 1914 and Milwaukee Sentinel, July 17, 1914.

1973] Keehn — Market Structure and Banking 153

Major Findings

Unit banking* predominated in nineteenth century Wisconsin and

this pattern has continued to the present. The number of banks

increased from 97 in 1870 to 776 in 1914 and the number of per¬

sons per bank declined from 10,782 in 1870 to 3,681 in 1900. The

prohibition of branching and poor communication and transporta¬

tion acted to limit market size, and were primarily responsible

for this pattern of small independent unit banks. Bank sector as¬

sets, loans and deposits, total and per capita, expanded substan¬

tially from 1853 through 1914, but the growth of the banking sec¬

tor was uneven, and private, state and national banks exhibited

different patterns with respect to the rate of growth of assets.

Changes in the total number of banks changed the structure of

county and local banking markets as well. The number of counties

with two or more banks increased from 13 to 61 between 1870

and 1900, while the number of cities and towns with banks in¬

creased from 31 to 221. The average banking market — -county

or city — remained quite concentrated by usual standards; 29 of

70 counties had less than four banks in 1900 and many banking

markets appear to have been monopolistic or oligopolistic.

Bank size and population of town where located were positively

related, reflecting the influence of market size on bank size (espe¬

cially in the absence of branching and in the face of high transfer

costs). The average national bank was larger than the average

state bank and tended to locate in larger cities and towns, reflect¬

ing, in part, the higher capital requirements of national banks.

Private banks tended to be smaller and to locate in places that

state and especially national banks found unprofitable. Average

national bank size increased substantially between 1870 and 1900,

but average size of state and private bank exhibited little increase

over the period.

A study of the regression coefficients indicates that structural

variables— -the number of banks or various measures of concen¬

tration in each local market, county or local — had little influence

on bank lending performance as measured by the loan : total asset

ratio. These results were fairly constant over time, did not change

much as additional variables were added or deleted from the equa¬

tions, and were similar whether counties or towns were used as

local markets. In general, differences between markets in the de¬

gree of local bank competition were of little help in explaining

inter-bank variation in loan ratios. These results conflict with the

hypothesis that structure exerts a measurable influence on bank

performance and are at odds with studies of the structure-per¬

formance relationship in banking in the 1950’s and 1960’s, which

usually found a statistically significant but small impact of struc-

154 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

ture on performance.7 These conflicting results can be reconciled

by examining entry conditions. Entry into banking markets in the

nineteenth century was much easier than in more recent times

(especially since 1933). 8 There was virtually no administrative

control, and while charter requirements did impose some restric¬

tions, these were not burdensome. Entry into banking was more

difficult than into other industries but not by much. This relative

freedom of entry meant that banks in apparently restricted mar¬

kets were faced with the possibility of new competitors and new

banks did, in fact, enter more and more markets over time. Poten¬

tial and actual entry tended to make most banking markets com¬

petitive despite appearances to the contrary, acted as a constraint

on the behavior of existing banks, and made bank performance in

different markets relatively insensitive to variations in local com¬

petitive conditions. This interpretation is consistent with the ob¬

served increase in the number of banks in the state from 97 in

1870 to 362 by 1900 (and 776 by 1914). It is also consistent with

economic theory which indicates that inability to block entry of

new Arms will weaken monopoly power of existing firms.

Apparently monopolistic or oligopolistic banking markets existed

throughout the period despite the rapid entry of new banks, but

these do not appear to have been the result of direct restrictions

or the restrictive features of the National Banking Acts. The con¬

tinued existence of markets with one or a few banks and/or high

concentration ratios can be explained by product differentiation

advantages of existing banks and by limited market size which

tended to limit entry of new banks in many cases. New banks did

enter local markets when these expanded in size to accommodate

additional firms. The restrictive features of a national bank char¬

ter tended to force national banks to locate in cities and towns of

above some minimum size but the prohibition of state bank notes

did not affect the entry of state and private banks after about

1880 when deposit banking developed rapidly.9

Organizational form should exert a measurable influence on

7 For example see : Richard C. Aspinwall, “Market Structure and Commercial Bank

Mortgage Interest Rates’’, The Southern Economic Journal, Vol. XXXVI, (April

1970), No. 4, pp. 376-384; Eric Brucker, “A Microeconomic Approach to Banking

Competition’’, Journal of Finance, Vol. XXV, (December 1970) No. 5, pp. 1133—1141;

Franklin R. Edwards, Concentration and Competition in Commercial Banking: A Sta¬

tistical Study (Boston, Federal Reserve Bank of Boston, 1964) ; George G. Kauf¬

man, “Bank Market Structure and Performance : The Evidence from Iowa”, The

Southern Economic Journal, Vol. XXXII (April 1966) pp. 429—439; Paul A. Meyer,

“Price Discrimination, Regional Loan Rates and the Structure of the Banking In¬

dustry”, The Journal of Finance, Vol. XXII, (March 1967) No. 1, pp. 37-48; Al-

marin Phillips, “Evidence on Concentration in Banking Markets and Interest Rates”,

Federal Reserve Bulletin, (June 1967), pp. 916-926.

8 Sam Peltzman, “Entry in Commercial Banking”, Journal of Law and Economics,

Vol. VIII, (October 1965), pp. 11-50.

9 These conclusions are at variance with those of Sylla, “Federal Policy”, pp. 6 5 7 —

686.

1973]

Keehn — Market Structure and Banking

155

individual bank loan ratios. National and state bank regulations

were designed to alter bank portfolio decisions usually in the di¬

rection of greater safety, and therefore should lower bank lending

below the level desired, if unregulated. The specific hypothesis

is that national banks should have lower ratios than state banks

because of more restrictive regulations, while private banks should

have higher loan ratios than state banks because of freedom from

virtually all regulation and control. The regression coefficients in¬

dicate that national banks did tend to invest less in loans than

state banks, ceteris paribus. This might have been partially re¬

sponsible for the excellent safety record of these institutions, but

the impact on the economic development of the local area, the state

and nation is not clear. Private banks tended to have lower loan

ratios than state banks, ceteris paribus; this unexpected result

probably relates to the unsophisticated management and small

size which limited a private banker’s ability to diversify and offset

the advantage of virtually no regulation.

The most consistent and significant influence on individual bank

lending was the capital or net worth to total asset ratio. Portfolio

theory suggests that net worth and loan ratios should be positively

related because a high net worth ratio allows a bank to assume

more risk in the form of loans in striving for higher yield. The

regressions indicate that the ratios were in fact negatively asso-

ciated and the coefficients were large and significant (at the five

percent level) in all but 1880. Net worth ratios reflect differences

in risk between banks; bank size and net worth ratios were in¬

versely related indicating that small banks were riskier than

larger units on the average. Minimum capital requirements also

forced some small banks to maintain higher than desired net worth

ratios given the level of earning and total assets.

Other financial variables exerted a mixed influence on individual

bank lending. The impact of absolute bank size was somewhat un¬

stable but there is an indication that larger banks had higher loan

ratios, reflecting (possibly) a greater ability to diversify and to

take advantage of possible economies of scale. A high ratio of in¬

terest earning to total deposits was associated with marginally

higher loan ratios, indicating that banks tended to lend a greater

share of these deposits, ceteris paribus. There is no indication that

increased competition in local markets was associated with a higher

percentage of total deposits earning interest, as has been suggested

by proponents of the prohibition of interest payments on demand

deposits and the limitations on the rate paid on time deposits.

Efforts to account for differences in demand or scale between

markets were not very successful. Demand measures were gen¬

erally not significant and the coefficients quite small, probably re-

156 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

fleeting the use of inappropriate measures more than the actual

unimportance of demand factors per se. Location size (popula¬

tion), population change, market assets and market assets per

capita were all tried as a measure of the scale of the market.

Several of the variables were correlated with other independent

variables which contributed to the insignificant results.

Additional information on bank performance was developed

from the income and expense data reported for national banks

beginning in 1870. The data are divided into Wisconsin country

national and Milwaukee national banks but are not available on

an individual bank basis. These data are used to examine addi¬

tional areas of bank performance that have received considerable

attention from economists recently. Costs per unit of earning as¬

sets of Wisconsin national banks appear to have declined up to

some minimum level of output and then remained relatively con¬

stant over a wide range and there is little evidence in support of

the notion of economies of large scale. The rate of return on in¬

vestment in national banking remained relatively high and appar¬

ently above rates on alternative investments of comparable risk.

Above normal profits and relative ease of entry combined to attract

an increasing number of new banks into the market. This increased

entry of banks of all types evidently was unable to substantially

reduce returns to equity because demand for banking services was

shifting rapidly at the same time.

Various interest rate data confirm that substantial differences

in interest rates charged on loans existed within the state and

between Wisconsin and other areas in the nineteenth century. It

has been suggested that these differentials reflected differing de¬

grees of monopoly power in various markets and that the decline

in the differentials between areas resulted from institutional

changes, improving the mobility of funds which, along with the

erosion of entry barriers, increased bank entry and competition

in many local markets. The regression results suggest that inter¬

market differences in structure had little impact on individual bank

performance as measured by the loan ratio. Differentials in inter¬

est rates are best explained by the existence of positive and high

transfer costs between markets. Institutional changes, including an

increasing number of banks and the development of new financial

institutions, acted to lower transfer costs, broadly defined, but were

only one of several factors tending to do so.10 Improvements in

transportation and communication, for example, acted to reduce

transfer costs and therefore interest rate differentials between

regions, so these differentials would have declined somewhat even

in the absence of various institutional changes.

10 This conclusion differs from that of Davis, “The Investment Market”, pp. 355-

399.

LA QUERELLE DU CID: CLASSICAL RULES

OR POLITICAL EXPEDIENCY?

Edmund Roney

Ripon College —

Ripon

For years historians and scholars interested in the development

of the drama in France have puzzled over the curious set of cir¬

cumstances that, in 1637, led the recently established French

Academy to officially condemn Pierre Corneille’s Le Cid, the finest

and most popular play that had yet been produced in France.

Among the many questions to which “la querelle du Cid” has given

rise, the most fascinating has been, why did Cardinal Richelieu

revise his attitude toward the play during the summer of 1637?

Shortly after the initial production of Le Cid, Richelieu requested

two command performances of the play at his own palace. When

it was published, he permitted Corneille to dedicate it to his

favorite niece, then Madame de Combalet (later the Duchesse de

Rambouillet) . The ennoblement of Corneille’s family occurred in

January of 1637 with no objection from the Cardinal. In March,

a pamphlet war broke out. Many pamphlets attacked Le Cid and

its author, while some defended both. One of the attackers, George

de Scudery, suggested that the matter be turned over to the newly

formed French Academy for its resolution. According to an early

account, Histoire de VAcademie frangaise, written in 1653 by Paul

Fontanier Pellisson and l’Abbe d’Olivet, the Academy was reluctant

to take up the matter because its rules restricted it to scrutinizing

only the work of its own members and Corneille was not a mem¬

ber in 1637. The book cites a letter written by Corneille dated

June 13, 1637 to M. de Boisrobert, one of Richelieu’s secretaries

and an official of the Academy, in which Corneille, after clarifying

his objection to having the Academy criticize his play, stated:

The gentlemen of the Academy may do what they please, since you write

me that MONSEIGNEUR would be pleased to see their judgment. If that

would entertain His Eminence, then I have nothing to say.1

This comment of Corneille’s was evidently deemed sufficient to

give the Academy jurisdiction in the affair. Pellisson states that,

1 Pellisson et d’Olivet, Histoire de VAcademie francaise. Livet, ed. Paris: Didier

et Cie., 1858, pp. 86-89.

157

158 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

when the Academy still resisted undertaking the work, Richelieu

instructed one of his servants to

Tell the gentlemen that I wish this thing to be done, and that I will

esteem them as they esteem me.2

When the Academy finally took up the matter, it was assigned to

Jean de Chapelain, who, in a letter to Boisrobert dated July 81,

1637, wrote that a “judgment” of Le Cid had been sent to Riche¬

lieu prior to that date. This letter refers to a personal interview

between Chapelain and Richelieu, which, in the later light of Pel-

lisson’s description of the same event, demonstrates that Chape¬

lain was being subjected to a considerable amount of personal

pressure regarding his role in the affair. Chapelain commences the

letter with a reference to the critique he had written of Le Cid

in the Academy’s name. He mentions a number of criticisms that

he had received from Richelieu on it and asks Boisrobert to relay

his answers to those criticisms, should he catch the Cardinal in a

propitious moment of leisure. He protests that the public will take

the Academy for a poor judge, if it fails to find anything good to

say about Le Cid?

Pellisson reports that, according to the Academy Registers,

Chapelain first presented his notes on June 30, 1637, and that it

was ordered that his work be combined with that of messieurs de

Bourzeys and Desmarests. He states that the Academy Registers

do not show that this was done, but that Chapelain then delivered

the manuscript in his own writing to Richelieu.4 This document

was returned to the Academy with marginal notes both in the hand

of Richelieu and in that of Citois, his personal physician. Riche¬

lieu’s final comment was that the work needed a few handfuls of

flowers, or compliments.5 The Academy then turned the manuscript

over to a committee composed of Messieurs de Serizay, de Cerisy,

de Gombauld and Sirmond on July 17, 1637. When the galley

proofs of this revision were sent to Richelieu, they evidently dis¬

pleased him to such an extent that he had the printing stopped

immediately and asked Chapelain and the committee to report to

him immediately. This is undoubtedly the interview referred to

above by Chapelain in his July 31 letter to Boisrobert. Pellisson

records that Chapelain and two of the committee were advised

of the Cardinal’s wishes in no uncertain terms.

2 Ibid.

3 Lettres de Jean Chapelain, Tamisey de Larroque, editor, 2 vols., (Paris: Im-

primerie nationale, 1880-1883), Vol. I, p. 156.

4 Pellisson, op. cit., p. 91.

5 Collas, George, editor. Les Sentimens de V Academe franc a-ise sitr la Tragi-comedie

du Cid ; d’apres le Manuscrit de la main de Chapelain conserve a la Bibliotheque

Nationale avec les corrections, une introduction et des notes. Paris : Librairie Alphonse

Picard, 1912.

1973] Roney — La Querelle Du Cid 159

Richelieu spoke so intensely to Chapelain that he ivent so far

as to seize the tassels of his collars from time to time, as one is

apt to do unconsciously ivhen, in talking with another, one has a

strong desire to convince him of something .6

Chapelain’s July 31, 1637 letter to Boisrobert assumes a much

clearer aspect in view of this incident, which occurred on the same

day. There is no need to search for a letter from Boisrobert bear¬

ing the criticisms Chapelain attempts to answer in his letter

because Chapelain evidently received them at first hand on that

very day from Richelieu himself. That is why he timidly requests

that Boisrobert communicate his answers to the Cardinal’s criti¬

cisms “in a moment of leisure.”

There appears to be little doubt that, after his initially favor¬

able reaction to Le Cid, Richelieu became concerned with the quar¬

rel that followed and influenced the Academy to condemn the play.

The puzzling question is, Why? How could the first minister of

France have become involved in a literary quarrel at a time when

he was vigorously prosecuting foreign wars on three fronts and

engaged with domestic conspiracies? Some understanding of the

background of the problems with which Richelieu was confronted

in 1637 is necessary to the elucidation of this question.

Throughout the entire period of Richelieu’s tenure as first min¬

ister of Louis XIII, he was faced with serious domestic and foreign

problems. The internal unity and security of France were period¬

ically endangered by the relative independence of the Protestant

seaports from royal authority, the power of many French nobles

to resist or obstruct the power of the King, and dissensions within

the Royal Family itself. These dangers were intensified by the

external threat of the House of Hapsburg, whose dominion in

Spain, Italy, Austria, Germany and the Netherlands at times con¬

stituted virtual encirclement of France. The tendency of some sea¬

ports to side with England in periods of crisis made the eventual

isolation of France a clearly forseeable prospect. Richelieu attacked

these problems with great skill. He secured France’s flank through

his successful completion of the siege of La Rochelle. He reduced

the insurrectionary power of the French nobility through his

destruction of unnecessary internal fortifications and further dis¬

ciplined their tendency toward lawlessness through a strict en¬

forcement of his edict against duelling. To counter the military

actions that Spain was carrying out in the Valtelline, Padua, Man¬

tua, Franche Comte, the Netherlands and Wallenstein’s victories

in Bohemia, Gustavus Adolphus of Sweden was encouraged and

subsidized to combat the Hapsburgs in Germany through the

treaty of Barwalde.

0 Pellisson, op cit., p. 92.

160 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

While implementing these policies, Richelieu was subjected to

the severe harassment of constant plotting against not only his

own power, but the crown of Louis XIII. In the same year that he

moved against the independence of the feudal aristocracy, 1626,

the revolt of the “Conspiracy des Dames” broke out. Led by the

King’s brother, Gaston d’Orleans, it was supposedly a reaction to

Louis’ plan to wed Gaston to Mile. Montepensier. In his Histoire

de France (1756) le pere Daniel states that one of the conspira¬

tors in this plot, Henri de Talleyrand, the Marquis of Chalais,

attempted to implicate Queen Anne in the conspiracy. His con¬

fession indicated that the aim of the plot was to depose the King,

break off his marriage to Anne of Austria, and marry her to

Gaston d’Orleans.7 Daniel states that this was the main reason for

the prolonged aversion Louis XIII conceived for Anne, which

lasted, even after the birth of an heir, until his death. On his death

bed, when the Queen is said to have requested M. de Chavigny to

tell the King that she had never been guilty of what had been

inputed to her in the “affaire de Chalais,” the King responded,

“In my present state, I am obliged to pardon her, but I am not

obliged to believe her.”8

On November 12, 1630, the famous “Day of Dupes” incident

took place. The King’s mother, Marie de Medicis, who had helped

Richelieu to power and then turned against him, combined with

Anne of Austria to attempt to persuade Louis XIII to dismiss

Richelieu. They almost succeeded, but at the last moment, the King

decided to retain Richelieu. This provoked Gaston d’Orleans to

head another revolt, this time to free the King and the country

from the tyranny of Richelieu. It was crushed at Castelnaudry.

Following the death of Gustavus Adolphus at Lutzen in 1632,

and the conclusion of the Treaty of Prague in 1635, whereby the

Emperor Ferdinand of Austria was reconciled with most of the

German princes, Richelieu decided that the time had arrived for

France to commence open participation in the Thirty Years War.

Hostilities were accordingly commenced against Spain in Italy and

the Netherlands on May 29, 1635. At first the war went well for

the French, but in the summer of 1636, after receiving reinforce¬

ments from Austria, the Spanish invaded France from the Nether¬

lands, crossed the Somme River, took Corbie and threatened Paris.

On August 22nd the King led his army out of Paris and re-took

Corbie on November 14, 1636. The Spanish retired across the

Somme, but fighting continued on French soil through January

and February of 1637. (It seems indeed curious that at this same

time, a play celebrating a legendary Spanish hero should com¬

pletely capture the hearts of Paris.)

7 Le Pere Daniel, Histoire de France , 1756, t. XIII, p. 515.

8 Ibid.

1973]

Roney — La Querelle Du Cid

161

On November 21, 1636, six weeks before the first performance

of Le Cid, Gaston d’Orleans again revolted and fled to Blois. In

league with the Comte de Soissons, the aim of this plot was to

assassinate Richelieu. At the last minute, the King’s brother backed

out of the plot, but it was only after strong intimidation that he

was finally reconciled with the King on February 8, 1637. In the

summer of 1637, a similar reconciliation was effected with the

Comte de Soissons to prevent his becoming a tool of the Spaniards.

To further complicate matters, the Queen was discovered to be

in correspondence with the enemies. This correspondence might

have been innocent enough and of a purely personal nature, since

they were her relatives. Her brother, Phillip IV, was the King of

Spain and another brother, the Cardinal Infanta, commanded the

Spanish and Austrian armies in the Netherlands. Because of the

war, any communications with her relatives of which the King

was not advised had to be considered suspect. Suspicion was fur¬

ther increased by Anne’s intimacy with the Duchesse de Chevreuse,

an implacable enemy of Richelieu, who had been closely associated

with the attempts of the Queen Mother in exile, Marie de Medicis,

to subvert the power of France.

Richelieu and the King had been aware of this correspondence

for some time. They had learned that it was conducted from the

Convent of the Val-de-Grace, where the Queen was in the habit

of retiring for her religious devotions. Her letters were carried

by her personal messenger, La Porte, to the secretary of the Eng¬

lish ambassador, who then transmitted them to Brussels. By this

means, she communicated with the Marquis de Mirabel, the Cardi¬

nal Infanta, the Duke d’Olivares, the Queen of England, the Duke

of Lorraine and the Spanish court. In the last week of July, just

prior to his interview with Chapelain about Le Cid, Richelieu had

intercepted a letter from the Marquis de Mirabel to the Queen, but

he had been unable to intercept the Queen’s response. The King

now ordered the Chancellor and the Archbishop of Paris to make a

detailed search of the Queen’s apartments in the Val-de-Grace

convent. On the 12th of August La Porte was arrested; on the

13th the Val-de-Grace was searched, and on the 17th, the Queen

was interrogated, confessed, and was reconciled with the King.

During the period covered by “la querelle du Cid ,” from January

to December, 1637, Richelieu was extremely preoccupied with

affairs of state. It seems incredible that he could have been in any¬

way connected with the literary dispute. The first reference to

Richelieu in connection with it appears in one of the pamphlets

of the dispute written by Corneille, the Lettre Apologitique which

mentions the three performances of Le Cid at the Louvre and its

two performances at the “Hostel Richelieu.” Two performances in

162 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Richelieu's own palace was clearly a high mark of personal favor

on the part of Richelieu. If he had seen anything objectionable

about the play at this time, he would never have singled it out for

such an honor. To this may be added the facts that he did nothing

to prevent the ennoblement of Corneille’s family and did not object

to Corneille’s dedication of the play to his niece, Madame de

Combalet. The spurious legend that he was personally jealous of

Corneille because of the enormous success of Le Cid is flatly contra¬

dicted by the facts of his initial acceptance of the play.

A more plausible legend is that Richelieu opposed Le Cid because

it presents the practice of dueling in a favorable light. If this were

so, one would suppose that his opposition would have been immedi¬

ate. He is not known to have protested the favorable references

to duels in many other plays of the period, including his own

L’Aveugle de Smyrne. His edicts against dueling were not directed

against stage duels, which invariably involved affairs of honor,

but to brawling breaches of the King’s peace. Had Le Cid contained

anything of this nature, Richelieu’s animosity toward it would

have been displayed in January, when he witnessed it, and not six

months later.

The next connection between Richelieu and Le Cid appears in

Corneille’s letter to Boisrobert dated June 13, 1637, indicating

the Cardinal’s desire to see the matter turned over to the Academy.

There is no indication that he was hostile to the play at this time.

The marginal notes in the hand of Richelieu and Citois on

Chapelain’s first draft of Les Sentimens de V Academie francaise

sur le Tragi-comedie du Cid are also relatively noncommittal.

It is not until Chapelain’s July 31, 1637 letter to Boisrobert that

it appears that Richelieu had developed a strong partisan interest

in the quarrel. Pellisson’s account of the interview between Chape-

lain and Richelieu on that date substantiates this sudden preoccu¬

pation on Richelieu’s part with a condemnation of Le Cid . The

obvious question is, what happened at that time to awaken Riche¬

lieu’s animosity toward the play? In the last week of July, 1637,

Richelieu had intercepted incriminating correspondence between

Queen Anne and the Marquis de Mirabel. This led directly to the

Val-de-Grace incident. Could there have been any connection be¬

tween this event and Richelieu’s attitude toward Le Cid?

To substantiate this possibility, some connection would have to

be found between the Queen and the play. In the opinion of the

court historian, Jean Sirmond, the play had caused the Queen to

instigate the ennoblement of Corneille’s family.9 It is possible that

this was only an unusual reflection of the extent to which the play

e Le Souhait du Cid in Gaste, La Querelle du Cid , (Paris: H. Welter, 1819), p. 186.

1973]

Roney — La Querelle Du Cid

163

pleased the Queen, as it did many others in Paris. It is also possible

that she or her supporters saw in the play, which was adapted

from a Spanish original, a sympathetic presentation of a problem

very similar to the one in which the Queen found herself. The main

dramatic problem in Le Cid focuses on Chimene, once Rodrigue

decides to avenge his father. Chimene’s problem then becomes a

question of whether her primary loyalty is to her family honor or

to Rodrigue. She does not arrive at any really satisfactory solution

to her difficulty, but, in the course of coping with it, the play gives

a very thorough exposition of the problem.

Queen Anne had problems too, as previously indicated. Their

source was a situation basically similar to that of Chimene, i.e., a

conflict between her father's family and her husband. There are

obvious differences in detail between the character Chimene and

the Queen. Chimene is not married to Rodrigue as the Queen was

to Louis XIII, and Rodrigue fights Chimene’s father, while Louis

was at war with his Queen’s brothers. The situation is not at all

unusual historically. Many queens have found themselves in similar

circumstances. Anne’s sister-in-law, Henrietta Marie, Queen to

King Charles I of England was caught up in a comparable conflict

of interests during the same period.

To an audience trained by '‘Romans a clef” to discover allusions

to contemporary events in literary works,, it would have been a

simple matter to discover analogies between Chimene and Queen

Anne. In Le Souhait du Cid , Sirmond links both of their names in

answering Le Cid’s critics. “I am one,” he writes, “who hates those

who do not love Chimene, and infinitely honors she who, by pro¬

curing the author’s nobility, has judged her favorably.”10

If supporters of the Queen had commenced to draw analogies

between Chimene and Queen Anne, this would certainly explain

Richelieu’s sudden desire in the last week of July, 1637 to see the

play condemned. It was precisely at this same time that he dis¬

covered evidence leading him to suspect the Queen of treason. This

also would explain his initial favorable reception of the play and

the subsequent continuance of Richelieu’s patronage to Corneille.

Obviously he had no personal animosity toward the play or its

author. What he was concerned with was the political use that the

Queen’s supporters might attempt to make of the unusual artistic

success of Le Cid and the notoriety of the quarrel that followed it.

He was far too absorbed in major affairs of state to take more than

a passing interest in the quarrel until it became apparent that his

enemies might be able to use it for their own political advantages.

So, he took steps to neutralize it. Even then, his manner of han¬

dling it was discreet. He did not permit himself to become embit-

10 Gaste, loc. cit.

164 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

tered against Corneille. He knew that Corneille could not be held

responsible for political interpretations that others might read into

his play. The most discreet way to dispose of the issue was to turn

it over to the Academy and make sure that they did not fail to

condemn Chimene’s behaviour as immoral, which they eventually

did.

One of the results of the quarrel and its culminating document,

Les Sentimens de VAcademie francaise sur le Tra gi-come di e du

Cid, was the popularization of the “rules” of French neo-classical

drama to the extent that by 1640, they were generally known,

accepted and followed by French dramatists, and this led directly

to the great seventeenth century period of neo-classicism. It would

not, however, be accurate to say that the primary nature of the

quarrel was a dispute over these “rules.” Although discussion of

the rules entered into some of the pamphlets, the majority of them

were concerned with the expression and refutation of jealousy,

envy and personal abuse. This was the primary nature of the dis¬

pute up to the final week of July, 1637. At that time, the concur¬

rence of a major political event with the quarrel gave strong

political overtones to it. When Queen Anne’s treasonous activities

were disclosed, the possibility that the play might be considered

as a plea on her behalf led to its being condemned. Richelieu’s

motivation in securing this condemnation from the Academy was

therefore entirely political. He was far too busy with affairs of

state to concern himself seriously with the rules of drama or the

petty squabbles of his poets. It was his purely political interest that

pressured the Academy against its will to turn out Les Sentimens.

In all probability, it would never have been produced without the

Cardinal’s insistence because not only were the members of the

Academy adverse to such an undertaking, but Corneille himself

was strongly opposed to it, and the laws of the Academy required

an author’s consent before consideration of his work. Without the

pressure that Richelieu applied to Corneille through Boisrobert,

even the flimsy pretext of consent that the Academy was forced to

act upon would have been lacking.

THE EFFECTS OF HARVESTING AQUATIC

MACROPHYTES ON ALGAE1

Stanley A. Nichols

University of Wisconsin —

Madison

ABSTRACT

A study in the University Bay of Lake Mendota was under¬

taken to determine whether control of aquatic macrophytes, by

means of harvesting, would lead to an increased algal problem. A

significant increase in algal biomass was found 1-year after 2-

years of harvesting, in shallow water areas (depth 1 m). This

increase was concomitant with a decrease in macrophytes. In the

deeper water (average depth 1.3 m) , a drop in macrophyte

biomass was accompanied by a drop in algal biomass. Productivity

differences were also found between the shallow and deep water

areas. In the deep water, peak productivity for algae and macro¬

phytes occurred at nearly the same time. Peak macrophyte produc¬

tivity was accompanied by low algal productivity in shallow water.

INTRODUCTION

The heavy growth of rooted aquatic plants with their attached

filamentous algae constitutes a distinct problem for use of the

lakes in the Madison, Wisconsin area. In proposing management

solutions to the problem, questions must be answered about the

relationships between the algae and the macrophytes. Is there a

mutualistic relationship between them, an antagonistic relation¬

ship, or no relationship? If there is a mutualistic relationship, a

treatment method that would eliminate one member of the pair

should eliminate the other. If the relationship is antagonistic, a

treatment which discourages one member may encourage the other

member. If there is no relationship between the two, one is dealing

with two separate problems.

A quick review of the literature would support the hypothesis

that there is an antagonistic relationship between the two. Hasler

and Jones (1949) conducted experiments to determine if dense

growths of large aquatic plants, in small silo ponds, had a signif-

1 Arboretum Journal paper No. 83, August, 1971.

165

166 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

icant inhibiting* effect on the growth of phytoplankton and rotifers.

With the greatest density of macrophytes, the phytoplankton and

rotifer populations were minimal. Goulder (1969) substantiated the

work of Hasler and Jones with data on the macrophytes and algae

in two ponds in northeast England. In the pond with substantial

amounts of macrophytes, algal productivity rates were very low,

while in the macrophyte free pond, the algal productivity rate

remained high.

Langhans (from Hasler and Jones, 1949) proposed the idea that

an antibiotic may be secreted by the large aquatic plants which

inhibit the growth of algae. Curtis (from Hasler and Jones, 1949)

suggested that perhaps large plants were favored by early spring

conditions and emerge before the plankton. Embody (1928), Ben¬

nett (1942), Wiebe (1934) and later Fitzgerald (1968, 1969a,

1969b) maintained that higher aquatic plants and phytoplankton

are in direct competition with each other for nutrient materials.

Certainly other causes such as different pH requirements, lower

light threshholds for macrophytes, and bacterial action might also

be factors.

The objectives of this study were to determine whether control

of aquatic macrophytes would lead to an increased algal problem,

as predicted by previous workers.

DESCRIPTION OF THE AREAS

This study was undertaken in University Bay, a small bay on

the south side of Lake Mendota, Dane Co., Wisconsin. It lies

between Picnic Point and the University of Wisconsin campus.

The bay is 0.8 km wide and 1.2 km long, with an area of approx¬

imately 106 ha. Lake Mendota has been the site of intensive

limnological investigations for many years. The limnological con¬

ditions are, therefore, well documented. Poff and Threinen (1962)

should be consulted for a review of the limnological conditions

found in the lake. Two specific areas within University Bay were

selected as study sites (Fig. 1). The first site, Stand Z, lies in

shallow water, uniformly 1 m deep, inside a sandbar which would

hinder general circulation with the lake. Stand Y is deeper, with

a depth ranging from 1.0-1. 6 m, and is outside the sandbar.

METHODS

The study was designed to test both the long and short term

effects of harvesting. It lasted 3-years and was carried out in con¬

junction with a larger study of harvesting on macrophyte plants.

For this study, two 30 m by 30 m plots were constructed (Stand

Z and Stand Y), with a 100 m2 sections deleted in the upper right

1973]

Nichols — Effects of Harvesting on Algae

167

FIGURE 1. Hydrographic map of University Bay showing stand locations.

corner of each plot (Fig. 2). The stands were delineated with

plastic coated line attached to looped concrete reinforcement rods.

Each stand was further divided into 100 m- plots. This peimitted

eight different treatments to be tested. After the first year s study,

each plot was divided into two 5 by 10 m subplots. One subplot

was treated, and the other was left as a control to study the ef¬

fect of one year’s harvest. At the beginning of the third year, the

control and treatment plots were reversed so the effect of two

years harvest could be studied.

168 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 2. Plot location and treatment method in each stand, 1968. In 1969

the treatment methods were reversed, i.e., 1A was given the treatment that

1AA was given in 1968. Each plot is 5 by 10m. J — June, JL — July, A — August,

S — September.

The harvest intensities varied from one to three times during

the summer. One plot was harvested three times, once each in

June, July, August. Two plots were harvested twice, one during

June and July and the other during July and August. Three plots

were harvested once either in June, July or August. Any plot

might then be harvested once, twice, or three times during the sum¬

mer of one or two years. Three areas were used as controls and

were not harvested, but were sampled. Figure 2 will better define

the plot size, sampling and treatment methods. It should be noted

that no plot was specifically harvested twice, once in June and

once in July, but this information could be obtained from the

1973] Nichols — Effects of Harvesting on Algae 169

plot harvested three times. Harvesting was accomplished by cutting

all the plant material in the 50 m2 plot, as close to the bottom as

possible, using a sickle or diver’s knife.

At the middle of each month and prior to each harvest, two

random, 1-m2 samples were taken from appropriate harvest and

control areas. The macrophytes and filamentous algae, were col¬

lected in each sampling area. The dry weight, per square meter,

for the macrophyte species and for filamentous algae was ob¬

tained by separating the samples, oven drying and weighing the

tissues.

RESULTS AND DISCUSSION

The short term effects of harvesting were tested with the Pear¬

son product-moment correlation coefficient. No significant corre¬

lations were found between the biomass of algae and the biomass

of macrophytes during the months immediately following treat¬

ment. This would indicate that a reduction of macrophyte biomass

did not have any significant effect on algal biomass during the

year of treatment.

The design was then analyzed to see if there was any signifi¬

cant increase in algal biomass with the long term reduction of

macrophyte biomass. Different treatments, for each month, by

stands, were compared for 1-year after 1-year’s harvesting, and

1-year after 2-year’s harvesting. A Student’s t-test for the signifi¬

cance between two means was used to compare the control with

each treatment by month. The results are summarized in Tables

1 and 2.

TABLE 1: MEAN BIOMASS OF ALGAE IN STAND Z, IN g/m2

June _

July -

August _

September

1-year after 1-year previous harvesting

0.1

28.1

0.1

0.1 0.1

1-year after 2-years previous harvesting

Differences between control and treatment significant at *90% and **95% level.

170 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 2: MEAN BIOMASS OF ALGAE IN STAND Y, IN g/m2

1-year after 2-years previous harvesting

Difference between control and treatment significant at *90% and **95% level.

From these data the following conclusions might be drawn.

In the shallow water stand (Stand Z) there was a significant in¬

crease in filamentous algae in areas that had been harvested heavily

for the previous 2-years. In this area it would appear there may

be an antagonistic relationship between algae and macrophytes.

By decreasing the macrophyte biomass in this area, for a long

period of time, a natural, biological control of filamentous algae

might be eliminated.

In the deeper water stand (Stand Y) an antagonistic relation¬

ship is not apparent. There is less pattern to the significant differ¬

ences and, when they occur, it appears that a decrease in macro¬

phyte biomass leads to a decrease in algal biomass. Here it would

appear that harvesting is a good control measure for both plant

types.

There is a large, but not statistically significant, difference be¬

tween some means. This is primarily due to the unusually large

variance found in these particular populations.

To investigate further the relationship between the two plant

types, the productivity of the algae was compared to the produc¬

tivity of Myriophyllum spicatum L., the major macrophyte species.

Here again a definite difference is noticed between the stands

(Fig. 3). In the deep water areas, the production rate peaks at

about the same time for both M. spicatum and algae. In the shallow

water area the case is just the opposite,

1973]

Nichols — Effects of Harvesting on Algae

171

FIGURE 3. Production rates of Myriophyllum spicatum L. and algae, 1969.

1 — Stand Z, algae. 2 — Stand Z, M. spicatum . 3 — Stand Y, algae. 4 — Stand Y,

M. spicatum. M — May, J — June, JL — July, A — August, S — September. Let¬

ters indicate mid-month.

CONCLUSIONS

From the data it appears that the relationship between the

algae and macrophytes is different between the two stands. These

differences lead to different management recommendations.

In the shallow water areas there appears to be an antagonistic

relationship between the algae and macrophytes. The competitive

action takes place in a period of active growth, macrophytes being

the most successful competitor. Nutrient competition, as proposed

by Fitzgerald (1968, 1969a, 1969b), would therefore be a likely ex¬

planation. If a prolonged intensive harvesting program was used

172 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

in these shallow water areas, an increase in algal biomass would

be expected.

In the deep water area the antagonistic relationship is not

apparent. Here any harvesting method that was beneficial for

reducing macrophyte biomass also appeared to be beneficial in

reducing algal biomass.

ACKNOWLEDGMENTS

The author wishes to express his thanks to Mr. David Nelson

for a basic literature review, to Dr. Grant Cottam for critical read¬

ing of the manuscript, and to Mr. Jere Mossier for starting the

experiment and collecting some of the initial data.

The work was supported in part by the United States Depart¬

ment of Interior as authorized under the Water Resources Re¬

search Act of 1964, Public Law 88-379, OWRR #B-019-Wis.,

Agreement No. 14-01-0001-1968, and the University of Wisconsin

Arboretum.

REFERENCES

1. BENNETT, G. 1942. Management of small artificial lakes. Bull. Ill. Nat.

Hist. Survey 22: 357-376.

2. COTTAM, G. and S. NICHOLS. 1970. Changes in water environment re¬

sulting from aquatic plant control. Tech. Report OWRR B-019-Wis., The

Univ. of Wis. Water Res. Center, Madison, Wisconsin, 27 pp.

3. EMBODY, G. C. 1928. Principles of pond fertilization. Trans. Amer. Fish.

Soc. 58: 19-22.

4. GOULDER, R. 1969. Interactions between the rates of production of

freshwater macrophytes and phytoplankton in a pond. Oikos 20: 300-309.

5. HASLER, A. and E. JONES. 1949. Demonstration of the antagonistic

action of large aquatic plants on algae and rotifers. Ecology 30: 359-364.

6. FITZGERALD, G. 1968. Detection of limiting or surplus nitrogen in algae

and aquatic weeds. J. Phycol. 4: 121-126.

7. - . 1969a. Field and laboratory evaluations of bioassay for nitro¬

gen and phosphorus with algae and aquatic weeds. Limnol. Oceanogr. 14:

206-212.

8. - . 1969b. Some factors in the competition or antagonisms among

bacteria, algae, and aquatic weeds. J. Phycol. 5: 351-359.

9. POFF, R. and C. THREINEN. 1962. Surface water resources of Dane

County, Wisconsin Cons. Dept., Madison. 66 pp.

10. WIEBE, A. 1934. Nocturnal depressions in the dissolved oxygen in fish¬

ponds with special reference to an excess of coarse vegetation and of fer¬

tilizers (Texas). Trans. Amer. Fish. Soc. 64: 181-188.

ANNOTATED LIST OF TRICHOPTERA

(CADDISFLIES) IN WISCONSIN1

Jerry L. Longridge and William L. Hilsenhoff

University of Wisconsin —

Madison

Caddisflies in Wisconsin were first studied by Vorhies in 1908.

He found larvae of 50 species in southern and central Wisconsin,

and suggested that probably 100 species or more were present in

the state. More recent studies by Ross (1938, 1941, 1944, and 1946) ,

Flint (1960), and Yamamoto and Wiggins (1964) list records of

112 species from Wisconsin. An intensive study of the Pine-Popple

River in northeastern Wisconsin yielded 149 species of caddisflies

(Longridge and Hilsenhoff 1972) and demonstrated that previous

records of Wisconsin’s Trichoptera were scattered and incomplete.

Because of the importance of caddisflies as food for fish and

indicators of water quality, a study was initiated in April 1970

to determine the distribution and abundance of the caddisfly

fauna in Wisconsin. Collections were made in nine 24-mile square

study areas (Fig. 1 and Appendix 1) that were selected as repre¬

sentative of the state on the basis of geographical location, soil

type, geology, and vegetation. Light-traps (Longridge and Hilsen¬

hoff 1972) were run at sites along selected streams in these areas

(Appendix 1) and adults were collected from banks of other

streams with a sweep-net. As a result 208 species are now known

from Wisconsin.

The records listed below are for adult male caddisflies, except

when noted otherwise. The study areas in which they were collected

are noted by abbreviations (N, NW, NE, etc.) from Figure 1. Col¬

lections from the Pine-Popple River system (PP) and from

counties outside the study areas are also listed. All specimens

have been preserved in 70% ethanol and deposited in the Uni¬

versity of Wisconsin Insect Collection.

List of Species

RHYACOPHILIDAE

Rhyacophila acropedes Banks — N and NE ; 10 June-26 Aug.

R. fuscula (Walker) — PP; 27 Aug.

R. vihox Milne — EC ; 28 May.

1 Published with the approval of the Director of the Research Division, College of

Agricultural and Life Sciences. Research supported in part by the Wisconsin Depart¬

ment of Natural Resources.

173

174 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 1. Locations of the nine 24-mile square study areas.

GLOSSOSOMATIDAE

Agapetus hessi Leonard and Leonard — NC; 25 June.

Glossosoma intermedium (Klapalek) — N, NW, NC, EC, SW, SE,

SC, PP and Sauk; 28 Apr. -2 5 Aug.

G. nigrior Banks— N, NW, NE, NC, WC, SC, SW and PP; 28

Apr. -13 Sept.

Protoptila erotica Ross — NW, WC, EC, SC, SW and PP; 10 June-

24 Aug.

P. maculata (Hagen) — EC and PP; 10 June and 15 July.

P. tenebrosa (Walker)— N, NW, NE, NC, WC, SC and PP; 6

June-18 Aug.

1973]

Longridge and Hilsenhoff — Caddisflies

175

PHILOPOTAMIDAE

Chimarra aterrima Hagen — NE, EC, SC, PP and Sauk ; 15 May-

13 Aug.

C. feria Ross— NW, NE, NC and PP ; 28 May-27 Aug.

C. oh scar a (Walker)— NW, NE, NC, WC, EC, PP, Langlade

and Sawyer; 21 May-27 Aug.

C . soda Hagen — NE, NC, EC and PP; 3 June-27 Aug.

Dolophilodes distinctus (Walker) — NW, NE, PP and Sauk;

4 Mar.-l Nov.

Wormaldia moestus Banks — Listed in Ross 1944.

PSYCHOMYIIDAE

CyrneUus marginalis (Banks) — NW, WC, SW, Dane and Pepin;

28 July-20 Aug.

hype diversa (Banks)— N, NW, NE, NC, WC, EC, SC, SW and

PP; 15 May-18 Aug.

Neureclipsis himaculatus (Linnaeus) — NC, WC, SC, PP and Por¬

tage; 8 June-13 Aug.

N. crepuscularis (Walker) — NW, NE, WC, SC, SW, PP, Dane

and Outgamie; 9 June-20 Aug.

Nyctiophylax vestitus (Hagen) — N, NE, NC, WC, EC, SC, SW,

SE, PP and Douglas ; 9 June-18 Aug.

Phylocentropus placidus (Banks) — N, NW, NE, NC, WC, EC, PP

and Vilas ; 9 May-27 Aug.

Polycentropus aureolas (Banks)- — N, SC and PP; 25 June-18 July.

P. centralis Banks — N and EC; 10 June and 7 July.

P. cinereus Hagen— NW, NE, NC, WC, EC, SC, SW, SE and PP ;

5 June-26 Aug.

P. confusus Hagen — NC, NE, PP and Sawyer; 9 June-15 July.

P. crassicornis Walker — WC; 15 June, $

P. flavas (Banks) — NE, SE and PP; 9 May-31 July.

P. glacialis (Ross) — Listed in Ross 1944.

P. interruptas (Banks) — NW, NC, WC, PP, Shawano and Wash¬

burn; 15 June-31 July.

P. nascotius Ross — Listed in Ross 1944.

P. pentus Ross — NC and PP ; 7 June-23 June.

P. remotas Banks— NW, NE, NC, WC, SC, SE and PP; 21 May-

17 Aug.

P. weedi Blickle and Morse — NC, SC and PP; 11 June-13 and Aug.

Psychomyia flavida Hagen — N, NW, NE, NC, WC, EC, SC, SE,

PP, Douglas, Langlade, Portage and Sauk; 9 May-13 Sept.

176 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

HYDROPSY CHID AE

Cheumatopsyche analis (Banks) — NW, NE, NC, WC, EC, SC, SE,

PP, Clark, Dodge and Vilas; 3 May-13 Aug.

C. aphanta Ross — EC and PP ; 10 June and 12 Aug.

C. campyla Ross — NW, NE, EC, SC, SW, SE, Dane and Rock;

21 May-20 Aug.

C. gracilis (Banks) — NW, NE, NC, SC and PP; 8 June-17 Aug.

C. minuscula (Banks) — NE, NC and PP; 10 June-15 July.

C. oxa Ross — NW, NE, NC, EC, SC, SW, SE, Outagamie and Por¬

tage; 15 May-17 Aug.

C. pasella Ross — NW, NE, NC and SW; 9 June-17 Aug.

C. sordida (Hagen) — NW and NE ; 20 June -17 Aug.

C. speciosa (Banks) — NW and WC; 15 July-18 Aug.

Diplectrona modesta Banks — NE, NC and EC; 9 June-11 June.

Hydropsyche arinale Ross — Waukesha; 19 Aug.

H. betteni Ross— NW, NC, WC, EC, SC, SW, SE and PP; 21 May-

12 Aug.

H. bidens Ross — NW and Dane; 11 July and 17 Aug.

H. bifida Banks— NW, WC, EC, SW, SE, PP and Barron; 21 May-

18 Aug.

H. bronta Ross — PP; 3 June-12 Aug.

H. cheilonis Ross — NW, NC, EC, SC, SE and PP; 7 June-13 Sept.

H. cuanis Ross — EC and SE ; 21 May and 10 June.

H. dicantha Ross — NC, PP and Price; 23 June and 19 July.

H. hageni Banks — NW and PP ; 28 May-17 Aug.

H. morosa Hagen — NE, NC, EC and PP; 28 May-11 Sept.

H. orris Ross — NW, WC, SC, SW, SE and Dane; 1 June-18 Aug.

H. phalerata Hagen — NW ; 17 Aug.

H. placoda Ross — NW, WC, PP and Dane; 19 June-17 Aug

H . recurvata Banks — WC, Oneida, Portage and Vilas; 21 May-18

Aug.

H. riola Denning — N, NC and WC; 23 June-18 Aug.

H. scalaris Hagen — EC and PP; 15 July and 10 Aug.

H. simulans Ross — WC, SC and SW; 3 May-18 Aug.

H. slossonae Banks— N, NW, NC, WC, EC, SC, SE, PP, Monroe

and Washburn; 21 May-15 Nov.

H. sparna Ross — NW, NE, NC, WC, SW and PP; 28 May-26 Aug.

H. vexa Ross — NE, NC and PP ; 27 May-29 June

H. walkeri Betten and Mosely — NE, NC and PP; 9 June-12 Aug.

Macronemum zebratum (Hagen) — NE, NC and PP; 21 June and

16 July.

Parapsyche apicalis (Banks) — N and SW ; 10 June-5 Aug.

Potamyia flava (Hagen) — WC, SW, SE, Dane and Sauk; 2 May-

3 Sept.

1973] Long ridge and Hilsenhoff — Caddis flies 177

|

HYDROPTILIDAE

Agraylea multipunctata Curtis — N, NW, WC, SC, SE and PP; 21

May-2 Sept.

Hydroptila ajax Ross — NW and EC; 10 Aug.-17 Aug.

H. albicornis Hagen — NE and PP; 9 June-12 Aug.

H. amoena Ross — N, SC and PP; 28 May-25 June.

H. armata Ross-— NW and SE; 21 May-17 Aug.

H. berneri Ross — NW, NE, SC and SW ; 5 Aug -24 Aug. ; $ ’s.

H. consimilis Morton — WC, SE and PP; 15 June-18 Aug.

H. grandiosa Ross- — NE, SE, EC and PP; 9 June-12 Aug.

ti. hamata Morton — PP; 29 May-12 Aug.

H. jackmanni Blickle — NC, WC and SW ; 11 June-23 June.

H. perdita Morton— NW, WC(, EC and SE; 21 May-18 Aug.

H. scolops Ross — NW, NE, NC, EC, SW and SE; 9 June-26 Aug.

H. spatulata Morton — NW, EC and SW; 10 June-18 Aug.; $ ’s.

H. ivaubesiana Betten — NE and PP; 28 May-13 Aug.

H. wyomia Denning — NE; 9 June-20 June.

Ithytrichia clavata Morton — NE and SC; 18 June-17 Aug.

Leucotrichia pictipes (Banks) — NW, WC and PP; 10 June-3 Aug.

May atrichia ayama Mosely— NW, NE, NC, WC and PP; 12 Aug.-

24 Aug.

Ochrotrichia spinosa (Ross) — Listed in Ross, 1944 from North

Lake, Wis.

0. tar satis (Hagen) — WC and SW ; 5 Aug -18 Aug.

Orthotrichia americana Banks — NW, NE, WC, EC, SC and Dane;

9 June-20 Aug.

0. cristata Morton — PP; 15 July.

Oxyethira forcipata Mosely — SE and PP; 10 June and 3 Aug.

O. pallida (Banks) — Listed in Ross 1944.

O. serrata Ross — PP; 10 June-15 July.

Stactobiella delri (Ross) — PP; 10 June-15 July.

8’. palmata (Ross) — PP; 9 June-23 June; $ ’s.

PHRYGANEIDAE

Agrypnia straminea Hagen — PP; 12 Aug.

A. vestita (Walker) — PP; 17 Aug.

Banksiola crotchi Banks — NC, SW and PP; 8 June-15 July

B. smithi Banks — PP; 12 Aug.

Fabria complicata (Banks) — PP; 10 June.

Hagenella canadensis (Banks) — NC, N and PP; 10 June-15 July

Oligostomis ocelligera (Walker) — NC; 28 May and 4 June.

Phryganea cinerea Walker — NW, NC, SE and PP; 8 June-17 Aug

P. sayi Milne — Listed in Ross 1944.

178 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Ptilostomis ocellifera (Walker) — NC and PP; 7 June-28 June.

P. semifasciata (Say) — NE, NC, WC and PP; 28 May-8 July.

LIMNEPHILIDAE

Anabolia bimaculata (Walker) — NC and PP; 15 July-12 Aug.

A. consocia (Walker)— N, NE, NC, WC, EC and PP; 12 June-25

Aug.

A. ozburni (Milne) — N, PP and Dane; 29 June-3 Aug.

A. sordida (Hagen) — Brown; 3 July; 9.

Apatania incerta (Banks) — Listed in Flint 1960.

A. zonella (Zetterstedt) — N; 7 July; 2.

Arctopora pulchella (Banks) — NE ; 28 June; 2.

Asynarchus montanus (Banks) — PP and Dane; 4 June-16 July.

Frenesia missa (Milne) — PP and Sauk; 19 Oct. and 5 Nov.

Glyphopsyche irrorata (Fabricius) — PP; 27 Aug.

Hesperophylax designatus (Walker) — N, NE, SC, SW, SE and PP;

27 Apr.-24 Aug.

Hydatophylax argus (Harris) — NW, NE, NC, WC and PP; 7

June-8 July.

Ironoquia lyrata (Ross) — N; 25 Aug.

I. punctatissima (Walker) — SC and PP; 13 Aug. and 19 Oct.

Leptophylax gracilis Banks — PP; 8 July.

Lenarchulus pulchellus (Banks) — PP; 29 June.

Limnephilus argenteus Banks — PP; 3 June-10 June.

L. canadensis Banks — NC and Clark; 23 June-11 July.

L. externus Hagen — PP; 13 Sept.

L. hyalinus Hagen — N, NC and PP; 12 Aug.-26 Aug.

L. indivisus Walker — WC and PP; 12 Aug.- 13 Sept

L. infernalis (Banks) — PP; 13 Sept.

L. janus Ross — PP; 14 July-12 Aug,

L. minusculus (Banks) — SE; 21 May

L. moestus Banks — N and PP; 10 June-12 Sept.

L. ornatus Banks — PP ; 28 May-12 Aug.

L. parvulus (Banks) — SE and PP; 21 May-29 June.

L. perpusillus Walker — N ; 7 July; 2 .

L. rhombicus (Linnaeus) — SW and PP; 10 June-29 June.

L. sericeus (Say) — N, NC, PP and Door; 10 June-13 Sept.

L. submonilifer Walker — N, NW, PP and Dane; 28 May-30 Sept.

N eophylax autumnus Vorhies — Listed in Ross 1944.

N. concinnus McLachlan — N and PP; 7 July and 13 Sept.

N. fuscus Banks — PP and Manitowoc; 18 May-1 Oct.

N. oligius Ross — NW, NC and Calumet; 17 Aug.-l Oct.

Onocosmoecus quadrinotatus (Banks) — N, NE and PP; 24 Aug.-l 3

Sept.

179

1973] Longridge and Hilsenhoff — Caddisflies

Platycentropus amicus (Hagen) — NW, NE, NC and PP; 17 July-

13 Sept.

P. radiatus (Say) — NC, EC and PP; 23 June-10 Aug.

Pycnopsyche agalona Ross — NC and PP; 26 Aug.-14 Sept.

P. guttifer (Walker)— N, NE, NC, EC and PP; 10 Aug.-l Oct.

P. lepida (Hagen)— N, NW, NE, NC, WC, EC, SC, PP and Grant;

12 May-13 Sept.

P. limb at a (McLachlan) — N, NE and PP; 12 Aug.-13 Sept.

P. scabripennis (Rambur) — NE and PP; 12 Aug.-l 3 Sept.

P. subfasciata (Say) — NW, NE, NC and PP; 12 Aug -13 Sept.

MOLANNIDAE

Molanna blenda Sibley — N, NW, NE, NC and PP ; 9 June-12 Aug.

M. flavicornis Banks — NW, NE, PP, Shawano and Washburn;

9 May-17 Aug.

M. tryphena Betten — N, NW, NE, NC, SC and PP ; 9 June-13

Sept.

M. uniophila Vorhies — NW, NE, NC, EC, SW, PP and Kenosha;

9 June-26 Aug.

ODONTOCERIDAE

Psilotreta indecisa (Walker) — NE and PP; 10 June and 20 June.

LEPTOCERIDAE

Athripsodes alagmus Ross— PP and Dane; 11 July and 15 July.

A. ancylus (Vorhies) — NC, WC and PP; 8 June-15 July.

A. angustus (Banks) — NW, EC and PP; 15 July-17 Aug.

A. annulicornis (Stephens) — NW, NE, NC and PP; 28 May-29

June.

A. arielles Denning — N and NW ; 18 June and 7 July.

A. cancellatus (Betten) — NC and PP ; 23 June and 15 July.

A. dilutus (Hagen) — NE, NC and PP; 3 June-15 July.

A. flavus (Banks) — Listed in Ross 1944.

A. erraticus Milne — PP; 28 May-10 June.

A. mentieus (Walker) — Listed in Ross 1944.

A. miscus Ross — N, NW, SC, Barron, Chippewa, Iron and Por¬

tage; 19 June-11 July.

A. punctatus (Banks) — NW, NC, EC, SW and Dane; 10 June-

20 Aug.

A. resurgens (Walker) — NE and PP; 9 June-12 Aug.

A. tarsi-punctatus (Vorhies) — NW, NC, WC, SC, SW, PP and

Dane; 8 June-18 Aug.

A. transversus (Hagen) — PP; 29 June-12 Aug.

180 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Leptocella albida (Walker)— NW, NE, WC and SE; 3 Aug.-24

Aug. ; $ ’s.

L. Candida (Hagen) — NW, NC, WC, SW and Dane; 11 June-26

Aug.

L. exquisita (Walker) — NW, EC, SC and Dane; 15 July-17 Aug.

L. pavida (Hagen) — PP; 26 Aug.-ll Sept.

Leptocerus americanus (Banks) — NW, WC, SC, SW, PP, Dane

and Rock; 11 June-13 Aug.

Mystacides longicornis (Linnaeus) — NW, NC, Kenosha and Sauk;

10 June-17 Aug.

M. sepulchralis (Walker)— NW, NE, NC, EC, SC, PP, Ashland,

Langlade and Pepin; 3 June-26 Aug.

Oecetis avara (Banks) — N, NW, NE, NC, EC, SC, SW, PP and

Price; 10 June-17 Aug.

O. cinerascens (Hagen) — NW, NC, WC, EC, SC, SE and Dane;

7 June-2 Sept.

O. immobilis (Hagen) — NC and SC; 13 Aug. and 26 Aug.

O. inconspicua (Walker)— N, NW, NE, NC, WC, EC, SW, SE,

PP, Dane and Pepin ; 21 May-2 Sept.

O. ochracea (Curtis) — PP; 15 July-12 Aug.

0. osteni Milne — NW and C; 13 Aug. and 17 Aug.

O. persimilis (Banks) — NW, NC and PP; 23 June-17 Aug.

Setodes incerta (Walker) — NC and PP ; 23 June-12 Aug.

S. oligia (Ross) — PP; 8 July-12 Aug.

Triaenodes aba Milne — Listed in Ross 1944.

T. baris Ross — PP; 29 June-15 July.

T. ignita (Walker) — NW ; 19 June.

T. injusta (Hagen) — NW, NC and PP; 19 June-12 Aug.

T. frontalis (Banks) — SC; 13 Aug.

T. marginata Sibley — PP; 15 July.

T. tarda Milne — N, NW, EC and SC; 10 Aug.-25 Aug.

T. sp. a — NC and PP; 23 June-15 July.

GOERIDAE

Goera stylata Ross — EC, SE and PP; 21 May-29 June.

LEPIDOSTOMATIDAE

Lepidostoma bryanti (Banks) — N, NW, NC, WC, EC, SW, PP and

Iron; 7 June-7 July.

L. costalis Banks — SC; 13 Aug,

L. griseum (Banks) — PP; 12 Aug.

L. sackeni (Banks) — NE, WC, SE, PP and Columbia; 3 Aug.-24

Aug.

L. togatum (Hagen) — NW, NE, NC, SC and PP; 5 June-13 Sept.

1973]

Longridge and Hilsenhoff — Caddisflies

BRACHYCENTRIDAE

181

Brachycentrus americanus (Banks) — N, NW, NE, NC, SW, SC

and PP ; 5 May-27 Aug.

B. lateralis (Say) — PP; 29 May-3 June.

B . numerosus (Say) — NW, SC, SW, SE, PP, Marathon and Sauk;

29 Mar.-3 June.

B. occidentalis Banks — -N, NE, WC, SC, SW, and PP; 29 Apr.-29

May.

Micrasema rusticum (Hagen) — NE, NC and PP; 5 June-23 June.

M. ivataga Ross — PP; 15 July.

SERICOSTOMATIDAE

Sericostoma distinctum (Ulmer) — PP; 8 July-12 Aug.

HELICOPSYCHIDAE

Helicopsyche borealis (Hagen)— N, NW, NE, NC, EC, SC, SW,

SE, PP, Oconto and Portage; 9 June-27 Aug.

Appendix 1. Location of study areas and streams, and dates in

1970 on which light-traps were run.

Northern (N) in Bayfield and Ashland Counties (R4-7W in the

south and all the Bayfield Peninsula north of T47N and east of

R7W) .

E. Fk. Cranberry R. (T50N, R7W, S-20) — 19 June, 7 July,

25 Aug.

Jet. Little Sioux R. and Big Sioux R. — -7 July, 25 Aug.

Jet. Sand Cr. and Racket Cr. — 19 June, 7 July, 25 Aug.

Siskiwit R. (T51N, R6W, S-35)— 7 July, 25 Aug.

Northwestern (NW) in Burnett and Polk Counties (T37-40N,

R15-18W, the St. Croix R. acting as the northwestern boundary).

McKenzie Cr. (T37N, R15W, S-36) — 18 June, 17 Aug.

St. Croix R. (T40N, R18W, S-30) — 18 June, 17 Aug.

Wood R. (T37N, R17W, S-30-14)— 18 June, 17 Aug.

Yellow R. (T39N, R16W, S-2)— 18 June, 17 Aug.

Northeastern (NE) in Forest, Florence and Marinette Counties

(T36-39N, R15-18E) .

Halls Cr. (T39N, R18E, S-30)— 24 Aug.

Johnson Cr. (T39N, R18E, S-14) — 20 June, 24 Aug.

Little Popple R. (T38N, R18E, S-23-24) — 24 Aug.

182 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Peshtigo R. (T36N, R16E, S-33)— 9 June.

South Branch Pike R. (T36N, R18E, S-36) — 20 June, 24 Aug.

Northcentral (NC) in Lincoln, Oneida, Price and Taylor Coun¬

ties (T33-36N, R2-5E) .

Averill Cr. (T33N, R5E, S-21-28) — 26 Aug.

Johnson Cr. (T35N, R5E, S-9-16) — 23 June.

Little Somo R. (T36N, R4E, S-24). 23 June.

North Branch Levitt Cr. (T43N, R2E, S-16) — 8 June, 11

June.

Ottertail Spring (T35N, R1E, S-36) — 8 June.

Scott Cr. (T36N, R4E, S-31) — 23 June, 26 Aug.

Sheepranch Cr. (T33N, R2E, S-15-22) — 8 June.

Sprit and Squaw Cr. (T34N, R4E, S-3) — 23 June, 26 Aug.

Wood R. (T33N, R3E, S-33) — 23 June, 26 Aug.

W estcentral (WC) in Buffalo, Dunn and Pepin Counties (T23-

26N, R11-14W except in the southwest corner where Lake Pepin

forms the border) .

Chippewa R. (T26N, R11W, S-4) — 18 Aug.

Chippewa R. (T25N, R13W, S-21) — 15 June,

Duscham Cr. (T26N, R12W, S-21) — 15 June.

Red Cedar R. (T26N, R12W, S-19) — 15 June, 18 Aug.

Rock Cr. (T26N, R11W, S-15) — 15 June, 18 Aug.

Spring Cr. (T24N, R13W, S-7) — 15 June, 18 Aug.

Eastcentral (EC) in Calumet, Fond du Lac, Manitowoc and

Sheboygan Counties (T15-18N, R7-10E).

Branch Onion R. (T15N, R21E, S-29) — 10 June, 10 Aug.

Mullet R. (T15N, R21E, S-5) — 10 June, 10 Aug.

Onion R. (T14N, R21E, S-10) — 10 Aug.

Brook Town Eaton (T18N, R21E, S-27-26) — 10 June.

Southcentral (SC) in Adams, Marquette and Waushara Coun¬

ties (T16-19N, R7-10E) .

Bird Cr. (T19N, R10E, S-33-34)— 13 Aug.

Carter Cr. (T19N, R8E, S-19-20) — 25 June.

Lawrence Cr. (T17N, R8E, S-32) — 5 May.

Mecan R. (T17N, R10E, S-28) — 25 June, 13 Aug.

Mecan R. (T18N, R9E, S-16)— 13 Aug.

Togatz Cr. (T17N, R9E, S-7) — 25 June, 13 Aug.

Southwestern (SW) in Crawford, Richland and Vernon Counties

(T9-12N, R1-3W, except along the south edge where the Wisconsin

River acted as the boundary).

1973]

Long ridge and Hilsenhoff — Caddis flies

183

Core Hollow Cr. (T10N, R2W, S-14) — 11 June, 5 Aug.

Melancthon Cr. (T12N, R1E, S-3) — 11 June, 5 Aug.

Pier Sprg. (T10N, R1W, S-13) — 5 Aug.

Pine R. (T10N, R1E, S-27)— 5 Aug.

Wisconsin R. (T9N, R1E, S-35) — 11 June, 5 Aug.

Southeastern (SE) in Jefferson, Rock, Walworth and Waukesha

Counties (T2-5N, R14-17E).

Brook Bluff Road (T4N, R17E, S-10) — 27 April, 21 May,

3 Aug.

Scuppernong R. (T5N, R16E, 8-24) — 21 May, 3 Aug.

Spring Whitewater L. (T3N, RISE, S-3) — 21 May, 3 Aug.

Sugar Cr. (T3N, RITE, S-15) — 21 May, 3 Aug.

ACKNOWLEDGMENTS

We wish to thank Dr. Herbert H. Ross, Dr. Glenn B. Wiggins,

Dr. Oliver S. Flint, Jr., and Mr. Toshio Yamamoto for their help

in the identification of species collected in this study. We also wish

to extend our appreciation to Mr. Steven Billmyer who assisted

in collecting many of the caddisflies.

REFERENCES

FLINT, O. JR. 1960. Taxonomy and biology of nearctic limnephilid larvae

( Trichop tera ) , with special reference to Eastern United States. Entomol.

Amer. 40: 1-20.

LONGRIDGE, J. L. and W. L. HILSENHOFF. 1972. Trichoptera (caddis¬

flies) of the Pine-Popple River. Wis. Dep. Nat. Resources Tech. Bull. #54,

pp. 20-30.

ROSS, H. H. 1938. Lectotypes of North American caddisflies in the Museum

of Comparative Zoology. Psyche 45: 1-61.

- . 1941. Descriptions and records of North American Trichoptera,

with synoptic notes. Trans. Amer. Entomol. Soc. 73: 125-168.

- . 1944. The caddisflies or Trichoptera of Illinois. Ill. Nat. Hist. Survey

Bull. 23: 1-326.

■ - . 1946. A review of the nearctic Lepidostomatidae (Trichoptera).

Ann. Entomol. Soc. Amer. 39: 265-291.

YAMAMOTO, T. and G. B. WIGGINS. 1964. A comparative study of the

North American species in the caddisfly genus Mystacid.es (Trichoptera:

Leptoceridae) . Can. J. Zool. 42: 1105-1126.

ORGANISMS, ESPECIALLY INSECTS, ASSOCIATED WITH

WOOD ROTTING HIGHER FUNGI (BASIDIOMYCETES)

IN WISCONSIN FORESTS

J. K. Ackerman and R. D. Shenefelt

Lakeland College — Sheboygan

University of Wisconsin —

Madison

ABSTRACT

A general survey of insects associated with macro-fruiting

bodies of forest fungi was made by collecting 511 lots of fungi

over a four year period which involved 112 species of fungi in

56 genera. Forty-seven of these were specifically associated with

wood and 30 cause damage of importance to the timber industry.

The study was undertaken primarily to determine whether or not

insects might be useful in control of these injurious forms. A total

of 25,379 organisms were obtained from the mushrooms and brack¬

ets, with 15,314 belonging to insects in 13 orders. The insects en¬

countered ranged from visitors to those totally dependent on the

fungus for development. The majority of organisms, though not

necessarily linked to the fungi, were capable of developing in them.

Parasites, predators and microorganisms formed a complete ecolog¬

ical community within one macro-fruiting body. Studies of life

cycles, population development, host destruction, host ranges and

seasonal movement of organisms into and out of fruiting bodies

were made.

To determine whether insects may play a role in regulating the

abundance of forest fungi in Wisconsin and to assess the possibil¬

ities of their being useful as biotic agents for control of wood-

rotters, a preliminary survey of the insects associated with the

fungi was conducted and their relationships with the hosts ex¬

amined. Basidiomycetes, especially the Agaricaceae, Hydnaceae,

Thelophoraceae (soft-bodied mushrooms) and Polyporaceae

(brackets) are the major agents causing heart-rot of trees. It has

been known for over 100 years that their sporophores provide

microhabitats for a variety of arthropods, and that many of these,

especially insects, actually feed upon fungal tissue.

185

186 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Several authors (Anderson 1936; Austin 1933; Barnes and Bux¬

ton 1953; Kessel and Kessel 1939a, 1939b; Pielou 1966; Pielou and

Matthewman 1966; Pielou and Verma 1968; Weiss 1920; Weiss

and West 1920, 1921) have listed insect genera and species found

on or in various fungi. A few have conducted ecological studies

on fungus-related fauna (Falcoz 1921, 1927, 1930; Graves 1960,

1965; Graves and Graves 1966a, 1966b, 1968, 1969; Heatwole

1968; Hubbard 1892; Lawrence 1967; Liles 1956; Pace 1967) and

a specialized vocabulary relating to the fungus microhabitat was

presented by Graves (1960).

MATERIALS AND METHODS

Over a four-year period (1966-69) 511 lots of fungi were col¬

lected in 21 counties (32 sites) in Wisconsin (Figure 1). In the

field the samples were placed in individual plastic bags and upon

return to the laboratory each was transferred to an ice cream

carton provided with a water wick [dental roll inserted through

the bottom] and a Saran-wrap cover. Paper towels were placed

in the cartons to help maintain humidity or to prevent accumula¬

tion of moisture in the bottoms. After six months at room tem¬

perature the fungi were subjected to cold treatment, one week at

40 °F followed by two to four weeks at 0°F, in an attempt to break

any resting or dormant state of the insects. Emerging organisms

were preserved in 80% ethyl alcohol, pinned or used in rearing.

Methods of rearing varied with the organisms involved. Larvae

of fungus gnats (Mycetophilidae) were successfully reared in

petri dishes containing Czapek’s Agar covered with a piece of filter

paper on which a piece of the fungus involved was placed. Petri

dishes containing a layer of slightly moist soil, half of which was

covered by a piece of filter paper, and containing a piece of fungus

were satisfactory for the rearing of Thalymus fulgidius Erickson

(Ostomidae). Glass vials, provided with strings in contact with

water and plugged with cotton, served for the incubation of

individual pupae.

THE HOSTS

Consideration was given to insects on both wood attacking and

non-wood attacking fungi as knowledge of the alternate hosts for

potentially valuable insects was essential. Table 1 gives the fungi

studied, those starred being important to the timber industry. All

of the fungi except Sarcoscypha (#172, 173) belong to the Basidi-

omycetes. The numbers shown for “Location by County” refer to

the numbers given in Fig. 1.

1973] Ackerman and Shenefelt — Wood Rotting Insects

187

0

Figure 1. Distribution of collection sites.

ORGANISMS ASSOCIATED WITH THE FUNGI

The animals occupying or visiting the fungi are presented in

the faunal check list. A total of 25,379 organisms was obtained

from the fruiting bodies. Of this number 10,165 were arthopods

other than insects, i.e., centipedes, millipedes, mites, and spiders,

or were snails. Insects, representing 13 orders, comprised the

remaining 15,214 specimens and were associated with 13 of the

17 families of fungi (Table 2).

vOl-

ec-

ion

Mo.

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

46

47

48

49

50

51

52

53

54

55

56

57

58

59

60

61

62

63

64

65

66

67

68

69

70

71

72

Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

TABLE 1. COLLECTION DATA FOR THE FUNGI.

Col¬

lec¬

tion

No.

73

74

75

76

77

78

79

80

81

82

83

84

85

86

87

88

89

90

91

92

93

94

95

96

97

98

99

100

101

102

103

104

105

106

107

108

109

110

111

112

113

114

115

116

117

118

119

120

121

122

123

124

125

126

127

128

129

130

131

132

133

134

135

136

137

138

139

140

141

Ackerman and Shenefelt — Wood Rotting Insects

TABLE 1. (CONTINUED)

189

Location by county**

5, 8, 11, 15

8, 11

8, 20

8, 10, 16

3, 11

2, 3, 16

10

5

12

10

5, 6, 10

1, 4, 8, 10, 16

2, 10

1, 5, 10, 12, 18

14

2, 5, 10, 12, 16

17

8

2, 10

8

5

1

10

2

8, 10

10

10, 11, 16, 17

2

2, 5, 6, 11, 16, 17

2, 10

8, 16

21

10, 16, 20, 21

20

5

1, 2, 10, 11

1, 10

10

2, 16

2, 8, 16

16, 21

10, 16

8, 16

10

9

2, 8, 12, 16

19

2

16

2

10

1, 2, 3, 5, 9, 10, 11, 12, 16

2

2, 6

7

2, 10

10, 20, 21

16

11

5, 10, 11, 16, 21

11, 16

2

16

2, 3

15

2, 16, 20

3

10

2, 5, 8, 10, 11, 16

10

5

190 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

TABLE 1. (CONTINUED)

^Indicates fungi considered important to the timber industry in Wisconsin (Newman 1919).

**Numbers under “Location by County” refer to numbers on Fig. 1

TABLE 2. RELATIONSHIPS OF THE INSECTA AND THE

FAMILIES OF FUNGI.

1973] Ackerman and Shenefelt — Wood Rotting Insects

191

FAUNAL CHECK-LIST

Listed here are all the animals obtained. The numbers refer to

the ‘‘Collection Number” given in Table 1.

AC ARINA (Mites)

Acaridae : Tyrophagus putrescentiae — 108

Belbidae: Relba sp. — 44

Camisiidae: Camisia sp. — 68

Caraboididae : Undetermined— -43

Ereynetidae : Undetermined — 169

Galumnidae : G alumna sp. — 49

Oribatulidae : Scheloribates sp. — 19

Parasitidae: Undetermined nymphs — 43, 108, 112, 113, 124, 134,

139, 128

Ologamasus sp. — 44

Pergamasus sp. — 10

Zerconidae: Zercon sp. — 50

ARACHNIDA (Spiders)

Agelenidae : — 1

Araneidae: — 2, 17, 46, 116

Clubionidae: — 2, 18, 139

Dictynidae: — 2, 17, 57, 62

Gnaphosidae : — 32

Hahiidae : — undetermined

Linyphiidae: — 15, 37

Lycosidae : — 17

Micryphantidae : — 9, 32, 46, 167

Pisauridae : — 55

* _ 1 V

Theridiidae 9, 17, 37, 38, 55, 57, 69

Thomisidae: — 9, 17, 34, 50

PHALANGIDA (Daddy-long-legs)

Undetermined -17, 40, 74, 124

CHILOPODA (Centipedes)

Geophilidae: Geophilus sp. — 25

Lithobiidae: Nadabius sp. — 38

Neolithobius mordax Koch — 99

Paitobius sp. — 2

192 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

DIPLOPODA (Millipedes)

Julidae : Cylindroiulus sp. — 46, 129

C. caeruleocinctus (Wood) — 113

Ophyiulus pilosus (Newport) — 25, 99

Nemastomatidae : Nopoiulus sp. — 113

GASTROPODA (Snails)

Anguispira alternata (Say) — 17, 27

I) erocer as laeve (Muller) — 10

D. reticulatum (Muller) — 2

Euconulus cher sinus (Say) — -17, 62

Vallonia parvula (Sterki) — 2, 50, 142, 162

Zonitoides nitidus (Muller) — 17, 42, 46, 50, 84

INSECTA (Insects)

BLATTARIA

Blattidae: Parcoblatta sp. — undetermined

COLEOPTERA

Anobiidae : Gator ama semibistriata Mance — 37

Lasioderma serricorne (Fabricius) — 9, 63, 84, 129

Anthribidae: Euparius marmoreus Oliver — 37, 38, 46, 65

Carabidae : Agonium sp. — 2

Carabus vinctus Weber — undetermined

Pterostichus sp. — 5

Chrysomelidae : Gastroidea polygoni (Linnaeus) — unde¬

termined

Ciidae: Ceracis sallei Mellie — 17, 31, 32, 37, 75, 164

C. singularis (Dury) — 32, 62, 164

C. thoracicornis (Ziegler) — 17, 31, 37

Cis americanus Mannerheim — 17, 38, 62, 139

C. fuscipes Mellie — 17, 25, 31, 32, 37, 50, 65

C. levettei (Casey) — 36, 37, 38, 50, 62, 66

C. subtilis Mellie— 17, 32, 37, 38, 65

Dolichocis manitoba Dury — 17, 38, 50

Malacocis brevicollis (Casey) — 32, 37,, 38, 50, 63

Octotemnus laevis (Casey) — -17, 31, 32, 38, 50, 73, 81

Rhopalodontus americanus Lawrence — 36, 57, 62, 124, 143

Sulcacis curtulus (Casey) — 32, 37, 38, 50

Cryptophagidae : Toramus pulchellus LeConte — 139

Cucujidae: Ahasverus advena (Waltl) — undetermined

Curculionidae : Sitophilus oryzae (Linnaeus) — 99

Elateridae: Undetermined — 86, 122, 124, 144

Endomychidae : Lycoperdina ferruginea LeConte — 162, 164,

165

1973] Ackerman and Shenefelt — Wood Rotting Insects

193

Erotylidae : Triplex sp. — 2

T. thoracicornis Say — 1, 2, 3, 53, 126, 132, 139

Mycetophagidae : Tritoma affinis Wickham — 2, 17

Melandryidae : Eustrophinus sp. — 2

Penthe obliquata (Fabricius) — undetermined

Teratoma sp. — 139

Nitidulidae: Glischrochilus quadrisignatus Say — 44

Phenolia grossa (Fabricius) — 46

Orthoperidae : undetermined — 53

Ostomidae: Thymalus fulgidius Erickson — 1, 17, 31, 33, 50,

52, 85

Tenebroides sinuata LeConte — 17

Ptiliidae : Pteryx duvalli Matthews — 37

Staphylinidae : Bryoporus sp. — undetermined

Lordithon sp. — undetermined

Mycetoporus sp. — 2, 17, 19, 52, 57, 74, 102, 103, 109, I'll,

116, 121, 124, 149, 162, 53

N eotrochus sp. — 88

Oxyporus femoralis Gravenhorst — 124

Stenus sp. — 20

T achinomorphus sp. — 2, 37, 46

Tribe Tachyporini sp. — 2, 149

Tachyporus sp. — 9

Tenebrionidae : Hoplocephalus viridipennis (Fabricius) — 62

Boletotherus cornutus Panzer — 37, 60, 62

Diaperis maculata Olivier — 2, 17, 38, 57, 62

COLLEMBOLA

Entomobryidae : Cyphoderus sp.— 2, 9, 32, 37, 44, 50, 63, 74,

114

Drepano cyrtus sp. — 57, 78

Folsomia elongata McGill — 30, 108, 139, 172

Lepidocyrtus pusillus Linnaeus — 14

Proisotoma minuta (Tullberg) — 57

Ptenothrix marmoratus (Packard) — undetermined

Salina sp. — 44, 50, 57

Tomocerus flavescens (Tullberg) — 57

Tomolonus sp. — 57

Poduridae: Podura aquatica Linnaeus — 2, 32, 37, 129, 139

Onychiurus subtenius Folsom — 30, 53, 57

Microgastrura sp. — 2, 31, 118

Sminthuridae : Sminthurides lepus Mills — 171

Sminthurinus niger (Lubbock) — 31, 42

S. quadrimaculatus maynardi Snider — 151, 173

S. quadrimaculatus quadrimaculatus (Ryder)- — 17, 44, 75

Sminthurius facialis Banks — 32, 62, 78, 105, 114, 173

194 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

CORRODENTIA

Thyrsophorus sp. — 32, 36, 37

Caecilius sp. — 36, 37, 78

DIPTERA

Anisopidae : Anisopus alternatus Say — 2, 4, 17, 36, 40, 80, 92,

124

Anthomyiidae : Anthomyia pluvialis (Linnaeus) — 2

Cecidomyiidae : Epimyia Carolina Felt — 50

Johnsonomyia rubra Felt — 31, 50, 116

Henria ( —Leptosyna) quercivora Felt — 116

Miastor metraloas Meinert — 15, 17, 25, 32, 37, 41, 50, 61,

63, 66, 73, 74, 76, 85, 107, 110, 124, 147

Mycodiplosis sp. — 5, 16, 17, 31, 32, 37, 38, 50, 53, 57, 61,

63, 75, 85, 86, 99, 122, 124, 135, 147

Mycophila lampra Pritchard — 20, 31, 32, 37, 46, 50, 53,

62

Neocatocha marilandica Felt — 15, 17, 31, 32, 50, 53, 62,

100, 116

Chironomidae : Procladius sp. — undetermined

Chloropidae : Eugaurax vittatus Sabrosky — 152

Oscinella sp. — 32

O. melancholica (Beck) — 4, 6, 9, 17, 32, 76, 86, 100, 108,

111, 124, 167

Tricimba sp. — 2, 17, 24, 37

T. spinigera Malloch — 4, 17, 27, 109, 116

Drosophilidae : Chymomyza sp. — 101

C. amoena (Loew) — 50

Drosophila sp.— 17, 38, 63, 6, 83, 85, 86, 90, 99, 101, 103,

109, 111, 115, 116, 124, 126, 144, 146, 147

D. busckii Coquillett — 2, 14, 46, 84, 92, 99, 102, 108, 116,

124, 146

D. duncani Sturtevant — 2, 15, 99, 116

D. falleni Wheeler — 4, 6, 9, 10, 15, 17, 20, 56, 82, 83, 85,

86, 89, 91, 101, 102, 103, 108, 109, 111, 114, 116, 118,

124, 125, 128, 135, 139, 144, 146, 147

D. putrida Sturtevant — 4, 17, 37, 57, 82, 124

D. recens Wheeler— 2, 99, 84, 50, 102, 124, 146, 152, 173

My codrosophila sp. — 17, 52, 86

M. dimidiata (Loew) — 17, 50, 52, 124

Muscidae: Cyrtoneuropsis rescita (?) — 2, 101, 102

Fannia canicularis Linne — 2, 6, 17, 46, 84, 86, 89, 107,

108, 110, 124, 125, 144, 146, 154

Graphomya maculata (Scopoli) — 2, 3, 89

Musca domestica Linnaeus — undetermined

Muscina assimilis Fallen — 89

1973] Ackerman and Shenefelt — Wood Rotting Insects

195

M. stabulcins (Fabricius) — 6, 84, 101, 102, 114, 146, 149,

46

Myospila meditabunda (Fabricius) — 89

Ophyra leucostoma Wiedemann — 17, 62, 162

Phaonia bysia (Walker) — 128, 149

Mycetophilidae : Coelosia sp. — 34

Cordyla sp. — undetermined

C. v olncr is Johannsen — 129, 135

Exechia perspicua Johannsen — 18, 44, 87, 88, 112, 133

E. quadrata Johannsen — 18

Megalopelma glahanum var. socium Johannsen — 2, 6, 34

Mycetophila sp. — 75, 129

M. caudata Staeger — 17

M. extenta Johannsen — 109

M. fisherae (Laffoon) — 10, 17, 44, 101, 124, 125, 128,

129, 132

M. jucunda Johannsen — 53, 129

M. scalaris Loew — 56, 93

M. socia Johannsen — 50, 124

M. trichonota Loew — 126

Mycomya (= appendiculata ) hirticollis (Say) — 5, 15, 57

M. ornata Meigen — 15

N euratelia desidiosa Johannsen — undetermined

Rymosia akeleyi Johannsen — 133

R. filipes Loew — 2, 32, 34

R. inflata Johannsen — 2, 17, 32, 36, 37, 52

R. triangularis Shaw — 18, 32, 34, 113, 121

Sciophila quadratula (Loew)— 6, 15, 46, 50, 56, 99, 118

Trichonota sp. — 101

T. triangularis Johannsen — 57, 75, 147

Trichosia sp. — 75

Otitidae : Undetermined — undetermined

Phoridae: Megaselia agarici Lintner — 76, 99, 124, 160

M. fungicola Coquillett — 31, 50, 73, 76, 82, 101

M. longipennis Malloch — 2, 3, 15, 17, 18, 31, 50, 53, 55,

84, 86, 88, 91, 101, 105, 108, 112, 113, 114, 115, 116, 118,

121, 124, 125, 128, 132, 135, 139, 143, 144, 145, 146, 149,

160, 172, 46

Platypezidae : Platypeza anthrax Loew — 1, 2, 4, 10, 25, 37, 52,

55, 72, 75, 84, 86, 97, 108, 111, 124, 137, 157

Scatopsidae : Scatopse fuscipes Meigen — 2, 37, 46, 50

Sciaridae: Bradysia sp. — 2, 17, 25, 31, 32, 37, 46, 50, 113

Lycoria oscellaris — 15

Tipulidae : Dricanoptycha septemtrionis Alexander — 53

Limonia indigence (Osten Sacken) — 31, 124

196 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

L. macateei (Alexander) — 53, 124

L. sociabilis Osten Sacken— 124

Faradelphomyia ( = Oxy discus) cayuga (Alexander) — ■

124

Phalacrocera sp. — 63, 86, 92, 110, 124

Ulomorpha sp. — 62, 124, 149

U. pilosella (Osten Sacken) — 124

Ula longicornis Dietz — 124, 149

U. paupera Osten Sacken — 62, 124, 149

HEMIPTERA

Anthocoridae : Asthenidea temnostethoides Reuter — 17

Orius sp. — 139

Xylocoris galactinus (Fieber) — 2

Aradidae : Aradus sp. — undetermined

Lygaeidae: Ischnorrhynchus resedae (Panzer) — 52

Miridae : Fulvius brunneus (Provancher) — 17, 32

Reduviidae: Empicoris sp. — 38, 55

HYMENOPTERA

Braconidae: Aspilota sp. — 2, 3, 17, 37, 80, 97, 124, 139, 149,

169

Ascogaster sp. — 38

Colast es polypori Mason — 57

Eubadizon sp. — 17

Meteor us sp. — 50

M. betulini Mason — 33

Neoblacus sp. — 32

Synaldis acutidens Fischer — 89

Diapriidae : Aneurhynchus sp. — undetermined

Eulophidae : Euplectrus sp.— 124

Hemiptarsenus sp. — 103

Stenomesius sp. — 32, 37

Formicidae: Lasius sp. — 17

Aphaenogaster (or Myrmica) — 101

Ichneumonidae : — 15, 18, 32, 35, 37, 75, 161

Perilampidae : Chrysolampus sp. — 2, 9, 32, 44, 50, 53, 56, 83,

126

Pteromalidae : Gastrancistrus sp. — undetermined

Moranila sp.— 32

Spalangia sp.- — 17, 62

LEPIDOPTERA

Geometridae : — 2, 88

Heliozelidae : Antispila sp. — 32

Tineidae : Homosetia sp. — 38, 40

Tinea rileyi Dietz — 31, 74

1973] Ackerman and Shenefelt — Wood Rotting Insects 197

ODONATA— 36

ORTHOPTERA

Locustidae : — 75

PLECOPTERA

Taeniopteryx nivilas Fitch — 32

THYSANOPTERA

Cryptothrips rectangularis Hood — 76

Hoplothrips sp. — 46, 53

H. major Hood — 53, 76

Haplothrips subtilissimus Haliday — 100

Roles of the Fauna

Complete ecological communities have formed within the macro-

fruiting bodies of the fungi, with organisms adopting the same

roles and exhibiting the same interrelationships as found on

broader scales elsewhere in our environment. The roles range

from that of the chance visitor (mycetoxene) to total dependence

on the fruiting body for development (mycetobiont) . An inter¬

mediate role is played by the mycetophile which can develop on

or in a fruiting body but is not restricted to it. In addition there

are parasites and predators and microorganisms present, all main¬

taining an intricate balance within the fruiting bodies.

INSECTA

BLATTARIA — In most cases roaches are apparently simply

visitors.

COLEOPTERA — Many beetles have been associated in the past

with various kinds of fungi and 18 families were encountered in

the Wisconsin forms examined. With a few exceptions, the beetles

preferred woody bracket fungi. Members of the Endomychidae,

Elateridae, Erotylidae and Melandryidae exhibited a preference

for the soft-bodied Cantharellaceae, Boletaceae and Agaricaceae.

Their presence in such fungi perhaps may be accounted for by

the fact of more moisture in these microhabitats than in the

brackets or puffballs. The staphylinids, which were both myce-

tophagic and predators, did not discriminate between kinds of

fungi, possibly because their presence depended on available prey.

Ciids were definitely associated with heart-rot causing fungi and

destroyed much of the fruiting bodies with which they were as¬

sociated. The ostomid, Thymalus fulgidius Erickson, was also

extremely efficient in destroying sporophores, especially those of

Daedalea unicolor.

COLLEMBOLA — The literature and ecological associations of

Collembola with fungi were reviewed by Graves (1960). Spring-

198 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

tails are common inhabitants of fungi and have caused consider¬

able damage in commercial mushroom beds (Austin 1933; May¬

nard 1951). They were common inhabitants of many of the fungal

types collected, being especially abundant in older brackets.

DIPTERA — It has been generally known for more than 150

years that many varieties of flies were in some manner associated

with fungi. In 1839 Dufour conducted an extensive study of dip-

teran-fungal associations (Buxton, 1960). In 1841 Canzanelli de¬

scribed life-cycles for some fungus-feeding flies and two species

of Sciaridae were reared on mushroom mycelia (Thomas, 1929).

However, even today very little is really known about these insects,

their specific biologies and the types of associations they enter

into with the fungi ; even their taxonomy is not well worked out.

This order was the most ubiquitous in the present study. Four¬

teen families of flies were reared from most of the fungus species.

With the exception of the Ciidae (Coleoptera) , fly larvae appeared

to effect the greatest destruction of their hosts. Even when they

occurred in relatively tough, leathery brackets the destruction

was noticeable. Most of the flies encountered were at least my-

cetophiles, with several tipulids, mycetophilids, platypezids, chlo-

ropids and scatopsids being mycetobionts. In most cases the life

cycles of the flies have become adjusted to fit with the life-span of

the fungal fruiting bodies. For example, several of the fungus gnats

(mycetophilids) completed their entire development in from two

to three weeks, that being the life-span of the mushroom fruiting

body in which they were developing. Apparently in some instances

rapid decline of the sporophores was associated with microorgan¬

isms introduced by the insects.

HEMIPTERA — The true bugs have been considered as chance

visitors to fungi. Some were predacious, e.g., anthocorids, nabids

and reduviids and appeared to be preying upon fungal inhabitants.

Others such as Aradus sp. and Fulvius brunneus (Provancher)

were fungus feeders and may be completely dependent on the

fungus for survival.

HOMOPTERA — The single cicadellid encountered was a visitor.

HYMENOPTERA — The Hymenoptera formed an integral part

of the fruiting body community. They parasitized mycetophages

and helped to keep their populations under control. Meteorus

betulini Mason (Braconidae) was found parasitizing Thymalus

fulgidius Erickson (Ostomidae: Coleoptera). Aspilota sp. (Bracon¬

idae) seemed to utilize phorid flies as hosts, as did Synaldis

acutidens Fischer. The ants appeared to be merely visitors.

LEPIDOPTERA — Members of this order, as a group, have

not been considered common fungus inhabitants, though some au=

1973] Ackerman and Shenefelt — Wood Rotting Insects 199

thors have found a few associated with fungi (Rehfous 1955;

Forbes 1923; Pielou 1966a, 1966b, 1968; Crumb 1956). However,

with the exception of the geometrid, the Lepidoptera encountered

appeared to be at least mycetophiles.

ODONATA — This was strictly a visitor.

PLECOPTERA — The plecopteron in a fungus was probably a

chance visitor.

THYSANOPTERA — Several species of thrips have been re¬

ported as being fungus feeders (Graves, 1960). A few are pre¬

dacious, but those observed during this study were feeding on the

plants. They were associated with relatively old, partially decayed

brackets in both wet and dry conditions.

CHILOPODA

This group of arthropods occasionally occurs in fungi and they

have been considered to be predacious (Williams, 1928).

DIPLOPODA

Diplopods are known to occur occasionally in fungi, being usually

saprophagous or herbivorous (Williams, 1928). Incidentally, all

the millipedes encountered had been introduced from Europe.

GASTROPODA

Snails are usually considered adventitious in fungal fauna. Six

species were found but their roles in the microhabitats are un¬

known. They appeared to scar the surfaces of soft brackets after

which small flies (midges, drosophilids) laid eggs in the areas

involved.

ACARINA AND ARACHNIDA

Spiders and mites form an important part in the fauna on fungi.

The predatory spiders find readily available prey of many kinds,

so it is not surprising that several families were encountered.

Many mites are phytophagous and others parasitic.

FOOD WEBS

The fungal habitat supports a highly complex community ex¬

hibiting intricate food relationships. Attempts to portray food in¬

terrelationships are usually incomplete because of a lack of data,

and those shown here form no exception. Figs. 2 and 3 illustrate

partial food webs occurring in wood fungi and soil fungi respec¬

tively. The food webs diagrammed are based primarily on the asso¬

ciations encountered during this study.

200 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FuItIus brunneus

Hoplothrlps

FIGURE 2. A partial food web for communities from wood associated fungi.

1973] Ackerman and Shenefelt — Wood Rotting Insects

201

* Lvooperdlnla f erruglneal

Triplex thoraclcomls

Collembola

Coleoptera Larvae

Aradidae

Mvcetoporus sp

Dlptera Larvae

Anlsopus alteraatus

Fannla canlcularls

Scatopse fusclpes

Oph.vra leucostoroa

Otltldae

Cordyla volucrls

Rhymosla f lllpes

Platypeza anthrax

Sclarldae —

Mlastor sp.

h.vcophlla

I"l)ros^>hC^la<^

Staphyllnldae Larvae

Nitldulldae Larvae

FIGURE 3. A partial food web for communities associated with soil fungi.

The wood substrate forms the basal portion of the food web

for wood-associated fungi. It is from this substrate that the fungi

draw their nourishment and eventually fruit. Their fruiting bodies

and spores then comprise the second level of the food web and are

the substrate which supports the faunal community. Fungus

feeders utilize the mycelia and spores. These animals are in turn

preyed upon by various predators and parasitized by mites and

Hymenoptera. A secondary level is also formed by microorganisms

growing in a fruiting body which in turn are fed upon by organ¬

isms sheltering in the fungus and which serve as hosts and prey

for parasites and predators. This same kind of picture may be de¬

veloped around soil fungi, except that the basal substrate is formed

by soil, leaf litter, etc.

202 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

SUCCESSIONAL STAGES

Ecological conditions within a fungal fruiting body do not

remain static throughout the life span of the fruiting body. Graves

(1960) provided a scheme for classifying the successional stages

of the conk microsere. The results obtained here support his

scheme, which worked well for the woody, bracket type of fungal

fruiting body. However, it did not apply to softbodied forms which

have a very short life span. In these, it appeared that most of the

organisms had already entered by the time the fruiting body was

ready to sporulate. From the collections examined it appeared that

only parasites and predators moved into the fruiting bodies after

sporulation had begun.

The stages occurring in soft fruiting bodies could be divided as

follows : Stage 1 — early development, prior to sporulation ; Stage

II — period of sporulation; Stage III — period of liquefaction and

decomposition.

During Stage I females seek out the fruiting bodies for oviposi-

tion. During this stage there may also be some initial larval devel¬

opment. During Stage II fly larvae complete their development and

enter the pupal stage. Beetles may also complete most of their

development during this time, but at a slower rate than the flies.

At this point many insects appear to leave the fruiting bodies to

pupate in the soil or leaf litter in the vicinity of the fruiting bodies.

During Stage III the fruiting body is completely destroyed and

many of the flies and beetles which had pupated emerge. These

successional stages in softbodied fungi may succeed one another

with extreme rapidity and the entire process be completed in a

matter of two to three weeks. Collembola, parasites, predators and

chance visitors can be found during both Stages II and III.

Not only did the fungal fauna change with the condition of the

fruiting body and the species of fungus involved, but it also

changed with the time of year in which the fruiting bodies were

collected. This is exemplified in Fig. 4 which depicts the seasonal

changes in fauna for 183 fruiting bodies of Daedalea confragosa

taken from the same collection site in Sauk County. The numbers

indicate the relative abundance of each organism during the

months involved. As shown, different orders become dominant at

different times, and within orders there is a succession of species.

While the successions appear to be quite definite, they are subject

to a large number of meteorological, physical, chemical and biolog¬

ical factors which it is not possible to analyze at the present stage

of the investigation or indeed without the aid of numerous, well-

controlled experiments directed toward that end. The study of

such successional series would be very important in choosing

insects for biological control programs. In all probability a series

1973] Ackerman and Shenefelt — Wood Rotting Insects

203

FEBRUARY

Cynipidae

Oscinella sp.

Rhymosia inflata

PARASITE

MYCETOPHILE

MYCETOPHAGE

MARCH

Stenomesius sp.

Cynipidae

Graphomyia sp.

Oscinella sp.

Miridae

Neocatocha marilandica

Mycophila lampra

Rhvmosta filipes

Collembola

Cis subtilis

Rhymosia inflata

_ 3 parasite

— MYCETOPHILE

MYCETOPH

MYC^OPHAGE

358

i

77

MAY

Cecidomyliidae |^7 MYCETOPHILE

Cis subtilis -------- MYCETOPHAGE

JULY

Oscinella sp.

Pteromalidae

Perilampidae

Mycetophilidae

Rhymosia triangularis

Cecidomy iidae

Antispila sp.

Octotemnus laevis

C orrodentia

Sciaridae

Cis fuscipes

Sulcacis curtulus

Collembola

Cis fuscipes

Malacocis brevicolis

Pteromalidae

Ceracis sinqularis

CeraciS Sallei

OCTOBER

NOVEMBER

H2

^12

— — — — — — — MYCETOPHAGE

PARASITE

rzs r ^Mvr ftp

/ PHAGE

156

FIGURE 4. Faunal succession in fruiting- bodies of Daedalea confragosa.

(Based on 183 fruiting bodies).

204 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

of insects, active at different times of the year within Stage I for

soft-bodied forms and Graves’ Stages I and II for bracket fungi

would be needed.

CONCLUSIONS

Fungal fruiting bodies form complete and highly complex

ecological communities. Several groups of insects showed potential

as possible biological control agents for some timber destroying

fungi and should be studied in greater detail. These include Ciidae,

Ostomidae (especially T. fulgidius) , Tenebrionidae (especially B.

cornutus) , Drosophilidae, Muscidae and Mycetophilidae. Detailed

studies of faunal successions related to stages of decay of the fruit¬

ing bodies and to season of the year should be conducted.

ACKNOWLEDGMENTS

Research supported by the Research Division, College of Agri¬

cultural and Life Sciences, University of Wisconsin and by the

Guido Rahr Foundation, the Wisconsin Department of Natural

Resources and the University Research Committee.

Deep appreciation and thanks are expressed to:

R. F. Patton, project advisor; M. P. Backus and D. Myron for

assistance and guidance in naming fungi; the many individuals

and organizations who so graciously aided through either identify¬

ing or verifying identifications of the insects : D. M. Anderson,

R. E. White, G. B. Vogt, P. M. Marsh, D. L. Wray, W. W. Wirth,

C. W. Sabrosky, G. C. Steyskal, J. M. Kingsolver, Wm. Shear,

D. M. Wheeler, T. E. Woodward, W. Robinson, R. P. Narf, B.

Branson, E. Cook.

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AUSTIN, M. D. 1933. The insect and allied fauna of cultivated mushrooms.

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BUXTON, P. A. 1960. British Diptera associated with fungi. III. Flies of

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CRUMB, S. E. 1956. The larvae of the Phalaenidae. Tech. Bull. U.S. Dept.

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VACATION RESORTS IN ONEIDA COUNTY

(WISCONSIN) A Study of 1950-1968 Trends and

Owner-operator Characteristics

L. G. Monthey and Daniel Zielinski

University of Wisconsin —

Madison and W aukesha

INTRODUCTION

Oneida County, located in the Northern Highland Region of

northeast Wisconsin, has long been part of a well-known, concen¬

trated resort area. Together with Vilas County, just to the north,

this prime recreation-resort district embraces almost 30% of

Wisconsin’s named lakes and has included over 25% of the state’s

vacation-resort establishments.

This is one of the most scenic counties of Wisconsin. Bedecked

with millions of evergreen trees and more than 1,100 sparkling

fresh-water lakes, Oneida County has attracted sportsmen, vaca¬

tioners, and recreational travelers since the 1880’s, when the

first “tourist resorts” were established here. A few of the p re-

1900 establishments are still in existence and still attracting vaca¬

tioners. Some of the earliest resorts were developed on the sites

of old logging or lumber camps and from pioneer farmsteads, when

the whole area was referred to as the “Cutover Country.” Their

location was also determined, to some extent, by the early high¬

way pattern (a meager “network” of dirt roads and sand trails)

and by railroad lines, which constituted the main transportation

system of that area prior to 1920 and continued to be important

for at least three decades thereafter. Rhinelander, the County’s

principal city and its county-seat town, has been an important

railroad center for approximately ninety years. It was only

natural that some of the original resort establishments would

locate on the larger, more attractive lakes near railroad stops

or reasonably close to the few good roads of that pre-automobile

era. Most of the resorts, however, were started during the 1920’s

and 1930’s, the period between the two world wars. A majority

of these are still in existence today.

A “landmark” study by C. W. Loomer and associates of the

University of Wisconsin showed that Oneida County had more

than 600 resort establishments in 1950. These investigators clas-

207

208 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

sified them by type of business, and they reported that 90%

were of the self-service (housekeeping) variety. Recent studies

by the authors and others have shown that, despite sizeable de¬

clines in total resort numbers, the proportion in the self-service

category — mostly seasonal resorts which usually provide little more

than lodging plus a boat — is still approximately 90%.

Despite its continued importance in the tourist-recreation in¬

dustry of Wisconsin, Oneida County has “lost” more resorts since

1950 than any other county in the state. From Loomer’s original

total of over 600, the number had dropped to 438 establishments by

the year 1968. A more detailed study of the nature and causes

of this trend seemed appropriate.

Why was the decline in numbers so much greater here than in

most other resort areas of the state? Also, why have the trends

here been somewhat different from the general trends in the

statewide lodging business? For example, Oneida County’s tourist¬

housing capacity dropped almost 14% during the 1960’s, while

the state’s over-all capacity (in Bedroom Units, or B.U.) declined

scarcely at all. Also, whereas the state showed a 15% increase in

the number of motel-type establishments during the 1961-68 pe¬

riod, Oneida County had a gain of 50% !

The basic data for this report were obtained from mailing lists

and inspection records of the Department of Health and Social

Services. The methods used to compile and classify the data on

resorts and other T-L establishments were similar to those em¬

ployed in previous studies (1, 2). The information on owner-

operator characteristics, opinions and related topics was obtained

through extensive field interviews and surveys conducted by

Zielinski in 1967 and 1968.

Detailed data on the number, type, seasonality, and bedroom-

unit (B.U.) capacity of Oneida T-L establishments, including the

changes between 1961 and 1968, are presented in a series of appro¬

priate tables and graphs. Several of these relate specifically to

resort-type establishments. Pertinent data, as well as significant

changes or trends derived therefrom, are discussed briefly. Part

II of this report presents a summary of the more interesting

findings in the owner-operator study.

RESULTS AND DISCUSSION

Oneida County, like many other resort areas of Wisconsin, has

undergone some substantial changes in the type and number of

tourist-lodging businesses since 1950, mainly within the last 10

years.

As previous studies have shown, the number of smaller Wis¬

consin establishments, mainly the tourist courts and cottage re-

1973] Monthey and Zielinski — Vacation Resorts 209

sorts, has declined rather markedly. The number of motel-type

establishments has generally increased. Statewide, there has been

a sizable increase in the number of establishments with more than

100 bedroom units and a sizable decrease in the number of those

with less than 20 B.U. However, despite a net decrease of about

12% in the total number of establishments, Wisconsin’s capacity

to house visitors has remained virtually unchanged since 1958,

when it was approximately 80,000 B.U. in total.

Oneida County has been, figuratively speaking, a “bellwether”

in the so-called resort industry of Wisconsin. The resort business

has showed more change since 1960 than any other segment of

the tourist-lodging industry, with the possible exception of the

motel business.

Changes in the resort business are characterized mainly by

losses in the number of establishments, particularly in the smaller

cottage-resort enterprises. Oneida County appears to be in the

forefront of this trend, since it has lost over 23% of its resort-type

establishments (and almost 19% of its resort-guest capacity)

since 1961, as this study will show.

First, however, let us examine the overall changes in the

tourist-lodging industry of Oneida County. The total number of

T-L establishments declined about 200 between 1950 and 1968,

and approximately 160 of these were identified as “resorts.” For

our purposes, a resort is defined as a T-L facility (excluding

campgrounds and trailer parks) that is located in or near a rec¬

reational or scenic area. However, in practice this definition is

difficult to use because of the many types of establishments that

would fit such a broad interpretation. Thus, we have relied upon

the owner-operator’s designation for his business. If he (or she)

labeled it as a “resort” or resort-type establishment, we have

placed it in this category also. The same rule was applied to the

other classifications of tourist-lodging establishments (e.g. motels)

used herein.

As mentioned earlier, the major changes or trends in the num¬

ber and type of T-L establishments, both statewide and in Oneida

County, occurred during the 1960’s. The 1961-1968 data for

Oneida County are summarized in Table 1 for the four main

classes of T-L establishments. During this eight-year period, the

County lost two hotels but gained 12 motels — reflecting a net in¬

crease of 211 B.U. for the two categories. Meanwhile, there was

a loss of 134 resort-type establishments and 16 other-type busi¬

nesses, resulting in a net decrease of 1,092 B.U. for these two

categories. Thus, Oneida County showed an over-all decrease of

140 T-L establishments and 881 B.U. during the period. This was

primarily in the resort category, where the loss of establishments

210 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 1. OVER-ALL SUMMARY CHANGES IN THE TYPE AND B.U.

CAPACITY OF ONEIDA COUNTY TOURIST-LODGING

ESTABLISHMENTS BETWEEN 1961 AND 1968.

ran 23% compared to 14% for the state as a whole. Taking all

T-L classes into account, the decreases were 20% and 12%,

respectively.

The T-L establishments of Oneida County are highly seasonal

in nature, running 90.2% in 1961 compared to a statewide average

of 74.3% for all Wisconsin T-L businesses that year. (A seasonal

operation, by definition, is one which stays open for business less

than nine months in a given year.) It is interesting to note that

the degree of seasonality among Oneida County establishments

changed only slightly during the eight-year period, dropping from

90.2% to 89.0%. In other words, the percentage of year-round

establishments increased from 9.8% to 11.0% in this county, com¬

pared to 25.7% to 26.8% statewide. Both changes are of small

magnitude but show the same general trend.

In terms of B.U. capacity, however, the trend toward year-round

operations has been somewhat more pronounced. In Oneida

County, the percentage of bedroom units available on a year-round

basis rose from 14.1% to 17.1% during the 1961-1968 period,

compared to 40.7% to 46.5% for the state.

As Table 2 indicates, however, Oneida County showed signifi¬

cant declines in both seasonal and year-round establishments of

the resort-type category. There were also net decreases in the B.U.

capacity in both cases, although the decline in seasonal B.U. was

considerably greater than that for year-round B.U. Table 2 shows

additional detail on the seasonality of various types of T-L estab¬

lishments in Oneida County. It is noteworthy that, although the

county had a net decrease of only 10% in year-round establish¬

ments during the 8 years (with even a small gain in B.U.

capacity), it lost nearly 21% of its seasonal T-L businesses, pri¬

marily the resort types.

With respect to the general trends in this county’s T-L industry,

it is of interest to compare what has happened here with the state¬

wide changes and trends during the same period of years. Tables

1973]

Month ey and Zielinski — Vacation Resorts

211

TABLE 2. CHANGES IN SEASONALITY AND B.U. CAPACITY FOR

THE VARIOUS TYPES OF TOURIST-LODGING ESTABLISH¬

MENTS IN ONEIDA COUNTY (1961-1968).

3 (a) and 3 (b) summarize information on the number and ca¬

pacity of various types of T-L establishments, with 1961-1968

state data and the county figures side by side.

First, we can readily see that the per cent changes vary con¬

siderably between this county and the state as a whole. Table 3 (a)

presents data on the changes in number and type of establish¬

ments. In Oneida County the net decrease of 10% in hotels of the

traditional type is less than one-half of the average statewide

decline. While hotels are of minor importance in the county’s T-L

economy, they are nevertheless “hanging on” better, and they

continue to provide over one-third of Oneida’s year-round lodgings

(B.U. capacity).

This County’s 50% gain in motel-type establishments during

the 8 years was over three times the statewide gain. Although

they still comprised only 6.3% of the Oneida’s T-L establishments

in 1968, the 36 motels already were providing more year-round

B.U. than 438 resorts during that year. In fact, the hotel-motel

community of Oneida County (less than 10% of the total busi¬

nesses) was offering 67% of the available year-round units —

which are so important to fall and winter trade!

With respect to the number of resort-type establishments, the

eight-year decrease in Oneida County was almost double the

average statewide decline — 23.4% compared to 13.9%. Almost

98% of those lost were seasonal cottage-type establishments, aver¬

aging about 7 B.U. per resort. The fourth category, “Other” es¬

tablishments, which comprises about one-seventh of Oneida Coun¬

ty’s T-L businesses, declined at a uniform rate of about 17%

both statewide and in the county during the eight years.

212 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 3 (a). CHANGES IN NUMBER AND TYPE OF TOURIST¬

LODGING ESTABLISHMENTS IN ONEIDA COUNTY COM¬

PARED TO STATEWIDE TRENDS FOR 1961-1968

All in all, Oneida County lost 19.7% of its T-L businesses be¬

tween 1961 and 1968, while the average statewide decline was

11.6%. Thus, Oneida County’s rate of decrease was 1.7 times

greater than the statewide rate, and accounted for approximately

one-sixth (15.7%) of all T-L business “drop-outs” in the entire

state during the 1960’s.

Table 3 (b) shows the changes in B.U. capacity for various

types of T-L establishments, including the total, for both county

and state during the 1961-1968 period. Here again, percentage

changes in Oneida County vary widely from the average state¬

wide changes. Whereas the total B.U. capacity of all Wisconsin

T-L establishments has changed very little (only —0.7% since

1960), Oneida County lost 13.7% — or about one-seventh of its

capacity.

Looking ahead, it is quite possible that new motel-type enter¬

prises, which have already shown significant increases in the

county, will fill this housing “gap” eventually. However, it may

take ten years or longer, because of concurrent declines in the

capacity of resort-type facilities. During the 1961-1968 period,

Oneida County motels gained approximately 230 B.U., whereas

TABLE 3 (b). CHANGES IN BEDROOM-UNIT (B.U.) CAPACITY FOR

VARIOUS TYPES OF LODGING ESTABLISHMENTS IN ONEIDA

COUNTY COMPARED TO STATEWIDE TRENDS FOR 1961-1968.

1973] Mont hey and Zielinski — Vacation Resorts 213

resort-type establishments lost nearly 1,000 B.U. Meanwhile,

county hotels and other-type establishments (the two remain¬

ing classes) showed a net decrease of about 130 B.U. or 14.7%.

These figures, taken together, account for the overall decrease of

881 B.U. during the eight years.

It is interesting to note the compositional changes in Oneida

County’s T-L industry during the 1960’s and how these changes

differ from comparable statewide trends. Figure 1 illustrates the

compositional changes for state and county establishments in

graphic form.

The percentage makeup of the B.U. composition in the county

differs more widely from state averages than for establishments

only. In 1961, only 18% of Oneida County’s tourist-housing

capacity was in non-resort facilities compared to 50% for the

state. Eight years later, the corresponding percentages were 23%

and 55%, respectively. The biggest change, of course, was in the

percentage of total B.U. provided by motel-type establishments.

This segment of the industry increased by 55% for the state as

a whole, while it virtually doubled in Oneida County. However,

only one-tenth of the county’s visitor-housing units were pro¬

vided by motels in 1968, compared to about 25% of the statewide

total. Figure 2 illustrates the above changes and differences in

B.U. capacities.

Although the trend toward a higher percentage of year-round

B.U. is quite evident in both the state and county data, the pro¬

portion of such units (in relation to the total B.U.) was much

smaller in Oneida County than for the state in both 1961 and

1968. This relationship is portrayed graphically in Figure 3,

which suggests a need for more year-round accommodations in

the Oneida area.

If recent past trends continue, these new year-round establish¬

ments are likely to be of the motel or motel-hotel type. Only a

handful of new resorts have been built in Oneida County during

the 25 years since World War II. However, quite a few of the

older places — perhaps as many as 50 percent— have either ex¬

panded or “modernized,” or both, since 1950. Despite this fact,

the number of small- and medium-sized resorts has dropped

sharply since 1961. Actually, the rate of decline has been accel¬

erating since 1964, as the statistics show when we examine them

on a year-by-year basis for both the county and the entire state.

Table 4 (a) shows the 1961-1968 changes in the number of

resort-type establishments by size class for both Oneida County

and the state. No attempt was made to separate the resorts into

type categories, e.g. self-service (“housekeeping”) vs. American

Plan, European Plan, etc. However, recent observations and sur-

214 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Number of Establishments

FIGURE 1. Changes in the establishmental composition of the Tourist-Lodging

Industry between 1961 and 1968, comparing Oneida County trends with state¬

wide changes.

Wisconsin (Statewide) Oneida County

1973]

215

Month ey and Zielinski — Vacation Resorts

Total Bedroom Units (thousands)

r\> c* -p- vji o>

o o o o o o

- 1 — — - f — - 1 - 1 - r

Hotels

Motels

Resorts

Others

Total Bedroom Units (thousands)

-* rv> -P" ' vji o^

1 - — - f ’ i r J t

Hotels

Motels

Resorts

Others

FIGURE 2. Changes in the Bedroom-Unit composition of the Tourist-Lodging

Industry between 1961 and 1968, comparing Oneida County trends with the

statewide changes.

Wisconsin (Statewide) Oneida County

216 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

19 6 8

FIGURE 3. Proportional-scale graphs showing the per¬

centage of year-round accommodations (B.U.) in both

1961 and 1968, comparing statewide data with that for

Oneida County (see inset).

1973] Monthey and Zielinski — Vacation Resorts 217

veys have shown that less than 10% of the resort-type establish¬

ments — both Oneida County and statewide — provide full restau¬

rant services, although 17% of Wisconsin’s 4,100 resorts had a

restaurant permit in 1968. Presumably, about half of this group

obtained the annual permit in order to sell coffee, sandwiches, and

other snacks to their clientele.

As mentioned earlier, Oneida County lost almost one-fourth

of its resort-type establishments between 1961 and 1968, com¬

pared to a statewide decrease of 14%. However, there was no

consistent trend among the various resort-size categories, except

that large resorts (30 B.U. and higher) showed the smallest

numerical and percentage declines, and the medium-sized resorts

(10 to 29 B.U.) showed somewhat smaller declines than the small¬

sized group (1 to 9 B.U.). These two general trends pertain to

both the county and statewide figures during the 8 years. The

net decrease in Oneida County (135 establishments) showed the

following 1961-1968 losses : 99 small-sized resorts ; 35 medium¬

sized ; 1 large-sized. It is interesting to note that the first two

figures are almost exactly 20 percent of the statewide losses in

these two major size groups. In the large-sized category (30 B.U.

and up), Oneida County dropped one and the state gained one —

or virtually no change.

The small-sized resorts were divided into two categories; (1)

those with less than 5 B.U. and (2) those with 5 to 9 B.U.

[Table 4 (a)]. This separation brings out an important difference

between the county’s trend and the statewide change. The num¬

ber of very small resorts in Oneida County dropped at a rate

three times greater than the statewide decrease between 1961 and

1968, while the decline in the 5 to 9 B.U. group was almost identi¬

cal to the statewide decline. Why? Probably the most likely reason

is that the older and smaller resorts in this “mature,” highly

scenic recreation area suddenly became a hot commodity in the

real-estate market during the 1960’s. Several realtors verified this

supposition and further stated that these very small resorts were

not only the least profitable ones to operate but they also carried

the smallest price tags. Hence, more of them were put up for

sale, and there were more potential buyers in this lower-price

range. Many of these establishments (or the land areas they

once occupied) have since been converted to residential uses,

either to substantial summer dwellings or to year-round homes.

This county has been a leader in second-home developments since

1960.

There is one other significant difference in these data on trends

for the county and the state. The medium-sized group (10 to 29

B.U.) showed a 1961-1968 decline of 17.7% in Oneida County

218 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

compared to a statewide decrease of 11.4%. This category was

also separated into two groups; (1) those with 10 to 19 B.U.,

and (2) those with 20 to 29 B.U. In comparing county vs. state¬

wide trends for 1961-1968, the first group followed the state¬

wide pattern rather closely, showing losses of 16.7 % for the

county and 12.5% for the state. However, when we examine the

20 to 29 B.U. segment, we note a loss of 22.2% for the county

and of only 5.5% for the state. Aside from the fact that group 2

is much smaller numerically than group 1, there is some difficulty

in explaining the difference between county and state for this

medium-to-large category. One possible reason could be the ap¬

parent high mortality rate in small American-Plan resorts, par¬

ticularly those which had a limited capacity and served fewer

than 75 guests in total, since 1960. Perhaps the greater loss of

medium-size resorts in Oneida County is associated with a higher

percentage of these “ailing” establishments; this needs to be

checked out in more detail.

Table 4 (b) is similar to the foregoing one, except that it

shows the resort trends in bedroom units rather than number of

establishments for each size category. It would appear that Oneida

County's larger resorts have decreased in average size, while

statewide statistics reflect a 10% increase in average size (from

43 B.U. to 47 B.U.) for resorts in this category.

The other significant change in resort B.U. capacity involves

the small-resort category (less than 10 B.U.). Statewide, the

B.U. capacity of these small operations declined 16%, while the

county totals showed a 25% drop. As Table 4 (b) shows, most of

the county’s B.U. loss in this group involved the very small es¬

tablishments (1 to 4 B.U.).

TABLE 4 (a). CHANGES IN NUMBER OF RESORT-TYPE ESTABLISH¬

MENTS (BY SIZE CLASS) IN ONEIDA COUNTY (1961-1968), COM¬

PARED TO STATEWIDE CHANGES FOR THE SAME PERIOD.

1973] Monthey and Zielinski — Vacation Resorts 219

TABLE 4 (b). CHANGES IN B.U. CAPACITIES FOR RESORT-TYPE ES¬

TABLISHMENTS (BY SIZE CLASS) IN ONEIDA COUNTY, COMPARED

WITH STATEWIDE CHANGES FOR THE SAME PERIOD (1961-1968).

Part II Owner-operator Study

In view of the foregoing data and the trends derived there¬

from, one might reasonably ask the following questions:

Why have numerous resorts gone out of business in this scenic,

long-established resort area? What social, personal, and economic

factors seem to have a bearing on this striking decline in resort

numbers? What type of operators, in general, have been managing

Oneida County resorts in recent years? What happens to the land

and buildings used by these establishments after they are closed?

In an effort to get some answers to these questions and to

gather more facts on the traditional “resort industry,” several

researchers have conducted a variety of studies in this area. The

work of Zielinski (1966-1970) in Oneida County is appropriate

to this report, since it includes data from active resorters, former

operators, town assessors, realtors, and county officials.* The

information in this section of the paper is based to a considerable

extent on his findings and conclusions.

A group of 83 “retired” resorts was studied by interviewing

the former operators and/or their spouses. This group repre¬

sented about one-half of the county’s 162 resort “drop-outs” dur¬

ing the 1950-1967 period. These people were asked why their

establishments went out of business. Their responses to this

question are summarized in Table 5.

As can readily be seen, most of the reasons are of an economic

nature. Even the leading answer given, i.e. old age, illness or

death of the operator (or spouse), suggests economic considera¬

tions too, since it appeared that no other person was able or

willing to take over and continue the resort as a business. It is

* Zielinski’s full report has been submitted to the Department of Geography, Uni¬

versity of Wisconsin — Madison, as Thesis for a Ph.D. degree.

220 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 5. THE REASONS GIVEN BY OPERATORS FOR THE SALE OR

DISSOLUTION OF SPECIFIC RESORTS IN ONEIDA

COUNTY DURING THE PERIOD 1950-1967.

*In some cases, the information was obtained “second hand” from town assessors or nearby resort

operators.

also likely that many of these entrepeneurs viewed their resort,

not only as a business, but also as a personal service for guests

who had become good friends down through the years. Often¬

times, this type of a clientele will tend to “go its own way,” when

death occurs and/or ownership changes.

When these resorts went out of business, they were frequently

subdivided for other uses. About 70% of these 162 Oneida County

resorts were either parcelled or fully subdivided for use as sum¬

mer cottages or year-round homes. Approximately one in ten be¬

came private “clubs” for the family and its special guests, while

another one-tenth had old facilities removed and the land cleared

for subsequent redevelopment (usually by the next owner). The

final group of inactive resorts, about 9 percent, remained in es¬

tate or was otherwise idle.

In order to shed light on certain other questions, particularly

the owner-operator characteristics of current establishments, Zie¬

linski conducted an interview-survey of 103 Oneida County re¬

sorts during 1967. This sample was carefully selected so as to

be quite representative of the county’s resort industry.

A significant number of the 103 resort operators interviewed

were found to be relatively “new” owners. One in eight had

owned their establishments 3 years or less, and almost one-third

reported ownership of less than 7 years. On the other hand, al¬

most two-fifths (39%) of them had owned their present resort

property for 16 years or longer. The latter group included 25

owner-operators who had built their present facilities ; some dur¬

ing the late 1940’s and early 1950’s, but others in the 1930’s or

earlier. Thus, it appears that 70% of the resort operators in

Oneida County would fall into one of two main categories, which

are almost equal in size : ( 1 ) short-term owners — 6 years or less ;

1973]

Month ey and Zielinski — Vacation Resorts

221

(2) long-term owners — 16 years or longer. The mean length of

ownership for the 103 respondents was 13.3 years.

About one-fourth of the owner-operators surveyed had originally

purchased (or otherwise acquired) their present resort property

without a concrete recreational or business objective in mind.

When interviewed, this group mentioned such things as (1) a

good retirement project, (2) a healthful outdoor life, (3) a good

investment opportunity, and (4) a chance to live in Northern

Wisconsin, as their reasons for acquisition.

Relatively few “young” people were found in this sizable

sample of Oneida County operators, and a fairly high percentage

of them were either retired or semi-retired. Their ages ranged

from 33 to 80 with a median age of 57 years. Of the 100 who

answered this query, 47% were 61 years or older, 28% were

68 or older, and 6% were 74 or more. One out of 10 said they

were semi-retired and 29% indicated full retirement. Neverthe¬

less, a high percentage of the group reported supplemental income

through outside employment or from other sources.

Only 26% of these operators said that the resort business was

their major occupation on a full-time basis. Almost one-half

(46%) indicated that it was their principal occupation but said

they had part-time employment elsewhere. Over one-fourth of

them (28%) had a major occupation other than the resort busi¬

ness. Most of the part- or full-time work was located in or near

Rhinelander, the largest city of the area, and it is interesting

to note the types of “off-resort” employment involved. Personal

services (27%) and factory work (25%) accounted for over one-

half of the job opportunities utilized. Other types were: business

and retail sales — 14.6%; professional and clerical — 12.5%; con¬

struction — 8.3%; logging and local government — 4.2%; others —

8.3%.

Of the 103 owner-operators interviewed in this 1967 survey,

almost two-thirds lived on the resort property throughout the

year. In fact, there appears to be a definite trend toward year-

round residency. Virtually all of these resorts (98%) were owner-

operated establishments, and there was definite absentee owner¬

ship in only one case.

The prior occupations of present resort operators were de¬

termined in this study, and they were found to vary greatly.

Almost one-half of the group (48.4%) had a background in fac¬

tory work or in business (mostly retail trade). The following

list shows the principal prior vocations of 95 Oneida County

operators :

222 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Factory workers (hourly) _ 26.3

Business and retail trade _ 22.1

Personal services _ 14.7

Construction _ 7.4

Sales and advertising _ 7.4

Professional (includes teaching) _ 7.4

Military and administrative _ 6.4

Farming and odd jobs _ 4.2

* Others _ 4.1

100.0

* Oddly enough, only 2 operators said they had been associated with another tourist-

type establishment before buying their own business.

When queried as to their sources of professional help or in¬

struction in resort operations, three-fourths of these operators

said they had received none. Of the remainder, 24 indicated some

help from University Extension (three who said “resource agent”

are included), while three others mentioned commercial sources.

Within this group, 22 stated that the instruction they received

dealt with either modernization of facilities or housekeeping and

management methods.

This group of 103 resorters was also asked to list the skills

and personal qualities required for operating a successful resort

business. Surprisingly, only 14 suggested business and manage¬

ment skills as necessary ingredients. Two-thirds of the operators

said it was largely a matter of good personal relationships with

the clientele. About two-fifths mentioned the quality of being a

“jack of all trades” (repair and maintenance), while one-eighth

stressed cleanliness. Only two operators felt that prior experience

in the hospitality business was important — probably the same two

who had mentioned their own touristry experience in answering

a prior question!

An interesting comment on this matter of prior occupation

came from a leading official of the Better Resorts Association, a

Rhinelander area resort group that stresses quality service and

good facilities. He said : “Most people in the resort industry know

little or nothing about running an effective resort operation. They

tend to be poor resort managers, even if they were previously

successful in some other type of business.”

Several studies during the early 1960's have indicated that a

resort business was a relatively poor investment, characterized by

low returns in relation to the inputs of labor and money. A re¬

port by Fine and Tuttle, in 1962, for example, indicated that

38% of Wisconsin's vacation-resort enterprises operated at a loss

1973]

Monthey and Zielinski — Vacation Resorts

223

in 1961 ; of those which showed a profit, three-eighths netted

under $2,500. A 1965 report by Staniforth and others of resorts

in the Oneida County area indicated that all measures of income

related to the resort business were low. After deducting costs,

only 25% of the enterprises showed any net return for the labor

and management contributed by the operator, his family, and

other employees (if any). However, the net worth for most of

the resorts studied was rather high, because of the appreciation

in value of lakeshore properties and the low-debt situation of

resort owners themselves.

The following Table 6 indicates the 1966 gross income figures

for 75 of the 103 resorts included in this study.

TABLE 6. GROSS INCOME FOR 1966, AS REPORTED BY

75 RESORT OPERATORS

These data support the conclusion that low income does indeed

characterize the resort industry. In 1966, about 70% of the 75

resorts in this study had a gross income under $5,000 ; and over

half of these (36%) grossed less than $2,000 for the year. The

most likely reasons for this low-income situation appeared to be

as follows ;

(1) the extremely short operating season ;

(2) the small size of resort ;

(3) the low weekly rate charged ;

(4) the lack of supplemental resort income

(other than cottage rentals).

Although Oneida County resorts in this study were open for

business an average of 17 weeks per year, their “effective season”

was generally about 10 to 11 weeks. In fact, most of the operators

questioned said that their family trade (the so-called “bread and

butter” business) started on or after June 21, when school vaca-

224 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

tions begin, and ran through Labor Day (a total of less than 80

days) .

Three-fourths of the resorts surveyed did not have any signifi¬

cant supplemental income besides cottage rentals.* The average

annual per-cottage income for 60 of these resorts came to $605,

or $55 per week for an 11-week season. This group had a mean

cottage number of 4.7 and a mean income per resort of $2,856.

In this study, resort size could not be related, in any significant

way, with the income per cottage. Some small resorts had a rela¬

tively high per-cottage income, while some of the larger estab¬

lishments had a low general figure. For example, resorts with

three cottages per establishment had a range of annual income

per cottage all the way from $97 to $833, while those resorts with

10 or more cottages had income ranging from $417 to $1,100 per

cottage.

With self-service (housekeeping) resorts in Oneida County,

there seemed to be no relationship between gross resort income and

the amount of advertising expenditure per establishment. Further¬

more, there was only a weak relationship between the B.U.

capacity of self-service resorts and the size of their advertising

budgets. In general, those with more than 10 B.U. spent about

twice as much for advertising as those with fewer than ten. But

the mean expenditure for advertising in 1966, based on inter¬

views with 86 of the 103 resorts, was only $160 per establishment.

Only three of these resorts topped the $1,000 level, and 16 spent

nothing on promotion. One used television commercials, two used

radio services, and only eight used newspaper advertising in

their own name. Most of the promotional expense involved high¬

way signs, brochures, and association or chamber of commerce

dues.

In addition to small advertising budgets, Oneida County resorts

allocated a relatively small amount of money for maintenance and

repairs in 1966. On the average, only $900 a year was spent

by 43 resorts in the sample survey. Almost one-half of the es¬

tablishments spent less than $400 for repairs and four-fifths spent

less than $1,000 each in that year.

When asked about their plans for the improvement of existing

(1967) facilities, over three-fifths (63%) of the resort operators

said that they had no such plans. Of the 94 establishments answer¬

ing this question, 25 volunteered the information that they were

thinking about selling their resorts and therefore were not plan¬

ning to improve their facilities.

In this regard, two major handicaps have been the high cost

* The principal source of additional income lies with food and beverage sales. Only

23% of the resorts in this study had either a beer bar or a full liquor license.

1973] Monthey and Zielinski — Vacation Resorts 225

of such improvements and the inability of resort establishments

to obtain long-term loans from their local lending agencies. This

has made it most difficult to build, purchase, expand or modernize

a resort — or even to maintain the quality of the plant. Conse¬

quently, almost no new resorts are now being constructed, and

there is very little expansion and/or modernization of existing

facilities in this County at present.

This study found that almost 90% of the establishments sur¬

veyed were in operation before 1950, and about 70% of the cot¬

tages were built before that date. No new resorts were constructed

during the 1961-67 period, and only a few new cottages were

added. The average land acreage per resort was found to be 32

acres.

Quality classes were established for the 103 resorts studied

in 1967, with the help of information from the town tax assessors.

A summation of these “ratings” is as follows :

Class “A” — Deluxe (modern, excellent condition) _ 7.0%

Class “B” — Modern (fair to good condition) _ 49.0

Class “C” — Semi-Modern (poor to fair condition) _ 37.0

Class “D” — Sub-Modern (very poor condition) _ 7.0

In general, the operators of these establishments tended to rate

their own places higher than did the assessors or the researchers

making this study.

Within Oneida County, there was a considerable variation

among the townships with respect to changes in resort numbers

and capacity between 1950 and 1967. Of the 20 governmental

Towns in the county, 15 lost resorts, two showed no change, and

only one had a net increase during the period ; two Towns had no

resort establishments. Eleven Towns had 20 or more resorts

apiece in 1950, and 9 of these still had 20 or more establishments

in 1967. Two of the largest Towns in area, Minocqua and Three

Lakes, have also been the most important in Oneida’s resort in¬

dustry. Each of them had more than 90 resort-type establishments

in 1950, but both showed sizable decreases by 1967.

Table 7 gives data to show trends for the 11 Towns which had

20 or more resorts apiece in 1950. These Towns contained ap¬

proximately 85% of the county’s resorts in both 1950 and 1967.

They varied considerably in the degree of change that took place

over the 18 years. Only the Town of Nokomis showed a net in¬

crease in resort establishments. Ten of the 11 “resort towns”

showed significant declines, but the net decreases ranged from

8.7% in the Town of Woodruff to 38.5% in the Town of Three

Lakes.

It is interesting to compare the latter with the Town of Minoc¬

qua, the other “big” resort town in Oneida County. According

226 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 7. CHANGES IN RESORT NUMBERS AND B.U. CAPACITY FOR

THE LEADING RESORT TOWNS OF ONEIDA

COUNTY (1950-1967).

*This figure is undoubtedly low. since many of the very small (non-commercial) resorts with one or

two cottages did not get tallied during the 1946-1955 period.

to the Board of Health information, each of them had over 90

resort-type establishments in 1950. But, whereas Three Lakes lost

almost 39% of theirs during the period 1950-1967, Minocqua

showed a more modest decrease of about 14%. In terms of bed¬

room units, Three Lakes dropped 29% compared to 14% in

Minocqua.

Zielinski believes that the relative increases in real estate taxes

and property assessments during the 1960’s were important fac¬

tors in the above differences. For example, the Town of Three

Lakes had the highest ratio (0.25) of real estate taxes to gross

resort income for any resort town in the county. Moreover, this

town had the highest assessment ratio (102.2% in 1966) in the

county, almost a four-fold increase over the assessment ratio

(28%) in 1963. This may be why nearly one-half of the town’s

resort operators interviewed in the 1967 survey were planning on

selling their establishments. Almost all of those interviewed re¬

ported increases of several hundred dollars in their 1966 real-estate

tax, compared to the year before.

To summarize this comparison, the Town data showed that

the ratio of resort “dropouts” to resorts still in operation (1966-

1967) was twice as high for the Town of Three Lakes as for

the Town of Minocqua. At the same time, the ratio of real estate

taxes to gross resort income was four times higher in the Town

of Three Lakes than in the Town of Minocqua (0.25 compared to

0.06). This could well be a major factor in the decision of some

owner-operators in the Three Lakes area to go out of the resort

business.

1973] Monthey and Zielinski — Vacation Resorts 227

The primary reason for the large real estate tax increases in

the county during the years following 1960 has been the dramatic

appreciation of market values in lakeshore property. Prominent

realtors in both Rhinelander and Three Lakes reported that lake

frontage in their areas that was selling for $2 to $8 per “front foot”

in 1950 was moving at $40 to $50 by 1967. Since 1960, the rate

of value appreciation of “good” shoreline property has averaged

about 10% a year.

Another factor in tax increases has been the tendency on the

part of local assessors to raise the property assessment, which

had ranged from 20 to 40% of market value, toward the 100%

level throughout northern Wisconsin. This trend toward full valu¬

ation, prompted by state tax authorities, has led to major adjust¬

ments, mostly upward, in the real estate taxes for lakeshore

property. This is particularly true in those townships where a

high percentage of the mercantile tax base consists of riparian

land, particularly lakeshore developments.

Because of the strong demand for recreational real estate

(especially water-based lands), either for private use or long-term

investment, the value of lakeshore property continues to spiral

upward. In recent years, many people have felt sufficiently “well

off” to pay the market price for lakefront lots, despite high taxes,

in order to build a summer cottage, retirement home, or year-

round residence. Others have acquired such property as an invest¬

ment for the future.

On the other hand, many resort establishments that are holding

a large piece of high-tax shoreline, and operating a low return

business thereon, are being encouraged to sell or subdivide their

holdings, usually at a handsome profit. To many resorters, this

has been an attractive alternative to the growing problem of

mounting costs and high property taxes. Oftentimes the price

offered to them for the resort is 50% to 150% higher than their

total investment to date, and this has promoted both real-estate

activity and resort sales.

Resort conversions of this kind have caused a gradual disap¬

pearance of the traditional resort industry in some areas, and

this appears to be the case in several Oneida County townships

at present. Looking ahead, this changing usage and function of

commercial resort-recreation lands could become a major trend

throughout Northern Wisconsin.

REFERENCES

1. MONTHEY, L. G. 1964. The resort industry of Wisconsin. Trans. Wis.

Acad. Sci. Arts Lett. 53(A); 79-94.

2. MONTHEY, L. G. 1970. Trends in Wisconsin’s tourist-lodging industry.

Trans. Wis. Acad. Sci. Arts Lett. 58: 71-99.

MAJOR CAUSE OF ALGAE AND WEEDS IN

LAKE MENDOTA

M. Starr Nichols

University of Wisconsin —

Madison

ABSTRACT

Because of low partial pressure of carbon dioxide in the

atmosphere and the resulting low total solubility, atmospheric

carbon dioxide is not the major source of carbon for algal

and plant growths in hard water lakes. Bicarbonates of calcium

and magnesium in the lake water are the major source. This is

shown by the lowering of the hydrogen ion concentration of the

lake water during extensive growths, causing a rise in pH to 9.5

to 10.0 and above following rapid growth during hot weather. At

times the pH rise causes precipitation of marl deposits in cal¬

careous waters.

The perennial problem of the summer growth of algae and

weeds in Lake Mendota has caused much speculation and comment.

That it is a real problem is attested to by many lake-edge dwellers

and users of this lake for recreational purposes.

The cause is rooted in the process of green plant growth. The

many contributory causes often are spoken of as pollution by

man, and rightfully so in some respects. Green plants grow in

lake waters in the same manner as grass grows on our lawns,

trees grow in the forest, and the corn crop grows in the farmer’s

field. These plant growths, both in water and on land, are im¬

portant to man as the source of the oxygen we breathe and the food

we eat.

The sun is the energy source to produce these growths of green

plants. Photosynthesis is the name of the process by which the

plants capture solar energy. The chlorophylls in algae and aquatic

weeds, using solar energy, convert carbon dioxide and lake water

into extensive growths of plant tissue and at the same time lib¬

erate pure oxygen from the process. The chemical expression of

this conversion is given in the following equation : C02 + 2HL>0 +

chlorophylls + solar energy - > HCOH + H20 + 02 + potential

energy. The HCOH represents the simplest organic compound of the

photosynthetic process. This conversion expressed in pound units

229

230 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

means that 44 pounds of carbon dioxide and 18 pounds of lake

water are converted into 30 pounds of organic plant growth repre¬

sented by HCOH in the equation and 32 pounds of free oxygen

gas. This oxygen becomes, therefore, a part of the oxygen in the

lake or released to the atmosphere which we breathe. It is this

process that maintains the oxygen in the atmosphere at 20.99

per cent by volume. Note that no organic wastes or “pollution”

are involved as such in the process.

SOURCES OF CARBON DIOXIDE

Some of the carbon dioxide used by the green plants growing

in Lake Mendota, comes directly from the air by simple solution.

The quantity of carbon dioxide present in the air by volume is

320 parts per million parts of air. Because of its small vapor pres¬

sure, 0.243 millimeters of mercury (Henry’s Law), only small

amounts become dissolved in the lake water. At 20 C and average

air pressure, only 0.55 parts per million by weight of carbon

dioxide will be dissolved in water. The total amount in the atmos¬

phere is so very great as to constitute by weight about one pound

over every square foot of surface of the earth both land and

water (1) or about 13,000 tons per square mile.

Carbon dioxide in the air is in equilibrium with the carbon

dioxide in all surface water — lakes, rivers, and oceans — on the

earth’s surface. It is from this large reservoir of carbon dioxide

in the atmosphere plus that produced by decay of plant and ani¬

mal residues that green plants with chlorophylls and solar energy

are able to fix, in usable energy packets, the complete energy

food for life on this planet, and the oxygen to use it.

A major source of usable carbon dioxide for the algae and

weeds growing in Lake Mendota water, is found in its alkalinity.

This alkalinity consists of the bicarbonates of calcium and magne¬

sium represented by the following formulae: Ca(HC03)2 and Mg

(HC03) 2 respectively. The ~HC03 of these formulae represent H20

and C02 and it is a part of this C02 which becomes available for

the production of growths of algae and weeds. From the alka¬

linity of 160 pounds per million pounds of lake water there is

available 44 pounds of carbon dioxide for every million pounds of

Lake Mendota water. Since there are 478,370,000 cubic meters of

water in Lake Mendota (2) which expressed in pounds would be

1,000,000,000,000 pounds = 500,000,000 tons. In this 500,000,000

tons of water there would be 22,000 tons of carbon dioxide avail¬

able from bicarbonates for photosynthesis of algae and weeds.

Since 44 pounds of carbon dioxide under the action of chloro¬

phyll and solar energy could produce 30 pounds of algae and weeds,

we have in Lake Mendota waters sufficient carbon dioxide in form

1973] Nichols — Cause of Algae in Lake Mendota 231

of bicarbonates to produce more than 15,000 tons of algae and

weeds. If only carbon dioxide dissolved from the air was avail¬

able, less than 200 tons of algae and weeds could be produced.

The 15,000 tons divided by the acreage on Lake Mendota, 9,728

acres (15.2 sq. miles (2)), gives a crop of 1. 8 tons per acre —

equivalent to a small crop of alfalfa on land.

Note. As the bicarbonate (carbon dioxide) is used by the lake

plants, there is a continuous addition of carbon dioxide from the

air and from carbon dioxide from decay processes. The lowering

of the hydrogen ion concentration of lake water to pH 9 or 10

or higher shows the rapid decrease of bicarbonate ion to approach

only normal carbonates of calcium and magnesium to be present

at times in late summer, when algal growth has been rapid. It

is at this stage of growth that marl formation takes place. During

times of slow growth or no growth, the hydrogen ion concentra¬

tion rises and the pH drops to 7.8 or 8.0 and the bicarbonates of

calcium and magnesium are at least partially formed again.

SOURCES OF OTHER FOOD FOR ALGAE AND WEEDS

Not mentioned in the above conversion equation for photosyn¬

thesis are the 3 pounds of nitrogen, 0.33 of a pound of phosphorus

as compounds, and trace quantities of magnesium and other min¬

erals in solution in the Mendota lake water.

Lake Mendota is partially spring-fed with water containing

nitrate nitrogen. Streams such as Yahara River, Token Creek,

Six-Mile Creek, Pheasant Branch, Merrill Springs Creek and Uni¬

versity Creek furnish drainage water containing bicarbonates,

both nitrogen and phosphorus compounds and small amounts of

other minerals. Rain and snow contribute 8 to 10 pounds of nitro¬

gen in form of compounds each year for every acre of Lake Men-

dota’s surface, or a total of more than 40 tons of actual nitrogen.

Attempts are being made to curtail the input to the lake of

nitrogen and phosphorus from farm operations but the seepage

(base flow) flow found by Minshall, Nichols and Witzel (3)

would add slightly more than one pound of nitrogen per acre

per year from the 89,600 acres (140 square miles) of land

draining into Lake Mendota; this represents 44 tons of nitrogen

as fertilizer loss from farm practice. In addition to this nitrogen

from seepage flow, the amount of phosphorus in farm loss from

seepage flow was found to be 0.09 pounds (1 pound of phos¬

phorus is equivalent to 4.6 pounds P205)* From 140 square miles

of drainage entering Lake Mendota this loss would amount to

four tons of phosphorus per year. The above values were calcu¬

lated from a survey of drainage seepage of Southwestern Wis-

232 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

consin farm land in which sampling was made from streams drain¬

ing 643 square miles during the year 1969. Samples were obtained

when there was no flood flow and represented the basic flow or

seepage flow. Samples were obtained from 38 streams with flows

ranging from two cubic feet per second (over 1.22 million gal¬

lons per day) to 31 cubic feet per second (19 million gallons

per day). Over 500 separate analyses were made.

The amount of fertilizer nitrogen and phosphorus from surface

drainage will be considerably more than from seepage flow. From

another research project by the same workers (4) it was found

that the average loss per acre per year from surface drainage

was 2 pounds of nitrogen and 0.46 pounds of phosphorus. These

values, applied to Lake Mendota drainage waters, would add 89

tons of nitrogen and 20 tons of phosphorus. The drainage areas

of these experiments were planted to corn and were fertilized

with manure only at the time of planting. No cultivation was

used. The percentage of the land in Dane County planted to corn

was given by the Dane County Agricultural Agent, Mr. Thomas

O’Connell, as about 34 percent on the average. Out of 600,000

acres of farm land, 204,000 acres would be planted to corn.

While the values of seepage flow should properly apply to the

drainage into Lake Mendota, the runoff of surface flow might

be less accurate when calculated from a small experimental plot

of 0.1 acre of crop land, as the basis for Lake Mendota fertilizer

input. The values over three calendar years were rather uniform,

however, varying only from 1.25 to 2.8 pounds per acre per year

for nitrogen fertilizer and from 0.3 to 0.55 pounds per acre per

year for phosphorus.

AVERAGE VALUES OF ELEMENTAL

COMPOSITION OF THE BIOSPHERE

The biosphere includes both animal and vegetable living bodies.

The average composition of body substance of the biosphere on

a dry weight basis is given in the following Table 1, calculated

from results given by Edward Deevey Jr. (5).

TABLE 1. ELEMENTS PRESENT IN BIOSPHERE (DRY WEIGHT)

Element

% wt. Element

°7c wt.

Hydrogen _ 0.66

Oxygen _ 52.30

Carbon _ 40.00

Nitrogen _ 0.50

Calcium _ 0.38

Potassium _ 0.26

Silicon _ 1.21

Magnesium _ 0.87

Phosphorus _ 1.26

Sulfur _ 0.45

Aluminum _ 0.56

Total _ 98.55

1973] Nichols — Cause of Algae in Lake Mendota 233

The small percentage not accounted for in this table would in¬

clude elements not determined such as iron, manganese, boron,

zinc, selenium, vanadium, chromium, molybdenum, iodine, fluorine,

copper, sodium and cobalt. It is to be noted that 93 percent of

the weight of growth consists of carbon, hydrogen and oxygen,

the principal constituents of plant carbohydrates such as starch

and cellulose-type compounds of algae and weeds.

Nitrogen and sulfur are essential elements in proteins, and phos¬

phorus is an essential element in metabolism of animal and plant

tissue. One atom of magnesium is needed for each molecule of

chlorophyll Silicon is needed in the shells of diatoms (algae).

Calcium is a principal skeletal element of animals.

SOURCES OF CALCIUM AND MAGNESIUM BICARBONATES

IN LAKE MENDOTA WATER

The natural recycling in soils of waste organic matter such as

animal manures, corn stalks, straw, weeds and other waste or¬

ganic matter by aerobic and anaerobic decay converts the carbon

in these wastes to carbon dioxide. The conversion may be rapid

or slow depending on the availability of soil nutrients and micro¬

bial activity. The carbon dioxide formed by decay is dissolved

by rain water and the carbonic acid formed attacks the calcium

and magnesium carbonates present in the soil naturally or added

as agricultural lime, and as this carbon dioxide charged water

passes deeper, it dissolves the carbonates present in the rock strata

to form the soluble bicarbonates of calcium and magnesium.

These bicarbonate waters are intercepted for our domestic water

supply and the drainage water from soil and springs often con¬

tain large amounts of bicarbonates, as are found in Lake Men¬

dota. Madison city water intercepted at depths of from 200 to

1,000 feet by our deep wells contains nearly double the bicarbo¬

nates of the water of Lake Mendota.

SUMMARY

1. Carbon dioxide and water have always been the principal

quantity of food materials used by algae and weeds and, in fact,

all plant growth; they contribute more than 90 percent of the

weight of these growths in terms of dry weight of tissue.

2. In the absence of carbon dioxide or a deficiency from sources

other than the atmosphere, the low partial pressure of carbon

dioxide in the air limits the rate and quantity of solution of this

gas in lake water to the extent that large and extended blooms

of algae and weed growths would be materially limited. In lakes

with a low bicarbonate content, carbon dioxide could be a critical

factor for extensive growths of algae and weeds.

234 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

3. In the case of Lake Mendota there is an ample supply of

carbon dioxide available in the bicarbonates, while the air acts

only as a slowly available reservoir.

4. Minerals such as nitrogen, phosphorus, sulfur and traces

of many other elements must be in adequate supply in these bi¬

carbonate waters, if growths of algae and weeds occur in sufficient

quantity to become a nuisance.

5. There seems to be an adequate supply of phosphorus and

nitrogen in Lake Mendota water from drainage into the lake from

farm land, through precipitation (rain and snow), and from

springs flowing into the lake at the bottom. Nitrogen and phos¬

phorus are also furnished by a rapid recycling of these elements

and their compounds. Phosphorus is available for metabolic ac¬

tivity of growing algae. Both phosphorus and nitrogen become

readily available through active decomposition of dead algae and

weed substance by organisms of decay. These decay processes

are active continuously both day and night and summer and win¬

ter (but less active in cold temperatures).

6. While the use of fertilizers are necessary for world food

production, agriculturists are cognizant of the losses by land

drainage. Contour plowing is one of the steps already taken by

many farmers to retain fertilizers not immediately used by plants,

and to prevent erosion losses. Current practices for manure han¬

dling also help to keep drainage levels low.

REFERENCES

1. BOLIN, BERT. 1970. The carbon cycle. Sci. Amer. 223 (Sept.) : 124.

2. NEESS, JOHN C., WILLIAM W. BUNGE, JR. 1956. Unpublished manu¬

script of E. A. Birge on the Temperature of Lake Mendota. Part 1.

Trans. Wis. Acad. Sci. Arts and Lett. 45: 193-237.

3. MINSHALL, NEAL, M. STARR NICHOLS, and S. A. WITZEL. 1969.

Plant nutrients in base flow of streams in southwestern Wisconsin.

Geophys. Union, Water Resources Res. 5: 706-713.

4. MINSHALL, NEAL, STANLEY A. WITZEL, and M. STARR NICHOLS.

1970. Stream enrichment from farm operations. Proc. Amer. Soc.

Civil Engin., Sanit. Engin. Div. 96: 513-524.

5. DEEVEY, EDWARD JR. 1970. Mineral cycles. Sci. Amer. 223 (Sept.) :

148.

ISOLATION AND CHARACTERISTICS OF A POSSIBLE

ALLELOPATHIC FACTOR SUPPORTING THE DOMINANT

ROLE OF HIERACIUM AURANTIACUM IN THE

BRACKEN -GRASSLANDS OF NORTHERN WISCONSIN

Dana S. Dawes and N. C. Maravolo

Lawrence University—

Appleton

Possible allelopathic effects of Hieracium aurantiacum were in¬

vestigated to elucidate their role in stabilizing the bracken-grass¬

lands. These areas, where Hieracium is common, displayed im¬

paired invasion by the tree species characteristic of the northern

forests.

Extraction and chromatography of soil taken from Hieracium

stands in Vilas County, Wisconsin contained seven phenolic com¬

pounds. Extraction of sand in which Hieracium had grown showed

these same seven substances. Three of these were tentatively iden¬

tified as ferulic acid, vanillic acid, and umbelliferone. Bioassays

with Abies balsamea, Lactuca saliva, and Pinus strobus consistently

revealed the presence of several active inhibitors of both germina¬

tion and hypocotyl growth. At least two of these were non-

phenolic. Other active fractions were correlated with the presence

of phenolic compounds.

Abies and Pinus seeds planted and grown in sand which had pre¬

viously supported Hieracium had normal germination and shoot

growth; however, survival was greatly reduced in both species.

The possible physiological action of the phenolic inhibitors is

considered, particularly their role in hormonal control and mineral

nutrition.

INTRODUCTION

In northern Wisconsin there is a prairie community dominated

by bracken fern (Pteridium aquilinum) , and the grasses Danthonia

spicata, Bromus kalmii, Agropyron trachycaulum, and Oryzopsis

asp eri folia. Curtis (1959) defines this as the bracken-grassland

(Fig. 1). Following glaciation, most of this area was forested.

These openings probably originated with the intensive logging and

subsequent fires in the late 1800’s. Since that time, much of the

cleared area has undergone natural succession, invasion of trees,

235

236

Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FIGURE 1 (Top) — Bracken-grassland in Vilas County, Wisconsin.

FIGURE 2 (Bottom) — Hieracium aurantiacum, common orange hawkweed

1973]

Dawes and Mar avoid — Bracken-Grasslands

237

and gradual return to xeric or dry-mesic forest. The bracken-

grassland has remained free of trees. One of the proposed explana¬

tions for this stability is the introduction by established plants of

a chemical agent detrimental to the other species (Levy, 1970).

This allelopathic agent could affect either the germination or the

growth of other species, or both. This, in turn, would hinder the

migration, ecesis, or competition of these plants so as to retard

succession (Muller, 1966). Allelopathy may also alter the species

composition by only allowing the establishment of tolerant species

(Whittaker and Feeny, 1971).

Curtis suggested that several important bracken-grassland spe¬

cies may affect the community in the former way. One of these was

Hieracium aurantiacum, common orange hawkweed (Fig. 2). Al¬

though Hieracium aurantiacum was not considered a dominant

species by Curtis, he found it in 63% of the areas studied. Levy

(1970), working with three similar community types in the same

region of northern Wisconsin, found frequency values for Hiera¬

cium of 100, 100, and 87%.

Curtis suggested that Hieracium obtained complete local domi¬

nance through production of a potent allelopathic agent. Guyot

(1957) observed a similar pattern of dominance in another mem¬

ber of the same genus, H. pilosella. It has been reported that water

soluble substances which inhibit the germination and growth of

other species have been extracted from soil in which this species,

as well as two other Hieracium species had grown (Becker and

Guyot, 1951a and b).

This evidence suggests that allelopathic agents may be present

in the soil near Hieracium and that these compounds may play an

important role in preventing the establishment of trees in the

bracken-grassland. This would account for the preservation of the

pioneer community, but no conclusive evidence supporting the sug¬

gestion exists.

The most common method of demonstrating allelopathic effects

employs a bioassay, based upon defined responses by biological

material exposed to suspected compounds. An assay analyzing

the germination and growth of selected seeds was used in this

study. The most meaningful bioassays for allelopathy are those

which employ plant species likely to be found near the suspected

plant in the field. Preliminary work was done in and around the

bracken-grasslands to determine the species of trees most likely to

invade the areas.

Soil was removed from Hieracium stands, extracted, and assayed

for the presence of allelopathic substances. To show conclusively

that these compounds were produced by Hieracium alone, a similar

238 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

extraction was done of washed sand in which isolated Hieracium

plants had grown.

Whittaker (1971) suggested that allelopathic substances gener¬

ally fall into two major chemical groups, each associated with a

particular macroclimate. In arid regions, terpenes and related

volatile inhibitors predominate (Whittaker, 1971; Muller, 1965;

Muller, Lorber, and Haley, 1968; Muller and del Moral, 1966).

Phenolic compounds are prevalent in more humid areas (Whit¬

taker, 1971; Whittaker and Feeny, 1971). The bracken-grasslands

areas in Wisconsin are located in an area with high humidity, and

moderately high rainfall (105 days per year with at least 0.01

inches of rain, Curtis, 1959). It seemed probable that if allelo¬

pathic agents were being produced by Hieracium, they were

phenolic.

MATERIALS AND METHODS

During September, 1971 a survey was made of 12 bracken-

grasslands, primarily in Vilas County, Wisconsin. At each site,

the primary tree species surrounding the area were identified, and

the nature of the upper soil horizon determined.

At a large bracken-grassland near Boulder Junction, Wisconsin

several 6250 cm3 soil samples were taken from under heavy stands

of Hieracium. The samples were kept on ice, returned to the labo¬

ratory and then frozen to slow microbial decomposition. Prior to

use these samples were air-dried at room temperature for a mini¬

mum of three days.

Intact H. aurantiacum plants were collected from a site near

New London, Wisconsin. The plants were rinsed with tap water,

and transplanted to quartz sand, which had been washed in 2M

HC1 followed by several rinses of distilled water. The plants were

watered twice weekly with Hoaglund’s nutrient solution and at

other times with tapwater.

The removal of the phenolic compounds from the soil and the

sand was based on a modification of the technique developed by

McPherson, Chou, and Muller (1971). This process is based on

the principle that phenolic compounds are more soluble in base

than in acid; and that when subsequently extracted in a diethyl

ether-acid partition, the phenols will be more soluble in the ether.

Phenols are readily recovered from ether by drying.

A 100 g sample of air-dried soil or sand was first screened to

remove plant material and large soil particles. One hundred-fifty

milliliters of 95% ethanol were added and the pH of the solution

adjusted to 13.0 with 2M NaOH.

The soil-alcohol suspension was placed in a mechanical shaker

for 24 hr. The pH was periodically readjusted to 13.0. After re-

1973]

Datves and Maravolo — Bracken-Grasslands

239

moval from the shaker, the suspension was centrifuged for 10 min

at 1,500 x g to remove suspended soil particles. The pH of the

supernatant fluid was adjusted to 7.0 with 2M HC1, and the fluid

then dried for 24 hr at 65 C. The residue was ground to a powder

and added to a flask containing 100 ml each of 2M HC1 and diethyl

ether. The ether layer, containing the phenols, was separated and

dried at room temperature. This residue was redissolved in 10 ml

of methanol for chromatography.

The methanolic solution was applied to 7.5 x 57 cm strips of

Whatman #3 chromatography paper. The phenols were separated

by descending chromatography in 2% aqueous acetic acid (v/v).

After the solvent front had reached the bottom of the paper, the

chromatogram was removed and dried overnight at room tempera¬

ture.

Abies balsamea and Pinas strobus seeds were obtained from

F. W. Schumacher Company (Sandwich, Massachusetts). Before

use, the seeds were vernalized to break dormancy, by refrigerat¬

ing the seeds at 4 C for at least two months.

The bioassays for allelopathic agents were a modification of the

technique of McPherson, et al. (1971). Each chromatogram was

divided into 10 5.0 cm fractions and each fraction into 25 cm2

squares. These paper squares were placed on sponge pads (6 x 6 x

1 cm) in petri plates (Muller, 1965). Seeds from Abies, Pinas or

Lactuca were placed on top of the paper. Another sponge was

placed over the seeds to insure contact with the paper. Untreated

Whatman #3 paper served as a control. After adding 20 ml of

distilled water, the petri plates were sealed with tape to prevent

desiccation. Three replicates were made of every experiment and

incubated at 18 C and 85% relative humidity during the bioassay.

The duration of the bioassay varied with the species used. Abies

and Pinas seeds were treated for three weeks; Lactuca for four

days. Lactuca seeds were used to provide a rapid estimation of

general toxicity.

At the end of the treatment period, the total numbers of germi¬

nated and ungerminated seeds were determined and the mean

germination percentage calculated. Hypocotyl (root) length of

each germinated seed was determined to the nearest 0.1 cm for

the two tree species, and average values were calculated.

The effects of possible soil-borne allelopathic agents on later

seedling growth were investigated by using sand in which Hiera-

cium had grown as a substrate for seedlings. Similar sand which

had not supported Hieracium was used as a control. After the

intact Hieracium plants had been carefully removed, 10 tree seeds

of either Abies or Pinas were placed 1 cm deep in the sand of

each pot. Five replicates were made of each. These seeds were

240 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

maintained in the greenhouse and watered twice weekly with

Hoaglund’s solution.

After three months, total germination and mean height (near¬

est 0.1 cm) were determined. A determination was also made of

the number of seeds that had germinated but later died. These

data were compared with the total germination to measure the

degree of seedling mortality.

The identification of phenols in the Hieracium soil extracts es¬

tablished by spraying the chromatogram first with 20% aqueous

Na2C03 and then with 25% (v/v) aqueous Folin-Ciocalteau re¬

agent (Krebs, Heusser, and Wimmer, 1969). The characterization

of the individual phenols was based on three additional criteria:

fluorescence under ultraviolet (UV) light of wavelengths 253 nm

and 366 nm, and Rf values. As each chromatogram was examined,

the location, and color and intensity of each phenolic fraction was

noted, and identified as phenol A to G.

Many phenols have been implicated in allelopathy. These include

caffeic acid (del Moral and Muller, 1969), para-coumaric acid

(Muller, et al., 1968; del Moral and Muller, 1970; Hanawalt, 1971),

ferulic acid (del Moral and Muller, 1970; McPherson, et ah, 1971),

hydroquinone (McPherson, et al., 1971; Hanawalt, 1971), para-

hydroxybenzoic acid (McPherson, et al., 1971; McCalla, 1971),

syringic acid (McPherson, et al., 1971; McCalla, 1971), 7-hydroxy-

coumarin (umbelliferone) (McPherson, et al., 1971), and vanillic

acid (McPherson, et al., 1971; McCalla, 1971).

The fluorescence and Rf values in 2% acetic acid of samples of

each of these eight common allelopathic phenols were established

and compared with those of the substrate isolates.

RESULTS AND DISCUSSION

Populus tremuloides , P. grandidentata, Betula papyrifera, Abies

balsamea, Pinus strobus, and P. resinosa occurred most frequently

in the forests immediately surrounding the bracken-grasslands.

The two Populus species were the only trees that had invaded the

grasslands to any extent. Some Abies and a very few Pinus strobus

were seen. The upper soil horizons varied from loamy sands to

sandy clay loams.

The results of the Lactuca seed bioassay (Fig. 3) showed de¬

creased germination in all fractions. Germination was greatly re¬

duced in fractions 1, 3, 4, 6, and 7. It was also reduced, to a lesser

extent, in fraction 9. Fraction 6 had the lowest germination per¬

centage. Figures 4 and 5 depict the results of the Abies bioassay

of chromatogram fractions. The presence of fractions 1, 5, 7 and

10 slightly enhanced germination. Fractions 4 and 9 reduced ger¬

mination slightly, and fractions 3, 6, and 8 reduced germination

1973]

Dawes and Maravolo — Bracken-Grasslands

241

100

80

60

40

20

1

3 4

6 7 8 9 10

FRACTION

FIGURE 3. Effects of chromatographic fractions on the germination

of Lactuca sativa after four days.

by a large degree. Again fraction 6 had the lowest per cent germi¬

nation. There was no germination in fraction 2 for this species.

The Pinus strobus bioassay (Fig. 6) reduced germination in all

samples. Germination was inhibited by more than 50% by frac¬

tions 1, 3, 4, 6, and 10. Fractions 3 (Rf 0.2-0. 3), 6 (0.5-0. 6) and

9 (0.8-0. 9) inhibited germination in all three species.

All fractions except 3 reduced hypocotyl growth of Abies, com¬

pared with controls. The most inhibitory fraction was 9; 5 and

242 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FRACTION

FIGURE 4. Germination of Abies balsamea on chromatogram fractions

after three weeks.

6 also reduced growth by at least 20%. Pinus hypocotyl growth

was enhanced by fractions 2, 4, 8, and 9, but markedly reduced

in 1, 5, and 10.

Sand that had supported Hieracium for six weeks slightly en¬

hanced Abies germination, but reduced that of Pinus (Table 1).

Mean heights of Abies and Pinus were reduced by 8 and 11%

respectively. Study of Abies seedling mortality revealed that, ah

1973]

D caves and Maravolo — Bracken-Grasslands

243

FIGURE 5. Hypocotyl growth of Abies balsamea seeds exposed to

chromatogram fractions for three weeks.

though 90% of the control seedlings survived for three months,

only 62% of the experimental seedlings did. Pinus survival reached

80% in the controls, but only 27% in the experimental seedlings.

Seven phenols were detected with the Na^COs/Folin-Ciocalteau

spray in extracts from both field soil and washed sand in which

tiieracium had grown. The Rf values in 2% acetic acid and UV

characteristics for these isolates are summarized in Table 2 ; char-

244 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FRACTION

FIGURE 6. Germination of Pinus strohus seeds exposed to chromato¬

gram fractions for three weeks.

acteristics for eight known allelopathic phenols appear in Table 3.

On this basis, three of the seven phenolic isolates were tentatively

identified. The mean Rf for ferulic acid was 0.36; that of phenol

A, 0.34. Both bands were blue or light blue under both UV wave¬

lengths. Umbelliferone (7-hydroxycoumarin) , Rf 0.57, resembled

phenol D (Rf 0.60) under UV. Vanillic acid, Rf 0.63, and isolate

E, Rf 0.63, both absorbed short (253 nm) UV, but E was not de-

1973]

Dawes and Maravolo — Bracken-Grasslands

245

TABLE 1— PER CENT GERMINATION, AVERAGE HEIGHT, AND

SEEDLING SURVIVAL AFTER THREE MONTHS OF

A. BALSAMEA AND P. STROBUS

TABLE 2— CHARACTERISTICS OF UNKNOWN PHENOLIC COM¬

POUNDS ISOLATED FROM HIERACIUM SUBSTRATE

detected.

tected in long* (366 nm) UV. Perhaps this reflected concentration

differences.

Chromatographic fraction 6, which strongly inhibited germina¬

tion in all three species, included phenolic isolates C and D (um-

belliferone) . The inhibition by fractions 4 and 5 correlated with

the phenols A and B respectively. Compounds E and F appeared

in fraction 7, and G in fraction 8. Although fractions 3 and 9 in¬

hibited germination in Lactuca, Abies, and Finns, neither re¬

sponded positively to the Na2C03/Folin-Ciocalteau reagent. The

inhibitory effect of phenols on germination has been well docu¬

mented (Abdul-Wahab and Rice, 1967; McPherson, et al., 1971).

246 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

Key to abbreviations used, abs — absorbs light, It — light.

The physiological mechanism through which the phenols affect

germination is less clearly understood.

The correlative influence of hormones is paramount in mediating

germination. Gibberellin activity is especially important in seeds

possessing complex mechanisms of dormancy. Exogenously ap¬

plied gibberellins can break seed dormancy in several species.

Increasing levels of endogenous gibberellins have been associ¬

ated with release from dormancy (Wareing, 1969). If the in¬

hibitors isolated in this study affect gibberellins, they would in¬

directly influence germination. Corcoran (1971) demonstrated

that when several tannin-related plant phenols were applied to

intact plants, they acted as gibberellin antagonists; they negated

the effect of exogenously applied gibberellins. This same principle

may be expressed during germination, causing the observed de¬

crease in germination.

Fractions 1, 5, 6, and 10 inhibited hypocotyl growth of the two

tree species; Abies growth was not inhibited to as great of an ex¬

tent as Pinus, Because fraction 10 contained no Folin-Ciocalteau

positive compounds, the inhibition there must be due to a non-

phenolic compound. The origin compound (fraction 1) gave a pos¬

itive reaction to the reagent and showed some biological activity.

Fractions 5 and 6 also reduced growth. These latter fractions con¬

tained three phenolic compounds.

As with germination, the inhibitory effects of phenols on hypo¬

cotyl growth has been well demonstrated. Vazquez, et al. (1968)

have shown that umbelliferone inhibited the root growth of sev¬

eral woody species. Wang, et al. (1967) observed that vanillic

and ferulic acids inhibited both root and shoot growth of several

agricultural species. McPherson, et al. (1971) demonstrated that

1973]

Daives and Maiuvolo — Bracken-Grasslands

247

several phenols, including- umbelliferone, ferulic acid, and vanillic

acid suppressed hypocotyl growth of Lactuca sativa.

The physiological basis for this response is probably also based

upon hormonal balance. The most important hormone involved

with root growth is auxin (Thimann, 1969). Phenols are well

known to interfere with auxin balance by either inhibiting or

enhancing auxin oxidation (Waygood and MacLachlan, 1961).

Consequently, some phenols increase auxin levels, others decrease

them (Tomaszewski and Thimann, 1966). This situation seems

to be expressed by Pinus (Fig. 7). Fractions 2, 4, and 9 elicited

greatly increased hypocotyl growth. The lack of any detectable

phenolic activity in fraction 2 suggests that the enhancement in

this fraction might be due to a non-phenolic compound. The inhi¬

bition by fraction 10 was also apparently due to a non-phenolic

substance.

Germination of Abies and Pinus seeds in Hieracium sand was

not reduced; total growth was reduced slightly (Table 1). Seed¬

ling mortality was increased by more than three times in Pinus

and four times in Abies . Germination followed by death has been

reported in Panicum reptans and Euphorbia renif ormis when

these seeds are exposed to allelopathic agents in the soil (Holm,

1971). Buchholtz (1971) postulates that the intact plant may be

affected because nutrient uptake is affected by allelopathy. Knypl

(1970) found that coumarin, a compound closely related to these

phenols, caused chlorosis in cucumber cotyledons. This chlorosis

was overcome by the addition of KC1 to the substrate. The enzy¬

matic and structural role of inorganic ions in plant growth makes

these responses understandable. If seed-stored ions are quickly

consumed during germination and no supplement made available,

death would occur within the course of the bioassay.

These experiments have shown that Hieracium aurantiacum

releases, through some means yet unelucidated, at least seven

phenolic compounds, as well as several non-phenolic ones, into the

soil. Several of these compounds are detrimental to the germina¬

tion, growth, and survival of Abies balsamea and Pinus strobus.

Several other genera important to the bracken-grasslands demon¬

strate some degree of allelopathic activity: Agropyron (Levy,

1970) ; Antennaria (Woods, 1960) ; Aster (Woods, 1960) ;

Erigeron (Woods, 1960) ; Pteridium aquilinum (Levy, 1970;

Whitehead, 1964) ; and Solidago (Woods, 1960). This widespread

occurrence of allelopathy probably has a profound influence on

succession and partially explains the stability of the bracken-

grasslands.

The principal trees invading these areas were Populus grandi-

dentata and P. tremuloides. Abies balsamea and Pinus strobus

248 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

FRACTION

FIGURE 7. Effect of chromatogram fractions on hypocotyl growth of

Pinus strobus after three weeks.

occurred with less frequency. This may, to some extent, reflect the

influence of allelopathy. Except under special conditions, the two

Populus species do not grow from seed. Instead, reproduction

usually derives from suckers, shoots which develop on horizontal

roots of established trees. Examination of the possible inhibitory

effects of allelopathic agents on aspen sucker growth, while an

intriguing problem, is beyond the scope of these studies.

1973] Dawes and Maravolo — Bracken-Grasslands 249

Of the tree species used in these studies, Abies was least affected

by the allelopathic agents. Seed germination, seedling growth, and

survival were all suppressed less than in Finns (Fig. 4-7, Table 1).

This was supported by field observations. While both of these

species occur in low numbers, Abies is much more common than

Finns. A similar report of succession paralleling tolerance to

allelopathic agents has been made by Rice (1971).

Certainly other important processes affect invasion of bracken-

grasslands by trees. Many of these areas possess strong relief, and

form ideal basins for cold air drainage, forming frost pockets. The

lower leaves of aspen growing in these areas are often injured by

frost. Abies , which invades the grasslands in small numbers, is

the most frost-resistant tree in the region. A substrate consisting

of a dense mat of plant litter is often present, which could prevent

other seeds from reaching the soil. Locally high temperature may

also retard tree growth. Community dynamics may also be in¬

fluenced by small mammals and birds. The relative importance of

each of these factors probably varies from one locale to the other

(Levy, 1970). In some rare areas, all of them may combine to

exclude trees.

With techniques for elucidating allelopathic effects so well devel¬

oped, the phenomenon of allelopathy and its role in bracken-

grassland stability warrant further investigation. Conclusive iden¬

tification and quantification of the phenolic and non-phenolic

inhibitors produced by Hieracium are needed, as well as further

evidence of allelopathy by the other genera implicated. A determi¬

nation of the physiological modes of action of these compounds is

also needed.

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TOMASZEWSKI, M. and K. V. THIMANN. 1966. Interactions of phenolic

acids, metallic ions, and chelating agents on auxin-induced growth. Plant

Physiol. 41 : 1443-1454.

VAZQUEZ, A., J. MENDEZ, M. D. GESTO, E. SEOANCE, and E. VIEITEZ.

1968. Growth substances isolated from woody cuttings of Salix viminalis

L. and Ficus caprica L. Phytochem. 7 : 161-167.

WANG, T. and T. CHUANG. 1967. Soil phenolic acids as plant growth

inhibitors. Soil Sci. 103 : 239-246.

WAREING, P. F. 1969. Germination and dormancy, p. 606-671. in M. B. Wil¬

kins (ed.) The Physiology of Plant Growth and Development. McGraw-

Hill, New York.

1973]

Dawes and Maravolo — Bracken-Grasslands

251

WAYGOOD, E. R., and G. A. MacLACHLAN. 1961. Inhibition and retarda¬

tion of the enzymatically catalyzed oxidation of indole-3-acetic acid,

p. 149-166. in Plant Growth Regulation. Iowa State University Press,

Ames.

WHITEHEAD, D. C. 1964. Identification of p-hydroxybenzoic, p-coumaric,

and ferulic acids in soils. Nature. 202: 417-418.

WHITTAKER, R. H. 1971. The chemistry of communities, p. 10-18. in Bio¬

chemical Interactions among Plants. National Academy of Sciences,

Washington.

WHITTAKER, R. H. and P. P. FEENY. 1971. Allelochemics : chemical inter¬

actions between species. Science. 171: 757-770.

WOODS, F. W. 1960. Biological antagonisms due to phytotoxic root exudates.

Bot. Rev. 26: 546-569.

DETERMINATION OF MERCURY LEVELS IN WISCONSIN

RESIDENTS: A PROJECT INVOLVING SECONDARY

SCHOOL STUDENTS

Leon M. Zaborowski

University of Wisconsin —

River Falls

ABSTRACT

One of the many questions concerning mercury in the environ¬

ment is the extent of mercury contamination in humans. To

determine the level of contamination of Wisconsin residents, hair

samples were obtained from 200 high school students throughout

the state. After decomposition of the hair, mercury content was

found by flameless atomic absorption. Hair shampoos were tested

as possible sources of mercury.

INTRODUCTION

A great deal of concern has been generated in the past few years

as a result of the pollution of much of the environment by mercury

(1-6). Since 1950 there have been a number of instances of acci¬

dental mercury poisonings from the ingestion of contaminated

food. However, these were considered to be localized problems and

it was not until 1970 that the general public came to realize the

widespread nature of this problem. It is now known that a good

percentage of the lakes and rivers in this country contain signifi¬

cant amounts of this element and that regular consumption of fish

from contaminated waterways can lead to elevated and possibly

dangerous levels of mercury in humans.

A recent report by the Wisconsin Department of Natural Re¬

sources (7) pointed out the contamination of the Flambeau, Wis¬

consin, Chippewa, and Menominee Rivers and Lake Superior by

mercury. (Fig. 1). Based on this information it has been suggested

that people avoid eating fish from these waters more than once a

week. Because of the widespread nature of this highly toxic en¬

vironmental pollutant and its long half-life in the human system

(roughly two months), it is important to determine the level of

mercury contamination in man. To find out what has been happen¬

ing in the State of Wisconsin, contact was made with at least one

253

254 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

high school chemistry class in every county in the state to get four

samples per class for analysis.

Testing people for mercury can be done in a number of ways.

Samples of urine or blood can be analyzed; however, obtaining

200 to 300 of these samples through the mail is not practical.

Sampling people by use of a lock of hair is more feasible. Hair

readily picks up mercury from the body because of the presence

of sulfhydryl groups in its protein. Reaction of these groups with

dimethyl mercury, an extremely toxic form of mercury which is

now believed to be formed in the environment by the action of cer¬

tain bacteria on elemental mercury, can be written as :

2RSH »+ (CH3)2Hg - >- Hg(SR) 2 + 2CH4

1973] Zaborowski — Mercury Levels in Students 255

Because hair does not interact chemically with the body once it

grows out of the scalp, it constitutes a calendar of body chemistry.

However, it seems reasonable to assume that the average person’s

contact with mercury is relatively constant; thus any section of

hair should give a representative result.

The analysis of the hair, or any other appropriate material, for

mercury can be done by flameless atomic absorption. Prior to

analysis the sample is decomposed in concentrated sulfuric acid and

hydrogen peroxide followed by treatment with water and potassium

permanganate. After decomposition of excess permanganate with

hydroxylamine hydrochloride the Hg2+ is reduced to elemental

mercury by reaction with Sn2+ :

Sn2+ + Hg2+ - ^ Sn4+ + Hg°f

At this point air is bubbled through the sample solution and the

mercury vapor is carried through the cell and the instrument read.

EXPERIMENTAL

A Coleman MAS-50 Flameless Atomic Absorption Instrument

was used (any commercial or home-made spectrophotometer can

be used ; however, there must be ample room in the lamp compart¬

ment for placement of the absorption cell (8-10). For home-made

instruments a BOD bottle or glass acid bottle, of about 300 ml

volume, with a ground glass neck fitted with a fritted glass bubbler

can be used as a sample bottle).

The following chemicals were needed: 98% H>S04 ; 30% H;02 ;

0.15% NHoOH-HCl solution; 10% SnCL in 0.5 N H2S04. The

following mercury free solutions were needed (available from

Coleman Instruments) : 5% KMn04; 18 N H2S04; 4 N HN03;

0.15% NH20H-HC1; 10% SnCL ; mercury standard containing

1 jug Hg per ml.

The analyzer was standardized over the range to be used (0.1

to 3.0 ng is reasonable) by preparing samples with known total

mercury and noting the per cent transmittance. The Coleman

mercury free reagents were used to prepare the standard solutions.

The desired microgram amount of mercury standard was pipetted

into the sample bottle followed by the addition of 115 ml of distilled

water, 5 ml each of Coleman HN03 and Coleman H2S04. The solu¬

tion was then treated drop wise with Coleman 5% KMn04, with

swirling, to a “permanent” purplish-red color. Five ml of Coleman

NHoOH-HCl solution was added with swirling. After this addi¬

tion, if the sample did not turn clear in about 15 seconds,

NH2OH-HCl crystals were added until an uncolored solution was

obtained. Finally, 5 ml of Coleman SnCl2 solution was added and

the bubbler immediately inserted, making sure that the stopper

256 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

provided a good seal. The lowest per cent transmittance (% T)

was recorded.

Since mercury free reagents were not used (except for the

KMn04) on all the collected samples (for the sake of economy),

a blank was run to determine the mercury content of the laboratory

reagents. Twenty-five ml of concentrated H2S04 and 2 ml of 30 %

H202 were added to the sample bottle and warmed in a water pan

on a hot plate at 100 C for 4 hours, followed by cooling to room

temperature in an ice bath. Then 100 ml distilled water was slowly

added, in increments, while cooling. The solution was treated drop-

wise with Coleman 5% KMn04 to the “permanent” purplish-red

color and NH2OHHCl added, followed by SnCl2 as directed above,

and the reading taken.

The accurately weighed (to the nearest 0.0001 g) 0.1-0.2 g

sample of hair was used (roughly the amount in an average spit-

curl). It had been freshly washed, as hair sprays, tonic, etc. could

introduce a weight error. To the sample, in a 50 ml volumetric

flask or other suitable container, was added 25 ml concentrated

H2S04 and 2 ml of 30% H202, followed by warming to 60-70 C in

a water pan on a hot plate for % hr. The solution should finally

be pale yellow. After cooling, the solution was carefully washed

into a sample bottle with 100 ml of distilled water, cooled and

treated dropwise with the KMn04 solution to a “permanent”

purplish-red color. The analysis was completed with addition of

NH2OH*HCl and SnCl2 as described in the standardization section,

and the reading taken.

Once the standard samples had been run the transmittance data

were converted to log (1/%T) and plotted vs micrograms of

mercury, with a straight line resulting. The micrograms Hg in the

blank and in the samples were then determined from the graph.

The micrograms Hg in the blank was subtracted from the micro¬

grams Hg in each sample, the difference divided by the sample

weight to yield ppm Hg in the sample.

RESULTS

It is difficult to state a definite upper limit of safety regarding

the mercury level in humans. In addition, different individuals are

affected to a varying degree by the same mercury level. Ten parts

per million of Hg in hair is sometimes used as a guide, higher con¬

centrations indicating above normal contact with mercury. For

example, dental assistants have shown mercury levels of 12-45

ppm. A large sampling of the general public would show concen¬

trations of up to 7 ppm as common and persons who eat sea food

regularly as high as 40 ppm. Mercury poisoning occurs at about

1973]

Zaborowski — Mercury Levels in Students

257

PPM Hg (1970

FIGURE 2

175 ppm and above. Some symptoms of poisoning include loosening

of the teeth; tingling of hands, feet, or lips; ataxia; constricted

visual field; and emotional disturbances (11, 12).

The results of analyzing over 200 Wisconsin students showed

that the vast majority of them fell in the normal range with the

mode falling between 0 and 2 ppm (Fig. 2). Ten samples gave

results in the range of 21 to 68 ppm with one high value at 137

ppm. Some of these high results are in question as they came from

small hair samples, thus increasing the possibility of experimental

error. It is interesting to note that many of the hair samples high

in mercury came from people living close to one of those state

waters reported to contain fish high in that element. For example,

two samples from Wausau yielded values of 22.3 and 57.2 ppm.

One source of error in testing hair for mercury was the possibility

of contamination from mercurials, if present, in some hair sham¬

poos which might react with the hair surfaces. Tests of fifteen

different brands showed thirteen to contain essentially no mercury,

while two brands had 1.5 and 3.1 ppm Hg, levels low enough to

be considered insignificant.

258 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

REFERENCES CITED

1. AARONSON, TERRI. 1971. Mercury in the environment, Environment,

13: 16.

2. BOYLE, ROBERT H. 1971. The catch is, should you eat it? Sports Illus.

35 (July 12) : 46.

3. Environment Staff Report. 1971. Mercury in the air, Environment 13: 28.

4. GOLDWATER, LEONARD J. 1971. Mercury in the environment, Sci.

Amer., 224 (May) : 15.

5. GRANT, NEVILLE. 1971. Mercury in man, Environment 13: 2.

6. Mercury: Anatomy of a pollution problem, 1971 Chem. Eng. News, 49

(July 5) : 22.

7. KLEINERT, STANTON J., and DEGURSE, PAUL E. 1971. Mercury

levels in fish from selected Wisconsin waters, Research Report 73,

Wisconsin Department of Natural Resources.

8. HATCH, W. RONALD, and OTT, WELLARD L. 1968. Determination

of submicrogram quantities of mercury by atomic absorption spec¬

trophotometry, Anal Chem. 40: 2085.

9. KLEIN, DAVID H. 1972. Some general and analytical aspects of environ¬

mental mercury contamination, J. Chem. Ed., 49, 7.

10. RECHNITZ, G. A. 1962. Simplified atomic absorption spectrophotometer,

J. Chem. Ed., 39: 475.

11. STECHER, P. G. (Ed), The Merck Index, 8th ed., Rahway, New Jersey:

Merck and Company, Inc., 1968, p. 662.

12. WEAST, R. C., Handbook of Chemistry and Physics, 47th ed., Cleveland,

Ohio: The Chemical Rubber Company, 1966, p. B-120.

NITRATE CONTENT OF WELL WATER IN

WEST-CENTRAL WISCONSIN

Milan W. Wehking, James W. Pavlik,

Paul Strege and Dawn Gilles

University of Wisconsin —

River Falls

ABSTRACT

As a matter of local concern, a survey of wells in Pierce and St.

Croix Counties in west-central Wisconsin was undertaken. A total

of 124 wells in the two county area were included in the survey.

The nitrate content of these well waters was in the range 0-124

p.p.m. with an average value of 23.0 p.p.m. Of the wells sampled 20

(16.5%) were found to contain nitrate in excess of 45 p.p.m.

Examination of the data acquired on the water from these wells

revealed an inverse relationship between nitrate content and well

depth. The average depth of wells yielding water containing less

than 45 p.p.m. nitrate was 249 feet. Wells containing more than

45 p.p.m. averaged 188 feet. Further examination of the data

revealed that 26.3% of the wells with depths of 150 feet or less

yielded water with a nitrate content in excess of 45 p.p.m. In con¬

trast, only 12.2% of the wells with depths greater than 150 feet

exceeded this value.

The relationship observed was not perfect in that some very

deep wells contained large amounts of nitrate, while some shallow

wells were nearly nitrate-free. It is quite obvious that other factors

influence the entry of nitrate into underground waters.

The nitrate content of drinking water has aroused concern

among public health officials in recent years. This is particularly

true where ground water is serving as the source of drinking

water. It has been well documented that wells improperly located

with respect to barnyards, feedlots, cesspools or septic tanks are

subject to contamination by nitrogen from these sources (2, 3, 6,

7, 8). Agricultural fertilizers, especially those applied in the solid

form, are a potential source. All nitrate salts are water soluble

and the nitrate ion readily percolates through the soil. Much dis¬

agreement exists on the contribution that commercial fertilizers

259

260 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

make to the nitrate content of ground waters. In certain instances,

inorganic fertilizers clearly seem to be responsible for large quan¬

tities of nitrate in ground waters (5, 11). In nearly all of these

cases, however, it appears that overapplication of fertilizer or

application on very porous soils was responsible. An extensive

study carried out in Missouri gives some indication of the con¬

tribution of fertilizers to the nitrate content of ground waters (4),

at least in that geographical region. This investigation showed the

contribution of inorganic fertilizers to be low. On the other hand,

animal wastes or anhydrous ammonia may be applied to cropland

as a source of nitrogen. If these forms are used, it is necessary

that the nitrogen undergo oxidation to nitrate, if these sources

are to be effective plant fertilizers. The necessary oxidation is

catalyzed by enzymes which are produced by bacteria normally

present in soils. Nitrosomonas catalyzes the oxidation of ammonia

to nitrite which in turn is oxidized to nitrate via a reaction

catalyzed by Nitrobacter. These bacteria are autotrophic, synthesiz¬

ing organic carbon compounds from carbon dioxide, water and

getting their energy by the oxidation of ammonia and nitrite. As

a result, all nitrogen applied as fertilizer represents a potential

source of nitrate.

We must not, of course, overlook some of the more obvious

sources of nitrate. Decomposing vegetation represents a vast

potential source of nitrogen which eventually becomes nitrate.

The contribution from this source depends upon the amount of

organic nitrogen present, rate of decomposition, amount of vege¬

tative covering, rate of nitrogen uptake by this vegetative covering,

and quantity of rainfall.

It has also been shown that nitrate can be leached from geological

formations. Chalk and Keeney (1) reported that limestone con¬

tained significant quantities of nitrate. As the two county area

covered by the present investigation is underlain by limestone, this

also represents a potential source. Assignment of any portion of

the nitrate content of ground water to any specific source is difficult

and cannot be done reliably. Due to this difficulty, it appears the

controversy over inorganic fertilizers in this respect has just

begun.

Both humans and domestic animals are adversely affected by

excessive amounts of nitrate in drinking water. The U.S. Depart¬

ment of Interior Water Quality Criteria set 10 p.p.m. nitrate-

nitrogen (or 45 p.p.m. as ~N03 ion) as the permissible level in

drinking water for humans but set the desirable level at “virtually

absent’' (9). No standards for animals were established.

Nitrate ion per se is not harmful to the human body. The

presence of nitrate reductase-producing intestinal organisms, how-

1973] Wehking, Pavlik , Strege and Gillen — Nitrate Content 261

ever, leads to the formation of nitrite which is harmful. Members

of the coliform group of bacteria are capable of reducing nitrate to

nitrite and are believed to be responsible in the case of infants,

especially those whose diet is high in carbohydrates. When ab¬

sorbed into the blood stream, the nitrite bonds to the hemoglobin

forming nitrohemoglobin. This form of hemoglobin cannot carry

oxygen and its presence causes the victim to show symptoms of

oxygen deficiency. Nitrite may also oxidize the ferrous (Fe+2)

iron of hemoglobin to ferric (Fe+3) iron. The resulting methemo-

globin is incapable of carrying oxygen.

The primary objective of this investigation was to determine the

amount of nitrates present in well water in Pierce and St. Croix

Counties, Wisconsin. Geologically, these two counties are predomi¬

nated by Prairie du Chien dolomites, Platteville-Galena limestones

and St. Peter sandstones ; all three being of Ordovician age. These

Ordovician formations are underlain by Cambrian sandstones

from which major towns of these counties tap their water supplies.

These sandstones are being tapped from three hundred to four

hundred feet below the surface and consequently should have low

nitrate levels. The occurrence of glaciation after the deposition

of these Ordovician and Cambrian sediments, however, dumped

much glacial till down to depths of one hundred feet in places.

Many of the local farm wells are tapping this till due to its high

pore space. In this situation it might be anticipated that nitrate

levels would be higher than that of water found in rock formations

below.

As west-central Wisconsin is predominantly agricultural, the

ground water receives nitrates from the variety of sources men¬

tioned previously. It was the intent of this investigation to find

the proportion of wells containing excessive amounts of nitrate

and to determine if any relationship existed between nitrate con¬

tent and geographical location of the well and/or well depth.

EXPERIMENTAL

A total of 124 wells were included in this survey. The wells were

chosen randomly and all were within the boundaries of Pierce

and St. Croix Counties, Wisconsin. Water samples were collected

after the pump had been in operation for a few minutes to avoid

collecting water which had been standing in the well pipes for a

time. The samples were placed in dark containers and returned

to the laboratories for analysis. Information on the depth of the

well was also obtained at the time of sample collection.

Nitrate analyses were performed by the Brucine Method for de¬

termination of nitrate as described in Standard Methods for the

262 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

Lxamination of Water and Waste Water (10). Absorption meas¬

urements were made with a Spectronic 20 spectrophotometer at

410 nm. A calibration curve was prepared with a series of solu¬

tions containing known concentrations of reagent grade KN03. The

concentration of nitrate in the well water samples was then deter¬

mined directly from this calibration curve.

Duplicate analyses of each sample were performed to ensure

reproducibility. Every fifth sample analyzed was a solution of

known nitrate concentration. This was done to determine the re¬

liability of results obtained for the well water samples. The results

are expressed as p.p.m. ~N03, i.e. nitrate ion.

During the course of the investigation concern arose over the

effect of iron (Fe+2 and Fe+3) on the results of the nitrate

analyses. In several of the samples, a small amount of sediment

appeared after the sample vials had been allowed to stand undis¬

turbed. It was determined that this sediment contained a great

deal of iron. A series of solutions was therefore prepared contain¬

ing 22 p.p.m. nitrate and various concentrations of ferrous or

ferric iron. The effect of soluble iron on the results of the nitrate

analysis is demonstrated by the results in Table 1.

The effect of soluble iron on the results of nitrate analyses per¬

formed by the Brucine Method can be observed in the results

tabulated in Table 1. Presence of Fe was unimportant until the

concentration exceeded 10 p.p.m. As none of the well water samples

contained this much iron, it was of no further concern during

this investigation.

TABLE 1. EFFECT OF FERROUS AND FERRIC IRON ON

NITRATE DETERMINATION

1973] Wehking, Pavlik , Strege and Gilles — Nitrate Content 263

A summary of the data obtained during this investigation is

given in Table 2. Further information as to the frequency with

which specific nitrate concentrations occurred can be gained from

Fig. 1. An examination of this figure reveals that the nitrate con¬

tent of these well waters varied widely, with the majority contain¬

ing concentrations of less than 20 p.p.m. Nevertheless, 20 (16.5%)

of the wells yielded water containing nitrate in excess of 45 p.p.m.

Of the 124 wells included in this survey, information as to the well

depth was available on 68 of the wells. To determine if these were

representative of the total, a comparison of results obtained was

made (Table 2). This table shows that the mean, median and

range of nitrate concentrations found were nearly identical, indi¬

cating no difference between the two groups of wells.

The data acquired from those wells for which depths were known

were examined for relationship between well depth and nitrate

content. The data are summarized in Table 3 and reveal that such

a relationship does indeed exist. This table indicates that as well

depth increased, the average nitrate content decreased. This is by

no means a perfect correlation, as some of the deepest wells con¬

tained excessive nitrate, while some very shallow wells (<50 feet)

were nearly nitrate free. This might be expected, however, in an

area underlain by limestone, as faults in the subterranean struc-

TABLE 2. SUMMARY OF DATA

TABLE 3. RELATIONSHIP BETWEEN WELL DEPTH

AND NITRATE CONTENT

264 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

ture, of which there are many, allow surface water to reach depths

of several hundred feet quite rapidly. This table does indicate, how¬

ever, that a deep well is less likely to contain large amounts of

nitrate. Further evidence of this can be observed by an examina¬

tion of Table 4. Excessive quantities of nitrate are found with

DISTRIBUTION OF RESULTS

PPM NO"

3

FIGURE 1

TABLE 4. FREQUENCY OF EXCESSIVE NITRATE (OVER 45 P.P.M.)

1973] Wehking, Pavlik, Strege and Gilles — Nitrate Content 265

much greater frequency in wells which are less than 150 feet deep

than in those which are deeper (Table 4).

No relationship was found to exist between geographical loca¬

tion and nitrate content of wells. Those wells containing more than

45 p.p.m. nitrate were found to be distributed fairly evenly

throughout the two county area.

SUMMARY

In summary, one would have to say that no conclusions can be

reached from the results of this study as to the origin of nitrate

in ground water. It was, however, clearly evident that shallow

wells (<150 ft) have a greater probability of containing excessive

quantities of nitrate (i.e. in excess of 45 p.p.m.) than do deep wells

(>150 ft).

REFERENCES

1. CHALK, P. M. and KEENEY, D. R. 1971. Nature, 229, 42.

2. DEUTSCH, M. 1963. U. S. Geol. Surv. Water Supply Paper 1691, 71 pp.

3. GILLHAM, R. W. and WEBER, L. R. 1969. Jour. Water Pollution Con¬

trol Fed., 41, 1752.

4. KELLER, W. D. and SMIGH, G. E. 1967. Geol. Soc. Amer. Spec. Paper

No. 90, 59 pp.

5. “Poisoning1 the Wells,” Environment, J anuary-February, 1969, pp. 16-23,

45.

6. SMITH, G. E. 1965. Mo. Agr. Expt. Sta. Spec. Rept. 55:42-52.

7. SMITH, G. E. 1967. Amer. Assoc. Adv. Sci., No. 85, pp, 173-186.

8. STEWART, B. A., VIETS, F. G., JR., HUTCHINSON, G. L., and KEM¬

PER, W. D. 1967. Environ. Sci. Tech., 1, 736.

9. United States Department of Interior, Federal Water Pollution Control

Administration, Report of the Committee on Water Quality Criteria, U.S.

Dept. Int., 234 pp., April 1968.

10. Standard Methods of the Examination of Water and Wastewater, 12th

ed., 1965. American Public Health Association, Inc.

11. WARD, P. C. 1970. Nitrate and Water Supply: Source and Control,

Twelfth Sanit. Engin. Conf. Proc., Engin. Pub. Off., pp. 14-26, Univ. of

Ill., Urbana.

(

AN ORDINATION OF CORTICOLOUS LICHEN

COMMUNITIES IN THE POPPLE RIVER

BASIN OF NORTHERN WISCONSIN1

James A. Jesberger

University of Saskatchewan

Saskatoon, Saskatchewan

INTRODUCTION

Lichen communities of Wisconsin have been investigated over

broad geographic regions by Hale (1955) and Culberson (1955).

These and other studies by Barkman (1958), Beals (1965a), Brodo

(1961), Coleman et al. (1956), and Pearson and Lawrence (1965)

suggest that lichens are ecologically sensitive. This would seem

to indicate that an ecological investigation of them might best

be approached by an intensive examination of a small, environ¬

mentally uniform region. The Popple River Basin provides such

an area. It is small and has a comparatively uniform temperature.

The presence of the river and the nightly summer fog keeps the

humidity uniform during most of the growing season. The en¬

vironmental complexes presented by different forest types were

variables that could be controlled somewhat in selection of the

sites to be sampled. The vegetational and physical features of the

region are discussed in other papers of the Popple River series

and in Curtis (1959).

The present analysis of the lichen communities of the watershed

is on the basis of: (1) interspecific association of species of li¬

chens; (2) comparison of similarities and interrelationships among

the lichen communities on different tree species and in different

stands; (3) examination of the environmental specificity exhibited

by the lichen species.

FIELD METHODS

Ten forest communities along the Popple River in Forest and

Florence Counties, Wisconsin, were sampled in August, 1968. Each

stand was sampled by the quarter method (Cottam and Curtis,

1956), using five points per stand except in stand 2, where ten

1 This is paper No. 8 in the series “Studies on the Pine-Popple Rivers Area of

Northeastern Wisconsin.

267

268 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

points (40 trees) .were used. At each point the basal area and

identification of each of the trees sampled was recorded and an

estimate of the per cent canopy was made. Lichens were collected

from belt transects twenty inches wide encircling the tree, one

located at the base and one centered at breast-height (1.4 m.) on

the tree. The stands were picked for their accessibility and close¬

ness to the river, i.e., within a distance of one quarter of a mile.

The following is a list of the tree composition of each stand and

its Continuum Index value (C.I.) (Curtis, 1959) . The Continuum

Index value is calculated by multiplying the importance value of

each tree species in the stand (a value obtained by integrating

size of individuals, number of individuals, and ubiquity of dis¬

tribution) by the adaptation number of the species (an assigned

ranking from 1 to 10 depending upon association with bur oak

or sugar maple) . The resulting weighted values are then summed

to give an index value which ranges from 300 for a pure stand

of bur oak to 3000 for a pure stand of sugar maple. These represent

a continuum from xeric to mesic habitats and the C. I. index num¬

ber of each of the ten following stands will give an idea of the

approximate position of the stand along this continuum.

STANDS : Stand 1 : Lowland swamp, canopy 70%, C.I. 1254 ;

Thuja occidentalis (35%), Abies balsamea (24%) , Picea glauca,

Acer rubrum, Fraxinus nigra, and Betula lutea. Stand 2 : Low¬

land site, canopy 90%, C.I. 2714; with Acer saccharum (70%),

Betula lutea (10% ) , Ulmus americana (10% ) , and Os try a vir-

giniana. Stand 3: Upland site, C.I. 588; with Pinus resinosa

(80%) , Betula papyrifera, and Populus tremuloides. Stand 4 :

Lowland site, dense ground layer of ferns, canopy less than 50%,

C.I. 1573 ; with Prunus serotina (40%, Picea glauca (15% ) , Pop¬

ulus tremuloides (25% ) , Acer rubrum (15% ) , and Betula lutea.

Stand 5 : Upland site, canopy less than 50%, C.I. 1567 ; with Pinus

resinosa (50%) , Ulmus americana (25%), Prunus serotina, Abies

balsamea, and Acer rubrum. Stand 6 : Lowland site, dense ground

cover of ferns, C.I. 1375 ; with Populus tremuloides ( 90 % ) , Prunus

serotina, and Picea glauca. Stand 7 : Mixed, dry site, C. I. 1627 ;

with equal amounts of Pinus strobus, Picea glauca, Populus tremu¬

loides, and Betula papyrifera. Stand 8 : Lowland site, dense

growth of ferns and Viburnum, C.I. 1350 ; with predominantly

Populus tremuloides and some saplings of Ulmus americana and

Fraxinus nigra. Stand 9 : Upland site, little ground cover, canopy

90%, C.I. 2508 ; with Acer saccharum (60% ) , Ulmus americana

(15%), Tilia americana (15%), and Populus tremuloides. Stand

10 : Lowland swamp, C.I. 1405 ; with Thuja occidentalis (25%) ,

Abies balsamea (55% ) , Acer rubrum (10% ) , and Picea glauca.

1973] Jesherger — Lichen Communities in Popple River

269

STATISTICAL METHODS

The lichens collected at breast-height and base were treated

separately. Frequency per stand was calculated in percentage of

belt transects on which the lichen occurred in the stand sampled.

The frequency per species of tree was calculated by the per cent

with which each of the 99 species of lichens occurred on a given

species of tree in the total of 220 trees studied. The lichen species

with frequencies greater than 2.3% are given in Table 1 together

with their frequencies at the base and at breast-height on the 12

most abundant tree species. The frequencies of the less frequent

species are given later in the paper.

Cole’s index values of interspecific association (Cole, 1949)

were calculated for those species attaining a frequency greater

than or equal to 2.3% and are given in the upper right of Tables

2 and 3. These values will range from +1 for a perfect positive

association (the one species always associated with the other)

to a —1 which would mean that the two species were never found

together. A 0 value would mean that they were randomly asso¬

ciated. As an example in Table 2 Caloplaca cerina gives a value

of 1.0 with Caloplaca holocarpa and was thus always associated

with it, but with Gr aphis scripta it gives a value of —1.0, indi¬

cating that the two were not found together. On the other hand

Caloplaca cerina is randomly associated with Candelaria concolor

as shown by the value 0 in the appropriate position in Table 2.

To test the statistical significance for all joint occurrences for

which the Cole’s index values were computed, the Chi-square values

with Yates Correction Factor (Yates, 1934) were calculated by

the usual 2x2 contingency table. These Chi-square values are

presented at the lower left of Tables 2 and 3. A Chi-square value

of 3.841 or higher is significant at the 5% level, and one of 6.635

or higher is significant at the 1% level.

An index of similarity, a percentage measure of the vegetational

affinities between stands or trees, ranging from 0 to 100, was

calculated for the 12 most common tree species and the ten dif¬

ferent stands. The index was calculated according to the formula

2 w"

— k a discussion of which can be found in Beals (1960, 1965a)

or Bray and Curtis (1957). A comparison of the lichens occurring

at the base and those occurring at breast-height in the stands was

also made using the index of similarity. Chi-square values with

Yates Correction Factor were calculated to test the statistical sig¬

nificance of joint occurrence or lack of joint occurrence of the lichen

species of Table 1 with the common tree species of the study. The

270 Wisconsin Academy of Sciences, Arts and Letters [VoL 61

TABLE 1. FREQUENCIES AT BREAST-HEIGHT AND BASE ON THE

12 COMMON TREE SPECIES OF THE 48 LICHEN SPECIES

SHOWING THE HIGHEST FREQUENCY IN THE STUDY.

UPPER VALUES GIVEN FOR EACH LICHEN ARE ITS

FREQUENCY AT BREAST-HEIGHT ON THE TREE,

THE LOWER VALUE DESIGNATES ITS FRE¬

QUENCY AT THE TREE BASE.

(Table I continued on next page)

1973] Jesherger — Lichen Communities in Popple River

271

Table I. (Continued from preceding page, with headings abbreviated.)

results are shown in the glyphs in Figs. 1 and 2. To read these

glyphs first examine the legend at the bottom of Fig. 1. As an

example take Parmelia sulcata (P. sul.) toward the center left

of the figure. The glyphs indicate positive association with Acer

rubrum , negative association with Populus tremuloicles, Prunus

serotina , and Acer saccharum when the glyphs are read in clock¬

wise fashion. Likewise the glyphs on Rinodina halei (R. h.) at

the lower right indicate negative association with Populus tremu-

loides and positive association with Acer saccharum and Ulmus

americana.

SPECIES INTERRELATIONSHIPS

The ordinations in Figs. 1 and 2 were derived by the methods

used by Beals (1965a, 1965b). The axes of the ordination are not

in themselves significant but are only useful to graph the points.

It is the interrelationship of the points alone that is useful. The

closer the points are grouped together the more closely the species

are associated. The more distant the points, the more negative is

the association of the species. For example on Fig. 1 there is a

closely grouped number of points in the center with such species

as Ramalina fastigiata, Buellia punctata, Parmelia subaurifera,

Parmelia aurulenta, Lecanora coilocarpa, Usnea comosa, etc.

272 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 2. MATRIX OF COLE’S INDEX VALUES AND CHI-SQUARE

VALUES WITH YATES CORRECTION FACTOR FOR LICHEN

SPECIES WITH A FREQUENCY GREATER THAN OR EQUAL

TO 2.3% AT BREAST-HEIGHT ON THE TREE. COLE’S

INDEX VALUES ARE IN UPPER RIGHT HALF, CHI-

SQUARE VALUES ARE IN LOWER LEFT HALF.

Lichen

species

- »-

o o w b x -i -i

5 n

•H

S3

o

3

a

o

+>

s

r—S

E

3

1973] Jesberger — Lichen Communities in Popple River 273

274 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

TABLE 3. MATRIX OF COLE’S INDEX VALUES AND CHI-SQUARE

VALUES WITH YATES CORRECTION FACTOR FOR LICHEN

SPECIES WITH A FREQUENCY GREATER THAN OR EQUAL

TO 2.3% AT THE BASE OF THE TREE. COLE’S INDEX

VALUES ARE IN UPPER RIGHT HALF CORNER, CHI-

SQUARE VALUES ARE IN LOWER LEFT HALF.

20.

21.

22.

23.

2l+.

25-

26.

27.

28.

29.

30.

31.

32.

33.

3k.

35.

36.

37.

36.

39.

1+0.

O. pulicaris 0

P. auruienta .07

P. caperata

P. orinita

P. galbina 6.77

P. rudecta 1.25

P. saxatilis

P. subaurifera 1.87

P. sulcata 9.00

P. laevigata 0

P. multipuncta 0

P. ciliata

f. ciliata

?. ciliata

f.fibrillosa 8.91

P. grisea 1.1+

P. orbicularis ,

f. orbicularis ‘i‘°3

P. orbicularis

f . rubr opulchra

1.15

R. fastigiata 0

R. halei .95

R. pyrina 0

U. comosa 0

X, polycarpa .38

1. 86

0

.01

.21+

.52

.81+

0

.15

.1*

.33

1.98

8.7

1.97

0

1.35

o

0

.08

1+1+ 1.55

o o

0

kh

.10

.55

o

2.91 1.17

3.75 .18

o

0

.06

-.05

o

.05

o

0

'.hh

o

0

.20

.33

0

.61+ .93

2.1+0 7.93

2.53 .02

0

.67

0

.58

• Hi

2.59

0 0 .08

0 0 .05

.82 .17 11.08

3.75 5.37 7.58

0 3.16 1.38

0 2.3U 12.H+

.1+1 .56 1.12

3.05 .18

.81+ .05

0 10.19

o

2.76

0

1.67 12.30

0 0

.21+

0

.83

2.59 6.91

0 0

.25 0

0 12.20

.51 13.79

i.ol+ 1+5.60

5.03 17.22

0 8.75.

1.1+9 0

1.79 1.39

0 .98

0 0

.57 .38

0

.17 1.92

0

1.72 6.76

.01+ .97

1.1+1+ 9.12

.16 .87

.20 1.03

.08 2.15

.93 1.55

0 .07

0

.08

0

.29

0

.06

0

.37

.52 1,80

25

0 .1+8

0 0

.11+ ,30.1+6

0 .32

0 .03

.22 1.1+1+

0 6.67

.08 2.92

.16 .56

0 0

0

.02

0

1.01

0

3.81+

8.95

0

.01

0

1.01

0

.06 3.51 13.

0 .28

.33

.0!+

• 36

0

.17 18.02 I+.78 6.63

0

.1+8

.37

0 11

0

0 15.

2.05 20

.17 5.52

0 .23

,11+ 0

.03 2.62

.58 1.1+9

.02 3.25

.16 0

1.11

.1+2

0

1.1+0

0

.75 2.25

.07

1.22

.28

.17

O

0

.18 .60

5.57 3.99

.07 1.73

0 0

0 .22

.59 0

0 11.15

0 .10

.1*7 .69

5.07 1+.31

I.02 13,

.59 8.03 6

.22 .11+

0

1.26 1

6.11

.13

.1+1

0

39.79 1.57

.03 0

2.81+

.12

0

.1+9

0

,80

,1+8

,81

,66

,21+

0

.33

.17

.79

0 8

0

0 8,

0 2.

0

0 13.

0

51 .20

05 1.38

61 .01

13 .06

05 0

02 28.67

25 .01+

1973] Jesherger — Lichen Communities in Popple River-

275

polycarpa

276 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

«r

A . c.

B.s. b.1.

i-t

C.cer ,

L . ve .

E ,m.

P . orb . rub .

i<

L .va .

^ ;

A

H.p.

1 P . sax .

C.s. P . c i 1 . c i 1 .

P .gr

B.c.

P . su 1 .

R;f‘* iB.p.

1% 1 L,CV • P.au.

.sub. *» * ^ R-P* ®

•j* ix p C.conc.

L) . c . P • ga • P>C1- 1 .f.fib.

C.p.

P.r.

**0.p.

P . cr .

P .mu 1 .

C . cr . ^

L.s.

*

BASE

f ^

C .com . * c .ch

P . cap.

G.s.

P .or b . orb ,

t-

R.h.

FIGURE 1. Two dimensional ordination of lichen species at the tree base

on the basis of their Cole’s index value. Glyphs indicate statistically signifi¬

cant positive or negative association with the common tree species. Position

and shape of glyphs for the different tree species and the sign indicating

negative association are given in the table below the ordination. See the text

for further directions. Initials of lichen species are from lichens of Table 1.

1973] Jesberger — Lichen Communities in Popple River

277

FIGURE 2. Two dimensional ordination of lichen species at breast-height

on the trees on the basis of their Cole’s values. Symbols the same as in

Figure 1.

showing closer association to each other than to the group of

species toward the top of the ordination. The latter group encom¬

passed by Bacidia suffusa , Caloplaca cerina, Physcia ciliata, and

Physcict orbicularis presents another such group. Although an

ordination only approximates the species interrelationships that

are present in the ecosystem (Beals, 1965b), examination of Figs.

1 and 2 reveals a number of significant species groups or micro¬

environments.

Considering the parameters that control the distribution of the

various lichen species and the extreme ecological sensitivity of the

lichens to these parameters, it seems logical to assume that species

which occur close together on the ordination do so because of simi¬

lar microenvironmental specificity. Microenvironmental specificity

is that property of a lichen species which restricts its distribution

to microenvironments in which the physical and biological proper-

278 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

ties satisfy the physiological requirements of the lichen through¬

out the year, especially during critical periods of growth in the

spring and fall.

A comparison of Figs. 1 and 2 shows that species interrelation¬

ships at breast-height are similar to those found at the base.

Some of the lichen species exhibit a high degree of microenviron¬

mental specificity as shown by their negative association with

several tree species, and positive association with only one tree

species. Other lichen species find suitable microenvironments on

several tree species. The lichens on Populus tremuloides suggest

that several microenvironments may exist in environmental com¬

plexes occupied by P. tremuloides. This is shown, for example,

by the spacing between two groups of lichen species that show

positive association with Populus tremuloides. One group may be

found toward the upper right in Fig. 1 and more widely spaced

but toward the upper right in Fig. 2 and is composed of Caloplaca

cerina, Physcia ciliata f. ciliata, and Physcia grisea. The other

group is composed of Xanthoria poly car pa and Physcia ciliata f.

fibrillosa and is toward the center in Fig. 1 and toward the right

center of Fig. 2. Other examples of such situations can be seen on

examination of Figs. 1 and 2. Microenvironmental specificity is

further considered in the Discussion.

DISTRIBUTION OF MICROENVIRONMENTS

The ordination of Figs. 3, 4, 5 and 6 were derived from the

index of similarity values according to the method used by Beals

(1960, 1965a) and Bray and Curtis (1957). They provide a spatial

ordering in three dimensions based on a simple expression of the

amount each tree or stand has in common with all the other trees

or stands. As before, the axes of the ordination are not in them¬

selves significant. It is the interrelationships of the points that

are meaningful.

In Figs. 3 and 4, the stand interrelationships are different be¬

tween the base and breast-height (e.g. stands 3, 4 and 5 are far

apart when the base communities are plotted, and close together

when the breast-height communities are plotted). This suggests

that the microenvironments at the base and at breast-height are

not necessarily correlated. In Figs. 5 and 6, the pattern of the

species is remarkedly similar, suggesting that gross environ¬

mental factors are operating similarly for the base and the breast-

height.

Stands occurring close together on the ordination would be ex¬

pected to possess environmental complexes with closely related

microenvironments. Thus an analysis of the distribution of rare

species among stands would give a greater degree of biological

1973] Jesherger — Lichen Communities in Popple River

279

FIGURES 3, 4, 5 and 6. Three dimensional ordination of stands (Figures

3 and 4) and trees (Figures 5 and 6) on the basis of their lichen communities.

Radii of the circles indicate magnitude of position in the 3rd axis. Initials

for tree species are for those tree species of Table 1.

validity to the ordination and support the previous statement.

Supporting this reasoning is the observation that a rare species

is so because it exhibits a high degree of specificity for a micro¬

environment that is rare. Thus, if two stands have a larger num¬

ber of rare species in common, it can be assumed that their en¬

vironmental complexes provide similar microenvironments. The

proximity of Stands 1 and 10 in Figs. 3 and 4 suggests that they

possess similar environmental complexes with closely related mi¬

croenvironments. Analysis of the occurrence of rare species shows

that these stands have seven rare species in common. Conse¬

quently their arrangement in the ordination has increased bio¬

logical validity. This idea does not hold true for all comparisons

280 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

of position in the ordination. Stands 2 and 9 have no rare species

in common. This, however, could be due to the age difference of

the stands. The trees of Stand 2 have an average basal area twice

that of trees from Stand 9. Curtis (1959) stated that as a woods be¬

comes more mature, its interior environment changes. The two

stands have only seven common species in common. Stand 5, which

is placed in the middle of both ordinations (see Figs. 3 and 4), has

one of the most diverse lichen floras and also is the most diverse

stand with regard to tree composition.

An analysis of the spatial arrangement of trees species was

attempted in order to explain the positioning of tree species that

was arrived at in Figs. 5 and 6. The arrangement is close to that

found by Curtis (1959), when he ordinated the trees of the forests

of northern Wisconsin.

When the occurrence of the rare species on the trees is com¬

pared with the ordinations of the tree species, the occurrences

of the rare lichens are not necessarily correlated with the prox¬

imity of the tree species on the ordination. A difference can be

seen in the ordination position of some of the tree species, when

ordinations at breast-height and base are compared. This is prob¬

ably a result of the environmental complex at the two levels vary¬

ing more sharply in some stands than in others. Trees that occur

close together in Figs. 5 and 6 can be seen to be close together in

the glyphs on the ordinations of Figs. 1 and 2.

ADDITIONAL OBSERVATIONS

Acer saccharum supported the greatest number of species,

45, followed by Populus tremuloides, 42, Ulmus americana, 39,

Thitja occidentalis , 38, and Prunus serotina, 36. Pinus resinosa,

one of the most abundunt tree species, had only 28 species of

lichens on it. Stand 1 with 49 species supported the most species

of lichens, followed by Stand 5 with 48, Stand 4 with 45 and

Stand 10 with 42. Stands 2, 6, 8 and 9 had the lowest num¬

ber of lichen species with 25-27. These stands were also the lowest

in diversity of trees.

Thuja occidentalis and Acer saccharum were found to support

the greatest number of rare species, 15 and 14 respectively. Nine

of those on T. occidentalis were unique to Thuja , and 4 of the 6

on Populus tremuloides were unique to Populus. Every stand had

at least one rare species that was unique to it except for Stands

3 and 6. All stands had at least two rare species, with Stands 1

and 10 having the greatest number, 15 and 12.

A definite preference for environmental complexes is suggested

in the distribution of the different lichen growth forms. Stand 1

was found to support the most fruticose species at breast-height,

1973] Jesberger — Lichen Communities in Popple River 281

9, while Stand 10 supported the most fruticose species at the

base, 10. Stands 6, 8 and 9 had no fruticose species at the base,

while Stand 2 had only one. Stand 1 supported the greatest num¬

ber of crustose species, 14 at each level. Within Stand 5 the

greatest number of foliose species were found, 22 at breast-height

and 18 at the base.

Of the 99 species found, 65 occurred at both levels while 19

were restricted to breast-height. Calculation of index of similarity

values showed that Stand 3 was the most uniform in composition

from base to breast-height, having an index value of 91. Stand 7

exhibited the greatest difference between the two levels with an

index of only 56. Stand 5 had the most species in common be¬

tween the two levels, 26. The index of similarity between the base

and breast-height for all the trees was 70. The above observa¬

tions will be further evaluated in the Discussion.

DISCUSSION

Microenvironmental specificity can be seen as the controlling

force in the distribution of lichen species. Though the glyphs in

Figs. 1 and 2 suggest that the lichens are distributed according

to specificity for a particular tree species, Culberson (1955)

found that on examination of the physical characteristics of the

bark, the results obtained could not fully explain the distribu¬

tion of lichen species. The microenvironment inhabited by the

lichen results from the interaction of the physical and biological

factors of the environmental complex in which the tree occurs.

Microenvironmental specificity being a property of a lichen

species, it can be assumed that no two species have exactly the

same microenvironmental specificity. Hale (1955) in examining

the distribution of lichens along the continuum (Curtis, 1959)

found that no two species had identical peaks on the continuum,

i.e., no two species had the same amplitude of tolerance to the

environmental complexes occurring along the continuum, even

though they did overlap in varying degrees.

A wide variation was observed in the number of lichen species

that a particular tree species or stand supported. The reason for

this is either that a stand or tree supports a large number of

lichen species because it possesses a large number of microenviron¬

ments, or else a large number of the species present have ampli¬

tudes of tolerance that overlap. The real situation probably re¬

sults from a combination of the two situations.

From these results it can be suggested that future investiga¬

tions of distribution patterns should be directed at delimiting

the nature of the parameters that control lichen distribution, and

investigation of the amplitude of tolerance of the individual lichen

282 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

species to each of these parameters. If the above approach is used,

a better understanding of the complex nature of microenviron¬

mental specificity may be achieved.

ADDITIONAL SPECIES REPORTED

Those species previously unreported from Wisconsin are indi¬

cated by*.

The following species were found on rocks in and along the

Popple River: Arthonia sp., Bacidia albescens (Kremp.) Zw.*,

Caloplaca f randans (Th. Fr.) Oliv., Catillaria chalybaeia (Borr.)

Mass., Dermatocarpon fluviatile (G. Web.) Th. Fr., Lecanora

caesiocinera Nyl., Lecanora polytropa (Ehrh.) Rabenh., Lecidea

cineroatra Ach., Rinodina ioivensis Zahlbr., Sphinctrina gelasinata

(With.) Zahlbr.*, Staurothele clopima (Wahlenb.) Th. Fr., Rhizo-

carpon sp.

The following species were found on burned decomposed stumps

along the Popple River: Alectoria nidulifera Norrl., Bacidia chlo -

rococca (Grawe) Lett., Cladonia fimbriata (L.) Fr., Cladonia

gracilis (L.) Willd., Cladonia verticillata (Hoffm.) Schaer., Leca¬

nora symmicta (Ach.) Ach., Lecidea berengeriana (Mass.) NYL.,

Lecidea carnulenta (Tuck.) Fink*, Lecidea scalaris (Ach.) Ach.

The following species were found on the ground and on moist

moss-covered logs : Cladonia grayi Merr., Cladonia mitis Sandst.,

Cladonia rangiferina (L.) Wigg., Peltigera canina (L.) Willd.,

Peltigera horizontalis (Huds.) Baumg., Peltigera polydactyla

(Neck.) Hoffm., Stereocaulon tomentosum Fr.

The following species were found on trees in the study area or

on trees along the river. Frequencies are given for those lichen

species that were found on trees used in the statistical analysis

of lichen communities. Alectoria americana Mot. 1.3%, Anaptychia

palmatula (Michx.) Vain, Anaptychia pseudospeciosa Kurok. 5%,

Arthonia radiata (Pers.) Ach., Bacidia bacillifera (Nyl.) Fink

1.2%*, Bacidia fuscornbella (Hoffm.) Bausch. 1.2% Bacidia in-

compta (Borr.) Anzi. 0.9%, Bacidia schweinitzii (Tuck.) Schneid.

0.5%, Bacidia sphaeroides (Dicks.) Zahlbr. 0.9%, Bacidia stig-

matella (Tuck.) Zahlbr. 1.8*, Biatorella microhaema Norm.

0.5%*, Buellia disciformis (Fr.) Mudd, Caloplaca aurantiaca

(Lightf.) Th. Fr. 1.8%, Cetraria ciliaris Ach. 1.3%, Cetraria halei

Culb. 1.4%, Cetraria oakesiana (Tuck.) 0.5%, Cladonia botrytes

(Hag.) Willd. 0.5%, Cladonia capitata (Michx.) Spreng. 0.5%,

Cladonia crispata (Ach.) Flot. 0.9%, Cladonia fimbriata (L.) Fr.

1.3%, Collema conglomeratum Hoffm. 0.5%, Conotrema urceola-

tum (Ach.) Tuck. 1.8%, Haematomma cismonicum Beltr. 0.5%,

Lecania dimera (Nyl.) Th. Fr. 1.3%*, Lecanora coilocarpa (Ach.)

Nyl. 2.0%, Lecanora hypoptoides Nyl. 1.4%*, Lecanora pallida f.

1973] Jesberger — Lichen Communities in Popple River 283

pallida (Schreb.) Rabenh. 1.4%, Lecanora pallida f. rubescens

Imshaug and Brodo, Lecidea cadubriae (Mass.) Hedl.*, Lecanora

sambuci (Pers.) Nyl. 0.5% *, Lobaria quercizans Michx. 0.9%,

Lobaria pulmonaria (L.) Hoffm. 0.9%, Micarea prasina (Fr.)

Korb. 1.3%, Ochrolechia pallescens (L.) Mass. 0.9%, Pachyphiale

fagicola (Hepp.) Zw. 0.5%*, Parmelia bolliana Mull. Arg. 0.9%,

Parmelia f randans Nyl. 0.5%*, Parmelia ulophyllodes (Vain.)

Sav. 0.9%, Peltigera aphthosa (L.) Willd. 2.0%, Peltigera canina

(L.) Willd. 0.5%, Peltigera polydactyla (Neck.) Hoffm. 0.5%,

Pertusaria amara (Ach.) Nyl. 0.9%, Pertusaria pertusa (L.)

Tuck. 1.4%, Pertusaria pustulata (Ach.) Duby 0.5%, Physcia

millegrana Degel. 1.3%, Physcia orbicularis f. albociliata Thoms.

0.9%, Physcia stellaris (L.) Nyl. 1.3%, Physcia tribacoides Nyl.

0.9%, Pyrenula nitida Weig. 0.9%, Pyxine sorediata (Ach.) Mont.

0.9%, Rinodina ascociscana Tuck. 0.5%, Usnea cavernosa Tuck.

0.45%, Usnea dasypoga (Ach.) Rohl. 0.45%, Usnea glabrescens

(Nyl.) Vain. 0.9%, Usnea substerilis Mot. 0.45%, Usnea trichodea

Ach., Xanthoria fallax (Hepp.) Arn. 1.3%.

FUTURE CONSIDERATIONS

The species of lichens present in the Popple River Basin indi¬

cate that no air pollution has occurred in the area studied (Skye,

1968). Many of the species present are dependent on the main¬

tenance of the present environmental complexes and could be ex¬

pected to change their distribution as the environmental complexes

change. Such man-made catastrophes as fire and cutting could

also alter the environmental complex and cause a change in the

lichen communities.

ACKNOWLEDGEMENTS

I wish to thank Dr. J. W. Thomson for his generous assistance

in taxonomic problems and in the preparation of this manuscript

and also for the encouragement that he offered throughout the

study. I also wish to thank Frank Barth for his help in the col¬

lection of field data, Dr. Edward Beals for the advice he offered

on statistical and ordination techniques and for critically reading

the manuscript, and Dr. Grant Cottam for the suggestions he of¬

fered throughout the study and for critically reading the manu¬

script.

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BEALS, EDWARD W., 1960. Forest bird communities in the Apostle Islands

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- , 1965b. Species patterns in a Lebanese Poterietum. Vegetatio 13:

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BRAY, J. R., and J. T. CURTIS, 1957. An ordination of the upland forest

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BRODO, I. M., 1961. A study of the lichen ecology in central Long Island,

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COLEMAN, BABETTE BROWN, W. C. MUENSCHER and D. R. CHARLES,

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COTTAM, GRANT, and J. T. CURTIS, 1956. The use of distance measures

in phytosociological sampling. Ecol. 37 : 451-460.

CULBERSON, W. L., 1955. The corticolous communities of lichens and bryo-

phytes in the upland forests of northern Wisconsin. Ecol. Monogr. 25:

215-231.

CURTIS, J. T., 1959. The vegetation of Wisconsin: an ordination of plant

communities. Madison, Wisconsin.

HALE, M. E., 1955. Phytosociology of corticolous cryptogams in the upland

forests of southern Wisconsin. Ecol. 36: 45-63.

PEARSON, L. C., and D. B. LAWRENCE, 1965. Lichens as microclimate

indicators in northwestern Minnesota. Amer. Midi. Nat. 74: 257-268.

SYKE, ERIK, 1968. Lichens and air pollution. Acta Phytogeographica

Suecica. 52.

YATES, F., 1934. Contingency tables involving small numbers and the Chi-

square test. J. Roy. Stat. Soc. Suppl. 1: 217-235.

ABOUT THE AUTHORS AND THEIR PAPERS

FRANK L. KLEMENT, Professor of History at Marquette University,

Milwaukee, Wi, is “without doubt the best informed student of the

American Civil War in Wisconsin. Characteristically . . . this work is

another example of his digging out an unusual vignette in ground that

has been combed fine by historians ... To the scholars and buffs who

have spent hours in the Gettysburg literature and trod over the rolling

field and woodlands Element’s manuscript reawakens memories of the

somber November day when President Lincoln participated in the dedi¬

cation ceremonies . . . Professor Element has turned up some new

material about the battlefield that will honor Wisconsin so long as men

remember the Iron Brigade.”

Such are the words of the reviewer, John Patrick Hunter, him¬

self a Civil War buff and President of the Civil War Round Table of

Madison. Our readers will value his judgment and enjoy Prof. Element’s

paper the more for knowing that he and Mr. Element are friends.

JOHN B. RAY, Professor of Geography-Geology at the University of Wis¬

consin — Whitewater has struck a timely note ! Now we can understand

why Chile contests the presence of American fishing boats and we will

be alerted to the much more important problems of Ocean Space with

which the Geneva Convention will have to grapple in the spring of 1974!

MAURICE E. PERRET, Professor of Geography at the University of Wis¬

consin — Stevens Point contributes some very personal geography — the

antithesis of Ocean Space geography. Wisconsin readers, many of us

with ethnic and cultural backgrounds of those very groups in Portage

County, will look around us for survival of cultural features in our

own counties. And as The Bicentennial of 1976 approaches we can take

pride in the pioneer heritage of our State and Nation.

JOHN M. GILLETT AND THEODORE S. COCHRANE. Dr. Gillett is

now of the National Museum of Canada, Botany Division, Ottawa,

Ont., Canada. This is paper No. 836 of the Plant Research Institute,

Canada Department of Agriculture. Dr. Cochrane is Curator I of the

Herbarium, Department of Botany, University of Wisconsin — Madison.

The paper is also No. 63, as part of a longtime series on the Flora

of Wisconsin, initiated in the 1920’s under Prof. Norman Fassett and

currently under direction of Prof. Hugh litis. For the information

of our botanical readers many of the earlier papers in this series have

been published in the Transactions.

A. W. SCHORGER, Emeritus Professor of Wild Life, University of Wis¬

consin — Madison. It is sad to record that this is the last paper of Arlie

W. Schorger to appear in the Transactions. It was received and ac¬

cepted before his death on May 26, 1972, and is virtually untouched

by the Editor. We are glad to publish it as he wished, and we call

attention to his many and valued works which have appeared in

Transactions.

285

286 Wisconsin Academy of Sciences, Arts and Letters [Vol. 61

CARL E. KROG, Assistant Professor of History at the Marinette Campus,

University of Wisconsin — Marinette. This lively account of the early

days in Marinette is interesting, as we look forward to the Bicentennial

year 1976. Is there nothing new in Wisconsin? Prepaid medical care,

hospitals struggling to survive in a sparsely settled area, concern over

safe drinking water and contaminated foods are problems today as they

were in 19th Century Marinette. Your Editor (a bacteriologist) agrees

that “Germs” should be the first word of the title, since they precede

these problems!

R. V. SMYTHE AND H. C. COPPEL. Prof Coppel and his associate Dr.

Smythe, now entomologist at the Wood Products Laboratory, Gulf¬

port, Miss, have provided a fascinating record of confrontation be¬

tween termites and thief ants. To some of us it comes as a surprise

that termites are found in the wild in Wisconsin, but, if so, it is well

to know that their enemies are not far away.

PETER S. WENZ, Professor of Philosophy at the University of Wisconsin —

Stevens Point has provided a thoughtful and logical explanation of a

seeming discrepancy in Plato’s writings of Socrates. Few citizens of

today would go as far as Socrates in viewing our relation to our

government nor would we think of the citizen’s relation to the state

in terms of philosophy. But then we are not Socrates and it is good

to have Prof. Wenz to analyze for us.

D. C. ROUSAR AND A. M. BEETON. Drs. Rousar and Beeton are both

on the staff of the Center for Great Lakes Studies, University of Wis¬

consin — Milwaukee. This paper is typical of the comprehensive studies

now going on concerning the Great Lakes, particularly Lake Michigan.

It will be from such data that the pollution status of Lake Michigan

will be ascertained and hopefully controlled before it is too late.

THOMPSON WEBB III. Dr. Webb is now of the Department of Geological

Sciences, Brown University, Providence, R. I. To an amateur like

this Editor, it has always been amazing that pollen grains are pre¬

served, let alone are recognizable in the debris of lake sediments. But

fortunately they are so and thus analysis of their numbers and kinds

is helpful in assessing the changes induced by man’s settlement of the

area. Every bit of evidence should be examined as it may bear upon

the rate of such change. Hopefully then we may see a way to resist, to

stabilize the desirable diversity that once existed.

RICHARD H. KEEHN, Assistant Professor of Economics, Division of So¬

cial Sciences, University of Wisconsin — Parkside, Kenosha, Wi, has as¬

sembled the banking history of Wisconsin for a period before the memory

of most present bankers. But the paper tells more than the history —

it shows the effects of market structure including the very local market

of many of the early banks. To those interested in bank regulation,

both state and federal, there should be interesting points bearing upon

the reasons for present regulations.

1973]

About the Authors and Their Papers

287

EDMUND RONEY, Associate Professor of Drama at Ripon College, Ripon,

Wi, has given us a tantalizing story of the controversy over the play

Le Cid and in his covering letter he said it was “one chapter of a de¬

tailed analysis of the causes and nature of La Querelle du Cid.” One

wonders (and looks forward to future papers of Prof. Roney) what

more can there be?

STANLEY A. NICHOLS is a Specialist in the Environmental Resources

Unit of the Extension Division, University of Wisconsin — Madison,

and as such has some important things to say to the public. Seeing

the overgrowth of aquatic weeds, we might clamor for cutting them

ruthlessly, as we do the terrestrial weeds in our gardens. It takes an

environmentalist to see in perspective the large weeds and the tiny

but potent algal weeds. When he concludes from a 3-year study, that

the macrophyte: algal relations are different in shallow vs deeper water,

it gives one pause. The problem is complex.

JERRY L. LONGRIDGE AND WILLIAM L. HILSENHOFF of the De¬

partment of Entomology, University of Wisconsin — Madison, are un¬

doubtedly writing for the professional entomologist but indirectly for

the fisherman and the rest of us, when they contributed this annotated

list of caddisflies — “food for fish and indicators of water quality.” One

is impressed by the precision of their records of time and place of the

catches of each of the 208 species, and by the last simple statement of

their text: “All specimens have been preserved in 70% ethanol and

deposited in the University of Wisconsin Insect Collection.”

J. L. ACKERMAN AND R. D. SHENEFELT. Miss Jacqueline Ackerman

is now Instructor at Lakeland College, Sheboygan, Wi, and Prof. Shene-

felt is a member of the Department of Entomology of the University

of Wisconsin — Madison. With the present day interest in ecosystems

and the scarcity of real knowledge of the food of many lower animals,

this paper assumes greater reader appeal than it would, say 5 or 10

years ago. The intricate pattern of basidiomycetes and their wood sub¬

strate and of the insects and their basidiomycete substrates form an

incredibly complex web. Truly the insects, whether visiting or preda¬

tory or parasitic, can set up a small world of their own within a

single macrofruiting basidiomycete. And we who do not know such

things would call that mushroom wormy!

L. G. MONTHEY AND DANIEL ZIELINSKI, respectively Extension Spe¬

cialist in Recreation Resources Center at the University of Wisconsin —

Madison and Instructor in Geography, University of Wisconsin Center —

Waukesha have dug out the facts on the Oneida County resort situa¬

tion in a very thorough manner. The people of Oneida County and those

trying to improve the economy and suggest better and wiser use of

the beautiful Oneida area will find much of interest in this record.

288 Wisconsin Academy of Sciences , Arts and Letters [Vol. 61

M. STARR NICHOLS. The readers will be interested to know that Dr.

Nichols is Emeritus Professor of Sanitary Chemistry, long retired and

now living in a retirement home (Oakwood Lutheran Home on the

Mineral Point Road, Madison, in case his friends wish to write or

visit him). It is typical of Dr. Nichols to puzzle over hard problems

and to see in them aspects that others neglect. Who is to know who is

right about the major cause of superfluous growth of Algae? At least

Dr. Nichols’ reasoning is sound and perhaps will stimulate some algolo-

gist to bring forth the supporting data. May we all in retirement be

as interested in current problems as is Dr. M. Starr Nichols!

DANA S. DAWES AND N. C. MARAVOLO. Dr. Nicholas Maravolo and stu¬

dent Miss Dana Dawes of the Department of Biology, Lawrence Univer¬

sity, Appleton, Wi, have put to the test of chemistry what in other years

might have remained a chance observation that orange hawkweed had

something to do with maintaing the bracken openings in a pine forest.

Thus this bit of ecological theory is now backed by the comparatively new

technic of chromatography in chemistry. Truly ecology is multi-disci¬

plinary.

LEON M. ZABOROWSKI. With all the talk about mercury in the Wis¬

consin environment, Dr. Zaborowski, Professor of Inorganic Chemistry

at the University of Wisconsin — River Falls, did something about it,

and in the process involved some cooperating students. Chemistry stu¬

dents in at least one high school in every county of the state were

asked to sacrifice their hair, but only a lock of it, to be analyzed for

mercury !

MILAN W. WEHKING, JAMES W. PAVLIK, PAUL STREGE AND

DAWN GILLES. Prof. Wehking, Associate Professor of Chemistry and

his students, Mr. Pavlick and Miss Gilles, at the University of Wisconsin —

River Falls have pointed to another area where the well water may con¬

tain 45 p.p.m. nitrate, presumed to exceed the safe level. This is not

surprising. Wells in other counties show a similar range, when the facts

are known from actual analyses.

JAMES A. JESBERGER, now of the Department of Biology, University

of Saskatchewan, Saskatoon, Sask., Canada, was invited to contribute this

paper as representing Nonvascular Plants in the Wild Rivers Series.

With apology to Dr. Jesberger for the long delay in publication, it is

hereby designated No. 8 in the Series. This too is a paper for the spe¬

cialist but this Editor no doubt can speak for the rest of the Academy

readers in admiring the precise detail of the work.

WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS

OFFICERS

President

Louis W. Busse

University of Wisconsin — •

Madison

309 Pharmacy Building

President Elect

Richard W. E. Perrin

9825 Concordia Ave.

Milwaukee, Wi 53222

V ice-Presid en t — S cien ces

Robert E. Esser

University of Wisconsin — •

Parkside

1001 S. Main Street

Racine, Wi 53403

V ice-President — A rts

Robert E. Gard

University of Wisconsin — •

Madison

719 Lowell Hall

Madison, Wi 53706

1972-73

V ice-President — Letters

Donald Emerson

University of Wisconsin — •

Milwaukee

Garland Hall

Milwaukee, Wi 53201

T reasurer

Edward Schneberger

6818 Forest Glade Court

Middleton, Wi 53562

Secretary

Jean Cronon

5601 Varsity Hill Drive

Madison, Wi 53705

Librarian

Jack C. Clarke

University of Wisconsin — •

Madison

4232 Helen White Hall

APPOINTED OFFICIALS

Executive Director and

Editor — Wisconsin Academy Review

James R. Batt

W.A.S.A.L. Office

1922 University Ave.

Madison, Wi 53705

Editor — Transactions

Elizabeth McCoy

University of Wisconsin — Madison

Department of Bacteriology

Director — Junior Academy

LeRoy Lee

W.A.S.A.L. office or

2215 Wood Road

Middleton, Wi 53562

F. Chandler Young

Norman C. Olson

William B. Sarles

Adolph A. Suppan

John W. Thomson

Walter E. Scott

ACADEMY COUNCIL

PAST PRESIDENTS

Aaron J. Ihde

J. Martin Klotsche

Carl Welty

Henry A. Meyer

Robert J. Dicke

Stephen F. Darling

Joseph G. Baier

Ralph N. BuckstafT

Katherine G. Nelson

Otto L. Kowalke

Henry A. Schuette

includes the above officers, officials and past presidents.

Sou • 13

uj 1\*J 0,3

rRANSACTIONS OF THE

WISCONSIN ACADEMY

OF SCIENCES, ARTS

AND LETTERS

;

LXII— 1974

Editor

ELIZABETH McCOY

TRANSACTIONS OF THE

WISCONSIN ACADEMY

Established 1870

Volume LXII

TOWARD A NEW MATURITY: THE WISCONSIN ACADEMY IN

ITS SECOND CENTURY Presidential Address 1

Louis W. Busse

A HEMLOCK RELICT ALONG LAKE MICHIGAN,

SHEBOYGAN COUNTY, WISCONSIN 11

Thomas Foster Grittinger

THE ROTH CASE: THE BURGER COURT

AND JUDICIAL RESTRAINT 19

Warren R. Wade

WILD SOILS OF THE PINE-POPPLE RIVERS BASIN 37

Francis D. Hole

THE NOVEL AS A VEHICLE TO TELL THE STORY OF

THE MENOMINEE INDIAN 51

William Steuber

LAKE WINGRA, 1837-1973: A CASE HISTORY OF HUMAN IMPACT 57

Paul C. Baumann, James F. Kitchell, John J. Magnuson

and Terrence B. Kayes

A HISTORICAL SKETCH OF THE EVOLUTION OF ENERGETICS

Robert T. Balmer

95

THE ORIGIN OF THE WORD “DOLLAR” — THE NAME

OF OUR UNIT OF ACCOUNT 111

Edward E. Popp

HISTORY OF BIOLOGICAL CONTROL ATTEMPTS AGAINST

INSECTS AND WEEDS IN WISCONSIN 115

J. W. Mertins and H. C. Coppel

NUTRIENT SOURCES FOR LAKE MENDOTA— 1972 133

William C. Sonzogni and G. Fred Lee

AN EVALUATION OF THE USE OF THE EEG TECHNIQUE

TO DETERMINE CHEMICAL CONSTITUENTS

IN HOMESTREAM WATER 165

Jon C. Cooper, G. Fred Lee, and Andrew P. Dizon

STUDIES ON THE Ca, Mg and Sr CONTENT

OF FRESHWATER CLAMSHELLS 173

G. Fred Lee and William Wilson

HYDROLOGY AND TROUT POPULATIONS OF COLD WATER

RIVERS OF MICHIGAN AND WISCONSIN 181

G. E. Hendrickson and R. L. Knutilla

TEMPERATURE OPTIMUM OF ALGAE LIVING IN THE OUTFALL

OF A POWER PLANT ON LAKE MONONA 195

Thomas D. Brock and James Hoffmann

A CORRECTION IN SET THEORY 205

William Dilworth

THE FISCHER COLLECTION 217

James La Malfa

A PRELIMINARY SPATIAL ANALYSIS OF QUALITY OF LIFE

IN MILWAUKEE COUNTY, WISCONSIN 227

Richard A. Karsten, Harold McConnell, and Thomas D. Patterson

PRIMES AND FAREY SEQUENCES 245

Arthur Marshall

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN.

NO. 64. ADOXACEAE— MOSCHATEL FAMILY 247

Theodore S. Cochrane and Peter J. Salamun

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN.

NO. 65. DIPSACACEAE— TEASEL FAMILY 253

Peter J. Salamun and Theodore S. Cochrane

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN.

NO. 66. CYPERACEAE II— SEDGE FAMILY II.

The Genus Cyperus — The Umbrella Sedges 261

Brian G. Marcks

ANTIMYCIN— BEYOND TELEOCIDE 285

Mary Ellen Antonioni

ARBOVIRAL ANTIBODY SURVEY OF WILD MAMMALS

IN SOUTHEASTERN WISCONSIN 303

Omar M. Amin and Wayne H. Thompson

DISTRIBUTION AND ECOLOGICAL OBSERVATIONS OF

WILD MAMMALS IN SOUTHEASTERN WISCONSIN 311

Omar M. Amin

SAMPLING AND ANALYSIS OF BOTTOM SEDIMENTS

OF SOME WISCONSIN LAKES 327

Laverne C. Strieker and Robert N. Cheetham, Jr.

STUDIES ON AQUATIC OLIGOCHAETA IN INLAND

WATERS OF WISCONSIN

Richard P. Howmiller

A HISTORY OF THE VEGETATION OF EAU CLAIRE

COUNTY, WISCONSIN

William J. Barnes

EDITORIAL POLICY

The Transactions of the Wisconsin Academy of Sciences, Arts and Letters

is an annual publication devoted to original, scholarly papers, some preference

being given to the works of Academy members. Sound manuscripts dealing

with features of the State of Wisconsin and its people are especially desirable,

although other papers of merit are also acceptable. Subject matter experts

will review each manuscript submitted.

Contributors are asked to submit two copies of their manuscripts to the

Editor. The manuscripts should be typed on 8V2 x 11" bond paper. The title

should be at the top of the page, typed in capital letters throughout. The

author’s name and brief address should appear below the title and toward the

right hand margin. It should be typed with capital-lower case style and the

address portion underlined for italics. Each page of the manuscript beyond

the first should bear a page number and author’s name for identification;

e.g. 2-Brown, 3-Brown, etc. A note on separate sheet, submitted with the

manuscript, should identify the author with his institution, if appropriate,

or with a personal address for the Editor’s use in correspondence.

The style of the text should be that of scholarly writing in the field of the

author but to minimize printing costs the Editor requests that the general

form of the current volume of Transactions be examined and followed so far

as possible. For the Science papers an ABSTRACT is requested. Documentary

Footnotes may be useful, especially for the Arts and Letters manuscripts, and,

if needed, should be typed as a group on separate sheets and numbered for

citation in the text. Such FOOTNOTES will be published at the end of the

text in place of BIBLIOGRAPHY. For BIBLIOGRAPHY of Science papers

or of any papers at the author’s choice, references should be assembled alpha¬

betically and typed on sheet or sheets as needed; citations in the text are

usually by author and date in parentheses. The style of references will be

standardized as in the current volume, to promote accuracy and reduce print¬

ing costs.

The cost of printing the Transactions is great. Therefore excessively long

papers will not ordinarily be accepted. In the rare instance of such a long

paper or of some especially costly printing or illustrations, the author may

be asked to subsidize publication.

Galley proofs and manuscript copy will be forwarded to the senior author

for proof reading prior to publication; both should be returned to the Editor

within ten days.

Authors of papers which appear in Transactions will be provided by the

Editor with one complimentary copy of the volume in which their work ap¬

pears. Reprints can be made at publication or at later time by offset printing

and are available in multiples of 100. Prices and forms for the orders will be

sent to authors at time of publication or are obtainable from the Academy

office upon request. Single copies only are available to other persons at 10

cents per page for papers or $5.00 for a volume of Transactions from the

W.A.S.A.L. office at 1922 University Ave., Madison, Wis. 53705.

Papers received on or before November 15 will be considered for publication

in the 1975 volume of Transactions. Manuscripts should be sent to the Editor

at the W.A.S.A.L. office above.

LOUIS W. BUSSE

51st President

WISCONSIN ACADEMY OF SCIENCES, ARTS

AND LETTERS

TOWARD A NEW MATURITY: THE WISCONSIN ACADEMY

IN ITS SECOND CENTURY

Presidential Address

Louis W. Busse

President 1972-73

Ripon, Wisconsin

April 27, 197S

President-elect Perrin, Members of the Council, Colleagues of the

Academy, and Friends :

On the First of February, 1870, J. W. Hoyt and 104 of Wiscon¬

sin’s finest scholars, governmental leaders, and noted citizens cir¬

culated about the state a “Call for a Meeting to Organize.” The

Wisconsin Academy of Sciences, Arts and Letters was about to be

born. It was to be, in the words of the authors of the Call, . . an

institution that shall be of great practical utility and lasting honor

to the State.”

For a number of years the idea for such an institution had

been in the hearts and minds of many. As the language of the Call

indicates, the founders of the Academy were motivated by the

premise that “. . . the prosperity and power of a State depend not

more upon its material resources than upon the culture of its

people and the extent of their knowledge of nature and man.” They

incorporated this belief within the charter through the listing of

such purposes as:

1. Researches and investigations in the various departments of the mate¬

rial, metaphysical, ethical, ethnological and social sciences;

2. A progressive and thorough scientific survey of the State, with a

view to determining is mineral, agricultural and other resources;

3. The advancement of the useful arts, through the applications of science,

and by the encouragement of original invention;

4. The encouragement of the fine arts by means of honors and prizes

awarded to artists for original works of superior merit;

5. The formation of scientific, economical and art museums;

6. The encouragement of philological and historical research, the collection

and preservation of historic records, and the formation of a general

library; and

7. The diffusion of knowledge by the publication of original contributions

to science, literature and the arts.

It was, as can be seen, an undertaking cast in a philosophy and

purpose of substantial magnitude. That the Academy was to suc-

1

2 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

ceed in the fulfillment of much of what it had been charged to

accomplish, given its financial limitations, is a high tribute to its

leadership and to its membership. Through the course of nearly

all its first century of existence, the Wisconsin Academy achieved

despite, not because of, its monetary resources. The question must

always remain: What proud paths might we have traveled and

realized what goals had there been a more adequate fiscal base?

I submit to you, however, that a comparable question must be

answered today. While the coffers of the Academy are far from

overflowing, the fact remains that, thanks to the generosity of such

as our late colleague, Dr. Harry Steenbock, our friend and fellow

member, Dr. Elizabeth McCoy, and others, we are now in a posi¬

tion to realize goals dreamed of by our illustrious founders and to

travel some of the paths chartered by these people. We can no

longer plead lack of monetary resources for future failures.

In looking toward a new maturity, there are several ways to

look at the Academy. One of them might be to consider the

Academy as a 100 years old adolescent, who never quite reached

maturity because of a shortage of one essential hormone — that of

fiscal resources (money). Suddenly in 1969 he received a $950,000

injection — and now, as a rapidly growing youth, is trying vigor¬

ously to find himself and to mature.

Or, we can look at ourselves as a 100 year-old institution or

organization, seeking renewal with the vitality of a youngster

stimulated by the faith of several individuals in our goals and

objectives and their generosity to help us achieve or make progress

toward these goals. When we mention the vitality of a society or

institution or organization we must include the vitality of its indi¬

vidual members. The society and its members are one and the same.

A society or an organization decays when its institutions and indi¬

viduals lose their vitality. As I have viewed the Academy over

these past three years, I see members of long standing and new

members demonstrating a new vitality and interest which speaks

well for our future.

You know, one of the fascinating things I have found in the

Academy is its resilience — its constant ability to renew itself. When

organizations and societies are young, they are flexible and fluid,

not yet paralyzed by rigid specialization. As they age, however,

vitality diminishes, flexibility gives way to rigidity, creativity

fades in this environment and finally there results a loss of capacity

to meet challenges from unexpected directions. I view the Academy

as now at the beginning of a new maturing period. This is neces¬

sary for we cannot and should not remain forever with the con¬

fusion of youth. The question to which the Academy will have to

seek the answer is : How can it become a mature organization and

1974] Busse — Second Century of Wisconsin Academy 3

still retain the flexibility and adaptability characteristic of youth

which permit the ever renewing process, when, in fact, the proc¬

esses of maturing are essentially those that reduce flexibility and

adaptiveness.

I do not have the answer, of course, but I would assume that the

first step would be to recognize the facts : to know the difference ,

that is, the difference between maturity and rigidity or lack of

flexibility. So, I would urge our future leaders and especially our

members to constantly be on the alert for the appearance of those

disastrous side effects of the maturing process and to treat these

quickly as they appear. In this kind of an organization, the ever-

renewing process can take place, and then what matures is an

organization or framework within which continuous innovation,

renewal and rebirth can occur. I think this is what we all want for

the Academy and with your help we can develop this kind of

organization.

Now, before an organization can proceed toward its major

objectives and program goals, it must have a certain stability. It

must have its machinery in working order. It must solve its day-

to-day operating problems and get out from under these uncer¬

tainties. It must get out of the confusion of youth.

I thought tonight I might make a portion of my message to you

in the form of a “report from your President.” This is not to set

a precedent for future presidents. However, your Academy has had

three busy years-— two under the leadership of Norman Olson and

F. Chandler Young, and this past year with me and Dick Perrin.

These were good years for me and I greatly appreciated the oppor¬

tunity to serve with all of these people.

I would like to indicate some of the significant events or hap¬

penings over this past year for I think they reflect stages in the

maturing process that the Academy is now going through, and

they give proof to the claim that we are getting our house in order

and going on to greater accomplishments.

THE FIRST EVENT

Of great significance to the Academy was our success in clarify¬

ing our tax status with the Internal Revenue Service. As some of

you know, due to errors in previous reporting to the Revenue Serv¬

ice we were classified as a private foundation and our income from

our investments, thus, would be taxable. This classification was an

intolerable situation for the Academy. Therefore, competent legal

counsel was retained and the Academy’s income, operating

expenses, and contributions were properly identified and estab¬

lished. A brief was prepared and submitted to the I.R.S. and just

last month we were notified of our new classification as a tax-

4 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

exempt foundation. This had to be established in order for the

Academy to continue to develop to its full potential. It pleases me

to be able to report our success in this effort. The amount of staff

time which went into this effort was enormous and how our Direc¬

tor was able to keep anything else going is testimony to his effort.

In this relation also I must acknowledge the help of Walter Scott

and express my appreciation for the tremendous help he was in

leading us and supplying us with information on the Academy.

In order to protect the Academy from any further mistakes in

this area and because of the increasing activities of the Academy

which may have legal implications needing interpretations, I have

recommended to the Council (and they have approved) the con¬

tracting of legal services on a retainer basis. We have secured the

services of the law firm of Werner, Lathrop, Heany, which handled

our tax case.

One of the other events which gave me real pleasure was to

observe and work with Bob Dicke and our Facilities Committee.

Bob was persistent and insistent that a basement office in a lumber

yard was no place for a distinguished organization such as ours.

Bob believed in the “shock technique.” He first came to the Council,

as many of you know from our meeting last fall, and presented a

plan which involved an expenditure of some $350,000. First, he got

all of us to agree that our location at that time just was

not acceptable and then in turn he agreed that maybe $350,000

was a little high and that he and his Committee would look again.

So, when he came back with a request to the Academy to purchase

a $70,000 building, which was on University Avenue and just off

campus, we were relieved to get off that easily.

I am especially enthused with our new headquarters building

at 1922 University Avenue, which the Academy has purchased and

is remodeling for its needs. It was purchased with capital gains

from our investment trust for $67,500. Our distinguished member,

Dr. Elizabeth McCoy, suffered through this agonizing period with

us, and very early in our deliberations offered us a location on her

farm or estate along with a contribution of $22,500 to remodel a

building there. This we considered very seriously but hesitated

only because of the distance from the campus and central city.

However, when she saw Bob’s and my pleasure at the opportunity

of the University Avenue site she couldn’t resist and said, “This

is what you need; it’s the right location; and it is fitting for the

Academy.” After another two seconds she said, “What’s more, I’ll

round it off to $25,000 toward the purchase or remodeling, which¬

ever you wish.” Ladies and Gentlemen, that’s why you have a home,

a headquarters building, an image if you will, and one in which

you may feel happy and comfortable for many years.

1974] Busse — Second Century of Wisconsin Academy 5

It’s a charming* little building*, a delightful place for the staff to

work in, and an extremely pleasant place to visit. I know you will

be pleased and I urge you to come and see it and experience the

revitalization, the renewed effort that this facility has stimulated

in all of us. We will be moving* in on Tuesday, May 1 and although

it will not be completely furnished, we might be able to find you

a chair to sit down on.

The acquisition of the building and the contracting for remodel¬

ing and redecorating services has taken a tremendous number of

hours of staff time this past month, so if some of the daily routine

or deadline schedules were not met, please bear with us for another

month or so.

Another event, with rather severe implications for the Academy,

was the establishment by the Governor of a task force on the status

of the arts and humanities in Wisconsin with Dean Adolph Suppan,

a past president of the Academy, as its Chairman. The task force

did hold several meetings in the state and in Madison, but for some

reason neither I nor our staff were made aware of the work of

this committee until just a few months ago. The recommendations

of the Committee were already formulated and the Governor

already had in his budget the funds for the establishment of a State

Board for the Arts and Humanities. This board is to disperse the

funds contributed to the arts in Wisconsin by the National Endow¬

ment for the Arts and thus will relieve the Arts Council of the

state of this responsibility.

As many of you know, this situation resulted in the resignation

of Mr. Lauch as Executive Director of the Arts Council and the

publishing in the Milwaukee Journal of an interview with him in

which he denounced the work of the task force and took serious

exception to the creation of this new board or state agency.

Even though it was late to intervene, I felt that the implications

for the Academy were so great that appearance was necessary in

opposition to the bill at the hearing before the Joint Finance Com¬

mittee. Mr. Batt and I appeared there and spoke in opposition and

presented the Committee with much material relating* to the

Academy’s activities in the Arts and Humanities. Needless to say,

the Committee was very surprised there was such an organization

closely connected to the State by charter, already performing these

functions, and seemed pleased to be made aware of this alternative.

I can inform you that the staff of this committee considered three

alternatives for the responsibility:

1. To leave in the hands of the Arts Council;

2. To place in the hands of the Academy of Sciences-Arts-Letters ;

3. To follow the Governor’s recommendation for a new State Board.

6 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Mr. Davis, of the Committee, informed us that the Committee is

recommending going along with the Governor’s proposal.

In spite of this, the Academy will continue to maintain a vigor¬

ous program for the Arts and Letters, and will work with the

Board, if and when one is created.

Two other areas in which progress has been made in the past

two years has been in membership and publications — the vitality

and life blood of our organization.

We have increased our membership by approximately 300 and

quadrupled our life memberships. The new membership directory

has been published and it makes an impressive appearance. Mem¬

bership drives will continue and in the meantime all of us as indi¬

viduals can help tremendously by each bringing in a new member

next year. Doubling our membership next year would be an

encouraging step.

I am particularly pleased with the improvement of our publica¬

tions this past year and I hope that you are too. The TRANSAC¬

TIONS has always been an exemplary publication. Now, the

ACADEMY REVIEW has been expanded and takes on a new look,

and I think does the Academy proud. The last two issues were

particularly good and carried a good balance between our three

cultures.

In addition, you now have received an issue or two of the

ACADEMY TRIFORIUM, our new monthly newsletter, designed

to keep you informed of Academy developments in a more up-to-

date manner. I think it is interesting and will help to keep the

Academy in the forefront of our minds and our interest.

Publications, to my mind, are the life blood of the Academy and

will determine its future. I think it is essential that the Academy

have an extensive publishing program for it is only through this

mechanism that we can really create and maintain the image and

the status we seek. Our publications will determine whether the

mass of the membership continues to remain in the Academy or

to withdraw for lack of contact with our programs and objectives.

This is costing the Academy money ; however, early results seem

to indicate it will also bring in money. Nevertheless, the budget for

1973-74 will show a significant increase in the item for publica¬

tions. In addition, this cannot be accomplished without increased

manpower and so our budget will show that we have acquired the

full time services of Miss Monica Janigh. One could say that we are

doing it with womanpower instead of manpower.

Seriously, I think our staff, Jim, Nancy, Marie and Monica, are

doing an excellent job in their areas of Academy responsibility and

we owe them a vote of thanks for their dedication.

1974] Busse — Second Century of Wisconsin Academy 7

These accomplishments, I like to think, are in the category of

getting our house in order :

1. Our status as a tax-exempt organization

2. A house befitting our image with adequate space and facilities to do

our job

3. Publications reflecting our progress toward our goals and objectives.

4. Membership buildup

Significant progress has been made in all of these this year.

We are now just in the process of assessing our future role and

goals, hopefully exhibiting the vitality, flexibility, and adaptabil¬

ity necessary to assure our success. There are two things I would

like to mention in this regard.

First is the reorganization of our Junior Academy, which has

been growing rapidly under the direction of Mr. LeRoy, Lee. More

high schools are participating, more students are participating,

more institutes for both school and summer months, are being

organized. I think over 600 students have been at the various

institutes this past year.

In our reorganization, we will be dropping the Junior Academy

nomenclature and thus not have a separate organization. Instead,

we will have a youth program in which these students are associate

members of the Academy which will make them feel more a part

of us-— an example of our adaptability and flexibility. I look for

this aspect of our activity to mushroom over the next years and

I am sure that it will require a full-time director for this program

in a few years. As an example, the Academy, through the efforts

of LeRoy and Jim, is the recipient of a grant of $7,500 from the

Atomic Energy Commission to sponsor institutes on our energy

crisis at the high school level. There is no end to the kinds of pro¬

gramming the Academy can sponsor and develop for these young

people. However, as we involve ourselves more with the young

crowd, or to put it the other way, as we take the young people more

into our establishment, we must be prepared to accept change and

to have the framework which will accept and permit constructive

changes.

Second, I would like to mention some of our efforts to decide our

future roles and objectives. In this respect I would like to express

my appreciation to Bob Gard and Don Emerson for their contribu¬

tions. In the role of stimulators , particularly in the Arts and Let¬

ters, the Academy is embarking on the following pathways:

1. The Council has authorized an Academy literary award of $5,000 for

the best book published by a Wisconsin author or author with Wisconsin

heritage, to be given in 1976 as a part of the Bicentennial celebration.

2. We are giving citations in “The Theatre” category and Professor Gard

informs us he has already had the pleasure of presenting one award on

8 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

behalf of the Academy before an audience of 600-700 people in Stevens

Point, who were thrilled at the recognition bestowed upon them by our

organization.

3. We are also establishing a Citation for the best book in Science pub¬

lished by a Wisconsin author and it will be presented by and for the

Wisconsin Library Association.

4. Our Fall Gathering probably will be held in Ft. Atkinson this Sep¬

tember 22 and 23 in conjunction with the Wisconsin Regional Writers

Association and the dedication of the Wisconsin Authors Hall of Fame.

I mention all of these items to you because they indicate to me

two things : ( 1 ) the Academy is actively seeking to find its role

in its relationship to the specialty organizations making up the

Arts and Letters; and (2) is attempting to bring them into our

Academy home and to nourish them as individuals in any way we

can. I think this is marvelous progress, and much credit should go

to our Vice Presidents and to our Executive Director.

In this same vein, Dick Perrin will be appointing a committee to

study the feasibility and mechanism by which the Academy could

enter into a cooperative arrangement with the U. W. Extension

Division in support of a faculty member who would work at the

Academy offices. This person would study the role of the Academy

as a co-ordinator for all of the Arts and Letters organizations in

the state and develop a procedure by which the activities of these

organizations could come under the umbrella of the Academy. In

addition, he will determine how the Academy can help these organ¬

izations to fulfill their goals and objectives. If this were to come

about and prove to be successful, I can visualize a similar role in

the Sciences and with this kind of work power the Academy should

be able to fulfill the goals and objectives so forcefully stated in our

Charter of 1870. I do look for this to happen!

In closing, let me say these have been three wonderful years.

I hope I may continue to serve the Academy in the future. As you

know, Wisconsin is one of the few Academies of Science which

encompasses the Arts and Letters as well. As a pharmacist and

scientist one of the real pleasures of this opportunity to serve you

was the chance it gave me to meet and know so many people in the

other cultures, and to know and understand the working of the

mind of the artist and the writer. I hope similarly it gave the artist

and the writer a closer view of the thinking and the orientation

of the minds of the scientists and professionals. I no longer think

of the artist and writer in an adversary relationship and I hope

they too can begin to see professionals and scientists in non¬

adversary relationships.

I think it is extremely important to stable progress in our

Academy that these adversary relationships between these cultures

be reduced and that this be replaced by mutual understanding. It

1974] Busse — Second Century of Wisconsin Academy 9

seems to me that this is a particularly significant role for the

Academy to perform, and since it is an organization of all three

cultures it is ideally suited to perform it. Members of the Academy,

if the Academy can accomplish this for me, I know it can accom¬

plish it on a much greater scale for the people of our state. I urge

you to make this a major effort in the next years.

It has been an honor and a pleasure to serve you.

And now it is my privilege and pleasure to turn the reins of the

Academy over to your President-elect, Richard Perrin, who now

becomes your President for the coming year. I hope I give the

Academy to you in as good a state as it was when it was turned

over to me by President Young. I know the Academy will be in

good hands. Members, your new President, Mr. Richard Perrin.

A HEMLOCK RELICT ALONG LAKE MICHIGAN,

SHEBOYGAN COUNTY, WISCONSIN

Thomas Foster Grittinger

University of Wisconsin — Center

Sheboygan County — Campus —

Sheboygan

Hemlock ( Tsuga canadensis (L). Carr.)* had been reported

along Lake Michigan as far south as Sheboygan County by sur¬

veyors (Goder, 1955). The southernmost stands mapped along the

lake are listed as relicts in Manitowoc County; however a few

scattered hemlocks can still be found within 50 feet of the shore¬

line in Sheboygan County (Goder, 1955). This report is an analysis

of a well established stand composed mostly of hemlock, found east

of Oostburg in Sheboygan County (Fig. 1). Despite the southern

location of this northern community and its disjunct nature, it is

rich in species and the hemlock is vigorous.

This hemlock stand lies on old dunes between the more recent

dunes to the east and the swampy swales covered with lowland

hardwood to the west. Farther west a steep bluff of Nippissing

age separates the agricultural uplands from the forested lowlands

along Lake Michigan. The bluff appears to provide a wind shadow

from the prevailing dry westerly winds. East of the hemlock the

younger dunes support white pine (Finns strobus L.) , white cedar

( Thuja occidentalis L.) , white birch ( Betula papyrifera Marsh.),

and green ash ( Fraxinus pennsylvanica Marsh, var. subinteger-

rima (Vahl) Fern.). This forest buffers the hemlock community

from the easterly and northeasterly winds off Lake Michigan. The

hemlock stand thus receives protection from both the prevailing

summer winds and the winter storms.

The proximity to Lake Michigan has a major impact on the

climate. Lake effects such as a cooling in the spring and early

summer when onshore breezes are common, a high frequency of

local fogs, postponement of the first killing frost in the fall, delay

of bud opening in the spring, as well as persistence of snow cover

near the shore in the winter when the lake acts as a cloud gener¬

ator and a heat and moisture source are well known (U.S. Dept.

Comm., 1961 ; Bruncken, 1910 ; Whitford and Salamun, 1954 ; and

Lyons, 1970). These climatic effects account for the southward

extension of northern forests along the lake. Northern species have

* The nomenclature used here is according- to Fernald (1950).

11

12 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

FIGURE l

been recorded as far south as Grant Park in Milwaukee County

(Bruncken, 1910; and Whitford and Salamun, 1954).

SAMPLING METHODS

The quarter method was used to examine tree composition and

structure in the stand. The trees reported include stems over 4

inches in diameter at breast height (d.b.h.). Quadrats 4x4 meter

square were used for shrubs and saplings, and 1x1 meter quad¬

rats were used for herbs and seedlings. Saplings included stems

with diameters of 1 to 4 inches, seedlings with diameters below

1 inch, and shrubs, multistemmed woody species that generally

do not reach tree size. Quarter points and the quadrats were

placed at 50 foot intervals along compass lines. To cover most of

the stand, three compass lines were used with a total sample of

50 quarter points and quadrats of each size. Relative frequency,

relative density, and relative dominance were calculated for tree

species and these values were summed to give an importance value

(Cottam and Curtis, 1956). Quadrat data provided relative fre¬

quency and frequency.

RESULTS AND DISCUSSION

The community includes many species typical of the northern

coniferous forests and even the boreal forest (Curtis, 1959) ; see

1974] Grittinger — Hemlock Relict Along Lake Michigan

IB

Tables 1, 2, and 3. For example, hemlock, beech ( Fagus grandif olia

Ehrh.) , red-berried elder (Sambucus pubens Michx.) , Solomon’s

Seal ( Polygonatum pubescens (Willd.) Pursh), and club-moss

(Lycopodium lucidulum Michx.), all species that achieve their

highest presence values in the northern mesic forest, are common

throughout this community. White pine, red maple (Acer

rubrum L.), white birch, and wild sarsaparilla (Aralia nudi -

caulis L.) , characteristic of the northern dry-mesic forests, are

present; white pine is especially common on the dune ridges. The

northern wet-mesic forests contain yellow birch (Betula lutea

Michx.), black ash (Fraximus nigra Marsh.), and white cedar.

Yellow birch is found throughout the stand; black ash and white

cedar are frequent in the low, moist depressions between dune

ridges. Species of the boreal forest, mountain maple (Acer spi-

catum Lam.), large-leaved aster (Aster macrophyllus L.) , blue-

bead-lily (Clintonia borealis (Ait.) Raf.), wild lily-of-the-valley

(Maianthemum canadense Desf.) , star-flower ( Trientalis borealis

Raf.), fly-honeysuckle (Lonicera canadense Bartr.), and twisted-

stalk (Streptopus roseus Michx.) are also present in this stand.

Viola Selkirkii Pursh., which occurs in this community, is noted

by Fassett (1959) as present along the northern borders of Wis¬

consin, “south to Clark and Door Counties, and in cold canyons

at Devil’s Lake and the Dells of the Wisconsin River”. Many other

species found here with low frequencies also exhibit their highest

presence values in northern Wisconsin. Northern species seen but

not encountered in the quadrats include goldthread (Coptis groem

landica (Oeder) Fern.) and bunchberry (Cornus canadensis L.).

Northern plant communities are reasonably frequent near Lake

TABLE 1. PHYTOSOCIOLOGICAL DATA ON TREE SPECIES1

Average distance between individuals = 13.6 feet Trees/acre - 235.6

Quarter method, 50 points used

2 Importance value = relative frequency + relative density + relative dominance

14 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 2. FREQUENCY VALUES FOR SHRUBS AND SAPLINGS

TABLE 3. FREQUENCY VALUES FOR SEEDLINGS AND HERBS

Michigan in Sheboygan County. This forested strip consists largely

of white pine and hardwoods on mesic sites or black ash and white

cedar on lower areas. Hemlock is infrequent and appears to be a

disjunct species.

In this community hemlock represents by far the most important

tree species, with an importance value of 138.2 (Table 1). Other

1974] Gritting er- — Hemlock Relict Along Lake Michigan 15

trees found include in descending order of importance : red maple,

yellow birch, white birch, sugar maple (Acer saccharum Marsh.),

beech, white pine, red oak ( Quercus rubra L.), basswood ( Tilia

americana L.), green ash, white cedar, black ash, and mountain

ash ( Pyrus americana (Marsh.) DC.). This is in contrast to the

stands examined by Stearns (1951) where sugar maple, with a

relative frequency of 89%, a relative density of 41.8%, and a rela¬

tive dominance of 28%, had the greatest importance value; hem¬

lock followed the sugar maple with a relative frequency of 67.6%,

a relative density of 22.3%, and a relative dominance of 23.8%.

Numerous large trees gave hemlock high relative dominance. Old

hemlock stumps in the stand suggest the possible ages, of these

large trees. Three stumps ranged in age from 118 years for a 24

inch diameter stump to 184 years for a 29 inch stump. However

estimation of hemlock from stem diameters is subject to error,

largely because hemlock tolerates suppression (Curtis, 1959).

Stearns (1951) found that hemlock may experience several periods

of suppression before finally reaching a position of dominance in

the canopy. In this community, hemlock attains a greater size than

other species save for white pine. The white pine also has a high

relative dominance but is not present as seedlings, or saplings,

although it is present in various sizes as trees. Although no hem¬

lock seedlings were encountered in the quadrats (Table 3), they

were often seen on old stumps (Fig. 2).

Hemlock shows a high relative density, compared to relative

frequency. Favorable sites for seed germination are rotting stumps

and logs, and tip-up mounds; all of these microhabitats are com¬

mon in this community. The stumps resulted from the infrequent

removal of a few trees. Relatively shallow root systems, and loose

sandy soil account for the high percentage of windthrown hemlock.

The importance of windthrow in creating sites for germination

and in opening the canopy for hemlock has been documented

('Coder, 1955). Reproduction on stumps is seen in this community

(Fig. 3). Thus hemlock creates its own environment for reproduc¬

tion (Coder, 1961). Since the microhabitats so created tend to

occur in specific limited situations, clumping of trees inevitably re¬

sults. The presence of dense island-like groupings of hemlock was

noted by Stearns (1951).

Hemlock is considered preferred deer food (Swift, 1946) ; how¬

ever, the deer population is low in this area perhaps because many

of the lakeshore cottages and nearby homes are occupied through¬

out the year. Hemlock, if browsed, has little chance for recovery

(Curtis, 1959). Elsewhere along Lake Michigan, deer populations

16 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

FIGURE 2

have grown rapidly and browse lines are visible in places within

Milwaukee County (Forest Stearns, personal communication).

In addition to hemlock, other trees appear capable of continuing

in the community under present conditions. Both red maple, and

especially sugar maple, in all size classes are present throughout.

Yellow birch was found in many sizes, with many exhibiting the

prop rooted condition mentioned by Curtis (1959) ; this condition

is the result of seedling development on rotting logs and stumps

with the subsequent decomposition of the log or stump. In spite

of this unique method of early development, yellow birch reproduc¬

tion is very successful on moist, mineral soil surfaces (Curtis,

1959), and heavy windfall or partial cutting will permit an in¬

crease in yellow birch reproduction (Stearns, 1951). White birch

apparently owes its presence to localized disturbances, since it is a

gap-phase tree found in small openings in the forest (Curtis,

1959). Beech can be expected to continue reproducing in this com¬

munity; this species maintains itself by vegetative reproduction

(Ward, 1956).

The origin of this community is tied to long term lake level

changes, to the development of sand dunes and their stabilization,

and to continuing disturbances. After the moving dunes are colo¬

nized and stabilized by vegetation, an orderly process of plant

succession takes place, which eventually terminates in a mesic

1974] Gritting er — Hemlock Relict Along Lake Michigan

17

FIGURE 3

forest (Cowles, 1899; Olson, 1958a and b; and Curtis, 1959).

However since 800 to 1000 years are required to progress from

initial stand to a climax mesic forest, such uninterrupted succes¬

sions are very rare (Curtis, 1959). Disturbance is indicated by

white pine, white birch, and red oak. The presence of white pine

as scattered individuals of uneven age in a mixture of hemlock and

climax hardwoods is due to relatively small openings caused by

windfall or some other purely local influence (Nichols, 1935). The

presence of continuing disturbance in this community is further

demonstrated by hemlock and yellow birch which require wind-

produced clearings in order to compete with sugar maple in a

mixed stand of sugar maple, hemlock, and yellow birch (Stearns,

1949). Stearns (1951) further suggests that where hemlock exists

18 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

in clans or dense groups, it is probably due to mass establishment

following some occurrence which not only opened up the forest

canopy but which also coincided with favorable seed years and

favorable climatic conditions. The preference of pure stands of

hemlock for very moist situations may be related to the increased

frequency of heavy windfall in shallow rooted trees and with the

moisture requirements for seed germination (Stearns, 1951).

ACKNOWLEDGEMENTS

The author is indebted to Dr. F. W. Stearns for advice and

encouragement.

REFERENCES

BRUNCKEN, E. 1910. Studies in plant distribution. 9. The shore of Lake

Michigan. Bull. Wis. Nat. Hist. Soc. 8:145-157.

COTTAM, G. and J. T. CURTIS. 1956. The use of distance measures in phyto-

sociological sampling. Ecology 37:451-460.

COWLES, H. C. 1899. The ecological relations of the vegetation on the sand

dunes of Lake Michigan. Bot. Gaz. 27:95-117, 167-202, 281-308, 361-391.

CURTIS, J. T. 1959. The vegetation of Wisconsin. The University of Wis¬

consin Press. Madison. 65/ pp.

FASSETT, N. C. 1959. Spring flora of Wisconsin, 3rd ed. The University

of Wisconsin Press. Madison. 189 pp.

FERNALD, M. L. 1950. Gray’s manual of botany, 8th ed. American Book

Co. New York. 1632 pp.

GODER, H. A. 1955. A phytosociological study of Tsuga canadensis near

the termination of its range in Wisconsin. Ph.D. thesis, University of

Wisconsin.

GODER, H. A. 1961. Hemlock reproduction and survival on its border in

Wisconsin. Trans. Wis. Acad. Sci. Arts Lett. 50:175-182.

LYONS, W. A. 1970. Numerical simulation of Great Lakes summertime con¬

duction inversions. Proc. 13th Conf. Great Lakes Res. 1970. Internat.

Assoc. Great Lakes Res. 369-387.

NICHOLS, G. E. 1935. The hemlock-white pine-northern hardwood region of

eastern North America. Ecology 16:403-422.

OLSON, J. S. 1958a. Lake Michigan dune development. 2. Plants as agents

and tools in geomorphology. Jour. Geol. 66:345-351.

OLSON, J. S. 1958b. Lake Michigan dune development. 3. Lake-level, beach,

and dune oscillations. Jour. Geol. 66:473-483.

STEARNS, F. W. 1949. Ninety years change in a northern hardwood forest

in Wisconsin. Ecology 30:350-358.

STEARNS, F. W. 1951. The composition of the sugar maple-hemlock -yellow

birch association in northern Wisconsin. Ecology 32:245-265.

SWIFT, E. 1946. A history of Wisconsin deer. Wisconsin Conservation De¬

partment Pub. 323. Madison. 96 pp.

U.S. Department of Commerce, Weather Bureau. 1961. Wisconsin Clima¬

tological Data.

WARD, R. T. 1956. The beech forests of Wisconsin — changes in forest com¬

position and the nature of the beech border. Ecology 37:407-419.

WHITFORD, P. B. and P. J. SALAMUN. 1954. An upland forest survey of

the Milwaukee area. Ecology 35:533-540.

THE ROTH CASE: THE BURGER COURT AND

JUDICIAL RESTRAINT

Warren R. Wade

Univei'sity Wisconsin —

Stout Menomonie

When the United States Supreme Court announced its decision

in Roth v. The Board of Regents of the State Colleges1 on June

29, 1972, it made a ruling of landmark significance in the law

governing academic personnel matters. The Court held that non-

tenured faculty members have no constitutional right to a statement

of reasons or to a hearing in cases regarding non-retention.

The Roth case was surrounded by a great deal of notoriety and

publicity which caused many individuals and groups, in addition

to the plaintiff, Dr. David Roth, an Assistant Professor at Oshkosh

State University, to become interested in its outcome. Several

other non-tenured faculty members whose contracts had not been

renewed by the Wisconsin State University System were anxiously

awaiting the Court ruling. At the same time, college presidents,

vice-presidents, deans, and department chairmen throughout Wis¬

consin quivered at the mere mention of the possibility that the

lower federal courts’ decisions (which maintained that non-

tenured faculty were constitutionally entitled to a statement of

reasons and a hearing in a non-retention case) would be sustained.

Although most college administrators in the rest of the country

probably had not heard of the Roth case, several state and national

associations, representing their interests, had filed amici curiae

briefs on behalf of the defendant-appellant Wisconsin State Uni¬

versity Board of Regents, arguing that detrimental results would

occur if the lower court decisions were not reversed.2

The Roth case is unique because it marked the first time that

the Court had addressed itself to the issue of what procedural

rights the Constitution guarantees a non-tenured teacher. This

paper approaches Roth as a case study attempting to identify the

kinds of factors and events capable of generating a legal contro¬

versy of sufficient magnitude to cause the nation’s highest judicial

1 408 U.S. 654 (1972).

3 Briefs were filed by the following- : the Board of Governors of State Colleges and

Universities of Illinois ; the Board of Regents of Regency Universities of Illinois, the

Board of Trustees of Southern Illinois University ; the American Association of State

Colleges and Universities ; and the American Council on Education and the Associa¬

tion of American Colleges ; by Attorneys Albert E. Jenner, Chester T. Kamin, and

Richard T. Franch, 135 South La Salle Street, Illinois 60603.

19

20 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

tribunal to deem it worthy of a hearing. The development of the

controversy is traced from its origin in late November, 1968 to its

final adjudication on June 29, 1972 and includes a description of

the original plaintiff, Roth, the charges made against him, and

an analysis of the three federal court opinions in the case.

EVENTS LEADING TO THE NON-RETENTION DECISION

The events leading to the non-retention of Dr. David Roth in

the Political Science Department at Oshkosh State University, Osh¬

kosh, Wisconsin (now known as the University of Wisconsin —

Oshkosh), were extremely complicated, as evidenced by the con¬

flicting interpretations of Roth’s actions as well as the events

surrounding the case. Although several of the charges against Roth

can be accurately documented, many diverse opinions were offered

as to their verity or falsity. Out of the numerous interviews with

many of the parties involved, including Roth himself, and through

extensive examination of reports, legal briefs, newspaper articles,

and federal court opinions, this writer has attempted to make a fair

judgment as to why Roth’s contract was not renewed for the 1969-

1970 academic year in addition to the important legal and consti¬

tutional implications for state control of educational disciplinary

matters.

The events leading to Roth’s non-retention initially arose out of

the suspension of ninety-four Oshkosh black students after a con¬

frontation involving the occupation of President Roger Guiles’

office on November 21, 1968. The black students were protesting

against President Guiles’ alleged insensitivity to their demands for

more black instructors, additional black studies courses, a black

culture center, and better counseling services for black students.

President Guiles, arguing that it was administratively and finan¬

cially impossible for him to do so, refused to agree to the demands

whereupon one student yelled, “Do your thing.” With this, bedlam

broke loose. His desk and files were overturned and their contents

scattered around the room. Typewriters were thrown on the floor,

tables were overturned, draperies were torn, and a calculator was

thrown out the window, causing damage in excess of $7,000. All

ninety-four students were arrested and immediately suspended

from classes.

Although Roth had not been involved with black students on the

Oshkosh campus prior to the disturbance in the president’s office,

in the two or three weeks following it he became the center of

attention. One Oshkosh faculty member maintained that Roth acted

more like a student leader than a faculty member during these

1974]

Wade — The Roth Case

21

weeks. Indeed, Roth, by his own admission, said that he gave ad¬

vice and support to numerous student groups.

On December 2, 1968, a meeting was scheduled between the

Oshkosh administration and the parents of the suspended black

students. Roth admitted dismissing his class in International Rela¬

tions which was scheduled at the same time, and he encouraged his

students to join him at the meeting. Three days later Dr. Raymond

Ramsden, Vice-President of Academic Affairs, sent Roth a note

indicating that he wanted to discuss this action with him. Roth

refused to respond to the Vice-President’s request.

Between the end of Thanksgiving recess and Christmas vacation,

Roth was accused by the Oshkosh Administration of not teaching

material pertinent to his International Relations course. Several

students claimed in depositions that fifty to seventy-five per cent

of the class time between these periods was devoted to discussing

the black student riots and related matters. Roth freely admitted

this, arguing that the Oshkosh events constituted a “microcosm of

international conflict.” Arthur Darken, Dean of Letters and Science

and an international relations scholar, admitted this was a good

analogy. But he maintained that Roth greatly overused what should

have been an illustrative point in one lecture.

On Friday, December 20, 1968, the Wisconsin State University

Board of Regents convened in special session to make a final de¬

cision on the academic status of the ninety-four suspended students

in response to a court order issued by United States District Judge

James Doyle.3 Roth led a group of students and faculty in present¬

ing a petition to the Board asking for the reinstatement of the

ninety-four students. When the petition was being presented to the

Board an incident allegedly occurred that compounded David Roth’s

problems at Oshkosh. He was accused of saying, “Here it is, you

racist pigs.” No administrator or faculty member interviewed at

Oshkosh admitted to having heard Roth make the statement. Roth

himself denied it. However, two members of the Board of Regents

maintained this is what he said. Whether or not he actually made

the statement is therefore unclear. As one high administrative

official at Oshkosh stated, “I have heard from reliable sources that

he did say it, and I have heard from equally reliable sources that

he didn’t say it.”

At any rate, no mention of this particular allegation was made

in the official memorandum from Dean Darken to President Guiles

and Vice-President Ramsden which recommended that Roth not

be retained.4 Instead, Dean Darken emphasized the fact that Roth

3 See Marzette v. McPhee, 294 F. Supp. 562 (1968).

‘Dated January 28, 1969, and reprinted as Exhibit C in the Appendix to the appel¬

lant State of Wisconsin’s Brief at pp. 120-126.

22

Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

was scheduled for three classes on Friday, December 20, during

which time he was in attendance at the Board meeting. He stated :

... if this is to be overlooked or simply ‘winked at’ because Dr. Roth was

casting himself in the role of a ‘liberal spokesman’ the door is opened to

all professors to cut their classes whenever they believe one of their

personal or social objectives would be served in this way. Faculty are

engaged to teach and it is simply not acceptable that they fail to meet

classes for such reasons as prevailed here.5

Roth admitted that he did not teach his classes on the day that

the Regents were meeting in Oshkosh. However, he maintained he

was absent on December 20 due to illness. He went instead to his

office and the Board meeting because he thought that these de¬

mands on him would be less than in the classroom. The administra¬

tion, needless to say, viewed this as a gross breach of professional

ethics and responsibility.

Dean Darken was also disturbed by several public statements

Roth made to the press. Darken said they “indicate a very unschol-

larly approach to the truth and the search for knowledge that make

it doubtful he has the qualities of scholarship desirable in a faculty

member.” The Dean also indicated that Roth said: “Many of us

feel that the authoritarian and autocratic structure of this univer¬

sity is no longer tolerable,” and “The state universities will not be

able to keep good professors if they are told they can’t teach this

or that in their classes.” Darken argued that Roth never made any

attempt to specify what was authoritarian and autocratic nor did

he mention any instances where any faculty were told “what to

teach or what not to teach.”6

Furthermore, according to Dean Darken, Roth advised students

not to go to an informational meeting to learn of the progress being

made on the demands of the black students. He was reported to

have said, “We won’t talk to any Mickey Mouse committee.”

Darken concluded :

If a college professor cannot be expected to encourage a rational approach

to a problem of our day, who can be expected to do it? . . . What is

involved here is not the freedom of the faculty member to explore com¬

peting views in search of the truth in his class or research but the use

of unsubstantiated allegations of a grave nature in a tense and possibly

inflammatory campus situation. This behavior is not consistent with what

the University has a right to expect from a faculty member in the way

of scholarship.7

Finally, Roth did not give a two-hour written final examination

in his International Relations course for freshmen, which is

required at Oshkosh, despite an official directive from the Vice-

5 Ibid., p. 121.

0 Dean Darken’s memorandum, pp. 122-123.

7 Ibid.

1974]

Wade — The Roth Case

23

President for Academic Affairs,8 Instead, Roth used a group proj¬

ect which had been done earlier in the semester in lieu of a final.

Dean Darken’s memorandum stated:

The final examination, especially in lower level and freshman courses

serves an important purpose and it is not tolerable for a first year faculty

member in one of his first teaching assignments to cavalierly decide he

won’t give a final examination to students and breach a major all-univer¬

sity policy that is in the interest of the students.9

PROCEDURES LEADING TO THE NON-RETENTION

DECISION

The procedures leading to the non-retention of Dr. David Roth

began on Dcember 17, 1968 when the Political Science Tenure

Committee (composed of the department chairman, who was not

tenured, and the four tenured members of the department) voted

unanimously to retain all of the non-tenured faculty members,

including Roth. Apparently, there were some reservations about

retaining Roth, but the general feeling was that there wasn’t suffi¬

cient evidence of misconduct to warrant non-retention at the time

of the December 17 meeting which, of course, was prior to the

December 20 incident at the Board of Regents meeting.

Prompted by the December 20 incident, however, as well as by

some additional public statements made by Roth early in January,

and also by his failure to give a final examination, Dean Darken

asked the four tenured members of the department to meet in his

office on Friday, January 24, 1969. At this time the Dean informed

them of his recommendation not to retain Roth. He then told them,

if they chose to do so, they had until 4:00 p.m. Monday, January

27, to reconsider their recommendation in light of Roth’s actions

since the December 17 meeting. On Monday the committee cast one

vote for retention and two against it, with two abstaining. Roth

was informed of his non-retention for the 1969-1970 academic year

both by telephone and by registered letter on January 30.

DAVID ROTH IN THE FEDERAL COURTS

On February 14, 1969, through attorneys provided by the Wis¬

consin Chapter of the American Civil Liberties Union, David Roth

filed suit in the Federal District Court for Western Wisconsin,

Judge James Doyle presiding. In the complaint it was alleged that

Roth’s contract had not been renewed merely because he had exer¬

cised his constitutional rights of freedom of speech on matters of

8 Vice-President Ramsden wrote, in a memorandum to all faculty, dated December

17, 1968 : “Every class, with exceptions listed below, will end with a 2-hour final writ¬

ten examination to be given during- the time arranged for it in the schedule. If you

are of the opinion that a two-hour final written examination is inappropriate for a

course not listed, please make this known to me by 4 :00 p.m. Friday, December 20.’’

0 Dean Darken’s memorandum, p. 120.

24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

university policy and practice. Furthermore, it argued that to

refuse to renew his contract without providing him a statement

of the charges made against him and an opportunity to respond

to them at a formal hearing was a denial of his constitutional right

to due process of law.

Judge Doyle unexpectedly held this case under advisement for a

whole year, finally handing down a decision on March 12, 1970.

Judge Doyle agreed with Roth’s complaint and ordered:

Substantive constitutional protection for a university professor against

non-retention in violation of his First Amendment rights or arbitrary non¬

retention is useless without procedural safeguards. I hold that

minimal procedural due process includes a statement of the reasons why

the university intends not to retain the professor, and a hearing if the

professor appears at the appointed time and place.10

He further stated that if the defendants, that is, the university

officials, elected not to abide by the above order requiring a state¬

ment of reasons and a hearing, they

. . . shall be required, on or before June 1, 1970 to offer the plaintiff a

contract as a member of the faculty of the university for the academic

year 1970-1971, on terms and conditions no less favorable to him than

those contained in his contract for the academic year 1968-1969.11

The Wisconsin Attorney General applied to the Seventh Circuit

Court of Appeals for a stay of execution of the lower court’s deci¬

sion pending appeal, which was routinely granted. The case was

argued before the Seventh Circuit Court of Appeals on December

5, 1970. The Seventh Circuit Court issued its opinion on July 1,

1971, sustaining Judge Doyle’s decision. Speaking for a divided

court, Judge Thomas Fairchild agreed that a hearing and a state¬

ment of reasons were constitutionally required. He said :

Although the principle announced by the district court applies by its

terms to all non-retention decisions, an additional reason for sustaining

application in the instant cases, and others with a background of con¬

troversy and unwelcome expressions of opinion is that it serves as a

prophylactic against non-retention decisions improperly motivated by

exercise of protected rights.13

Chief Circuit Judge F. Ryan Duffy dissented on the grounds that

the decision would have the effect of making the federal courts the

“final arbiters of all similarly situated cases.” Duffy added:

In my view, the state’s interest in preserving a workable system of tenure

which includes, almost by definition the ability to select freely and

maturely its non-tenured teaching personnel, far outweighs any expectancy

which the plaintiff David Roth might have had in continued employment

at Wisconsin State University ... I further believe that the majority’s

10 310 F. Supp. 972 (1970), 978.

11 Ibid., p. 980.

12 446 F. 2d. 806 (1970), Circuit Judge Otto Kerner sided with Judge Fairchild.

1974]

Wade — The Roth Case

25

holding is both unprecedented and represents an unwarranted intrusion of

the Federal Judiciary into state educational systems.1'1

The stage was set, then, for an appeal to the United States

Supreme Court, and the Board of Regents decided almost imme¬

diately to go ahead with an appeal.

THE SUPREME COURT DECISION

The United States Supreme Court granted certiorari on October

26, 1971. 14 Oral arguments in the case were presented on January

18, 1972, and a decision was handed down on June 29, 1972. 15

The opinion of the court was delivered by Justice Stewart and

joined in by Justices White, Blackmun, and Rehnquist. Chief Jus¬

tice Burger filed a concurring opinion. Justices Douglas, Brennan,

and Marshall dissented. Justice Powell did not participate in the

decision.

Stewart's opinion began with a brief statement of the facts, a

review of applicable state statutes, and a summary of the lower

court opinions. Clearly, according to Stewart's opinion, Roth did

not have any tenure rights to continued employment. At that time

all teachers in the Wisconsin State University System were gov¬

erned by statutory tenure which said that a teacher acquired tenure

after being employed continuously for four years.16 A non-tenured

teacher under Wisconsin law was entitled to nothing more than

his initial one-year appointment. State statutes definitely gave uni¬

versity officials complete discretion over whether or not to renew

a non-tenured teacher's contract. While the Board of Regents rules

did not provide any protection for a non-tenured teacher whose

contract was not renewed, the Court indicated that they did pro¬

vide for a hearing and a statement of reasons if a teacher was

dismissed during the school year before the term of his contract

was up.17

According to the Court, the main issue was whether Roth “had

a constitutional right to a statement of reasons and a hearing on

the University’s decision not to rehire him for another year. We

hold that he did not.”18 The remainder of the opinion is devoted

to a statement of the rationale for this conclusion. While the Court

did not disagree with Judge Doyle's standard of simply weighing

the interest of the plaintiff in having his contract renewed against

the university's interest in refusing to renew his contract, it held

that it was necessary to go beyond this. Before weighing the inter-

13 Ibid.

14 404 U.S. 909 (1971).

15 408 U.S. 593 (1972).

1(1 1969 Wisconsin Statutes, Chapter 37.31.

17 408 U.S. 567 (1972).

18 Ibid., 569.

26 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

ests, the Court held that it was necessary to first determine “the

nature of the interest at stake/’19

In order to determine “the nature of the interest at stake” the

Court specified that it had to be decided whether or not the interest

was specifically protected under the Fourteenth Amendment’s pro¬

tection of liberty and property. The Court first considered the

liberty aspect. It stated that liberty included:

not merely freedom from bodily restraint but also the right of the indi¬

vidual to contract, to engage in any of the common occupations of life,

to acquire useful knowledge, to marry, establish a home and bring up

children, to worship God according to the dictates of his own conscience,

and generally to enjoy those privileges long recognized ... as essential

to the orderly pursuit of happiness by free men.20

If the university had made charges against him which could seri¬

ously damage his position in the community by implying, for

example, that “he had been guilty of dishonesty and immorality,”

then, the Court concluded, that would have been a completely dif¬

ferent situation. The Court would also have disapproved if the

university had in some way placed a stigma on him that took away

his freedom to continue engaging in his profession.21 However,

the Court decided that the university had done neither of these

things. In finalizing its discussion of the liberty issue the Court

held :

Hence, on the record before us, all that clearly appears is that the

respondent was not rehired for one year at one university. It stretches

the concept too far to suggest that a person is deprived of ‘liberty’

when he simply is not rehired in one job but remains as free as before

to seek another.22

The Court also maintained that the plaintiff had not proven that

the decision not to renew his contract was based on his free speech

activities. The Court stated : “Whatever may be a teacher’s rights

of free speech, the interest in holding a teaching job at a state

university, simpliciter, is not itself a free speech interest.”23

Next, the Court’s opinion discussed the property issue, admitting

that an individual’s property rights or interests may take a wide

variety of forms. The Court cited several cases defining those prop¬

erty interests. For example, the Court said that it had held that

welfare benefits are a property interest safeguarded by procedural

due process.24 Similarly, the Court stated that it has held that

public college professors cannot be dismissed from their positions

19 Ibid., 570-571.

20 Meyei's v. Nebraska , 262 U.S. 390 (1922), 399, cited by Justice Stewart at 408

U.S. 564 (1972), 572.

21 408 U.S. 564 (1972), 573-574.

22 Ibid., 575.

23 Ibid.

2* Ibid.

1974]

Wade — The Roth Case

27

without procedural due process25 and that they cannot be sum¬

marily dismissed during the terms of their contracts.26 Moreover,

the Court has maintained that even a substitute teacher who had

been employed two months could not be dismissed because of her

refusal to sign a loyalty oath.27 From these decisions, the Court

argued that a definition of property interests emerged :

To have a property interest in a benefit, a person must have more than

an abstract need or desire for it. He must, instead, have a legitimate

claim of entitlement to it. It is a purpose of the ancient institution of

property to protect those claims upon which people rely in their daily

lives, reliance that must not be arbitrarily undermined. It is a purpose

of the constitutional right to a hearing to provide an opportunity for a

person to vindicate those claims.38

The Court then made the distinction that property interests are

not created by the Constitution but by laws or rules enacted or

understandings promulgated by states. Hence, the plaintiff-respond-

ent’s property interest in having his contract renewed at Oshkosh

State University was created and limited by the terms of his

appointment which specifically secured his interest in employment

from September 1, 1968 to June 30, 1969.29 Indeed, the Court indi¬

cated the terms “made no provision for renewal whatsoever.”

Thus, it concluded its assessment of the property interest and the

entire opinion by saying:

In these circumstances, the respondent surely had an abstract concern

in being rehired, but he did not have a property interest sufficient to

require the University authorities to give him a hearing when they

declined to renew his contract of employment.

IV

Our analysis of the respondent’s constitutional rights in this case in

no way indicates a view that an opportunity for a hearing or a statement

of reasons for non-retention would, or would not, be appropriate or wise

in public colleges and universities. For it is a written Constitution that

we apply. Our role is confined to interpretation of that Constitution.

We must conclude that the summary judgment for the respondent

should not have been granted, since the respondent has not shown that

he was deprived of liberty or property protected by the Fourteenth

Amendment.30

DISSENTING AND CONCURRING OPINIONS IN ROTH

Justices Brennan and Marshall filed separate dissenting opinions.

They argued for a broader definition of liberty and property, say¬

ing that both plaintiffs were entitled to a statement of reasons and

25 Slochower v. Board of Education, 350 U.S. 551, cited at 408 U.S. 564 (1972), 577.

28 Weiman v. Updegraff, 344 U.S. 183, cited at 408 U.S. 564 (1972), 577.

'2' Connell v. Higginbotham, 403 U.S. 207, cited at 408 U.S. 564 (1972), 577.

28 408 U.S. 564 (1972), 577.

29 408 U.S. 564 (1972), 577-578.

30 Ibid., 578-579.

28 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

a hearing on the disputed issues.31 Justice Douglas in a more

lengthy opinion argued that a statement of reasons and a hearing

were necessary to guarantee individual rights against arbitrary

action. He stated:

When a violation of First Amendment rights is alleged, the reasons for

dismissal or for non-renewal of an employment contract must be ex¬

amined to see if the reasons given are only a cloak for activity or atti¬

tudes protected by the Constitution.33

Chief Justice Burger wrote a concurring opinion for the express

purpose of emphasizing one central point that he felt had been

obscured in the majority opinion. That point was that disputes

between a state educational institution and one of its teachers is a

matter of state concern and state law. Burger raised the doctrine

of abstention, as he has done in many other opinions since being

appointed to the Court. He concluded :

If relevant state contract law is unclear, a federal court should, in my

view, abstain from deciding whether he is constitutionally entitled to a

prior hearing, and the teacher should be left to resort to state courts on

the questions arising under state law.33

ASSESSMENT OF ROTH DECISION

The Roth case was an extremely complicated controversy arising

out of unusually explosive circumstances. As is true in any

emotionally-charged crisis, whether it be an automobile accident,

a robbery, or a riot, reasonable men can view the same factual

circumstances and come to completely opposite conclusions. This

was certainly true of the Roth case, and it became more and more

apparent with every individual interviewed, and every document,

report, newspaper article, or legal brief. A deep schism was created

over the personality of David Roth between those who regarded

him as politically naive and those who saw him as a political

revolutionary.

To the people of Oshkosh, Wisconsin, Roth presented himself

as a true revolutionary bent on the complete destruction of the

university and the American political system. His public state¬

ments, which were widely published in the press in Oshkosh and in

Madison and Milwaukee, lent a good deal of substance to that col¬

lective impression.

One faculty member characterized Roth as a “bright, personable,

abrasive bastard.” Another observed that:

when he set his mind to it he could be very charming, but when he got

tired-up he was a real fireeater. When there was controversy he seemed

31 408 U.S. 564 (1972), 587-592, 604-605.

33 Ibid., 579.

33 Ibid., 582.

1974]

Wade — The Roth Case

29

to thrive on the smoke of the battle. He had a sort of ‘Dr. Jekyll-Mr.

Hyde personality/

Roth characterized himself as having been “very immature and

naive” throughout the whole Oshkosh crisis. He felt that “speaking

out against injustice” was his responsibility as a member of the

academic community:

My non-retention resulted from a personality conflict because I embar¬

rassed the administration by criticizing their policies and their lack of

sensitivity to blacks. This was perceived by them as disloyalty to the

university and being in favor of violence.

During the four hours this writer spent with Roth, he certainly

gave the impression of being naive and trusting. For example, in

the middle of the evening he suddenly announced that he had to

leave to meet a friend but that I was free to go through all of his

files on my own. He was gone for over two hours. Needless to say,

this was quite surprising in that I was a total stranger to him.

Throughout the interview he came across as a very personable,

calm, and likable individual. Indeed, the evening was a very pleas¬

ant one.

However, if Roth had been more politically astute he could very

easily have taken many of the same stands, while remaining within

the bounds of professional ethics and responsibility as well as civil

discretion. In fact many of the things that he did and said (and

allegedly did and said) demonstrate such incredible naivete that

one might begin to question whether he might not have been

deliberately projecting this as a facade in the hope that it might

advance his cause.

After all, David Roth was definitely not a provincial person

from the homogeneous socio-economic political environment of a

rural community. He had graduated from Claremont Men’s College,

a small liberal arts college with a reputation for academic excel¬

lence located in Claremont, California (on the eastern edge of Los

Angeles County). He had gone to the University of California Law

School at Berkeley for two years. He had lived in cosmopolitan

San Francisco, while obtaining a master’s degree from San Fran¬

cisco State College. He had traveled extensively in the Philippines

and the Far East, while doing research for his doctoral disser¬

tation at the Claremont Graduate School, Claremont, California.

Finally, he had spent a year in Berkeley, California doing research,

prior to coming to Oshkosh. With this kind of background he should

have been able to discern the legitimate modes and boundaries of

dissent in a more sophisticated manner than he demonstrated in

the 1968-1969 academic year at Oshkosh.

Whatever David Roth’s intent was, however, an extensive in¬

vestigation of the circumstances and events surrounding and lead-

30 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

ing up to the federal litigation in the Roth case leaves little doubt

that the Oshkosh administration acted in a reasonable manner.

They made the decision not to renew Roth's contract mainly because

this was the most rational action they could take. Had they allowed

Roth to remain on the faculty, they would have been taking a

calculated risk of his escalating his protest actions and thus in¬

flaming an already dangerous situation. Roth's actions in dismiss¬

ing classes on two separate occasions in order to pursue his own

political objectives, his extensive use of the classroom podium to

discuss irrelevant material, his failure to give a final examination,

and his refusal to honor a request from the Vice-President of

Academic Affairs for a meeting all forced the Oshkosh adminis¬

tration to refuse to renew his contract.

IMPLICATIONS OF THE ROTH DECISION

The United States Supreme Court's decision in Roth was char¬

acteristic of the reasonable restraint the Burger Court has

demonstrated in matters of federal-state relationships. Its stance

regarding judicial activism is quite different from that of the

Warren Court. If this case had come before the Court one or two

years earlier, a vastly different decision would probably have been

reached. However, the majority opinion rightly recognized the

difficulties likely to be created when a federal court involves itself

in matters that might more prudently be left to the states.

One problem that would probably have occurred if the lower

court decision in Roth had been sustained is that an administrator

would be extremely reluctant to decide against renewing an indi¬

vidual's contract if he knew that this would oblige him to file formal

charges and go through a formal hearing. By the nature of the

case the definition of good teaching is rather subjective and precise

documentation of bad teaching is very difficult. After all, a teacher

can always be defended by such arguments as (1) every instructor

has some bad days, (2) he was not feeling well, (3) some students

think he is wonderful, or (4) whatever his faults, he is improving.

The burden of proof in a formal, adversary hearing procedure thus

falls on the administrator instead of the teacher. As a practical

matter probably very few contracts would go unrenewed given

this kind of situation. How many administrators could reasonably

be expected to non-retain a professor when the benefits of his non¬

retention might very well be overshadowed by the bitter rhetoric

and controversy caused by the hearing? If, as a matter of policy,

an administrator did decide to proceed with hearings under such

circumstances, he would probably have to keep a dossier on every

faculty member, dating and documenting every incident that might

1974]

Wade — The Roth Case

31

conceivably bear upon a teacher’s competence. This is a prospect

that would not be welcomed by many, especially not by those who

are the strongest advocates of extensive hearing procedures.

Moreover, if Roth had been upheld, faculty members who would

ordinarily be releasable would have an incentive to create ambigu¬

ous and volatile situations which would make it very difficult, if

not impossible, to remove them. A dissident faculty member could,

for example, set himself up as an outspoken defender of a minority

group in order later to be able to accuse the administrators of being

racist or bigoted for not renewing his contract, and in many re¬

spects Roth did exactly that.

Justice Douglas’ dissenting opinion in Roth is a classic example

of the limits of adjudication arising from our judicial adversary

system. The Court simply does not have the required investigatory

powers similar to those available to state (and national) legisla¬

tures. For example, his opinion flippantly glosses over the events

and circumstances, in the case that could not be covered adequately

in the appellate briefs and oral arguments. This is how he charac¬

terized Roth’s behavior:

Though Roth was rated by the faculty as an excellent teacher, he had

publicly criticized the administration for suspending an entire group

of 94 black students without determining individual guilt. He also

criticized the university’s regime as being authoritarian and autocratic.

He used to discuss what was being done about the Black episode; and

one day, instead of meeting his class, he went to the meeting of the

Board of Regents.31

As a result, he concluded that the district and circuit court de¬

cisions should be sustained.

The problem with his capsulization is that is grossly oversimpli¬

fies a very complex set of circumstances and events. It completely

ignores the intensity and the magnitude of Roth’s actions and the

context in which they occurred. The fact that Roth attended a

Board of Regents meeting is portrayed as a petty irregularity

instead of the blatant breach of professional responsibility that it

seemed to be according to most reports. Moreover, it ignores the

other occasion when he failed to meet his classes, that is, when he

attended the meeting between the Oshkosh president and the

parents of the black students. It also fails to note the fact that he

violated an explicit university rule by failing to give a written

final examination. And it fails to mention his insubordination when

he cavalierly spurned the request to meet with his Vice-President

for Academic Affairs.

Finally, if the lower courts’ decisions had been sustained, the

traditional distinction between tenured and non-tenured faculty

408 U.S. 564 (1972), 576.

32 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

would have been obliterated. Instant tenure, with all its ensuing

rights and privileges, would have been created for all faculty

merely upon signing an initial employment contract. Such a rule

would have been in direct contradiction of accepted practice on

the part of 1,080 colleges and universities with 300,000 faculty

members and 6,000,000 students.35 It would have meant that all

non-tenured, non-retained faculty members could have appealed

their cases.

The amount of administrative and faculty time that would have

had to go into investigating, documenting, and holding hearings

would have been staggering. In the Wisconsin State University

System, for example, there were some 206 non-tenured faculty

contracts that were not renewed for the 1970-1971 academic year

alone.36 The cost of paying the investigating administrators’

salaries as well as the legal expenses incurred would have been

overwhelming. The federal courts, of course, would not have ever

had to face these particular consequences.

In short, the Roth case raised this basic question: is there any

rational justification for distinguishing between the rights enjoyed

by tenured faculty and those to which non-tenured faculty should

be entitled ? Why should one group of faculty members have greater

rights than another? In criminal law the granting of certain pro¬

cedural rights to one group of alleged criminals, while granting a

different set to another, would be considered unconstitutional be¬

cause it would be a denial of “the equal protection of the law.”

In Roth Judge Doyle and the Seventh Circuit Court of Appeals

said in effect that there was really no rational basis for making

such a distinction, although they never specifically discussed

whether the distinction was constitutionally valid or even rationally

defensible. By so doing they seemed to make educational disci¬

plinary cases more closely akin to criminal proceedings and, in the

process, take away the right of a professional group to organize

and regulate their profession according to their own free choice.

The primary reason for differentiating between the rights of

tenured faculty and non-tenured faculty is, of course, that only

performance or experience can provide a sound basis for the pre¬

sumption by an institution that a new teacher has attained a degree

of excellence deserving of tenure. Initial appointments cannot

ordinarily be made on grounds providing sufficient assurance that

a person will reach this standard of excellence. For this reason it

35 Amici Curiae in support of the State of Wisconsin’s Petition for a Writ of Cer¬

tiorari, p. 2, written by the Honorable Robert W. Warren, Attorney General of Wis¬

consin, Assistant Attorney General Charles A. Bleck, and Assistant Attorney General

Robert D. Martinson, October Term, 1971, all at 114 East, State Capitol, Madison,

Wisconsin, 53702.

30 Ibid., p. 13.

1974]

Wade — The Roth Case

33

is inherently unfair to assume that a non-retention decision, made

early in an instructor’s affiliation with an institution, represents a

denial of academic freedom rather than a reasonable assessment

of his ability as a teacher and a scholar.37

If the Supreme Court had affirmed the Roth decision, the federal

judiciary would have been allowed to expand its role in school

disciplinary cases, and where some federal judges might have gone

from this point, if given encouragement, is anyone’s guess. Public

policy would have gradually ceased to reflect the wishes of the

people as expressed through their elected representatives on state

and local governing boards. Instead, it would have reflected the

judgment of a few appointed federal judges far removed from the

express wishes of the people.

Fortunately, the Supreme Court made a decision in Roth that

promises to arrest the continued expansion of the federal judiciary

in educational disciplinary cases. The Court seems to be moving

in the direction of giving the job of policy-making back to state

and local governments not only in the area of educational discipline

but also in other areas where continued expansion of judicial

policy-making occurred during the Warren Court era.

There is no doubt that local and state governing boards, as well

as the voters who either directly or indirectly select them, occasion¬

ally make decisions that by most rational standards of measurement

are wrong. But in a democratic republic such as the United States,

do they not have the right to be wrong? And do they not have the

right to be free from having a decision superimposed on them by

federal courts against deliberate policy decisions they have made?

Then, too, careful analysis of the two lower court decisions in the

Roth case demonstrates that it is at least an open question whether

they will be any less reasonable in their decisions than some federal

judges have been.

To the extent that federalism has been challenged by the lower

federal court decisions in Roth and in numerous other educational

disciplinary cases at both the secondary and higher educational

levels, federal judges (and to some extent the legal profession

itself) have done a disservice to American society. They have

raised relatively minor injustices into constitutional issues which

have strained the federal system. If the Burger Court had not

reversed the district and circuit court’s decisions in Roth and hence

reversed the trend established in educational disciplinary cases

in the later 1960’s, the state and local governments’ ability to con¬

trol educational policy in their own schools and colleges would have

continued to decrease. Further erosion of state control of educa-

37 William S. Van Alstyne, “Tenure, A Summary, Explanation, and Defense,” AAUP

Bulletin, LVII (September, 1971), pp. 332-333.

34 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

tional disciplinary matters could have contributed to the death of

federalism which the framers of the Constitution believed vital to

the cause of freedom.

REFERENCES

JENNER, ALBERT E., JR., KAMIN, CHESTER T., and FRANCH,

RICHARD T. Brief of the Board of Governors of State Colleges and

Universities of Illinois, the Board of Regents of Regency Universities of

Illinois, the Board of Trustees of Southern Illinois University, the

American Council on Education and the Association of American Colleges

as Amici Curiae, in David F. Roth v. The Board of Regents of State

Colleges in the United States Court of Appeals for the Seventh Circuit.

Chicago: United States Law Printing Co., 1970.

MARZETTE V. McPHEE, 294 F. Supp. 562 (1968).

ROTH V. BOARD OF REGENTS OF STATE COLLEGES, 310 F. Supp. 972

(1970), 446 F. 2d 806 (1971), 408 U.S. 654 (1972).

FINAL REPORT OF THE SPECIAL PROBLEMS COMMISSION, P. J.

Thompson, Chairman. Released by Wisconsin State University — Oshkosh,

December 15, 1969. (Mimeographed)

STEINGLASS, STEVEN H., and REYNOLDS, ROBERT L. Brief of Ap¬

pellee in David F. Roth v. Board of Regents of State Colleges, in the

United States Court of Appeals for the Seventh Circuit . Madison,

Wisconsin: Gilman Press, Inc., 1970.

- . Brief of the Respondent to a Petition for a Writ of Certiorari to

the United States Court of Appeals for the Seventh Circuit, in the Board

of Regents of State Colleges v. David F. Roth, in the Supreme Court of

the United States. Madison, Wisconsin: Gilman Press, Inc., 1971.

WARREN, ROBERT W., and BLECK, CHARLES A. Brief of the Appellants

in David F. Roth v. The Board of Regents of State Colleges, in the

United States Court of Appeals for the Seventh Circuit. Madison, Wis¬

consin, State of Wisconsin Bureau of Documents, 1970.

• - , and MARTINSON, ROBERT D. Petition for a Writ of Certiorari

to the United States Court of Appeals for the Seventh Circuit, in the

Board of Regents of State Colleges v. David F. Roth, in the Supreme Court

of the United States. Madison, Wisconsin: State of Wisconsin Bureau of

Documents, 1971.

ADDENDUM

After this paper went to press, a federal court jury in Madison,

Wisconsin, on November 9, 1973, awarded David Roth, now an

assistant professor of political science at Purdue University, $6,746

in damages because, in their judgment, his constitutional rights

of free speech were violated. The damages were assessed against

Oshkosh administrators, President Roger Guiles, Vice-President

Raymond Ramsden, Dean Arthur Darken, and two members of

the Oshkosh Political Science Tenure Committee, David Chang and

Charles Goff, all of whom participated in the decision not to retain

Roth.

It must be understood that Roth was awarded these damages in

a civil suit which he hied after the United States Supreme Court

1974]

Wade — The Roth Case

35

in 1972 had ruled on the procedural question that the due process

clause of the Fourteenth Amendment did not require that a non-

tenured professor be given a hearing in a non-retention proceeding.

Hence, it must be emphasized that the civil suit for damages, con¬

cerning the substantive question as to whether Roth’s constitutional

right of free speech had been violated, has no direct legal bearing

on the procedural decision handed down by the Supreme Court.

In spite of the jury’s verdict to the contrary, I must respectfully

disagree and stand by my own conclusions which are based upon

an extensive investigation of the events and related constitutional

issues.

WILD SOILS OF THE PINE-POPPLE RIVERS BASIN1

Francis D. Hole

University of Wisconsin — Madison

and

Wisconsin Geological and

Natural History Survey

The preservation of wild rivers involves the protection of the

surrounding ‘‘wild” soils. The usage of the term is new as applied

to soils, but the concept is old. Wild soils are those which still

function as integral parts of their native ecosystems. They have

not been domesticated through drastic human manipulation of plant

and animal populations and of the soils themselves, with resulting

modifications in local cycling of nutrients and water. It is note¬

worthy that modern soil classification does not adequately

recognize the difference between a wild soil and its cultivated

counterpart, both of which are given the same name! This is an

ecological error, which may be corrected by recognizing wild and

domesticated variants of soils.

The purpose of this paper is: 1) to summarize and interpret,

in the context of the ecosystems of the Pine-Popple Rivers Basin,

field and laboratory data obtained on soils of the area by the Wis¬

consin Geological and Natural History Survey, particularly during

the study in Florence County in 1958-1961 (Hole et al, 1962) ;

and 2) to delineate some future studies of wild soils to which the

Basin is well suited.

LITERATURE REVIEW AND METHODS

The Pine-Popple Rivers Basin has been clearly delineated in

maps recently published by Mills (1972) and Becker (1972), show¬

ing what portions of Forest and Florence Counties are included in

it. Table 1 traces changes in terminology for the five general kinds

of soils that have been recognized in the Basin since 1885. For an

explanation of the technical classification terms given in paren¬

theses for 1962 in Table 1, and throughout Table 2 and Fig. 2, the

reader is referred to the new U.S.D.A. soil classification scheme

(Soil Conservation Service, 1973). The terms used in the legend of

Fig. 1 of this paper are simplified.

1 This is paper No. 9 in a series of studies of the Pine-Popple Wild Rivers area of

northeastern Wisconsin ; the publication of which began in volume 60 of these

Transactions.

37

TABLE 1. SOME DESIGNATIONS OF FIVE GENERAL KINDS OF SOILS IN THE PINE-

POPPLE RIVERS BASIN

38 Wisconsin Academy of Sciences, Arts and Letters

[Vol. 62

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2 Whitson et al., 1916 (Soil map of Northeastern Wisconsin).

3Whitson et al., 1921 (Soil map of Northern Wisconsin).

4Hole et al., 1962 (Soil map of Florence County, Wisconsin).

TABLE 2. A CLASSIFICATION OF MAJOR SOIL SERIES IN THE PINE-POPPLE WILD RIVERS BASIN1

1974]

Hole — Wild Soils of Pine-Popple Rivers Basin

39

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40

Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

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1974] Hole — Wild Soils of Pine-Popple Rivers Basin

41

Field methods used were those for general or reconnaissance

soil mapping. These are described in the Soil Survey Manual (Soil

Survey Staff, 1951). Soils were observed at intervals of about one

half to two miles. Laboratory analyses are reported elsewhere

(Hole et al., 1962) .

Bouma and Hole (1971) presented evidence that two portions of

a body of soil may behave very differently, where one portion is

undisturbed under native vegetation and the other is managed for

agricultural purposes. “Management-induced structural changes

can strongly alter hydraulic properties of a soil. These changes

may be of sufficient magnitude and continuity to consider recog¬

nition of hydraulic soil phases of established soil series. .

SOILS AND SOILSCAPES OF THE PINE-POPPLE

RIVERS BASIN

Soils

The soils occupy the area of the “vast unbroken forest” lying

on either side of the “thin blue thread” of each wild river (Mills,

1972). Of the forty-two kinds of soils (classified at the soil series

level) and two kinds of miscellaneous land types (Alluvial soils and

Granitic Rockland) described in the soil survey report for Florence

FIGURE 1. General soil map and block diagram of the Pine-Popple Wild

Rivers Basin. Legend: C = clayey soils (Hibbing and Ontonagon); F = loca¬

tion of the village of Fence; LA = loamy soils including rock outcrops and

Ahmeek loams; LE i= loamy soils including Pence and Iron River loams and

sandy loams; P t= peat; Sa = sandy soils (Omega, Vilas); Si = silty soils

(Stambaugh, Goodman, Fence, Bohemian). Dotted lines are soil boundaries

adapted from Hole et al., 1968. Large lakes are labeled L.

42 Wisconsin Academy of Sciences, Arts and Letters

[Vol. 62

FIGURE 2. A classification chart for major great groups of

soils in the Pine-Popple Wild Rivers Basin. The Haplorthods,

Eutroboralfs and Udorthents are well drained members of

their respective orders. Haplaquods, Haplaquepts and all

Histosols are poorly to very poorly drained under natural

conditions. See Table 2.

County, twenty-six soils (Table 2 and Fig. 2) and one land type

are shown in Figs. 3 through 6. Nineteen of these soils are Spodo-

sols (Podzols), which essentially consist in cross-section (profile

view as shown in Fig. 6 for Vilas, Crivitz, Pence, Hiawatha, and

Au Train soils) of four main horizons (layers) : surface organic

horizons (the 01 horizon is forest litter; the 02 is humus) ; a

bleached pale mineral topsoil (A2 or albic horizon) ; a dark brown

subsoil (Bhir or spodic horizon enriched in humus and iron) and

yellowish brown to reddish brown glacial drift (C horizon: initial

material). Wetland soils include peats (Histosols) and poorly

drained mineral soils (Aquepts, Aquolls) which have black topsoil

(Al) and grey (anaerobic and gleyed) subsoil. Some clayey soils

(Gray Wooded soils: Eutroboralfs), including the Ontonagon and

Hibbing series, have a subsoil enriched in clay (Bt or argillic hori¬

zon) instead of iron and humus.

1974] Hole — Wild Soils of Pine-Popple Rivers Basin

43

portions of the Basin. The forest cover on these wild soils is not shown. (Fig.

courtesy of the Wisconsin Geological and Natural History Survey.)

FIGURE 4. Silty soils (Si in Fig. 1) and sandy soils (Sa) of the Pine-

Popple Wild Rivers Basin. Goodman silt loam is over glacial till, Stambaugh

silt loam is on glacial outwash sand and gravel, Fence (and Tipler) silt

loam on acid glacial lake beds, and Bohemian (and Brimley and Bruce)

silt loam on calcareous lake beds; Vilas sand (and associated Randville)

on deep outwash sands. The forest cover on these wild soils is not shown.

The unlabeled upland on the extreme right is occupied by Ontonagon soils

in this idealized sketch. (Fig. courtesy the Wisconsin Geological and Nat¬

ural History Survey.)

44 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

It is interesting that the Basin contains many upland soils that

are “hybrids”, that is consisting of a Spodosol (Podzol) over a

Eutroboralf (Gray Wooded) all in a single soil profile (Beaver,

1966). The horizon sequence in these “double” soil profiles is

typically: 01, 02, A2, Bhir, A'2, B't, C. The prime symbol (') in

the fifth and sixth horizons indicates that they are in the second or

lower sequence. These two horizons are commonly fragipan hori¬

zons in the Pine-Popple Rivers Basin. The term fragipan (Olson

and Hole, 1967) denotes compactness and resistance to water move¬

ment and root penetration. Double soil profiles (bisequal soils) with

two light colored A2 horizons and two somewhat dark B horizons

are shown in Figs. 4, 5 and 6 for soils named Fence, Tipler,

Bohemian, Brimley, Superior, Ulby, Padus and Stambaugh. Theo¬

ries about the formation of the soils are described elsewhere (Hole

et at., 1962; Buol et al., 1973). Soil profiles, like those illustrated

in the sketches, took thousands of years to form and represent an

impressive environmental record. In deciphering this record, re¬

searchers look for evidence of the influence on soil of different for¬

est types and climates, past and present, as discussed by Milfred

et al. (1967) and Buol et al. (1973).

HIBBING ONTONAGON RUD- PICK- BERG- SUPER- MANISTEE UGLY MENOMINEE

silty clay siltyctay YARD FORD LAND IOR loamy sandy loamy sand

loam loam silty silty clay sand sand loam

clay clay loam

loam loam

FIGURE 5. Clayey soils (C of Fig*. 1) include Hibbing (on glacial till) and

Ontonagon (on lake clays) with associated more poorly drained soils (Zim,

Tromald; Rudyard, Pickford, Bergland). Where sandy coverings occur, Su¬

perior and associated sands and loamy sands are found. Vilas and Hiawatha

sands are very deep (Sa of Fig. 1). The forest cover on these wild soils is not

shown. (Fig. courtesy the Wisconsin Geological and Natural History Survey.)

1974] Hole — Wild Soils of Pine-Popple Rivers Basin

45

OMEGA VILAS CRIV- PENCE feDUS STAMBAUGH HIAWATHA AU TRAIN

sand sand ITZ sandy loam silt loam sand sand

loamy loam

sand

FIGURE 6. Loamy soils (LE, Fig. 1) are chiefly Pence sandy loam and the

deeper Padus loam* with peat (P, Fig. 1) in deep kettles; and Vilas sand and

Emmert gravelly sandy loam on esker ridges (of which there are about 200

in the Pine-Popple Basin). The forest cover on these wild soils is not shown.

(Fig. courtesy the Wisconsin Geological and Natural History Survey).

Soils capes

Soils are observed not only in the two-dimensional exposures

called soil profiles, but also in geographic bodies and patterns of

soil bodies on the landscape. A soilscape is the portion of a land¬

scape consisting of the soils (Buol et al., 1973). Fig. 1 shows in

a very general way the arrangement of major soil groupings in the

Pine-Popple Rivers Basin. The eastward change that Mills (1972)

observed from the canoe, from “gravelly, richer soils of the upper

river ... to poor sand in the lower stretches” is indicated in the

Figure by the dotted boundary between silty (Si) or loamy (LE)

soils and sandy (Sa) soils. Soilscapes are shown in much greater

detail in the idealized sketches of Figs. 3 through 6. Throughout

the Basin droughty soils (such as Vilas and Emmert) and wetlands

(such as Bergland and Bruce mineral soils and peat) are typically

in close proximity (within a mile or less). Tree-tip mounds and

associated pits are common microtopographic features in the forest

(Gaikawad and Hole, 1961).

46 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Factors of soil formation. The bedrock of the Basin consists of

Precambrian volcanic and metamorphic rocks, some of which are

exposed in the river channel bottom, as noted by Mills (1972).

Glacial drift as much as 250 feet thick was deposited over the bed¬

rock by continental glaciers, the last of which melted away from

the Basin about 11,000 years ago or soon thereafter (see Bryson

and Wendland, 1967). Coarse wind-blown silt was laid down in a

blanket as much as a foot or two thick locally. Land forms include

moraines, drumlins, eskers, outwash and lacustrine plains. Since

glaciation, soils have formed from these materials under shifting

continental climatic zones and biotic communities.

The biotic factor has been the most dynamic. Coniferous and

mixed forest plant communities and associated fauna have left their

impress on the soils of the Basin, as Wilde et at. (1949) and Hole

et al. (1962) have discussed elsewhere, and as research in the area

can be expected to elucidate in the future.

Soil processes

Soils of wild river forest lands perform important functions in

the hydrologic and nutrient cycles. Water is absorbed and stored

in soil, from which the vegetation draws vast quantities and

transpires them during the growing season. The relatively small

portion of the annual rainfall that is not used by vegetation perco¬

lates to the water table, and moves to bogs, lakes, and rivers (Hole,

1974). Some small brooks flow for considerable distances almost

out of sight below a mat of roots of hemlock, cedar and other hy¬

drophilic plants. With the spring melting of snow and frozen sur¬

face soil on uplands, before deciduous trees have leafed out, some

water moves laterally (down-slope) through the upper soil horizons

over the fragipan.

Except where rivers form cutbanks (which are comparatively

rare in the Basin), the soils are shielded by vegetation from run¬

ning water and they serve to filter out sediment, even much fine

colloidal material. On the other hand, the aqueous extracts of the

living plants, decaying wood and soil organic matter in the forest

do contribute dark humic material to the wild rivers, giving them

their characteristic yellowish brown tint. Black and Christman

(1963) found that waters colored by organic materials from wilder¬

ness lands are surprisingly uniform over the continent and typically

contain fulvic acids (about 90% of the organic fraction) and hymato-

melanic and humic acids, ranging in all from 10 to 120 grams per

liter with seasonal variability. These materials occur as negatively

charged colloidal particles, as well as dissolved substances (Birge

and Juday, 1934) . Iron (0.1 to 2.0 ppm in water) is commonly asso¬

ciated with the organic matter. These data are not from the Pine-

1974] Hole — Wild Soils of Pine-Popple Rivers Basin 47

Popple Rivers Basin but probably represent conditions in the lakes

and streams there. The humus in wild rivers stimulates the growth

of algae somewhat, perhaps because it carries nutrient elements

in exchangeable form.

Nutrient cycling involves: 1) both mineral and humus portions

of the soil, in which is present a complete complement of elements

needed by native plants ; and 2) the above-ground forest vegetation.

The first category has always been considered to include fallen

logs, branches, needles, leaves, and fruiting bodies. These, along

with residues of associated decomposers, make up the surficial or¬

ganic horizons of the soil. However, the bulk of the forest consists

of dead wood of the standing boles and large branches. This wood

differs from that of fallen logs of the soil proper only in the higher

proportion of relatively undecomposed material. It is difficult to

categorize this bulky dead organic matter as waste, because it con¬

tributes so importantly to the forest community by giving both

mechanical support and nutrients to the living portion. Mills (1972)

observed during his canoe trip that “a barn overgrown by popple

decays into oblivion.” That oblivion includes tangible wild soil. A

roadside picnic grounds built by the Depression era Civilian Con¬

servation Corps near the Highway 139 bridge over the Popple

River was by 1970 overgrown with popple, grass and briars. “We

observed nature struggling to heal the scars inflicted by man.”

OPPORTUNITIES FOR STUDIES OF WILD SOILS

“No other geographic region has such a variety of life forms

and such an ability to manifest the effect of environmental factors

as does the forest” (Wilde et al., 1949). Our knowledge of inter¬

actions between soils and native plants is far from complete. Most

effort in soil survey has gone into the characterization and mapping

of soils for agricultural, silvicultural and engineering purposes.

Priority has been given to the demands for food and fiber for peo¬

ple and domestic animals, and materials for their shelter; the

demand for stable soil conditions for the support of roads, buildings

and pipelines ; and the demand for soil conditions suitable to absorb

and filter water and, locally, liquid wastes. The soils that research¬

ers have observed to a depth of four or five feet in cultivated fields

(which occupy little of the area of the Pine-Popple Rivers Basin)

are in an ecological sense “decapitated” soils. They have lost the

root mass of the original forest and the humus and forest litter

layers, as well as the standing, three-storied forest itself. They

have been made more compact than they were, and very probably

conserve water and nutrients less efficiently than they did before

the impact of man’s activities (Bidwell and Hole, 1965). Accel-

48 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

erated soil erosion has occurred on cultivated fields by water and

wind, and on slopes above road-cuts, by slump during thaws in

spring.

The unusual opportunities that the Pine-Popple Rivers Basin

offer for the study of wild soils may be indicated by the following

topics.

(1) Relationships between individual species of plants and soils.

We know that well developed Spodosols (Podzols) commonly occur

under hemlock trees (Tsuga canadensis), (Milfred, et al, 1967),

but need to investigate less obvious influences of other plant species

on soil genesis in natural forest ecosystems of the Basin.

(2) Relationships between plant communities and soils. We

know that mixed hardwood-conifer forest litter supports more

earthworm (Lumbricus terrestris) activity than do pure coniferous

stands. This is part of the reason for the presence of an Al horizon

in the mixed forests and its absence in many coniferous stands.

Many such synecological details of soil genesis need to be worked

out in the Basin.

(3) Relationships betiveen individual species of animals and

soils. Birds bring to soils grit regurgitated from their crops and

nutrients in the form of excreta. Soils under roosting sites of crows

( Corvus sp.), for example, receive notable deposits of these two

kinds of materials. The work of beavers ( Castor sp.) in flooding

low-lying soil bodies alter soil properties over considerable areas,

and these changes await serious study. One could list many species

of animals whose influence on soils has not been documented

carefully.

(4) The influence of human activity on soils. Comparative soil

studies are needed between fields, pastures, second growth forests,

selectively cut forests, and undisturbed forests. Of particular in¬

terest would be : 1) differences from site to site in the water budget,

which could be measured by recently developed physical methods

(Bouma et al., 1973) , and 2) effects of different methods of harvest¬

ing forest products in areas adjacent to the Rivers (Boyle and Ek,

1972).

CONCLUSION

The wild soils of the Pine-Popple Rivers Basin function to sup¬

port forests and thus to influence the natural temperature, chemis¬

try and steady flow regimes characteristic of these wild rivers. A

general soil survey has been made of the watershed, particularly in

the Florence County portion, but detailed soil ecological work has

scarcely begun. The opportunities for long-term research on wild

soils are excellent in the Basin.

1974] Hole — Wild Soils of Pine-Popple Rivers Basin 49

LITERATURE CITED

BEAVER, ALBERT J. 1966. Characteristics and genesis of some bisequal

soils in eastern Wisconsin. Ph.D. Thesis. University of Wisconsin,

Madison.

BECKER, GEORGE. 1972. Annotated list of the fish of the Pine-Popple Basin.

Trans. Wis. Acad. ScL, Arts and Lett. 60:309-329.

BIDWELL, O. W., and F. D. HOLE. 1965. Man as a factor of soil formation.

Soil Sci. 99:65-72.

BIRGE, E. A., and C. JUDAY. 1934. Particulate and dissolved organic matter

in inland lakes. Ecol. Monographs 4:440.

BLACK, A. P., and R. F. CHRISTMAN. 1963. Characteristics of colored sur¬

face waters. Am. Waterworks Assn. J. 55:753-770.

BOUMA, J., and F. D. HOLE. 1971. Soil structure and hydraulic conductivity

of adjacent virgin and cultivated pedons at two sites: a Typic Argiudoll

(silt loam) and a Typic Eutrochrept (clay). Soil Sci. Soc. Amer. Proc.

35:316-319.

BOUMA, J., W. A. ZIEBELL, W. G. WALKER, P. G. OLCOTT, E. McCOY,

and F. D. HOLE. 1973. Soil absorption of septic tank effluent. Inform.

Circ. 20, Wis. Geological and Nat. Hist. Survey, University of Wisconsin-

Extension, Madison.

BOYLE, J. R., and A. R. EK. 1972. An evaluation of some effects of bole and

branch pulpwood harvesting on site macronutrients. Canad. J. Forest Res.

2:407-412.

BRYSON, R. A., and W. M. WENDLAND. 1967. Radiocarbon isoclines of

the retreat of the Laurentide ice sheet. Tech. Rept. 35, Dept. Meteorology,

Univ. Wisconsin, Madison.

BUOL, S. W., F. D. HOLE, and R. J. McCRACKEN. 1973. Soil genesis and

classification. Ia. State Univ. Press.

CHAMBERLIN, T. C. 1883. Geology of Wisconsin, Vol. I (with soil map,

1882). Commissioners of Public Printing.

GAIKAWAD, S. T., and F. D. HOLE. 1961. Characteristics and genesis of a

Podzol soil in Florence County, Wisconsin. Trans. Wis. Acad. Sci., Arts

and Lett. 50:183-190.

HOLE, FRANCIS D. 1974. Wisconsin soils, (in press, Univ. Wis. Press). Wis.

Geological and Nat. Hist. Survey. University of Wisconsin-Extension,

Madison.

HOLE, FRANCIS D., G. W. OLSON, K. O. SCHMUDE, and C. J. MILFRED.

1962. Soil survey of Florence County, Wisconsin. Wis. Bui. 84, Soil series

59, Wis. Geological and Nat. Hist. Survey, University of Wisconsin-

Extension, Madison.

HOLE, F. D., M. T. BEATTY, C. J. MILFRED, and A. J. KLINGELHOETS.

1968. Soils of Wisconsin (Wall Map, Scale 1:710,000). Wis. Geological

and Nat. Hist. Survey, University of Wisconsin-Extension, Madison.

MILFRED, C. J., G. W. OLSON, and F. D. HOLE. 1967. Soil resources and

forest ecology of Menominee County, Wisconsin. Bui. 85, Soil series 60,

Wis. Geological and Nat. Hist. Survey, University of Wisconsin-Exten¬

sion, Madison.

MILLS, J. 1972. Canoeing on the wild rivers Pine and Popple. Trans. Wis.

Acad. Sci., Arts and Lett. 60:239-263.

OLSON, G. W., and F. D. HOLE. 1967. The fragipan in soils of northern

Wisconsin. Trans. Wis. Acad. Sci., Arts and Lett. 56:174-184.

Soil Conservation Service. 1974 (in press). Soil Taxonomy of the National

Cooperative Soil Survey. USDA. Gov. Printing Office, Washington, D. C.

Soil Survey Staff. 1951. Soil Survey Manual. USDA Handbook No. 18. Gov.

Printing Office, Washington, D. C.

50 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

WHITSON, A. R., W. J. GEIB, CARL THOMPSON, CLINTON B. POST,

A. L. BUSER, L. R. SCHOENMANN, and ARTHUR E. TAYLOR. 1916.

Reconnaissance soil survey of northeastern Wisconsin. Bui. 37, Soil Series

12, Wis. Geological and Nat. Hist. Survey, University of Wisconsin,

Madison.

WTHITSON, Y., J. DUNNEWALD, and CARL THOMPSON. 1921. Soil survey

of northern Wisconsin. Bui. 55, Soil Series 27, Wis. Geological and Nat.

Hist. Survey, University of Wisconsin, Madison.

WILDE, S. A., F. G. WILSON, and D. P. WHITE. 1949. Soils of Wisconsin

in relation to silviculture. Pub. No. 525-49. Wis. Conservation Dept.,

Madison.

THE NOVEL AS A VEHICLE TO TELL THE STORY

OF THE MENOMINEE INDIANS

William Steuber

Madison

What is to be done about the American Indian? The question

has never been successfully answered. It has been a puzzle ever

since Columbus reached the Caribbean and thought he had found

India. The American Indian was an annoying problem to the early

pioneers, to the westward settlers, to the United States Army, to

every president and Congress from George Washington through

Richard Nixon. The Indian question has been particularly trouble-

some in recent years with the take-over at Alcatraz, at Wounded

Knee, and with the occupation of the Bureau of Indian Affairs in

our nation’s capital.

Wisconsin still puzzles over the Indian much as our forefathers

did before we became a state. Currently the Menominee are in

the headlines. Two decades ago they were prosperous, second only

to the oil-rich Oklahoma Indians. Under pressure from Congress,

they voted to terminate their reservation and become free citizens,

supporting their own county. Today they are poverty ridden, and

have successfully pleaded to again become wards of the federal

government. The processes are now underway to disestablish

Menominee County and to reestablish the Menominee Indian

Reservation.

There are no easy answers to Indian problems, either among the

Indians themselves, or in the Congress of the United States, Wis¬

consin, or anywhere.

The American Indian is still a generous natural resource. We

made fortunes from him, giving him beads and brandy for his

beaver. We traded cheap promises for good land. We put his profile

on our pennies and our nickels, but let him own very few of them.

We are not finished merchandising Indians. We are now capital¬

izing our guilt, our compassion, our social consciousness in a flood

of belated attention with varying motives. Jane Fonda and Marlon

Brando have been typically loud in Indian causes. One wonders

how much of this is truly for the Indian, how much for their own

publicity.

Or take so many of the present books that seem to speak for the

Indian. The accounts seem so stylized. Find an old Indian. Take

51

52 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

a tape recorder. Ply him with whiskey. Let him cry and sing his

heart out. Then take your notes and your tapes, rewrite his story

into the current pattern of popular guilt, twist his halting words

into your ideology, into your sense of timing, into your telling

phrase, your witticism, your creation of something intimate, some¬

thing significant.

One becomes highly suspect of ghost-written accounts, as-told-to

presentations, two-person authorships and their ability to bulge,

perhaps rupture, the honest line between fiction and non-fiction.

Such dubbed-in writing may appear to present solid and substan¬

tial fact which in reality may be highly manipulated and distorted

by the views, motivations, aims and skills of the co-author. A

strong compulsion to take such liberties may be based on present

reader preference for non-fiction, and better sales, over true novels.

Nevertheless, the novel is a writing form of historic power and

honesty. In the hands of a Charles Dickens, an Upton Sinclair, a

John Steinbeck, its created characters become more alive than

actual people. Social problems are shown in the glare of truth, and

a great strength is born to help solve those problems.

The Menominee Indians today are a troubled minority people,

protesting white dominance more intensely now than at any other

time in history. In order to understand and appreciate the prob¬

lems of the Menominee Indian today, one should know how he lived

before the white people overwhelmed him. One should know his

basic weaknesses in the presence of a different culture, his beliefs,

his fears, how he handled his world when he alone walked it.

The Great Lakes Menominee, before the white man, were a small

and prosperous tribe. They had wild rice, maple sugar, beaver,

sturgeon, bear and deer, wild fowl and passenger pigeon, in such

plenty that food was no problem. They had abundant time to

contemplate the mysteries, to instruct their young in countless

legends and traditions, and to perform the endless rituals that

would please the spirits.

All Indians were dominated by spirits. Every rock, tree, bush,

wind, animal, stream, lake, storm or fire was a spirit. Christian

missionaries, recognizing no such spirits, seemed to the Indian

to be hopelessly insensitive. Only when God was described as the

Great Spirit, could spiritual communication begin.

In the dream fast, a young person would go alone for three or

four days without food, water, or shelter until in his parched

delirium a spirit would come to be guardian and confessor to him

for the rest of life. Elation and sense of great power came.

No Indian tribe north of Mexico had any kind of fermented or

distilled liquor. When they were first treated to it by the white

1974]

Steuber — Novel Tells Story of Indians

53

man, they felt the same elation, the same power, the same com¬

munion with all their spirits as their dream fast had given them.

They quickly looked upon liquor as something good, something

holy. With this extra appeal, in addition to the pleasure to the

palate and the warming glow, Indians readily became alcoholic.

Indians before the white man came had no cows, goats, or sheep.

Yet every child had milk for the first three years of his life. He

nursed many months after he had learned to run and talk, but

once he was weaned he never tasted milk again in all his life. Milk

was for babies and only for babies. When the settlers came and

brought their cows, and the Indians saw grown children and white

adults drink milk they were horror stricken. To them it was a filthy

abomination, a perversion of the worst kind. They turned their

heads in utter disbelief and disgust.

White traders took Indian wives. They were a great help to the

trader in getting him accepted among the tribes. It was a great

honor for an Indian woman to be so chosen, and some became so

proudly arrogant nobody could stand them.

But when the settlers came, as desperately as they needed

women, no settler would ever take an Indian wife because they

were no help at all.

They could not fry an egg.

They could not set a table.

They could not make a bed.

They could not milk a cow.

All primitive people become highly agitated by thunder and

lightning. Thunder quickly becomes a part of their mythology.

This was as true for the ancient Greeks and early Norsemen as

for the American Indian, whose thunderbird is traditional with

every tribe.

Even today, with all our modern sophistication, we too pay

homage to thunder and its company. We pay high premiums for

wind and hail insurance. Public utility companies and cities install

lightning protection and keep emergency crews on standby during

storms. We all keep our radios on during a tornado watch. There

still are many farm families who get everybody up in the middle

of the night to sit out a storm.

Try to imagine then, the fear that thunder and lightning could

bring to an uninformed but imaginative Indian, crouching under

bark slabs that leaked, that blew off with the wind, under an angry

heaven striking at him with all the fury of the universe. His fear

of lightning and thunder so overwhelmed him that he could find

peace only through the mystic power of his medicine man. Only

when the medicine man stripped naked and ran chanting all around

54 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

the camp were the thunderers appeased and the camp made safe

again.

Over and over again, Indian spokesmen tell us that Indians are

not interested in the white-man’s ambitions, or white economics,

or white life styles, nor even white definitions of freedom or

equality. The red man tells us that he does not want to imitate any

other race, or be like any other people. He says he wants only to

be an Indian and to be left alone so that he can be an Indian.

Certainly one must accept that as a noble and a simple request.

To be a brother to the deer and the bear, to be in accord with every

tree and babbling brook, to walk as a natural creature of the

woodland, to live in intimacy with all the spirits of the rocks and

rains and winds, to live only with what Mother Nature sets before

one, surely this is the ideal life.

This dream to go back fascinates, and is symbolic to young

idealists, to the human-rights people, the ecologists, the nature

scientists. It certainly must appeal to every thoughtful citizen who

holds dear his own freedom.

All the natural riches of wholly Indian North America — vast

beds of wild rice, maple groves bleeding rich sap, streams alive

with sturgeon and beaver, the great plains black with buffalo, the

woodland skies darkened with wild fowl, berries from spring to

fall in every marsh and upland, grapes and nuts beyond all gather¬

ing. With all this boundless treasure, Nature working alone took

more than three square miles of land to support each Indian.

If the Indians today could live primitively by themselves, it

would be the highest priced living the earth has ever known. We

would have to remove 197 million other Americans. We would

have to take down our cities, bulldoze our highways back into the

soil, abandon our farms, and return everything to the native

prairies, marshes, and forests. Only then could an Indian live his

ideal life as an Indian.

This is the dilemma the Indian creates for himself and for us.

This is why the Indian himself makes it so difficult to help him.

This is why, after four hundred years of trying we still cannot

answer the Indian question either to his satisfaction or to ours.

Nature can renew, nature can replenish, nature can go on no

matter how violently she has been insulted, but nature cannot go

backward. Only in fiction, only in make-believe, can we go back

to relive the days that once were.

GO AWAY THUNDER, a novel published in 1972, is a tale of

the Menominee Indians many generations before Columbus. It is

wholly sympathetic to the Indian viewpoint. As fiction, it can go

1974]

Steuher — Novel Tells Story of Indians

55

back, it can re-live the times when the Menominee were the giants

of the Great Lakes woodland.

Through nearly three hundred years of the fur trade, the wood¬

land Menominee homeland on the north and west shores of Lake

Michigan remained undisturbed, its culture intact. Once the Euro¬

pean demand for beaver hats declined there was no economic

reason whatsoever for a white man to stay in Menominee country.

Nothing much was asked of the Menominee until lumbermen

took notice of the finest stand of white pine the world has ever

seen on their ancestral lands. American enterprise, needing lumber

and lots of it for expanding America, easily negotiated treaties

to take the Menominee land. By that time we had already passed

the middle of the 19th century, our nation was approaching the

astonishing age of one hundred years, and we were affluent enough

to have money to put into carefully detailed studies of the people

who were here before we were.

The Bureau of American Ethnology as a part of the Smithsonian

Institution began a series of cultural studies of Indian tribes, a

series of studies continuing for many decades. The remarkable

Menominee, essentially living the same as their ancestors long

before the first white man in America, were studied and docu¬

mented, their chiefs and medicine men explaining their traditions

and values.

From these detailed ethnic studies, from the much earlier ex¬

plorers’ and traders’ notes, from the early missionary documents

it is possible to piece together a story of Menominee life when

Indians alone occupied the northern woodland. With this authentic

documentation, GO AWAY THUNDER was written. It recreates

one year of early Menominee tribal life in the long long parade

of mankind. The title, GO AWAY THUNDER expresses an awe,

a fear, a plea to the unknown, a prayer to the mysteries by the

primitive American Indian. GO AWAY THUNDER is also a

symbolic cry toward the coming of the white man, the most awe¬

some thunder of all.

This has been my work to find out about earlier Indians for a

better understanding of the Indian of today. I have now com¬

pleted, as a following novel entitled BEAVER, BRANDY, BEADS

AND BELLS, the amazing story of their first contact with white

civilization.

Later, after much more research, I intend to put into a third

Menominee novel the continuing conflict between these proud

Indians and white culture today.

I

LAKE WINGRA, 1837-1973:

A CASE HISTORY OF HUMAN IMPACT1

Paul C. Baumann, James F. Kitchell,

John J. Magnuson and Terrence B„ Kayes

University Wisconsin — Madison

ABSTRACT

Lake Wingra is a shallow, eutrophic lake bordered on one side

by the City of Madison and on the other by the University of Wis¬

consin Arboretum. This paper recounts the history of the biological

and hydrographic changes which occurred in the lake between 1837

and 1973. In the early 1900’s, the area, depth and drainage patterns

of the lake and adjoining wetlands were extensively altered by the

activities of man. Early management practices (e.g. introduction

of carp in the late 1800’s, fish rescue and stocking operations dur¬

ing the 1930’s and carp removal programs during the 1930’s, 1940’s

and 1950’s) had marked and often unexpected effects. The lake

once had extensive bordering marshes, diverse aquatic macrophyte

communities, and large northern pike and yellow perch populations.

It now has few connecting marshes, dense stands of Eurasian water

milfoil and is dominated by stunted bluegill and yellow bass. The

number of macroinvertebrate species has been reduced ; many

larger forms have been eliminated. Since 1969, Lake Wingra and

its watershed have been a major study site of the International

Biological Program. Hopefully, ecosystem concepts derived from

this project and the lessons of history will facilitate more ecolog¬

ically sound management decisions.

INTRODUCTION

During the 1960’s, the effects of cultural eutrophication on fresh

waters became increasingly apparent to scientists and the general

public (Hasler, 1973). Other planned and accidental disturbances

by man have proven highly important in producing major changes

in aquatic ecosystems. Among these, physical changes produced by

dredging, draining and damming, introduction of non-native

species, and activities associated with intensive recreational use

(e.g., rough fish removal, weed cutting, sport fishing) have sig¬

nificantly altered many aquatic systems. All have occurred at Lake

Wingra.

1 Contribution No. 141, US-IBP, Eastern Deciduous Forest Biome and journal paper

No. 86, University Wisconsin Arboretum.

57

58 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

Lake Wingra, Dane County, Wisconsin, is a shallow, hard-water

lake with a surface area of 140 hectares and a maximum depth of

6.4 meters (Poff and Threinen, 1962). A dense littoral macrophyte

community almost completely dominated by one species, Myriophyl-

lum spicatum (Nichols and Mori, 1971) , classifies the lake as highly

eutrophic according to the scale of Swindale and Curtis (1957).

“Whole-basin” mixing probably provides sufficient nutrients to sup¬

port a rich algal flora throughout the growing season. Fish and

invertebrate populations are also dense. Recreational use of the lake

is intensive.

The lake has been the site of substantial scientific investigation

because it is near the University of Wisconsin and its Arboretum.

A chronology of investigations prior to 1968 concerning Lake

Wingra is as follows: Birge (1891), Juday (1914), Cahn (1915),

Pearse (1916, 1918), Pearse and Achtenberg (1918), Pearse

(1921), Domogalla and Fred (1926), Wright (1928), Tressler

(1980), Tressler and Domogalla (1931), Leopold (1937), Juday

(1938), Frey (1940), Noland (1951), Threinen and Helm (1954),

Neess, Helm, and Threinen (1957), Helm (1958, 1964), and Sachse

(1965). Unpublished data and a variety of maps are in the files

of the University of Wisconsin Arboretum Committee and the Uni¬

versity of Wisconsin Archives. Noland (1951) summarizes much

of the earlier records of the Wisconsin Department of Natural

Resources (unpublished data in Water File).

The Lake Wingra basin is now under intensive investigation as

an Eastern Deciduous Forest Biome site of the International Bio¬

logical Program (IBP) (Adams et al., 1972). This project did not

officially start until July 1, 1970, but preliminary investigations

began in 1968. As part of the Analysis of Ecosystems Project,

studies are being conducted to “secure an understanding of our

emerging man-dominant ecosystems” and for “development of

mathematical models and simulation of complex events in the eco¬

system over extended periods of time.” One major reason for this

site’s selection is that the “Lake Wingra watershed is representa¬

tive of the situation existing in many watersheds throughout the

eastern United States for it is markedly influenced by the activities

of man.” The site “offers an opportunity to assess relative impor¬

tance of city versus natural areas,” being bordered to the north

by the city of Madison and to the south by the University of Wis¬

consin Arboretum (Anonymous, 1969).

The purpose of our paper is to collect, in one narrative, the

diverse information available on Lake Wingra. We briefly describe

the white man’s settlement of the watershed, depict the hydro-

graphic and biological history of the lake, and qualitatively evalu¬

ate changes related to human impact.

1974] Baumann , Kitchell, Magnuson, Hayes — Lake Wingra 59

SETTLEMENT

Lake Wingra and the extensive marshlands surrounding it were

used by the Winnebago Indians as a major hunting and fishing

ground as late as 1925. “Winnebago tradition had it that the

swamp would take over entirely when the last red man left the

shores” (Sachse, 1965).

In 1837 Moses Strong came to the Madison area to make surveys

for establishment of the new state capitol. At that time some 500

to 1,000 Winnebago Indians were camped around Lake Wingra.

The Winnebago liked to camp by the lake because of the “great

abundance of fish in its waters and game on its shores” (Brown,

1915). After the financial crash of 1857, Levi B. Vilas, owner of

an enormous tract north and south of Lake Wingra, sold several

parcels of land. In 1865 Frank M. Grady settled a tract southwest

of the lake, and “by the seventies a few isolated homesteads dotted

the north shore” (Sachse, 1965).

During these early years, Dead Lake (later the Winnebago name,

Wingra [Duck], was adopted) was relatively inaccessible. In 1841

only Indian trails penetrated to the lake, and an old territorial road

(now Fish Hatchery Road) skirted the southeastern marshland. In

her “Map of Lake Wingra before 1915,” Sachse shows many of

Madison’s major streets in approximately their present-day loca¬

tions, and shows trails following the north and south margins of

the marshland.

Aside from agricultural activities, major physical alterations of

the watershed did not begin until 1905, when dredge and fill opera¬

tions began in the construction of Vilas Park. Between 1911 and

1920 the Lake Forest Land Company made its abortive attempt to

build a “model suburb” in the southern lowlands. Urbanization of

the north shore followed the sporadic development of Nakoma and

westward expansion of Madison, beginning about 1910 (Sachse,

1965).

Throughout the early years, Wingra’s rich populations of musk¬

rat, waterfowl, game fish and turtles were heavily exploited. Since

then the lake, its biota and surrounding marshland have been dra¬

matically altered by the influence of man. In his 1934 reminiscences

of the 1870’s, L. B. Rowley stated the point simply: “. . . it isn’t

the same lake at all” (Sachse, 1965).

HYDROGRAPHY

From earlier scientific papers, unpublished theses, public rec¬

ords, maps and accounts of private citizens, Noland (1951) col¬

lected detailed information about the hydrographic history of the

Wingra basin. His accounts did not, however, reconstruct a picture

of the basin’s original nature.

60 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

The Original (ca. 1900) Shoreline and Wetlands

Reconstruction of Wingra’s original hydrography is difficult

because scientific investigations were not done before the period of

maximum hydrographic change, and early accounts are conflicting

and unclear. The lake’s surface has been reported to be 217 hec¬

tares (Pearse and Achtenberg, 1918), 81.0 hectares (Juday, 1938),

133 hectares (Noland, 1951), and 140 hectares (Poff and Threinen,

1962). An example of semantic confusion is the frequent but unde¬

fined use of the word “marsh.” This term describes a variety of

conditions. To avoid further confusion, we use the fresh-water

classifications described by Smith (1966).

Our attempt to reconstruct the original shoreline and surround¬

ing marshland is a stepwise alteration of present-day maps

accounting for known and inferred hydrographic changes in the

past.

Figure 1 indicates present shorelines, local geographic refer¬

ences, and other physical characteristics of the area adjacent to the

lake. The lake is 262.9 m above sea level, 1 m higher than Lake

Monona (United States Geological Survey, 1959). According to

Nichols and Mori (1971), the present spillway dam (Fig. 1) main¬

tains a 0.6 m head between lake level and Murphy’s Creek. The

creek falls the remaining distance to Lake Monona. The dam and

McCaffrey Drive isolate Gardner Marsh from the lake proper. The

channels in this wetland measure 0.3 to 0.6 m below lake level (our

observation) and drain into the creek above Fish Hatchery Road.

Except at the outlet, a strip of high ground separates marsh from

creek ( Fig. 1 ) . The northwestern two-thirds of the wetland is pri¬

marily a mixture of shallow and deep fresh marsh. The southeast¬

ern third is dominated by shrub swamp, which becomes increas¬

ingly dry toward the outer margins. The ponds and channels in

Gardner Marsh are open freshwater areas, although vegetation is

dense in some parts. Both Redwing Marsh and the marsh west of

Edgewood Bay (Fig. 1) are deep fresh marsh, but become rapidly

shallow shoreward. The western wetland, including Wingra Marsh

and Wingra Fen (Fig. 1), is a patchwork of fresh meadow and

shrub swamp. Shallow fresh marsh borders the lake’s southwest¬

ern shore and merges with a narrow bank of deep fresh marsh

toward open water. The lake’s water originates from surface run¬

off, springs and, along the urbanized shore, storm sewers and drain

pipes (Noland, 1951). Spring water enters the lake directly and

via Ho-nee-um Pond and Gorham, Nakoma, Big Spring, East

Spring and Marshland creeks (Fig. 1).

Early accounts (Birge, 1891; Cahn, 1915; Juday, 1914; Rowley,

1934 in Sachse, 1965) agree that the entire shoreline was marshy

and marsh areas were more extensive. They also suggest that the

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra

61

.5

0

JL

1 KM

J

FIGURE 1. Lake Wingra, adjacent wetlands, and major streets ( _ ) in

early 1970’s. Combined from field observations and maps by the U.S. Geo¬

logical Survey (1958) and the University of Wisconsin Arboretum Committee

(no date) .

lake had a greater area. Data collected by Noland (1951) provide

two reasonable explanations. Early water levels were higher, about

1.3 m above Lake Monona or about 0.3 m above the present level.

The outlet was located just above the bridge at Fish Hatchery

Road (Fig. 1), and upper Murphy Creek and much of Gardner

Marsh were probably part of the original lake.

A hypothetical shoreline (Fig. 2A), taking account of the above

information, was made by designating a point above Fish Hatchery

Road as the original outlet and by drawing a line midway between

the lake’s present shoreline at 262.9 m and the 263.4 m contour line

determined from a U.S. Geological Survey Map (1959). Earth-

moving operations were assumed not to have appreciably altered

the contours.

Additional information substantiates the proposed shoreline in

Fig. 2B. Both Rowley (Sachse, 1965) and a map made by Moritz

62

Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

N A

FIGURE 2 A. Pre-1900 reconstruction of Lake Wingra, its wetlands and

littoral zone. Scaled to map by U.S. Geological Survey (1959).

in 1904 (this map is currently in Wisconsin Historical Society

Archives) indicate that Gardner Marsh originally extended to

what is now Fish Hatchery Road. The road probably marks

the outer boundary of the original lake’s emerging vegeta¬

tion. Higher water must have covered the southern end of the

strip now separating Gardner Marsh from Murphy Creek. Because

land-fill operations created the peninsula at Vilas Park (Juday,

1914; Noland, 1951), the original shoreline must have been con¬

tinuous from Murphy Creek along the northern edge of the present

Vilas Park Lagoons (Figs. 1 and 2A). The higher water probably

covered much of Edgewood Point. An incomplete survey done in

1834 (Wilcox, 1936) confirms the proposed shoreline of the eastern

half of the lake. Two spillway dams maintain a 0.3 m head between

Ho-nee-um Pond (Fig. 1) and the lake (Sachse, 1965). A soils map

by Retzer (1950) suggested that the island separating pond from

lake was formed by dredge spoils, so the hypothetical shoreline was

placed along the western edge of the pond.

The area deeper than 2.0 m on the present contour map (Fig. 3)

likely represents the original pelagic zone of the lake and is indi¬

cated as such in Fig. 2A. Maximum depth was probably about 4.3

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra

63

N

B

i _ i - 1 - - - 1

I .5 0 1 KM

i _ I _ i _ i

FIGURE 2B. An early (1904) map of Lake Wingra redrawn from an original

by Moritz. Depth contours are given in feet. Streets identified are those with

same names today.

m in the area midway between what is now Edgewood Bay and the

southern shore (Fig. 2B). The remainder of the basin was largely

a gradual and regular slope to all shorelines north and south with

extensive shallow littoral zones to the east and west.

The wetland originally surrounding Lake Wingra “covered a

wide area in all directions except the adjoining high land on the

north side and a wooded knoll on the southwest about twenty acres

in extent” (Rowley (1934) in Sachse, 1965). Water from springs

in and around the margin of the wetland drained into the lake.

In the northeast, the marshland’s outer edge was probably defined

by Reynolds Spring, which was situated “just south of the old

Dividing Ridge” (Noland, 1951) and had a creek as its outlet. In

the west and southwest the wetland extended “to Monroe Street

and Nakoma Road” and took “in the flat part of what is now the

Nakoma Golf Grounds” (Fig. 1) (Rowley (1934) in Sachse, 1965).

Apparently the original wetland extended almost up to the 266.6

m contour line (Fig. 2A). The “Map of Lake Wingra Before 1915”

64 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

THE LAKE WINGRA DRAINAGE BASIN

including- approximate depth contours of the lake at 1 m intervals.

(Sachse, 1965) verified the proposed outer margin. The area from

this wetland margin to proposed shoreline was likely a mixture of

fresh meadow and shrub swamp. Cahn (1915) reported that the

Western wetland was covered with up to 30 cm of water in May

and was muddy through summer and autumn.

! Moritz, a geologist, surveyed the lake and site of Vilas Park

prior to construction. His map (1904) indicates that the original

northeastern shoreline roughly corresponded with what is now

Vilas Park Drive (Fig. 1) and that the eastern region of the lake

was a broad, shallow area extending to Fish Hatchery Road. We

located Moritz’s map after having reconstructed the lake based on

contour lines for Fig. 2A. In general, the margins of the littoral

zone area proposed in our reconstruction are confirmed by Moritz’s

map suggesting that the area of the lake was approximately 265

hectares. The current area of the lake is 137 ha. (see Huff et al.,

1973, for a detailed summary of present physical characteristics).

Thus, prior to major construction activity, the area of Lake Wingra

was approximately 1.9 times greater. Including the surrounding

marsh and wetlands, the area was about 3 times greater than that

at present.

1974] Baumann, Kitchell, Magnus on, Hayes — Lake Wingra 65

Major Changes to Shoreline and Wetlands

The assumption made in Fig. 2A, that the 263.4 m contour line

has not changed appreciably, is probably correct. But some changes

have occurred and are documented below, primarily from Noland

(1951). The east and southeast end of the original lake and wet¬

land was most altered. This area now includes the lowland east of

Murphy Creek, the full length of the creek, Gardner Marsh, the

sections east of the marsh to Park Street and south beyond Carver

Street as well as the south and east sides of the lake proper (Figs.

1 and 2A).

In 1905 and 1906 (Fig. 1, present streets as landmarks), a chan¬

nel was dredged from the bridge of Fish Hatchery Road in an arc

along Wingra Drive to a point south of Randall Avenue. A wooden

lock and spillway dam were erected below the bridge to maintain

the lake at its original level. In 1907 and 1908 the creek’s entire

length below the bridge to Lake Monona was dredged. Though

documentation was not found, spoils from these operations were

probably used to fill adjacent wetlands.

Between 1914 and 1920 the Lake Forest Land Company

attempted to build a “model suburb” in the southern wetland

beyond the 263.4 m contour line. At this time, a dike (McCaffrey

Drive dike) must have been built from the south shore and south¬

east to the outlet to isolate the present Gardner Marsh from Mur¬

phy Creek and the lake proper (Fig. 1). Channels, dug in the

marsh and wetland to the south, emptied into the creek below the

old spillway dam. This lowered the water level of the marsh below

the lake level and diverted water from four southern springs away

from the lake. In 1917 the dike was breached at the southeast cor¬

ner of the lake, effecting a 1.0 m fall in the lake’s level and a dra¬

matic reduction of surface area. The present lock and spillway

dam (Fig. 1) were built and the dike was closed in 1919. The lake

filled to its current level about 1.0 m above that of Lake Monona.

Murphy Creek was then dredged to levels approaching Lake

Monona’s to further facilitate drainage of marsh and wetland.

Beginning in 1915, a dredge working 12 hours a day, spring

through fall (Sachse, 1965), removed bottom materials from the

lake near its south shore. By autumn of 1916, a trench 9.3 m deep

had been dug. This has partly filled to the present depth of 6.4 m

(Fig. 3) which is currently the maximum depth in the lake (Poff

and Threinen, 1962). The areas above and below the present dam

were dredged, producing Outlet Bay and Murphy Creek Wide¬

spread. Spoils may have been used to close and strengthen McCaf¬

frey Drive dike.

In spite of all the above effort, little solid land was produced.

The Lake Forest Land Company was bankrupt in 1922.

66 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Development of the University of Wisconsin Arboretum and

urbanization have affected the east and southeast end of the orig¬

inal basin. A soils map by Retzer (1950) indicated extensive filling

east of Gardner Marsh between Fish Hatchery Road and Park

Street (Fig. 1). Construction of Wingra Drive and McCaffrey

Drive raised shores along their routes. In 1936, channels in Gard¬

ner Marsh were redredged (Sachse, 1965), but since then, sedimen¬

tation has partly filled them.

The northeast end of the original basin was greatly altered by

the construction of Vilas Park in 1905, 1906 and 1913 (Noland,

1951). The area now in the outer part of Edgewood Bay (Fig. 1)

was dredged to over 4.0 m. Marsh and wetland north of the present

Vilas Park Lagoons were filled and the shore raised. Fill was also

used to construct the peninsular part of the Park. Vilas Park

Lagoons were dredged to 1.6 m in the center. They were redredged

in 1941, and the sediments deposited on an island in the center.

Removal of groundwater by new wells combined with dredging

and filling destroyed five springs in the original marsh. The depth

in Edgewood Bay rapidly decreased as materials eroded from

shore. The outer part of the bay is now about 1.2 m (Fig. 1).

The west end of the original basin was altered when Ho-nee-um

Pond was built in 1938 and 1939 (Fig. 1). Material dredged from

the pond site was used to form an island between pond and lake.

Spillway dams were constructed at each end of the island, and the

pond’s water level was established 0.3 m above the lake. Dredged

material was also deposited west of the pond and along the lake’s

shore north of the pond (Sachse, 1965).

Along the north shore, 16 springs have ceased to flow. This

probably resulted from reduced replenishment of ground water

owing to the construction of buildings, streets and storm sewers.

Runoff from the northern and western regions of the drainage

basin now reaches the lake primarily by the storm sewers (Fig. 3).

Current land use patterns have been summarized by Cullen and

Huff (1972). Excluding the lake proper, more than 20% of the

watershed has been altered to the extent that it is impervious to

water. Buildings occupy 6.3% ; roads, 8.6% ; drives, 2.3% ; parking

lots, 1.5% ; and sidewalks, 1.0%. Natural forests, primarily in the

University Arboretum, comprise 16.0%, while lawns, parks, ceme¬

teries and golf courses make up the remaining 63.6%. Although

the latter are collectively termed “natural” land areas, they reflect

the impact of human use. Selection for fast growing grasses, orna¬

mentals and trees has resulted in a 30% greater productivity in

comparison to equal-aged forests within the Arboretum (Lawson,

Cottam and Loucks, 1972). Further, landscaping of lawns, parks

and gardens and applications of fertilizers ultimately increase the

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 67

nutrient load of runoff waters entering storm sewer systems

(Huddleston, 1972).

Summary

In about 1900 Lake Wingra had a shallower maximum depth

and covered about twice the area that it now does. Surrounding

marsh was more extensive which, in combination with the original

lake, provided a wetland and lake area nearly three times greater

than that at present. These changes have been directly caused by

dredging, draining and construction of dams and dikes. Other

hydrographic effects of man on the lake were induced by altera¬

tions of soils, vegetation, ground water sources and surface water

runoff patterns.

AQUATIC MACROPHYTES

The Original Vegetation

Wetlands of various types surrounded Lake Wingra in the late

1800’s. Tamaracks were abundant around the southern shore until

1870. The soil of the East Marsh is composed of peat, from which

tamarack branches, logs, and cones have been recovered. Although

no good descriptive accounts from this early period exist, the

assumption can be made that the bog flora usually associated with

tamaracks was probably also present (Irwin, 1973).

Broad margins of marsh still surrounded Lake Wingra around

1900. Horsetails (Equisetum) , sedges (Carex) and wild rice

(Zizania aquatica) were probably included in the wide margin of

“weeds” described by Rowley (Sachse, 1965) along with cattails

(Typha latifolia and T. angustif olia) and a bulrush (Scirpus

validus), which are now the dominant emergent forms (Nichols

and Mori, 1971). Dense growths of Chara were interspersed be¬

tween areas of wild rice and reeds (Birge, 1891). Wild celery

(Vallisneria americana) was particularly abundant and received

special note in Rowley’s accounts (Sachse, 1965). Almost the entire

lake bottom was covered with water plants of various kinds (Birge,

1891). Accounts after 1900 agree that the lake’s entire shoreline

was “marshy,” that submerged macrophyte beds were extensive,

and that wild rice was profuse (Cahn, 1915; Juday, 1915; Leopold,

1937; and Rowley (1934) in Sachse, 1965).

Other native Wisconsin species currently found in Lake Wingra

that were likely present when the first white settlers arrived in¬

clude four species of small floating plants (Lemna minor, L. tri-

sulca, Spirodela polyrhiza and Wolffia columbiana) , two species of

water lilies (Nuphar variegatum and Nymphaea tuberosa) and a

group of submerged macrophytes including : Potamogeton pectina -

tus, P . nodosus, P. richardsonii, P. zoster if or mis, P. foliosus, P.

1

68 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

\

i . • .

natans, Zanichellia palus tr is, Najas flexilis , Anacharis canadensis,

Heter anther a dubia, C eratophyllum demersum, Utricularia vul¬

garis and Ranunculus longirostris (Nichols and Mori, 1971).

Type, abundance and distribution of aquatic macrophytes have

changed dramatically over the past century. Many aspects of these

changes can be attributed to man.

Major Changes in the Vegetation

In the southeast regions of the original lake, changes in littoral

vegetation were related to dredging and filling. The net result was

the transformation of a wide littoral area, probably rich with wild

cfelery, coontail (C eratophyllum demersum) , pondweeds (Potamoge-

ton spp.), water lilies, wild rice, bulrushes, cattails and sedges into

Gardner Marsh, which is dominated by cattails. Channels in this

marsh and Murphy Creek now contain dense growths of Eurasian

water milfoil ( Myriophyllum spicatum) , though water lilies are

still common.

In the northeast, construction of Vilas Park by dredging and

filling reduced the broad margin of weeds and cattails described by

Rowley (Sachse, 1965). A sand beach was built on the peninsula

and is still maintained. Mechanical harvesting and chemical treat-

meht are used to prevent growth of aquatic plants around Vilas

Park and the outlet to Murphy Creek (Nichols and Mori, 1971).

Considering the entire lake, changes in water level had particu¬

larly disturbing effects on aquatic vegetation. In 1917 the level

was dropped 1.0 m; in 1919 it was raised 0.7 m to its present

height, 1.0 m above Lake Monona. Fluctuation of water level and

overwinter drawdowns are commonly used to inhibit growth of

aquatic plants (Black, 1968; Beard, 1969). The net reduction of

water level by 0.3 m also substantially reduced the lake’s littoral

zone.

In addition to hydrographic changes, introduction of exotic

species altered the vegetation in the lake. The carp (Cyprinus

carpio) was introduced into waters connected to Lake Wingra in

the late 1800’s, was common by 1915, and was a dominant by 1930

(see FISHES of this paper). Carp destroy vegetation directly by

uprooting plants, and indirectly by increasing turbidity and silta-

tion (Cahn, 1929; Cole, 1904; Cahoon, 1953; Black, 1946; and

Threinen and Helm, 1954). Carp have an especially deleterious

effect on wild celery and wild rice (Cole, 1904). Records of the

Wisconsin Department of Natural Resources (unpublished data

in the Water File) indicate that Lake Wingra was nearly denuded

of aquatic macrophytes during the period of carp dominance from

the late 1920’s through 1955. From 1936 through 1955, an inter¬

mittent carp removal program was conducted by the Wisconsin

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 69

Conservation Department with the cooperation of the University

of Wisconsin (reviewed by Helm, 1958, and more fully described

in FISHES of this paper) . By 1956, biomass and species diversity

of aquatic plants, especially water milfoil, coontail and pond weeds

had increased (Helm, 1958). Carp removal and subsequent macro¬

phyte growth have not reduced turbidity because dense phytoplank¬

ton populations have characterized the lake since 1956 (Helm,

1958; Koonce, 1972).

Owing to hydrographic disturbances, introduction of exotics, or

a combination of factors, several native species disappeared from

Lake Wingra. Wild celery, Potamogeton freisii, P. illinoinensis,

P. amplifolius and P. praelongus have not been collected later than

1929 (Nichols and Mori, 1971). The decline of wild rice has not

been documented, but it is no longer present.

The Eurasian water milfoil now dominates the macroflora of all

Madison lakes, including Lake Wingra. It was introduced to the

Chesapeake Bay region of North America around 1900 and re¬

ported in Wisconsin by 1936 (Nichols and Mori, 1971). Although

its appearance and increased abundance in Lake Wingra are not

documented, Helm (personal communication) recalls that milfoil

was not the dominant macrophyte in the late 1950’s. Rather, pond-

weeds were most abundant after carp removal.

The Present

Gardner (or East) Marsh vegetation at present is composed of

a variety of terrestrial and aquatic plants with shrubs invading

the open areas and trees located along the edge. Seven major com¬

munities exist, including tree, shrub, wet meadow, aster-solidago,

calamagrostis, cattail, and nettles. The large nettle community

thrive on burnt peat soil, and was apparently established after

the original marsh had been dried and destroyed by fire (Irwin,

1973).

Presently, five littoral communities occur in Lake Wingra as fol¬

lows (Nichols and Mori, 1971) : shallow water Myriophyllum,

68% ; deep water Myriophyllum, 13% ,* Potamogeton-Myriophyllum,

17%; Nuphar, 5%; Nymphaea, 2%. Dominant emergents are

Typha latifolia, T. angustifolia and Scirpus validus. The littoral

zone extends to 2.7 ± 0.4 m, a depth primarily dictated by light

penetration, and covers almost one-third of the lake’s surface area.

Since water milfoil can grow from depths greater than 4 m, a ma¬

jor reduction in turbidity might allow stands to develop throughout

the lake. Native pondweeds and coontail are largely limited to

depths less than 2 m.

The dense growths of littoral vegetation have considerable in-

70 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

fluence on the lake. They provide shelter from predation for many

juvenile fishes, thereby increasing survival and population densities

of panfish species in particular (Andrews and Hasler, 1942). Storm

sewer waters are literally “filtered” by littoral vegetation. The

lake is thus buffered from rainfall-induced pulses of allochthonous

nutrients (MaeCormick et ah, 1972). Finally, dense macrophytes

provide substrate, cover and food for many aquatic invertebrates

which can ultimately be associated with distributional dynamics

of fishes (Baumann and Kitchell, 1974).

In summary, many of the present physical and biological condi¬

tions of Lake Wingra are directly related to changes in the aquatic

macrophyte component since the carp removal program.

PHYTOPLANKTON AND NUTRIENTS

The status of phytoplankton communities and nutrients are not

available for Lake Wingra around 1900, but dominant forms listed

for 1928 by Tressler (1930) generally correspond to present ob¬

servations (Koonce, 1972). Thus, changes in the assemblages of

phytoplankton do not appear to have accompanied the dramatic

changes in the macrophyte associations. Differences in analytical

methods prevent quantitative comparisons of phosphorus concen¬

trations reported by Tressler and Domogalla (1931) with those

determined by Koonce (1972) or Kluesner (1972). It appears, how¬

ever, that phosphorus concentrations in open water have been

reduced over recent time rather than increased. In addition, data

summarized by Isirimah (1972) suggest that nitrogen concentra¬

tions have not markedly increased since 1928 (Table 1).

In spite of these negative data, the trophic status of Lake Wingra

undoubtedly has been influenced by man’s activities over the past

century. The present storm sewer systems provide 80-90% of the

total annual phosphorus loading in direct runoff to the lake (Klues¬

ner, 1972) rather than through flow exposed to retention-by-soil

processes. Allochthonous particulate carbon input (leaves, twigs,

etc.) are similarly diverted from terrestrial decomposition. For

example, more than 1 metric ton (dry weight) per year comes from

the Manitou Way drainage, which represents less than 10% of the

total storm sewer system (Gasith, et ah, 1972; Gasith and Hasler,

1973). As mentioned before, vertical mixing in this shallow lake

probably maintains sufficient nutrients in the water column to

support high algal production throughout most of the growing

season. With little or no direct supporting data to the contrary,

we can only speculate that high productivity also characterized

Lake Wingra prior to human perturbations of the drainage basin.

The historic use of the area as a major hunting and fishing ground

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 71

TABLE 1. COMPARISON OF NITROGEN CONCENTRATIONS IN LAKE

WINGRA BASED ON ISIRIMAH (1972, TABLE 1). DATA FOR

1928-29 ARE FROM TRESSLER AND DOMOGALLA (1931),

FOR 1960 FROM CLESCERI (1961), AND FOR 1970

FROM KLUESNER (1972)

(Sachse, 1965) may lend some credence to this idea. Yet, the pres¬

ent algal productivity (Koonce, 1972) seems too high for an un¬

modified lake with a small forested watershed and, in 1928, when

the first studies were done, the lake had already undergone tre¬

mendous man-induced alterations.

INVERTEBRATES

Prior to 1900, Birge (1891) provided a list of cladoceran zoo¬

plankton found in Lake Wingra that included 48 species (Table 2).

This diverse fauna included a number of large open water species

such as Daphnia pulex and D. SchocUeri. Although substantial

changes in the taxonomy of cladocera have been made since Birge’s

time, the list can be validly compared with the lake’s present

cladoceran fauna.

Data on benthic invertebrates are available only as early as 1929,

when Tressler (1930) was active on the lake. The sub-littoral

benthos was represented by a diverse assemblage of macro¬

invertebrates that included aquatic insects, water mites and mol-

lusks. The littoral benthos was diverse also, but was dominated by

the amphipod Hyalella azteca which constituted nearly 90% of

the macro-invertebrates.

Dramatic changes have occurred in the communities of clado¬

ceran zooplankton and in the benthic invertebrates.

Only 23, or one half, of the cladocerans present in 1891 are still

present in lists (Table 2) prepared by Teraguchi (1970) and White

and Hasler (1972). Those no longer present include several

72 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 2. CLADOCERAN SPECIES RECORDED FROM LAKE WINGRA

BY BIRGE (1891) AND MORE RECENT STUDIES

(HASLER AND WHITE, 1971)

Daphnia spp. and Ceriodaphnia spp. that are relatively large or¬

ganisms. Only 4 new species have been added to the list.

Intensive sampling in 1970-1972 of the benthos and the fauna

on littoral macrophytes revealed an invertebrate fauna dominated

by small chironomids. No live mollusks, no relatively large aquatic

insects, and only two individuals of Hyalella were noted among the

tens of thousands of invertebrates identified by Peterson and

Hilsenhoff (1972). Perhaps more importantly, Hyalella rarely

has been discovered in the hundreds of fish stomachs analyzed to

date from 1970-1971 samples (Magnuson and Kitchell, 1971; Bau¬

mann and Kitchell, 1974), whereas they were common in fish

stomachs from 1954-1956 (Helm, 1958).

Few data are available that allow us to determine the time course

of the above changes during the last 80 years or to .relate those

changes to known causes. However, intense fish predation on larger

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 73

invertebrates may well explain a rather recent decline of larger

cladocerans and benthos in Lake Wingra. Dense panfish populations

produced virtual or real extinctions of similar species from ponds

through size-selective predation (Hall et ah, 1970). Hyalella is still

abundant in the littoral zones dominated by Eurasian water milfoil

in neighboring Lakes Mendota and Monona and remains as a domi¬

nant food of many fishes (El Shamy, 1973; Brauer et ah, 1972).

Our experience suggests that panfish populations in the latter two

lakes are not as dense as in Lake Wingra.

Intensive fish predation on aquatic invertebrates is the best docu¬

mented but not necessarily the sole explanation for the decrease

in diversity of cladocerans and functional extinction of Hyalella in

Lake Wingra. We speculate that these dramatic changes occurred

since the mid-1950’s.

FISHES

The following represents a journey through time, from the Indian

fishing grounds of the 1800’s to the children’s panfish lake of 1973.

The fish community will be examined from early records (1837-

1904), the period of major hydrographic change (1905-1925),

a period of biological adjustment (1926-1936), a period of new

species introductions (1937-1949), a period of major carp re¬

movals (1950-1957), and a second period of biological adjustment

(1958-1973). When possible, explanations for changes in com¬

munities or populations will be attempted.

As a graphic summary of the dynamics, we estimated the timing

of introductions and extinctions of major fish species (Fig. 4).

In addition, we reconstructed their relative abundances on a similar

time scale and related changes in abundance to major perturbations

by man (Fig. 5) . Information for these reconstructions was derived

primarily from surveys by biologists from the University of

Wisconsin — Madison and carp seinings by the Wisconsin Conser¬

vation Department.

Period of Early Records (18S7-190U)

Fish were referred to as being plentiful in the early years. Our

interpretation of the species list mentioned by Rowley (1934) is:

longnose gar (Lepososteus osseus), northern pike (Esox Indus),

black bass (Micropterus salmoides) , sunfish ( Lepomis spp.) , crap-

pies (Pomoxis spp.) and “yellow bass,” a local term at that time

for the smallmouth bass (Micropterus dolomieu). Pickerel were

also mentioned, but the term seemed to be used interchangeably

with northern pike. Earliest scientific investigations did not find

any true pickerel (Esox niger or E. americanus).

74 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

-ARGE pisgivores

BOWFIN

LONGNOSE GAR

NORTHERN PIKE

LARGEMOUTH BASS

SMALLMOUTH BASS

WALLEYE

N. PIKE x MUSKELLUNGE

BROWN TROUT

RAINBOW TROUT

BROOK TROUT

CHANNEL CATFISH

MEDIUM PREDATORS

YELLOW PERCH

BLUEGILL

PUMPKINSEED

GREEN SUNFISH

BLACK CRAPPIE

WHITE CRAPPIE

BROWN BULLHEAD

YELLOW BULLHEAD

BLACK BULLHEAD

WHITE BASS

YELLOW BASS

SMALL PREDATORS

BLUNTNOSE MINNOW

GOLDEN SHINER

BROOK SILVERSIDE

OMNIVORES

CARP

BUFFALO FISH

COMMON WHITE SUCKER

NORTHERN REDHORSE

SPOTTED SUCKER

970

INTRODUCED SPECIES

KNOWN PRESENT

BELIEVED PRESENT

FIGURE 4. The fish community of Lake Wingra from 1890 to 1973,

reconstructed from the literature showing presence and absence.

A biological study by Marshall and Gilbert (1905) in 1902-1903

mentioned four fishes from Lake Wingra: black crappie (Pomoxis

nigromaculatus) , bluegill (Lepomis macrochirus) , largemouth

bass (Micropterus salmoides ) , and yellow perch (Perea flavescens).

Undoubtedly other species were native, but these conclude the docu¬

mented list of fishes from early records (Fig. 4). Smaller panfish

and forage species were seldom referred to individually by name.

These species will be discussed in the next section.

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 75

FIGURE 5. Relative abundance of major fishes in Lake Wingra from 1890

to 1973, reconstructed from the literature.

76 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

In the late 1800’s, northern pike and bass were large in size and

sufficiently abundant to support a recreational fishery and a food

fishery for the early settlers as well as the Winnebago (Brown,

1984; Sachse, 1965).

The first perturbation by man was the introduction of carp

(Cyprinus carpio), an exotic from Europe. Between 1885 and 1897,

3,947 were released into the Yahara River system (Frey, 1940).

Carp were first noted in Lake Wingra by Dr. Samuel H. Chase

during the late 1890’s, but they were not common then (Leopold,

1937).

Period of Major Hydrographic Changes (1905-1925)

Limnological investigations by A. S. Pearse and his associates

and by A. R. Cahn occurred between 1914 and 1916, in the midst

of the most radical hydrographic manipulations to the lake and

almost 20 years after carp were first present. They viewed the fish

community as similar to that of the late 1800’s, but one whose

characteristics were already starting to change. Several new

species, introduced to the lake in the early 1900’s, also contributed

to these changes.

A diverse community of fish predators inhabited the lake

(Fig. 4). They included northern pike, smallmouth bass, large-

mouth bass, longnose gar, and bowfin (Amia calva) (Pearse and

Achtenberg, 1918). All were probably native.

In addition, walleye (Stizostedion vitreum) were stocked

(Fig. 4) in 1900-1903, 1905-1909, 1912, 1916, 1921, and 1922.

Noland (1951) assumed that walleye were native to the lake, but

Helm (1958) argued that they were not because classical spawning

sites of wave-washed gravel were lacking. Preigel’s (1970) obser¬

vation of walleye spawning in flowing deep marshes suggests that

alternative spawning sites were present in early Lake Wingra.

Since 2,655,000 walleye had been stocked eight years prior to any

species other than carp, and since they were not mentioned prior

to 1900 even though they were a favored game fish, and since they

were not caught in gill nets set by Pearse and Achtenberg (1918),

we conclude that they were not native to the lake.

The most abundant predator (Fig. 5) was northern pike, which

was twice as frequent in gill nets as the basses (Pearse and Ach¬

tenberg, 1918). Although northern pike are more vulnerable to gill

nets than bass, early accounts of Lake Wingra’s fishery seem to

agree that northern pike had the larger populations. Largemouth

bass were only slightly more numerous than smallmouth bass.

Longnose gar were quite abundant in 1914 (Brown, 1915).

In 1922, the Wisconsin Conservation Department recorded that

3,000,000 “pickerel” fry had been stocked in Lake Wingra (Lake

1974] Baumann , Kitchell, Magnuson, Hayes — Lake Wingra 77

Wingra file, 1972). They were most likely northern pike as inter¬

preted by Noland (1951) ; none of the later surveys have recorded

true pickerel present in the lake.

Medium predators on invertebrates and fish (Fig. 4) included

the yellow perch, black crappie, bluegill and pumpkinseed (Lepomis

gibhosus) (Pearse and Achtenberg, 1918). All were probably

native. Green sunfish (Lepomis cyanellus) were not mentioned by

Pearse and Achtenberg (1918) nor by Cahn (1915), but were

found in the 1950’s and are often common to this type of habitat.

We speculate that they were native but not common.

Yellow perch was the most common fish in the lake (Fig. 5).

Pearse and Achtenberg (1918) caught with gill nets 14 times more

yellow perch than the next most common species. They were small,

mean length ca. 15 cm, which may explain lack of their mention

in early fishing records. Interestingly, fingerling perch were

stocked in 1915 and 1917 despite their abundance and apparent

poor growth.

Pearse (1918) believed black crappie were more abundant than

bluegill and pumpkinseed, based on inshore dipnet catches, but

bluegill were slightly more frequent in his gillnets (Fig. 5).

The brown bullhead, Ictalurus nebulosus, was abundant (Cahn,

1915; Pearse and Achtenberg, 1918). While Noland (1951) be¬

lieved these were misidentified yellow bullheads (Ictalurus natalis),

this seems unlikely because Pearse was a limnology professor and

a professional fisheries expert employed by the Bureau of Commer¬

cial Fisheries. The brown bullhead was likely a native.

Small predators on invertebrates (Fig. 4) included golden

shiners (Notemigonus crysoleucas) , abundant in the gillnets set in

the lake (Pearse and Achtenberg, 1915) ; and, in the springs,

spring pools and streams, the blackchin shiner (Notropis hete -

radon) and blacknose shiner (Notropis cayuga) (Cahn, 1915) ; the

bluntnose minnow (Pimephales notatus) (Pearse, 1918) ; the cen¬

tral mudminnow (Umbra limi) (Cahn, 1915; Pearse, 1915); the

banded killifish (Eundulus diaphanus) (Cahn, 1915; Pearse, 1916) ;

the brook stickleback (Galea inconstans) (Cahn, 1915; Pearse,

1918) ; and the johnny darter (Etheostoma nigrum). While most

of these were probably native, some may have been introduced by

fishermen using them as bait.

The introduced carp had by that time become common (Cahn,

1915; Pearse and Achtenberg, 1918).

Period of Biological Adjustment (1926-1936)

Other than by a growing sport fishery, Lake Wingra was little

altered by new human influences from 1926-1939. Instead, the fish

community was adjusting to earlier hydrographic manipulations

78 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

and introductions of walleye and carp. Carp rose to dominance by

the 1980’s and a state supported program of carp removal began.

In 1936 the Wisconsin Conservation Department made two sein-

ings in Lake Wingra to remove carp and other rough fish. The

seinings also provided a valuable record of changes occurring be¬

tween University studies in the 1910’s and in the 1950’s. The large-

mesh seines only provided estimates of larger fishes, and even these

estimates must be inaccurate owing to the technical difficulties

involved in a large seine haul. Although the same fish predators

inhabited the lake as during the early 1900’s (Fig. 4), a marked

decline in abundance of northern pike had occurred (Fig. 5). Based

on the two seine hauls, longnose gar was the most abundant preda¬

tor (2,500 caught) ; black bass (1,100 caught), and walleye (1,000

caught) were also abundant. As before, largemouth bass were

slightly more abundant than smallmouth bass. Bowfin and north¬

ern pike were uncommon, with only 11 northern pike captured

(Noland, 1951).

The decline of northern pike (Fig. 5) was most likely a direct

result of reproductive failures owing to the hydrographic changes

of the 1910’s which had all but eliminated spawning marshes con¬

nected to the lake (Figs. 1 and 2A). Northern pike spawn in early

spring in marsh areas, usually only 30-60 cm deep. The increase in

walleye, a potential competitor, and the decline in yellow perch, a

favored food, may also have contributed to the decline. The abun¬

dance of walleye was probably a result of stocking 2,466,000 fry

between 1928 and 1930. Approximately 10,000 black bass finger-

lings had also been stocked in 1930.

Longnose gar and bowfin were in 1936 considered “rough fish”

and those caught in the seines were not returned to the lake. Long¬

nose gar are primitive fish with low reproductive potential and

would be significantly reduced by seining. Thus seining contributed

further to a decline of fish predators in Lake Wingra.

A major change in abundance among the medium predators on

invertebrates and fish (Fig. 4) was also detected with the seine

hauls. Crappies (Pomoxis) — 40,000 caught, sunfish (Lepomis) —

20,000 caught, and a new species, the white bass (Morone chrysops)

— 1,500 caught, were most common and bluegill was the primary

sunfish (Juday, 1938). White bass was an introduced species with

900 stocked in 1917 and 1000 in 1933. The yellow perch, which was

the dominant fish species of the early 1900’s, was no longer even

common. Another new species, the bigmouth buffalofish (Ictiobus

cyprinellus) , was caught for the first time. Juday (1938) reported

300 from the hauls. Noland (1951) believed bigmouth buffalofish

were native. Pearse did not mention them in his rather thorough

study, but they might have been present because the marsh area

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 79

between Lakes Wingra and Monona would appear to be a good site

for spawning. On the other hand, they may have immigrated to

the lake in 1917-1919 when the southeast dike was open to Lake

Monona.

The introduced carp, the stimulus for all the seining, was abun¬

dant (Fig. 5) and 6,000 were caught and removed. By weight,

their 20,000 kg exceeded the weight of all other species caught.

Period of Intense Species Introductions (1937-1949)

By the 1920’s, three new species had been intentionally intro¬

duced: the carp in the 1880’s, the walleye, first in 1900, and the

white bass, first in 1917. Between 1935 and 1945, the Wisconsin

Conservation Department stocked 20 to 23 different species of fish

depending on what species the categories of sunfish and bullhead

contained. Some of these were new to the lake. Even three cold

water species of trout, totally unsuited to Lake Wingra, were

stocked from 1934 until 1941 (Fig. 4). The total array of fishes for

stocking came both from hatcheries and from fish rescue

operations.

Fish rescue, intensely practiced, was unique to the upper Mis¬

sissippi drainage basin in the 1930’s and 1940’s. After periodic

floodings of the Mississippi River, large number of fish were often

isolated from the river and trapped in temporary ponds when the

river waters receded. These fish were “rescued” by federal authori¬

ties, held in hatcheries and later stocked. Mr. Charles Lloyd of the

Wisconsin Conservation Department, who was interviewed by No¬

land (1951), stated that fish species other than those actually

recorded had most likely also been stocked. Also, prior to 1941,

federal fish rescue and transfer operations were carried out inde¬

pendently of the state, and private individuals or organizations

could apply for a shipment of fish and stock it wherever they

wished (Noland, 1951). Records of introduction would not exist.

The continuation of the removal program in 1944, 1945 and

1949, and fyke and gill nettings by Noland and Neess from 1944-

1947 provide fairly good indication of the composition of the fish

community in the mid-1940’s. All seining data are from Noland

(1951). He originally obtained them from the Wisconsin Conser¬

vation Department’s records. Carp and bowfin were removed from

the 1944, 1945, and 1949 seinings. After 1945, longnose gar were

returned to the lake if caught.

The fish predator community (Fig. 4) still contained longnose

gar, largemouth bass, northern pike, walleye, and bowfin, but the

smallmouth bass was rare and a new organism, the northern pike-

muskellunge hybrid (E. Indus x E. muskelunge) , was present in

low numbers.

80 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

The relative abundance (Fig. 5) can be judged from 1944 and

1945 seine hauls. The 2-year combined hauls yielded 1154 longnose

gar, 297 black bass (probably largemouth), 170 northern pike and

162 walleye.

Overall, fish predators seemed less abundant than in 1986. In

addition, longnose gar were declining rapidly, as only 154 were

taken during the 1945 haul. Thereafter, they were no longer

removed in seining operations. Both walleye and bass populations

had dropped since 1936, even though walleye had been stocked in

1940 and 1943 (7,000,000 fry being stocked in 1943 year alone),

and largemouth bass had been stocked in every year from 1937 to

1944. Largemouth bass declined again through 1949 (Helm, 1958).

Perhaps the large numbers of walleye fry stocked in 1943 were too

small to be abundant in the seine hauls one and two years later,

but they were also uncommon in the 1949 seine hauls. The north¬

ern pike were more abundant than in 1936 and were probably the

result of the stocking that occurred in 1940, 1941 and 1942, as indi¬

cated by their average weight of 1.5 kg. Since no further stocking

was conducted, northern pike, without significant spawning areas,

started to decline again in the late 1940’s. The northern pike-

muskellunge hybrids were stocked in 1940, 1945, 1946 and 1948.

They did add to the sport fishermen’s catch and were caught in low

but steady numbers during the seine hauls. The collapse of the

smallmouth bass population has not been explained, but might have

been related to the expanding populations of medium predators on

invertebrates and fish. Noland (1951) suggested from his observa¬

tions that illegal removal of bass from above the dam during the

spawning season may have contributed to the decline.

The community of medium predators on invertebrates and fish

increased during the late 1930’s and the 1940’s. In addition to black

crappie, bluegill, pumpkinseed, white bass, and yellow perch, two

new species were noted — the yellow bass (M or one mississippiensis)

and the white crappie (Pomoxis annularis) (Fig. 4).

Marked changes occurred in the relative and absolute abun¬

dances of the above species (Fig. 5). As a group, they were more

abundant than in 1936. The magnitude of the catch during 1944

was much greater than expected by the Conservation Department

(Noland, 1951). Dr. Black had intended to have all individuals

counted, but the haul took two days to empty and during the night

large groups of fishes other than carp were allowed over the net.

Regardless, Dr. Black estimated that in 1944, the following adult

fish were caught: 350,000 white crappie, 100,000 black crappie,

50,000 bluegill, 40,000 yellow bass, and 10,000 white bass. In addi¬

tion, pumpkinseed had a relatively large population and yellow

perch were still present, but low in abundance.

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 81

The high numbers of white crappie and yellow bass in the catch

were especially surprising, since they had not been reported before.

Based on records from Buffalo Lake reported by Helm (1958), yel¬

low bass are favored by the water quality and vegetation changes

that go along with high carp abundance. Perhaps some of the crap-

pies caught in the 1936 carp seine hauls were white rather than

black crappie. Regardless, white crappie reached their large popu¬

lation level (and greatest abundance) within approximately ten

years, and this entire group of seven fishes, excluding yellow perch,

had become abundant. Even so, this period was characterized by

intensive stocking of some abundant species and a continually

declining fish predator population.

Many fish of these seven species were stocked intentionally or

unintentionally during fish rescue operations — crappies in 1940,

1941 and 1943, bluegill in 1939, 1940, 1941, 1943 and 1944, white

bass in 1940 and 1943, and yellow perch in 1938, 1939 and 1940.

White crappie and yellow bass were not recorded specifically in the

stockings, but crappies stocked were not noted to species.

In addition to the native brown bullhead, two species in the cat¬

fish family were noted (Noland, 1951) for the first time : the yellow

bullhead, and the channel catfish (Ictalurus punctatutus) (Fig. 4).

Unidentified bullheads were stocked in 1930, 1939, 1942, 1943 and

1945, totaling 40,000 fingerlings, 10,000 yearlings and 2,600 adults.

The bullheads were present in moderate numbers in the seine

hauls and most were yellow rather than the brown bullhead (Fig.

4) . Channel catfish never flourished (only one to three were caught

per seine haul during the 1940's).

Three other new species common in the Mississippi River were

first found in the 1944 and 1945 seine and fyke net studies: the

common white sucker (Catostomus commersonii) , the spotted

sucker (Minytrema melanops) , and the northern redhorse (Mox-

ostoma macro lepido turn) (Fig. 4). Six adult suckers were recorded

as stocked in 1940. Common white suckers apparently reproduced :

75 were caught in 1944 and 120 in 1945. Thereafter they declined.

Northern redhorse persisted in low numbers (one to three per seine

haul) for a number of years, but only one spotted sucker was

caught in a single seine haul.

Bigmouth buffalofish were caught, but only in low numbers (Fig.

4). Carp remained abundant (Figs. 4 and 5), with about 3,000

caught in the 1949 haul.

Entering the 1950's, the lake contained at least nine species not

present prior to the 1880’s. These species included not only the

abundant carp, but also the most abundant and fourth most abun¬

dant zooplankton and macro-invertebrate feeders — the white crap¬

pie and yellow bass. In addition, the black crappies and bluegill

82 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

had increased in numbers. Fish predators were declining, with the

primary predator — the northern pike — apparently requiring con¬

tinual artificial replenishment, since its original spawning areas

were reduced or eliminated.

Period of Intense Carp Removals (1950-1959)

An intensive and effective carp seining program was instituted

by the Wisconsin Conservation Department between 1953 and

1955 because earlier efforts had not reduced the population to low

enough levels. After each of 14 seine hauls in the 1950’s, the num¬

ber of carp remaining in the lake was calculated with mark and

recapture techniques and by the depletion of the residual popula¬

tion (Neess, Helm, and Threinen, 1957). Records from these sein-

ings for large fish and a major study by Helm (1958) using trawls,

fyke nets and small mesh seines for smaller species provide good

data on the fish community during the early and mid-1950’s.

The same large fish predators were present (Fig. 4) as in the

1940’s, but, among the predators, only largemouth bass were

increasing in abundance (Fig. 5). Largemouth bass entered the

1950’s at low to moderate population levels. Seinings in the early

1950’s revealed a sharp increase in their population (Lake Wingra

Files, Wisconsin Department of Natural Resources, 1971). Large

to good hatches occurred from 1954 to 1956 (Helm, 1958). Scien¬

tific collections and angling indicated further increases in the late

1950’s.

As many as 480 longnose gar were caught in three seine hauls

in 1952-1953, but by 1955 none were caught in six hauls (Lake

Wingra Files, Wisconsin Department of Natural Resources, 1971).

Helm (1958) noted that they continued to decline even though

returned to the lake after each seine haul. Thus longnose gar joined

the smallmouth bass as an extinct or rare native predator in Lake

Wingra (Figs. 4 and 5). Smallmouth bass were listed as present

by Helm (1958), but none were mentioned from any of the seine

hauls.

The low populations of both walleye and northern pike con¬

tinued to decline. Based on seine hauls, the northern pike fell to a

new low (Fig. 5). According to Helm (1958), marshes along north

central and western shorelines had virtually been filled in by nat¬

ural sedimentation in recent years to even further restrict spawn¬

ing to three limited areas, only one of which was large enough for

many young. Northern pike was the only species stocked in the

1950’s — 4,500, largely from the 1956 hatch, were stocked in Janu¬

ary, 1957. Helm stated some limited natural reproduction of wall¬

eye was occurring, but the lake did not have any traditional spawn-

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 83

ing habitats. Walleye were seldom caught by sport fishermen or in

the seines.

Bowfin and northern pike-muskellunge hybrids were caught in

low numbers in the seinings throughout the early 1950’s. The

hybrids, last stocked in 1948, were still present in 1955.

The same medium predators on invertebrates and fish were pres¬

ent as in the 1940’s (Fig. 4). In order of decreasing abundance

(Fig. 5) they were: white crappie, bluegill, black crappie, yellow

bass, and yellow perch. White bass, pumpkinseed and green sunfish

were low in abundance, and hybrids with their close relatives were

noted.

Populations of white crappie, bluegill, and black crappie were

observed in Helm’s careful study to fluctuate from year to year

(Fig. 5). Three trends were perhaps evident, though: an increas¬

ing abundance of bluegill from 1954 to 1958, a decline in white

bass, and a decline in the welfare of the yellow bass population.

Bluegill seemed to be favored by the increased vegetation after

carp removal, with good hatches in 1954-1956. Yellow bass were

abundant in 1953 and early 1954, but high mortalities occurred

during 1954. In a letter dated June 27, 1955, referring to carp

removals, Elmer Herman, then Area Coordinator for the Fish

Management Division of WCD, wrote: “We are already observing

great increases in the amount of rooted vegetation, decided dete¬

rioration of the yellow bass and increases in the number of bass

and bluegills” (Lake Wingra File, Wisconsin Department of

Natural Resources, 1971). Yellow bass numbers were at a low

point in late 1954 and 1955. By 1957, yellow bass numbers had

substantially recovered, but the new population of yellow bass was

noticeably stunted (Table 3). Although the numbers of yellow bass

had returned to a high level, their biomass remained a great deal

less (Helm, 1958).

By this time three bullheads were present : the yellow bullhead,

the brown bullhead, and the black bullhead (Ictaiurus melas). The

TABLE 3. AVERAGE TOTAL LENGTHS OF YELLOW BASS BEFORE

AND AFTER INTENSIVE CARP REMOVAL PROGRAMS IN

LAKE WINGRA. ADAPTED FROM HELM (1958)

Total Body Lengths (mm)

Before Carp After Carp

Removal Removal

Age (Years) (1945-1946) (1954-1957)

One 179 135

Three 232 160

84 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

yellow bullhead was most common. Helm (1958) still listed channel

catfish, common white sucker, and northern redhorse as present.

Bigmouth buffalofish were more abundant in the seine hauls than

earlier: 71 in 1952-1953, 125 in 1954, and 45 in 1955 (Lake Wingra

File, Wisconsin Department of Natural Resources, 1971).

Carp were greatly reduced during the 1950,s. From the mark-

recapture data, the carp population was estimated at 27,837 fish

prior to the start of the control program in 1953. By the end of

1953 (two seine hauls), the number had been reduced to 23,321.

By the end of 1954 (six seine hauls), the estimate was 5,585, and,

by the end of 1955 (six seine hauls), 2,698. Not all mortality was

from fishing. Natural mortality losses (especially over winter)

approximately equaled the yield to seining. Also notable was the

lack of reproduction by the carp population. Follow-up studies in

1956 consistently failed to find any young carp, indicating a further

decline in the population (Neess, Helm, and Threinen, 1957). Helm

(1958) computed the number of carp in Lake Wingra as of March,

1957, at 685 to 2,174 fish.

A comprehensive list of small predators on invertebrates was

also presented by Helm (1958). These included the species men¬

tioned by Cahn and Pearse in the 1910,s, except for the mudminnow

and the blacknose shiner. The golden shiner was at a moderate

population level in the lake. Four new species were reported: the

lake emerald shiner (Notropis atherinoides) , the satinfin shiner

(N. analostanus) , the central common shiner (N. cornutus), and

the Iowa darter (Etheostoma exile). These may have been over¬

looked by the earlier investigators, may have invaded the lake

during 1917-1919, or they may have been introduced as bait min¬

nows. Helm does not state that any of these species were abundant,

only that they existed.

The fish community entered the 1960’s with few carp, a declining

predator population except for the largemouth bass, and an abun¬

dant and diverse community of medium predators on invertebrates

and fish. The lake had begun to respond to the absence of carp and

rooted aquatic macrophytes were expanding in the littoral zone.

Second Period of Biological Adjustment (1968-1973)

In this period no major modifications by man were initiated and

the lake’s fauna again had time to come into a new equilibrium.

An intensive study of the fishes primarily through the Interna¬

tional Biological Program (see INTRODUCTION) from 1966

through 1973 documented fish community structure 11 to 18 years

after the completion of carp removal, 35 years after the massive

species introductions, 50 years after large reductions in adjoining

1974] Baumann , Kitchell , Magnuson, Hayes — Lake Wingra 85

marsh land, and 100 years after white settlers first inhabited the

watershed.

Fishes were sampled by trawls, fyke nets, electrofishing, and

beach seine (Magnuson and Kitchell, 1971; Baumann, 1972;

Churchill and Magnuson, 1972; El Shamy, 1973; and Baumann

and Kitchell, 1974). Population estimates by mark and recapture

were begun in 1972. In all activities, approximately 500,000 fishes

were captured, identified and enumerated.

The fish predator community only included largemouth bass and

northern pike (Fig. 4) as far as any practical influence on the

ecosystem is concerned. Walleye, longnose gar and bowfin were

rare. Smallmouth bass were absent entirely from the catches.

Largemouth bass were the most abundant predator (Fig. 5) and

were the third most common species taken by electrofishing in

1970. Angling pressure was low, but good catches of large bass

were not uncommon. Young fish were frequently taken in samples

from the littoral zone indicating successful reproduction.

Northern pike continued to decline since stocking ceased in the

late 1950’s. They were rare in anglers’ catches and young were

rare in any sampling gear. Fyke nets during early spring, 1973,

captured a few northern pike representing several size classes

(Churchill, personal communication). Redwing Marsh (Fig. 1) has

provided the only suitable spawning habitat in recent years.

Two adult walleye were caught since 1970. They probably are

limited by suitable spawning sites and represent residuals from

past stocking. Only an occasional bowfin was caught and two long-

nose gar were caught in a fyke net in 1972. Smallmouth bass were

occasionally reported by fishermen in the early 1960’s, but none

have appeared in surveys or in the sport fishery since then.

The medium predators on invertebrates and fish remained diverse

and, in decreasing order of abundance, included bluegill, white

crappie and yellow bass, black crappie, pumpkinseed, yellow perch,

and green sunfish (Figs. 4 and 5). White bass had disappeared.

The abundance (Fig. 5) of several species had changed markedly

since carp removal and the redevelopment of dense littoral macro¬

phytes. Bluegill and pumpkinseed both increased greatly and yellow

bass increased slightly. Both white and black crappie decreased

in numbers. Yellow perch remained stable but low. Green sunfish

had apparently increased to a low but noticeable population level.

Bluegill were by far the dominant fish species, comprising well

over one half of all fish caught during 1970 and 1971. Large num¬

bers of young-of-the-year bluegill indicated successful hatches in

every year except 1972. Churchill (personal communication) re¬

calculated population numbers for the total lake (Churchill and

86 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Magnuson, 1972) to account for sampling bias and marking mor¬

tality. Yearling bluegill were estimated during May, 1972, to have

been approximately 8,000,000 fish; subadults (11+) about 800,000;

adults (III+ to V+) about 340,000.

Bluegill were numerous, but stunted (Fig. 6) or characterized

by poor growth (El Shamy, 1973; Kuczynski, unpublished) com¬

pared with earlier years (Helm, 1958). The reduced growth began

to appear in 1955-1957, immediately after completion of the carp

removal, and by 1970-1972 was even lower (Fig. 6). At annulus I

total lengths seemed comparable, but by annulus V the average

length had declined by 25% over the 25 years since 1945-1946. In

terms of body weight this translated to a reduction of approxi¬

mately 70%.

Yellow bass adults were estimated by Churchill (personal com¬

munication) to be about 50,000 in May, 1972, and were second only

AN NULUS

FIGURE 6. Body growth curves for bluegill in Lake Wingra before, during

and after the successful carp control program of the early 1950’s.

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra 87

to bluegill in numbers caught during 1970 surveys, with 9,983

caught. The yellow bass after carp removal immediately began to

stunt (Table 3) and remained stunted through the 1960’s. Surveys

in 1968-1973 rarely captured yellow bass larger than 175 mm.

No white bass were captured from 1968 to 1973 ; they were most

likely ecologically replaced by yellow bass with the last members

genetically swamped out of existence by the hybridization noted

in earlier periods. A similar replacement has been occurring in

the other Madison lakes (Wright, 1968).

White crappie adults were estimated (Churchill, personal com¬

munication) to be approximately 50,000 fish in May, 1972, and over

4,700 were captured in the 1970 surveys. Adults of this open water

centrarchid appeared stunted, usually no longer than ca. 190 mm.

Black crappie were the most common fish in samples from the

littoral zone in 1970 — 1,114 out of 4,308 fish captured. On the other

hand, only 492 of the 42,160 fish caught by trawls in the pelagic

zone were black crappie. Overall they were the fifth most common

species caught in the surveys, with 1,606 captured. As noted by

Helm (1958) , the two crappie species spatially segregate with black

crappies concentrated in the littoral zone and white crappie con¬

centrated in the pelagic zone. In the 1965-1973 surveys, adult black

crappie were also stunted, usually no longer than ca. 185 mm.

On rare occasions we have observed large white crappies and

black crappies (ca. 300 mm-400 mm) , suggesting that a few become

large enough to feed routinely on young fishes.

Pumpkinseed adults and subadults in May, 1972, were estimated

to total 28,000 by Churchill (personal communication), and were

fourth in abundance in the 1970 survey, with 2,950 captured. While

this may be an overestimate of their actual relative abundance,

they were certainly among the five most abundant species and were

apparently favored by the carp removal. Also in the 1970 surveys,

195 yellow perch and 30-40 green sunfish were caught.

All three bullheads remained (Fig. 4), but by the late 1960’s

black bullheads and yellow bullheads were equally abundant in

catches and the native brown bullhead was rare. No channel catfish

were caught or reported during the period; they have probably

become extinct in this lake.

Neither bigmouth buffalo nor northern redhorse were reported

during the period and were likely extinct in this lake. Common

white sucker were captured singly and infrequently. They exist

in low numbers.

Carp populations were moderate and had not returned to high

levels after the removals in the early 1950’s. Only 58 of the 46,500

fish captured in 1970 were carp. With only several exceptions, all

88 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

carp were large, mature fish even though the gear would more likely

have captured smaller fish. During one 24 hour electrofishing sur¬

vey in shallow areas during September, 1970, no young-of-the-year

or yearling carp were caught, even though 1,200 young-of-the-year

bluegill were captured (Baumann, 1972). Apparently reproduction

by carp has been relatively unsuccessful in recent years, but we

cannot exclude the possibility that they may be slowly increasing.

After first being noted in the lake just prior to 1900, they were not

common until the 1910’s nor abundant until the late 1920’s or early

1930’s.

Small predators on invertebrates in the lake proper included

golden shiner, silversides and bluntnose minnows. Golden shiners

were abundant — 280 were caught in the 1970 survey. Silversides

were also caught in moderate numbers.

While the surveys from 1968-1973 did not concentrate on the

springs, spring pools and tributaries, the mudminnow was again

caught in the tributaries. Surprisingly, this is one of the two

smaller species mentioned by Cahn (1915) that Helm (1958) did

not find. Some of the other small species listed by Helm (1958)

may still be present but are not abundant in the lake proper.

The fish community of Lake Wingra proper entered the mid-

1970’s with 20 species of fish. Of these, only 12-14 species were

probably native to the lake. In the period of adjustment after carp

removal, carp remain low in abundance, predators remain low in

abundance, and invertebrate feeders are characterized by poor

body growth. The lake is not a major protein source to the city

around its shores but children enjoy a high catch rate of small

bluegill, pumpkinseed, yellow bass, and crappies during summer

vacations. Fishing through the ice in early winter provides some of

the better sport fishing.

SUMMARY AND CONCLUSIONS

Man’s activities in the Lake Wingra basin dominate its history.

Indians used the region as a major hunting and fishing ground.

Homesteads did not appear in the Wingra watershed until the late

1860’s, and the area remained isolated past the turn of the century.

From 1905 to 1920, major physical alterations of lake and sur¬

rounding wetland were made. Urbanization of the north shore fol¬

lowed the development of Nakoma and Madison from about 1910.

The original hydrography of Wingra before alteration by man

was different than now. Water level was about 0.3 m higher.

Gardner Marsh and upper Murphy Creek were originally part of

an extensive shallow area at the east and southeast end of the lake.

Shoreline extended beyond the lowland, now part of Vilas Park.

1974] Baumann, K itch ell, Magnuson, Hayes — Lake Wingra 89

Wetland surrounding the lake also covered a wider area. Maximum

depth was about 4.3 meters.

In 1905 man began to engineer changes in the hydrography of

the basin. Years of activity at the east and southeast end of the

lake resulted in the isolation (from the lake) and partial drainage

cf an area, now Gardner Marsh. Murphy Creek was channeled and

lowered below lake level after the present spillway dam was built.

From 1917 to 1919, the lake's level was lowered about 1.3 m below

the original; reduction of surface area resulted. After dam con¬

struction, the lake was raised 1.0 m to its present level. Material

dredged from the lake and creek were used to fill southern wetland.

Dredging created a deep trench off of the south shore. This has

partly filled, giving the lake its present maximum depth of 6.4 m.

Construction of Vilas Park reduced surface area and altered the

northeast shoreline. After 1920, the only major hydrographic

change in the lake proper was. the formation of Ho-nee-um Pond.

Lake and contiguous wetland areas were reduced by a factor of

three. Urbanization within the basin resulted in the disappear¬

ance of 28 springs; major changes in soil and vegetation types;

and the development of a storm sewer system that carries surface

water and allochthonous matter to the lake's periphery from much

of the drainage basin.

Original aquatic macrophyte vegetation around 1900 differed

in type, abundance and distribution from the present. Cattails

and bulrushes dominated shallow areas; wild rice abounded in

slightly deeper water. Submerged vegetation included water celery

and pondweed.

Dredging operations destroyed vegetation directly and indirectly.

Fluctuations in water level had detrimental effects. Hydrographic

alterations reduced the area available for littoral growth. From

the late 1920's to the mid-1950’s, carp nearly denuded Lake Wingra

of macrophytes. After an extensive carp removal program, vege¬

tation returned, but it was of a different nature than the original.

Lake Wingra's vegetation is now heavily dominated by a non-native

species, Eurasian water milfoil (Myriophyllum spicatum) . Wild

rice, wild celery, and several other native species are no longer

present.

Major changes in habitat and heightened predation pressure

from increasing fish populations resulted in substantial reductions

of invertebrate populations. Many species of zooplankton are no

longer found in the lake while others, and some macroinvertebrates

such as Hyalella azteca, have been reduced to virtual extinction.

Fish populations in Lake Wingra have changed continually.

Species native to the lake in 1900 included bluegill, pumpkinseed,

90 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

black crappie, largemouth bass, smallmouth bass, longnose gar,

yellow perch, northern pike, bowfin, golden shiner, brown bullhead,

and possibly the bigmouth buffalo and walleye. Largemouth bass,

yellow perch, northern pike and walleye populations have declined.

Longnose gar, smallmouth bass, bowfin and bigmouth buffalo are

actually or functionally extinct from the lake.

Changes in the native fish fauna can be partly explained by the

reduction of littoral areas due to hydrographic alterations and

the activity of carp. Selective fishing and the introduction of exotic

fish also played a role. The fish rescue-and-transfer operation was

an unfortunate outgrowth of the early conservation ethic, which

was gaining popularity during the 1930’s in Wisconsin. Reclaiming

these fish was considered beneficial in itself. The benefits of stock¬

ing fishes in a lake was an accepted truth of the time, and little

effort was made to correlate specific species with habitat require¬

ments. Among the 23 species stocked into Lake Wingra, two —

yellow bass and white crappie— have become abundant.

Both largemouth bass and bluegill populations have recovered,

since the carp population was controlled in the 1950’s. The bluegill

responded by becoming the dominant species in the lake. This in¬

crease together with the establishment of white crappie and yellow

bass have produced the large, stunted panfish population which

characterizes the present sport fishery.

Other contributors to the increase in panfish populations were,

first, the decline of native fish predators and, recently, the dense

aquatic vegetation that even further reduced the effectiveness of

those predators remaining. The depletion of both the gar and the

northern pike appeared to be independent of carp abundance per se.

Gar were reduced by rough fish removal aimed primarily at carp,

but the presence of a large carp population did not seem detri¬

mental to the gar population. Northern pike decline is closely linked

to destruction of suitable spawning habitat.

Wingra, like most lakes, has never been managed as an eco¬

system, but rather individual problems have been attacked one

at a time. To diversify the angling and reclaim fish, new species

were introduced. Little thought was given at the time as to whether

these species could thrive or to whether they would negatively in¬

fluence native species. Marshlands were drained for park and land

development, and the reproductive success of higher predators was

severely reduced. Carp were first introduced into our waters, then

became a nuisance and were actively removed. Again, the sec¬

ondary effects of carp removal were not anticipated.

To be effective, management plans must consider the entire fish

community and the total ecosystem. The historical perspectives

described in this paper serve as illustration of the complex inter-

1974] Baumann, Kitchell, Magnuson, Hayes — Lake Wingra

91

actions and response capabilities of a total ecosystem. An integrated

whole ecosystem approach is essential in establishing ecologically

sound management. Although resources are often not available to

make an overall ecosystem study, sound management would seem

to require consideration of as many interactions and secondary

effects as time, funding, and contemporary knowledge and tech¬

niques permit. As always, hindsight is better than foresight, which

probably explains why the next chapter of this article is not yet

written.

ACKNOWLEDGEMENTS

We thank Dr. Grant Cottam, Dr. John C. Neess, Dr. Wayland E.

Noland and Mr. C. William Threinen for reviewing preliminary

drafts of this paper. Ms. Patricia Tennis and Mr. John C. Gryskie-

wicz provided diligent technical support in assembling materials

and information.

Support supplied in part by the Eastern Deciduous Biome, Inter¬

national Biological Program, funded by the National Science

Foundation under Interagency Agreement AG-199, 40-193-69 with

the Atomic Energy Commission — Oak Ridge National Laboratory.

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A HISTORICAL SKETCH OF THE

EVOLUTION OF ENERGETICS

Robert T. Rainier

University Wisconsin

— Milwaukee

INTRODUCTION

Let ns begin by eliminating, as much as possible, the word “ther¬

modynamics” from our vocabulary. For we are not interested here

in the “dynamics of heat” in systems which are, in every way,

static, or what amounts to the same thing, quasistatic. Let us begin

anew and, without trying to coin new words or phrases, accurately

designate the ideas we propose to explore. We may easily title what

up to this point has been called “classical thermodynamics” with

simply “static energetics.” Since energetics is defined as “the sci¬

ence of energy,” static energetics is then to be taken as “the science

of energy in energetically static systems.” And the concepts which

of late are becoming known by the unfortunate, cumbersome and

inaccurate label of “irreversible” or “non-equilibrium” thermody¬

namics simply accept the title “dynamic energetics.” Dynamic

energetics is defined as “the science of energy in energetically

dynamic systems.” Thus energetics, like mechanics, consists of a

study of statics and dynamics. But whereas in mechanics one is

concerned with only the state of forces and motions of the system,

in energetics one is concerned with the total energy state of the

systems. Thus, in some sense, energetics includes mechanics as a

subset in the hierarchy of science.

Now that we have a common terminology base, we can begin

to trace the thread of energetics through time. My ultimate goal

here is to establish, both historically and logically, the place of

energetics as an axiomatic science which will eventually replace

both classical and irreversible (or whatever) thermodynamics as

a fundamental body of man’s knowledge.

ANTIQUITY TO THE RENAISSANCE

The modern scientific concept of “energy” dates only from the

19th century, and the word, with the spelling used here, seems not

to have been in use at all before about 1580. 1 Etymologically, the

word “energy” is of Greek origin, coming from the roots en — “in”

plus ergon — “work”,2’3 and seems to date from around the Platonic

95

96 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

era.4 The original Greek “cvepyei a,” was rendered into English5 as

“energia,” with this spelling dominating until about A.D. 1600.

Samuel Johnson,6 in his unique dictionary of the English language,

states :

“I have not been able to trace our word [energy] to any author before

Bacon [1626]. Sidney, in his Defence of Poesy, written soon after 1580,

shows that it was not then in use; for he there introduces, in its stead,

‘energia, as the Greeks call it.’ ”

A brief review of English dictionaries published from the time

of Bacon reveals Thynne7 (1599) using “energye,” Cockeram8

(1623) using “energie,” and Phillips9 (1658), Coles10 (1696), Ker¬

sey11-12 (1702 and 1708), Bailey13 (1730) and Sheridan14 (1788)

all using “energy.”

The meaning of the word “energy” (or “energia”) up to the

19th century remained essentially unchanged from that of the orig¬

inal Greek usage. Aristotle (ca 384-322 B.C.) defined energia as

meaning activity or actuality,15 commonly using these concepts

interchangeably.16 His usage, and that of the civilizations that fol¬

lowed him, implied a concept that is much more general than the

simple “in-work” etymological translation. However, Aristotle’s

rendering of energia as some form of undefinable “potential” is a

characteristic of the concept of energy today. “The capacity for

performing work” is the modern scientific definition of energy, and

capacity and potential are synonymous here.

Dictionaries of the 17th and 18th centuries consider energy as

synonymous with such words as strength, force, power, vigor and

efficacy. If we attempt to trace the energy concept into pre-Grecian

civilizations, we therefore must look for the concepts of work,

strength, power and force. Indeed, there are Assyrian words17 and

Egyptian hieroglyphics18 with exactly these meanings. However,

pre-Grecian antiquity seems sadly devoid of eloquent philosophers

whose works have survived to provide us with a tool for the

accurate translations of abstract concepts. It is probably true that

without such philosophers there is no need for words that are more

than simply descriptive,19 and thus abstract ideas will not naturally

evolve easily in such cultures.20

DESCARTES, LEIBNIZ AND VIS VIVA

Rene Descartes (1596-1650), latinized as Renatus Cartesius, in

his Principles of Philosophy (1644 ) 21* 22 made reference in his laws

of motion to the divine conservation of the “quantity of motion”

of a body. To Descartes this quantity was the volume of the body

times its velocity.23 In 1686 Gottfried Wilhelm Leibniz (1646-

1716) 24 published a strong refutation of Descartes’ position. Leib-

1974] Balmer — Sketch of Evolution of Energetics 97

niz felt that the force of a body in motion was uniquely different

from Descartes’ “quantity of motion.” His position was that:25

“. . . the forces are in compound proportion to the bodies (of the same

specific weight or density) and the generating heights of the velocities . .

Thus Leibniz held that the force of motion was proportional to the

square of the velocity. Further, he felt that there were two general

classifications of forces: living forces (dynamics) and dead forces

(statics). In 1695 he concludes:20’27

“Force is twin. The elementary force, which I call dead because motion

does not yet exist in it, but only a solicitation to motion, is like that of a

sphere in a rotating tube or a stone in a sling.

“The other is the ordinary force associated with actual motions, and I

call it living .”

This was for formal beginning of what was to be known for the

next 200 years as the “vis viva” (living force) controversy.28

Stated simply, the vis viva controversy is this : What is the force

of motion, mv or mv2? In a way, it is similar to the question : Which

is more fundamental, mv or mv2? These questions have never been

resolved. In 1747 the then 22 year old Immanual Kant29 recognized

the hopelessness of resolving vis viva.

The controversy involves two abstract concepts, force and mo¬

tion. Soon after the controversy started, scientists began to feel

that the whole argument could be settled simply by an arbitrary

definition. D’Alembert (1717-1783) is credited by many early

historians of science with ending the dispute in 1743 in his Traite

de Dynamique by referring to the controversy as:30’31

a: “. . . dispute of words too undignified to occupy the philosophers any

longer.”

It is, however, evident that the dispute was neither simply one of

definitions, nor was it ended by D’Alembert.32 There are many

examples in the literature following D’Alembert, wherein the con¬

troversy is taken up both philosophically and experimentally.33 It

was regurgitated most recently in the famous Ostwald-Boltzmann

energetics debate of 1895.34

The controversy has never been resolved and the question of the

ultimate fundamentality of energy vs. momentum is still occa¬

sionally raised today.35 However, when one thinks of motion today,

he thinks of F = ma, and not of energy. Though true that the dis¬

pute remains unresolved, the champion of mechanics, momentum,

seems to have won by popular vote. Perhaps because mechanics

was more abstractly axiomatic and more mathematically developed

than early energetics and thus was more functional, or perhaps

because man just feels more at home with mechanical concepts.

But whatever the reason, energetics was off to a poor start.

98 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Interest in the controversy finally dwindled in the late 19th cen¬

tury. Vis viva does not appear in the scientific literature later than

Thomson and Tait’s Treatise on Natural Philosophy in 1867. 36

Scientific jargon changed somewhat in the late 19th and early

20th centuries; today the force of motion is simply not discussed

by physicists. The word “force” has been reserved, in general, for

the Newtonian meaning as being derived from an acceleration.37

Momentum (Latin for “motion”) was the name finally given to

the quantity mv. The term which eventually received, for a short

time, the label vis viva was mv2. In 1804 Young38 began calling

mv2 “energy,” and Rankine39 in 1853 dubbed it “actual” or “sen¬

sible” energy. Its final name “kinetic energy,” was given by Thom¬

son and Tait40 in 1862.

Thus energetics was born in controversy, in opposition to the

worst possible opponent, the established monarch, mechanics. There

were two crucial events which would determine the superiority of

the stronger. The first was the consequences of the rise and fall

of the caloric theory of heat; the second was the tremendous suc¬

cess of what was later to be called, “statistical mechanics.”

HEAT

The failure of the caloric theory of heat was a crippling blow

to the 19th century energetists. Though they alone were not respon¬

sible for inflating the theory to a position of scientific importance,

when it collapsed under the strain of attacks by experimentalists

they bore the consequences and suffered the humiliation of a not-

so-obvious error. The fall of the caloric theory and the preceding

century and a half of confusion caused by the vis viva debate forced

the energetists onto their scientific knees. The final defeat of the

19th century energetists came from the statistical mechanics of

the Boltzmann atomists. They claimed that many of the results

of static energetics could be derived from statistical mechanics.

Thus energetics was reduced to mechanics, and therefore it con¬

tained nothing new or unique.

By the end of the 19th century the word “energetics,” connoting

at that time vis viva and caloric, both lost battles, had fallen into

disrepute and ultimate disuse. It is only now, almost a century

later, when all of these scientific calamities have been forgotten,

that the word is once again becoming popular.

Since the dawn of science there have been two rival theories on

the structure of matter : the atomlst versus the continuumist. And,

like the vis viva controversy, this dispute over the nature of matter

goes unresolved today.

This may surprise some, for who today can deny the existence

of atoms ? But a continuumist will say, “give me your most funda-

1974]

Balmer — Sketch of Evolution of Energetics

99

mental atomic particle, one which you are convinced is made up

of no others, and I will ask you — of what does this particle con¬

sist ?” “The answer can only be: a continuum.” And an atomist

will reply, “The question of the internal structure of an atomic

particle is meaningless — it does not enter into the microscopic or

macroscopic behavior of nature, and therefore is of no interest.”

And there the debate stagnates.

This antipathetic dichotomy reveals itself most clearly in the

modern “duality principle” of the structure of matter. Here we

experimentally observe that atomic particles sometimes behave like

mechanical particles, and sometimes as a continuum. This para¬

dox! al situation is analogous to the position of the caloric theory

as a result of the friction experiments of Rumford and Davy at

the beginning of the 19th century.

The caloric theory of heat was a natural branching in the theory

of the ether (or aether). The word “ether” is also of Greek origin

( mQrjp ) and means “clear sky” or “upper air.”41 Since Aristotlean

physics forbade the existence of a vacuum, it was necessary to fill

all space with something, something that behaved like a fluid. Soon

it was realized that light streaming through the clouds must some¬

how be associated with that imponderable fluid that fills all space.

The analogy was quickly drawn between the light-ether and the

sound-air systems. Thus light, like sound, was explained as vibra¬

tions of the propagating medium. And what about heat? If one

holds his hand up to a fire, can he not feel the heat pouring out?

There was no doubt that heat was also associated with the ether.

Descartes was the first to begin to quantitize and to assign

properties to the ether. Before his time the ether was used only

in the Aristotlean sense as a space filling fluid. In matters of ether,

Descartes was an atomist. He postulated that it was made up of

minute particles that were continually in motion. But since there

could be no empty space whatsoever (i.e., no vacuum), the motion

of these particles was complex indeed, with a moving particle

always entering the space which was simultaneously being vacated

by another particle.42

Descartes’ concepts of light were soon replaced by those of R.

Hooke, and Hooke’s by Newton’s and Newton’s by Huygen’s, and

so on until the modern theory of light as being the result of subtle

particles, now called quanta, was evolved.

By the middle of the 18th century, light was considered to be

composed of minuscule “corpuscles,” which move through the

ether. About this same time a third branch of the ether concept

was developing in the area of electricity. By 1750, the experimental

work with electric “currents” led to the introduction of an electric

fluid, the “effluvia,” which had its own properties. But before 1800

100 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

the laws of charge, attraction and repulsion had been discovered,

and effluvia was discarded in favor of the original ether, which was

now thought to be decomposable into the subtle fluids necessary to

explain all electrical phenomena.

A fourth branch of the ether concept was developed in 1679 by

George Ernst Stahl.43 Stahl invented the word “phlogiston”

(Greek: flame) to describe his fire principle of combustible mate¬

rials. All such materials were supposed to be composed of a calx

(ash) in combination with phlogiston. During combustion the

phlogiston escaped from the material leaving only the ash behind.

Lavoisier is usually credited with delivering the death blow to

Stahl’s theory some 100 years after its introduction.

There were, of course, other manifestations of the ether. The

magnetic fluid was one. And then there was perhaps the most

subtle ether of all, the “vital spirits” contained by living organ¬

isms. The elusive vital spirits were all that separated life from

death, like the soul.

At first heat and light were taken to be the same corpuscular

entity. However, the discovery of the “green house effect,” in about

1750, established that these two phenomena could not be simply

different manifestations of the same fluid. In 1789 Lavoisier44

introduced the word “calorique” to denote the ether of heat. Dur¬

ing the second half of the 18th century caloric was generally con¬

ceived to be the substance which occupied the interstices between

the particles of ponderable matter (atoms).

Near the end of the 18th century Rumford and Lavoisier estab¬

lished that the temperature of a body has no measurable effect on

its weight. Rumford and Davy then established that caloric was not

a conserved quantity as had been previously thought.45 This was

the beginning of the fall of caloric.

The atomists had a ready explanation. There had been an atomic

theory of heat for some time. To the atomists, heat was the result

of vibrations of the minute particles of matter. The rubbing of

two objects together simply caused these minute particles to

vibrate faster, and heat was directly proportional to the particles’

vibration.

In 1824 Sadi Carnot, a 28 year old French military engineer,

using the principles of hydraulics as a model for the flow of heat

(and accordingly the then known inaccurate conservation of caloric

principle) wrote his only publication, “Relexions sur la Puissance

Matrice de Feu,” in which he established the principle that the

thermal efficiency of a thermal energy conversion device was max¬

imized when the “flow of heat” was between bodies whose tem¬

peratures differ by only an infinitesimal amount.46 At the time

Carnot wrote his article he believed heat to be a fluid, caloric. But

1974] B aimer — Sketch of Evolution of Energetics 101

Carnot’s views on heat apparently changed before his death in

1832. A few of his undated notes were translated and published

in 1878, 47 in which he seems to have adopted the atomists’ position.

He writes :

“Heat is simply motive power, or rather motion which has changed

form. It is a movement among the particles of bodies. Whenever there is

a destruction of motive power, there is at the same time production of

heat in quantity exactly proportional to the quantity of motive power

destroyed. Reciprocally, whenever there is destruction of heat, there is

production of motive power.

“We can then establish the general proposition that motive power is

in quantity invariable in nature — that is, correctly speaking, neither ever

produced or destroyed. It is true that it changes form — that is, it pro¬

duces sometimes one sort of motion, sometimes another, but it is never

annihilated.”

We could be retrospectively generous and conclude from this

quote that Carnot was the first to actually state the conservation

of energy principle. However, the complete formulation of this

principle was not carried out until the second half of the 19th

century.

The primitive concepts of metabolic energy balances on physio¬

logical systems were apparent in the early experiments of Sanctus

Sanctorius (1561-1636) . 48 Although his “insensible perspiration”

embodied considerably more than simple perspiration, his approach

to the balance concept of metabolic energy was unusually advanced.

The first to correctly enunciate the conservation of energy prin¬

ciple correctly was a German physician, Julius Robert Mayer.49

Mayer’s interest in energy stemmed from then popular physiolog¬

ical studies on metabolism. As a result he performed no experi¬

ments himself but he utilized the results of the experiments of

others. This luxury of Mayer’s was to cost him most of the credit

for the discovery during his lifetime. Mayer published in Annalen

der Chemie und Pharmacie (1842) his ideas concerning the inter-

convertability of heat and work. His value for the mechanical

equivalent of heat was about 470 Btu/ft lbf.50

In 1843, Ludvig August Golding,51 unaware of Mayer’s work,

presented a paper to the Danish Academy in which he reported

original experiments which established the existence of the pro¬

portionality between heat and work. He did not, however, calculate

this factor of proportionality.

At about this same time James Prescott Joule was performing

extremely well designed experiments to determine the mechanical

and electrical heat equivalents.52 Joule, due to his experimental in¬

sight and excellence, received much acclaim for his work. He spent

nearly forty years of his life working on various aspects of heat

equivalents.

102 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

With this single equivalence between work and heat energetics

blossomed forth a complete and independent science. In 1848 Wil¬

liam Thomson (Lord Kelvin) unearthed Carnot’s ideas and devel¬

oped the absolute temperature scale.53 Clausius (1850) then re¬

moved the conservation of caloric premise from Carnot’s formula¬

tion and thus established for the first time the complete generality

of Carnot’s results.54 Thomson was the first to state both principles

of energetics together. In 1851 he wrote:55

‘‘The whole theory of the motive power of heat is founded on the two

following propositions, due respectively to Joule and to Carnot and

Clausius.

“Prop. I (Joule). When equal quantities of mechanical effect are pro¬

duced by any means whatever, from purely thermal sources, or lost in

purely thermal effects, equal quantities of heat are put out of existence,

or are generated.

“Prop. II (Carnot and Clausius). If an engine be such that, when it is

worked backwards, the physical and mechanical agencies in every part of

its motions are all reversed; it produces as much mechanical effect as

can be produced by any thermo-dynamic engine, with the same tempera¬

tures of source and refrigerator, from a given quantity of heat.”

In 1855 Rankine56 presented the first complete axiomatic study

of the science of energy in his “Outlines of the Science of Ener¬

getics.” He begins with a detailed discussion of abstractive vs.

hypothetical scientific methods, and continues with a series of

carefully formulated definitions of such terms as substance, prop¬

erty, mass, work, etc. On page 214 he defines the Science of Ener¬

getics as :

“. . . a science whose subjects are, material bodies and physical

phenomena in general, . . .”

He then continues, on page 218, with the “First Axiom” of ener¬

getics :

“All Kinds of Work and Energy are Homogenous.”

By which he meant that “energy is transformable and transfer¬

able.”

On page 219 he states the “Second Axiom” of energetics :

“The Total Energy of a Substance cannot be Altered by the Mutual

Actions of Its Parts.”

That is the principle of the conservation of energy.

And on page 220 he states the “Third Axiom” of energetics :

“The Effort to Perform Work of a Given Kind, Caused by a Given

Quantity of Actual Energy, is the Sum of the Efforts Caused by the Parts

of that Quantity.”

Unfortunately, Rankine’s axiomatic approach did not find favor

among his contemporaries. By the beginning of the 20th century

the subject was called “thermodynamics,” and its governing prin-

1974] B aimer — Sketch of Evolution of Energetics 103

ciples were called “laws.” The main influence for this terminology

seems to be the writings of Clausius.

Also by the beginning of the 20th century the concept of “heat”

had taken on a new meaning. Heat was neither caloric nor was it

the motion of atoms. Heat was abstracted, it now represented that

energy which is transferred across a system boundary due to a

temperature difference.57 The vibratory motion of the atoms was

now regarded as “innerer Arbeit,” (interior work) by Clausius;58

as “intrinsic energy,” by Perkins;59 as “l’energie interne,” (inter¬

nal energy) by Poincare.00 Today it is universally known as

internal energy.

Like vis viva, the meaning of the concept of heat cannot be estab¬

lished by an arbitrary definition. Caloric was originally presumed

to have weight, but no weight change with caloric change could

be measured. However, the modern theory of relativity predicts

that heat does have weight. In 1930 Tolman61 published “On the

Weight of Heat and Thermal Equilibrium in General Relativity,”

in which he shows that all the energy of a system has the property

of inertia. Thus if an amount of heat, aQ, is added to a system its

weight will increase, AW, by an amount given by:

AW = AQ (-iC) ,

where g is the acceleration due to gravity and C is the velocity of

light. Consequently if a body has 1 Btu of energy given to it via a

temperature difference between the body and its surroundings, then

the weight of the body will increase by about 1.2 X 10~13 Ibf

(5.5 X 10~n gm). This value is too small to be detected, even today.

But this may not always be the case. What a different history ener¬

getics would have had if caloric had been measurable. Thus

caloric’s last and perhaps greatest success came 150 years too late.

THE INFLUENCE OF MECHANICS

Mechanics has always been the favorite son of science. It became

the ruler of architecture — it created the pyramids of Egypt. It

became necessary in agriculture — it reaped the Roman wheat. It

became useful in commerce — it carried produce to and from foreign

ports. Thus human intuition in mechanics has been developed by

10,000 years of the necessity of invention.

In antiquity energy was too abstract a concept to be developed

completely ; what little energetics there was found itself hopelessly

entangled in the mechanics of the day. It was not until the 18th

century that energetics began to evolve as an independent entity.

But energetics was not looked upon as a welcome addition to

mechanics, and by the 19th century, as Gillispie02 puts it, “ener-

i04 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

getics became the opposite pole to mechanics.” Energetics has not

significantly changed its polarity since then. As an axiomatic study

it seems to have been stillborn with the work of Rankine. The

bastard child — thermodynamics — survived, and as it grew it

became evident that it was retarded.

The dynamic aspects of mechanics became more or less axio-

matized with the work of Aristotle (ca. 384-322 B.C.). Although

he was attempting to emulate nature, he realized this was a formid¬

able task and satisfied himself with principles which were abstract

enough so as not to be violated by observations.

Aristotlean dynamics reigned dominant, though inaccurate, for

almost 2000 years. It was finally discarded in the 17th century

largely as a result of Torricelli’s63 (and others) work with

vacuums. The dynamics of Galileo and Newton became firmly en¬

trenched during the 18th century, and today we do not speak of

“laws of nature” in dynamics, but instead we speak of Aristotlean,

or Galilean, or Newtonian, or Einsteinian dynamics (or rather of

the axioms contained in these studies).

The static aspects of mechanics began to become axiomatized

with the work of Archimedes (ca. 287-212 B.C.). His entire theory

of hydrostatics was founded upon the following two axioms:64-65

“Postulate 1

Let it be supposed that a fluid is of such a character that, its parts

lying evenly and being continuous, that part which is thrust the less is

driven along by that which is thrust the more; and that each of its parts

is thrust by the fluid which is above it in a perpendicular direction if the

fluid be sunk in anything and compressed by anything else.

“Postulate 2

Let it be granted that bodies which are forced upwards in a fluid are

forced upwards along the perpendicular [to the surface] which passes

through their centre of gravity.”

Thus, by the 17th century an axiomatic mechanics ruled science.

During the 18th century the omnipotent “mechanical philosophy,”

became deeply entrenched in science. One of the greatest successes

of this philosophy was the reduction, at the end of the 19th cen¬

tury, of the concepts of energetics to principles of statistical

mechanics, and, in particular, the rendering of Clausius’ illusive

“entropy” as simply mechanical probability.

The unrealistic zeal of science over the mechanical philosophy

essentially smothered the new energetic approach. It emphasized

the failure of energetics and at the same time ignored the short¬

comings of the theories built on the concept of force. The fall of

caloric was a catastrophe, but when the space-filling ether of the

mechanists finally dissolved under the influence of Michelson and

Einstein,66 hardly anyone noticed.

1974] B aimer' — Sketch of Evolution of Energetics 105

It is evident that the influence of mechanics on the philosophies

of science was, and is, considerable.

ENERGETICS SINCE RANKINE

Between the work of Rankine in 1855 and that of Caratheodory07

in 1909 very little contribution was made to the axiomatization of

the science energy.

Plank,08 in his 1897 V orlesungen iiber Thermo dynamik, recog-

ized that there were “essential difficulties ... in the mechanical

interpretation of the fundamental principles of Thermodynamics/’

and he decided to develop the subject in such a way that:

“. . . it does not advance the mechanical theory of heat, but, keeping

aloof from definite assumptions as to its nature, starts direct from a few

very general empirical facts, mainly the two fundamental principles of

Thermodynamics.”

Although his intentions were good, his approach was not abstract

enough to be considered a truly axiomatic study. He could not com¬

pletely divorce himself from the mechanical world of the cyclic

heat engine and perpetual motion.

Rankine axiomatized the conservation of energy principle and,

in 1909, Caratheodory, a mathematician, axiomatized the Clausius-

Carnot entropy principle. Catatheodory’s aim was to establish an

equivalent mathematical axiom for the “second law of thermody¬

namics” which would be independent of any mechanical devices

(such as heat engines). He was completely successful, and conse¬

quently developed the following statement of the axiom:09

“In the neighborhood of any arbitrary initial state P0 of a physical

system there exists neighboring states wThich are not accessible from P0

along quasistatic adiabatic paths.”

In 1917 Tolman70 introduced today’s concepts of “extensive” and

“intensive” properties, terms which have been parroted by text¬

book authors ever since, without bothering to carry along the orig¬

inal meanings. Tolman was attempting to introduce the fundamen¬

tals of measure theory into energetics. This problem does not arise

in mechanics, since it is built upon our most primitive concepts of

measure: mass (and/force), length and time. However, energetics

requires, in addition, the measurement of fundamentally more ab¬

stract quantities of energy (internal, mechanical, chemical, nuclear,

electrical, etc.), temperature, and entropy. Tolman postulated that

only extensive properties were additive mathematical measures,

and thus only they could serve as fundamental quantities. An

intensive property (such as temperature), therefore, cannot be a

fundamental entity.

106 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Fowler and Guggenheim71 introduced the so-called “zeroth law

of thermodynamics’’ in 1939. 72 It is unfortunate that this terminol¬

ogy has caught on, for this, even more than the other so-called

“laws,” is no law. Its value to energetics is apparently recognized,

but few authors have understood its true meaning. It is nothing

more or less than the transitive property (mathematical) which is

a necessary condition for the existence of an equivalence relation.73

If the reflexive and symmetric properties are added to the zeroth

law statement,71 then we have defined “thermal equilibrium” as an

equivalence relationship.

More recently the axiomatic approach has been adopted by

Landsberg75’ 76 (1956), Falk and Jung77 (1959), Callen78 (1960),

Tiza79 (1961), Fong80 (1963), and Truesdell and Coleman81'83

(1966). Of this group only Landsberg, Truesdell and Coleman are

mathematicians, and their developments, as far as they go, are

necessarily the clearest. Coleman’s work includes the “functional”

concept with which Noll84 had such success in describing material

deformation phenomena.

Since Caratheodory energetics has largely been ignored by

creative scientists, except for a few, mostly recent, some of whom

are mentioned above; nearly all thermodynamics texts written in

the last four decades have been written by nut and bolt engineers,

men more interested in applications than meanings.

Thomson and Clausius would be puzzled if they read a text of

today, for there is hardly a single concept, phrase, or term that

was not known to them over a century ago. Have we made no

progress in the axiomatization of this subject? I think we have

not come far, mainly due to the apathy of a technologically oriented

science. But perhaps through a little concerted effort the axiomat¬

ization of the statics and dynamics of energetics can be completed,

and the subject can be reborn with Rankine’s original expectations

realized. Then the science of energy along with the science of force

can take its rightful place as a part of man’s fundamental

knowledge.

NOTES AND REFERENCES

1. MURRAY, J. A. H. (Ed.) A New English Dictionary on Historical Prin¬

ciples. Claredon Press, Oxford, 1897.

2. NEILSON, W. A., KNOTT, T. A., and CORHART, P. W. (Eds.) Web¬

ster's New International Dictionary. G. and C. Merriam Co., Spring-

field, Mass., 1957

3. However, it should be realized that the first mathematical definition for

the scientific meaning of “work” seems to have been Poncelect’s defini¬

tion for mechanical work in 1826 as force times the distance moved in

the direction of the forces (Millikan, Roller, and Watson, Mechanics,

Molecular Physics, Heat and Sound, M.I.T. Press, Cambridge, Mass.,

p. 65, 1965). Thus the use of the word “work” here refers only to the

abstract concept.

1974]

Balmer — Sketch of Evolution of Energetics

107

4. JOHNSON, S. A Dictionary of the English Language, Second Edition,

London, 1827.

5. The word “energy” in modern French and German is “energie,” and in

modern Italian is “energia.”

6. JOHNSON, S. Op. Cit.

7. MURRAY, J. A. H. Op. Cit.

8. COCKERAM, H. The English Dictionarie (1623). Scolar Press Ltd., Mem

ston, England, 1968.

9. PHILLIPS, E. The New World of English Words (1658). Scolar Press

Ltd., Menston, England, 1969.

10. COLES. An English Dictionary, London, 1696.

11. KERSEY, J. A New English Dictionary (1702), Scolar Press Ltd., Men¬

ston, England, 1969.

12. - . Dictionarium Anglo -Britannicum (1708). Scolar Press Ltd.,

Menston, England, 1969.

13. BAILEY, N. Dictionarium Britannicum (1730). George Olms Verlag,

Hildesheim and New York, 1969

14. SHERIDAN, T. A General Dictionary of the English Language (1780).

Scolar Press Ltd., Menston, England, 1967.

15. APOSTLE, H. G. Aristotle's Metaphysics. Indiana Univ. Press, Blooming¬

ton, pp. 145-159, 1966.

16. WOLFS ON, H. A. Cresca’s Critique of Aristotle. Harvard Univ. Press,

Cambridge, p. 526, 1929.

17. Assyrian: emugu = power; dunnu = power, strength; imuku power,

force.

18. BUDGE, E. A. W. Egyptian Language. Routledge and Kegan Paul Ltd.,

London, 1966.

19. For example, the concepts of potential and kinetic energy can, in very

elementary terms, be considered as height and motion respectively.

20. One of the most modern scientific definitions of energy is that of P. W.

Bridgman {The Logic of Modern Physics. Macmillan Co., New York, p.

Ill, 1927). Like Newton’s “perfectly meaningless” definition of mass as

a quantity of matter, Bridgman would have us take energy as a property

of a material system.

21. DESCARTES, R. Principia Philosophiae. Amsterdam (1644).

22. BLACKWELL, R. J. Isis 57:220, 1966.

23. Later, this concept was quickly Newtonianized to be the mass (vis inertia)

of the body times its velocity.

24. LEIBNIZ, G. W. Acta Eruditorum. Leipzig (1686).

25. DUGAS, R. A History of Mechanics. Editions du Grifton, Neuchatel-

Switzerland, p. 220, 1955.

26. LEIBNIZ, G. W. Specimen Dynamicuum. 1695.

27. DUGAS, R. Op. Cit. p. 221.

28. Also known as vis matrix (force of bodies in motion).

29. RABELO, G. Kant. Oxford Univ. Press, London, 1963, p. 4.

30. D’ALEMBERT, J. Traite de Dynamique. Gauthier-Villars, Paris, 1921.

31. HANKINS, T. L. Isis 56:282, 1965.

32. LANDAN, L. L. Isis 59:131, 1968.

33. SCOTT, W. L. Isis 50:199, 1959.

34. BOLTZMANN, L. Lectures on Gas Theory (Trans, by S. G. Brush). Univ.

California Press, Berkeley and Los Angeles, p. 14, 1964.

35. GAGGIOLI, R. A., SCHOLTEN, W. B., LEWIS, J. L., and OBERT, E. F.

Proc. 4th Southeastern Symp. on Thermal Sciences, Univ. of Tenn.,

Tullahoma, 1968.

108 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

36. THOMSON, W., and TAIT, P. G. Treatise on Natural Philosophy . Clare*

don Press, Oxford, p. 163, 1867.

37. The equation, F = ma was not known to Newton. The precise mathematical

relationship implied by Newton was derived by Euler in 1744, almost

twenty years after Newton’s death (see Truesdell, C. A. Essays in the

History of Mechanics. Springe r-Verlag, New York, 1968)

38. YOUNG, T. Nat. Philos, viii, 1845.

39. RANKINE, W. J. M. Miscellaneous Scientific Papers (W. J. Millar,

editor). Charles Griffin and Co., London, p. 203, 1881.

40. THOMSON, W., and TAIT, P. G. Good Words (Charles Dickens, ed.).

October, 1862.

41. MURRAY, J. A. H. Op. Cit.

42. WHITTAKER, E. A History of the Theories of Aether and Electricity.

Harper and Brothers, New York, p. 6, 1960.

43. PARTINGTON, J. R. A History of Chemistry. Vol. 2, MacMillan, London,

1961.

44. LAVOISIER, A. L. Traite Elementaire de Chimie. 1789.

45. Rumford, with his well planned cannon boring experiments, and Davy’s

experiments of rubbing two pieces of ice together until they melted,

demonstrated that caloric was not carried from one body to another as

a conserved quantity, but that all bodies have an inexhaustible supply

of it.

46. This is the concept of “reversibility.” Carnot, N. L. S., Reflexions sur la

Puissance Motrice du Feu. Bachelier, Paris, 1824.

47. PRESTON, T. The Theory of Heat. London, p. 676, 1919.

48. PERKINS, J. F. in Handbook of Physiology. Sect. 3, Vol. 1, Chap. 1,

American Physiological Society, 1964.

49. MAYER, J. R. Annalen der Chemie und Pharmacie 42:233 (1842).

50. The currently accepted value is 778.26 Btu/ft lbf.

51. COLDING, L. A. Oversigt 8:1, 1856.

52. JOULE, J. P. Philosoph. Mag. 23:263, 347, 435, 1843.

53. THOMSON, W. Proc. Cambridge. Phil. Soc. 1:69, 1848.

54. CLAUSIUS, R. Ann. Phys. 79:368, 500, 1850.

55. THOMSON, W. Trans. Roy. Soc. Edinb. 20:264, 1851.

56. RANKINE, W. J. Op. Cit., pp. 209-228.

57. BADGER, P. H. Equilibrium Thermodynamics . Allyn and Bacon, Boston,

p. 39, 1967.

58. CLAUSIUS, R. Mechanische Wdrmetheorie. Druck and Verlag, Braun¬

schweig, p. 33, 1864.

59. PERKINS, H. A. General Thermodynamics. John Wiley and Sons, New

York, p. 20, 1916.

60. POINCARE, H. Thermodynamics, (G. Carre, ed.) Paris, p. 55, 1892.

61. TOLMAN, R. C. Phys. Rev. 35:904, 1930.

62. GILLESPIE, C. C. The Edge of Objectivity. Princeton Univ. Press, Prince¬

ton, N. J., p. 360, 1967.

63. MIDDLETON, W. E. K. Isis 54:11, 1963.

64. HEATH, T. L. The Works of Archimedes. Macmillan Co., New York, pp.

253-300 1897.

65. MILLIKAN, R. A., ROLLER, D., and WATSON, E. C. Mechanics ,

Molecular Physics, Heat, and Sound. M.I.T. Press, Cambridge, Mass.

1965, p. 297.

66. HOLTON, G. Isis 60:133, 1969.

67. CARATHEODORY, C. Math. Ann. 67:355, 1909.

68. PLANK, M. Treatise on Thermodynamics. Dover Pub., New York, p. viii,

1955.

1974]

B aimer — Sketch of Evolution of Energetics

109

69. ZEMANSKY, M. W. Heat and T her mo dynamics. McGraw-Hill, New York,

p. 173, 1957.

70. TOLMAN, R. C. Phys. Rev. 9:237, 1917.

71. FOWLER, R. H., and GUGGENHEIM, E. A. Statistical Thermodynamics.

Cambridge Univ. Press, New York, 1939.

72. The zeroth Law Statement is: “if an object A is in thermal equilibrium

with B (Ta — Tb), and if A is in equilibrium with C, then B and C

are always in equilibrium with each other” (see: Redlich, 0. Rev. Mod.

Phys. 40:556, 1968).

73. The zeroth Law Statement must be preceded by the words “In a set of

thermodynamic objects S,” in order to conform to the meaning of

equivalence in modern set theory (see, Stoll, R. R. Set Theory and Logic.

W. H. Freeman and Co., San Francisco and London, p. 29, 1963).

74. In the terminology of the zeroth Law Statement these would take the form :

a) Symmetric:

If object A is in thermal equilibrium with object B(TA — Tb), then

object B is in thermal equilibrium with object A (TB — TA).

b) Reflexive:

Every object A in S (see note 73) is in thermal equilibrium with

itself (Ta — Ta).

75. LANDSBERG, P. T. Rev. Mod. Phys. 28:363, 1956.

76. - . Thermodynamics , Interscience Pub. Co., New York, 1961.

77. FALK, G., and JUNG, H. Encyclopedia of Physics. Vol. Ill, Pt. 2,

Springer- Verlag, Berlin, 1959.

78. CALLEN, H. B. Thermodynamics. John Wiley and Sons, New York, 1960.

79. TIZA, L. Ann. Phys. 13:1, 1961.

80. FONG, P. Foundations of Thermodynamics. Oxford Univ. Press, New York,

1963.

81. TRUESDELL, C. A. Modern Developments in the Mechanics of Continua.

(Proc. 1965 Internatl. Conf. on Rheology) (S. Eskinazi, ed). Academic

Press, New York, 1966.

82. - . Irreversible Aspects of Continuum Mechanics , (H. Parkus and

L. Sedov, eds.) . Springer-V erlag, New York, 1968.

83. - - . Rational Thermodynamics. McGraw-Hill, New York. 1969.

84. NOLL, W. Arch. Rat. Mech. Anal. 2:197, 1958.

■

THE ORIGIN OF THE WORD “DOLLAR”—

THE NAME OF OUR UNIT OF ACCOUNT

Edward E. Popp

Port Washington

In order to conduct business transactions without direct barter¬

ing and to be able to record the exchange values of the items and

to express and record the exchange values of all goods, currencies,

and services, it is necessary to have a unit to serve such purposes.

It is necessary to have a common denominator unit which a person

can apply to any item in order to express his idea of the exchange

value of that item.

Because the unit used to express exchange value is used in ac¬

counting, it is called the unit of account. In the United States the

word “dollar” is the name of that unit.

The word “dollar” had its origin in Bohemia in 1519 when a

Count Schlick, a silversmith, who lived in St. Joachimsthal, Bo¬

hemia, made a large silver coin. He gave the coin the name of the

village in which he lived, i.e., he called the coin a St. Joachimsthaler.

Just as Henry Ford called the automobile which he made a Ford.

Because St. Joachimsthaler was a long name, people began to

call the coin a thaler . In other German speaking areas it was called

a taler. Other countries also made a similar coin with a similar

name. In Denmark it was called a daler, in Holland a dalder, in

Italy a tallero, in Poland a talar, and in Spain a dollar.

After Spain discovered large deposits of silver in Mexico, it used

much of that silver to make silver dollar coins. These coins were

used as currency for the vast trade which took place between the

Spanish American colonies and the English American colonies.

It so happened at that time that the Spanish American colonies had

an unfavorable balance of trade with the English American colo¬

nies. The result was that a large number of the Spanish silver

dollar coins passed into English American hands and circulated as

the predominant currency in the English American colonies.

As time passed the English Americans began to express the

exchange value, i.e., the price of goods and services in dollars instead

of in English pounds, as was their previous custom.

When the exchange values of goods and services were expressed

and recorded with the word “dollar” rather than with the word

“pound,” the word took on an additional meaning. It took on the

meaning of the word “pound.” The word “dollar” was still used

111

112 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

as the name of the large Spanish silver coin, but it also was used

as the name of a unit with which to express exchange value.

By the time the English American colonies became independent

from England the word “dollar” was so well established in its

usage that on July 6, 1785, the United States Congress made the

word, dollar, the official name of the unit of account which was to

be used in the United States. The Congress called it the official

monetary unit, perhaps because the Congress later intended to

authorize the minting of a silver coin with the exchange value equal

to the value expressed by the dollar unit already in use.

That took place in 1792 when the Congress authorized the mint

to make the first silver dollar coins. The price of silver at that

time was $1.29 per troy ounce. With the silver at that price it was

determined that 371.25 grains of silver had an exchange value of

one dollar.

The government declared the coin to be legal tender for a one

dollar payment of public and private debts. The coin then had two

exchange values :

1. The exchange value of the 371.25 grains of silver.

2. The one dollar legal tender value.

Both values were equal at that time, but they did not remain

equal. Between 1794 and 1874 only a small number of silver dollar

coins .were minted because the exchange value of 371.25 grains

of silver was above the legal tender value of the coin.

However, in 1874 large deposits of silver were discovered in

Nevada. The result was that the market value of the 371.25 grains

of silver in the coin became less than the one dollar legal tender

value of the coin. Generally speaking, from 1876 to 1964 the market

price of the silver in the silver dollar coin was less than one dollar.

At times it was less than fifty cents. With the price of silver today

(April 28, 1973) at about $2.25 per ounce the market value of the

371.25 grains of silver may be about $1.75.

We mention these historical facts just to show that the exchange

value of the dollar coin is not now and has not been for many years

equal to the amount of exchange value expressed by the unit of

account called dollar. In other words, the dollar coin and the unit

of account called dollar are two different things.

Let us illustrate the point we are trying to make by comparing

the dollar unit used to express exchange value, i.e., the unit of

account, with a unit we use to express volume. Let us say, the

gallon. If a person says that he has a gallon jug, the word “gallon”

is used as an adjective. It describes the noun, jug. If a person says

that he has a gallon of water, the word “gallon” is used as a noun.

It is the name of a unit we use to express a certain volume. A per-

1974]

Popp — Origin of the Word “ Dollar ”

113

son does not say, I have one gallon, because a gallon as a unit to

express volume does not exist as a physical thing. The unit called

gallon exists only as a concept.

The same is true for the unit we call a dollar. When the word

“dollar” is used as the name of the concept or device we use as

a common denominator unit with which we express our idea of the

exchange value of goods, currencies, and services, it is used as a

noun. When the word “dollar” is used to describe a Federal Reserve

note or a coin, it is used as an adjective.

A close study will teach us that the dollar unit of account by

itself has no exchange value, because it is not a physical thing. It

is a unit concept. It has meaning only when it is applied to some¬

thing. Just as the word “gallon” has no meaning unless it is applied

to something. A person cannot buy a gallon. He cannot carry a gal¬

lon. He cannot store a gallon. But a person may buy, carry, or store

a gallon of something.

In like manner a person does not receive dollars from a bank.

He does not carry dollars in his pockets. He does not store dollars

in his safe. But a person may receive, carry, or store dollars’ worth

of currency (coins or Federal Reserve notes), because items of cur¬

rency are physical things. They are not concepts.

Also, when we hear the expression, “the dollar lost value,” we

should know that the statement does not tell us what is meant.

We know that the dollar unit did not and could not lose value,

because the unit being a concept has no exchange value to lose.

What is meant by the expression, “the dollar lost value,” is that

the currency, the exchange value of which is expressed in dollar

units, lost exchange value.

The question might be asked, for what purpose is the dollar unit

used? The dollar unit was used and is used to express the ratio of

the exchange value of one item with the exchange value of another

item. For example, a dollar bill has a specific amount of exchange

value, one dollar’s worth, when it is used as a payment for taxes.

But when it is used to buy goods or services it will have the amount

of exchange value to which the buyer and seller mutually agree at

the time of each transaction.

As an illustration, let us say, a person offers to sell his potatoes

for a one dollar bill per bushel; the one dollar bill will have the

exchange value of one bushel of potatoes. If at a later date, he offers

to sell his potatoes for two one dollar bills per bushel, each dollar

bill will then have the exchange value of only one-half bushel of

potatoes. The dollar unit was used to express the two different

ratios of the exchange values of the dollar bills and the potatoes.

114 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

We should note that the exchange values of both, the dollar bills

and the bushel of potatoes, changed from the first transaction to

the second transaction. The dollar unit was used to express the

different ratios of exchange values of the dollar bills and the bushel

of potatoes at both transactions.

When the people in the early days of our country decided to use

the amount of the exchange value of the silver dollar coin as the

amount of exchange value to be expressed with the dollar unit,

the unit expressed the specific amount of exchange value that 371.25

grains of silver had at that time.

The silver dollar coin was intended to be the embodiment of the

concept unit called a dollar. It remained so only so long as the price

of silver did not increase or decrease. When the price of silver

changed the unit called a dollar might just as well have been called

a plain unit or a point. Because it served only as an abstract unit

with which the exchange value of one item was expressed in its

relation with the exchange values of other items.

However, the word “dollar” continued to be used as the name of

our unit to express exchange value. So the exchange values of all

goods and services are expressed in the ratio of the exchange value

of each item with the exchange value of each other item with the

abstract unit called a dollar.

Once the ratio of the exchange value of each item is well estab¬

lished in its relation to the exchange values of other items, the

exchange values of all items including items of currencies, could

be expressed with an abstract unit called a point, just as well as

with the unit called a dollar.

In other words, we could say that a bushel of potatoes has the

exchange value of one point's worth of the currency and that one

point's worth of the currency has the exchange value of one bushel

of potatoes. If we did that, then it would be easy to understand

that our common denominator unit, that is, our unit of account,

is an abstract unit without exchange value in itself.

HISTORY OF BIOLOGICAL CONTROL ATTEMPTS

AGAINST INSECTS AND WEEDS IN WISCONSIN

J. W. Mertins and H. C. Coppel

University Wisconsin — •

Madison

Biological control may be defined generally as the directed use

of biotic agents, or their products, for the suppression of living

organisms detrimental to man.

The procedure has a long history, dating back to the domestica¬

tion of the cat for rodent control by the ancient Egyptians. Al¬

though sporadic trials and experiments were undertaken in the

interim, it was not until 1888 that biological control, as we know

it today, was established as a valid method of pest suppression.

The importation of the vedalia beetle, Rodalia cardinalis (Mulsant) ,

from Australia to California for the control of the cottonycushion

scale, I eery a purchasi Masked, a citrus pest, is considered the turn¬

ing point in our thinking about insect control, and is one of the

classical stories in biological control (Doutt, 1964).

The following history of biological control work in Wisconsin,

includes attempts against 1 weed and 10 insect pests (listed alpha¬

betically) , and several lesser case histories. The resulting overview

is a compendium of scattered published accounts and a large body

of previously unpublished data from the files of our colleagues.

Cirsium arvense (L.) Scopoli

CANADA THISTLE

The Canada thistle is an introduced noxious weed, naturalized

to North America from Eurasia. It was established first in Canada

by early 17th century settlers, who probably transported it in con¬

taminated seed (Gilkey, 1957) . Shortly thereafter it was discovered

in New England, possibly an independent introduction, and has

since spread throughout the northern half of the United States and

in Canada from Quebec to British Columbia (U.S. Department

of Agriculture, 1971). The most serious problems occur in the Lake

States, Great Basin, and Pacific Northwest. Worldwide, (7. arvense

is rated as one of the most serious threats to crop production. Ef¬

forts toward biological control of this thistle have occurred chiefly

in Canada (Peschken, 1971). The most conspicuous natural enemies

already occurring there were identified as a leaf bettle, a weevil, a

115

116 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

fruit fly, and a rust disease, none of which hold the plant below

economic densities. Biological control workers found 80 insect

species feeding on C. arvense in Europe (Zwolfer, 1965). A leaf

beetle, Altica carduorum Guerin-Meneville, a weevil, Ceutor-

hynchus litura (F.), and a fruit fly, Urophora cardui L., were

selected for further study because of their apparent host specificity

and climatic suitability. The 2 beetles were subsequently released,

but the gall-forming fly is still under study in laboratory culture.

Colonies ranging in size from 24 to 1,160 individuals of A. cardu¬

orum were released in Ontario, British Columbia, Nova Scotia,

and Alberta during 1963-1968 (Peschken, 1971). Permanent estab¬

lishment apparently did not occur, although a small colony survived

for 3 years at Lacombe, Alberta, before disappearing. Establish¬

ment was probably prevented by heavy predation of the immature

stages and excessive adult dispersal. Colonies of 22-150 adult

C. litura were released at 4 sites near Belleville, Ontario, in 1965

and 1967. It appears that establishment has occurred on only 1 site,

a heavily thistle-infested pasture, where 230 beetles were released

in 1967.

The release of A. carduorum against Canada thistle was the

first attempt at biological weed control in Wisconsin (Wisconsin

Department of Agriculture, 1968a) . Approximately 200 adult bee¬

tles were caged for 1 week over a thick patch of thistles in a pasture

near Rewey in southwestern Iowa County. The cage was subse¬

quently removed, but periodic recovery attempts in the area have

been unsuccessful (Wisconsin Department of Agriculture, 1969),

perhaps due to excessive dispersal or unsuitable climate.

Coleophora laricella (Hiibner)

LARCH CASEBEARER

The larch casebearer, a European defoliator of larches, occurs

throughout the range of its host in eastern North America and in

localized infestations in the West. Although about 70 insect parasite

species have been reared from C. laricella in North America, serious

outbreaks and damage were prevalent until the introduction of

several European parasites (Sloan, 1965). In eastern North

America biological control has progressed to where only minor

damage occurs, but western populations apparently still lack effec¬

tive natural enemies, and heavy defoliation, resulting in radial

growth loss, is common.

Biological control of larch casebearer in Wisconsin is one of the

most successful of such attempts to date. Casebearers were first

recorded from Wisconsin by MacAloney (1939). Wisconsin Conser¬

vation Department (now Department of Natural Resources) annual

1974] Martins and Coppel — History of Biological Control 117

forest insect surveys indicated heavy populations during the period

1951-1956. In the period 1956-1964 populations were reported as

“very light to light'’ statewide, and since 1964, the survey has not

reported on (7. laricella. The reduced importance of C. laricella as

a pest of larch is traceable in all likelihood to the introduction of

2 species of parasites in August, 1953. A total of 189 male and 158

female Agathis pumila (Ratzeburg), a braconid, and 208 male and

235 female Chyrsocharis laricinellae (Ratzeburg), a eulophid, were

released at 2 sites in northcentral Wisconsin. About half of the total

number of each species was released in the Hugo Sauer Nursery

near Rhinelander, and the other half in a stand of 35-foot tamarack

at the west end of Thunder Lake Swamp, 1 mile west of Three

Lakes. These parasites originated from the Dominion Parasite

Laboratory, Belleville, Ontario, Canada. A smaller, indeterminate

number of parasites originating from casebearer hosts collected in

Michigan had also been released at these sites earlier in the season

(R. D. Shenefelt, personal communication). Both species became

established, and through capture/rerelease and natural dispersal,

have spread over the entire state. Sloan (1965) estimated the aver¬

age annual spread of A. pumila after its introduction as equivalent

to over 29 miles/year. Within 3 years of the first introductions

populations of larch casebearers had dropped to subeconomic levels

statewide and have remained essentially so ever since.

Of the two species, A. pumila is the most important, accounting

for 31-98% of the total parasitization in various localities studied

(Sloan and Coppel, 1965). C. laricinellae usually accounts for less

than 5% of the parasitized hosts. Parasitization resulted in 19-68%

host mortality in the various collections studied.

The parasites, C. laricinellae and A. pumila, were also the sub¬

jects of trial exportation from Wisconsin in a cooperative agree¬

ment with R. B. Ryan, U.S. Forest Service, Corvallis, Oregon. In

June, 1972, laboratory rearings of field collected casebearers

produced 45 C. laricinellae and 77 A. pumila for shipment. The

survivors of the insects shipped were to be used for laboratory

propagation, and the subsequent progeny released against case¬

bearers in the Pacific Northwest.

Diprion similis (Hartig)

INTRODUCED PINE SAWFLY

The introduced pine sawfly first appeared in North America in

a Connecticut nursery in 1914, and probably arrived on ornamental

stock imported from Holland (Rowher, 1916). It is now distributed

through most of the range of its preferred host, Pinus strobus L.,

in eastcentral North America. The heaviest populations and most

118 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

serious defoliation occur at present in the Lake States, particularly

Wisconsin and Minnesota. Few, if any, parasite importations were

made specifically against D. similis, but benefits have accrued from

releases made against associated sawfly species, and some recoloni¬

zations of parasites within the continental area have been directed

against this species.

Mertins and Coppel (1971a & b) reported 30 species of insect

parasites reared from D. similis cocoons in Wisconsin. Five of these

account for 94.3% of the parasitization, and all but 1 of the 5 are

of European origin. The hymenopterous parasites, Monodontomerus

dentipes (Dalman), Dahlbominus fuscipennis (Zetterstedt) , and

Eupelmella vesicularis (Retzius), were associated with Wisconsin

sawfly populations almost from the time of their discovery in 1944,

and probably moved into the state with the host. It also appears

that these species entered North America accidentally prior to later

purposeful introductions. In August, 1953, 3,500 Diprion sp.

cocoons parasitized by D. fuscipennis were obtained from Belleville,

Ontario, and set out at 3 widely separated points in Wisconsin by

R. D. Shenefelt. Although the parasites were liberated against 3

other sawfly species, it is probable that some individuals attacked

D. similis cocoons as well. Both D. fuscipennis and M. dentipes have

been reared from field collections of D. similis cocoons, and re¬

colonized within the state to areas where they were less abundant.

L. vesicularis, which frequently functions as a hyperparasite, has

not been relocated to new areas. The fourth European species,

Exenterus amictorius (Panzer), a wasp which attacks D. similis

in Wisconsin, was originally brought to North America by the

Canadian government (McLeod et ah, 1962). It has never been

released in Wisconsin, but apparently has spread here from release

points in southern Ontario. Mertins and Coppel (1968) documented

the arrival and increasing abundance of E. amictorius, and its rela¬

tionship with M. dentipes. These 2 species account for 87 % of the

annual mortality caused by parasites and are the most important

species present. The suggestion (Mertins and Coppel, 1968) that

E. amictorius might displace M. dentipes as the principal parasite

of D. similis has now proven itself in fact. E. amictorius is now by

far the most prevalent parasite reared from cocoons of the first

sawfly generation, although M. dentipes continues to account for

slightly greater mortality in the second (overwintering) genera¬

tion. At least 11 hymenopterous species have been reared hyper-

parasitically from E. amictorius in Wisconsin (Mertins and Coppel,

1973), and these may possibly reduce the effectiveness of the

species below its potential. After establishing the presence of an

effective larval parasite (E. amictorius) and an effective cocoon

parasite (M. dentipes), Coppel (1962) and Mertins (1967) sug-

1974] Mertins and Cojjpel — History of Biological Control 119

gested introduction of an egg parasite, as well. The European

eulophid, Dipriocampe diprioni (Ferriere) might be a likely candi¬

date.

The parasites, E. amictorius and M. dentipes, were also the

subjects of exportation from Wisconsin in a cooperative effort

with A. T. Drooz, U.S. Forest Service, Research Triangle Park,

N.C. In the summer of 1969 adult E. amictorius were field collected

in Wisconsin and shipped to North Carolina for release; forty-eight

parasites were released near Elizabeth City against a pine sawfly,

Neodiprion excitans Rohwer. Between 1970 and 1972, laboratory

rearings of field collected D. similis cocoons produced 1,900

E. amictorius and 26,350 M. dentipes for shipment. The survivors

of the insects shipped were released in North Carolina, Virginia,

and Florida against the following pine sawflies : Neodiprion pratti

pratti (Dyar), N . excitans, N . hetricki Ross, and N. lecontei

(Fitch) .

Research on the sex pheromone of the introduced pine sawfly

as a potential tool for biological control has also centered in Wis¬

consin (Coppel et ah, 1960; Casida et ah, 1963; Jones et ah, 1965;

Mertins, 1971), and work on chemical identification of the sub¬

stance is progressing (Jewett, 1971). A mathematical model

describing the potential effects of pheromone trapping on sawfly

populations was devised by Mertins (1971). It predicts the theo¬

retical annihilation of an isolated sawfly population in 3-4 genera¬

tions of trapping. Experimental traps are under development and

a pilot study is planned to test the feasibility of the technique in

the field.

Hyper a postica (Gyllenhal)

ALFALFA WEEVIL

The alfalfa weevil was probably introduced to the New World

from southern Europe about 1900. Injury was first detected near

Salt Lake City, Utah, in 1904, and the insect has since spread to

most of the alfalfa growing areas in the West. It is also now widely

distributed in the East, and is considered the most important insect

enemy of alfalfa (Metcalf et ah, 1962) . Clausen (1956) reviewed

the early beneficial insect introductions which began in 1911. Of

more than a dozen species introduced, only Bathyplectis curculionis

(Thomson) , an ichneumonid larval parasite, became established

in sufficient degree to be effective. It is reportedly found throughout

the range of H. postica in the West, and in most areas attains a

high rate of parasitization, frequently over 90%. With the weevil’s

invasion of the eastern United States in 1951 came renewed interest

in importation of natural enemies, and several additional parasite

species were introduced (Brunson and Coles, 1968). In the East,

120 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

B. curculionis again appears to be a significant mortality factor,

but other species also contribute additional increments.

The alfalfa weevil was first discovered in Wisconsin in 1966 in

Kenosha County. It was considered a severe threat to alfalfa

production in the state, which ranks first in the nation in alfalfa

acreage (about 3 million acres) . Indirectly, the state dairy industry

was thus threatened. B. curculionis moved into the state along with

the host, and was first collected in 1968 emerging from 8% of the

weevils collected (Wisconsin Department of Agriculture, 1968b).

The abundance of B. curculionis varies from area to area, but its

distribution generally coincides with that of the weevil, and “it has

probably been responsible for slowing the increase of alfalfa weevil

populations in Wisconsin” (Wisconsin Department of Agriculture,

1971b). Parasitization reached 62% in some fields in the southern

tier of counties in 1971, where the highest weevil populations have

been found. Despite the presence of the parasite, alfalfa weevil

now occurs in all except the 13 northernmost counties of the state.

In supplement to the naturally occurring B. curculionis , J. W.

Apple and J. A. Litsinger released 691 additional parasites in

Kenosha and Rock Counties in 1969 and 1970. The parasites were

provided by the U.S. Department of Agriculture, Moorestown, N.J.,

which, along with personnel at Wooster, Ohio, also provided 6 other

hymenopterous species for release against the weevil in Wisconsin :

2,578 Microctonus aethiops (Nees) in Kenosha, Rock, Dane and

Sauk Counties, 1969-1972; 41 M. colesi Drea in Kenosha County,

1969; 190 M. stelleri Loan in Rock County, 1970; 1,603 Tetrastichus

incertus (Ratzeburg) in Kenosha and Columbia Counties, 1967 and

1972, respectively; 1,256 Bathyplectes anurus (Thomson) in Keno¬

sha, Rock, and Dane Counties, 1969-1971; and 431 B. stenostigma

(Thomson) in Kenosha and Rock Counties, 1969-1970. Despite

these releases, the only parasite known to be established on the al¬

falfa weevil in Wisconsin is B. curculionis . Recently a secondary

ichneumonid parasite, Gelis sp., has been reared in very small

numbers from Wisconsin B. curculionis . At present it is of little

consequence, but its occurrence bears watching because of potential

detrimental effects it may have on the primary parasite population.

Mortality to B. curculionis eggs through host encapsulation has

occurred in the western states, but it has not been found in Wis¬

consin to date.

Neodiprion lecontei (Fitch)

REDHEADED PINE SAWFLY

The redheaded pine sawfly is one of the most important native

forest insects defoliating young hard pines in eastern North

America (Benjamin, 1955). During the 19th century the insect

1974] Merlins and Coppel — History of Biological Control 121

was only of minor significance on park and shade trees, but with

the advent of extensive pure pine plantations in recent years, many

local and widespread outbreaks have occurred. The saw fly has been

recorded from nearly every state east of the Mississippi River,

adjacent Canada, and the 5 states adjacent to the Mississippi on

the west. Benjamin (1955) listed 58 species of parasitic and preda¬

tory insects known to be associated with N. lecontei in eastern

North America, but remarked on the lack of emphasis on employing

biological agents in population management of this sawfly. Releases

of only 2 parasite species have been directed against N. lecontei in

the United States (Dowden, 1962). In 1932, 120 Drino bohemica

Mesnil (Diptera: Tachinidae) were liberated in Pennsylvania; no

recoveries were made. From 1940 to 1946 more than 1.3 million

Dahlbominus fuscipennis were released in Alabama, Tennessee,

New York, and Michigan. Few recovery attempts were made, but

D. fuscipennis was established on N. lecontei in Lower Michigan.

To our knowledge, the only biological control attempt against

N. lecontei in Wisconsin took place in August, 1953, when 2,600

sawfly cocoons parasitized by D. fuscipennis were set out against

it (and the larch sawfly, Pristiphora erichsonii) in Burnett County

just west of McKenzie Lake. Labeled specimens in the U.W. Insect-

arium confirm that D. fuscipennis is established on the redheaded

pine sawfly in Wisconsin.

Neodiprion sertifer (Geoffrey)

EUROPEAN PINE SAWFLY

The European pine sawfly was first discovered in New Jersey in

1925. Currently its range extends through scattered infestations

in most all the states between New Jersey and Maine on the east

and Iowa and Missouri on the west, including the southeastern

portion of Ontario as well. The Canadian government has released

large numbers of parasites against the sawfly, and also developed

the use of an effective polyhedral virus disease introduced from

Europe. Colonies of the parasites as well as the virus were obtained

from Canada and released in the United States, particularly in New

Jersey where severe defoliation was observed (Dowden, 1962). Of

7 species liberated, 3 Hymenoptera are established : Dahlbominus

fuscipennis , Exenterus abruptorius (Thunberg) , and Pleolophus

basizonus (Gravenhorst) .

The nature of the first recorded appearance of N. sertifer in

Wisconsin is shrouded in uncertainty (D. M. Benjamin, personal

communication). A single larva determined as this species by

several authorities was discovered amongst a group of Neodiprion

nanulus Schedl larvae collected near Arkdale (Wood County) in

122 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

1955. No other specimens were found at that time or later, but to

offset the potential threat of infestation, approximately 23,000

D. fuscipennis were released, the area was treated with a quantity

of the polyhedral virus obtained from the U.S. Forest Service in

Columbus, Ohio, and sprayed with DDT. It is probable that the

parasite release was useful, insofar as D. fnscipennis was estab¬

lished on N. nanulus. In the spring of 1972 a verified infestation

on N. sertifer was discovered in southern Walworth County, and

parasite or virus releases are contemplated for 1973.

N eodiprion swainei Middleton

SWAINE JACK PINE SAWFLY

The Swaine jack pine sawfly has been one of the most important

pine-infesting sawflies in eastern Canada for many years. It was

first noted in the Lake States in the early fifties, and is now widely

distributed in Upper Michigan, northcentral Minnesota, and Wis¬

consin. The Canadian government released 2 parasites, the ichneu-

monid, Pleolophus basizonus, and the tachinid, Drino bohemica,

against N. swainei, and both have been recovered (McLeod and

Smirnoff, 1971). Two other species, Exenterus amictorius and

Dahlbominus fuscipennis, released against the European spruce

sawfly, Diprion hercyniae (Hartig), have also been recovered from

N. swainei . None of these parasites occur in large numbers. Smir¬

noff (1967) described the use of a polyhedral virus in extensive

field tests in Canada. The virus was very effective and persisted

in the population for several years.

The sawfly has occurred in outbreak conditions in southwestern

Wisconsin, and one parasite release against it was made by R. D.

Shenefelt in August, 1953. Six-hundred sawfly cocoons parasitized

by D. fuscipennis were set out in an infested jack pine stand 1 mile

west of Arena in Iowa County. The parasite became established,

but before its effectiveness was determined the sawfly population

collapsed.

Ostrinia nubilalis (Hiibner)

EUROPEAN CORN BORER

The European corn borer was probably introduced accidentally

to North America about 1909 in broomcorn shipped from Italy or

Hungary (Metcalf et ah, 1962). Since the time of its first detection

near Boston, Massachusetts, in 1917 it has become one of the most

destructive insect pests of corn. The present distribution of the

corn borer covers practically all the major corn-growing areas of

the United States, including all of the states north of Florida,

Louisiana and Oklahoma, and east of Montana, Wyoming and

Colorado. Parasite importation investigations were begun almost

1974] Mertins and Coppel — History of Biological Control 123

immediately by the U.S. government, and between 1920 and 1938

approximately 3 million individual parasites were imported from

Europe and the Orient, representing 24 species of Diptera and

Hymenoptera. Although 6 or 7 of these species became permanently

established in the Northeast where they were released, only 2, the

tachinid, Ly della thompsoni Herting, and the braconid, Macrocen-

trus grandii Goidanich, became sufficiently abundant or widely

distributed to be of appreciable value (Clausen, 1956). M. grandii

is the dominant species in New England where parasitization up

to 52% has been observed, and L . thompsoni has accounted for

mortalities of 10-45% in the East and 45-75% in the northcentral

states.

As the borer continued to spread westward across the corn belt

during the 1930’s and 1940’s, entomologists continued to move its

parasites into the new areas as well. The insect was first discovered

in Wisconsin near Sheboygan in 1931, and beginning in 1942 para¬

sites provided by the U.S. Department of Agriculture were released

in the state by the Wisconsin Department of Agriculture and the

University of Wisconsin, Department of Entomology. The first year

about 3,300 individuals were released, over half of which were

M. grandii. During the 8 year period 1942-50, over 109,800 para¬

sites of 5 different species were imported from the Northeast and

released in Wisconsin against the borer (Wisconsin Department

of Agriculture, 1971a). More than 80% of these were M. grandii,

but also included were L. thompsoni, and the hymenopterans,

Horogenes punctorius (Roman), Sympiesis viridula (Thomson),

and Chelonus annulipes Wesmael. Recoveries were subsequently

made of all 5 parasite species, but C. annulipes apparently did not

become permanently established. The dipteran, L. thompsoni, for

several years occurred in large numbers, reaching a peak parasiti¬

zation rate of 42 % in 1950, and a lesser peak of 20% in 1958.

However, by 1964 it had disappeared from fall corn borer surveys

in Wisconsin as well as in other midwestern states, and it has not

been recovered in significant numbers since. The status of S. viri¬

dula is currently unsure because it is usually not encountered in

survey work due to its host selection habits. It is believed to occur

in low numbers. H. punctorius and M. grandii are currently the

most frequently encountered introduced parasites in Wisconsin,

accounting for parasitization of 2-5 % between them on a statewide

basis. Parasitization is highest in the southwest area where it gen¬

erally reaches 12-15%. It is also interesting to note that M. grandii

was not recovered in significant numbers on a consistent basis until

1959, 9 years after it was last released in the state (J. W. Apple,

unpublished data).

Although the introduced parasites established in Wisconsin on

124 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

the European corn borer do not account for a great deal of mor¬

tality, they add to that caused by several of the 30 species of

recorded native parasites. The most important of these in Wisconsin

are the tachinid flies, Aplomya caesar (Aldrich) and Pyraustomyia

penitalis (Coquillett) . Other natural mortality factors aiding in

reduction of corn borer populations and infestations are a protozoan

disease organism, Perezia pyraustae Paillot, and numerous preda¬

tors including coccinellids, chrysopids, nitidulids, and a number of

birds, especially the woodpeckers (Baker et ah, 1949).

Pristiphora erichsonii (Hartig)

LARCH SAWFLY

The larch saw fly is apparently a true Holarctic species as its

range follows most of the circumpolar distribution of its host trees

in the genus Larix. However, the insect was not reported in North

America before 1880 when it was discovered in Massachusetts

(Drooz, 1960), and the nature of its distribution before that time

is unsure. Many widespread and serious outbreaks have occurred

since 1880 covering virtually all of the Canadian provinces and

the northern tier of states where larches grow. The highly success¬

ful ichneumonid parasite, Mesoleius tenthredinis Morley, against

the larch sawfly was introduced to Canada from England in

1910-11. By 1950, parasitization rates of up to 90% were recorded

and average rates were 55-75%. A few small releases of

M. tenthredinis were made in the U.S. in Michigan, Minnesota,

New Hampshire, and Massachusetts, but it is unlikely that they

affected the spread of the parasite (Dowden, 1962). However,

M. tenthredinis has spread effectively into the Lake States and

New England from Canadian releases (Drooz, 1960). Unfortu¬

nately, the importance of M. tenthredinis has decreased greatly in

central Canada and the Lake States ; an immune reaction has been

developed by the host against the parasite egg, resulting in encapsu¬

lation and prevention of development.

The natural spread of M. tenthredinis into Wisconsin resulted

in reported parasitization levels of about 40% by 1935 (Drooz,

1960), but by 1954 the rate of effective parasite attack was reduced

to 3% or less by encapsulation. The only active biological control

attempt against larch sawfly in Wisconsin was carried out by R. D.

Shenefelt in August, 1953. Sawfly cocoons parasitized by Dahl -

bo minus fuscipennis were obtained from Canada and set out in

two areas: about 2,600 cocoons 1 mile north of Solon Springs in

Douglas County, and about the same number in Burnett County

just west of McKenzie Lake, in an area also infested by Neodiprion

lecontei.

1974] Mertins and Coppel — History of Biological Control 125

Scolytus multistriatus (Marsham)

SMALLER EUROPEAN ELM BARK BEETLE

The smaller European elm bark beetle is an introduced pest of

elms first observed near Boston, Massachusetts in 1909 (Baker,

1972). Were it not for the fact that it is the major vector of the

Dutch elm disease fungus, Ceratocystis ulmi (Buisman) C. Moreau,

the beetle would be of little concern, for it usually breeds only in

weakened or unhealthy trees. However, the beetle-fungus associa¬

tion has spread over most of the United States and southern

Canada. The disease is also present in areas beyond the distribution

of the introduced vector where it is transmitted by a native elm

bark beetle. Six parasite species were known from S. multistriatus

in North America before any importations were attempted (Bush¬

ing, 1965). Although insufficient in numbers to control the beetle,

3 of the species are fairly common, and 2 of these, Cheiropachus

colon (L.) and Entedon leucogramma (Ratzeburg), are European

hymenopterous parasites established in the United States at an

early date by unknown means (Kennedy, 1970). In September,

1964, the first shipment of a braconid parasite, Dendrosoter

protuherans (Nees), was received from France by the U.S.D.A.

Laboratory colonies were maintained at Michigan State University,

East Lansing, and the Northeastern Forest Experiment Station,

Delaware, Ohio, and in 1966 releases were made on field plots in

Ohio and Missouri ; the parasite was apparently successfully estab¬

lished (Kennedy, 1970), and a small population is also established

in Detroit, Michigan (Kennedy, personal communication).

The first recorded occurrence of S. multistriatus in Wisconsin

was in 1952 (Wisconsin Department of Agriculture, 1956), but it

had probably been present for several years. Dutch elm disease was

not found until 1956. The previously mentioned common parasite

species have also been found, but have not adequately controlled

bark beetle populations in Wisconsin. Consequently, in July, 1967,

Milwaukee County Forestry Department personnel in cooperation

with U.S.D.A. entomologist B. H. Kennedy released a total of 248

male and 609 female D. protuherans on 5 beetle-infested parkland

trees. Samples taken from 3 of the trees in August, 1967, indicated

initial establishment of the parasite with a recovered : released

ratio of nearly 1:1 (Kennedy, personal communication). However,

the following year a stream channelization project almost com¬

pletely destroyed the release area, and samples from 2 of the small

trees remaining failed to produce D. protuherans (Kennedy, per¬

sonal communication). It is not known if the parasite established

itself beyond the initial release site.

126 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Synanthedon pictipes (Grote and Robinson)

LESSER PEACH TREE BORER

The lesser peach tree borer is a native insect pest of peach,

cherry, and plum trees. Although most common in the South, it

ranges from southern Canada to the Gulf of Mexico, and from the

Great Plains east to the Atlantic coast (Gilbertson, 1934). Branch

and tree mortality frequently occurs when large larval populations

girdle the affected parts. Several native parasites, a fungal disease,

and various insect and bird predators account for some natural

mortality, but borer populations are seldom held in check by these

agents because of their well protected habitat (King, 1917).

Although peaches and plums are not raised commercially in

Wisconsin, sour cherries are a major orchard crop in the state, and

S. pictipes may well be the most important pest of cherry in com¬

mercial plantings (Koval, personal communication). In 1969 per¬

sonnel of the U.S.D.A. Deciduous Fruit Insect Investigations Labo¬

ratory at Vincennes, Indiana, began initial surveys on Washington

Island, Door County, Wisconsin, to determine the feasibility of

suppressing borer populations with a sex pheromone trapping pro¬

gram (Wong et ah, 1971). Extensive field work was begun in 1970

in cooperation with C. F. Koval and M. G. Karandinos, Department

of Entomology, University of Wisconsin, and by 1972 a trapline

of about 3,000 virgin female-baited traps was in operation continu¬

ously during the flight season. After the 1972 trapping program

it was estimated that a population reduction of 42% had been

attained (Koval, personal communication). Plans call for continua¬

tion of these studies, and details are being worked out for the

addition of a proposed concurrent sterile male release program.

Several other programs are in progress or under consideration

utilizing the unique advantages of Washington Island’s relatively

isolated insect populations. In 1971, a trapline of 18,000 traps was

maintained by U.S.D.A. personnel to test and compare effectiveness

of synthetic pheromones and virgin females of the codling moth,

Laspeyresia pomonella (L.) . Additional similar investigations in¬

volved the redbanded leaf roller, Argyrotaenia velutinana (Walker),

and a sterile male release program against the horn fly, Haematobia

irritans (L.) , is scheduled for 1973.

MISCELLANEOUS EFFORTS

In addition to the preceding documented case histories, a number

of minor or little known efforts and suggestions for biological con¬

trol have been made in Wisconsin. A few of these have come to our

attention and are included.

1974] Mertins and Cojjpel — History of Biological Control 127

In 1971, C. F. Koval of the Department of Entomology reared

a number of specimens of the eulophid parasite, Encarsia formosa

Gahan, from greenhouse whiteflies, Trialeurodes vaporariorum

(Westwood), collected in a University of Wisconsin greenhouse.

This was the first record of the parasite in Wisconsin, and its ac¬

tivities in the greenhouse continue to be monitored. Although

T. vaporariorum does not survive Wisconsin winters outside, it does

build up large populations during the summer months in orna¬

mental and floral gardens replanted each year from greenhouse

stock. Therefore, an undetermined number of E. formosa were

recolonized from the greenhouse to a horticultural garden on

campus where the whitefly was a problem. Recovery collections

made later in the summer showed establishment of the parasite at

a parasitization rate of about 25-30%.

A number of studies have been carried out in Wisconsin on the

use of microbial pathogens for insect control, with varying results.

Walgenbach and Benjamin (1965) field tested the bacterium Bacil¬

lus thuringiensis Berliner (Thuricide® 90T) for suppression of

pine tussock moth larvae, Dasychira plagiata (Walker), on jack

pine. The treatment caused no significant reduction in numbers of

5th-6th instar larvae against which it was applied. In at least one

instance, however, practical recommendations providing for the

use of such materials on a commercial basis have resulted (Libby

and Wade, 1971). Certain commercially available preparations

of B. thuringiensis var. alesti successfully suppressed populations

of cabbage caterpillars, including Pieris rapae (L.) , Plutella

xylostella (L.) , and Trichoplusia ni (Hiibner) , under Wisconsin

conditions (Libby and Chapman, 1971). J. W. App)e (personal com¬

munication) found that 4 applications of Dipel®, a commercial

preparation of B. thuringiensis, provided 80% protection of sweet

corn from European corn borer in Wisconsin, compared to 97%

protection with the recommended treatment of 4 applications of

Sevin® insecticide.

Finally, we must include mention of the many dedicated amateur

naturalists, organic gardeners, and other concerned citizens who

have attempted .to reduce their use of pesticides around the home

garden and farm by releasing unknown numbers and kinds of insect

parasites and predators obtained from any of at least 20 commer¬

cial insectary sources. For example, records of only one such com¬

pany, Rincon-Vitova Insectaries, Inc. of Rialto, California, indicate

shipment of the following insects to Wisconsin during the last 5

years: 58,000 eggs of the green lacewing, Chrysopa carnea

Stephens; 528,000 Tricho gramma spp. egg parasites; 25,000

hymenopterous parasites including Tachinaephagus zealandicus

Ashmead, Spalangia endius Walker, and Muscidifurax raptor

128 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Girault and Sanders for pestiferous flies; 200 of the lady beetle,

Cryptolaemus montrouzieri Mulsant; and 200 convergent lady bee¬

tles, Hippo damia convergens Guerin-Meneville (L. Wainscott, per¬

sonal communication). One other organization reports shipping

coccinellids to Wisconsin, and a third company sent Trichogramma

to more than 25 customers in recent years. Also, the Chinese

mantis, Tenodera aridifolia sinensis Saussure, has been introduced

into several areas of the state, although the status of its establish¬

ment is unsure. At least 4 organizations responding to our ques¬

tionnaire have sold mantis oothecae to Wisconsin customers. One

or 2 specimens of this mantis are submitted by local citizens to the

Milwaukee Public Museum nearly every year (K. MacArthur, per¬

sonal communication) ; an ootheca containing infertile eggs was

collected in 1967 by one of the authors (J.W.M.) from a weedy

area of the University of Wisconsin Arboretum in Madison; and

the progeny from about 15 oothecae were released in a vegetable

garden in Amery, Wisconsin in 1971.

SUMMARY DISCUSSION

It is evident from the information presented that the utilization

of applied biological control in Wisconsin is as yet relatively

meager. None of the case histories described involved an extensive

program including recognition of the problem, investigation of the

pest and its ecology, biological and ecological studies of the bene¬

ficial organism (s), collection, importation and mass rearing of the

biotic agent (s), field releases, and follow-up recovery studies to

determine establishment and spread of introduced biotic agents.

That is not to say that the biological control approach cannot be

followed without involved ecological investigations and methodol¬

ogy. Simmonds (1972) recently discussed this problem thoroughly,

alluding to the practical, temporal, and economic considerations

involved, and concluded it is better to make a few expedient intro¬

ductions which may be of value than to do nothing at all. A case

in point is the biological control of the larch casebearer in Wiscon¬

sin, the most successful introduction program on record in the

state. Although the program was not conducted on a grand scale,

with great financial outlay or exhaustive investigations, the results

were significant. The non-classical approach used in the lesser

peach tree borer trapping program on Washington Island makes

it difficult to compare with the other classical programs discussed,

but, in its own way, this study will probably result in an exhaustive

set of data relating to objectives, procedures, and results.

The complete ecologically based program is by far the preferred

approach when it is feasible. For a number of years the forest

1974] Mertins and Coppel — History of Biological Control 129

entomology research group (D. M. Benjamin, H. C. Coppel, D. M.

Norris, R. D. Shenefelt) of the Department of Entomology has

conducted basic studies on the ecology and natural control factors

affecting forest insect pests in Wisconsin, thus providing a sound

basis upon which future biological control programs can be planned.

Information has accumulated on Acrobasis rubrifasciella Packard,

Choristoneura famif erana (Clemens) , C. pinus Freeman, Dasineura

balsamicola (Lintner) , Dasychira plagiata (Walker), Dendrotettix

quercus Packard, Glyptoscelis pubescens (F.) , Hylobius spp., Pis-

sodes strobi (Peck), Reticulitermes flavipes (Kollar), Rhyacionia

buoliana (Schiffermliller) , Tourney ella numismaticum (Pettit and

McDaniel), and others.

The classical biological method of pest suppression has practical

and desirable advantages for many pest species. First, it is often

self-sustaining. Once effective introduced beneficial agents have

been established, no further efforts or expenditures may be required

to avoid economic losses caused by the pests. In other cases, periodic

(early in the season or generation every year) or inundative (much

in the manner of an insecticide) releases of beneficial agents may

be necessary to restrain pest populations. Secondly, the self-

sustaining nature of an effective biotic agent may alleviate the

necessity for routine periodic chemical applications with their asso¬

ciated problems of residues, pest resistance, broad spectrum tox¬

icity, and environmental contamination. In conjunction with these

advantages are certain problems which in some instances must

be given consideration. Chemical pest suppression usually gives

quick and highly visible pest relief. Biological control requires a

certain lag time until the biotic agent establishes its effectiveness,

and although it is present and active, it is not always readily

observable to the untrained eye. This fact is exemplified by a con¬

sideration of the thousands of insect species which share the

environment with man, but are not economic pests. Natural enemies

of these potential pests are omnipresent and keep their numbers

in check, frequently without receiving due credit. Even though a

pest species population has been successfully reduced to tolerable

or subeconomic levels by an introduced biotic agent, pest numbers

are still subject to fluctuation from year to year. Complete popu¬

lation control is rarely obtained. Instead a desirable balance is

sought between detrimental and beneficial species numbers. In some

high value crops, such as fruits and vegetables, consumer demand

for virtually unblemished produce will not allow even an occasional

high insect population to occur, and in these instances the grower

may have to resort to alternative methods of suppression.

Although chemical and biological controls are sometimes in direct

opposition, they are not always mutually exclusive. The coordinated

130 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

use of the two methods at the same time, or for that matter, the

harmonious use of any two or more pest suppressive methods may

be termed integrated control, or integrated pest management, and

currently such systems are receiving increased attention (National

Research Council, 1969; Northeastern Forest Experiment Station,

1971). Research along these lines is also underway in Wisconsin,

and a number of reports have appeared (Coppel and Norris, 1960;

Oatman, 1966 ; Eckenrode, 1970).

ACKNOWLEDGEMENTS

Much of the information in this report is from, previously un¬

published data assembled here with the kind assistance of a num¬

ber of our colleagues, many of whom are acknowledged in the text.

Special appreciation for records and information is due to: J. W.

Apple, D. M. Benjamin, K. S. Hagen, B. H. Kennedy, T. H. Klahn,

C. F. Koval, D. M. Norris, Jr., and R. D. Shenefelt.

This project was supported by the College of Agricultural and

Life Sciences, University of Wisconsin, Madison, and in part by

the Wisconsin Department of Natural Resources through the School

of Natural Resources, and the University of Wisconsin Research

Committee of the Graduate School with funds supplied by the Wis¬

consin Alumni Research Foundation.

BIBLIOGRAPHY

BAKER, W. A., W. G. BRADLEY, and C. A. CLARK. 1949. Biological con¬

trol of the European corn borer in the United States. U.S. Dept. Agr.,

Tech. Bull. 983, 185 pp.

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NUTRIENT SOURCES FOR LAKE MENDOTA— 1972

William C. Sonzogni

University Wisconsin — •

Madison

and

G. Fred Lee

University Texas — •

Dallas

INTRODUCTION

It has been several years since Lee et al. (1966) published an

estimation of nutrient loadings to Lake Mendota (the paper was

revised in 1969, but only with regard to the figure for the annual

nutrient input from nitrogen fixation). Since that time new data

and information concerning certain nutrient sources has become

available. In addition, changes in the population and land use of

the watershed have occurred. In view of the recent diversion of

sewage effluent which normally entered tributaries to the lake, it

is imperative that the sources, amounts and types of nutrients be

estimated as well as possible, if the effect of this diversion is to

be properly evaluated. It is the purpose of this paper to update the

estimation of the total amounts of nutrients entering Lake Men¬

dota as well as to provide some quantitative information on the

forms of nutrients annually entering the lake from different

sources.

LAKE CHARACTERISTICS

Lake Mendota, perhaps the most studied lake in the world, is the

largest of the Madison lakes, which form a chain along the Yahara

River. Formed as a result of moranic damming during the most

recent ice age, it is classified as a hard-water, eutrophic lake ac¬

cording to most standards. The drainage area of Lake Mendota

is composed mostly of fertile farm land and urban area. The hypo-

limnetic waters become devoid of oxygen during summer stratifi¬

cation and during late winter oxygen depletion occurs in the bottom

water. Excessive weed growth and periodic algal blooms create

offensive conditions during the summer months.

The lake is located in south-central Wisconsin, which has a

semi-humid climate. The monthly average temperatures range be¬

tween 19 F and 73 F (Dane County Planning Commission, 1971).

Precipitation averages about 30 inches each year, while evapora¬

tion amounts to about 24 inches a year.

133

134 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Lake Mendota has a volume of about 128 x 109 gal. (486 x 106

m3) and a surface area of about 9,700 acres (3940 ha.). The

maximum depth of the lake is 84 ft (25 m), while the mean depth

is about 40 ft (12 m). The tributary flow into the lake is on the

order of 100 cfs (Burgy, 1950; Cleasby, 1951), the highest monthly

inflows being recorded during spring thaw. A synopsis of limno¬

logical information on the lake may be found in Frey (1963) .

Land Use

Land usage within the Mendota basin was estimated from maps

and current land use information provided by the Dane County

Planning Department. All incorporated land area within the water¬

shed was considered to be urban. The remaining land was

categorized as rural, marshland, or woodland, except for land

designated as residential or land used for transportation, utility or

communication purposes. Such land was counted as urban drain¬

age. Marshland and woodland were estimated from a map prepared

by the Planning Department.

Table 1 compares the land use estimates of Lee et al (1966)

with those of this study. The only significant difference is with

urban area and woodland. The increased urban drainage area may

be the result of recent urban sprawl. The decreased woodland area

in the revised estimate is probably due to the fact that only the

major woodland areas were counted in the study. Small clumps

of woodland of only a few acres could not be differentiated from

rural land.

ESTIMATED NUTRIENT SOURCES

Base Flow

Estimates of the nutrient contribution from base flow (portion

of total tributary flow which is derived from ground .water) are

summarized in Table 2. It can be seen that the few studies that

TABLE 1. PREVIOUSLY PUBLISHED AND REVISED ESTIMATE OF

LAND USE WITHIN THE LAKE MENDOTA WATERSHED

1974] Sonzogni and Lee — Nutrients for Lake Mendota

135

TABLE 2. BASE FLOW

Soluble Total Total

Source Ortho — P P NH 4+ — N NO 3“— N Org — N N

lbs/ acre/year

Minshall et al _ _ 0.1 _ _ _ 1.1

(1969)

Comment: Southwestern Wisconsin streams; includes minor wraste discharge

Zitter (1969)__. _ _ _ _ 0.13 0.17 0.27 0.65 1.0 1.9

Comment: Six Mile Creek-South (a Mendota tributary)

Gardner (1971) _ 0.06 0.15 _ 1 . 13 1 0.22 1.35

Comment: Includes some waste water discharge. Based on data of Belter and Calabresa (1950)

■Inorganic nitrogen

have been made are in surprisingly good agreement despite the

different ways in which the estimates were obtained and the differ¬

ent study areas. When examining Table 2 (and future tables) it

should be noted that one Ib/acre is approximately equal to one

kg/ha.

Minshall et al. (1969) estimated the quantity of nutrients car¬

ried by base flow annually during a two-year study of streams in

southwestern Wisconsin. Lake Mendota tributaries were deemed

unsuitable for determining base flow on account of possible pollu¬

tion from domestic and industrial sources. For this reason, the

authors decided to study base flow in southwestern Wisconsin, a

predominately rural area whose streams receive little such pollu¬

tion. Minshall et al. (1969) felt that the nutrients carried by base

flow in these streams would be roughly comparable to the nutrients

transported by base flow in the Madison area.

Minshall et al. (1969) reported that the average annual base

flow in the drainage areas of southwestern Wisconsin was 0.360

cfs/mi.2 Cleasby (1951) and Burgy (1950) found the inflow to

Lake Mendota to be approximately 95 cfs per year. If one takes

the watershed of Lake Mendota to be about 250 mi,2 then the

average base flow is 0.38 cfs/mi.2 This suggests a comparable base

flow rate on a unit area basis. Minshall et al. (1969) also found

that the annual nutrient input from base flow per acre of drainage

was generally constant for the different streams studied and varia¬

tions appeared independent of flow. Thus, the results of Minshall

et al. (1969) are quite likely applicable to Lake Mendota.

Zitter (1969) studied a portion of the watershed of Six Mile

Creek-South, a tributary to Lake Mendota. The drainage area is

predominantly agricultural (dairy farms and mixed crops). Using

automatic sampling equipment and continuous flow measuring

devices, the annual nutrient contribution from the watershed due

to base flow and surface runoff was estimated. Although Zitter’s

136 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

data did not explicitly separate base flow from surface runoff, he

observed that about 75% of the annual nutrient losses occurred

during periods of high runoff, suggesting that base flow accounted

for about 25% of the nutrients carried by the stream.

Gardner (1971) attempted to separate base flow from total run¬

off from the tributary nutrient input data for Lake Mendota,

compiled by Belter and Calabresa (1950). Gardner (1971) summed

cumulatively the nutrient quantities (soluble ortho-P, total-P, or-

ganic-N, inorganic-N) from all the tributary sources and plotted

them versus time. He then used the rate changes to roughly dif¬

ferentiate between base flow and runoff during the four seasonal

periods. During periods when no runoff was thought to have oc¬

curred, the rate of nutrient input was relatively constant and

significantly lower than during periods when runoff was known to

occur. From these periods of low, constant input an assessment

of the base flow nutrient contribution was made. The results in¬

cluded any municipal and industrial wastes discharged into the

tributaries.

Since the data for base flow in Table 2 implicitly includes small

but significant nutrient contributions from other sources (i.e.,

small waste water inputs, barnyard drainage, septic tank seepage

and others) which are difficult to separate from the nutrients in

stream flow arising solely from ground water flow, the values listed

in Table 2 may overestimate the base flow input. For this reason,

the values reported by Minshall et al. (1969), which were the

minimum values reported in Table 2, will be used to reestimate the

nutrient loadings from base flow. The forms of nutrients in base

flow would be expected to be predominantly N03~— N and soluble

ortho-P.

If a tributary drainage area of 123,000 acres (Belter and Cala¬

bresa, 1950) is assumed for Lake Mendota, an average annual

loading of 12,000 lbs of phosphorus and 135,000 lbs of nitrogen can

be predicted, using the data of Minshall et al. (1969) . The previous

estimated contribution of base flow, based on the amount of ground

water thought to be entering the tributaries directly (see section

on Ground Water Seepage ), was 79,000 lbs/year of nitrogen. No

estimate of the phosphorus input from base flow was made in the

previous study. Thus, in the case of nitrogen the new estimate

about doubles the old one.

An important point to consider is the denitrification of ground

water as it emanates from marshy areas and contributes to the

base flow of streams. Lee et al. (1971) found significant denitrifi¬

cation reactions, which result in the conversion of nitrate nitrogen

to nitrogen gas, occurring in the Mendota watershed. It is also

possible that the opposite reaction, nitrogen fixation, could occur

1974] Sonzogni and Lee — Nutrients for Lake Mendota 137

within a marsh (a study is currently underway at the University

of Wisconsin Water Chemistry Program to determine the signifi¬

cance of nitrogen fixation in the marsh environment) . These re¬

actions would have to be accounted for if the nitrogen contributed

by base flow were estimated from the amount of ground water

entering tributaries directly, as was done in the Lee et al. (1966)

report.

Rural Runoff

Several studies have been conducted in the Madison area which

have estimated the average annual nutrient loading from rural or

agricultural runoff. The results of these studies have been sum¬

marized in Table 3.

Witzel et al. (1969) measured the amount of surface runoff

from several agricultural watersheds near Fennimore, Wisconsin,

in 1967. Based on the data obtained they found the contribution

of surface runoff to be 3.6 lbs of total-N/acre/year and 1.1 lbs of

total-P/acre/year. However, because the 1967 winter runoff in the

area was about twice normal because of one heavy rain in January,

they estimated the nutrient losses for a typical year to be only

2 lbs of total-N/acre/year and 0.61 lbs of total-P/acre/year, as¬

suming that nutrient losses were directly proportional to runoff.

TABLE 3. NUTRIENT LOADING FROM RURAL RUNOFF-

ESTIMATES FOR SOUTHERN WISCONSIN AREA

Soluble Total

Source Ortho — P P NH 4+ — N NO 3- — N Org — N Inorg— N Total N

lbs/acre/year

Witzel et al _ _ 0.6 _ _

(1969)

Comment: Estimated for normal year — Southwestern Wisconsin

Witzel et al _ _ 1.1 _ _

(1969)

Comment: Measured during study

Zitter (1969) _ 0.39 0.53 0.85 1.50

Comment: Measured runoff — Mendota tributary

Zitter (1969)-.-- _ 0.60 0.80 1.3 2.3

Comment: Estimated for normal year

Gardner (1971) _ 0.08 0.15 _ _

Comment: Based on Belter and Calabresa (1950)

Sawyer (1947) _ 0.08 0.40 - -

Comment: Uncorrected for base flow

Sawyer (1947) _ _ 0.30 _ _

Comment: Corrected for base flow estimated by Minshall et al. (1969)

Average* 2 _ 0.33 0.6 _ _

'Correction made assuming base flow all inorganic N

2 Averages computed from Sawyer’s values corrected for base flow

3 Assume soluble ortho — P equals 50 % of average total — P

4 Difference between average total — N and average inorganic — N

138 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Zitter (1969) studied a portion of the watershed of Six Mile

Creek-South. As mentioned previously, surface runoff and base

flow were not measured separately, but about 75% of the annual

nutrient losses occurred during periods of high flow and thus was

considered to be the result of surface runoff. Zitter (1969) rea¬

soned that his data probably underestimated the normal nutrient

contribution of the watershed due to the below normal winter

precipitation the year of his study. Because of the lack of snow,

manure was spread on bare ground or on a very thin layer of snow.

Consequently, Zitter (1969) concluded that there probably was not

sufficient water at spring thaw to leach all the nutrients from the

manure and carry them into the stream. Zitter (1969) therefore

multiplied his runoff results by 1.5, as he felt the resulting figure

would be more representative of nutrient losses during a normal

year.

Sawyer (1947), during a study of the Madison lakes in the mid-

1940’s, determined the nutrients in the agricultural drainage to

Lakes Monona, Waubesa and Kegonsa. The data shown in Table 3

represent the average for the three lakes studied. No attempt was

made to differentiate between base flow and runoff. Sawyer’s re¬

sults minus base flow (as estimated in the previous section) are

also included in Table 3.

As discussed previously, Gardner (1971) used the data of Belter

and Calabresa (1950) to separate base flow and rural runoff from

total nutrient input. Belter and Calabresa (1950) estimated the

total tributary input to Lake Mendota from October 1, 1948 to

October 1, 1949. In their study continuous flow measurements were

made, but samples for nutrient analysis were obtained only every

two to three weeks. In calculating total nutrient inputs they as¬

sumed a constant concentration of nutrient between sampling

periods.

Two other studies of the tributary nutrient input to Lake Men¬

dota were conducted in the late 1940’s, but their results have not

been included in Table 3. Bartsch and Lawton (1949) measured

the input of nutrients to Lake Mendota via the lake’s tributaries

over a two year period from 1945 to 1947. Flow measurements

were not made on a continuous basis, but just at the time of

sampling which was irregular. Their total flow measurements were

low compared to measurements made by others (Burgy, 1950 and

Cleasby, 1951) despite normal precipitation during the years of

their study. This suggests that their tributary nutrient input

values, which included base flow and some municipal and industrial

wastes, were low (Fruh and Lee, 1965). Thus, the results of

Bartsch and Lawton (1949) have not been included in Table 3.

Emelity and Hanson (1949) also measured the total tributary

1974] Sonzogni and Lee — Nutrients for Lake Mendota 139

nutrient input to Lake Mendota between July 1, 1948 and May 1,

1949. Again, they did not attempt to differentiate between base

flow, runoff and waste water inputs. Although they used continu¬

ous flow data, they collected samples only every three weeks and

assumed the concentration of nutrients remained constant between

sampling periods. Although Emelity and Hanson’s (1949) study

was not for a full year, they found higher inputs than estimated

by Bartsch and Lawton (1949) for a full year.

Thus, the Mendota tributary studies conducted by Bartsch and

Lawton (1949), Emelity and Hanson (1949) and Belter and Cala-

bresa (1950) all obtained their samples on a periodic basis. They

then assumed that the concentrations of nutrients remained con¬

stant between sampling dates. However, it has been shown that a

very large amount of nutrients may be carried off during periods

of high flow (Kluesener, 1965; Lee, 1969; Zitter, 1969; Taylor et

ak, 1971). This would be especially important for manure spread

on frozen ground, as a large pulse of nutrients would be carried

into the receiving stream during a spring thaw. Zitter (1969)

found that nutrient loss from rural land was highest during a thaw

in February. A heavy June rain also resulted in runoff and high

nutrient loss.

Studies on Black Earth Creek by the University of Wisconsin

Water Chemistry Program have demonstrated the need for basing

the sampling program on stream discharge rather than arbitrary

time frequency. Figure 1 shows the discharge-conductance data

for Black Earth Creek, (outside Lake Mendota Basin) during the

spring thaw in 1965 (Shannon and Lee, 1966). The stream had a

base flow from groundwater of approximately 20 cfs for March 29

and 30. During the period of March 30 to April 2, the discharge

increased to a peak of 150 cfs on April 1, then gradually receded.

Flow during this period was primarily due to snow melt. On April

2 it started to rain; discharge increased to approximately 150 cfs

on April 5. After this date, the discharge gradually decreased,

approaching base flow after April 13. The specific conductance

of the water showed the typical inverse relationship with dis¬

charge, with the minimum in specific conductance approximately

corresponding to the maximum in discharge. The snow melt and

precipitation runoff were low in dissolved salts and diluted the

relatively hard groundwater base flow.

Figure 2, on the other hand, shows that maximum phosphate

concentration corresponded to the maximum discharge (Shannon

and Lee, 1966). As shown in this illustration the snow melt and

rainfall runoff contained more phosphorus than did the base flow.

This phosphorus was probably derived from manure. The con¬

densed phosphates in the figure were composed of the nonfilterable

PHOSPHORUS CONCENTRATION mg P/l DISCHARGE - cfs

140 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

DATE

FIGURE 1. Conductance and discharge of Black Earth Creek during

spring 1965. (After Shannon and Lee, 1966)

DATE

FIGURE 2. Phosphorus concentrations of Black Earth Creek during

spring 1965. (After Shannon and Lee, 1966)

CONDUCTANCE - jumho./cm

1974] Sonzogni and Lee — Nutrients for Lake Mendota 141

phosphorus that is measured by mild acid hydrolysis. No known

sources of domestic or industrial pollution were upstream from this

sampling station; the condensed phosphates, therefore, were prob¬

ably organic phosphorus compounds. As expected, an appreciable

amount of total phosphorus transported by the stream during

spring thaw was particulate phosphate.

The same relation of phosphorus and nitrogen concentration to

periods of high discharge was found at all times of the year; the

major proportion of these elements were transported during pe¬

riods of high discharge. Kluesener (1965) found that 65% of the

ammonia nitrogen carried annually by Black Earth Creek was car¬

ried in just five days during spring thaw. It is important, there¬

fore, to establish a sampling program in which the frequency of

sampling increases as the discharge increases. Sampling a stream

once every three weeks or even more frequently may not be

representative and could result in gross error in computing the

amount of plant nutrients contributed by the stream.

Thus, it is likely that the early studies of the Madison lakes may

have underestimated nutrient loadings. A possible exception would

be the results of Belter and Calabresa (1950), who claimed that

samples were collected at peak flows as well as dry weather flows.

Hence, the data of Belter and Calabresa (1950), as reported by

Gardner (1971), have been included in Table 2.

Table 3 shows that the reported contribution of total-P varies

widely, from about 0.1 to 1.0 Ib/acre/year, with an average value

of 0.58 lb/acre/year. Soluble ortho-P ranges from about 25 to 75%

of the total-P, based on the limited data available. Thus, assuming

an average loading of 0.6 lb/acre/year for total-P, and assuming

soluble ortho-P comprises about one half the total-P, an average

value of 0.3 lb/acre/year for soluble ortho-P is obtained.

Total-N ranges from about 1.0 to 10 lbs/acre/year (Table 3)

with an average value of approximately 4.5 Ib/acre/year. Inor-

ganic-N ranges from about 0.5 to 6.0 lbs/acre/year with an aver¬

age of about 2.8. The organic-N values range from about 0.5 to

5.0 lbs/ acre/year. If an average inorganic-N value is subtracted

from the average total-N value, a value of 1.8 lbs/acre/year is

obtained for organic-N.

The results from Table 3 may be compared with nutrient load¬

ings for rural lands from other parts of the country. Table 4 lists

the results of several North American studies conducted in areas

with climates somewhat similar to Wisconsin. When comparing

these numbers, one must be aware of the wide variability between

experimental methods, soil characteristics, mode of drainage,

topography, percolation, and infiltration rates, as well as many

142 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

other factors. In most cases total tributary input was measured so

that base flow and sometimes municipal and industrial waste inputs

are included in the nutrient loading figures. At best, the loadings

listed in Table 4 are only roughly comparable, but they should

prove valuable for delineating a possible range of values.

The results of some literature studies have not been compiled

in Table 4. The results of Weidner et al. (1969) and Sylvester

(1961) are generally an order of magnitude higher than the re¬

sults in Table 4. Owens (1970) studied the tributary input of

English rivers draining rural lands and found that the total-N

contribution ranged from 0.5-20 lbs/acre/year and the total-P

contribution varied between 0.5 and 2.1 Ibs/acre/year. These

numbers include base flow but the nutrient concentration from

sewage was subtracted. The results of Owens (1970) for England

are generally higher than are most North American studies.

Returning to Table 3, it can be seen that the mean value for

total-P, 0.6 lb/acre/year, is somewhat higher than that found by

most workers in Table 4. The higher value for Wisconsin might

TABLE 4. NUTRIENT LOADING FROM RURAL RUNOFF: ESTIMATES

FROM NORTH CENTRAL STATES AND SOUTHERN CANADA

Soluble Total

Source & Place of Study Ortho — P P NH4+ — N NO 3' — N Org — N Inorg — N Total — N

lbs/acre/year

Engelbrecht & Morgan _ _ 0.2 _ _ _ _

(1961) Illinois

Comment: Total — P actually ortho =*= hydrolyzable P; includes base flow and waste discharges

1974] Sonzogni and Lee — Nutrients for Lake Mendota 143

best be explained by the high concentration of dairy farms in the

watershed and the then prevalent practice of spreading manure

on frozen ground. Minshall et al (1970), Corey et al (1967) and

Hensler and Attoe (1970) have discussed the high nutrient yield

from manured land. Both Zitter (1969) and Witzel et al. (1969)

attributed their high total-P values for rural runoff (Table 3) to

the spread of manure on frozen ground.

Nutrient budget estimations based on small scale watersheds

could have low reliability if extrapolated to cover large areas

(Armstrong and Rohlich, 1970). For example, it is possible that

not all of the phosphorus which enters a stream in runoff will be

carried to a lake. Soil particles suspended during high flow could

sorb phosphorus and at some point be deposited along the stream

before they reach the lake or ocean. Such a phenomenon has been

cited for the high natural fertility for some of the lands in large

river basins, such as the lower Nile River. This phenomenon is not

thought to be of any importance in the Lake Mendota drainage

basin.

Lee (1972) discussed another mechanism by which the soluble

inorganic P and N present in a stream at one location may be

transformed into non-available (refractory) N and P at some down

stream location. A small but definite percentage of the N and P

taken up by macrophytes and algae is not remineralized in natural

water systems. Based on the various studies it appears that 0.1

to 1.0 lb of total-P /acre/year should provide a range within which

the true average value is likely to be found. The range of total-N

values for the Madison area agrees reasonably well with those

values in Table 4. Data for the different individual forms of nitro¬

gen and phosphorus in Table 4 are rather limited, but generally

fall in the range estimated for the nutrient forms in Table 3.

The nutrient loading from rural runoff originally estimated by

Lee et al. (1966) is compared to the estimates from this report

in Table 5. Lee et al. (1966) used the data of Eck et al. (1957)

and Midgley and Dunklee (1945) to arrive at their numbers.

Although it was not completely clear in their report, the estimates

published by Lee et al. (1966) were for soluble forms only. It can

be seen that the new estimates (Table 5) are considerably higher

than those of Lee et al. (1966), especially for nitrogen. However,

as shown in Tables 3 and 4, there were no studies that found such

a small contribution of nitrogen from rural runoff as Lee et al.

(1966). Thus, it appears that the estimate for soluble nitrogen

made by Lee et al. (1966) was low.

More information is really needed on the specific sources of

nutrients in rural runoff. The relative significance of manured

144 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

TABLE 5. PREVIOUSLY PUBLISHED AND REVISED ANNUAL

NUTRIENT LOADING TO LAKE MENDOTA FROM

RURAL RUNOFF

1 Average value from Table 2 plus predicted range in parenthesis

2 Based on a rural watershed of 115,000 acres

3 Average plus predicted range in parenthesis

4To the nearest 1,000 pounds

land, different types of cropland, rainfall, etc., as sources of nutri¬

ents in rural areas should be determined.

Lee et al. (1966) attempted to determine the component of the

rural runoff nutrient loading which originates specifically from

manured land. This estimate was derived by assuming each of the

approximately 20,000 cows in the watershed produces 15 tons of

manure in a year and that one-half of this manure is applied dur¬

ing the months of November through March, so that about 150,000

tons will be applied on frozen ground. This amount of manure

would cover 15,000 acres if applied at the rate of 10 tons/acre. Lee

et al. (1966) then used the data of Midgley and Dunklee (1945),

which indicated that about 3 lb of N and 1 lb of P were lost from

a 10 tons/acre application of manure on an 8% slope in Vermont,

to calculate that about 45,000 lb of N and 15,000 lb of P would be

lost from manured land in the Mendota watershed. If these very

rough figures are compared to the revised estimates in Table 5, it

can be seen that about 45% of the soluble phosphorus and 15%

of the inorganic nitrogen in rural runoff could be derived from

manured lands. It should be kept in mind that this estimate is of

low reliability and that more studies on the sources of rural runoff

are needed. To be sure, manured lands are a significant contributor

to the fertility of agricultural runoff in the Madison, Wisconsin

area.

Urban Runoff

The annual amount of nutrients entering Lake Mendota per

acre of urban drainage was estimated from Kluesener (1972). The

study consisted primarily of intensive sampling for short periods

1974] Sonzogni and Lee — Nutrients for Lake Mendota 145

of time on a small urban section of the watershed of Lake Wingra,

one of the Madison lakes. It is felt that this study concurrently

presents the best estimate of the nutrient content of urban runoff

for the Madison area. Kluesener (1972) has compared the results

of his study with previous studies of urban runoff and found gen¬

eral agreement with most of the previous studies in the area.

A comparison of the estimates of Kluesener (1972) and those

of Lee et al. (1966) for urban runoff is given in Table 6. It can

be seen from the revised estimates that more nutrients are entering

Lake Mendota from urban runoff than previously thought.

Although Kluesener (1972) did not determine the relative

amounts of nutrients derived from various urban activities, he did

point to the importance of leaves and seeds from trees as phos¬

phorus sources in urban runoff. Recently, Co wen and Lee (1971)

followed up Kluesener’s preliminary investigation on the role of

leaves as a source of phosphorus in urban drainage. They found

that inorganic phosphorus was rapidly leached out of leaves col¬

lected in early autumn. Even leaves weathered by rain water or

lake water contained significant amounts of leachable phosphorus.

In general, they found that each gram of leaves may potentially

fertilize many liters of water above the critical concentrations

of phosphorus often cited as causing excessive growths of aquatic

plant or algae in natural waters. Hence, leaves and seeds and

flowers must be considered as a major contributor to the relatively

high concentrations of phosphorus found in urban drainage.

Atmospheric Precipitation and Dry Fallout

Kluesener (1972) measured the nutrient content of rain and

snowfall as well as dry fallout during his study of Lake Wingra.

If it is assumed that his results are typical of the Madison area,

TABLE 6. PREVIOUSLY PUBLISHED AND REVISED ANNUAL

NUTRIENT LOADING TO LAKE MENDOTA FROM

URBAN RUNOFF

'Based on an urban watershed of 10,000 acres for Lake Mendota

2 To the nearest 1,000 pounds

146 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

his data may be used to reestimate this source of nutrients to Lake

Mendota, assuming’ an average rainfall of about 30 inches.

A comparison of the estimates based on Kluesener (1972) and

those of Lee et al. (1966) for atmospheric precipitation and dry

fallout are presented in Table 7. The revised estimates when dry

fallout and atmospheric precipitation are totaled are significantly

higher than the contribution originally published.

Groundwater Seepage

Unfortunately, very little information is available on the ground-

water nutrient input to Lake Mendota by direct seepage into the

lake. The previously published estimates (Lee et al., 1966) were

based primarily on a report by Cline (1965). Cline (1965) esti¬

mated that 80% of the total atmospheric precipitation on the

watershed is consumed by evapotranspiration and that almost none

of the ground water is lost from the watershed by underflow. If

the average annual atmospheric precipitation amounts to about 30

inches/year, then about 6 inches will be divided between runoff

and percolate. Assuming about 2 inches will run off the surface,

an average of about 4 inches, or about 65 cfs over the entire

watershed, should enter the ground as percolate each year. Cline

(1965) thus estimated that about 30 cfs of this enters the lake

directly each year, the rest contributing to the flow of surface

tributaries (base flow). The roughness of these estimates was

emphasized in the original report by Lee et al. (1966). Further,

C. L. R. Holt, Jr. (personal communication), district geologist of

the U.S. Geological Survey, has indicated that the previous esti-

TABLE 7. PREVIOUSLY PUBLISHED AND REVISED ANNUAL

NUTRIENT LOADING TO LAKE MENDOTA FROM

ATMOSPHERIC PRECIPITATION AND DRY FALLOUT

1 Based on a lake surface of 9,730 acres

aTo the nearest 1,000 pounds

1974] Sonzogni and Lee — Nutrients for Lake Mendota 147

mates based on Cline (1965) for total ground .water input to the

lake are not very accurate. Nevertheless, using this estimated

ground water inflow and a concentration of 2.5 mg/1 of NO^~— N

(based on the data of Eck, 1958, and Domogalla et al., 1926) and

0.01 mg/1 P (based on the phosphorus concentration found in a

well on the western shore of the lake), Lee et al. (1966) concluded

that about 171,000 lbs of NCL- — N and 600 lbs of P enter the lake

via ground water seepage each year.

Unfortunately, even if the ground water inflow to the lake and

the chemical composition of the ground water at some point up

gradient from where it emerges into the lake were known with

some degree of certainty, a reliable estimate of the amounts of

nutrients contributed via the ground water could not be easily

obtained (Lee and Kluesener, 1971). This is due to the fact that

information must be available on the manner in which the ground

water enters the lake. If entry is primarily through sand, sub¬

merged springs, or other areas in which there is little or no lake

sediment, then most of the nutrients present in the ground waters

would be transported to the lake. However, if the entry is through

the sediments of the lake, it can be expected that at certain times

of the year some of the phosphorus present in the sediments

through which the water enters would be removed by sorption

reactions in the sediments. At other times, it is possible that this

ground water might tend to displace interstitial waters, which may

be high in phosphorus, into the lake. In the case of nitrogen as

N03— N, it is reasonable to expect that in those areas of the sedi¬

ments where the oxygen concentration is low, there should be sig¬

nificant denitrification which would tend to reduce the amounts of

available nitrogen that enter the lake water from that which was

predicted based on total ground water input.

Recently, Keeney et al. (1971) investigated the importance of

denitrification in Lake Mendota sediments to the nitrogen budget

of the lake. Based on experiments with samples of surface sedi¬

ments from 7 and 14 meters, they estimated that about 63% of the

nitrate-nitrogen entering Lake Mendota will be denitrified. Thus,

they estimated that the nitrogen entering Lake Mendota via ground

water seepage to be only 63,000 Ibs/year vs. the original estimate

of 171,000 lbs/year.

The approach used by Keeney et al. (1971), however, may

grossly overestimate the significance of sediment denitrification as

a process reducing the amount of nitrogen input into lakes. Based

on the hydrology of the Lake Mendota basin and in particular the

ground waters around the lake (see Dane County Planning Com¬

mission, 1971), it would be expected that significant amounts of

shallow ground water enter the lake through the sides of the lake

148 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

near the surface. Furthermore, considerable amounts of ground

water are entering the lake through fissures in limestone outcrops,

as evidenced by divers working under the direction of the senior

author. Ground water entering through the sides of the lake and

through fissures would probably not be exposed to the anoxic sedi¬

ments which would lead to rapid denitrification as reported by

Keeney et at. (1971). If ground water entered the lake through

the deeper sediments under anoxic conditions, it would be correct

to expect rapid denitrification, as demonstrated earlier by Brezonik

(1965) ; however, it is unreasonable to reckon that significant

amounts of ground water enter Lake Mendota through these deep

sediments.

It is almost impossible to utilize any measurements of the rates

of denitrification in sediments to obtain a reliable estimate of how

much of the nitrogen entering a lake is lost through this reaction

without very good information on the hydrology of the ground

water, and in particular how the ground water enters the lake.

Presumably some denitrification of ground water does occur so

that the ground water input of nitrogen would probably be less

than the original estimate of Lee et al. (1966) report. Brezonik

and Lee (1968) have shown that denitrification in the anoxic

waters is a relatively small but significant sink in the nitrogen

budget for Lake Mendota. About 62,000 lbs of nitrogen were lost

from the lake hypolimnion during the summer of 1966. This would

also tend to reduce the input of nitrogen from ground water. How¬

ever, without a better understanding of the hydrology of ground

water seeping into the lake, it is not felt justifiable to change the

original estimate (Lee et al, 1966) of the amounts of nitrogen and

phosphorus contributed by ground water seepage into the lake.

The lack of information of the relative nutrient contribution of

ground water seepage to the total nutrient budget is one of the

greatest deficiencies of the proposed nutrient loading estimations

for Lake Mendota. This is especially true for nitrogen. Because of

the relatively high concentrations of nitrate present in ground

water in the Madison area, ground water must be considered as a

potentially important source of nitrogen to Lake Mendota.

Nitrogen Fixation

The amount of nitrogen fixed in Lake Mendota was most recently

studied by Torrey (1972). Nitrogen fixation was based on the ace¬

tylene reduction method, assuming a ratio of 3:1 for the rate of

acetylene reduced to nitrogen reduced. Estimates of nitrogen fixa¬

tion in Lake Mendota in the past were based on limited experiments

using fixation of 15N2.

Torrey (1972) reported that about 88,000 lbs of nitrogen were

1974] Sonzogni and Lee — Nutrients for Lake Mendota 149

fixed each year during 1970 and 1971 and this value will be used

to reestimate nitrogen fixation in Lake Mendota. This compares

with the estimate previously published by Lee et al. (1966) of

80,000 lbs of N/year ; subsequently, it was found that an error had

been made in the computation of this figure and in 1969 the figure

was revised to 2000 lbs of N/year.

Torrey (1972) found that most of the nitrogen was fixed by blue

green algae in the surface waters during the summer months when

combined nitrogen available for algal growth is low. However,

some fixation was found to occur in the sediments of Lake Men¬

dota. While this was thought to be small compared to the total

fixation that occurs in the surface waters, this fixation would tend

to counteract to some degree the denitrification reactions that

potentially take place as a result of anoxic conditions in some of

the sediments. Finally, it should be mentioned that the results of

Torrey (1972) do not include nitrogen fixation occurring in marsh

land bordering the lake or in littoral areas. Current studies by

Lonergan, graduate student, U. W. Water Chemistry Program,

show that potentially significant amounts of nitrogen fixation occur

in marshes.

Marshland

Bentley (1969) studied several marshes in the Madison vicinity

in order to estimate their nutrient contribution. On a yearly basis

he found that the nutrient input to a marsh was approximately

equal to its nutrient output. Apparently, nutrients flowing into

marshes from surface sources during most of the year are held by

the marsh. However, during spring thaw all of the nutrients stored

by the marsh are released with the high spring flows. Thus, based

on the study of Bentley (1969), the marshland within the Mendota

watershed is not thought to be nutrient sink, when a full year cycle

is considered.

Studies of the amounts of nutrients derived from drained

marshes by Bentley (1969), Amundson (1970) and Lee et al.

(1971) have shown that the drainage of a marsh can potentially

represent a major source of nitrogen and phosphorus. Laboratory

studies have shown that large amounts of inorganic nitrogen and

phosphorus can be leached from drained marsh soil in a period of

several years. Since there has been no significant drainage of

marshes in the Lake Mendota basin during the past five years, it

is felt that the large amount of marsh drainage that has occurred

previous to this time is not presently contributing consequential

amounts of nutrients to the lake. Therefore, no attempt will be

made in the current estimate of nutrient sources to include the

potential effects of previously drained marshes.

150 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Woodland

Kluesener (1972) has reviewed the possible contribution of

nutrients in runoff from woodland. In general, he concluded that

surface runoff from woodland would be insignificant. Kelling

(1972) found that prairies and forests in the Madison vicinity

were able to infiltrate rain applied with a sprinkling infiltrometer

at the rate of 5 in/hr. Hence, little surface runoff would be expected

in these areas. However, there might be a contribution of nutrients

to base flow sustained by groundwater in the forested regions,

although it would probably be less than in rural areas. Because

the highly cultivated Mendota watershed has few remaining

wooded areas of any size, the total nutrient contribution from

woodland will be considered negligible.

Waste Water Effluents

The nutrients contributed annually to Lake Mendota from the

waste water effluents of several small municipalities, as well as sev¬

eral industries, were estimated by Lee et al . (1966). Some of the

sources were eliminated late in 1971 when certain waste waters

were diverted to the Madison Metropolitan Sewage District system

so that they ceased to enter Lake Mendota. However, in order to

properly evaluate the significance of the diversion, it is necessary

to reestimate, based on population increases, and other new infor¬

mation, the quantities of nutrients contributed by waste water

sources immediately prior to diversion. Thus the nutrients con¬

tributed by waste waters in 1970 (prior to diversion), as well as

the sources remaining following the diversion, will be discussed.

The Morrisonville Sanitary District currently operates a sewage

system consisting of a two-cell stabilization pond. The Wisconsin

Department of Natural Resources (1971) reported that only on

one occasion was there noted an overflow to the Yahara River, a

Mendota tributary. The district is not thought to be a significant

nutrient source at the present time.

The Village of DeForest has a trickling filter treatment plant

which, up until December, 1971, handled domestic waste from the

village in addition to wastes from the Forest Milk Company.

Treated wastes were discharged to the Yahara River. A survey

(Wisconsin Department of Natural Resources, 1971) in 1970

showed a daily total waste volume of 200,000 gal/day and an esti¬

mated BOD of 167 lbs/day. The plant was reported to be badly

overloaded. In the winter of 1965 Lee et al. (1966) reported a

daily total waste volume of about 135,000 gallons, of which the

milk plant contributed about 35,000 gallons containing a BOD load

of 553 lbs/day. In October, 1962, a 24 hour survey found a total

daily flow of 74,800 gallons containing 174 lbs/day of BOD; how-

1974] Sonzogni and Lee — Nutrients for Lake Mendota 151

ever, the milk plant was not operating during the survey. Accord¬

ing to decennial census reports, the village population increased

from 1,223 in 1960 to 1,911 in 1970.

The Town of Windsor Sanitary District No. 1 currently1 operates

a trickling filter treatment plant which discharges into the Yahara

River. In 1960 the estimated population of the district was 350.

In 1969 the population of the district was estimated at 550 (Wis¬

consin Department of Natural Resources, personal communica¬

tion). A 24 hour-study on September 9 and 10, 1969 found that

the treatment plant removed only 67.7% of the BOD and 57.5%

of the suspended solids (Wisconsin Department of Natural Re¬

sources, 1971). The 24 hour study also revealed the following

nutrient data:

The flow at this time was about 50,000 gal/day and the BOD was

30 Ibs/day.

The Village of Waunakee, up until December, 1971, employed

a trickling filter to treat sanitary sewage from the community.

A 24 hour survey in the summer of 1962 revealed a flow of 148,500

gal/day which contained a BOD of 212 Ibs/day and the following

concentrations of nitrogen: organic N, 7.4 mg/1; NH4+— N, 15.6

mg/1; N02~— N, 1.64 mg/1; N03~— N, 3.64 mg/1 (Lee et al,

1966). A survey in 1970 showed a flow of 220,000 gal/day and a

BOD of 121 Ibs/day (Wisconsin Department of Natural Resources,

1971). In 1960, the Village of Waunakee had a population of 1,611,

while the 1970 census indicated the population had increased to

2,181.

The Waunakee Cheese Factory disposes of its milk wastes in a

seepage lagoon within the Village of Waunakee. Sodium nitrate

has been added at times to aid in odor control. The Waunakee Can¬

ning Company also disposes of wastes seasonally by means of spray

irrigation. It has been reported (Wisconsin Department of Natural

Resources, 1971) that runoff from the spray irrigation sometimes

enters the drainage course.

1 Since preparation of this report this waste water has been diverted to the

Madison Metropolitan sewage system.

152 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

In Madison the Avenue Car Wash rinse .water is no longer a

nutrient source as it was diverted to the sewage line and thus

around the Madison lakes prior to 1970. However, a new nutrient

source not reported by Lee et at. (1966) is the Pure Oil Truck

Plaza, which is located along the interstate highway system. The

plaza discharges its treated domestic wastes to Token Creek, a

tributary to Lake Mendota. The wastes are treated by a package

aeration plant and are chlorinated before discharge (Wisconsin

Department of Natural Resources, 1971). The truck stop dis¬

charges 10,000 gal/day of treated effluent, which has a BOD of

about 5 lbs/ day. Since the volume of domestic waste water in the

United States averages 100 gpdc (Fair et al, 1968) a population

equivalent of about 100 is calculated.

Since little additional information is available on the industrial

sources of nutrients, the original estimate will continue to be used

minus the contribution (625 lbs/year of phosphorus) from the

Avenue Car Wash. Thus, the estimated total contribution from

industrial waste sources for 1970 was about 17,000 lbs of nitrogen

per year and 9,000 lbs of phosphorus per year as shown in Table 8.

These values will be assumed to represent total-N and total-P,

although the form of the nutrients was not specified by Lee et al.

(1966). Since the municipally treated wastes from the Forest Milk

Company comprise most of the industrial loading, it is probable

that organically combined forms of the nutrient were important.

Hence, half of the total-N and total-P will be estimated to have

been in an organic form.

The contribution of nutrients from treated domestic wastes was

computed by Lee et al. (1966), from a per capita annual contribu¬

tion of 1.7 lbs of phosphorus and 7 lbs of nitrogen. Although it

TABLE 8. INDUSTRIAL WASTE SOURCES ENTERING

LAKE MENDOTA— 1970

(After Lee et al. (1966)

Total _ 9,300 16,800

1974] Sonzogni and Lee — Nutrients for Lake Mendota 153

was not specified in the original report (Lee et al ., 1966), these

numbers referred to soluble-orthophosphate and inorganic

nitrogen.

In order to gain insight as to the quantity of the different forms

of nutrients in the effluents, the previously discussed results of

24 hour nutrient surveys of the Waunakee and Windsor treatment

plant effluents may be utilized. Assuming that the concentrations

reported were typical, the annual per capita nutrient contributions

from these sources have been estimated. The results are listed in

Table 9. The Nine-Springs Sewage Treatment Plant, which pro¬

vides primary and secondary treatment for wastes from Madison,

has an effluent with an average annual per capita contribution of

8.5 lbs of inorganic nitrogen and 3.5 lbs of soluble phosphorus

(Mackenthun and Ingram, 1967).

Based on these limited data, the following annual per capita

contributions will be used to reestimate the quantity of nutrients

supplied by treated sewage effluent: soluble ortho-P, 3 Ibs/cap/yr;

total-P, 4.5 Ibs/cap/yr; inorganic-N, 6 Ibs/cap/yr; organic-N,

2 Ibs/cap/yr; total-N, 8 Ibs/cap/yr. Using these values and the

latest census information or population estimates, the 1970 annual

contribution of nutrients from treated domestic wastes has been

computed and the results are presented in Table 10.

The revised estimates of total nutrients contributed by waste

water in 1970 are compared to the originally published contribu¬

tion in Table 11. It can be seen that the revised estimates are

considerably higher than those previously reported for both phos¬

phorus and nitrogen.

One other waste water source must be considered, that being

the discharge from private domestic disposal systems (septic

TABLE 9. ANNUAL PER CAPITA NUTRIENT CONTRIBUTIONS FROM

THE WAUNAKEE SEWAGE TREATMENT PLANT AND THE

WINDSOR TREATMENT PLANT (SANITARY DISTRICT #1)

Waunakee1 2 Windsor'

lbs/capita/year

1 Based on a 24 hour study in 1962

2 Based on a 24 hour study in 1969

154 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 10. ESTIMATED NUTRIENT LOADING TO LAKE MENDOTA

FROM TREATED DOMESTIC WASTES FOR 1970

Estimated Soluble Inorg Org

Source Population O — P T — P N N T — N

Waunakee _ 1,911 5,700 8,600 11,500 3,800 15,300

DeForest _ 2,181 6,500 9,800 13,100 4,400 17,500

Windsor _ 550 1,600 2,500 3,300 1,100 4,400

Truck Plaza _ 100 300 400 600 200 800

Total _ 4,742 14,100 21,300 28,500 9,500 38,000

TABLE 11. PREVIOUSLY PUBLISHED (LEE ET AL. 1966) AND 1970

ANNUAL NUTRIENT LOADING TO LAKE MENDOTA FROM

MUNICIPALLY TREATED WATERS

Lee et al. Revised1

(1966) (1970)

(lbs/year)

1 To the nearest 1000 pounds

tanks) . Septic tanks are generally a problem only if they become

plugged, which is likely to occur fairly frequently in non-sandy

soils, found in much of the Mendota basin. When this occurs sur¬

face discharge may result and nutrients in the effluent can be

transported via overland flow to the lake. Lee et al. (1966) esti¬

mated the annual discharge by assuming a population of 500

having malfunctioning septic tanks and by using the same nutrient

loading rate as used for treated domestic waste. Again, the form

of the nutrients supplied by this source was not specified by Lee

et al. (1966). In revising this source, no organic forms will be

considered. The per capita contribution will be assumed to be the

same as for municipally treated domestic sewage. Therefore, again

estimating a population of 500 (this is perhaps an overestimation,

the number of malfunctioning septic tanks being unknown), the

source should provide about 1500 lbs of soluble ortho-phosphorus

and 3,000 lbs of inorganic nitrogen.

The total annual nutrient loadings as of 1970, from the three

types of waste effluents — municipally treated domestic wastes,

privately treated domestic wastes, and treated industrial wastes —

1974] Sonzogni and Lee — Nutrients for Lake Mendota 155

TABLE 12. PREVIOUSLY PUBLISHED AND 1970 ANNUAL NUTRIENT

LOADING TO LAKE MENDOTA FROM WASTE WATER EFFLUENT

Lee et al.

(1966) Revised1

(lbs/ year)

1 To the nearest 1000 pounds

are compared to the previous published estimates of Lee et al

(1966) in Table 12. The revised estimate represents more than a

50 percent increase over the previously-predicted loading.

Beginning in December, 1971, the Madison Metropolitan Sewer¬

age District completed an extension of their system which trans¬

ports the waste water of Waunakee and DeForest to the Nine

Springs treatment plant so that the waste effluent from these

communities is now diverted around Lake Mendota. Therefore,

with the completion of the extension to Windsor, which is sched¬

uled for the near future, all major waste water sources will be

eliminated. The estimates of the relatively insignificant waste

water nutrient sources remaining after diversion are presented in

Table 13.

TABLE 13. REMAINING WASTE WATER NUTRIENT SOURCES FOR

LAKE MENDOTA FOLLOWING ELIMINATION OF TREATED

WASTE WATER DISCHARGES FROM WAUNAKEE,

DeFOREST AND WINDSOR

2,000 6,900

Total

TABLE 14. ESTIMATED NUTRIENT SOURCES FOR LAKE MENDOTA— REVISED

(PRIOR TO DIVERSION OF WASTE WATER EFFLUENTS)

156 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

inorganic nitrogen

Negligible

1974] Sonzogni and Lee — Nutrients for Lake Mendota 157

Summary of Revised Estimates

The revised annual nutrient contributions from the various

sources (prior to the diversion of waste water effluents) have been

summarized in Table 14. It can be seen that the revised annual

contributions of nitrogen and phosphorus have been more than

doubled, as compared with the old estimates.

In the case of phosphorus, the per cent contribution of the vari¬

ous sources changed little. The major changes were in rural run¬

off, which increased from 42% to 50%, and in municipal and in¬

dustrial waste water, which decreased from 36% to 23%. In the

case of nitrogen, the major changes were in rural runoff and

ground water seepage. The contribution of rural runoff increased

from 11% to 42%, while the ground water contribution from

direct seepage decreased from 36% to 14% in the revised estimate.

In computing the annual contribution from rural runoff, average

lbs/acre values from Table 3 (discussed in a previous section) were

used to compute total nutrient input and the per cent estimated

contribution in Table 14. However, because rural runoff appears to

be the major nutrient source, a possible range of values was also

tabulated in Table 14 (see previous section on rural runoff). Even

if the minimum values suggested were used, the percentage of

nutrients contributed by rural runoff would still be significant and

the sum total nutrient input in lbs/year would remain greater than

the previous estimate of Lee et al. (1966). For example, if the

minimum predicted contribution of total phosphorus from rural

runoff (as listed in Table 14) were used to predict the annual phos¬

phorus input, the total input would decrease to 81,000 lbs/year,

still higher than the previously published estimate. However, the

total phosphorus contribution of municipal and industrial wastes

would increase to nearly 40%, while the rural runoff contribution

would decrease to about 20%. On the other hand, if the maximum

possible contribution for rural runoff were used to compute the

total phosphorus input, rural runoff would be by far the dominant

source of phosphorus.

Estimated nutrient sources for Lake Mendota after the diver¬

sion of waste water effluent from the communities Waunakee, De-

Forest, and Windsor are given in Table 15. It can be seen that the

diversion will decrease the phosphorus loading by about 20% and

the nitrogen loading by about 4%.

Finally, it should be emphasized that the revised estimates are

predicated on many assumptions and are, at best, still only rough

approximations. This is especially true of the total nitrogen load¬

ing, since the estimate of nitrogen influx due to groundwater seep¬

age, although listed as a major nitrogen source in Table 15, is based

on very limited information and is subject to considerable error.

158 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 15. ESTIMATED NUTRIENT SOURCES FOR LAKE MENDOTA

FOLLOWING ELIMINATION OF TREATED WASTE WATER

DISCHARGES FROM WAUNAKEE, DeFOREST AND WINDSOR

Waste water discharges _

Urban runoff _

Rural runoff _

Atmospheric precipitation on lake surface

Dry fallout on lake surface _

Ground water seepage _

Base flow _

Nitrogen fixation _

Woodland runoff _

Marsh drainage _

1 Negligible.

% Estimated Contribution

T-P T-N

2

14.5

63

2

6.5

1

11

01

0

1

6

43.5

6

11

14

11

7.5

0

Nonetheless, the revised estimates are believed to be the most cur¬

rent and reasonable estimates based on the data available.

Control of Nutrient Sources

The most easily controlled sources of nutrients are the point

sources, such as the discharge of waste water. In effect, the major

point sources for Lake Mendota were controlled with completion

of the sewage interceptor which diverted waste water from Wau-

nakee and DeForest to the Nine Springs Plant. Further elimina¬

tion of waste water sources, such as the Town of Windsor sewage

effluent or the effluent from the Pure Oil Truck Stop, will have a

relatively minor effect on the total nutrient flux to the lake. In fact,

the diversion of waste water from Waunakee and DeForest re¬

sulted, in the case of phosphorus, in a nutrient input reduction of

only about 20%. This is not to say that the elimination of these

sources is not of value. On the contrary, their elimination serves

as a preventive measure to avoid future degradation of the water

quality of the lake. This degradation could ensue if waste water,

which would steadily increase in amount as a result of the rapidly

expanding population on the north and west side of the lake, were

to continue to be discharged into the lake.

Since nutrients entering Lake Mendota are derived to a sig¬

nificant extent from a variety of sources (such as urban storm

water drainage and agricultural runoff), increased attention should

be given to the control of these diffuse sources of phosphorus. Be¬

cause of their diffuse nature, chemical treatment of these sources

will be extremely expensive, quite probably prohibitive. Rather

1974] Sonzogni and Lee — Nutrients for Lake Mendota 159

than attempting to collect and treat drainage waters from urban

and agricultural areas, a more fruitful approach might be to study

in detail specific sources of nutrients in urban and agricultural

areas, and then attempt to control the specific source at its origin.

For example, it is known that urban storm water drainage con¬

tains large concentrations of phosphorus. At the present time,

essentially nothing is known about the specific sources of phos¬

phorus in the urban environment. The mass balance approach needs

to be made in a number of urban communities throughout the

United States in order to determine the relative significance of

lawn fertilization, gasoline combustion, dust fall, terrestrial plants,

etc., as sources of phosphorus in urban areas.

As mentioned previously, Kluesener (1972) noted that the high¬

est concentrations of phosphorus in the urban stormwater drain¬

age were associated with the leaf fall period during the fall and

Cowen and Lee (1971), in a follow-up study on leaves as a source

of phosphorus, found that large amounts of phosphorus are readily

leachable from dead leaves. It appears, based on these preliminary

studies, that more effective leaf pick-up during the fall might min¬

imize the amounts of nutrients derived from this source in urban

stormwater drainage.

Agricultural sources of nutrients could be controlled through

education of the farmer, in terms of when to apply nutrients, what

concentrations to apply and how to control soil erosion and other

conditions that tend to promote high nutrient fluxes from farm¬

land. The manure problem that exists in Wisconsin and other

nearby states can be controlled by having the farmer store the

manure in tanks over the winter period and then spread it on the

land after the ground has thawed in the spring.

One of the methods beginning to be used for controlling agri¬

cultural and urban stormwater drainage is zoning for land use. In

Wisconsin there has been recent action by the Dane County Plan¬

ning Commission which is specifically designed to prevent urban

development in Lake Mendota’s watershed because of the fear that

the conversion of farm land to urban areas would increase the

nutrient flux to this lake. Further, Illinois is considering legisla¬

tion which prevents certain types of fertilizer application on lands

with a slope greater than a certain degree. Another example of

this type of development is the current legislation being considered

on the Lake Mendota drainage basin. The proposed guidelines

would prevent the winter spreading of manure from dairy opera¬

tions and the draining of marshland for agricultural or urban

use. It is likely that these developments in Wisconsin and Illinois

will set the pattern for similar developments throughout North

America.

160 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

It is of interest to examine the potential effect of future urbani¬

zation of Lake Mendota’s watershed. The Dane County Planning

Commission (private communication, 1972) has estimated that the

urban area within the Lake Mendota watershed will roughly double

in size by 1990 (the increase will vary somewhat according to the

growth rate projection used). Assuming the urban area doubles

by 1990 at the expense of rural land, an estimate of the total phos¬

phorus loading can be obtained using loading factors of 0.6

Ib/acre/year (Table 3) for rural runoff and 1.0 lb/acre/year for

urban runoff (Table 6) can be made, if other nutrient sources

remain at the same rate as present. In this manner it is predicted

that urban runoff will contribute 28% of the annual total phos¬

phorus loading, while rural runoff will contribute 51% in 1990.

These percentages compared with 14.5 and 63% respectively, for

1972 (Table 15), indicate that the sum total annual phosphorus

loading from all sources should increase from the present rate of

109,000 pounds per year to 115,000 pounds per year in 1990, an

increase of about 6%. The total loading predicted for 1990 is still

less than the total loading estimated for 1970 (prior to the 1971

diversion). Therefore, based on the above analysis, it does not

appear that doubling the urbanized area within the Lake Mendota

watershed will have a major effect on the water quality of Lake

Mendota.

It should be noted that not all conversions of agricultural lands

to urban areas would necessarily result in an increase in nutrient

flux, since some rural areas, such as heavily manured lands, could

conceivably result in little or no decrease in the nutrient flux when

the land is converted to urban areas. On the other hand, the effect

of urbanization would likely be of greater magnitude than reported,

if the new urban areas were at one time marshland, since, as was

discussed previously, the drainage of marshes results in a very

significant release of nitrogen and phosphorus to the drain water

(Lee et al., 1971). This release probably takes place over a period

of several years and it is estimated that marshes of the type found

in southeastern Wisconsin would yield on the order of 40 pounds

of phosphorus per acre.

Nutrient Availability

In addition to determining the amounts of nutrients derived

from various types of activities of man in urban and agricultural

areas, studies must be initiated to determine what part of the

nutrients from these sources is or can be made available for aquatic

growth. For example, some urban and agricultural activities con¬

tribute large amounts of total phosphorus ; however, a large por-

1974] Sonzogni and Lee — Nutrients for Lake Mendota 161

tion of this phosphorus is in a form, i.e., particulate, organic, etc.,

that is not immediately available for the growth of algae. A much

better understanding must be achieved on the aqueous environ¬

mental chemistry of particulate, inorganic, and organic forms of

nutrients in order to ascertain the real significance of the various

forms of nutrients derived from urban and agricultural areas in

stimulating the growth of aquatic plants and algae. Studies along

this line are currently underway at the University of Wisconsin

Water Chemistry Program.

In the case of phosphorus, about the best that could be done

at this time is to state that in most instances the available phos¬

phorus is somewhere between soluble orthophosphate and the total

phosphorus. The fraction of organic and particulate phosphorus

that becomes available in natural waters is extremely important

in designing meaningful eutrophication control programs for dif¬

fuse sources of nutrients. There is little point in attempting to

control particulate forms of phosphorus when it is known that the

phosphorus would not become available for algal growth under

the conditions existing in the receiving waters. The potential avail¬

ability of the nitrogen species are even less well known.

ACKNOWLEDGEMENT

This investigation has been principally supported by the En¬

vironmental Protection Agency, Grant No. R-801360 and by the

University of Wisconsin Department of Civil and Environmental

Engineering. Also, we wish to acknowledge the assistance given by

Dr. P. Uttormark, Associate Scientist, University of Wisconsin,

in providing a bibliography that he had prepared on the amounts

of nutrients derived from runoff. Finally, the cooperation of the

Dane County Planning Commission, which provided land use in¬

formation, is appreciated.

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AN EVALUATION OF THE USE OF THE EEG TECHNIQUE

TO DETERMINE CHEMICAL CONSTITUENTS IN

HOMESTREAM WATER

Jon C. Cooper

University Wisconsin — Madison

G. Fred Lee

University Texas — Dallas

and

Andrew E. Dizon

National Marine Fisheries Service

Honolulu — Hawaii

INTRODUCTION

A large body of literature now exists that points to olfaction

in migratory fish as the important sense in orientation near and

in the homestream (Collins et al., 1962; Fagerlund et aL, 1963;

Groves et al., 1968; Hasler, 1960a, 1960b, and 1966; Hasler and

Wisby, 1951). It is hypothesized that salmon can store odor infor¬

mation about the homestream and use these cues upon the homing

migration.

The chemical or chemicals involved in homestream cues are

probably of a very low concentration and perhaps a complex mix¬

ture. Efforts to determine these chemicals have met with some

success. Fagerlund et al. (1963) found that a portion of these

chemicals is volatile, although the non-volatile portion may play

some part in orientation for the fish. Hasler (1966) reported that

the active fraction was organic, heat labile and volatile at 25 C.

Idler et al. (1961) concluded that the material was neutral, dialyz-

able and heat labile.

Hara et al. in 1965 reported that homestream water perfused

through the nares of salmon produced a characteristic high ampli¬

tude wave in an electroencephalographic (or EEG) recording from

the olfactory bulb. The EEG, they suggested, might be used to study

on a physiological basis the olfactory hypothesis.

Since the EEG had been looked on as a possible bioassay for

individual homestream recognition (Oshima et al., 1960a and b;

Ueda et al., 1967), it was felt that the EEG might provide useful

information about homestream chemicals that were needed for

homing. Because the technique is quite recent, this problem was

first approached by repeating earlier experiments by Fagerlund

et al., Hasler, and Idler et al. Although the active fractions de-

165

166 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

scribed by these workers may not be the same as the stimulants in

the EEG experiments, it was felt that there should be some

correspondence in the groups of chemicals described in the two

types of experiments. This paper reports on an investigation de¬

signed to evaluate the feasibility of using EEG as a means of de¬

tecting the chemicals present in homestream water that are

responsible for the homing of coho salmon in a Wisconsin stream

bordering on Lake Michigan.

METHODS

Adult spawning coho salmon (Oncorhynchus kisutch) that had

homed to a tributary of the Ahnapee River, Algoma, Wisconsin

in the fall, 1970, were used in these studies. They were trapped

in the stream on the same day as the experiments and brought back

to a temporary laboratory, where they were held in city tap water.

The testing procedure was similar to that of Dizon et al. (1973) .

The fish were anesthetized with tricain methanesulphonate

(MS222, 0.01) and immobilized with gallamine triethiodide

(flaxedil, 2 mg/kg body weight). A portion of the brain was ex¬

posed by means of a dental drill ; a platinum coated stainless steel

electrode (Transidyne General) was inserted in the olfactory bulb.

EEG responses evoked by test samples were amplified with a Bio¬

electric Instruments (model DS2c) and recorded on a Hewlett

Packard model 7712B oscillagraph. Figure 1 presents a typical

recorder trace showing the background and stimulus response. A

second channel of the oscillagraph was equipped with an integrat¬

ing preamplifier, so that the integration of the EEG could be

recorded. The integrator sums the voltages in the positive part of

the wave form. The slope of the line obtained from the integrator

can be used to quantify the EEG records. To standardize the

response, the slope of the integration of a response to each sample

was divided by the slope of the integration of the response to home-

0.50 MV

FIGURE 1. Example of electroencephalographic output (hand wash).

BACKGROUND STIMULUS

RESPONSE

10 sec

1974] Cooper, Lee and Dizon — Use of EEG Technique 167

stream water. An F-test was used to test the significance of the

responses. Each sample was introduced in a random order one time

each trial. Trials were repeated 4 times. Algoma city tap water was

used to rinse the nares between trials.

Water samples from the homestream were fractionated in a

variety of ways. Factors considered were : the variability of the

response of a fish to a given sample and the pH, molecular weight,

and volatility of the sample. Additional studies on the chemical

nature of the homestream dealing with carbon filtration, dialysis,

and Sephadex chromotography were not conclusive and will not

be reported here. Experiments with the ionic strength and concen¬

tration of the homestream water will not be reported for the same

reason. For further details consult Cooper (1971).

In the first experiment, two samples of stream water, one taken

directly from the stream and one stored at 5 C for one day, were

used to test two fish for their variability in response.

In the second experiment, a group of samples of different pH

were prepared by adding sodium bicarbonate (0.01M) to Algoma

city tap water and adjusting the pH of the solutions with sulfuric

acid or sodium hydroxide, as needed, to pH 5, 6, 7, 8, and 9. One

fish was used in this experiment.

In a third experiment, homestream water was filtered through a

glass fiber filter and 0.45 micron and 0.22 micron pore size Millipore

filters. Four fish were used in this experiment.

Finally, in a fourth experiment, homestream water was frac¬

tionated by means of a vacuum distillation apparatus at 6 mm Hg

pressure at 20 C. This equipment consisted of a Snyder-ball column

and a water- jacketed condenser . A thermometer was positioned at

the top of the column to observe the temperature of the distillate.

A dry ice acetone bath was used to trap the distillate. It took 3.5

hours to reduce 750 ml to 250 ml under these conditions. Two fish

for each of 2 distillations were used in this experiment.

For the experiments reported here, a total of 10 fish and 15

water samples were used.

RESULTS

The results of the first experiment (Figure 2 and Table 1) indi¬

cate that the variability in fish response as reflected in the standard

deviation was roughly 25%. Data from different animals cannot

be compared directly, i.e. the data from coho 144 and 146 cannot

be pooled, although the ranking for samples for each fish can be

compared.

In the second experiment, filtration through glass fiber filter,

0.45 micron or 0.22 micron pore size Millipore filters did not seem

168 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

day at 5 C (3).

TABLE 1. REPLICATE SAMPLES OF RAW STREAM WATER AND

STREAM WATER STORED AT 5 C

(Mean and standard deviation of the integration of response to the stimuli

expressed as the per cent of the integration of the response of a reference

sample of stream water.)

1974] Cooper, Lee and Dizon — Use of EEC Technique 169

to affect the samples. The amplitude of responses to these samples

was roughly the same before or after filtration (Table 2).

The salmon responded most strongly to acid pH (4 and 5). They

responded less strongly to basic pH (8 and 9) than to neutral pH

(7) (Figures).

In the fourth experiment, it is clear that there is a higher ampli¬

tude response to the non-volatile portion than to the volatile portion

of the water distilled at 20 C (Table 3).

TABLE 2. EEG RESPONSES TO WATER FILTERED THROUGH GLASS

FIBER FILTER, 0.45 AND 0.22 MICRON PORE SIZE

MILLIPORE FILTERS

Coho Coho Coho Coho

FIGURE 3. EEG response to homestream water adjusted to pH, 5, 6, 7, 8,

and 9 (mean and standard deviation).

170 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 3. VACUUM DISTILLATION OF HOMESTREAM WATER.

DUPLICATE RUNS ON EACH FISH FOR TWO SETS OF

DISTILLATION SAMPLES

(Mean and standard deviation of the integration of the response to the stimuli

expressed as the per cent of the integration of the response of a reference

sample of stream water.)

DISCUSSION

One problem in interpreting the data is that one cannot make

inter-fish comparison. This may be due to difference in handling of

the fish. Coho salmon were taken out of the trap at one time on

the day of the experiment. It is possible that fish had been in the

trap for different lengths of time for a few minutes to several days.

Ueda et al. (1967) have pointed out that fish kept in a holding tank

for a week became unresponsive to their homestream water; this

may be due to an advance stage of sexual “Ripening” (as Fager-

lund et al, 1963 suggests) or a gradual acclimation of the fish to

the homestream water odor.

The results of the third experiment show that the homestream

odor has a molecular weight under one million, the approximate

size of material retained on a 0.22 micron pore size Millipore filter.

Since none of these filters seems to add an odor to the homestream

(or subtract one from the water), they are a useful technique for

cleaning up the extraneous matter in the water ; there was a large

quantity of material left on the filters after the experiments. The

final experimental results indicate that the stimulatory portion of

the water is non-volatile at 20 C, since there is a higher response

to the residue than to the distillate.

The second and fourth experiments mentioned above leave little

doubt that the EEG technique can be used to detect difference in

water samples; it is possible to determine whether the character

of the water sample has been changed by the experimental pro¬

cedures. However, since the standard deviation in response is quite

large, the technique may not be useful in detecting subtle differ¬

ences between samples.

A second underlying problem that may limit the usefulness of

the EEG technique, is that it is not possible to distinguish between

1974] Cooper, Lee and Dizon — Use of EEG Technique 171

two kinds of responses from the fish. A sample that represents a

danger or avoidance reaction, such as handwash (a sample in which

the hand has been dipped) will give the same amplitude response,

as far as one can tell from the EEG, as a sample that is stimula¬

tory, such as the homestream water. Therefore, it is impossible

to tell whether the distillation experiment, for instance, produced

residues important in homing or whether merely concentrated

“avoidance” fractions were formed. Indeed, the stimulatory frac¬

tions observed with the EEG technique may not be related in any

simple manner to either the stimulatory fraction found in be¬

havioral work or those fractions actually necessary for homing.

Perhaps in the future, any experimental results obtained with the

EEG technique should be confirmed with a behavioral bioassay.

Until these problems are overcome, any future work that at¬

tempts to utilize the EEG to determine homestream constituents

may be of limited value.

ACKNOWLEDGEMENT

We wish to acknowledge the assistance of A. D. Hasler and Ross

Horrall of the University of Wisconsin Laboratory of Limnology.

Their assistance in this study was greatly appreciated.

This work was supported by training grant number 5T1-WP-22

from the Environmental Protection Agency, the Department of

Civil and Environmental Engineering, University of Wisconsin.

BIBLIOGRAPHY

COLLINS, G. B., P. S. TREFETHEN, and A. B. GROVES. 1962. Orienta¬

tion of homing1 salmon. Am. Zoologist 2:399-400.

COOPER, J. C. 1971. The determination of chemical constituents by EEG

of homestream water of coho salmon ( Oncorhynchus kisutch), M.S. Thesis,

University Wisconsin — Madison.

DIZON, A. E., R. M. HORRALL, and A. D. HASLER. 1973. Long-term

olfactory “Memory” in coho salmon, Oncorhynchus Kisutch. Fish. Bui.

71:315-317.

FAGERLUND, U. H. M., J. R. McBRIDE, M. SMITH and N. TOMLINSON.

1963. Olfactory perception in migrating salmon. III. Stimulants for adult

sockeye salmon (Oncorhynchus nerka) in homestream waters. J. Fish.

Res. Bd. Can. 20:1457-1463.

GROVES, A. B., G. B. COLLINS, and P. S. TREFETHAN. 1968. Roles of

olfaction and vision in choice of spawning site by homing adult chinook

salmon ( Oncorhynchus tshawytscha) . J. Fish. Res. Bd. Can. 25:867-876.

HARA, T. J., K. UEDA, and A. GORBMAN. 1965. Electroencephalographic

studies of homing salmon. Science 149:884-885.

HASLER, A. D. 1960a. Homing orientation in migrating fishes. Ergebn. Biol.

23:94-115.

HASLER, A. D. 1960b. Guideposts of migrating fishes. Science 132:785-792.

HASLER, A. D. 1966. Underwater Guideposts. Univ. Wisconsin Press, 155 pp.

172 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

HASLER, A. D. and W. WISBY. 1951. Discrimination of stream odors by

fishes and its relation parent stream behavior. Am. Nat. 85:223-238.

IDLER, D. R., J. R. McBRIDE, R. E. E. JONES, and N. TOMLINSON.

1961. Olfactory perception in migrating salmon. II. Studies on a labora¬

tory bioassay for homestream water and mammalian repellent. Can. J.

Biochem. Physiol. 39:1575-1584.

OSHIMA, K., W. E. HAHN, and A. GORBMAN. 1969a. Olfactory discrimina¬

tion of natural water by salmon. J. Fish. Res. Bd. Can. 26:2111-2121.

OSHIMA, K., W. E. HAHN, and A. GORBMAN. 1969b. Electroencephalo-

graphic olfactory responses in adult salmon to waters traversed in the

homing migration. J. Fish. Res. Bd. Can. 26:2123-2133.

UEDA, K., T. J. HARA, and A. GORBMAN. 1967. Electroencephalographic

studies on olfactory discrimination in adult spawning salmon. Comp.

Biochem. Physiol. 21:133-143.

STUDIES ON THE Ca, Mg, AND Sr CONTENT

OF FRESHWATER CLAMSHELLS

G. Fred Lee

University Texas— Dallas

and

William Wilson

University Wisconsin — Madison

ABSTRACT

The relationship of Ca, Mg and Sr in Lakes Mendota, Fox and

Trout in Wisconsin and clamshells collected from these lakes shows

a negative correlation between shell and water Sr/Ca ratio. The

Mg/ Ca ratio showed considerable scatter and in general a direct

relationship between shell composition and water composition.

INTRODUCTION

As part of a study on the feasibility of using the Ca, Mg and Sr

content of freshwater clamshells as indicators of paleohydrologic

conditions, it was necessary to investigate the relationship of fresh¬

water clamshell and water composition. Since river and stream

chemical composition varies with discharge, it was decided to use

lake clams, since the water composition should be relatively con¬

stant. A literature review on the relationship between clamshell

and water composition has been presented by Lee and Wilson

(1969).

EXPERIMENTAL PROCEDURE

The Ca, Mg and Sr contents of lake water and clamshells were

determined by atomic absorption spectrophotometry according to

the procedure described by Lee and Wilson (1969) . The clams were

taken alive by wading along the beach in waist-deep water. The

shells were cut into sections and ashed to separate into laminar

layers. Only the laminar layers were analyzed. Except as noted, the

laminar layers were analyzed as a group.

RESULTS

A preliminary phase of this study was concerned with the deter¬

mination of the variability of the calcium content of lake clams.

This was undertaken because there was no information on their

173

174 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

chemical components, as all previous work has been concerned

with specimens found in rivers, streams and marine environments

(Nelson, 1963, 1967; Odum, 1957; Thompson and Chow, 1955).

Sixteen clams of the same species (Lampsilis siliquoidea rosacea)

were collected live from Lake Mendota. The laminar layers of each

of the shells were analyzed for their calcium content. The sample

had a mean and standard deviation calcium content per gram of

shell weight of 409 ± 6 mg/g. As with other aragonite shells, lake

clamshells are principally calcium carbonate with the addition of

trace elements.

The total sample of lake clams (N = 75) yields a mean and

standard deviation of 413 ± 10 mg Ca/g of shell weight. When

this is compared with a series of river clams analyzed by Nelson

(1963), where 23 clams had a mean and standard deviation of

400 ± 1.4 mg Ca/g of shell, the difference between the means is

statistically significant (calculated t = 10.8 with 109 degrees of

freedom). Further, Nelson (1963) compared the means of river

clams with the data of Thompson and Chow (1955) on 64 marine

shells, which had a mean and standard deviation of 392 ± 1.2 mg

Ca/g of shell. This gave a statistically significant t of 4.3 with 85

degrees of freedom. The differences between lake, river and marine

shells (in a system that is almost pure calcium carbonate) , would

suggest that the environment is an important factor in the secre¬

tion of elements within the clamshell. Therefore, the clams of 1

species from 1 lake appear to have an essentially constant calcium

concentration. However, the data have indicated that clams may

vary according to their environmental situation. Because of this

variability, calcium analyses have been included for every sample.

This procedure insures a more accurate measurement of the ratios

of calcium with other cations.

Studies were conducted to examine the variability of shell among

lake clams. A live clam (Lampsilis siliquoidea rosacea) was taken

from Lake Mendota; its shell after ashing was separated into

areas shown in Fig. 1. The morphological parts were analyzed

separately. Examination of the results, as presented in Table 1,

shows that differences occur between layers within each part of

the shell. For example, the Sections 3-7 (laminar layers) have

a mean and standard deviation Ca, Mg and Sr concentration of

406 ± 15 mg/g, 59 ± 15 ,/xg/g and 146 ± 17 yg/g, respectively.

Differences occur between parts of the shell. The peripheral layers

(Sections 9-14) contain about 70% as much Sr as Sections 3-7.

This variability found in lake clams in both concentration and

atomic ratios was similar to that reported by Nelson (1964) for

river clamshells.

1974] Lee and Wilson— Freshwater Clamshell Studies

175

FIGURE 1.

Cross section of clamshell analyzed. Sections 1-7 were the

laminar layers from inside toward outside.

The laminar layers are secreted as annual growth layers. The

peripheral layers are often damaged, encrusted with foreign mat¬

ter, and the parts near the umbo usually show signs of abrasion

and wear. Because of the large differences found between various

morphological parts of the shells, all further analyses have been

confined to the laminar layers. These layers shown in Fig. 1 as

Sections 3-7 represent that part of the shell least likely to be

affected by post-depositional factors and, therefore, would most

closely reflect the living clam’s chemical milieu at the time of depo¬

sition of each laminar layer.

Two shells (Lampsilis siliquoidea) , from each of 3 Wisconsin

lakes, were collected to examine the composition of the laminar

layers in a multiple sample and from different environments. The

specimens were separated into laminar layers and analyzed for

Ca, Mg and Sr content. The data in Table 2 show that the laminar

layers of the clams from each lake have marked differences in

TABLE 1. COMPOSITION OF LAKE MENDOTA CLAMSHELL

(Lampsilis siliquoidea rosacea)

176 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 2. CONCENTRATION OF CALCIUM, MAGNESIUM AND

STRONTIUM IN THE LAMINAR LAYERS OF LAKE CLAMSHELLS

Lampsilis siliquoidea

composition. This variability displays the typical heterogeneous

distribution of trace elements in mollusks. Even in this very small

sample, there appear to be differences in the atomic ratios from

one lake to the next, although the samples are too small for com¬

parative purposes.

The variability found in this study is like that mentioned previ¬

ously (Curtis and Krinsley, 1965), where a large number of

samples must be studied to define any relationship between shell

and water composition. Similar conclusions must be reached as

a result of this study. A large number of clams of 1 genus and

preferably of 1 species must be analyzed from a number of lakes

of differing composition before a relationship can be defined.

Inasmuch as the nature of this work is concerned with the mean

chemical composition of groups of clams in relationship to their

environment, all further analyses were conducted on all the lami¬

nar layers as a whole. The pretreatment of the shell was modified,

such that after ashing the laminar layers were separated from

the other parts of the shell. The layers were then dissolved as a

group in HC1 and analyzed.

Three Wisconsin lakes were utilized in this study on the basis

of the availability of clamshells and water samples. In this group

1974] Lee and Wilson — Freshivater Clamshell Studies 177

there are 2 hard-water lakes (Fox and Mendota) and 1 soft-water

lake (Trout). A total of 29 clamshells from Trout Lake, 30 from

Fox Lake and 16 from Lake Mendota, all of the species Lampsilis

siliquoidea, have been analyzed for Ca, Mg and Sr. The water ratios

are represented by the median summer values. Although large

amounts of data are available for Lake Mendota water composi¬

tion, only the summer values were available for Fox and Trout

Lakes. These values are more closely comparable for the purposes

of this study and also coincide with the expected period of maxi¬

mum growth of the clams in these lakes.

The Sr/Ca ratio of clamshells versus water, as shown in Fig. 2,

demonstrates a strong negative correlation, with a correlation

coefficient (r) of —0.83.

The Sr to Ca atomic ratio in shells is constrained because all

shells contain essentially constant amounts of calcium. It should

be pointed out that the mean Sr/g of shell increases with increas¬

ing hardness of the water. Even though Dodd (1965) demonstrated

a temperature dependence for Sr in marine pelecypods, this does

not appear to be a factor for freshwater clamshells used in this

FIGURE 2. Relationship of atom ratios of lake clamshell and water

composition. Mean and standard deviation indicated by point and bar.

178 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

study. The temperature regimes for Fox Lake and Lake Mendota

are approximately the same ; while Trout Lake, located in northern

Wisconsin, is about 2°C cooler. Therefore, the atomic ratio of

Sr/Ca in these freshwater clamshells is dependent upon the ele¬

mental composition of its environment.

The Mg/Ca atomic ratio of clamshells versus water shows a

weak positive correlation coefficient (r) of 0.4. This is the result

of large variances within each sample and is consistent with the

results obtained from the archaeological sites studied by Lee and

Wilson (1969). It is not clear why there should be such a high

degree of variability within a single species in a relatively constant

environment.

DISCUSSION

Clams are integrators of their environment. The chemical com¬

position of the shell should reflect the chemical composition of the

water. Yet, this study has shown that for the three lakes examined

there is an inverse relationship between the Sr*/Ca ratio in the

lake water and in the laminar layers of a clamshell. Part of this

variability is due to the vital effects of the clam. The study of the

single Lake Mendota clam where the shell was sectioned into vari¬

ous parts showed that there was almost as much variability be¬

tween parts in a single shell as was found between shells taken

from a beach in a single lake or shells taken from different lakes.

It is clear that the simple chemical composition of the water does

not control shell composition. Other factors must also have an

important function in determining the small differences observed

in shell composition. Some of the variability may be due to the

water itself. Although a lake is, in general, a fairly homogeneous

chemical system, it must be pointed out that the near-shore en¬

vironment is the zone of great biochemical activity and its com¬

position is much more subject to change than the open water in

most lakes. The clam’s immediate environment is often very

complex because of the active photosynthesis and respiration tak¬

ing place. Many of the clams collected from Lake Mendota and

other lakes have thick growths of the alga Cladophora attached to

them. The water taken in by the clam must pass through or near

this algal mat. During periods of light, CaC03 precipitation is

likely in hard-water lakes due to the photosynthesic pH increase.

Some of the clams taken from some lakes had 1-2 mm thick layers

of precipitated CaC03 on the outside of the shell. In the dark

the pH of the clam’s intake water should drop significantly due to

the respiration of the attached algae, recycled clam outlet water

and the respiration of microorganisms working on the detritus

1974] Lee and Wilson — Freshwater Clamshell Studies 179

that accumulates on the bottom in the vicinity of the clam. While

collecting the clams in Lake Mendota, it was noted that they often

could be located by finding the wave-sorted detritus arising from

dead leaves, twigs, etc. The reduced pH that results from respira¬

tion of organisms would be expected to cause solution of CaC03

precipitates. In addition to obtaining the Ca, Mg and Sr from the

water, the clam also may take in particulate CaC03 that arises

from photosynthetic precipitation as described above and detrital

CaC03 arising from parts of other organisms. This particulate

CaC03 could be dissolved and later incorporated in the shell.

Although few studies have been conducted on the chemical char¬

acteristics of the near-shore environment of lakes, it might be

suspected that it would show not only diurnal and seasonal changes

but also annual changes due to difference in the amounts of

macrophytes and attached algae. Evidently, based on the marked

variability of the chemical composition of annual layers of the

clamshells, it must be concluded that the clam is responding to

these changes. There is no single relationship between water and

shell composition and any study of the relationship must involve

large numbers of samples and detailed studies of the clam’s micro¬

environment.

ACKNOWLEDGMENTS

This investigation was supported by N.S.F. grant GP-5572X and

Training Grant No. 5T1-WP-22 from the Federal Water Pollution

Control Administration. In addition, support was given the project

by the University of Wisconsin Engineering Experiment Station,

the Department of Civil and Environmental Engineering, Water

Resources Center, and the U.S. Office of Naval Research. Special

appreciation is extended to R. Bryson and D. A. Baerreis for their

assistance on this project.

BIBLIOGRAPHY

CURTIS, C. D. and D. KRINSLEY. 1965. The detection of minor diagenetic

alteration in shell material. Geochimica et Cosmochimica Acta 29:71-84.

DODD, J. R. 1965. Environmental control of strontium and magnesium in

Mytilus. Geochimica et Cosmochimica Acta 29:385-398.

LEE, G. F. and W. WILSON. 1969. Use of chemical composition of fresh¬

water clamshells as indicators of paleohydrologic conditions. Ecology

50:990-997.

NELSON, D. J. 1963. The strontium and calcium relationship in Clinch and

Tennessee River mollusks, p. 203-211. In V. Schultz and A. W. Element,

Jr. (eds.) Radioecology . Reinhold, New York.

NELSON, D. J. 1964. Deposition of strontium in relation to morphology of

clam (Unionidae) shells. Verb. International Verhein. Limnology 15:

893-902.

180 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

NELSON, D. J. 1967. Microchemical constituents in contemporary and Pre-

Columbian clamshells. In S. I. Auerbach (ed.) Radiation Ecology. Oak

Ridge National Labs. 3849:43-100.

ODUM, H. T. 1957. Biogeochemical deposition of strontium. Inst. Marine

Science, Univ. of Texas, Austin. 4:38-114.

THOMPSON, T. G. and T. J. CHOW. 1955. The strontium-calcium atom ratio

in carbonate-secreting marine organisms. Deep-Sea Research, 3G: 20-39.

HYDROLOGY AND TROUT POPULATIONS OF

COLD WATER RIVERS OF MICHIGAN AND WISCONSIN1

G. E. Hendrickson

U. S. Geological Survey — •

Madison, Wis.

and

R. L. Knutilla

U. S. Geological Survey — ■

Okemos, Mich.

ABSTRACT

Statistical multiple-regression analyses showed significant rela¬

tionships between trout populations and hydrologic parameters.

Parameters showing the higher levels of significance were tem¬

perature, hardness of water, percentage of gravel bottom, per¬

centage of bottom vegetation, variability of streamflow, and

discharge per unit drainage area. Trout populations increase with

lower levels of annual maximum water temperatures, with increase

in water hardness, and with increase in percentage of gravel and

bottom vegetation. Trout populations also increase with decrease

in variability of streamflow, and with increase in discharge per

unit drainage area. Most hydrologic parameters were significant

when evaluated collectively, but no parameter, by itself, showed

a high degree of correlation with trout populations in regression

analyses that included all the streams sampled. Regression analyses

of stream segments that were restricted to certain limits of hard¬

ness, temperature, or percentage of gravel bottom showed improve¬

ments in correlation. Analyses of trout populations, in pounds per

acre and pounds per mile and hydrologic parameters resulted in

regression equations from which trout populations could be esti¬

mated with standard errors of 89 and 84 per cent, respectively.

INTRODUCTION

Trout populations in rivers respond to a variety of environ¬

mental factors, most of which are related to the hydrology of the

streams in which they live. In 1970, a preliminary analysis of

hydrologic parameters and trout populations for 16 stream seg¬

ments in Michigan showed a negative relationship between mean

annual maximum water temperatures and trout populations

(Hendrickson and Doonan, 1971). A negative relationship between

1 Publication authorized by the Director, U.S. Geological Survey.

181

182 Wisconsin Academy of Sciences, Aids and Letters [Vol. 62

I

the variability of streamflow and trout populations also was

suggested.

The present study, covering a larger area of Michigan and Wis¬

consin, was designed to analyze in greater detail hydrologic factors

that might influence trout populations. The study was also designed

to determine, if possible, the relative importance of each hydrologic

parameter on the populations. Hydrologic data were determined

for 112 stream segments in Michigan and Wisconsin for which

trout population data were available. Eighty-eight of these seg¬

ments (Fig. 1) were used to test for possible relationships between

trout populations and hydrologic parameters. Selection of the 88

segments was based entirely on accuracy and completeness of

hydrologic data. For those streams on which data were available

for many segments, some segments, generally alternate segments,

EXPLANATION

• Trout populations estimated by m a rk— an d- r ecap tu r e.

o Trout populations estimated by single survey.

FIGURE 1. Location of stream segments included in sampling.

1974] Hendrickson and Knutilla — Hydrology and Trout 183

were omitted from the analysis to avoid over-emphasis of any

particular stream.

HYDROLOGIC PARAMETERS

Hydrologic parameters that were used in this study describe

the character of a stream’s channel, bed, and banks, its streamflow

characteristics, and the quality of its water. Most of the data on

channel character were obtained by field mapping during the sum¬

mer of 1971. Data on streamflow were obtained from records for

stream gaging stations operated by the U.S. Geological Survey

and from discharge measurements that were available or were

made at the time of channel mapping. Data on water quality were

obtained from records of State agencies, the U.S. Geological Sur¬

vey, and from field analyses made at each stream segment.

Channel Character . Each stream segment was surveyed to de¬

termine its average width, depth, and surface area ; the type of bed

materials, as percentages of gravel, sand, or muck; and the per¬

centage of the stream bottom that had submerged vegetation. Also

determined were the average bank height, bank material, and

bank vegetation as percentage of hardwoods, conifers, brush, or

grass. Finally the percentage of the stream bottom which would

afford cover for trout, such as undercut banks, logs, boulders, and

overhanging brush was visually estimated.

Average gradients of the stream segments were determined from

topographic maps. Each reported gradient is the average computed

between topographic contours that cross the stream above and

below the measured stream segment. The actual gradient of the

segment may be higher or lower than the value so obtained.

Streamflow . For each stream segment the average discharge,

in cubic feet per second and cubic feet per second per square mile,

and parameters of low flow and flow duration were determined.

As an index of low flow, the median annual 7-day flow (7-day Q2)

was used. Flow-duration data included the discharges equaled or

exceeded for 10 and 90 per cent of the time.

Characteristics of streamflow were determined from stream-

flow records at gaging stations or by correlating discharge meas¬

urements made at each stream segment with records for gaging

stations. Because most discharge measurements were made when

streams were at base flow, estimates of low-flow parameters

(median minimum 7-day Q2 and 90 per cent duration) are more

reliable than those of high flow (10 per cent duration). Accuracy

of streamflow parameters also varies with the number of stream-

flow measurements available at each stream segment and with the

degree of correlation of the measurements with data for gaging

184 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

stations. The low-flow parameters for most stream segments

probably are accurate to about 20%. High-flow parameters prob¬

ably are accurate to about 30%.

The ratio of 10 per cent and 90 per cent duration discharges

was used as an index of the variability of streamflow. As an index

of the velocity of each stream segment, the median minimum 7-day

discharge was divided by the cross-sectional area of the segment.

The cross-sectional area, computed as the product of the average

width and average mid-channel depth, is greater than the actual

area, because the mid-channel depth usually is greater than the

average depth in the cross-section. Accordingly, the average veloci¬

ties tabulated are low. However, velocities are representative and

are believed to be comparable between streams.

Quality of Water. Because temperature was shown to be one

of the more significant water-quality parameters in the 1970

study, greater effort was made to obtain temperature data than

other water-quality data. Thermometers that register maximum

and minimum temperatures were installed in each of the stream seg¬

ments mapped, unless adequate temperature records were already

available at the site. Records of maximum and minimum tempera¬

tures were obtained for at least one summer month at almost all

sites. These records were correlated with temperature records at

gaging stations to obtain the mean annual maximum and mean

July temperatures. For most sites these temperatures are probably

accurate within 3 F.

Specific conductance, pH, and hardness of water were deter¬

mined in the field during the summer of 1971, usually during

base-flow conditions. Values of specific conductance probably are

accurate within about 10 micromhos; values of pH probably are

accurate to one-half pH units; and values of hardness probably

are accurate to about 20 milligrams per liter. Appearance of the

water (clear, colored, or turbid) was recorded, but numbers were

not assigned to these parameters and they were not included in

this analysis.

Another important water-quality characteristic is dissolved oxy¬

gen. One field analysis of dissolved oxygen was made at most

stream segments. However, because of diurnal variation in dis¬

solved oxygen, all values are not representative of the dissolved

oxygen content for each stream segment. Consequently, values of

dissolved oxygen were not used in the regression analysis.

TROUT POPULATION PARAMETERS

Trout population data were obtained from State fisheries

agencies in Michigan and Wisconsin. The accuracy of these data

1974] Hendrickson and Knutilla — Hydrology and Trout 185

varies widely because the purpose and method of inventory vary.

Estimates of trout in 39 stream segments were obtained by the

“mark and recapture” method. The accuracy of the results on these

segments is believed to be within about 30%. Estimates of trout

populations for other segments were obtained from one or more

stream surveys. The error in estimating these populations may

be 50% or more. However, it is believed that the errors are

random, and that the population estimates are not consistently

higher or lower than those obtained by “mark and recapture”

survey.

All trout population surveys used in this study were made in

late summer or early fall. Some of the streams sampled receive

annual plantings of trout. No attempt was made to exclude hatch¬

ery trout in the population estimates. However, large trout that

were obviously migratory spawners were excluded. Population

data were defined as number per acre, pounds per acre, number per

mile, pounds per mile, and pounds per acre foot.

RESULTS OF STATISTICAL ANALYSES

Statistical multiple-regression analyses were used to develop

relationships between trout populations and hydrologic parame¬

ters. The analyses provided a mathematical equation of the relation

between trout populations and selected hydrologic parameters.

The analyses also provided a measure of the accuracy of the de¬

fined relationships and the level of significance of each hydrologic

parameter in the relation.

Trout populations were correlated with hydrologic parameters

of channel character, fish cover, streamflow characteristics, and

water quality. For each combination of variables tested, the re¬

gression equation, multiple correlation coefficient, standard error

of estimate, and the significance of each independent variable were

calculated. Calculations were then repeated, eliminating the least

significant hydrologic parameter each time, until only the most

significant parameter remained. The procedure was repeated for

many combinations of hydrologic parameters and trout populations.

In the analyses it was assumed that the hydrologic parameters

were logarithmically related to trout populations in the linear

format. In each case this may not be entirely true. For example,

Benson (1953) has shown that the condition and growth of brook

trout in the Pigeon River (Michigan) were best when water tem¬

peratures ranged from 55 to 66 F. Also, as this and other studies

have shown, trout populations tend to increase as hardness and

specific conductance of water increase. However, it appears un¬

likely that high levels of hardness or specific conductance would

186 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

be increasingly favorable to trout populations. Other hydrologic

parameters also may have favorable ranges not yet demonstrated.

It is. believed that any deviation from the straight-line relationship

does not seriously affect the results of this study. Graphic plots

of trout populations and independent variable did not reveal any

divergency from the straight-line relationship. The probable reason

for this is that values for most parameters are within the favorable

range for trout. Also, values of many of the parameters evaluated

in the study affect trout populations in only one direction from

the favorable limits. For example, few of the streams sampled are

too cold in summer for maximum production of trout. Again, it is

unlikely that any of the streams sampled have hardness or specific

conductance greater than the optimum range for trout.

A summary of the hydrologic and trout population parameters

for which data were obtained are listed in Table 1. Shown in the

table are the degree of correlation between each of the parameters.

This simple correlation matrix was used to aid in selecting inde¬

pendent variables — variables that did not exhibit a high degree

of intercorrelation or interdependence — for regression analysis.

For example, specific conductance and hardness were not used in

the same regression analysis because of their high degree of cor¬

relation (0.97). Table 1 also shows the degree of correlation be¬

tween the dependent variables (last five rows) and each of the

independent variables. None of the independent variables, by itself,

shows a high degree of correlation with trout populations for the

units of population shown. The relation between pH and trout,

in pounds per mile, has the highest degree of correlation (0.47).

Analyses of trout populations in pounds per acre and pounds

per mile resulted in regression equations from which trout popu¬

lations could be estimated with approximately the same standard

error (89 and 84%, respectively). The equations that provide the

best relations, and for which independent basin parameters are

effective within 90-per cent confidence, are shown in Table 2.

Analyses of trout populations in pounds per acre-foot, number per

acre, and number per mile with several combinations of inde¬

pendent variables showed no strong correlation. In the analyses,

the standard error of estimate was generally in excess of 110%

(Table 2) .

To test the defined relationships for possible areal differences,

the difference between observed and computed values, termed

residuals, were calculated for each station analyzed. An analysis

of the residuals showed that the regression equations were un¬

biased areally. Reaches of some streams, and small local areas,

showed some differences or bias, but they were too small to warrant

separate analysis.

1974]

Hendrickson and Knutilla — Hydrology and Trout

187

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188 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

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1974] Hendrickson and Knutilla — Hydrology and Trout 189

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190 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

DISCUSSION

Trout populations appear to be limited chiefly by stream tem¬

perature, hardness of water, bottom materials, bottom vegetation,

variability of streamflow, and discharge per unit drainage area.

The relatively small correlation coefficients (less than 0.5) of single

hydrologic parameters with trout populations in all regressions,

using the total sample of 88 stream segments, suggest that popu¬

lations in a heterogeneous sampling of streams are not dominated

by a single hydrologic characteristic. In general, where sample

size was restricted to stream segments within certain limits of

hardness, temperature, or percentage of gravel bottoms, improve¬

ment in the standard error and multiple-correlation coefficients

were obtained over that where the total sample was used (Table 2).

This improvement reflects a greater homogeneity of the restricted

samples.

Temperature of water was one of the most significant hydrologic

parameters in almost all regression analyses in which it was in¬

cluded. The mean annual maximum temperature proved to be

slightly more significant than the average July temperature. Final

regression analyses were run using annual maximum temperatures

and annual maximums divided by 55 F. Using temperatures as

a ratio to the selected lower limit for trout (55 F) does not change

the accuracy in making population estimates from the regression

equations but puts the equations into a more usable form. Earlier

studies (Benson, 1953) have indicated that temperatures greater

than 68 F are above the optimum for brook trout. When the re¬

gression analyses were restricted to the 26 stream segments having

maximum temperatures of less than 68 F, temperature did not

significantly affect trout populations. However, when the regres¬

sion analyses were restricted to the 47 stream segments having

maximum temperatures of less than 72 F, temperature remained

the second most important parameter (after hardness).

Hardness was a significant hydrologic parameter in most of the

regression analyses, except for those restricted to streams having

specified ranges of hardness. When the regression was restricted

to the 62 stream segments having hardness greater than 120 mg/1

the parameter of hardness dropped out early in the regression.

This suggests that for hardness values greater than about 120 mg/1

trout populations may not improve as hardness increases. All of

the soft-water streams (streams having hardness of less than 120

mg/1) are in Michigan’s Upper Peninsula and adjacent areas of

northern Wisconsin. Most of the hard-water streams are in Michi¬

gan’s Lower Peninsula and in central and southern Wisconsin.

The generally smaller trout populations in the Upper Peninsula,

1974] Hendrickson and Knutilla — Hydrology and Trout 191

in comparison to those in the Lower Peninsula, reflect, in part,

this difference in hardness of the water.

The positive relationship of trout populations to specific con¬

ductance of water in five Pennsylvania streams was pointed out

by McFadden and Cooper (1964). Specific conductance is highly

related to hardness in most fresh-water streams.

Channel characteristics that appear to significantly affect trout

populations are bottom materials, bottom vegetation, and width-

to- depth ratio. Populations generally are greater in streams having

higher percentages of gravel bottom and bottom vegetation. The

relationship of width-to-depth ratios and trout populations gen¬

erally is positive where populations are expressed in pounds per

mile and negative where populations are expressed in pounds per

acre.

In almost all analyses percentage of fish cover showed a surpris¬

ingly poor correlation with trout populations. This may be due in

part to the subjectivity of estimates of fish cover. Also, it is pos¬

sible that fish cover generally is adequate on almost all streams

considered, and that cover significantly influenced populations only

in those short segments where great differences in cover and in

populations may be typical of the stream in general

The differences in multiple correlation coefficients with different

parameters of trout populations suggest that units of measurement

of population in either pounds per acre or pounds per mile are pre¬

ferred to pounds per acre foot or to number per acre or per mile.

(The unit of pounds per acre foot is the unit of pounds per acre

multiplied by depth of channel). Multiple-correlation coefficients

for pounds per mile are slightly higher than those for pounds per

acre, and the standard errors are slightly lower. Most individual

hydrologic parameters were also found to be more highly correlated

with pounds per mile than with pounds per acre. Possibly a

parameter of trout populations combining both area and length

might show a higher correlation with hydrologic parameters than

the units of trout populations used in these analyses.

Variability of streamflow (expressed as the ratio of the 10 and

90 per cent duration discharges, or as the ratio of the mean dis¬

charge and median minimum 7-day low flow) was also a significant

factor in several of the regression analyses. The negative relation¬

ship of variability of streamflow to trout populations was ex¬

pected, because it is generally recognized that “flashy” streams, do

not support large populations of trout. For these streams, avail¬

able cover is greatly reduced at low stages, and shallow depths

contribute to warming of the water. At high stages, erosion of

banks is likely to occur and sediment deposition may become a

problem.

192 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

A negative relationship between the ratio of the 10 and 90 per

cent duration discharges and specific conductance and hardness is

indicated by their relatively high correlation coefficients (—0.65

and —0.70, respectively). This relationship may be explained in

part by the fact that the more uniformly-flowing streams discharge

relatively large amounts of ground water whereas the more flashy

streams discharge relatively large amounts of surface runoff.

Ground water in the study area usually is harder and higher in

specific conductance than surface runoff.

Stream discharge per unit drainage area significantly affected

trout populations in several of the regression analyses. Populations

generally were greater where 90 per cent duration discharge or

7-day low flow were large. Streams having high discharge per unit

drainage area during periods of base flow generally are those that

discharge relatively large amounts of ground water. These streams

also generally have a more stable flow than those with a smaller

discharge per unit drainage area. A negative relationship between

the ratio of the 10 and 90 per cent discharge and the 7-day low

flow, in cubic feet per second per square mile (—0.67) is shown

in Table 1.

SUGGESTIONS FOR FURTHER STUDY

This is the first study known to the authors that attempts to

correlate a wide range of hydrologic parameters with trout popu¬

lations for a large number of stream segments. Although signifi¬

cant correlations were obtained, improvement in the accuracy of

hydrologic and population data would probably improve the results.

It is shown, for example, that when the regression analysis was

run using only streams for which trout population data were

based on the “mark and recapture” method a significant improve¬

ment in the results was attained. With this limitation the standard

error is 65% whereas analyses for streams having only single

survey population counts had a standard error of 95% (Table 2).

If more precise data were to be obtained, it may be desirable to

test the data for curvilinear relationships as well as straight-line

statistical analyses to see whether the correlations might be im¬

proved. It is also possible that more accurate data would show

relationships not demonstrated in the present study. For example,

significant relationships between trout cover and trout populations

may be found.

ACKNOWLEDGMENTS

This study could not have been accomplished without the help of

many fisheries specialists in Michigan and Wisconsin. Special thanks

1974] Hendrickson and Knutilla — Hydrology and Trout 193

go to Gaylord R. Alexander, David P. Borgeson, William H. Bullen,

and Howard Gowing, of the Michigan Department of Natural

Resources, and to Lloyd Andrews, Clifford Brynildson, Robert L.

Hunt, John W. Mason, and Thomas Thuemler, of the Wisconsin

Department of Natural Resources, and to J. H. Green of the U.S.

Geological Survey office, Madison, Wisconsin for his assistance in

preparing the manuscript for publication.

BIBLIOGRAPHY

ALEXANDER, G. R., and SHETTER, D. 1969. Angling statistics and post¬

season trout populations from nine lower Michigan Streams: Michigan

Dept, of Natural Resources, Inst, for Fisheries Research Preliminary

rept., 10 pp.

BALDWIN, N. S. 1957. Food consumption and growth of brook trout at

different temperatures: Trans. Amer. Fish. Soc. 86:323-328.

BENSON, N. G. 1953. The relationship among certain ecological conditions

and trout populations in the Pigeon River: Michigan Dept, of Conserv.,

Inst, for Fisheries Research Rept. No. 1369, 100 pp.

- . 1953. The importance of ground water to trout populations in the

Pigeon River, Michigan: Trans. Eighteenth N. American Wildlife Conf.,

March 9-11, 1953. Wildlife Management Inst., Wash. D. C., pp. 269-281.

- . 1954. Seasonal fluctuations in the feeding of brook trout in the

Pigeon River, Michigan: Trans. Amer. Fish. Soc., 83:8.

ELLIS, R. J., and GOWING, H. 1957. Relationship between food supply and

condition of wild brown trout, Salmo trutta Linnaeus, in a Michigan

Stream. Limnol. and Oceanog. 2:299-308.

FRY, F. E. J. 1951. Some environmental relations of the speckled trout.

Proc. N.E. Atlantic Fish. Conf., May 1951, 29 pp.

HENDRICKSON, G. E., and C. J. DOONAN. 1971. Hydrology and recreation

on the cold-water rivers of Michigan’s Southern Peninsula. U.S. Geol.

Survey open-file report, pp. 50-51.

KNUTILLA, R. L. 1967. Flow characteristics of Michigan streams. U.S.

Geol. Survey open-file report, 337 pp.

McFADDEN, J. T., and E. L. COOPER. 1964. Population dynamics of brown

trout in different environments. Physical Zool. 37:355-363.

MOYLE, J. B. 1956. Relationship between the chemistry of Minnesota surface

water and wildlife management: Jour. Wildlife Manag. 20:303-320.

NEEDHAM, P. R. 1969. Trout streams. 2 ed. Holden-Day, Inc. San Francisco,

Cal. 241 pp.

TEMPERATURE OPTIMUM OF ALGAE LIVING

IN THE OUTFALL OF A POWER PLANT

ON LAKE MONONA

Thomas D. Brock and James Hoffmann

University Wisconsin — ■

Madison

ABSTRACT

Temperature optima for photosynthesis v/ere measured for algal

populations living in the outfall of an electric power plant on Lake

Monona and were compared with the temperature optima of algae

living in a control area in the nearby Yahara River. The tempera¬

ture of the power plant outfall averaged about 8°C higher than

that of the Yahara River. Studies were carried out in both summer

and winter. In the winter, no differences in species composition

between the two study areas could be detected, Cladophora and

Ulothrix being the dominant algae. The temperature optima of

the populations from the two locations were the same, around 27 °C,

although the habitat temperatures at both locations were consider¬

ably lower. The only difference in response to temperature seen

between the two populations was that the population .at the outfall

was able to photosynthesize at higher temperature, still showing

high photosynthesis at 35 °C and detectable photosynthesis at 46 ° C,

a temperature at which the population from the Yahara River

showed no detectable photosynthesis. In the summer, the dominant

algae at the power plant outfall were Stigeoclonium and filamentous

blue-green algae (family Oscillatoriaceae) , whereas at the Yahara

River the algal population was almost exclusively Cladophora . The

temperature optima of both summer populations were the same,

31.5°C, only slightly higher than the mid-winter optima. Again, the

population from the power plant was able to photosynthesize at

higher temperature than the control population, showing quite

active photosynthesis at 42.5 °C, a temperature at which the popu¬

lation from the Yahara River was completely inactive. These results

are discussed in relation to the possible environmental impact of

power plants on Wisconsin lakes and rivers.

Electric power plants in many parts of the world use convenient

natural waters for cooling, returning the water to the environment

at temperatures higher than ambient. In the mixing zone of heated

195

196 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

water discharge, impact on aquatic organisms and ecosystems may

occur. One of the presumed consequences of man-made heating is

an increase in algal growth and a change in the algal species

present (Patrick, 1969). However, few studies have been carried

out to verify this assumption. Additionally, there have been no

studies to see if the algae developing in such warm water outfalls

have become adapted to the temperatures they are experiencing.

The availability of a convenient power plant outfall of the Madison

Gas and Electric Company on Lake Monona prompted the present

study. In a sense, this study supplements and extends a more de¬

tailed study which was done by Boylen and Brock (1973) in Yellow¬

stone National Park, measuring the effect of thermal additions

from the Yellowstone geyser basins on the benthic algae of the

Firehole River. In that study, it was shown that heating of the

river resulted in marked increases in algal growth rate and stand¬

ing crop, but that the algae were optimally adapted to the tempera¬

tures that they experienced. Further, the algae retained their

mid-summer temperature optima even during the time of the year

when the water temperatures were considerably lower due to runoff

of melting snow.

It was of interest to see whether algae living in a power plant

outfall would retain their mid-summer temperature optima

throughout the year. Consequently, the temperature optima of the

algae in the Lake Monona outfall were measured both in winter

and mid-summer, and were compared to the temperature optima

of algae from a control area, the Yahara River, which carries Lake

Mendota water into Lake Monona, but whose temperature is not

modified by power plant effluents. The results show that the algae

in both the power plant outfall and the Yahara River have tem¬

perature optima are no higher. The only clear distinction between

as with the Yellowstone algae, they retain their mid-summer optima

even during cold portions of the season. Despite the fact that the

algae in the power plant outfall experience throughout the year

temperatures higher than the algae in the control area, their tem¬

perature optima is no higher. The only clear distinction between

the two algal populations with respect to temperature is that the

algae in the power plant effluent are able to photosynthesize at

temperatures somewhat higher than the control algae. Additionally,

the species composition of the algae in the power plant effluent

shows minor differences from that in the Yahara River, although

it has not been shown that this is a direct effect of the heating

brought about by the power plant, since the two habitats may differ

in other, unmeasured ways.

1974] Brock and Hoffmann — Temperature of Algae 197

MATERIALS AND METHODS

The outfall from the Madison Gas and Electric Company plant

is at the foot of Blount Street, Madison, Wisconsin, immediately

adjacent to the Elks Club. The current created by the effluent is

very strong immediately adjacent to the outfall, but decreases con¬

siderably with distance. The actual flow pattern of the outfall is

influenced by wind and current, but there is an area immediately

north of the outfall where relatively constant current exists, and

samples were taken from this area. It was important to avoid areas

of extremely rapid flow because the scouring action of the water

prevented significant algal development. The control area on the

Yahara River was chosen so that the flow rate wras. similar to that

of the outfall area. In the winter, ice rarely forms in the Yahara

River and algal populations were always present. Temperature was

measured with a thermistor probe at the site of collection.

Algae were collected by removing rocks containing visibly green

material, placing these in plastic bags containing water from the

habitat, and returning these to the laboratory. The algae were

removed from the rocks by scraping gently with a brush, or by

picking algal filaments with forceps. Care was taken to maintain

the temperature of the algae at that of the habitat until the experi¬

ments were initiated. To avoid any changes after collection, experi¬

ments to measure the temperature optima of the algae were done

the same day the samples were collected.

The optimum temperature for photosynthesis was measured

using a radioisotope method, details, of which are described by

Boylen and Brock (1973) . The filamentous algae were cut into small

pieces with scissors and distributed in approximately equal amounts

in 5 ml glass vials containing 4 ml water from the habitat. The

vials were immersed in water baths at the desired temperatures,

allowed to equilibrate for 10 minutes, and 0.1 ml of NaH14C03

(10 /xCi/ml) injected into each vial. Light intensity was held con¬

stant at each temperature at about 250 foot-candles using fluores¬

cent lights. In addition to the experimental vials, vials incubated in

the dark were used. Uptake of radioactivity in the dark was much

lower than in the light, and the dark uptake values were subse¬

quently deducted from the experimental values.

The experiments were terminated by adding 0.5 ml of 40%

formaldehyde to each vial. An additional control in each experiment

was a vial to which formaldehyde was added at zero time, before

the isotope. Formaldehyde controls, always showed low uptake of

radioactivity. Uptake of radioactivity was linear with time over

several hours ; in most experiments incubation times of three hours

were used. Such incubation times were sufficiently long so that

198 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

significant incorporation of radioactivity occurred, yet short enough

so that secondary effects of incubation were avoided.

For processing, the contents, of each vial .were transferred to a

plastic centrifuge tube, homogenized with a Teflon homogenizer,

and 0.5 ml filtered through a membrane filter. The radioactivity

of the dried filter was then measured by liquid scintillation count¬

ing. The remaining material in each tube was centrifuged, the

pellet suspended in acetone, and homogenized again. The tubes were

allowed to stand overnight in the refrigerator, centrifuged, and the

chlorophyll content of the extracts measured in a Bausch and Lomb

Spectronic 20 colorimeter at a wavelength of 665 nm. The radio¬

activity per vial was then divided by the chlorophyll per vial, so

that each sample was normalized to the same amount of chlorophyll

In this way, variations in sample size in the original incubation

vials could be obviated. The details of these procedures are de¬

scribed by Brock and Brock (1967).

RESULTS

The measured temperatures of the power plant outfall and the

control station on the Yahara River over a one year period are

shown in Table 1. As seen, the power plant effluent was always

TABLE 1. WATER TEMPERATURE OF THE LAKE MONONA POWER

PLANT OUTFALL AND OF THE YAHARA RIVER NEAR ITS

ENTRANCE INTO LAKE MONONA

Average temperature difference between the outfall and the river over the

period of study, 8°C.

1974] Brock and Hoffmann — Temperature of Algae

199

warmer, and the average temperature differential was 8.0 °C. The

warmest temperature measured at the outfall, in mid- July, was

35.0 °C, and the coldest, in mid- January, was 10.2 °C. The winter

of 1973 was unusually mild, and ice left Lake Men dot a in early

March, so that the temperatures of the Yahara River in winter

were warmer than normal. In a normal winter when ice does not

leave Lake Mendota until April, the temperature of the Yahara

River would remain close to 0°C for three or four months.

Winter studies. No obvious differences in species composition

between the two study areas could be determined which could be

related to temperature differences. At both stations, Cladophora

and Ulothrix were dominant algae, associated with small numbers

of pennate diatoms. At the outfall, Stigeoclonium was found in

small numbers, and in the Yahara River Rhizoclonium was seen.

The temperature optima of algal samples collected on 1 February

1973 are shown in Fig. 1. As seen, the optimum for both algae

was about the same, around 27 °C, despite the fact that the habitats

of both populations must have been considerably cooler for at least

two or three months. The only difference in response to temperature

FIGURE 1. Effect of temperature on photosyn¬

thesis of algal populations collected at the power

plant outfall (temperature 11.2° C) and the Yahara

River (temperature 3.5° C) on 1 February 1973.

200 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

between the two populations was that the population at the outfall

was able to photosynthesize at higher temperature, still showing

high photosynthesis at 35 °C and detectable photosynthesis at 46 °C,

a temperature at which the population from the Yahara River

showed no detectable photosynthesis. Another experiment was done

with material collected on 13 February 1973. Again, both algal

populations had the same temperature optimum, but the optimum

was now a little lower, a broad peak being seen from 20-27 °C.

As before, the algal population from the outfall was able to photo¬

synthesize at a higher temperature than the population from the

Yahara River. The species composition on 13 February was similar

to that on 1 February.

Summer studies. Samples were collected on 14 August 1973, at

which time both habitats had been warm for at least 1.5 months.

At the power plant outfall, the dominant algae were Stigeoclonium

and filamentous blue-green algae of the family Oscillatoriaceae.

At the Yahara River, the population was almost exclusively

Cladoyhora. It is of interest that blue-green algae were never seen

in the winter, or at the Yahara River even in summer, and at no

time could blue-green algae have been considered dominant.

The temperature optima of the algal populations collected in

mid-summer are given in Fig. 2. As seen, the optima of both popu¬

lations are the same, 31.5°C, only slightly higher than the mid¬

winter optima illustrated in Fig. 1. Again, the population from

the power plant is able to photosynthesize at higher temperature

than the control population, and shows quite active photosynthesis

at 42.5 °C, a temperature at which the population from the Yahara

River is completely inactive. Another interesting comparison be¬

tween Fig. 1 and Fig. 2 is that the actual photosynthetic uptake

of C02 per unit chlorophyll is much higher in the summer than in

the winter algal populations. This presumably reflects the fact that

the winter populations are- in a physiologically inactive state, either

because of the low light intensity or the low temperature at which

they are growing. Despite these marked quantitative differences

between the populations from the two seasons, the temperature

optima are virtually identical.

DISCUSSION

The results of the present studies agree closely with the work

of Boylen and Brock (1973) in the Firehole River, Yellowstone

National Park, in showing that the temperature optima of algae

do not vary throughout the year even though the habitat tem¬

peratures do vary widely. The algae in the present study showed

virtually the same temperature optimum in mid-winter as in mid-

1974] Brock and Hoffmann — Temperature of Algae

201

FIGURE 2. Effect of temperature on photosyn¬

thesis of algal populations collected at the power

plant outfall (temperature 30.0° C) and the Yahara

River (temperature 24.0° C) on 14 August 1973.

summer, despite the fact that the habitat temperature was 15-25 °C

higher in summer than in winter. This suggests that, at least in

freshwater environments, algae adapted optimally to low tempera¬

tures do not occur. There is evidence in some marine environments

of winter algal populations adapted to low temperatures, being

replaced in summer by different algae adapted to higher tempera¬

tures (Feldmann, 1951). Conceivably, in freshwaters, algae op¬

timally adapted to low temperatures would have no low temperature

refuge during the warm summer months, would be killed off, and

hence would not be available to colonize during the subsequent

cold period. In marine environments, deeper cold habitats may be

available as low-temperature summer refuges.

202 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

A second conclusion from this study is that a population specif¬

ically adapted to the warmer waters of the power plant effluent

has not developed. Although it is true that the temperature differ¬

ential between the two habitats averaged only about 8.0 °C, Boylen

and Brock (1973) were able to observe adaptation of algal popu¬

lations to temperature differences even less than these. However,

even though the algae at the outfall did not have a temperature

optimum which was higher than the control, they did show some

ability to adapt to the warmer waters, since it was always observed

that the algae at the power plant outfall were able to photosynthe-

size at higher temperatures than the algae at the control location.

It was also noted in every case that photosynthesis per unit chloro¬

phyll was higher in the algae from the power plant outfall than in

the control algae, presumably because the former were able to

photosynthesize more efficiently. Conceivably, this more efficient

photosynthesis reflects an algal population which is growing under

more favorable conditions. This may be a temperature effect, but

it could also be a current effect, even though some care was taken

to select two habitats which had similar current patterns. Some

of the difference in the temperature responses of the two algal

populations could be due to differences in species composition, and

thus only indirectly be due to differences in temperature.

These results, together with those of Boylen and Brock (1973),

permit some prediction of the effect of thermal pollution on the

development of algal populations in freshwaters. First, since the

algae seem to be preadapted to warm temperatures even in the

winter, if a freshwater habitat is heated as a result of a new power

plant effluent, algal growth should increase in rate, probably result¬

ing in increase in standing crop. Although the present study did

not concern algal growth rates, the previous study (Boylen and

Brock, 1973) showed clearly that algal growth rate and standing

crop were higher in the heated than in the unheated water. Since

in both habitats, the algae are preadapted to the warmest tem¬

peratures found in the summer, it seems reasonable to predict

that increasing the temperature of the habitat artificially should

result in increased algal growth.

Second, at the temperatures observed in the present study, blue-

green algae never became dominant. This was true even though

temperatures at the power plant outfall reached 35.0 °C in mid¬

summer. As has been shown in previous work (Brock, 1967, Tansey

and Brock, 1972), eucaryotic algae are able to grow at tempera¬

tures up to 55-60 °C, and only at temperatures above this would

exclusively blue-green algal populations necessarily develop. How¬

ever, in most hot spring thermal gradients blue-green algae are the

1974] Brock and Hoffmann — Temperature of Algae 203

dominant or sole algal components at temperatures as low as 40 °C.

Only when temperatures have dropped below 40 °C are eucaryotic

algae seen to form extensive or dominant populations. Since in the

habitats under study in the present paper, the highest temperature

seen was 35 °C, it is not surprising that eucaryotic algae dominated

the populations throughout. Interestingly, some blue-green algae

were seen in the power plant effluent in mid-summer, at which time

the temperature was the highest, suggesting that the habitat tem¬

perature may have been becoming more favorable for the blue-

green algae. A reasonable prediction is that if the power plant

effluent should increase in temperature above 40 °C, the scale may

be further tipped in the direction of the blue-green algae and these

organisms may become dominant. Such increases in temperature

could conceivably occur as a result of increased power plant

loading.

From the point of view of thermal pollution, the state of Wis¬

consin is fortunate in that its natural waters are generally low

in temperature, so that increased temperature due to power plant

activity does not push the water temperature above the critical

40 °C point. In other parts of the country, where natural water

temperatures are higher in the summer, an increase of 8-10 °C

due to power plant activity might have considerably greater effect

in promoting blue-green algal development. At least at the moment,

there is no reason to believe that the power plant on Lake Monona

is having any troublesome effect on algal populations which develop

in its outfall.

ACKNOWLEDGEMENT

This work was supported by a research contract from the

Atomic Energy Commission (COO-2161-18) .

REFERENCES

BOYLEN, C. W. and T. D. BROCK. 1973. Effects of thermal additions from

the Yellowstone geyser basins on the benthic algae of the Firehole River.

Ecology 54:1282-1291.

BROCK, T. D. 1967. Life at high temperatures. Science 158:1012-1019.

BROCK, T. D. and M. L. BROCK. 1967. The measurement of chlorophyll,

primary productivity, photophosphorylation, and macromolecules in

benthic algal mats. Limnol. Oceanogr. 12:600-605.

FELDMANN, J. 1951. Ecology of marine algae. In Smith, G. M. (ed.)

Manual of Pliy colog y. Ronald Press, New York.

PATRICK, R. 1969. Some effects of temperature on freshwater algae,

p. 161-185. In Krenkel, P. A. and F. L. Parker (eds.) Biological aspects

of thermal pollution. Vanderbilt Univ. Press.

TANSEY, M. R. and T. D. BROCK. 1972. The upper temperature limit for

eucaryotic organisms. Proc. Nat. Acad. Sci. 69:2426-2428.

■? .

A CORRECTION IN SET THEORY

William Dilworth

Beloit , Wisconsin

PREFACE

The logic and assumptions which today comprise the ‘foundations

of mathematics’ often lead to paradox — the name given to a logical

but patently absurd conclusion. One may regard paradox with awe,

or one may look for underlying errors.

Orthodox mathematical belief today holds that we may take a

solid sphere of any fixed size, divide it into a few pieces, and then

reassamble those pieces into two solid spheres, each of the same

fixed size as the first. This theorem, due to S. Banach and A. Tarski

in 1924, has been acclaimed as a triumph of modern methods.

Logically similar notions weave through the “new mathematics”

taught everywhere today.

In the 1924 paper the authors depend explicitly on the work of

F. Hausdorff, who was in turn building on Georg Cantor’s theory

of sets. So either the sphere surgery can be done, and one equals

two, or we had better have another look at Cantor’s sets. Here I

present an analysis, made possible by modern semantics, of a cen¬

tral fallacy in Cantor’s theory. The reader will follow my argument

without difficulty if he understands that certain endless sequences

of fractions have finite limits; e.g., one-half plus one-quarter plus

one-eighth, and so on, never totals more than one, no matter how

far extended.

This paper expands on the following topics :

Numbers Generally . The Archimedean test for equality or in¬

equality of two quantities enables us to determine, no matter what

anyone may claim, whether some given form actually defines a

numerical value, or not.

Scalars. The adjective “real” has traditionally been applied to an

important technical class of numbers, confusing students and

promulgating philosophical haggles. The exact synonym “scalar”

number is adopted for its structural implications.

Nndless Convergent Summations. General examples of the “one-

half plus one-quarter” type of sequence are introduced, along with

the compact modern notation in which we can exactly express them.

We find that all of the scalar numbers can be so expressed, under

the summation symbol. On the other hand, decimals, while prac¬

tical and convenient, happen to be inadequate to this task.

205

206 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Given Numbers. To have a “given number”, as it is casually put

in the literature, some Jones must give it to some Smith. This

human condition limits the possibilities in interesting ways.

Permutations. This is the technical word for what we commonly

call arrangements, such as the way in which a set of books might

be arrayed along a short shelf, ordered say by authors, or by titles,

etc. Simple laws define how much total variety of arrangement is

possible given, say, twenty books and a shelf which will hold five.

i(New Permutations” . With the laws just mentioned, we are in a

position to refute immediately any claim that someone could pro¬

duce a new, unexpected, and unpredictable permutation within an

already defined system. From this framework of secure knowledge,

we are prepared to examine critically Georg Cantor’s famous

“proof” that the scalar number system is irreparably disordered

and disarrayed.

Cantor’s Diagonal Argument. This is the cornerstone of Cantor’

century of influence on mathematics. He claims always to generate

“new numbers”, not already in any conceivable list. From our

permutational point of view, his manipulations are not impene¬

trable. We and Cantor agree on the elements (books and shelf)

of a system. We then display to Cantor all permutations of the

books taken, say, three at a time (a permutation of length 3). In

order to find a “new” permutation, Cantor insists on using a greater

length — namely, four. Thereupon we display to him all permuta¬

tions of length four; he retreats to the claim that he should be

allowed five. Before modern semantic methods made the present

analysis possible, Cantor’s ability to produce “new” arrangements,

while unseen he increased the length of the arrays to which he had

access, appeared almost magical. His claims were accepted— never

explained.

Decimal Expansions. The illusion that Cantor has each time

come up with a “new number” involves a misreading of the decimal

expressions he uses. By a straightforward inspection of decimals,

as well as by a general professional consensus, many scalar num¬

bers cannot be expressed exactly in decimal form. He who fails,

through a lack of rigor, to remember the limitations of the decimal

system, may imagine that he sees in Cantor’s truncated decimal

forms the “objective real numbers”; he slides into Cantor’s subtle

mistake.

The literature contains many dire predictions that parts of

higher mathematics “would collapse” if any defect in Cantor’s

theory were ever found. My personal communications with pro¬

fessionals show that many of them share this fear. What are the

facts? Nothing solid in mathematics is going to collapse. Certain

1974]

Dihvorth — A Correction In Set Theory

207

passages in special arguments may have to be modified. New, pos¬

sibly fruitful, insights may result.

INTRODUCTION

Georg Cantor, a German mathematician of the late 19th century,

fathered the ‘theory of transfinite sets’. His remarkable ideas were

scathingly attacked by his contemporary, the illustrious Kronecker.

Poincare, considered the leading mathematician of the time, was at

first intrigued but later became bitterly disillusioned and declared

the thing to be a mathematical disease. But Cantor finally won

authoritative acclaim from Hilbert and Bertrand Russell, and car¬

ried the day. With rare exceptions, critics have been silent to this

day.

Direct results of Cantor’s theory now appear even in elementary

textbooks, and a long deductive chain runs from it to the ‘Banach-

Tarski’ spheres. In a roundabout but hopefully heuristic way, we

want to have a good look at the cornerstone of Cantor’s system.

We are concerned here only with the set-theoretic issues; the

quality and importance of some other work done by Cantor is, I

believe, beyond dispute.

Numbers generally. Archimedes first gave formal expression to

our common intuition as to what constitutes a number or, rather,

how we can tell in some case whether we have a number or not.

It is often called Archimedes’ principle ; it is in the strictest modern

sense an operational test. Briefly, if C and D are numbers, then

just one of the following three cases holds:

a) C is greater than D (C > D),

or b) C is equal to D (C = D),

or c) is less than D (C < D).

If, for whatever reason in a given case, we find that we cannot

determine which of these three condition holds, then it follows that

C, or D, or both, does not define a number.

It frequently happens, both in practice and in theoretical work,

that we cannot make the Archimedean test because, while we have

some information about a number, we do not have enough to define

it exactly. That is, we may know a range within which the number

lies, but nothing more. If C is “4 and a little more”, the test may

fail. Likewise if C is “4 + a remainder”. And particularly, if C is

given in an incomplete form, such as 4.32* ••, then it is in fact

impossible to determine whether C is greater than, equal to, or

less than 173/40, for example. The meaning of the familiar “ • • • ”

can become rather subtle, when we get into endless (infinite) se¬

ries. A small difference between C and D can make a lot of differ-

208 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

ence in results. If the expression “C — D” appears in a denominator,

then our answer may be negative, or positive, or meaningless,

depending upon that “little difference”.

The exact definition of a number need not be simply in digits.

Not only are operator symbols such as + and / regularly used in

defining numbers but purely verbal sentences or longer contexts

may be used. Anything we can put in mathematical symbols we

can also eventually manage to say. “A circle’s radius over its cir¬

cumference” defines in simple English exactly one number; by

routine operations we can actually compare “half the reciprocal

of pi” with any other given value, for equality or specific inequality,

as required.

The full scalar number system. All the numbers referred to here¬

under will be scalar numbers. One scalar number, or scalar, defines

exactly one point on a lineal scale which is endless both left and

right of a zero point and on which a unit length is assigned. A

scalar defines one point. Conversely, any fixed point on the line,

even if it has been defined only by geometric operations or other¬

wise, is associated with one unique scalar. Historically, in cases

where a geometric or algebraic definition came first, the develop¬

ment of the required numerical scalar became an urgent need. Such

needs have always been met and today the scalar number system is

considered complete. All modern theories in calculus and analysis

assume and depend upon this fact.

The foregoing statements imply that every scalar number is fully

defined, single-valued, fixed and constant. I will rigorously adhere

to that connotation. When a domain of definitions, or a range of

values, is meant, those terms, along with ‘function’ and ‘variable’

will be used. One may argue with considerable justification that

the multi-level structure of both mathematical and nonmathematical

language make some ambiguity inevitable. I will say that it is diffi¬

cult to be consistently clear but I will try anyhow.

A scalar number is additive in structure. Any two scalars add

to form a single scalar. From the theory of algebraic fields : Addi¬

tion is postulated, multiplication is repeated addition, subtraction

is inverse addition and division is inverse multiplication. As would

be expected, the latest forms of scalars to evolve are the most gen¬

eral and have an elaborate composition. However, an organized

hierarchy of transformations permits us to express even the earliest

and simplest scalars precisely in the latest general forms. For the

exact representation of scalars generally, two specific numerical

frames are available : Continued fractions, and endless series. It is

of interest that we can without loss of meaning interchange the

adjectives, to say endless fractions and continued series. Also, many

1974]

Dilworth — A Correction In Set Theory

209

people would prefer “infinite” to “endless” — one may consult the

dictionary and his own taste.

Endless convergent summations. Although the continued frac¬

tions are of profound number-theory interest, bounded summations

of endless series of diminishing fractional terms are more familiar

and fully adequate for our purpose. In the powerful notation of

modern mathematics, the notion of an endless convergent summa¬

tion condenses to :

n a.

(1) S, a scalar, = lim 2 -t-1.

i bi

The index i ranges over i = 1, 2, 8, • • *, to and including n. And

n has no upper bound. This is important. The symbol n and the

complex symbol n — > oo are used interchangeably in the literature,

which interchangeability I fully accept, in the above context. There

is a context which does not permit n and n — > oo to mean the

same thing, and that is when the series ai/bi is such that the

partial sums do not converge. In that case the “limit of the summa¬

tion” has no meaning, and the summation itself has a numerical

value only when n is fixed. In the case we are discussing, (1), the

series is defined as convergent and the summation has an upper

bound and a limit even when n exceeds any bound, and n then ex¬

ceeds a still higher bound, endlessly.

After such a paragraph, we need examples. Every so-called

‘transcendental’ scalar can be genetically expressed in the form

shown in (1), although the detailed structure of the fraction ai/bj

may be quite complicated. For a relatively uncomplicated example,

we have

n 1

(2) e (epsilon, base of nat. logarithms) = lim 2,-p - y~j.

The summation in (2) can be proven never to exceed 14/5, when

n increases without limit.

On the other hand, the innocent-looking fraction 1/i, when part

of the summation expression

n ^ co

(3) 5 i — , becomes part of a summation that never stops

i 1

growing ; it exceeds every bound ; hence it would be incorrect

to precede it with the term “lim” (limit), or to equate it with

any numerical or algebraical symbol whatsoever.

When it is understood that the series converges, we can call

OO

2 ti a closed or hound form, because when ti is given we have,

210 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

in a finite number of symbols, all the information necessary to

expand the series endlessly. But, of course, to get the closed form,

we had to use the finite abbreviation “i” to represent the basic,

perfectly regular, endless system of integers 1, 2, • • * . If i did

not represent a perfectly regular, pre-known system, it would then

00

be impossible to expand 5 U to produce a single, definite

i

numerical scalar value.

Given Numbers. Mathematics textbooks often use the phrase

“given the number x . Let us expand a little on this phrase, the

meaning of which is usually taken to be self-evident. To say that

someone, say Jones, has given a number to someone else, say Smith,

means first of all that they both understand and accept a language

and a symbol system. It also means that Jones knows techniques

by which he can expand that number to any required degree of

accuracy, say to n places, and that Smith, independently, can also

expand that number to as many places, and that the two expansions

will be identical, term by term. It may be easier, in this case or

that, to make decimal expansions, but it can always be done also

in common rational fractions. Note that Jones can never give Smith

the endless expansion itself — only some closed form. Jones and

Smith are limited by the conditions of human communication.

Surprisingly, many of the professional mathematicians whom I

have consulted over the years resist and refuse to acknowledge

the foregoing operational facts. I have been accused of “philosophiz¬

ing” about the matter. They have insisted that the endless expan¬

sion itself can somehow be completed, communicated, and that it

alone constitutes the scalar number. Moreover, the requirement

for regularity of development of the expansion is not generally

recognized. A professor of mathematics at Michigan, with whom

I had already discussed these points in person and established some

rapport, sent me a well-drafted letter declaring that the successive

digits of the endless decimal expansion of a number could be de¬

cided by a function on repeated throws of a pair of dice. He was

at first non-plussed when I pointed out that his procedure would

predictably produce different sequences each time it was performed

and so that it was, if anything mathematical, a variable rather than

a single, definite number. Soon, however, that reasonable man con¬

ceded, “By golly, you’re right!” Had he written a textbook before

that exchange, however, his “formula for producing a scalar num¬

ber” might very well have gone on to classes and, perhaps, to

posterity.

Permutation systems. A permutation is an arrangement of some

or all elements of a prescribed set. Repetitions of the same element

1974]

Dilworth — A Correction In Set Theory

211

may or may not be permitted. The word ‘sphere’ (s,p,h,e,r,e) is a

six-place permutation of letters from the 26 of the alphabet. One

character, e, is repeated. The total possible variety of such six-place

permutations is just 26°, or exactly 308,915,776, a large but clearly

finite number. Most of the arrangements do not spell words, of

course. The alphabet itself has a fixed, rather small number of

characters. Even the total variety of characters and signs available

for printing is limited. Obviously, an ‘infinite alphabet’ would

before long contain unrecognizable symbols and has in fact no

serious meaning.

Every expression defining a scalar number is a permutation

selected from a fixed ‘font’ of symbols, including digits and opera¬

tors such as 2, 7, +, /, etc. Each character in the set is a single,

uncompounded, discrete character. The word ‘permutation’ implies,

of course, that the characters gain added significance from their

position within the arrangement.

Consider now the two digits 0 and 1 of the binary system. It is

a universally acknowledged fact that these two characters, plus the

point can define every numerical value that is possible in the

standard decimal system. This similarity extends in particular to

the issue of endless expansions.

We regard the characters 0 and 1 from a permutational point of

view, neglecting for the moment their standing as scalar numbers

and digits. However, for continuity in the later argument, each

permutation is preceded by a point, so we write .0 1, .0 1 0, .1 1,

etc. Since the point appears in the same position in every case, it

has no bearing on the variety of possible permutations. Let us now

see what is the mathematical meaning of “all possible permuta¬

tions of 0 and 1, to n places, when n = 1, 2, 3, • • •, and so on with¬

out end.” We could start with n = 15, say, and then pick up the

smaller values of n in another order, but this would make sys¬

tematic examination of all the possibilities more awkward. So we

start more naturally with n = 1.

Let “pm.” stand for “permutation”. Then a little experimenta¬

tion quickly reveals the following structure :

212 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

There is a simple law describing the quantitative growth of per-

mutational variety in the above system. The number of elemental

characters is 2. Taking j as index for the number of places, the

number of pms. in each row is exactly 2j. It cannot be more. More¬

over, there is a simple law for the total number of pms. in all rows

up to and including the j-th, and that is just 2j+1 — 1. This last sum

shows that all the pms. in the system are technically countable , i.e.,

the pms. may be paired off with the integers i = 1, 2, 3, • • •. We

note here the elementary fact that the integers cannot be summed.

OO 00

That is, 5 i is a divergent series and has no total. Likewise, 3 23 is

i _ j

i

divergent, and indeed at a faster rate than 2 i. Nevertheless, the

OO

technical property of countability is preserved. We have sketched

the structure of any two-character permutational system. With

suitable substitutions in the algebraic variables, the same state¬

ments hold for the base-ten, decimal system.

“New” permutations. It 'happens', i.e., it is a consequence of our

standard, highly abbreviated system for writing integers, that

every permutation of the set of base-ten digits, without a point,

defines an integer. Some pms. are redundant, eg., 03 = 3, but

otherwise there are no "spelling rules" needed.

Referring again to the base-two system, suppose now someone

comes along and claims that he has a way of constructing a new

pm., one not included in the countable system we have developed

above. To common sense, this appears to be impossible. If his new

entry has one place, how can it be other than 0 or 1 ? Obviously, it

cannot. If his new entry has two places, how can it be other than

0 0, 0 1, 1 0, or 11? Obviously, it cannot. Common sense is right.

If his new entry has n places, how can it be other than one of the

2 11 pms. which develop in our countable system? It cannot . Only

one 'possibility' remains. Does this orderly, fully developed permu¬

tational system suddenly fail if n has no upper bound? Why

should it?

Still, someone has made the claim, not only that he can produce

a new pm. not in any list we can devise, but that he has also

thereby proved that pm. systems in general are unorderable, that

their members cannot be counted off one-for-one with the integers,

and that he thus demonstrates that he knows of an infinity beyond

infinity. If we try to count his pms., he says, the integers will

become exhausted. "Yes sir," the head of the mathematics depart¬

ment of a Univ. of Illinois section said matter-of-factly to my face,

"The integers will become exhausted." Relieve it or not, Georg

Cantor made these remarkable claims stick with the world's mathe-

1974]

Dilworth — A Correction In Set Theory

213

maticians of his time, and they stick unto this day. The effects of

the Cantorian grip on the professional mind have to be experienced

to be believed.

Cantor's diagonal argument . Here is how he proceeds. Cantor

invites us to make a list of permutations, and we are to take the

position that our list is so arranged as to include all possible pms.

Suppose we start:

1st pm. .0 0 0

2nd pm. .0 11

3rd pm. .0 10

• • • « • •

Now, Cantor reminds us, one pm. is defined as distinct from an¬

other if it differs from that other in any one specified place . We

agree to this. “So,” he says, “because the first digit of your first

pm. is 0, the first digit of my new pm. (usually called z) will be

not 0 but 1. Therefore, z is not the same as your first pm.” “Now,”

says Cantor, “the 2nd digit of your 2nd pm. is 1, so I make the

2nd digit of z to be 0. And so z is not your 2nd pm. Since the 3rd

digit of your 3rd pm. is 0, I make the 3rd digit of z as 1. There¬

fore z is not your 3rd pm. And so on. I win.” Historically and up

to this date, he has icon. The horrendous “alephs” of his endless

infinities thunder through the evening skies of academe “with

hooves of steel”, as the songwriter put it.

You will doubt that anyone could be deceived by the foregoing

brief and transparent manipulations. Nor were they. Cantor’s his¬

toric presentation was in subtly different terms. No discussion of

permutational structures preceded his demonstration. He insists

that he is dealing directly with the objects of the “real” (scalar)

number system, and that therefore their putative un-orderability

and un-countability are a matter of the gravest concern for all of

mathematics. He declared that his conclusions were forced upon him

by compelling logic, against his own will. He made it stick and, if

one believes him, one will also come to believe in the Banach-Tarski

spheres.

Let us go back to where Cantor says, “And so on.” Because, con¬

trary to Cantor’s colleagues and even his critics, we have taken

pains to analyze and understand in advance the orderly and end¬

less structure of our permutational system, and we do not have to

confuse it with “real numbers”, we are in a position to object to

his calm, “And so on.” “You, Cantor, have not seen our 4th pm.

yet. It has only three digits; there is not a “fourth” one to work

on; and your “new” pm. z = .1 0 1 certainly does appear in our

list.”

Cantor wants to wave aside our statements. “You don’t under¬

stand,” he says. “All such arrays of digits are endless. You have

214 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

only shown me three places. Every one knows that all real num¬

bers lead to endless expansions . . We ask, “Can’t we just talk

about the pms. first? All numbers are permutations of digits and

other symbols. Don’t these laws apply?” And at about this point

your Cantor (ian) will declare, “Right now I don’t care anything

about permutations. What I want to do is prove to you the correct¬

ness of the diagonal argument, and you are supposed to have writ¬

ten the real numbers all down here and only the endless decimal

form will do and closed forms and finite rationals and things like

that simply do not matter. They do not matter because they will

not help my argument which can only proceed, when you have

already written down all the endless decimals and so on.” I do not

exaggerate.

Now the refusal of the Cantorians to allow us to develop an

orderly listing of all scalars by taking one-symbol pms. first, then

two-symbol pms., etc., using whatever conventional digital and

operator characters are required, is truly ironic. That is because

Cantor himself had earlier made mathematical history by proving

that all of the first three orders of scalars, namely integers, ration¬

als, and radicals, could indeed be ordered, and counted, and all had

the same “type of infinity”. It is worth noting that expositors today

describe this feat of Cantor’s without ever mentioning that he did

it by systematically putting all of the shortest forms first ; then the

next longer, and so on. There is, as you may well conclude, no other

way of ordering an endless system. Whether subconsciously or not,

these writers manage to avoid displaying short vs long forms, by

sticking to extremely simple cases in ten-base numerals, or by

resorting to algebraic symbols which effectively conceal the lengths

of the corresponding numerals. At least a part of their interest is

clear : They have written many books giving many solemn proofs,

confirmations and consequences of Cantor’s argument that, alone

among all the scalars, the general form employing the summation

operator 2 cannot be ordered and counted. Their desire to believe

is strong, if not profound. The shibboleth, “The n-th digit of z is

different from the n-th digit of the n-th number in the list” acts

to paralyze the higher centers. Since the simple scalar numeral

“3” does not have a “n-th” digit the Cantorians “expand” it with

an infinity of zeroes after the point, for no other reason or purpose

than to sustain the illusions of the diagonal argument.

Decimal expansions. Let us look at the definition of an endless

decimal as “a definite scalar number”. First of all, the literature

is in fact replete with descriptions of the properties and limita¬

tions of decimal expansions which state that some scalar numbers

cannot be exactly defined by decimals. Yet, in other references, the

endless decimals are given genetic status as scalar numbers. The

1974]

Dilworth — A Correction In Set Theory

215

endless decimals may be regarded as the actual objects of the real

number system, one authority puts it. Now these declarations

involve flat contradictions, of course. We could say that mathe¬

maticians are only human, but that would explain nothing. There

is a reason, indeed a doctrinal reason, for this anomalous situation.

I want to say here that our modern system of decimal notation

is a truly marvelous mathematical structure, thousands of years

in development and brought to its present state only during the

Renaissance. (Shortly it will be refined a little more, when the

clumsy redundancy of “x Kb'” in so-called scientific notation will

be replaced by a suitably positioned single e for the power of ten.)

However, decimal fractions, wherein the issue of “expansions’'

arises, are only one of several possible forms for fractions, and of

course everyone who is taught arithmetic is taught common frac¬

tions, as well. Now it is an elementary but noteworthy fact that, if

we actually restrict ourselves to the ten digits and the point, many

common fractions can never be exactly shown or ‘given’ in decimal

form. Consider a/b = .2592 • • This cannot be unambiguously

solved for two integers a and b. The best we could say is that a/b

is equal to or greater than 162/625 and less than 2593/10,000. Of

course this might be good enough for many practical purposes but

is not significant in a theory of rational numbers. Recall that the

Cantorian diagonal argument implies that a difference in the n-th

digit is meaningful, no matter whether that might be the 10th digit,

or the billionth, or when n increases endlessly.

The integers a and b in the above example can be exactly defined

by the addition to the decimal system of a conventional but little

used symbol called the vinculum (or some equivalent). This is a

line over a set of decimal digits meaning that the permutation of

digits repeats endlessly in the expansion. In other words, the

vinculum is a symbol abbreviating the sentence preceding this one.

In many cases, writers modify and circumscribe a numerical ex¬

pression by a couple of paragraphs of special, one-purpose exposi¬

tory text and then appear to believe that all that complex of

meaning actually resides in the numerical itself. It reminds one of

the comedians’ convention where so many jokes were going about

that they were referred to as No. 29, No. 172, etc. When one fellow

heard “No. 17” he fell into a fit of laughter, since he had never

heard that one before! At any rate, the vinculum is an addition to

the set of ten digits and the point; it permits us to drop one digit

from our example and show a/b = .259, wherefrom a/b is exactly

7/27, and nothing else. Without the vinculum, or some symbolic or

linguistic substitution for it, the “endless decimal expansion” can¬

not define 7/27 or others of that type.

216 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

When it comes to so-called irrational numbers, like y7, the

deficiency of the decimal expansion is far more striking. There

is no equivalent of the vinculum for irrationals. Nor can any be

invented. For rationals, the vinculum delineates a pattern which

survives the transcription from the common fraction into the

decimal fraction form, and from this pattern the closed form a/b

can be exactly recovered. When \/7 is expanded in decimals, no

pattern survives, and hence the original expression cannot be un¬

ambiguously recovered. This is all perfectly conventional and well-

known, you can check it with anyone, and indeed if you were so

inclined could prove it to yourself with some effort and practice.

The loss of pattern in an irrational is the cost of expanding it

decimally; in practice of course it is often advantageous because

of the ease with which decimal approximations can be compared

with one another, combined with each other, etc.

Perhaps it is not obvious what pattern there actually is in an

irrational like V7. However, there are many ways to expand y7

in forms in which the pattern becomes clearly visible and is never

lost. Continued fractions, mentioned above, are one. Another is by

the expansion of (6 + 1) 1/2 by the binomial theorem. There are

indefinitely many different ways of expanding y7 by the two

systems mentioned, and there are other systems, too.

So the “endless decimal expansions” cannot give us even a simple

radical exactly. But the other systems, including the endless con¬

vergent summations, always provide closed expressions, not only

for radicals but for a still higher form of scalar number usually

called “transcendental”. These include trigonometric functions, for

example, so they are eminently ‘practical’ sorts of things.

I can assure you that every mathematician will concede the tech¬

nical accuracy of the foregoing. On some points he may have to

do some ‘figuring’, or long-recalling, but he will concede. As I have

pointed out, for mathematicians no less than for the rest of us does

a contradiction between tw'o propositions preclude their both being

carried in the same head.

So why have the “endless decimal expansions” (or binary expan¬

sions, for that matter) been raised to such a status as to be equated

with scalar numbers themselves? The answer, I believe, is now

clear. Because the open admission that closed forms for scalar

numbers, which forms can obviously be ordered and counted, are

available in other expansion systems but not in the decimal or

binary type, would lead rather quickly to the exposure of the

number-theory fallacies in the Cantorian diagonal argument and

the deductions made from it. Remember the spheres.

THE FISCHER COLLECTION

James La Malfa

University Wisconsin Center — •

Marinette County

The delivery, in November 1971, of the last of the Fischer Collec¬

tion to the Geology department of the Milwaukee Public Museum

represents the finale of a most interesting story. The Fischer collec¬

tion was donated by me and consists of 296 samples of pigment, dye,

minerals, oils and binder used in paint from prehistoric time to the

20th century. The collection also includes a complete Ostwald color

wheel in pigment, a post card from Wilhelm Ostwald to Dr. Martin

Fischer, Fischer's scrapbook and his notes from lectures on paints

delivered all over the United States.

The Fischer Collection has been valued at $4000 by Professor

Lawrence Rathsack, art instructor at the University of Wisconsin —

Milwaukee. Professor Rathsack, himself an expert on the subject

of paint technology, Mr. Joseph Emmuletti and Mr. George Gaens-

len of the Geology department, Milwaukee Public Museum, have

been most helpful in finding the Fischer Collection a permanent

home.

The prime motivation for my writing this article and placing

the collection where it is now housed, is to make available to schol¬

ars and artists a rich source of information on the history of paint.

Mr. Emmuletti, who presides over the collection, has assured me

that the items collected by Dr. Fischer, will be made available to

any interested party for study, examination and photography.

In August 1968, Mr. Louis Voight, Seminary Librarian at Wit¬

tenberg University, Springfield, Ohio, called me and asked if I

would like some paint. I was an art instructor at the time in Wit¬

tenberg’s art department and Louis was a friend and neighbor.

Although my specialization is drawing and sculpture, I replied in

the affirmative, more from curiosity than any other motive. I met

Louis in the Thomas Library basement and he showed me some

dirty cardboard boxes which, he said, the library was planning to

discard because they lacked sufficient storage space.

I opened one of the boxes of “paint” and observed a carefully

labelled collection of jars, capped and sealed with sealing wax. This

was, I reflected, most unusual paint ! Thus my curiosity was aroused

by my first glance at the pigment collection of Dr. Martin Fischer.

I was determined to uncover the reason for the collection and facts

217

218 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

about the man who assembled it. Every box marked “Fischer Col¬

lection” was moved to my studio where I proceeded carefully to

open and catalog each box and each item. The number of boxes

totaled fourteen, most containing notes in pencil by Dr. Fischer,

describing the contents of each.

Each group was assigned a letter, A to N, and I carefully copied

down the contents of every bottle. Copies of the resulting catalog

are in Mr. Emmuletti and Professor Rathsack’s hies. A perusal of

the catalog indicates the order of a careful, scientific mind. Box A,

for example, contains all of the whites and blacks ever used as

pigment. It is interesting to note that of twenty-two specimens of

white pigment, many, listed under proprietary names, are chemi¬

cally identical to other whites with different proprietary names.

For example, Chremmitz, Silver, Flake and Lead white are all lead

carbonate.

Of the ten samples of blacks, asphalt and asphaltum are the same.

Cork black and lamp black are identical, basically sooty carbon.

The two samples of blue-black are graphite. Although not true

blacks, Fischer included sepia and mummy as examples of warm

blacks which were utilized by 19th century artists. Sepia is squid

ink and mummy is literally that; ground up Egyptian mummies!

The last worth mentioning is bister (bitumin lake), an analine dye

co-precipitated on a clay base. Genuine bister is asphalt.

The fact of the matter is that each sample in this group and every

other sample in Dr. Fischer’s collection are there for a reason.

Either the pigment is historically significant because it was a bad

pigment or because it was a good pigment.

Box A also contains samples of mineral pigments used by Me¬

dieval and Renaissance artists. Box B contains lakes, madders and

various oils and varnishes used as binders. Box C contains stearates

and phenols, Box D some reds and browns, Box E a series of ochres

and yellows and Box F miscellaneous pigments.

Box G included a note by Dr. Fischer : “Color change from yellow

to red due to change in size of particles — 9 bottles.” These pigments

demonstrate the fact that, although chemically identical, the cad¬

mium yellow (light, medium, dark), oranges and reds depend upon

the size of the grind to determine their value. The coarser particles

appear darker.

Box H contains a mixed group of lake (analine dye) colors and

assorted types. Box I, is according to Dr. Fischer’s note, “Color

types — Earths.” Box J is labelled “A series of the natural sources

of the dye” and includes samples of madder root from which the

Egyptians first extracted a red dye called Alizarin Crimson, and

Cochineal, dried bodies of an insect which yields a brilliant red,

1974]

La Mai fa — The Fischer Collection

219

Carmine, introduced to European artists in the 17th century. There

are 16 samples in this group and one of Dr. Fischer’s lectures to

the Art Student League of New York, on sources of the dye, sug¬

gests the reason for the inclusion of these samples. In addition,

Box J contains 18 bottles labelled “Color type pigments — blue-

green series.” Boxes K and L contain a complete Ostwald color

wheel, in powdered pigment.

In order to puzzle out the series, I researched Wilhelm Ostwald.

Dr. Martin Fischer was a friend of Ostwald and co-authored several

books with his son, Wolfgang. I assembled biographical information

on Wilhelm Ostwald and obtained the book Basic Color: an inter¬

pretation of the Ostivald color system by Paul Thebald. A thorough

reading of the book gave me the key to Dr. Fischer’s series of coded

envelopes.

The color sphere is stored, in Box K, a multidrawer wooden box.

Each compartment contains 24 envelopes and a letter code. It was

now obvious what Fischer had done. Each of the compartments

contains all of the 24 colors which Wilhelm Ostwald placed around

his color wheel. The letters on each package indicate the black and

white content and can be assigned a position thusly on one triangle

leaf of a three dimensional Ostwald color sphere. Each group of

color packs in Dr. Fischer’s series are equal white and black circles

or horizontal circles from the Ostwald sphere. The white and black

content remains equal all around the 360° of the sphere but the

hue changes from #1-24 (yellow to green). If one were to add a

binder to these coded pigments, one could produce a complete color

sphere.

Box M contains “Gums and oils used for making water and oil

color” and 20 samples of “Minerals — Natural — Colored.”

Box N contains what Dr. Fischer called his 0 and S system.

This consists of four wooden frames, much like test tube racks.

The first is marked S and like the others, contains a series of pig¬

ments in glass tubes. In a conversation with Henry Levison,

founder of Permanent Pigments, Inc., and himself a color chemist,

Fischer’s 0 and S concept was discussed. “It was,” said Mr. Levi¬

son, “correct in theory, but not necessary. The 0 colors (oxides)

supposedly could not be mixed with vermilion, whereas the S (sul¬

fides) could be.” This follows one of Dr. Fischer’s conditions for

permanency of pigment which states that pigments should not

interact chemically with each other within a painting.

Vermilion, discovered by the Egyptians, is a brilliant red called

Cinnabar by the Renaissance artists and is sulphide of mercury.

Mr. Levinson continued, “The breakdown of Vermilion was due

more to interactions with foreign substances in the earths (ochres) ,

not the earths themselves. Thorough washing of the earths by the

220 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

Permanent Pigments, Inc. has eliminated any bad effects of earths

on Vermilion.”

The other’ three racks contain the remaining S and two 0 series

of pigment. Box N also contains panels of oil paint which Dr.

Fischer tested for fading with exposure to hydrogen sulfide ( H2S ) .

These very same panels are reproduced in Dr. Fischer’s book “The

Permanent Palette ” (Plate II p. 12 — Effects of air pollutants

on some pigments). Incidentally, plate 1, p. 38, “Effects of light

exposure on some pigments,” which shows the effects of ultraviolet

light, was also found in this group of lecture notes. It consists of a

series of strokes of water color pigment on paper, some exposed

to sunlight, the others not.

With the Fischer collection safely stored in my studio, I spent

the remainder of my stay at Wittenberg researching Dr. Fischer.

I have included a biographical sketch of Martin Fischer from the

Wittenberg University Publication “Alumnus”, in the Appendix.

It should be obvious that Martin Fischer was a man of many

talents and interests, well traveled and internationally known. He

was a prodigious correspondent and the Thomas Library at Witten¬

berg has, in its archives, virtually every letter Dr. Fischer received

and saved during his lifetime. The letters are housed in bound

containers by year and contain many insights into Dr. Fischer’s

personality and interests.

I began sifting through the letters from 1929-1930 because 1930

is the year during which the book “ The Permanent Palette” was

published. The letters from publishers beginning in January 1929

provided a running commentary on Fischer’s efforts to get his book

published. A letter dated January 3, 1929 from Bridgeman Publish¬

ing Company, a firm that specialized in art books, states “We are

reviewing your manuscript.” A letter from Bridgeman dated March

23, 1929 rejects the manuscript. A letter dated August 10, 1939

from publisher Charles Thomas, provides some interesting com¬

ments on art and artists. Thomas rejects the manuscript because

he feels it is too well written for an artist!

Dear Dr. Fischer:

One of my friends has the largest book decorating organization,

another is the second largest buyer of colors, in this country. “The

Permanent Palette” was read by both with great interest. Both are

artists, and employ some of the best artists in the country. Both feel

that the manuscript is written with the authority on the chemistry of

color, but express the belief that there will be a very limited sale for

the book, for the reason that they believe it is too scientific in charac¬

ter to hold painters. They say it is almost impossible to get artists to

take time or make the effort to undertake serious study of color. Louis

Kreiger, a Baltimore artist, who has worked out with the physics

department of the Hopkins, the most superior color Atlas that I have

1974]

La Malfa — The Fischer Collection

221

seen, as well as Priest, of the Bureau of Standards, deplores the lack

of attention of artists to scientific fundamentals. They want to be

absolute free lances, seek no guidance.

I hate to tackle a crowd that does not desire knowledge as much as

I would like to do the MMS for the sake of the subject it represents.

So I have taken the liberty of returning the manuscript, not be¬

cause it is not good, but because of the special difficulties to be

encountered.

Faithfully,

Charles C. Thomas

Eventually the book was published by the National Publishing

Society (Fischer, 1930), and contributed to a reform of certain

questionable practices within the art products industry. From the

Thomas Library archives I turned my attention to the Alumni book

collection. The library had two copies of “The Permanent Palette”,

one of which I copied and eventually gave to Professor Lawrence

Rathsack.

I read the book and reread it looking for a key to Fischer’s pig¬

ment collection. The book itself and its basic format are the frame¬

work within which the collection made sense. In the Introduction

Dr. Martin Fischer states his thesis; modern painting as a craft

is declining. Many 19th century painters were seduced by the

spectral colors of analine dye pigments (lakes) and other non¬

permanent pigments and their work has dulled in a relatively short

period. Fischer attributes this to the artists’ ignorance of the

chemistry of paint, and his book is an attempt to remedy the

situation.

Dr. Fischer’s medical interests led him to study the colloidal

suspension of water in the cell. Thus when he turned his attention

to paint, also a colloidal suspension, he approached his subject in

a scientific and analytical manner. According to Dr. Fischer, the

following factors affect color permanence: (a) light, (b) air, (c)

intermixture with other colors, (d) reaction with the binder (me¬

dium), (e) reaction with the ground (primer) on the support

(canvas, paper, plaster) and (f) cold or warmth. To Fischer’s

list I would add the use of non-permanent binders and pigment,

exposure to atmospheric pollution and shoddy craftsmanship as

the greatest destroyers of paintings in the last one hundred years.

PHENOMENA THAT AFFECT PIGMENT

The ultraviolet portion of sunlight is a strong bleaching agent.

It tends to break down all but the simplest pigments such as the

ochres or cadmium compounds. Obviously, the informed collector

will not hang his paintings in direct sunlight. However, even arti¬

ficial light will cause non-permanent pigments to fade.

222 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Virtually all manufacturers of paint test their products for light

fastness. Permanent Pigments, Inc. tests their paints in the follow¬

ing manner : panels covered with pure paint and tints are mounted

at a 45° angle. They are exposed for a six month period to outdoor

sunlight. Then they are tested for fading with a colorimeter. These

paint tests are run at Cincinnati, where the home plant is located

and at Miami, Florida where a branch plant is located. Interest¬

ingly, the samples at Cincinnati faded more than those at Miami,

although subjected to less intense sunlight and fewer hours expo¬

sure. The extreme fading at Cincinnati is said to be caused not by

sunlight but by pollution in the air. Samples are also tested indoors,

subjected to fluorescent light at a distance of six inches. Both pure

and tinted hues are tested because, in use, artists seldom use pure

colors. Tints tend to fade more quickly than pure colors.

Why this great concern with fading? Did the artists of past

centuries experience this difficulty? No, not at any rate until the

1850’s, for up until then all commonly used artists’ pigments, even

the synthetic ones, were permanent. The first organic synthetic

pigment was Mauve, created by William Perkins in 1856. This pig¬

ment was a dye derived from the destructive distillation of coal

tar. The dye is precipitated on a white earth (alumina) also called

white clay. The result is the impermanent analine lake. All lake

colors are dyes characterized by their intense spectral hue, a large

and complex molecule, and by their impermanence.

All of these analine lakes with the exception of Alizarin Crimson

proved the undoging of the artists who used them, attracted by

the brilliance of these lakes. Fischer mentions Charles Duveneck

and James Abbott McNeil Whistler as two artists whose paintings

experienced fading, to which list I might add many lesser known

artists, numbering in the hundreds.

Alizarin Crimson, (surrogate) Ci4H804, resembled and replaced

traditional Alizarin, which was first used by the ancient Egyptians.

Traditional Alizarin is a dye extracted from the root of the madder

plant. Its 19th century substitute proved to be the only early analine

dye to be light fast. The other lakes all faded and, even today, many

artists consider dye colors suspect.

Dr. Fischer mentions air as potentially destructive to paint. The

effect of pollutants in our atmosphere is well known in 1973. The

pollutants in question are sulfide, sulphur dioxide, sulfuric acid,

carbonic acid, and water. Sulfide will cause lead white to yellow or

blacken in the following manner: the hydrogen sulfide produced

when coal or gas is burned acts upon lead carbonate and lead hy¬

droxide components of white lead and changes them to lead sulfide

which can appear yellow, orange or black.

1974]

La Mai fa — The Fischer Collection

223

What then is a safe palette ? Dr. Fischer describes the permanent

palette on page 33, chapter 7:

Black-Ivory black

Blue-Ultramarine blue

Green-Chromium oxide (Viridian)

White-Zinc white

Yellow-Pale cadmium

Yellow-Middle cadmium

Orange-Cadmium

Red-Cadmium

May also use: Vermilion-Yellowish to

orange

Earths

Y ello w-ochre-Light

Yellow-ochre-Dark

Gold ochre

Raw and burnt sienna

Raw and burnt umber

Mars yellow (earth)

Mars red

Mars purple

These colors are : light resistant, fairly bright, and incapable of chemical

reaction with each other and elements in the air.

If additional colors are needed, these may he added: Blue-Cobalt, Blue-

Cerulean, Green-Cobalt, Violet-Cobalt and Green-Terra verde.

Dr. Fischer recommends, as a primer for linen, one part zinc

white with one part lead white cut with turpentine. His preferred

medium was raw cold pressed linseed oil, bleached by sunlight. He

cautions against the use of boiled linseed oil or of driers and

siccatives.

The preferred varnish would be pure cold pressed linseed oil or

as a second choice, resinous varnishes such as Copal, Damar, Mastic

or Amber. The recommended solvent for thinning oil paint is gum

spirits of turpentine. . >

As stated earlier, my purpose in writing this paper is to encour¬

age other scholars and students interested in artists’ paints to

pursue the subject, using an extremely valuable tool, the Fischer

Collection. Those wishing to examine the specimens should contact

Mr. Joseph Emmuletti of the Milwaukee Public Museum. I would

also be happy to supply additional information such as the catalog

of the collection and related data. In addition, researchers should

try to locate copies of Fischer’s “The Permanent Palette ” published

in 1930; see References.

Let me conclude with a brief aside. In January 1973, I received

a letter from Miss Ilo Fischer, Thomas Library, Wittenberg Uni¬

versity, stating that the last remaining samples of Dr. Fischer’s

collection had recently been uncovered in the library storeroom.

“Would I like them sent to me?” Indeed I would was my reply.

This mini-collection was then sent to me by Mr. Louis Voight.

I have merely hinted at many possibilities for further study re¬

garding paint and would like nothing better than to see these areas

researched in depth by other scholars.

224 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

THROUGH HISTORY WITH THE FISCHER COLLECTION

The following1 list summarizes some of the more important pigments as they

were introduced in various Asian and European epochs. Each italicized pig¬

ment is found among the specimens in the Fischer collection.

The first group illustrates some of the pigments used by prehistoric artists.

Y ellow-Limonite or yellow ochre — an iron oxide, Red Ochre — red iron oxide

The next sample was used in the ancient Middle East or Mesopotamia and

is Black , Bitumen or asphaltum

Egyptian artists used the following: Green, Malachite — copper carbonate,

Red, Hematite — red iron ore, Blue, Azurite — blue copper carbonate, Red,

Alizarin Crimson — a dye from the madder root, Bright red, Vermilion or Cin¬

nabar — mercury compound, Yellow, Orpiment — arsenic compound

During the Middle Ages and up to the Renaissance all of the above were

used plus the following: Black Ivory — charred ivory, White Lead — -lead carbo¬

nate, Indigo Purple — dye from the Indigo plant

The Baroque and Rococo periods saw the introduction of: Cobalt Blue —

ground smalt (Venetian glass), Red, Minium — red lead oxide, Naples Yellow —

antimoniate of lead, Orange Realgar — disulphide of arsenic, Cambode Yellow —

solid resin from Cambodia, Carmine Red — dye from the cochineal insect,

Verdigris Green — made by mixing fermented grapes and copper (copper

acetate) .

The first pigment to be synthesized from raw materials was discovered by

accident, Diesbach’s blue, in 1704. The color is now called Prussian blue or

Berlin blue after the nationality of its discoverer: Prussian Blue-— iron ferro-

cyanide.

Cobalt Blue followed in 1802 and is cobalt oxide plus aluminum oxide.

Ultramarine Blue was synthesized in 1824 and is thought to be colloidal

sulphur in a glassy matrix.

Viridian, a dark green, was synthesized by Guignet, a Frenchman, in 1838.

It also goes by the name Verte Emeraude, or chromium green, and is hydrous

chromic oxide.

Cerulean Blue was created in the 19th century by mixing Cobalt Blue with

tin oxide, a white pigment. It is actually a tint of Cobalt Blue.

Zinc White was also developed in the 19th century and is zinc oxide.

As stated before the first organic synthetic pigment was Mauve which is

precipitated on alumina, or clay. Mauve and other analine dye colors, as first

produced, proved to lack light fastness.

In the early 1900’s, two inorganic pigments were introduced which are light

fast. They were: Titanium W hite — titanium oxide and the Cadmium pigments

which are based on chemically pure cadmium sulfo-selenide. They range from

Cadmium Yellow to Orange and Red. All are identical except for the particle

size of the pigment. The finer sized are lighter.

REFERENCES

FISCHER, MARTIN, The Permanent Palette, 1930. National Publishing So¬

ciety, Mountain Lake Park, Md. and 1819 Broadway, New York City.

WOODY, RUSSELL, Painting With Synthetic Media, Reinhold Publishing

Corp., New York City.

RESOURCE PEOPLE

MISS ILO FISCHER, Research Librarian, Thomas Library, Wittenberg Uni¬

versity, Springfield, Ohio.

MR. HENRY LEVISON, Permanent Pigments Inc., 2700 Highland Avenue,

Cincinnati, Ohio.

1974]

La Malfa — The Fischer Collection

225

PROFESSOR LAWRENCE RATHSACK, University of Wisconsin— Milwau¬

kee, Art Department.

MR. JOSEPH EMMULETTI, Geology Department, Milwaukee Public Museum.

APPENDIX

“Alumnus”

May 1962

Wittenberg University

Springfield , Ohio

MARTIN H. FISCHER

Dr. Martin Fischer, widely-known and versatile man of science,

art, and letters who directed the Department of Physiology and

taught at the University of Cincinnati College of Medicine for 40

years, died January 19 at his Cincinnati home after a long illness.

He was 82.

Dr. Fischer received Wittenberg’s honorary Doctor of Science

degree in 1932.

Dr. Fischer was one of those rare teachers who sparked interest

in medicine and other areas of the mind in generations of students.

His wide capacity for friendship made him one of the city’s best

known and loved persons.

Dr. Stanley E. Dorst, ’19, ?48H, dean of UC’s College of Medicine,

called Dr. Fischer “a unique person.”

“Not only was Dr. Fischer a physiologist and brilliant lecturer,

he was also an artist of considerable ability and a man of litera¬

ture,” Dr. Dorst pointed out.

“His translations of Gracian’s Truth-telling Manuel’ is a classic

and his books oh the lives of Christian R. Holmes and William B.

Wherry are monumental works.

“During Dr. Fischer’s term as director of the Department of

Physiology, two special units were developed with his aid and en¬

couragement. These are the widely-known Kettering Laboratories

in the University of Cincinnati Medical Center and Tanners’ Coun¬

cil Research Laboratories on Cincinnati University’s main campus.

Both were outgrowths of experimental work then going on in the

Physiology Department.”

In physiology Dr. Fischer was known for his early work in the

investigation of the colloidal chemistry of body tissues and for his

textbook on physiology of alimentation. Best known of his medical

writing was the book “ Edema and Nephritis .”

Dr. Fischer and his wife were major benefactors of Wittenberg.

Mrs. Fischer who preceded him in death, bequeathed the University

$127,000. Additional bequests from Dr. Fischer will raise the total

to approximately $220,000.

226 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Through the years Dr. Fischer made numerous gifts to Witten¬

berg including rare books, first editions, maps, and works of art.

His total benefactions include about 2,500 volumes, 350 bound

periodicals, several hundred monographs on medicine and fine arts,

valuable pieces of art and a collection of medals from the Society

of Medalists.

He also gave the University a 700 year old Catholic Breviary

containing what is believed to be one of the earliest forms of the

Rosary. The embossed hand-lettered Breviary, illuminated in color

and gold leaf, is the most valuable single item in Wittenberg’s rare

book collection.

Dr. Fischer’s experiments in the pigments of oil paint resulted

in the art pigment industry in America. He gave Wittenberg the

sequence of pigments that he used for his experiments. His book,

“The Permanent Palette resulted from this research.

Among the paintings given to Wittenberg by Dr. Fischer were

some of his own works ; paintings representative of the beginnings

of Japanese fine art, including prints by the famed Japanese artist

Kunisada; oil paintings by Miss Dixie Seldon, landscape and por¬

trait painter of Cincinnati; and John Weis, a portrait painter.

Dr. Fischer had served as director of the Department of Physiol¬

ogy at the University of Cincinnati College of Medicine for 40 years

before retiring in 1950. His key contribution in the area of physiol¬

ogy was in the area of colloidal chemistry of body tissue.

A PRELIMINARY SPATIAL ANALYSIS OF QUALITY

OF LIFE IN MILWAUKEE COUNTY, WISCONSIN

Richard A. Karsten

University Wisconsin — Milwaukee

Harold McConnell

Florida State University — Tallahassee

Thomas D. Patterson

Northern Illinois University — DeKalb

INTRODUCTION

The United States is a metropolitan nation : two-thirds of its

population lives in urban counties comprising Standard Metro¬

politan Statistical Areas. Moreover, ninety per cent live in or within

commuting distance of these urban areas. Despite the fact that

urbanization is generally equated with economic growth, metro¬

politan areas are deteriorating physically, socially, and economi¬

cally as the urbanization processes continue. In fact, many failures

in urban planning can be traced to lack of understanding of rela¬

tionships between social factors or conditions and economic and

physical parameters of urban systems. In short, there must be an

optimization of what people deem important within a social context

at the same time that economic and physical parameters of urban

systems are directed toward meeting a number of explicit goals

(Michelson, 1970).

THE STUDY

The social environment of a metropolitan parcel is mirrored by

the socioeconomic health or quality of life of that area. This study

compares three quality of life attributes elicited from a mix of 42

socioeconomic variables for 43 geographic areas or neighborhoods

in a representative urban area, Milwaukee County, Wisconsin, to

determine how quality of life varies spatially. Where quality of life

differs from place to place, certain urban activities are supported,

others are elicited, and still others are depressed as stress develops

which may evolve into urban strain (Michelson, 1970). This study

will attempt to depict the spatial variation of potential urban strain

as it is reflected in quality of life in Milwaukee County.

Quality of Life Factors

Zwerdling (1973) suggests that the basic environmental prob¬

lems of urban life have, as their origins, blight and poverty which

227

228 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

are in turn reflected in the quality of life of city dwellers. Stellman

(1973) states that more immediate indicators of blight and poverty,

such as state of health, educational level, housing quality, income

level, and employment status, are the basic ingredients of the qual¬

ity of life of an urbanscape. The 42 socioeconomic health indices

analyzed here mirror these components in Milwaukee County. With

respect to the neighborhood level of aggregation of the study, as

Rose (1969; 1971) suggests, the neighborhood may be considered

to be a social area consisting primarily of persons of a single race

and having similar subcultural characteristics that operate and

interact in a complex system of social and economic constraints at

various levels. Furthermore, the neighborhood can be considered

to be a socioeconomic-spatial institution which is an outgrowth of

complex socio-psychological decisions which are influenced by the

operation of economic forces (Rose, 1969).

The Immediate Environment and Quality of Life

Much of the ongoing research concerning environmental matters

focuses upon man's activities and interactions with the physical

environment, e.g., the land, the water, and the ambient air. These

may be considered epigene factors as opposed to the hypogene

factors that comprise the human or cultural environment. These

sets of factors do, indeed, interact with one another in a dynamic

system of inter and mutual dependence. However, for convenience

they may be treated as separate entities. The epigene factors pri¬

marily constrain place utility of the generalized environment (Aus¬

tin, 1971) and are outside of the scope of this study, while the

hypogene factors are composed of “people organized and interacting

in a group (society) and a distinctive way of life shared by mem¬

bers of the group (culture)" (Commins and Fagin, 1954). In the

microcosm this concept may be scaled to the individual dwelling

unit which becomes the immediate society, while the neighborhood

becomes the culture.

In order to evaluate the quality of life (mental well-being and

social satisfaction) of man, it is implicit that his immediate en¬

vironment be considered in the context of the home as it is reflected

in his neighborhood or culture. Commins and Fagin (1954) con¬

sider that the immediate environment is of far greater importance

in human development and quality of life than the physical environ¬

ment (except for its extremes). Although the home and neighbor¬

hood are tangible entities, they should be considered the result of

societal fabrication (Marcus and Detwyler, 1972) . These societal

fabrications are mirrored in what Lynch (1960) calls the percep¬

tion phenomenon of the city.

Michelson (1970) has approached the quality of life in urban

1974] Karsten , McConnell and Patterson — Spatial Analysis 229

areas by reducing social life into five basic components: life style,

stage in the life cycle, social status, value orientation, and per¬

sonality. It is implicit in Michelson’s analysis that various measures

of socioeconomic health would reflect varying levels of quality of

life in urban areas.

Scale of Research and Data Collection

Beverstock and Stuckert (1972) have divided Milwaukee County

into community areas, using methodology developed by the Social

Science Research Committee of the University of Chicago (1963)

(see Figure 1). These community areas are treated as neighbor¬

hoods here, since their delineation was based upon criteria which

include internal homogeneity of various socioeconomic attributes.

Wolpert, et ah (1972) points out that although neighborhoods are

difficult to delineate, the neighborhood unit and the home or im¬

mediate environment are the most incisive units for urban analyses.

Although Michelson (1970) considers neighborhood to be a “slip¬

pery term,” he suggests that cohesive delineation of such units is

the most viable approach to urbanscape analysis.

The same authors (Beverstock and Stuckert, 1972) also col¬

lected and developed socioeconomic data from 16 U.S. Bureau of

the Census sources for the Milwaukee S.M.S.A. and its various

components, including the community areas or neighborhoods of

Milwaukee County. These data include general characteristics of

the population, housing, marital status, age, sex, race, families,

households, labor force, occupation, income, nativity, ethnicity,

vital statistics, infant mortality, mobility, and educational level.

Forty-two variables were selected from the above set which were

deemed to be measures of socioeconomic health for the 43 com¬

munity areas or neighborhoods of Milwaukee County and which

possess those attributes which reflect quality of life or the popula¬

tions’ mental well-being and degree of social satisfaction (see

Table 1). The County Institutions community area is not included

in this study.

The Analysis

As is characteristic with geographic data bases, none of the

indices is arrayed in accord with the normal probability law. Each

of the variables exhibited positive skewness and was transformed

into its common logarithm prior to the analysis of the data.

Principal Component Analysis. There are numerous redundancies

among the 42 incides which were deemed to describe the socio¬

economic health of Milwaukee County. Principal components analy¬

sis was employed to collapse this set of colinear variables into a

smaller number of basic dimensions which are the mutually inde-

230 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

FIGURE 1. Location of Milwaukee County community areas

(after Beverstock and Stuckert, 1972) 4

‘County Institution community area 40 is not included in this

study.

1974] Karsten, McConnell and Patterson — Spatial Analysis 231

LEGEND FOR FIGURE 1

Identified- Community Identified -

pendent (mathematically orthogonal) attributes of the variables.

Similar procedures were employed by Thompson, et al. (1962) in

a study of economic health in New York State and by McConnell

(1970) to analyze the socioeconomic health of large Illinois munici¬

palities. The method is described in brief in the following general

statement.

Overvieiv of Methods and Procedures. The straight-forward

objective of principal components analysis is to transform a set

of m standardized variables into a set of m orthogonal variables.

However, it is generally used to collapse m standardized variables

into a smaller set (p) of orthogonal variables or fundamental

attributes of the original variables. Denote the following:

Z — N X m matrix of standard scores

R — m X m matrix of Pearson product moment correlations

between the standard scores

P — m X p matrix of correlations between variables and

attributes

Pt — p X m transpose of P

P' — m X p varimax rotation of P

C — N X p matrix of standard scores on the attributes

232 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Each variable is standardized about its mean. Correlations be¬

tween the standardized variables are arrayed in an m X m matrix

(R) which is subjected to the principal component analysis to

transform the original indices into mutually exclusive attributes

or principal components :

( Zx , zx , . . . , zx \

\ 1 2 ill /

TABLE 1. VARIABLES AND PRINCIPAL COMPONENTS ANALYSIS

1974] Karsten, McConnell and Patterson — Spatial Analysis 233

is transformed into a new set of orthogonal vectors or principal

components

(Pi, P2, . . . , Pm)

Those p principal components whose eigenvalues (Aj) are 1.00 or

larger are usually considered the basic dimensions in R or the

fundamental attributes of the variables.

The eigenvalues in a principal components analysis are the latent

roots of the m characteristic equations which set the determinant

of R at zero:

Given Pu as the correlation between the ith standardized variable

and the jth principal component (or as the ith coefficient of the jth

component), a series of equations of the following form is

developed :

(2)

P"ll + P“21 + . • • + P2ml — Ai

P“l2 T P“22 T" • • • H~ p2m2 — A 2

P“lm + P22m + • • • + P2mm — Kn

The function of the first principal component is to resolve a max¬

imum amount of variance in R: succeeding principal components

resolve as much as possible of the remaining variance through m

iterations. Thus, it is necessary to maximize Ax initially and

Larangian multipliers are used to obtain

The first root is the eigenvalue corresponding to Pi and the sec¬

ond through mth roots correspond to P2, P3, . . ., Pm, respectively

and are the largest roots of the residual correlation matrices Ri,

R2, . . ., Rm— i, respectively. The coefficients (pu, p2X, . . pmi) of

Pi are obtained by scaling its normalized eigenvector by Ax :

(4) Pil = an (Ai) 1/2

The coefficients of P2, P3, . . . , Pm are obtained by similar opera¬

tions on the residual correlation matrices (R-PP1) until

p = m

m

2Aj = m

PP* = R

234 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

It has already been noted that the second root of (3) is the eigen¬

value corresponding to P2, and so on.

Varimax Rotation. Labeling of the attributes of R is facilitated

by transforming P to P', using a varimax criterion in which P is

rotated to that angle (0) in the plane which will maximize the

variance of the principal components, subject to maintaining each

variable’s communality (h2i). This has the effect of increasing the

absolute magnitude of those loadings (py’s) whose absolute values

are initially high and minimizing those whose absolute values are

initially low. Generally, only those principal components of P whose

eigenvalues are larger than 1.00 are rotated. Thus,

(5) h2i = | p2ij

Specifically, 0 is the angle of rotation from which the transfor¬

mation matrix

cos 0 — sm 0‘

(6) T =

_sin 0 cos 0.

may be developed to maximize the function

(7) V = m l [™ (Pu/hi)4 - l P2ij/h2i ) 2 ]

Scores on Principal Components. The m standardized variables

may be condensed into p orthogonal standardized variables. Evi¬

dently, the most useful algorithm is

(8) C = ZPA-1

where A-1 is the inverse of the diagonal-eigenvalue matrix. Zero

is of a larger order of magnitude than say, —1. In a spatial situa¬

tion, Cij is the standardized magnitude of attribute j at site i. Such

scores are distributed as the standard normal deviate with mean

zero and unit variance if the original data were normally

distributed.

The primary function of the analysis was to elicit the basic

attributes of the 42 transformed variables. Thus, a 42 X 42 cor¬

relation matrix was computed and collapsed into mutually exclusive

attributes. Despite the fact that several eigenvalues were larger

than one, it was deemed germane to examine only the first three

principal components since they resolve, respectively, 50.5 percent,

17.2 percent, and 8.3 percent of the variance in the original cor¬

relation matrix, or collectively, 76.0 percent of its variance. The

attributes were identified after varimax rotation and quantified

according to the algorithm given in (8) for each neighborhood for

mapping purposes.

1974] Karsten, McConnell and Patterson — Spatial Analysis 235

Interpretation of Attributes. The first principal component (Pi)

is interpreted as a positive indicator of poverty. Neighborhoods

with high positive scores (cu) on this attribute would be expected

to have low levels of achievement in such areas as education, in¬

come, quality and value of housing units. Such neighborhoods

would typically have high birth and fertility rates, comparatively

large numbers of families and individuals living below poverty

levels, low levels of home ownership by occupants, and fewer people

employed in the so-called “white collar” category than neighbor¬

hoods having near-zero or negative scores on this indicator.

The second principal component (P2) is deemed to be indicative

of the youthfulness of the population. Neighborhoods with high

positive scores on this attribute would be expected to have low

median ages for both males and females, comparatively large pro¬

portions of the population younger than the age of 20, small pro¬

portions older than 60, and associated low death rates.

The third principal component (P3) is identified as an ethnicity

attribute. Neighborhoods with high positive scores on this attribute

would be expected to have comparatively large proportions of the

population classified as foreign stock as well as small proportions

listed as Black. Positive scores would also be indicative of propor¬

tionately large numbers of males in the labor force, high propor¬

tions of the population completing high school, and most households

represented by both husbands and wives. Such characteristics are

those which are traditionally associated with the stereotyped ethnic

White neighborhood.

Maps of Scores on Principal Components

As the intensity of the shading increases, the presence of the

attribute measured by the principal component increases relative

to areas of less intense shading. No absolute values are attached to

either rates of occurrence of an attribute within an area or to

rates of change between areas. In other words, comparisons be¬

tween areas are of the “greater than”, “less than”, or “approxi¬

mately equal to” nature.

The map of scores on the first principal component (cu) depicts

a readily discernible gradient away from the center of the city:

high positive scores on the poverty attribute of the data are cen¬

tered on Halyard Park and Garfield, economically and socially

depressed sections of the county. Low and negative scores occur as

one moves away from the downtown area (Figure 2, Table 2).

The map of scores on the second principal component (ci2) indi¬

cates that the youngest populations are in the “core” of the city

(e.g., Garfield, Halyard Park, and Midtown) ; in the northwestern

236 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

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Attribute ( Poverty )

1974] Karsten, McConnell and Patterson — Spatial Analysis 237

TABLE 2. RANK ORDER OF NEIGHBORHOODS ON THE FIRST

SOCIOECONOMIC ATTRIBUTE (POVERTY) MAP CLASSES—

RANKED HIGHEST TO LOWEST

Halyard Park

Garfield

Midtown

Walker’s Point

The Valley

Grand Avenue

Riverside West

Kosciuszko

Juneautown

Lincoln Creek

Muskego Avenue

Tippecanoe

Silver Spring

Layton Park

Sherman Park

West Milwaukee

Old West Allis

Johnson’s Woods

Bay View

Cudahy

Lakeside

North Milwaukee

St. Francis

South Milwaukee

Lake

Oak Creek

Granville

West Allis Addition

Jackson Park

Franklin

Shorewood

Wauwatosa Avenue

Greenfield

Wauwatosa East

Wauwatosa West

Glendale

Greendale

Hales Corners

Brown Deer

Whitefish Bay

River Hills

Fox Point

Bay Side

portion of Milwaukee County (Granville, Silver Spring, and Brown

Deer) ; and in the southern portion (Oak Creek, Franklin, and

Greendale) (Figure 3, Table 3).

The map of scores on ethnicity (ci3) shows that high degrees of

this attribute are concentrated almost exclusively to the south of

the Menominee River. Highest scores are found in such neighbor¬

hoods as The Valley, Cudahy, St. Francis, Tippecanoe, Lake, Jack-

son Park, and Kosciuszko. Low and negative scores evidently

indicate one of two things; either a lack of people classified as of

foreign stock or the presence of large numbers of Blacks in the

population. This accounts for the fact that such neighborhoods as

Whitefish Bay and Halyard Park have similar scores on this com¬

ponent (Figure 4, Table 4).

TABLE 3. RANK ORDER OF NEIGHBORHOODS ON THE SECOND

SOCIOECONOMIC ATTRIBUTE (YOUTHFULNESS) MAP CLASSES—

RANKED HIGHEST TO LOWEST

238 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

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Socioeconomic Attribute (Youthfulness)

1974] Karsten, McConnell and Patterson — Spatial Analysis 239

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FIGURE 4. Distribution of Scores on the Third Socioeconomic

Attribute (Ethnicity)

240 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 4. RANK ORDER OF NEIGHBORHOODS ON THE THIRD

SOCIOECONOMIC ATTRIBUTE (ETHNICITY) MAP CLASSES—

RANKED HIGHEST TO LOWEST

The Valley

Cudahy

St. Francis

Tippecanoe

Lake

Jackson

Kosciuszko

West Milwaukee

Muskego Avenue

Layton Park

Oak Creek

Silver Spring

Granville

Old West Allis

West Allis Addition

Walker's Point

Bay View

Greenfield

South Milwaukee

Hales Corners

Brown Deer

Wauwatosa Avenue

Franklin

Sherman Park

North Milwaukee

Greendale

Johnson’s Woods

Wauwatosa West

Riverside West

Lakeside

Glendale

Juneautown

Wauwatosa East

Shorewood

Grand Avenue

Lincoln Creek

Midtown

Whitefish Bay

Garfield

Fox Point

Bayside

River Hills

Halyard Park

Quality of Life Index

A synthetic quality of life index was constructed using a linear

combination of each neighborhood’s scores on the three principal

components, i.e.,

(9) QLj zz — AjCii + A ^Ci2 + A 3Ci3

Since the eigenvalues are the coefficients of the scores, each com¬

ponent contributes to the magnitude of the index in an amount

directly proportional to the amount of the variance which it re¬

solved. Thus, the index is heavily weighted by the first principal

component, whose coefficient is given a negative sign since it meas¬

ures a poverty attribute. Nevertheless, interesting patterns emerge

when the index is mapped (Figure 5, Table 5). As with the maps

of component scores, only comparisons of the “greater than”, “less

TABLE 5. RANK ORDER OF NEIGHBORHOODS ON THE QUALITY

OF LIFE INDEX MAP CLASSES— RANKED HIGHEST TO LOWEST

Fox Point

River Hills

Bay Side

Whitefish Bay

Glendale

Wauwatosa East

Shorewood

Wauwatosa West

Wauwatosa Avenue

Hales Corners

Lakeside

Brown Deer

Greendale

North Milwaukee

Greenfield

West Allis Addition

Sherman Park

Johnson’s Woods

Jackson Park

Bay View

Franklin

Juneautown

Old West Allis

South Milwaukee

St. Francis

Layton Park

Lake

Granville

Oak Creek

Cudahy

Tippecanoe

West Milwaukee

Grand Avenue

Silver Spring

Muskego Avenue

Lincoln Creek

Kosciuszko

Riverside West

Walker’s Point

The Valley

Midtown

Garfield

Halyard Park

1974] Karsten, McConnell and Patterson — Spatial Analysis 241

- — + — • — 2 3~ — ~~ 4--" — •— — 4~ — — — #

I I

i 00000++++++++++++. .. . i

[ 000000+++++ ++++++. ••••••••• • I

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+ 000000000++ +++2++ . . . +

I €000800000 + + + +++ + . .X . I

I 00000000000+ + + + + +•••••••••« • l

I 00008000OU80+ ++++.••••••• 1 * • I

I 80008000480600 + + + • ••••••••• I

X €000000008 000000 + .... . 1

I 00800080008808000 ......... I

I 0000000080 0800088 ........ I

I 80000000080800800 . 1... I

l 0006000000004 8000 ......... I

+ ... ..08080800800000. . .v. .. . +

I •••••«• €600000+ ++00000 ... 1 • • I

I . .... + + ++ + + ++008008 . I

I ... ....... + + + 2 + ++0 04 0000...... I

1 ........... ++++++080000BBB. . X. I

2 ........... ++ ++++80008 Mill. ... 2

I ...... .1.. . .+++++86611111 588. •+ I

I . .1 . ++++++BBBBB5BBBBB++++ I

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i . .+++2++a5aaaaaao+++++ i

+ . . ++++aaaaaB5aaooo++ +

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I ••••••••••••« • ++B00008BOOOQQ I

I +++ . +++++++00000400BBaa I

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+ +++ ++ + 0 COD 00000000 O0O0BB5BBOO +

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I +++ ++ + ++++ + + +++++ 00000 00 3 000008 400 00 I

+ ++ 2 + ++ + +++ 2 ++ ++++0 0000 000 00 0086008 08 +

I +++ +++ + +++ + ++ + +++ 00000 000 000888800 88 I

I +++++++++++++++++000000000000860800 I

I ++ + + + 4- +++ + + + + + + + + + QLJOU QOO 00 000000000 I

I ++++++++++++++++++000000000000000000 I

5 +++ ++Q00++ + + + + + + + + 00000080000000 30 00 5

I +++OOOOOOOOOOOOO++e88066080OOOOOOOOO I

I 00000000000000000000800800000000000 I

I 00000000000000000008000000000000000 I

I 00000000000000000800008800000000000 I

+ 000 000 0G3 00 00 00000 80868800800000000 +

I 000000000000000008000088804000088000 I

I 0000000 000 0000 00008080 680006000880 00 I

I 0000000000000000008888800880008000080 I

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i oooooGoooQooooooo8e000ee00ee88880e0800 i

I 00000000000000000080000000080000008000 I

I I

— 1 — — — 2— + — — 3 - +— — 4 —

FIGURE 5. Distribution of Scores on the Quality of Life Index

242 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

than7', or “approximately equal to” nature are possible. Those

neighborhoods which might be deemed least advantageous or most

susceptible to urban strain from the socioeconomic standpoint form

two parcels separated by the commercial development of the Grand

Avenue neighborhood (Milwaukee’s most intensively commercial

area) . The northern parcel is comprised of Halyard Park, Garfield,

Midtown, and Riverside West. The southern parcel consists of The

Valley, Kosciuszko, and Walker’s Point. Those neighborhoods for

whom quality of life is evidently the highest also form two essen¬

tially contiguous groups; Wauwatosa Avenue, Wauwatosa East,

and Wauwatosa West in the west-central section of the county and

Shorewood, Glendale, Whitefish Bay, Bay Side, River Hills, and

Fox Point in the northeastern section.

CONCLUSIONS

This study has shown in a quantitative manner that there is

considerable spatial inequality in the quality of life in Milwaukee

County — that there is a large amount of neighborhood to neighbor¬

hood variation in its socioeconomic health. This is a condition which

is suggestive of potential urban strain, or as Harvey (1972) has

noted, tension between the spatial organization of the present

society and that spatial organization which might be demanded by

a new social order. One might advocate that such a spatial organi¬

zation must, in an economic and social context, generate a social

surplus product if urbanism is to survive. Hence, this study should

not only provide a data base for both public and private agencies

for research and planning, but also serve as a vehicle to promote

intensive social change in those areas of Milwaukee County deemed

lacking in the socioeconomic health or quality of life enjoyed by

other neighborhoods.

REFERENCES CITED

AUSTIN, C. M. 1971. Urban public facilities: location and impacts. Unpub¬

lished Ph.D. Dissertation, Department of Regional Science, The University

of Pennsylvania, Philadelphia. 193 pp.

BEVERSTOCK, F., and STUCKERT, R. P. (eds.). 1972. Metropolitan Mil¬

waukee fact book: 1970. Milwaukee Urban Observatory, Milwaukee, Wis.,

549 pp.

COMMINS, W. D., and FAGIN, B. 1954. Principles of educational psychology.

2nd ed., Ronald Press, New York, 795 pp.

HARVEY, D. 1972. Society, the city and the space-economy of urbanism.

Commission on College Geography, Resource Paper No. 18, Assoc. Amer.

Geographers, Washington, D. C., 56 pp.

LYNCH, K. 1960. The image of the city. M.I.T. Press, Cambridge, Mass.,

194 pp.

MARCUS, M. G., and DETWYLER, T. R. 1972. Urbanization and environ¬

ment in perspective: in Urbanization and environment: the physical

1974] Karsten, McConnell and Patterson — Spatial Analysis 243

geography of the city. T. R. Detwyler and M. G. Marcus (eds.), Dux-

bury Press, Belmont, Cal., p. 3-25.

McCONNELL, H. 19?0. Socioeconomic health of Illinois municipalities. Trans.

Illinois State Acad. Sci. 63:273-284.

MICHELSON, W. 1970. Man and his urban environment: a sociological

approach. Addison-Wesley, Reading, Mass., 242 pp.

ROSE, H. M. 1969. Social processes in the city: race and urbanism residen¬

tial choice. Commission on College Geography, Resource Paper No. 6,

Assoc. Amer. Geographers, Washington, D. C., 34 pp.

ROSE, H. M. 1971. The black ghetto: A spatial behavioral perspective. Mc¬

Graw-Hill, New York, 147 pp.

Social Science Research Committee. 1963. Local community fact book, Chicago

metropolitan area, 1960. Social Science Research Committee, Univ. Chi¬

cago, Chicago, p. xiii-xiv.

STELLMAN, S. D. 1973. Director, Center for Continuing Education & Com¬

munity Action for Social Service, University Wisconsin-Extension. Per¬

sonal communication.

THOMPSON, J. H., et al. 1962. Toward a geography of economic health: the

case of New York State. Ann. Assoc. Amer. Geographers, 52:1-20.

WOLPERT, J., et al. 1972. Metropolitan neighborhoods: participation and

conflict over change. Commission on College Geography, Resource Paper

No. 16, Assoc. Amer. Geographers, Washington, D. C., 51 pp.

ZWERDLING, D. 1973. Poverty and pollution. Progressive 37:25-29.

PRIMES AND FAREY SEQUENCES

Arthur Marshall

Madison

This paper is aimed at helping teachers of pre-college or fresh¬

man mathematics to demonstrate how prime numbers arise in the

natural (“counting”) number system and how prime numbers

can be “predicted” from examination of numbers less than them¬

selves.

We will show that primes are the result of adding the numerator

and denominator of certain “Farey Sequences” fractions ( a/b

with 0 < a < b, b g 2, and ( a , b) =1), starting with 1/2. (We

have just used the greatest-common-divisor symbol of number

theory literature.)

All letters stand for positive integers (sometimes called natural

numbers or counting numbers).

THEOREM: If (a, b) — 1, then (a, a + b) =1 and

(b, a + b) =1. Conversely, if 0 < a < n/2, and ( a , n) ~ 1, then

(a, n — a) =1 with a/(n — a) <1, and ( n — a, n) ~ 1, with

(n — a)/n < 1.

PROOF : If a = 1, the theorem is obvious.

If a > 1, then (using congruence notation of elementary number

theory) a = 0 (mod p) , where p is any prime dividing a. Because

(a, b) = 1, b 0 (mod p) , then, by addition of congruences

(a + b) 0 (mod p) , and the first assertion of the first part of

the theorem is proved. We follow an analogous procedure for q, any

prime dividing b, and the second assertion of the first part is

proved.

To prove the converse, we note that p, any prime dividing a,

cannot, by hypothesis divide n. Then (n — a) — n 0 (mod p)

and the first statement of the converse is proved. To prove the

second statement of the converse, we note that qf any prime divid¬

ing n, cannot, by hypothesis divide (n — a), because (n — a) =

—a (mod q) , and that completes proof of the theorem.

COROLLARY 1. All fractions in Farey sequences, lying between

0 and 1, can be obtained by successive additions of numerators and

denominators, beginning with 1/2, to obtain new fraction denomina¬

tors and taking each summand numerator and denominator as

numerators over the new denominators. (The proof is immediate

from the theorem.)

245

246 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

We now use the usual Greek symbol </> (phi) for the Euler

totient: $(n) is the count of numbers less than n which have no

common divisor with n greater than 1.

COROLLARY 2. For n > 2, there exist </> (n) /2 Farey fractions

of the type a/ (n — a) with denominators less than or equal to

(n — 1).

PROOF. Immediate from Corollary 1.

COROLLARY 3. For n > 2, , n is prime if, and only if, there

exist (n — l)/2 Farey fractions of the type a/ (n — a) with de¬

nominators less than or equal to (n — 1).

PROOF. By definition, n is prime if, and only if, n has just two

divisors: 1 and n, where n > 1. If there exist fewer than (n — l)/2

Farey fractions of the type a/ (n — a), with denominators less

than or equal to (n — 1), then n has at least one divisor lying be¬

tween 1 and n. If there exist just (n — 1)/2 Farey fractions of the

above type, our definition of primes is satisfied.

ALTERNATIVE PROOF. Directly from Corollary 2.

COROLLARY 4. There exist exactly (n — 1) Farey fractions

with denominator n, if, and only if, n is prime.

PROOF. For n = 2, this is immediately verified. For n > 2,

this follows immediately from Corollary 3 and our theorem.

ALTERNATIVE PROOF. Directly from our definition of prime

numbers.

The writer believes quick primality tests and factoring algorithm

programs can be written from this theorem and its corollaries, but

will leave that to computer specialists. Any such programs, of

course, would have to be competitive with those arising from Prob¬

lem E 2355 [1] and the more sophisticated programs discussed by

Collins [2].

1. Problem E 2355, Amer. Math. Monthly, 80:560-561. 1973. (Solution by R. J.

EVANS. Proposed by A. MARSHALL.)

2. G. E. COLLINS. Computer algebra of polynomials and rational functions.

Amer. Math. Monthly, 80:725-755, 1973.

Correction

Congruence symbol “ = ” and non-congruence symbol “=j|” in the

text of this paper should have three bars, but the correct type items

were not available.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN

NO. 64. ADOXACEAE — MOSCHATEL FAMILY

Theodore S. Cochrane

University Wisconsin — M ndison

and

Peter J. Salamun

University Wisconsin — Milwaukee

The Adoxaceae consists of a single worldwide, north temperate

species which occurs rarely in the United States, including Wis¬

consin. The family is usually included in the Rubiales, although

some authorities consider it to be more closely related to the Saxi-

fragaceae or possibly the Ranunculaceae. According to modern

opinion Adoxa is a specialized offshoot of the Caprifoliaceae (Cron-

quist 1968), the family in which it was listed in an earlier Wis¬

consin report (Wade and Wade 1940).

The distribution of the species, habitat information and dates

of flowering and fruiting were compiled from specimens in the

herbaria of the University of Wisconsin — Madison (WIS), Uni¬

versity of Wisconsin — Milwaukee (UWM) , University of Wisconsin

Center — Rock County (ULJ), Milwaukee Public Museum (MIL),

University of Iowa (IA), University of Michigan (MICH) and

the University of Minnesota (MIN) . Each dot on the map indicates

a specific location where a specimen was collected, while triangles

represent county records taken from the literature. Additional

records from Hartley’s unpublished “Flora of the Driftless Area”

(1962), Lakela’s Flora of Northeastern Minnesota (1965) and

Morley’s Spring Flora of Minnesota (1966) have been included.

The numbers within the squares in the lower left-hand corner of

Map 1 indicate the number of Wisconsin specimens noted that

were flowering or fruiting in the respective months. Specimens in

vegetative condition, in bud or with immature or dispersed fruits

are not included.

Grateful acknowledgement is made to the curators of the above

herbaria for loans of specimens and especially to Dr. Hugh H. litis

for discussing the material contained in this paper with the first

author.

ADOXACEAE J. G. Agardh Moschatel Family

A monotypic family of circumboreal distribution, extending

southward in North America to northern New York, northwestern

Illinois, South Dakota and in the Rocky Mountains to New Mexico.

247

248 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

jae° ’

ADOXA MOSCHATELLINA

1. ADOXA L. Moschatel, Muskroot

1. Adoxa moschatellina L. Maps 1, 2.

Moschatel, Muskroot

Low delicate perennial 5-15 (-20) cm high with musk-scented,

white-scaly rhizomes and an erect, simple, glabrous stem. Basal

leaves 1-3, long-petiolate, ternately compound, the ovate to obovate

or orbicular leaflets yellow-green, membranous, coarse-toothed to

3-parted, obtuse, glandular-mucronate. Cauline leaves an opposite

pair, 3-parted, smaller, less divided and shorter petioled than the

basal leaves. Inflorescence an angular , cymose head 6-8 mm in

diam, the slender peduncle often overtopped by the basal leaves.

Flowers small, 3-8, usually 5, regular or nearly so, perfect,

sympetalous. Calyx 2-4-lobed, persistent in fruit. Corolla U-6-lobed,

yellow-green, rotate, the lobes 1.7-3 mm long. Stamens twice as

many as the corolla lobes, in epipetalous pairs on the corolla tube

and alternate with its lobes, separate or partly united; anthers

1 -celled. Ovary semi-inferior, 3 -5 -celled; style short, deeply 3-5-

parted, the stigmas minute. Fruit a dry drupelet, green; nutlets

(1-) 3-5, lenticular, cartilaginous, 2.9 mm in diam. 2 n — 36 (54,

Japan) .

Rare and very local in cool moist woods under coniferous trees

on bare or mossy ground, sometimes in leaf mold, on shaded, usually

north-facing sandstone or limestone bluffs and talus slopes, rarely

in mesic hardwoods. Since Adoxa is of unusual biogeographic in¬

terest, all Wisconsin collections are cited herewith : Grant Co. :

Platteville (Smithyman s.n., 4 May 1896, MIL, WIS). [Pierce Co. :]

near Plum City, between Durand and Ellsworth ( Butters UhSU,

1974] Cochrane and Salamun — Moschatel Family Report 249

1 June 1924, MIN, UWM). Richland Co.: W-facing sandstone cliff

near Butternut School, sec. 34, TUN, R2R, 4 mi. N of Ithaca (Fas-

sett 26325, 2 June 1946, WIS). Rock Co.: limestone outcrops and

damp shaded slopes on S bank of Turtle Creek, with Tilia ameri-

cana, Garex alhursina , Erythronium albidum, Claytonia virginica ,

Isopyrum biternatum, Dicentra canadensis , D. cucullaria , Dentaria

laciniata , Miteila diphylla , etc., sec. 28, T3N [sic, for T2N], R14E

[3 mi. NE of Clinton] ( Musselman 3487, 2 May 1970, ULJ) ; same

location (Musselman 3505 , 6 May 1970, ULJ, UWM). Sank Co.:

small patch in liverwort and moss mat on moist N-facing sandstone

ledge along Raraboo River, shaded by Tsuga canadensis and Betula

alleghaniensis , sec. 20, T13N, R3E [1 mi. NW of La Valle] (Nee

& Feet 2234 , 18 July 1969, WIS) ; same location (Cochrane & Nee

5639 , 31 July 1973, WIS). Vernon Co.: Ontario (Fasseit 17963 ,

June 1936, WIS) ; talus at base of shaded sandstone ledge along

Kickapoo River, Wildcat Mt. State Park, sec. 14, T14N, R2W

(Hartley & Morrissey 9146 , 14 May 1960, IA, WIS) ; under conifers

just above sandstone ledges along Hemlock Trail, S end of Wildcat

Mt. State Park (Salamun & Matthias 2879 , 2 June 1970, UWM).

This species, not uncommon in northeastern Minnesota, should be

sought in the Lake Superior area of Wisconsin. Flowering 2 May

to 1 June; fruiting June to July.

The terminal flower is usually tetramerous, with 4 styles and 8

anthers, while the lateral ones are pentamerous or hexamerous,

with 5 or 6 styles and 10 or 12 anthers. Each pair of stamens

probably represents a single one which is divided nearly to the base

of the filament, with each segment bearing a half-anther. The pos¬

sibility that the calyx and corolla may constitute an involucre and

calyx contributes to the uncertain relationships of this family (cf.

Sprague 1927).

In an attempt to relocate Adoxa moschatellina the senior author,

accompanied by Michael Nee, visited on July 31, 1973, two water-

worn, upper Cambrian sandstone cliffs on which the species was

known to occur. The most conspicuous feature of such cliffs, which

are common in the Driftless Area, is the presence on many of them

of “relict” stands of conifers. These stands are intriguing, for they

are composed of species which are chiefly northern in distribution

and are rare or unknown in southern Wisconsin outside of that

famous region. The Butternut School station has long been known

to be botanieally rich, ever since Norman C. Fassett found there

both Adoxa and Ledum groenlandicum . Had he ventured a little

farther eastward to the north-facing portion of the cliff, unreach¬

able from below due to undercutting by Willow Creek, Fassett

would have encountered an idyllic cove on whose steep cool slopes

250 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

and cliffs, shaded by yellow birch and white pine, grow numerous

mosses and liverworts, including the rare sword moss (Bryoxi-

phium norvegicum) and a six-meter-square patch of Sphagnum.

The associated understory consists of a large number of typically

northern species : Equisetum scirpoides, Lycopodium selago, L.

lucidulum, L. ohscurum, Coptis groenlandica, Cornus canadensis,

Chimaphila umbellata, Gaultheria procumhens, and Mitchella

repens. The Adoxa was not found in 1973, and it is not known

whether or not colonies persist on this mile-long cliff, which bo-

tanically must be among the most remarkable ones in the Driftless

Area.

Mr. Nee discovered a population of Adoxa on the west end of

Horse Bluff, near LaValle in Sauk County, at the time of his first

visit. Although the plants are inconspicuous, the distinctive scintil¬

lating quality of their pale leaves and the absence of nearby vege¬

tation enabled us to find them again after a brief search. There are

only three small colonies, consisting of 21, 39, and 61 plants and

growing on narrow ledges one to two and one-half meters long

along the base of a small cove formed by low cliffs four meters

high. These sandstone ledges are relatively free of vegetation ex¬

cept for mosses, the liverwort Conocephalum conicum, and a few

plants of Pilea. The shallow mineral soil in which the plants grow

has an acid reaction, with a pH of 5.2. A fine stand of hemlock and

yellow birch occupies the steep rocky north-facing slopes. The

shrubs found in the stand show a resemblance to those of the

northern forests and include Taxus canadensis, Acer spicatum, and

Sambucus pubens. The conspicuous herbs in the nearby understory

are Laportea canadensis, Athyrium thelypteroides, and Dryopteris

austriaca var. intermedia. Certainly equivalent habitats are fre¬

quent in the Driftless Area, but most of them are not occupied by

Adoxa moschatellina.

The range descriptions given for Adoxa in the regional manuals

are incomplete, for Fernald (1950) omitted its occurrence in north¬

ern Illinois as did Gleason (1952) in northern New York. In south¬

ern Wisconsin and northwestern Illinois (that state’s only station

in Apple River Canyon) it is at the southeasternmost limit of its

range, except for two reports from Delaware Co., New York (Tay¬

lor 1913; Brooks 1960). House (1924, p. 655) suggested that the

one station known to him was an introduction, but surely plants

of this isolated locality high in the Catskills at 1,400 feet represent

native populations. The geographic distribution of Adoxa suggests

that in the Midwest it is a western immigrant that crossed the

Bering Straits and barely reached Wisconsin. With the exception

of New York it is totally lacking from eastern North America.

1974] Cochrane and Salamun — Moschatel Family Report 251

In the Midwest the distribution of Adoxa is correlated with, but

not restricted to, the “Driftless Area” of Wisconsin and adjoining

Minnesota, Iowa and Illinois (Map 2), a coincidence probably due

to the presence here of suitable habitats rather than to this famous

region’s being the plant’s Pleistocene refugium. In Europe, as in

America, the species occupies various highly disjunct locations in

the Caucasus and in Turkey, Italy, Spain and Morocco, indicating

that it is either easily dispersed or that it has survived successfully

as a relict from a cooler, postglacial period. It is important to note

that 1) it grows in clearly glaciated territory in Wisconsin, Minne¬

sota and east-central Iowa [but probably not southwestern Iowa as

shown by Hulten (1970, Map 104)], 2) other subarctic or boreal

species, namely Primula mistassinica ( cf . litis & Shaughnessy

1960) , Rhododendron lapponicum, Chrysoplenium ioense ( C. tetran-

drum) and Gymnocarpium robertianum also occur in the Midwest

in the region of the “Driftless Area,” 3) ferns, primroses and

saxifrages have small propagules, making long-distance, postglacial

dispersal possible and 4) the microenvironments of damp, cool,

rocky habitats would remain relatively little influenced by post¬

glacial vegetational changes occurring in the region, permitting

survival of the periglacial flora.

It is probable, on ecologic as well as geographic grounds, that

Adoxa is an old species whose present-day disjunct range, rather

than being the result of ease of dispersal and pioneer propensities,

is due likely to relictual survival in isolated microhabitats from

a cooler, interglacial or postglacial period. While now thought to

have been glaciated, the “Driftless Area” was never covered com¬

pletely by the later Pleistocene ice sheets. Certainly it was available

for plant occupation since early Wisconsinan time, if mostly over-

lain by ice of the Altonian Substage as believed by Black (1962),

and probably since the Nebraskan Stage. As developed by Butters

and Abbe (1953) and recognized by Cushing (1965), disjunct rari¬

ties, whether boreal (Adoxa), “cordilleran” (Mertensia paniculata) ,

or southern (Spiraea tomentosa var. rosea), should not be cited

(Fassett 1931; Schuster 1958; Curtis 1959) as members of a pre¬

glacial flora that survived continental glaciation in a “Driftless

Area” nunatak, nor should they be used even to imply that the

“Driftless Area” was a preglacial plant refugium.

REFERENCES

BLACK, R. F. 1962. Pleistocene chronology of Wisconsin. Geol. Soc. Amer.

Spec. Pap. 68:137 (abstract).

BROOKS, K. L. 1960. Adoxa and Uvularia in the Delaware County (N. Y.)

flora. Bull. Torrey Bot. Club 87:151-153.

BUTTERS, F. K., and E. C. ABBE. 1953. A floristic study of Cook County,

northeastern Minnesota. Rhodora 55:21-55, 63-101, 116-154, 161-201.

252 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

CRONQUIST, A. 1968. The Evolution and Classification of Flowering Plants.

Houghton Mifflin Co., Boston.

CURTIS, J. T. 1959. The Vegetation of Wisconsin. Univ. of Wisconsin Press,

Madison.

CUSHING, E. J. 1965. Problems in the Quaternary phytogeography of the

Great Lakes Region, in Wright, H. E., and D. G. Frey, eds., The

Quaternary of the United States. Pp. 403-416. Princeton Univ. Press,

Princeton.

FASSETT, N. C. 1931. Notes from the Herbarium of the University of Wis¬

consin — VII. Rhodora 33:224-228.

FERNALD, M. L. 1950. Gray's Manual of Botany, ed. 8. American Book Co.,

New York.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of the

Northeastern United States and Adjacent Canada. Vol. 3. Lancaster

Press, Lancaster, Pa.

HARTLEY, T. G. 1962. The Flora of the Driftless Area. Ph.D. Dissertation,

Univ. Iowa, Iowa City.

HOUSE, H. D. 1924. Annotated List of the Ferns and Flowering Plants of

New York State. New York State Mus. Bull. 254. Univ. State New York,

Albany.

HULTEN, E. 1970. The Circumpolar Plants. II. Dicotyledons. Sv. Vet. Akad.

Handl. Bd. 13, Nr. 1. Almqvist & Wiksell, Stockholm.

ILTIS, H. H., and W. M. SHAUGHNESSY. 1960. Preliminary reports on the

flora of Wisconsin. No. 43. Primulaceae — Primrose Family. Trans. Wis.

Acad. Sci. Arts Lett. 49:113-135.

LAKELA, 0. 1965. A Flora of Northeastern Minnesota. Univ. Minnesota Press,

Minneapolis.

MORLEY, T. 1966. Spring Flora of Minnesota. Printing Dept., Univ. Minne¬

sota, Minneapolis.

SCHUSTER, R. M. 1958. Boreal Hepaticae, a manual of the liverworts of Min¬

nesota and adjacent regions III. Phytogeography. Amer. Midi. Nat. 59:

257-332.

SPRAGUE, T. A. 1927. The morphology and taxonomic position of the Adoxa-

ceae. J. Linn. Soc. (Botany) 47:471-487.

TAYLOR, N. 1913. A plant new to the State of New York and the local flora

range. Torreya 13:78.

WADE, D. R., and D. E. WADE. 1940. Preliminary reports on the flora of

Wisconsin. No. 28. Caprifoliaceae — Honeysuckle Family. Trans. Wis. Acad.

Sci. Arts Lett. 32:91-101.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN

NO. 65. DIPSACACEAE— TEASEL FAMILY

Peter J. Salamun

University Wisconsin — Milwaukee

and

Theodore S. Cochrane

University Wisconsin — Madison

This study of the Dipsacaceae, represented in Wisconsin by four

Eurasian species which occasionally escape from cultivation, is

based on collections deposited in the herbaria of the University of

Wisconsin (WIS), University of Wisconsin — Milwaukee (UWM),

Milwaukee Public Museum (MIL), University of Minnesota

(MIN) and Wisconsin State University-— Oshkosh (WSO). We

thank the curators of these herbaria for loans of specimens, and

we gratefully acknowledge the cooperation of Mr. Floyd Swink of

the Morton Arboretum, Lisle, Illinois, who supplied data for an

additional map record.

Dots on the maps indicate exact locations where specimens have

been collected ; triangles represent county records added from

S wink’s Plants of the Chicago Region (1969). The numbers within

the map corner inserts show the amount of flowering and fruiting

material noted and indicate when the species may be expected to

flower or fruit in Wisconsin. Specimens with vegetative growth

only, buds, or with immature or dispersed fruits are not included.

The nomenclature and descriptive features generally follow Glea¬

son (1952) and Fernald (1950) ; however, more recent taxonomic

treatments of certain taxa are discussed in the text or cited in the

bibliography.

Dipsacaceae A. L. De Jussieu Teasel Family

Annual, biennial or perennial herbs with opposite or whorled,

exstipulate leaves and inflorescences of dense involucrate heads or

rarely short spikes. Flowers perfect, epigynous, more or less irreg¬

ular, subtended by a calyx-like involucel (epicalyx) which closely

subtends the ovary and often also a receptacular bract. Calyx min¬

ute, cupulate, a 4-5-toothed or divided into 5-10 conspicuous pap¬

pus-like segments. Corolla tubular, 4-5-lobed. Stamens (2-3) 4,

distinct, epipetalous near the base of the corolla tube and alternate

with its lobes, exserted, the anthers 2-celled, versatile, dehiscing

longitudinally. Ovary inferior, 1-celled, 2-carpellate but only one

253

254 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

developing-, with a solitary pendulous ovule; style filiform, the

stigma simple or 2-lobed, Fruit an achene enclosed within the

gamophyllous involucel and often crowned by the persistent calyx.

Embryo straight, the endosperm fleshy.

An Old World family of 8-12 genera and 250 species, mostly

Mediterranean, distinguished from the related Valerianaceae and

Caprifoliaceae by the unilocular ovary and the capitate inflo¬

rescence surrounded by an involucre of bracts, and from the Com-

positae by the distinct, exserted stamens, the pendulous ovule and

the fruit enclosed by an involucel. The “heads” are cymose aggrega¬

tions of 1-flowered heads whose flowers open first near the center

of the inflorescence, then simultaneously toward both the apex and

the base.

KEY TO GENERA

A. Plants prickly; heads ovoid to subcylindric, the involucral

bracts upcurved, and spine-tipped, 2-15 cm long; corollas all

alike; calyx 4-toothed or -lobed. _ 1. DIPSACUS.

AA. Plants not prickly; heads convex, hemispheric or subglobose,

the involucral bracts ascending to reflexed, not spine-tipped,

0.5-1. 5 cm long; corollas of marginal flowers expanded and

ray-like.

B. Receptacle with chaffy bracts; involucral bracts rigid

and brown, imbricate in several rows ; corollas cream to

yellow; calyx teeth ca. 25. _ 2. CEPHALARIA.

BB. Receptacle with dense hairs ; involucral bracts herbace¬

ous and green, in 1 or 2 definite rows; corollas lilac-

purple; calyx awns 8-16. _ 3. KNAUTIA.

1. DIPSACUS L. Teasel

Tall biennial or perennial herbs with stout prickly or rough-hairy

stems from taproots and opposite, sessile or connate, toothed or

pinnatifid leaves. Heads dense, ovoid to cylindric, with rigid, long-

tapering receptacular bracts and spine-tipped involucral bracts.

Flowers 8-15 mm long, the marginal ones not enlarged, subtended

by U- angled , obscurely k- toothed calyx-like involucels. Calyx tube

4-angled, adherent to the ovary, the limb cupulate, ^-toothed or

-lobed, ciliate but without appendages. Corolla funnelform, nearly

regularly 4-lobed, with whitish tube and pale purple lobes. Stamens

4. Ovary 1-celled ; style filiform, the stigma oblique, entire. Achenes

8-ribbed, enclosed by the involucel.

Native to Eurasia and North and East Africa, with 3 of the 12

species adventive or naturalized in North America.

1974] Salamun and Cochrane — Teasel Family Report 255

KEY TO SPECIES

A. Cauline leaves lanceolate to oblong-lanceolate, toothed and

often prickly on the margins, becoming entire upward and

commonly connate at the base. _ 1. D. FULLONUM.

AA. Cauline leaves pinnatifid or bipinnatifid, the margins more or

less bristly-ciliate, their bases confluent and forming a cup.

_ 2. D. LACINIATUS.

1. DlPSACUS FULLONUM L. Map 1.

Wild Teasel, Common Teasel

Dipsacus sylvestris Hudson of American authors

Dipsacus fullonum L. ssp. sylvestris Claph.

Stout biennial 0.5-2 (3) m tall, the stem striate-angled, prickly.

Basal leaves oblong to oblanceolate, arcuate, usually dying early in

the second season. Cauline leaves sessile, lanceolate to oblong-

lanceolate, to 3 (4) dm long, 4-10 cm wide, crenate-serrate, often

prickly -dentate on the margins and the midrib beneath, becoming

entire upward and commonly connate at the base . Heads ovoid to

subcylindric, 3-10 cm high, 3-5 cm wide, on long naked peduncles;

receptacular bracts oblong, abruptly tapering to stiff straight

barbed aivns exceeding the flowers; involucral bracts linear-

elongate, subulate, upcurved, some of them equaling or exceeding

the heads, 2-15 cm long, 2-5 mm wide. Calyx cup 1 mm high, silky-

pubescent. Corolla 8-12 mm long, the tube pilose, lavender (or

rarely white in forma albidus Steyerm.) . Achenes 4-7 mm long.

2 n - 16, 18.

Native to Europe, North Africa, and the Near East, now widely

naturalized throughout the United States and southern Canada, in

Wisconsin occasionally cultivated for ornament and adventive or

locally established along open roadsides and abandoned railroad

embankments, in fallow fields, low pastures, such as along Willow

Creek, Richland Co., with Verbena has tat a, Cirsium vulgar e and

Helenium autumnale ( Nee 2681, 27US, WIS), and persisting in

trash dumps and waste heaps near cemeteries and greenhouses.

Flowering July to September; fruiting September to October.

Fuller’s Teasel, Dipsacus sativus (L.) Honckney (D. fullonum

L. of many authors), differs from the above species in having

slightly stouter and strongly recurved receptacular bracts nearly

equaling the flowers, spreading involucral bracts not surpassing

the head and corollas 10-14 mm long. Grown for the “teasels” or

mature heads which are used in raising the nap (fulling) on

woolen cloth, it occasionally escapes in the vicinity of textile mills.

However, it has not been collected in Wisconsin. Several authors

256 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

have implied that the cultivated teasel with the recurved recep-

tacular bracts has been derived from the wild D. fullonum with

erect bracts. Hegi (1908) suggested instead that the cultivar was

selected originally from D. ferox Lois., a wild species of France,

Italy and Spain. The nomenclature of these two taxa follows that

adopted by Ferguson and Brizicky (1965).

2. Dipsacus laciniatus L. Map 2, dots.

Cut-leaved Teasel

Much resembling D. fullonum , but leaves irregularly and coarsely

pinnately -cleft, the margins bristly-ciliate, and the bases of the

paired cauline leaves confluent to form prominent cups. Involucral

1974] Salamun and Cochrane — Teasel Family Report 257

bracts lanceolate to linear-lanceolate, 2-8 cm long, 3-7 mm wide,

6-12 times as long as wide, shorter than to barely exceeding the

mature head . 2 n — 16, 18.

Native to southern Europe, southwestern Russia and Persia,

rarely adventive in North America, rare in south- and east-central

Wisconsin along roadsides, in disturbed marshes and in cemeteries

and dumps. This species, much rarer than the last, has been previ¬

ously reported in the state only from Rock Co. (Musselman et ah,

p. 187. 1971). The specimen cited there is Dipsacus fullonum, but

other authentic collections have been seen from Rock and several

other counties : Dane Co. : Madison, waste heap, Holy Cross Ceme¬

tery, Franklin Ave. and Regent St. ( Bergseng s.n., 30 Jul 1948,

WIS). Fond du Lac Co.: marsh S of Soo Line RR, sec. 6, T16N,

R17E (Furstenberg 151, 28 Nov 1969, WSO). Milwaukee Co.: Mil¬

waukee, 2 plants in field, Wanderer's Rest Cemetery ( Niss s.n., 27

Sep 1969, WIS). Rock Co. : roadside, Hwy. H, 1 mi. E of Indian-

ford ( Greene s.n., 31 Jul 1947, WIS) ; same location ( Curtis ,

Greene, and Sauer 1672, 2 Jul 1954, WIS). Portage Co.: sandy

waste area beside town dump, sec. 26, T25N, R10E ( Redmond UUl,

12 Aug 1971, WSO). Flowering late July to late September; fruit¬

ing dates probably the same as those for D. fullonum.

2. CEPHALARIA Schrad.

Annual or perennial herbs with simple or pinnately divided

leaves and flowers in globose heads on long peduncles. Involucral

bracts numerous, imbricate in several rows, not spine-tipped ,

shorter than the heads. Receptacular bracts more or less sharp-

pointed but not spine-tipped, shorter than the flowers. Involucel

4-angled, bearing U or 8 awn-like appendages at the apex. Calyx

limb shallowly cupulate, denticulate. Flowers similar to those of

Knautia, the corollas 4-lobed, the marginal ones sometimes enlarged

and ray-like. Fruit, 4-8-ribbed, prismatic and somewhat fusiform.

Native to the Mediterranean region, Central Asia and South

Africa, a few of the 60 species sometimes cultivated.

1. CEPHALARIA TATARICA Schrad. Map 2, circle.

Coarse perennial to 2 m tall, the stems bearing 3-5 heads,

retrorsepubescent and leafy below, glabrous and nearly naked

above. Cauline leaves deeply pinnately divided, the divisions decur¬

rent, elliptic and coarsely serrate to linear and entire. Heads 2.5-

4.5 cm wide, on peduncles 6-30 cm long. Corollas cream or yelloiv,

8-16 mm long, the marginal ones enlarged and ray-like. Involucral

258 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

bracts ovate, 5-10 mm long, rigid and brown, grading into oblance-

olate, abruptly stiff-acuminate receptacular bracts, these 8-12 mm

long, about equaling the corolla tube. Calyx teeth numerous, ca. 25,

0.5-1 mm long, densely villous. Achenes 6-9 mm long, pubescent,

with 8 ribs , these prolonged as subequal short awns (involucel

“teeth”) 0.7-1. 5 mm long. 2 n = 36.

Native to western Asia, possibly southern Russia, rarely culti¬

vated in flower gardens ; collected twice in Wisconsin and perhaps

reported here as an escape for the first time from the United

States : — Winnebago Co. : N edge of Oshkosh, Algoma Blvd., road¬

side adjacent plowed field near cemetery, sec. 10, T18N, R16E

( Harriman 3600, 18 Jul 1968 [fl, fr], WSO, fragment WIS) ;

spreading along road shoulder, same location ( Harriman 9209, 10

Jul 1973 [fl], WIS, WSO). The collector states that the plant is

“as well established as, say, Sonchus or Cirsiumf 9

3. Knautia L. Bluebuttons

Annual or perennial herbs with erect stems, pinnatifid leaves

and hemispheric to subglobose heads on elongate peduncles. Invol-

ucral bracts lanceolate, herbaceous, about equaling the heads. Re¬

ceptacle more or less densely hairy, without bracts . Corollas fun-

nelform to narrowly campanulate, the limbs more or less oblique

or bilabiate, 4-5-lobed, those of the marginal flowers often much

enlarged. Involucel strongly compressed, If -angled, the limb very

short, with only a few minute teeth at the summit. Calyx short,

with 8-16 elongate setaceous appendages. Stigma emarginate.

Native to Eurasia and North Africa, with 40 species.

1. Knautia arvensis (L.) Coult. Map 3.

Bluebuttons, Bluecaps

Scabiosa arvensis L.

Hirsute perennial 4-10 dm tall. Basal leaves oblanceolate, simple

or lyrate-pinnatifid. Lower cauline leaves usually only coarsely

toothed, the others deeply pinnatifid into 5-15 linear-oblong seg¬

ments, reduced upward. Peduncles 4-25 cm long. Heads 1.5-4 cm

wide, the outer involucral bracts ovate, the inner narrowly lance¬

olate, 8-16 mm long, herbaceous, about equaling the heads. Corolla

8-18 mm long, lilac-purple. Achenes Ip-ribbed, 5-6 mm long ,

strongly compressed-ellipsoid, pubescent, the apex truncate, min¬

utely denticulate, crowned by the calyx limb, its 8-16 awns 1.7-3. 6

mm long, eventually deciduous. 2 n — 20, 40.

1974] Salamun and Cochrane — Teasel Family Report 259

Native throughout Europe except for the extreme North ; in Wis¬

consin a sporadic escape from flower gardens, becoming locally

abundant in nearby fields, roadsides and waste places but possibly

not persisting. First reported here for the state, although some of

the following collections are old : — Ashland Co. : common along

roadsides 2 mi. SW of Glidden ( Courtenay s.n., 15 Jul 1962, MIL,

WIS). Forest Co.: one colony by roadside, with bracken, timothy

and dogbane, 1/2 mi. N of Hwy. 52 on Hwy. W, E side of Richard¬

son Lake ( Kruschke K-65-6U, 15 Jul 1965, MIL). Milwaukee Co.:

Whitefish Bay, edge of field ( Sorenson s.n., 20 Jun 1912, MIL) ;

Milwaukee, a garden escape, vacant lot on Hopkins St. % block S

of Villard Ave. ( Fuller F-U2-8U, 15 Jul 1942, MIL). Sawyer Co.:

colony extending ca. 50 yds. along gravelly roadside through Picea

glauca plantation, Forest Rd. 162 ca. 1 mi. N of Hwy. 70, sec. 5,

T39N, R3W ( Hansen 19Jf3, 5 Jul 1973, WIS). Fuller’s specimen

may have sprouted from garden refuse, since Malva alcea was col¬

lected at the same site (cf. Utech 1970). Flowering June to

September.

Many species of Scabiosa, especially S. atropurpurea L., Sweet

Scabious, are popular garden ornamentals that rarely escape from

cultivation. The features distinguishing them from Knautia include

the lanceolate involucral bracts, the 5 calyx setae, the chaffy recep¬

tacle, the cupulate involucel limbs, and the terete 8-fur rowed fruits.

None have been collected in Wisconsin.

BIBLIOGRAPHY

EHRENDORFER, F. 1962. Beitrage zur Phylogenie der Gattung Knautia

( Dipsacaceae ), I. Cytologische Grundlagen und allgemeine Hinweise.

Osterr. Bot. Zeitschr. 109:276-343.

- . 1963. Cytologie, Taxonomie und Evolution bei Samenpflanzen, in

Turrill, W. B., ed., Vistas in Botany 4:121-129. Pergamon Press, New

York.

• - . 1965. Evolution and karyotype differentiation in a family of flower¬

ing plants: Dipsacaceae, in Geerts, S. J., ed., Genetics Today , Proc. XI

Intern. Genet. Congr. 2:399-407. Pergamon Press, New York.

FERGUSON, I. K. 1965. The genera of Valerianaeeae and Dipsacaceae in the

southeastern United States. Jour. Arnold Arb. 46:218-231.

- and G. K. Brizicky. 1965. Nomenclatorial notes on Dipsacus fullonum

and Dipsacus saticus. Jour. Arnold Arb. 46:362-365.

FERNALD, M. L. 1950. Gray's Manual of Botany, ed. 8. American Book Co.,

New York.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of the

Northeastern United States and Adjacent Canada. Vol. 3. Lancaster

Press, Lancaster, Pa.

HEGI, G. 1908. Illustrierte Flora von Mittel-Europa. Vol. 6, I. J. F. Lehmann

Verlag, Miinchen.

MUSSELMAN, L. J., T. S. COCHRANE, W. E. RICE, and M. M. RICE.

1971. The flora of Rock County, Wisconsin. Mich. Bot. 10:147-193.

PARS A, A. 1943. Flore de L'lran. Vol. 3. Mazaheri Press, Teheran.

260 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

POHL, R. W., and E. P. SYLWESTER. 1962. Dipsacus in Iowa. Ia. Acad. Sci.

70:53, 54.

STEYERMARK, J. A. 1958. An albino form of Dipsacus sylvestris. Rhodora

60:174, 175.

SWINK, F. 1969. Plants of the Chicago Region. The Morton Arboretum, Lisle,

Illinois.

UTECH, F. H. 1970. Preliminary reports on the flora of Wisconsin. No. 60.

Tiliaceae and Malvaceae — Basswood and Mallow Families. Trans. Wis.

Acad. Sci. Arts and Lett. 58:301-323.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN

NO. 66. CYPERACEAE II— SEDGE FAMILY II.

The Genus Cyperus— The Umbrella Sedges

Brian G. Marcks

University Wisconsin — Madison

Cyperus , a vast genus of upwards of 700 species chiefly

distributed in warm temperate and tropical regions of the world,

is distinguished from other Cyperaceae by having, in combination,

strictly 2-ranked (distichous) scales and terminal umbellate in¬

florescences or heads. Since the group is technically difficult, exam¬

ples of inflorescences and their parts have been illustrated in Fig. 1

to aid in the understanding of some of the terminology used in this

report. The Wisconsin species flower from midsummer to early

fall and occur in dry exposed sandy areas or in various wetland

habitats, such as marshes and bogs, swales and low fields, ditches,

and lakeshores, riverbanks and streamsides. Several of our species

are notable for their widespread distribution. Cyperus aristatus,

C, esculentus and C. odoratus are semi-cosmopolitan in tropical and

temperate regions, while C. Engelmannii is widely distributed in

the New World. Cyperus strigosus and C. esculentus tend to become

weedy in low fields. Phenotypic dwarfing is common and particu¬

larly noticeable in depauperate specimens of the taller wetland

species, including C. strigosus, C. erythrorhizos, C. Engelmannii

and C. odoratus.

The present paper revises Greene’s (1953) preliminary partial

treatment of Cyperaceae in Wisconsin, including the genus Cyperus.

Distribution maps are based on specimens in the herbaria of the

University of Wisconsin, Madison (WIS), University of Wisconsin

— Milwaukee (UWM), Milwaukee Public Museum (MIL), Uni¬

versity of Minnesota (MIN), Field Museum of Natural History

(F), Wisconsin State University — Oshkosh (WSO), Wisconsin

State University — Stevens Point (WSP), and Northland College,

Ashland, Wisconsin (NC). Grateful acknowledgement is due to

the curators of the above herbaria for loans of specimens. The

results of my master’s and doctoral dissertations, population studies

of Cyperus section LAXIGLUMI in the United States (Marcks,

1967, 1972) have been freely incorporated.

Map dots represent exact locations. The map inset numbers

record flowering and fruiting dates as determined from specimens

housed at the University of Wisconsin, Madison (WIS). Plants in

261

262 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

A. Inflorescence

A. Spikelet

FIGURE 1. Inflorescences, spikelets, and achenes of Wisconsin Umbrella

Sedges labelled to illustrate parts and diagnostic characters: A) Cyperus

Schweinitzii; b) C. lupulinus ssp. macilentus.

vegetative growth only, in bud, or with dispersed fruit were not

included. Triangles in Iowa and Michigan represent county records

according to Gilly (1946) and Voss (1972). Most common names

are those used by Gilly (1946).

/

1. CYPERUS L. Umbrella Sedge, Flat Sedge, Nut-grass,

Galingale

Mostly low, cespitose annuals or loose-tufted perennials from hard

rhizomes or corm-like tubers ; culms 3-angled, solid, simple, usually

slender and erect, leafy near the base and bearing a leafy involucre

at the summit. LEAF blades conduplicate or flat, erect or spreading-

arcuate, usually scabrellate on the margins and dorsal midrib,

sometimes scaberulous on the ventral surface toward the apex ; leaf

sheaths glabrous, unpigmented or more commonly tinged reddish-

purple, hyaline ventrally, becoming fibrous-shredded. Involucral

1974]

Marcks — Sedge Family II Report

263

bracts blade-like, 3-several of unequal lengths, ascending, spreading

or reflexed. INFLORESCENCE terminal, an umbel-like aggrega¬

tion of simple or branched primary spikes (rays) with the central

spikes usually sessile or subsessile and the lateral spikes peduncled,

or the whole inflorescence contracted into a dense head. Peduncles

usually of unequal lengths, ascending or variously divergent, slen¬

der and smooth, the lower portion of each enveloped by a tubular

sheath (prophyll). SPIKELETS spirally arranged on the spike

axis (rachis), subremote to densely congested, divaricate, ascend¬

ing, radiate or digitate, ovate to oblong-linear, usually laterally

compressed, persistent or disarticulating at maturity ; rachilla

straight or zigzag, often winged on the lateral margins. SCALES

strictly 2-ranked, imbricate or subapproximate, persistent or de¬

ciduous, green-keeled. Flowers 3-many ; stamens 1-3 ; styles 2-3-

cleft. ACHENES lenticular or trigonous, beakless.

Key to Species

A. Achenes lenticular; styles 2-cleft; low annuals with strongly

flattened lustrous purplish-brown spikelets (Subgenus Pycreus,

Section SULCATI).

B. Styles exserted about 2 mm, persistent; scales loosely

imbricate, the laminae with a depressed membranaceous

dull patch surrounded by a broad lustrous purplish-brown

margin. _ _ 1. C. DIANDRUS.

BB. Styles exserted less than 1 mm, deciduous; scales closely

imbricate, the laminae firm, slightly convex, the surface

lustrous and purplish-brown throughout (rarely unpig-

mented). _ 2. C. RIVULARIS.

AA. Achenes trigonous; styles 3-cleft; habits various (Subgenera

Cyperus , Mariscus, Torulinium) .

C. Scales ending in a long recurved excurrent tip; strongly

aromatic dwarf annuals rarely more than 10 cm tall

(Section AMBILES) _ 3. C. ARISTATUS.

CC. Scales ending bluntly or in a straight or appressed tip;

nonaromatic annuals or perennials usually 10-100 cm tall.

D. Spikelets mostly divaricate at nearly right angles

(or slightly ascending) from a ± elongate rachis,

with only the uppermost and lowest spikelets

strongly ascending or deflexed ; rachilla winged along

lateral margins.

E. Scales 1.2-1. 5 mm long, densely aggregated and

tinged lustrous copper ; achenes ivory or pearly

white, 0. 8-1.0 mm long (Section FASTI-

GIATA) _ 4. C. ERYTHRORHIZOS.

264 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

EE. Scales 1.5-4. 5 mm long, variously colored;

achenes darker, 1.0-2. 2 mm long.

F. Spikelets flat or strongly compressed;

scales 3. 5-4.5 mm long, the laminae uni¬

formly yellow-tinged ; achene less than

half the scale length (Section STRIGOSI).

_ 5. C. STRIGOSUS.

FF. Spikelets not strongly flattened; scales

1. 5-3.0 mm long, the laminae golden-

brown to reddish-brown ; achene over half

the scale length, or female sterile.

G. Perennials bearing stolons with prom¬

inent membranaceous scales, each

terminated by a hard tuber; rachilla

not breaking into segments (Section

ESCULENTI). _

_ 6. C. ESCULENTUS.

GG. Annuals from fibrous roots; rachilla

breaking into short segments at ma¬

turity (Section FERACES).

H. Apex of scales overlapping su¬

perjacent scales. _

_ 7. C. ODORATUS.

HH. Apex of scales not overlapping

superjacent scales, giving the

spikelet a zigzag appearance.

_ 8. C. ENGELMANNII.

DD. Spikelets radiate to subdigitate from a short rachis

or ascending from an elongate rachis ; rachilla essen¬

tially wingless (Section LAXIGLUMI).

I. Achenes about twice as long as wide, the faces

flat or only slightly concave.

J. Inflorescence a single, sessile, dense hemi¬

spherical to nearly spherical head (rarely

with 1-2 additional pedunculate spikes) ; in-

volucral bracts commonly deflexed ; culms

smooth _ 9. C. LUPU LINUS.1

JJ. Inflorescence an open umbel with 1-8 pedun¬

culate spikes; involucral bracts ascending;

culms markedly scabrous on angles near the

apex. _ 10. C. SCHWEINITZII1

1 Hybrids between species 9 and 10 are common in Wisconsin (cf. Hybridization in

the G. lupulinus-C. Schweinitzii complex, p. 276).

1974]

Marchs — Sedge Family II Report

265

II. Achenes much less than twice as long as wide,

the faces markedly concave. _

_ 11. C. HOUGHTON II.1

1. Cyperus diandrus Torr. Map 1.

Low Cyperus

Rather soft, tufted or solitary low annuals 3-24 (-35) cm tall.

Culms rather weak, 2-26 cm long, smooth, the tufted forms some¬

times with secondary cushion-like inflorescences directly from the

base. Blades 1-3 mm wide, with membranaceous loose-fitting

sheaths enveloping the lower % of the culm; involucral bracts

blade-like, mostly 3, divergent. Inflorescence a subglobose cluster

of loose spikes, usually with 1-5 pedunculate rays up to 6 cm long

in addition. Spikelets subdigitate to pinnate from a rachis to 12

mm long, lance-oblong, flat, 5-40-flowered, 4-26 mm long; rachilla

persistent. Scales loosely imbricate, the achenes usually visible un¬

der the margins, 2. 0-2. 7 mm long, deciduous, nerved only on the

green keel, the tip appressed ; laminae with a dull depressed scari-

ous patch banded by a wide lustrous purplish-brown margin.

Stamens 2. Style 2-cleft nearly to the base, long-exserted (to ca 2

mm) beyond the scale, persistent. Achene lenticular, ellipsoid to

narrowly ovoid, ca 1.2 mm long, whitish to reddish-brown.

A wetland species ranging from North Dakota and Missouri

east to Quebec and Virginia, infrequent in Wisconsin on wet sandy

to muddy shores and banks of rivers, streams, and lakes, some¬

times in peaty soil of sedge mats, becoming rare northward ; often

confused with C. rivularis, but distinguished by its long-exserted,

persistent styles and the depressed scarious patch on each scale

lamina. Flowering from July to October; fruiting from late July

to late October.

2. Cyperus rivularis Kunth Map 2.

Shining or Brook Cyperus

Mostly low tufted annuals 3-25 (-32) cm tall, erect or spread¬

ing. Culms firm, smooth, 2-24 cm long, the plants often with sec¬

ondary cushion-like inflorescences directly from the base. Blades

0.5-2.0 mm wide, with rather tight-fitting sheaths enveloping less

than y3 of the culm; involucral bracts blade-like, 1-3, divergent.

Inflorescence a terminal umbel composed of a subglobose cluster

of loose spikes, with an additional 1-5 rays up to 6 cm long.

Spikelets subdigitate to pinnate from a rachis to 1 cm long, lance-

oblong, flat, 6-38-flowered, 4-23 mm long; rachilla persistent.

Scales closely imbricate, completely covering the achenes, 1. 8-2.2

266 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

mm long, nerved only on the green keel, the tip appressed ; laminae

firm, slightly convex, lustrous purplish-brown (rarely unpig-

mented). Stamens 2. Style 2-cleft to about the middle, exserted less

than 1 mm, deciduous . Achene lenticular, ellipsoid to narrowly

obovoid, whitish to reddish-brown, ca 1.2 mm long.

A common wetland species of wide distribution in temperate

North America, in Wisconsin in wet sand and mud of shores and

banks of lakes, rivers and streams, and in low fields, ditches and

marshes; easily confused with C. diandrus, but distinguished by

firm, closely imbricate scales with lustrous purplish-brown lami¬

nae and the absence of long-exserted, persistent styles. Flowering

from early July to early October; fruiting from August to mid-

October.

3. Cyperus aristatus Rottb. Map 3.

Awned Cyperus

C. inflexus Muhl.

C. aristatus var. inflexus (Muhl.) Kiikenth.

Mostly tufted, soft-based low delicate annuals 2-18 cm high,

from red fibrous roots, strongly aromatic, with the fragrance of

Ulmus rubra or Gnaphalium obtusifolium. Culms slender, flaccid,

smooth, sharply wing-angled. Blades 1-3 mm wide, from deep

purplish-red sheaths; involucral bracts blade-like, mostly 3, diver¬

gent. Inflorescence a dense hemispherical to subglobose head,

usually with 1-3 additional rays. Spikelets subdigitate from a short

rachis, somewhat flattened, 3-23-flowered, 2-9 mm long; rachilla

persistent. Scales 1.5-2. 5 mm long, the keel green, narrowed to a

long recurved excurrent tip; laminae 3-5-nerved, pale yellow to

rich copper. Stamen 1. Style 3-cleft. Achene trigonous, slenderly

obovoid, ca 0.8 mm long, pale brown.

Semi-cosmopolitan species of north temperate and tropical re¬

gions, occurring in the western two-thirds of Wisconsin in wet

sand and mud of river bars and banks, lakeshores, moist depres¬

sions and temporary pools. Flowering from July to October;

fruiting from mid-July to mid-October (n = ca 24, 28, 32; Murdy,

1968).

4. Cyperus erythrorhizos Muhl. Map 4.

Red-rooted Cyperus

Annuals from fibrous, caked blood-red roots . Plants erect and

solitary, 1-4 (-8) dm tall, or spreading from loose tufts, sometimes

dwarfed and only to 2 cm tall. Culms firm, smooth, coarsely

1974]

Marcks — Sedge Family II Report

267

268 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

3-angled. Blades soft, scabrous apically, pale beneath, 2-6 mm wide ;

involucral bracts blade-like, 3-9, spreading. Inflorescence a com¬

pound umbel, rarely simple or condensed, with a cluster of subses-

sile spikes and in addition 1-7 unequal divergent pedunculate rays

1-10 cm long. Spikelets crowded, mostly radiating horizontally

from an elongate rachis 5-30 mm long, slenderly oblong, 5-30-

fiowered, 3-20 mm long, ca 1 mm wide ; rachilla persistent, bearing

deciduous chaffy wings. Scales small and delicate, 1.2-1. 5 mm long,

densely aggregated ; keel broad and green ; laminae lustrous copper

to rich reddish-brown. Achene plano-convex, short ellipsoid to

ovoid, ca 0.8 mm long, lustrous ivory to pearly white.

Widely distributed wetland species of temperate North America,

in Wisconsin common in the southwest, absent in the northeast,

on wet sandy or muddy shores of lakes, rivers and streams and in

marshes and wet ditches. Flowering from mid- July to October;

fruiting from August to early November.

5. Cyperus strigosus L. Map 5.

Straw-colored Cyperus

Erect perennials 1-6 (-10) dm tall, the base becoming a hard

corm-like rhizome. Culms firm, smooth, acutely 3-angled. Blades 2-6

mm wide, flat, apically scabrous; involucral bracts blade-like, 3-8,

divergent, often greatly exceeding the umbel. Inflorescence a simple

or compound umbel with a cluster of subsessile spikes and 1-12

unequal, obliquely ascending pedunculate rays to 12 cm long.

Spikelets usually crowded, mostly radiating horizontally from a

± elongate rachis 4-28 mm long, linear-lanceolate, strongly flat¬

tened, 4-14-flowered, 5-18 mm long, ca 2 mm wide; rachilla decidu¬

ous, winged. Scales oblong-lanceolate, 3.5-A.5 mm long, loosely

imbricate, the keel green; laminae uniformly tinged yellow, the

margins spreading. Achene less than half the length of the scale,

trigonous and slender, 1. 6-2.2 mm x 0.5-0. 6 mm, purplish-brown.

A widespread and variable North American species, very com¬

mon in all but northeast Wisconsin, in wet sand and mud of

meadows, swales and sloughs, temporary pools and ditches, and

shores and banks of lakes, rivers and streams; sometimes weedy

in low fields. Flowering from July to October ; fruiting from August

to mid-October.

6. Cyperus esculentus L. Map 6.

Yellow Nut-grass, Chufa

Robust erect perennials from thickened bases, bearing soft

stolons with prominent scales, each stolon terminated by a hard

1974]

Marcks — Sedge Family II Report

269

tuber. Culms 2-9 dm high, acutely 3-angled, smooth and sub-

coriaceous at base, soft and scabrous at the apex ; involucral

bracts blade-like, 2-9, exceeding the umbel, spreading. Inflorescence

an open and often compound umbel of several subsessile spikes,

with several additional ascending pedunculate rays 1-10 dm long.

Spikelets mostly 4-ranked along an elongate rachis 1-3 cm long,

loosely aggregated and spreading nearly horizontally, barely flat¬

tened , 5-25 mm long, 5-28-flowered ; rachilla persistent, narrowly

winged. Scales overlapping , 2-3 mm long, yellowish-brown to golden

brown. Stamens 3, 1.2-1. 5 mm long. Achene trigonous, ellipsoid

to obovoid, rounded at the summit, 1.2-1. 6 mm long (sometimes

female sterile), lustrous tan to golden brown.

A widely distributed pantropical and warm temperate weed,

sparingly throughout Wisconsin in wet sand or mud of lakeshores

and riverbanks, and as a weed in low fields and roadsides. Flower¬

ing from July to October; fruiting from mid- July to mid-October

(2n = 108; Hicks, 1929; Darlington and Wylie, 1955).

7. Cyperus odoratus L. Map 7.

Coarse Cyperus

C. ferruginescens Boeekh

C. speciosus var. squarrosus Britt.

C. speciosus var. ferruginescens (Boeckl.) Britt.

C. ferax ssp. speciosus (Britt.) Kiikenth.

C. odoratus var. squarrosus (Britt.) Gilly

Fibrous-rooted annuals 2-60 cm tall, erect or dwarfed, spread¬

ing from loose tufts. Culms several, smooth, firm, acutely 3-angled.

Blades flat, apically scabrous; involucral bracts blade-like, 3-6,

spreading, sometimes greatly exceeding the umbel. Inflorescence

a compound umbel (or condensed into a loose head-like cluster),

with several loose subsessile spikes and up to 7 unequal divergent

pedunculate rays 1-6 cm long. Spikelets closely aggregated along

a ± elongate rachis 5-20 mm long, mostly spreading horizontally

or slightly ascending, slender and subterete, 4-20-flowered, 4-26

mm long; rachilla breaking into 1 -fruited sections at maturity,

winged. Scale with apex overlapping superjacent scale, 1. 5-2.0 mm

long, the keel green; laminae reddish-brown. Stamens 3, 0.3-0. 5

mm long. Achene trigonous-oblongoid, 1-1.5 mm long, ferruginous

to golden brown.

The entity from the interior of temperate North America,

formerly treated as C. ferruginescens Boeckeler but inseparable

from the semi-cosmopolitan tropical and subtropical C. odoratus

(O’Neill, 1942), is of infrequent occurrence in Wisconsin south of

270 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

y/ jr

CYPERUS ENGELMANNI

1974]

Marcks — Sedge Family II Report

271

the Tension Zone, in wet sand and mud of riverbanks, flats, sloughs,

lakeshores and marshes. The Wisconsin plants appear to belong to

var. squarrosus, which according to Gilly (1946) “is essentially

restricted to the Mississippi Valley, the Great Lakes area, and the

Western Gulf Coastal Plain. . . [as an] . . . interior and western

phase of the species, with the thinner, dull-brown to greenish

scales. . .” Often confused with C. Engelmannii and C. esculentus,

it is distinguished from the former by its subapproximate, over¬

lapping scales and from the latter by its lack of scaly stolons and

tubers and its much shorter stamens. Flowering from July to

September; fruiting from August to October.

8. Cyperus Engelmannii Steud. Map 8.

Engelmann’s Cyperus

Fibrous-rooted annuals 2-60 cm tall, erect or dwarfed and spread¬

ing from loose tufts. Culms smooth and firm, sharply 3-angled.

Blades flat, scaberulous apically, the sheaths light reddish-purple;

involucral bracts blade-like, 3-8, spreading, often greatly exceeding

the umbel. Inflorescence a compound umbel (or condensed into a

loose head-like cluster) with several subsessile spikes and up to

7 unequal divergent pedunculate rays 1-6 cm long in addition.

Spikelets closely aggregated along a ± elongate rachis 4-15 mm

long, mostly spreading horizontally or slightly ascending, slenderly

subterete, 6-18-flowered, 8-24 mm long; rachilla splitting into

1 -fruited sections at maturity, winged. Scales thin, remote and not

overlapping the superjacent scale (giving the spikelet a zigzag

appearance) , 1. 5-3.0 mm long; keel green; laminae golden brown

to reddish-brown. Achene trigonous, linear-oblongoid, ca 1.5 mm

long, brownish.

Widely distributed wetland species of tropical and warm tem¬

perate regions of the New World, in Wisconsin mostly in the north¬

west and southeast, in wet sand or mud of lakeshores, ditches and

marshes. Flowering from late July to early October; fruiting from

early August to mid-October.

9. Cyperus LUPULINUS (Spreng.) Marcks, comb. nov.

Slender stemmed Cyperus

Scirpus lupulinus Spreng. Mant. 1 :30. 1807. TYPE : Muhlenberg

from eastern Pennsylvania, in the Berlin Herbarium (B), photo¬

graph (WIS !) . The type sheet is a mixed collection; the plant on

the left is taken as the lectotype, since the plant on the right ap¬

pears to be Cyperus lupulinus ssp. macilentus.

Cyperus filiculmis of many authors, not of Vahl, 1806.

272 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Erect perennials 8-40 cm high, mostly in loose tufts from hard

subglobose corm-like rhizomes. Culms very slender, rigid and erect,

3-angled, 0.4-1. 2 mm thick near the apex. Blades pale green, nar¬

row, 1-3.5 mm wide, mostly conduplicate, scabrous-margined ; in-

volucral bracts 3-4, blade-like, mostly deflexed. Inflorescence a

single, sessile, dense hemispherical to nearly spherical head, 8-25

mm in diameter, rarely with 1-2 additional rays. Spikelets radiate

and densely congested, from S-U very short axes, 3-22-flowered,

3-22 mm long, 2. 5-4.0 mm wide; rachilla late-deciduous, essen¬

tially wingless. Scales oblong, 1.8-3. 8 mm long, readily deciduous;

keel green, short-mucronate ; laminae unpigmented or tinged red¬

dish-brown. Achene trigonous, narrowly ovoid to oblongoid, 1.4-

2.2 mm X 0.6-1. 1 mm, dark brown or black.

Cyperus lupulinus is usually included in C. filiculmis Vahl, an

endemic of the southern Atlantic and Gulf Coastal Plains of the

United States, but the two are distinguishable by several morpho¬

logical characters. The former has green rather than yellow scales,

wider achenes, and longer scale mucros. Moreover, biosystematic

studies (Marcks, 1972) have shown C. lupulinus to be essentially

allopatric in more northern and western regions and to have a

different chromosome number and phenolic profile pattern. These

data strongly support recognition of C . lupulinus as a species

separate from C. filiculmis.

The first legitimate epithet for this taxon is Scirpus lupulinus

Sprengel, dating from 1807. It is based on Muhlenberg 203 from

Pennsylvania (now in the Willdenow Herbarium in Berlin, num¬

ber 1221). A photograph of this sheet (WIS!) shows it to be

annotated in Sprengel’s hand as “Scirpus lupulinus n. s.” The sheet

bears two specimens, the left-hand one clearly referable to the

typical form of the species in the length of its scales (2. 8-3.0 mm

long), spreading scale margins and several-flowered spikelets, the

right-hand one probably referable to ssp. macilentus (described

below) on the basis of its 3-4-flowered spikelets and slightly shorter

scales (ca 2.5 mm long). Because the identification of the latter

specimen is questionable from the photograph, the specimen on the

left is taken as the lectotype for Sprengel’s name.

KEY TO SUBSPECIES

A. Scales 2. 5-3. 5 mm long, fitting loosely over the achene, the

margins spreading; spikelets 6-22-flowered.

9a. C. LUPULINUS SSP. LUPULINUS.

AA. Scales 1.8-2. 5 mm long, fitting firmly over the achene, the

margins tightly clasping; spikelets 3-5 (-7) flowered.

9b. C. LUPULINUS SSP. MACILENTUS.

1974]

Marcks — Sedge Family II Report

273

9a. Cyperus lupulinus (Spreng.) Marcks ssp. lupulinus Map 9,

Figs. 2 and 5, Table 1.

Mariscus glomeratus Bart. Prod. FI. Phil. 18. 1815, fide Horvat,

1941 (under C. filiculmis, sensu lato) .

Scirpus cyperiformis Muhl. Descr. 41. 1817. TYPE : Baltimore,

Maryland, Muhlenberg 143 sheet 2 in folder 46 (PH!).

Mariscus cyperiformis (Muhl.) Torr. Cat. PL N. Y. 14. 1819.

? Cyperus annuus Spreng. et Link, Jahr. 2 :83. 1820, fide Kukenth.,

1936.

FIGURES 2 AND 3. Distribution maps of Cyperus lupulinus ssp. lupulinus

and C. lupulinus ssp. macilentus.

274 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Cyperus etesios Schult. Mant. 2:114. 1824, fide Kukenth., 1936.

Cyperus Bushii Britt. Man. FI. N. U. S. 1044. 1901. Arkansas,

Sept. 28, 1894, Bush 619. HOLOTYPE: (NY) ; ISOTYPE (MO!).

Cyperus Houghtonii var. Bushii (Britt.) Kukenth. in Engler,

Pflanzenr. 20:470. 1936.

Widespread in the eastern United States on dry exposed sands,

gravels and rocky areas of prairies and oak openings, with geo¬

graphical dispersal centers in the Ozarks and Appalachians, in

Wisconsin rather rare, chiefly south of the Tension Zone on sand¬

stone and granite outcrops, gravelly areas and sand prairies

(Fig. 2).

More southwestern in distribution than ssp. macilentus, with

which it intergrades where sympatric in glaciated territory from

the upper Midwest eastward to the Atlantic Coastal Plain. Popu¬

lation studies (Marcks, 1972) have shown it to hybridize exten¬

sively in the Great Plains and lower Great Lakes region (including

southern Wisconsin) with Cyperus Schweinitzii (cf. p. 276).

TABLE 1. MORPHOLOGICAL COMPARISON OF CYPERUS LUPULINUS

SSP. LUPULINUS, C. LUPULINUS SSP. MACILENTUS,

AND C. SCHWEINITZII.

1974]

Marchs — Sedge Family II Report

275

A comparison of morphological characters of the latter taxon is

made with those of C. lupulinus ssp. lupulinus and C. lupulinus

ssp. macilentus in Table 1. Flowering from late June to October;

fruiting from mid-July to mid-October (n = ca 83 ; Marcks, 1972).

9b. Cyperus lupulinus (Spreng.) Marcks ssp. MACILENTUS

(Fern.) Marcks, stat. nov. Map 10, Figs. IB, 3 and 6, Table 1.

Cyperus filiculmis Vahl var. macilentus Fern., Rhodora 8:128.

1906. TYPE : Valley of the main Penobscot River, in sandy soil,

Penobscot Co., Maine, Fernald 2US. HOLOTYPE : Gray Herbarium

(GH !) ; ISOTYPES (B! C! LCU! PH! MO! US! WIS!).

Cyperus macilentus (Fern.) Bickn., Bull. Torr. Bot. Cl. 35:478.

1908.

Dry exposed sands, from the southern Appalachians to the At¬

lantic Coastal Plain of Virginia, northwestward in glaciated ter¬

ritory to Iowa and Minnesota (Fig. 3), in Wisconsin in sandy soil

of rock outcrops, roadcuts, abandoned fields, young pine planta¬

tions and of glacial lake beds and outwash plains.

Morphologically a highly reduced northeastern segregate of the

above typical subspecies largely confined to glaciated eastern North

America with an apparent center of dispersal in the northern Ap¬

palachians. Previously always included in synonomy with or as a

variety of C. filiculmis, an endemic of the southern Atlantic and

Gulf Coastal Plains, but specifically distinct on the basis of fewer

flowered spikelets, unpigmented scale laminae and shorter achenes.

Flowering July to October; fruiting mid-July to mid-October

(n = ca 83; Marcks, 1972).

10. Cyperus Schweinitzii Torr. Schweinitz’s Cyperus Map 11.

Figs. 1A, 4, 5 and 6, Table 1.

Erect loose-tufted perennials 2.5-9 dm high, from vertical, hard,

corm-like rhizomes 0.8-3 cm long, these branched horizontally and

bearing secondary shoots. Culms rigid, sharply 3-angled, scabrous

on the angles near the apex, 1-2 mm thick. Blades 2-8 mm wide,

pale green, scabrous-margined ; involucral bracts blade-like,

strongly ascending . Inflorescence an open umbel, with a cluster of

subsessile spikes and in addition 3-8 unequal, strongly ascending

pedunculate rays 1-12 cm long. Spikelets loosely disposed from an

elongate rachis 5-11 mm long, ascending, ovate to oblong-lance¬

olate, 3-12-flowered, 5-15 mm long, 3-4 mm wide; rachilla very

late-deciduous, essentially wingless. Scales firm, ovate, fitting

tightly over the achene, their keels green, each ending in a well

276 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

developed mucro 0. 5-1.0 mm long. Achene trigonous-ellipsoid,

2. 2-2. 6 mm X 1.2-1. 4 mm, brown or black.

A southern Great Plains element of sand hills and sand prairies

(Fig. 4), in Wisconsin rather common in dry exposed sands of

glacial outwash plains, river floodplains, dunes of Lake Michigan,

sandstone outcrops, and disturbed sands of roadcuts, sandblows

and pine plantations. Hybridizes extensively in Wisconsin with

C. lupulinus ssp. lupulinus and ssp. macilentus (cf. following

section). Flowering from mid- June to late September; fruiting

from July to early October (n = ca 83; Marcks, 1972).

HYBRIDIZATION IN CYPERUS SECTION LAXIGLUM1

IN WISCONSIN: THE C. LUPULINUS-C. SCHWEINITZI1

COMPLEX

Cyperus lupulinus ssp. lupulinus, C. lupulinus ssp. macilentus and

C. Schweinitzii, all members of the taxonomically difficult section

LAXIGLUMI, are often sympatric in dry exposed sands of prairie

blowouts, roadcuts, abandoned fields and young pine plantations,

sandstone outcrops and lake dunes in glaciated Eastern North

America. Wherever they occur together, they hybridize extensively

(Marcks, 1967, 1972; Marcks and litis, 1967). It is a fact of

greatest phytogeographic significance that many taxonomically

perplexing groups in glaciated eastern North America, such as the

1974]

Marchs — Sedge Family II Report

277

6.0

5.0

E

a

i

i—

Q

£

Li.

<

UJ

'4.0

3.0

2.0

1.0

0.0

FIGURE 5

lupulinus ssp.

V

*

t ^ * $)s

o_

0.9 1.1 1.3

CULM WIDTH (mm.)

K

¥

A f

'k V

Schweinitzii

1.5

1.7

1.9

Involucral Brae t Angle (°)

• less than 40

O more than 140

Culm Texture

(J) scabrous

<5 intermediate

O smooth

Ray Number

'b 5 or more

6 1-4

O none

Scale Mucro Length (mm)

i more than 0.5

6 0.3-0. 5

O less than 0.3

FIGURE 5. Scatter diagram of Wisconsin (WIS) and Illinois State Natural

History Survey (ILLS) herbarium specimens of the Cyperus lupulinus-C.

Schweinitzii complex collected in Mason and Tazewell Counties, Illinois (adja¬

cent counties in central Illinois), clearly showing intermediates between C.

lupulinus ssp. lupulinus (lower left) and C. Schweinitzii (upper right).

1.6

1.4 -

|i.2 -

Q

1.0

20.8

10

o

0.6

0.4 -

FIGURE 6

4

o o

o

o ©

lupulinus ssp .

macilentus

i

id i

; V

6

446

© 4

i'6

1

i y

*

k 4

4

6

4 4 4

4

4 4

V* *

6 \

Schweinitzii

0.0 ' 2.3 2.5 2.7 2.9 3.1 3.3

ACHENE LENGTH ♦ WIDTH (mm.)

3.5

3.7

3.9

Involucral Brae t Angle (°)

• less than 40

O more than 140

Plant Height (cm)

(!) more than 50

6 35-50

O less than 35

Culm Texture

*b scabrous

b intermediate

O smooth

Ray Number

i 5 or more

6 1-4

O none

FIGURE 6. Scatter diagram of mass collection ( Ma/rcks & Marchs 358, col¬

lected 2 mi. E of Big Flats, Juneau Co., Wis.) clearly showing intermediates

between Cyperus lupulinus ssp. macilentus (lower left, hollow circles and one

or no whiskers) and C. Schweinitzii (upper right, solid circles and three

whiskers).

278 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

C. lupulinus-C. Schweinitzii complex, owe their complexity to post¬

glacial hybridization of closely related species, often vicariads,

which evidently met after their long Pleistocene separation in

sometimes far removed “refugia” or “survivia” located south of

the glacial maxima (Hulten, 1937; litis, 1965, 1966, unpublished).

Notable examples which show this phytogeographic pattern and

which form often spectacular hybrid swarms in glaciated terri¬

tories include Actaea pachypoda and A. rubra (Kane, 1966; litis,

1966), Juniperus horizontalis and J. virginiana (Fassett, 1945),

Acer nigrum and A. saccharum (Desmarais, 1952), Helianthus

giganteus and H, grosseserratus (Long, 1961) and Gentianopsis

crinita ssp. crinita and G. crinita ssn. procera (litis, 1965).

Judging from the extraglacial distributions of these LAXI-

GLUMI taxa, C. lupulinus ssp. luvulinus survived glaciation in the

Ozarks (and perhans the Armalachians ; Fig. 2), C. luvulinus ssp.

macilentus on the Atlantic Coastal Plain and in the Appalachians

(Fig. 3) , and C. Schweinitzii in the southern Great Plains (Fig. 4) .

Postglacial migrations into once glaciated eastern North America

resulted in regions where two or three of these taxa are sympatric.

There, subsequent hybridization with backcrossing apparently

accounts for the complex intergrading variation patterns often

displayed by local populations of this complex (Figs. 5 and 6).

Postglacial hybridization of C. lupulinus ssp. lupulinus and

C. Schiveinitzii seems merely to have resulted in a blurring of

species lines and a great increase in population variability in re¬

gions of sympatrv. There may also be some introgression northward

in glaciated territory, since northern plants of either taxon com¬

monly vary slightly in the direction of the other taxon. In contrast,

hybridization between C. lupulinus ssp. macilentus and C. Schwein¬

itzii, aside from resulting in numerous highly variable hybrid

swarms, especially in areas of recent human disturbance, appears

to have resulted in the origin of a new species, C. Houghtonii,

within the last 10,000 years in naturally disturbed “Jack Pine”

(Pinus Banksiana) barrens of the upper Great Lakes region, a

hypothesis discussed more fully below.

Key to Hybrid Forms

A. Rachis 5-10 mm long; scales 3. 0-3. 5 mm long, fitting loosely

over the achene, the margins spreading. _lla. C. LUPULINUS

SSP. LUPULINUS X C. SCHWEINITZII.

AA. Rachis 2-5 mm long; scales 2.2-3. 2 mm long, fitting firmly

over the achene, the margins clasping the achene. _

_ lib. C. LUPULINUS SSP. MACILENTUS X

C. SCHWEINITZII.

1974]

MarcJcs — Sedge Family II Report

279

11a. Cyperus lupulinus (Spreng.) Marcks ssp. lupulinus X

C. Schweinitzii Torr. Map 12, Fig. 7.

Cyperus X mesochorus Geise, Amer. Midi. Nat. 15:249. 1934.

Cyperus Houghtonii var. uherior Kiikenth. in Engler, Pflanzenr.

20:469. 1936.

Usually similar to C. Schweinitzii , but with fewer rays, more

divergent involucral bracts and rays and only slightly scabrous

culms; sometimes with reduced seed set. These variable hybrids

occur in overgrazed or disturbed sands of sand prairies and sand

hills from the southern Great Plains northeastward in glaciated

territory to similar habitats in the lower Great Lakes region and

eastward more locally to eastern Pennsylvania (Fig. 7), in Wiscon¬

sin confined to disturbed sand prairies and outcrops south of the

Tension Zone. Hybrid scatter diagrams of local or regional popula¬

tions of this complex often show a great wealth of variously inter¬

mediate plants, suggesting crossing and backcrossing (Fig. 5).

Flowering and fruiting synchronously with the parents.

lib. Cyperus lupulinus (Spreng.) Marcks ssp. macxlentus

(Fern.) Marcks x C. Schweinitzii Torr. Map 13, Fig. 8.

Usually resembling C. Schweinitzii , but mostly smaller and more

slender with fewer rays, more divergent-spreading involucral

bracts, smaller aehenes, and only slightly scabrous culms; often

with low seed set. These highly variable hybrids center on the

region of parental sympatry in the Great Lakes region (Figs. 3, 4

and 7), in Wisconsin occurring on sandstone outcrops and in dry

exposed sands of roadcuts, sandblows of sand prairies, abandoned

fields and young pine plantations. Hybrid scatter diagrams of local

populations (including specimens of the highly fertile putative

stabilized hybrid, C. Houghtonii ) often show a rather complete

bridge of variously intermediate forms connecting the parental taxa

(Fig. 6). Flowering and fruiting synchronously with the parents.

Diakinesis and metaphase I configurations have several ring com¬

plexes which do not permit accurate chromosome counts (Marcks,

1972).

12. Cyperus Houghtonii Torr. Map 14, Fig. 9.

Houghton’s Cyperus

Erect perennials 6-45 cm high, in small loose tufts borne from

a cluster of hard, subglobose, corm-like rhizomes. Culms 3-angled,

smooth , 0.5-1. 5 mm thick at the apex. Blades 1-6 mm wide,

280 Wisconsin Academy of Sciences , Aids and Letters [Vol. 62

FIGURE 7

C LUPULINUS S5P LUPULINUS

C SCHWEINITZII

MAXIMA

7

3 -

FIGURE 8

C LUPULINUS SSP MACILENTUS

V. X

S C SCHWEINITZII

/ GLACIAL MAXIMA

_ -H

FIGURES 7 AND 8. Distribution maps of

X C. Schweinitzii and C. lupulinus ssp.

Cyperus lupulinus ssp. lupulinus

macilentus X C. Schweinitzii.

1974]

Marcks — Sedge Family II Report

281

scabrous-margined ; involucral bracts blade-like, divergent. In¬

florescence with a small dense cluster of terminal spikes and usually

also with 1-8 unequal, obliquely ascending pedunculate rays up to

8 cm long. Spikelets subdigitate from a short rachis 2-5 mm long,

3-18-flowered, 4-15 mm long, 2.5-3 mm wide. Scales rotund , 2. 0-2. 5

mm long; keel broad and green; laminae traversed by 3-4 promi¬

nent nerves, the margins spreading or clasping. Achenes broadly

ovoid, rounded at both ends, the sides markedly concave, 1.6-1. 9 X

1.0-1. 4 mm, dark brown.

A species narrowly restricted to dry exposed sands of “Jack

Pine” (Pinus Banksiana) barrens in glaciated eastern North

America (Fig. 9), in Wisconsin occurring mostly north of the

Tension Zone in pure sands of glacial lake beds and outwash plains.

Extensive population studies (Marcks, 1967, 1972), including

morphologic, cytologic and chromatographic analyses, indicate

C. Houghtonii to be of backcross hybrid origin to C. lupulinus ssp.

macilentus, selected out of postglacially formed hybrid swarms

of C. lupulinus ssp. macilentus and C. Schiveinitzii.

Natural disturbance resulting from wind throw and Are have

provided dry exposed sands suitable for establishment of plants of

the C. lupulinus ssp. macilentus-C . Schweinitzii complex in the

“Jack Pine” barrens essentially since glacial recession (Curtis,

1959). Because these glacial habitats are surely different and per-

FIGURE 9

C. HOUGHTONII

glacial maxima

FIGURE 9. Distribution map of Cy perns Houghtonii.

282 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

^CYPERUS LUPLJUNUS SSP MACILENTUS

CYPERUS SCHWEINITZII I

r ' — I i _|_ j

Me

haps somewhat intermediate to those in which the parental stocks

have had long adaptive histories south of the glacial maxima, one

might expect that new adaptive genotypes could have been selected

out of hybrid swarms in the naturally disturbed “Jack Pine” bar¬

rens. Stabilizing selection acting on hybrid swarms has thus

apparently allowed C. Houghtonii to evolve and exploit a new eco¬

logical niche to which neither of its parents is as well adapted.

The restriction of C. Houghtonii to glaciated territory with the

exception of a once collected and presumably adventive station

from Luray, Virginia, constitutes an anomalous distribution sug¬

gestive of a postglacial origin. With rare exceptions, all species

present in glaciated territories of eastern and central North Amer¬

ica have at least part of their range outside the Wisconsin glacial

maxima where we may assume they survived the Wisconsin glacia¬

tion. The rare exceptions include some species of Solidago and

other poorly understood microspecies of hybrid derivatives of large,

actively evolving genera, some microspermous species ( Thismia

americana , Cypripedium arietinum) , which may have been intro¬

duced by stratospheric dispersal from the Old World, various arctic

and boreal elements, which may have survived on or near the ice

sheet, and several Great Lake dune microendemics (Guire and

Voss, 1963; Johnson and litis, 1963; Mickelson and litis, 1966).

A postglacial hybrid origin for C. Houghtonii is thus suggested on

grounds of its narrow restriction to glaciated territory. Flowering

from July to October, fruiting from mid- July to mid-October

(n = ca. 84, 85, 86; Marcks, 1972).

1974]

Marchs — Sedge Family II Report

Acknowledgements

283

Field work in part supported by the Research Committee of the

University of Wisconsin on funds from the Wisconsin Alumni

Research Foundation.

Special thanks are due to the author’s major professor, Dr.

Hugh H. Utis, for his botanical appreciation of Cyperus and for

critical manuscript reading. Acknowledgement is also extended to

Mr. Theodore S. Cochrane for aid in preparing illustrations and

maps, for editing the paper and for seeing it through press.

BIBLIOGRAPHY

CURTIS, J. T. 1959. The Vegetation of Wisconsin. Univ. of Wisconsin Press,

Madison.

DARLINGTON, C. D., and A. P. WYLIE. 1955. Chromosome Atlas of Flower¬

ing Plants. George Allen & Unwin, Ltd., London.

DESMARAIS, Y. 1952. Dynamics of leaf variations in the sugar maples.

Brittonia 7:347-387.

FAS SETT, N. C. 1945. Juniperus virginiana, J. horizontalis, and J. scopulorum

— IV. Hybrid swarms of J. virginiana and J. horizontalis . Bull. Torrey

Bot. Club 72:379-384.

GILLY, C. L. 1946. The Cyperaceae of Iowa. Iowa State Coll. J. Sci. 21:55-

151.

GREENE, H. C. 1953. Preliminary reports on the flora of Wisconsin. No. 37.

Cyperaceae part I — Cyperus, Dulichium , Eleocharis, Bulbostylis, Fim-

bristylis \ Eriophorum, Scirpus, Hemicarpha, Rhynchospora, Psilocarya,

Cladium, Scleria. Trans. Wis. Acad. Sci. Arts, Lett. 42:47-67.

GUIRE, K. E., and E. G. VOSS. 1963. Distributions of distinctive shoreline

plants in the Great Lakes region. Mich. Bot. 2:99-114.

HICKS, G. C. 1929. Cytological studies in Cyperus, Eleocharis, Dulichium,

and Eriophorum. Bot. Gaz. 88:132-149.

HORVAT, M. 1941. A revision of the subgenus Mariscus found in the United

States. Contrib. Biol. Lab. Cath. Univ. Amer. No. 33.

HULTEN, E. 1937. Outline of the History of Arctic and Boreal Biota during

the Quaternary Period. Bokforlags aktiebolaget Thule, Stockholm.

ILTIS, H. H. 1965. The genus Gentianopsis (Gentianaceae) : transfers and

phytogeographic comments. Sida 2:129-154.

- . 1966. The western element in the eastern North American flora and

its phytogeographic implications. Amer. J. Bot. 53:634 (abstract).

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, Vernonieae, Cynarieae, and Cichorieae). Trans. Wis. Acad. Sci.

Arts, Lett. 52:255-342.

KANE, J. M. 1966. Hybrids of Actaea alba and A. rubra (Ranunculaceae)

in Wisconsin. M. S. Thesis, Univ. of Wisconsin, Madison.

KiiKENTHAL, G. 1936. Cyperaceae-Scirpoideae-Cypereae. In A. Engler,

Pflanzenr. 20:1-671.

LONG, R. W. 1961. Biosystematics of two species of Helianthus (Com¬

positae), II. Natural populations and taxonomy. Brittonia 13:129-141.

MARCKS, B. G. 1967. Population studies in the Cyperus filiculmis complex

of eastern North America. M. S. Thesis, Univ. of Wisconsin, Madison.

MARCKS, B. G. 1972. Population studies in North American Cyperus section

LAXIGLUMI (Cyperaceae). Ph.D. Thesis, Univ. of Wisconsin, Madison.

284 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

- - and H. H. ILTIS. 1967. Post-glacial hybridization of Cyperus

Schweinitzii and C . macilentus. Amer. J. Bot. 54:659-660 (abstract).

MICKELSON, C. J., and H. H. ILTIS. 1966. Preliminary reports on the flora

of Wisconsin No. 55. Compositae IV — Composite family IV (Tribes

Helenieae and Anthemideae) . Trans. Wis. Acad. Sci. Arts, Lett. 55:

187-222.

MURDY, W. H. 1968. Plant speciation associated with granite outcrop com¬

munities of the southeastern piedmont. Rhodora 70:394-407.

O’NEILL, H. 1942. The status and distribution of some Cyperaceae in North

and South America. Rhodora 44:43-64, 77-89.

VOSS, E. G. 1972. Michigan Flora Part I Gymnosperms and Monocots. Cran-

brook Inst, of Science, Bloomfield Hills.

ANTIMYCIN— BEYOND TELEOCIDE

Maiy Ellen Antonioni

University Wisconsin — •

Madison

ABSTRACT

Two species of clams, Lampsilis siliquoidea and Elliptio dilatatus ,

were exposed to each of 4 dosage levels of antimycin (5, 10, 12, and

15 parts per billion antimycin/water) at three temperatures (17 C,

22 C, and 27 C). Each temperature test constituted a “run” of 27

days, during which the organisms were observed and dieoff

recorded. The pH was a constant 7.5 and general water quality

constant. At run III (27 C), fluorescein dye (used as a tracer dye

with antimycin) was also tested alone and in combination with 15

ppb antimycin.

For both species, increase in dieoff as temperature increased was

significant for 12 ppb and 15 ppb, but not for 5 or 10 ppb. There

was an overall significant increase in dieoff as temperature in¬

creased. A comparison among 15 ppb antimycin, 15 ppb antimycin

plus fluorescein, and fluorescein alone, yielded the following re¬

sults: There was no significant difference in dieoff between the

15 ppb antimycin and 15 ppb antimycin plus fluorescein; there

was no dieoff for fluorescein dye alone.

The increase in dieoff with increasing temperature was sig¬

nificantly greater for L. siliquoidea than for E. dilatatus for

dosages 5 and 10 ppb. The total number of dead individuals after

27 days was significantly greater for L. siliquoidea than for E.

dilatatus at 22 C and 27 C, but not at 17 C.

INTRODUCTION

Reclamation of streams, ponds, or lakes is the control of “unde¬

sirable” species of fish with materials and methods for eradicating

such fish, either selectively or totally with subsequent replacement

by “desirable” species. Undesirable species are defined as those

with little or no commercial or sport fishing value; for example,

carp ( Cyprinus carpio) , green sunfish ( Lepomis cyanellus) ,

stunted yellow perch ( Perea flavescens) , etc.

A number of management methods for thus reclaiming streams

or lakes have been tried with little success; such methods include

drainage, seining, and trapping of undesirable species. More

recently, poisoning of such habitats with various teleocides has

285

286 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

been attempted with greater success, but not without added prob¬

lems of endangering non-target species of fish, other aquatic life

and non-aquatic life present in, or dependent upon, the aquatic

habitat.

One such teleocide is antimycin, which is being widely used pri¬

marily as a single management tool. This practice unwisely reduces

complicated aquatic ecosystems to the few component parts which

are able to withstand its effects. To date, few adequate studies

have been made regarding the effects of antimycin on non-target

organisms, on the reproductive success of organisms which do sur¬

vive, or on post-treatment productivity of ecosystems which have

been altered and simplified by its use. The present study is but one

of many needed to thoroughly evaluate antimycin as a teleocide.

At the University of Wisconsin in the Department of Plant

Pathology, Leben and Keitt (1948) noted that a culture of the

apple scab fungus Venturia inequalis, was markedly inhibited by

a white, slow-growing, actinomycete contaminant. An antibiotic

was isolated and discovered to be particularly effective against

fungi and hence was named antimycin.

A new phase of antimycin research began in 1963 when Derse

and Strong (1963) reported that fish were extremely sensitive to

the antibiotic. The Wisconsin Alumni Research Foundation, which

did much of the original research on antimycin as a teleocide,

licensed Ayerst Laboratories, New York, a division of American

Home Products Corporation, to produce and market antimycin.

This product, under the tradename of “Fintrol”, has been approved

for use in fresh water fish management and is registered as a

teleocide by the United States and Canadian governments. (Radon-

ski and Wendt, 1966).

There are three registered formulations of antimycin presently

available (Lennon, et al., 1971). Fintrol-5 consists of antimycin

coated on sand grains in such a way as to release the toxicant

evenly in the first 5 feet of water as the sand sinks; Fintrol-15

which releases it in the first 15 feet of depth, and a liquid, Fintrol

Concentrate, which was developed for use in very shallow running

waters and streams.

Experiments are being conducted on a fourth type of antimycin,

a cake form, that can be suspended in streams to release a specific

amount of the toxicant at a certain rate.

The qualities of antimycin which make it attractive as a tele¬

ocide can be summarized as follows : It is relatively specific to fish,

i.e., fish-killing concentrations are considered harmless to other

aquatic life, waterfowl and mammals. It is effective in small con¬

centrations against all life stages of fish, egg through adult. Its

respiratory inhibiting properties are irreversible at lethal dosages.

1974] Antonioni — Antimycin — Beyond Teleocide 287

It is odorless and colorless in water and does not repel fish from

treatment areas. Thus, it is the first fish toxicant to lend itself well

to partial or spot treatments of lakes. It can also be used somewhat

selectively against target species in certain circumstances (Radon-

ski, 1967; Burress and Luhning, 1969). Antimycin degrades rap¬

idly in water, usually within 4 days or sooner where pH is high.

When necessary, it can be detoxified in water by addition of potas¬

sium permanganate (Berger, et ah, 1969).

The treatment of the Rock River drainage system in Wisconsin

with antimycin is the largest reclamation project attempted thus

far using antimycin as a single management tool. Two hundred-

and-seventy-four miles of stream (East branch, Rock River) and

four impounded ponds were treated at the outset of the reclamation

in 1971. Another 226 miles of stream and over 30,000 acres of im¬

pounded waters (including the nationally known Horicon Marsh,

Dodge County, Wisconsin) were in the process of being treated as

of the date of this writing.

In a post-treatment survey of the East Branch of the Rock River

in Dodge County, it was discovered that mussels were apparently

one of the non-target species greatly affected by antimycin treat¬

ment (Bratley and Mathiak, 1971). Due to the unavailability of

the above reference, it is reiterated here in some detail. It can be

found in its entirety in the Environmental Impact Statement,

August, 1973, submitted to the State of Wisconsin by the Depart¬

ment of Natural Resources.

The study area was located on the East Branch of the Rock

River, a quarter mile south of Allenton, Wisconsin. It extended

upstream for 380 feet. Here, the stream was near the upper

reaches of the East Branch, ranging from 8 to 12 feet in width

and 6 to 18 inches in depth. The bottom was firm and gravelly,

though overlain with thick layers of silt.

Sampling was begun 8 days after antimycin treatment at which

time the water was very clear. The study continued for 40 days

after treatment, after which mortality was too low for further

sampling.

The entire area was traversed by two biologists walking abreast

upstream collecting all mussels lying on the stream bottom which

had died after treatment. Only those which still had parts of the

viscera intact were collected. Empty shells ivere not included in the

mortality counts. There was a total of 8 collections made within a

month (August 19, 1971-September 20, 1971). Near the down¬

stream end of the study area, a 25-foot section was measured off

wherein the bottom substrate was thoroughly hand-grubbed for

both live and dead mussels. The dead mussels were collected and

the live specimens were counted, identified, and returned to the

288 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

stream. This section was sampled at three separate times to deter¬

mine remaining species and numbers. Two other 25-foot sections,

one approximately a half mile and the other about three-fourths

mile upstream to the study area were hand-grubbed thoroughly

two months after treatment for live mussels to determine survival

in a different area. A one-inch wire mesh fence was placed across

the stream at either end of the study area for two days during the

time of peak mortality. The object was to catch any mussels which

may have drifted into or out of the study area and thus bias the

study. This method showed that drifting shells were insignificant.

The mussels were prepared for identification by removing the

viscera with a knife and cleaning the shells to remove algae and

other detritus adhering to the shells. Mr. Billy G. Isom, a mussel

expert from Mussel Shoals, Alabama, assisted in the field iden¬

tification of all individuals recovered.

Eleven species of clams were found to inhabit the 380-foot study

area, and, roughly approximated, made up a population of about

1200 individuals. Forty days after the treatment with antimycin,

approximately 62% mortality had occurred in this population. The

peak mortality rate for the population occurred 16-19 days after

the treatment, but four peak mortality periods were noted : imme¬

diately or shortly after treatment for Lampsilis siliquoidea (fat

TABLE 1. NUMBERS OF NEWLY DEAD CLAMS FOUND IN THE

380-FOOT STUDY AREA OF THE EAST BRANCH OF THE

ROCK RIVER AT ALLENTON AFTER ANTIMYCIN

TREATMENT*

Days Post-Treatment

*The treatment date was 8/11/71

1974] Antonioni — Antimycin — Beyond Teleocide 289

mucket) and Andonata grandis (floater) ; 16 days after for Am-

blema plicata (three-ridge) and Strophitus rugosus (squaw foot) ;

19 days after for Alasmidonta calceolus (slipper shell), Anodon-

toides ferussctcianus (cylindrical paper shell), and Lasmigona com -

planata (white heel-splitter) ; and 27 days after for Elliptio dilata-

tus (lady finger), and Fusconia flava (pig toe). Numbers of the

two remaining species, Alasmidonta marginata, and Leptodea

fragilis, were too small to place in any of these categories. Alas¬

midonta calceolus was eliminated from the study area by the 27th

day after treatment. Representatives of Elliptio dilatatus and

Amblema plicata survived best in the study area (See Table 1).

The study suggests: 1) that mussels are apparently affected by

antimycin; 2) that response rates are different for different

species; 3) that mussels are affected over a long period of time;

4) that the response is delayed (i.e. the mussels do not appear to

be immediately affected) ; and 5) that mussels are apparently

affected at fairly low dosages (i.e., at fish-killing concentrations).

It was the intent of the present study (in a controlled laboratory

experiment, eliminating field variables) to ascertain whether or

not the dieoff of mussels was a direct effect of exposure to anti¬

mycin and/or its tracer dye, fluorescein, or some other factor.

METHODOLOGY

Experimental Population

Due to the inability to collect large numbers of any one species

of mussel, two species were purchased from NASCO Biological

Supply Co. in Fort Atkinson, Wisconsin. Lamps ills siliquoidea and

Elliptio dilatatus were identified and chosen because of their avail¬

ability, recent collection from an untreated area (White River,

Wisconsin), and because both species are native to the Rock River

drainage system. Also according to Brat ley and Mathiak, 1971,

L. siliquoidea was relatively quickly affected following treatment

with antimycin whereas E. dilatatus did not succumb until con¬

siderably later; thus providing two ranges of tolerance to compare

when describing mortality patterns.

The mussels were transported to Madison, Wisconsin, and held

in a stream tank at 10 C for the duration of the experiment. The

holding tank was a large “living stream” aquarium with circulat¬

ing tap water cooled to 10 C. The organisms were not fed or main¬

tained in any other fashion ; this was in keeping with conditions

suggested by the NASCO Biological Laboratory.

Individuals were taken as needed, at random, for treatment.

All individuals of both species were of uniform size, shape, and

color, and appeared to be typical individuals for their species.

290 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

Experimental

The experiment was carried out at the Biotron, University of

Wisconsin, Madison, because of the necessity of maintaining

aquaria at constant temperatures. A large concrete water bath was

constructed for this purpose, with water of a given temperature

pumped into the bath and circulated around the aquaria which

were placed on angle irons to allow circulation underneath as well.

The aquaria were identical 20-gallon, slate bottom tanks, furnished

by the Nevin State Fish Hatchery, Madison, Wisconsin.

The aquaria were entirely covered with a plastic sheet which

was folded back only to make observations. Each tank was aerated

with a tube and airstone. The purpose of the cover was to prevent

evaporation as much as possible, as it was thought that addition

of water during the experiment could affect the level of the anti-

mycin in the treatment aquaria (i.e. evaporation of the water

would alter the parts per billion ratio initially ascribed to each

aquarium). The study area was brightly lighted with fluorescent

tubes automatically controlled to provide 12 hours of “daylight”

and 12 hours of darkness. The room was kept at a constant 10 C in

order to simplify the mechanics of providing the necessary water

bath temperatures, it being easier to heat the water bath against

a cool gradient than to cool it against a warmer room.

Experimental Conditions

Twelve individuals of each species were exposed to each of 4

treatment levels of antimycin at three temperatures (each tem¬

perature series constituting a “run” of 27 days) at a constant pH

of 7.5, and observed for dieoff on a daily basis.

The dosage levels used were as follows : Dosage level Control =

0 ppb (parts per billion antimycin/water) ; Dosage level 2 = 5

ppb; 3 = 10 ppb; 4 = 12 ppb; and 5 = 15 ppb. The temperatures

were 17 C for Run I, 22 C for Run II, and 27 C for Run III. Addi¬

tional conditions for the third temperature run are described

below.

In order to conserve mussels, it was decided to run the tracer

dye tests at Run III only (27 C) . Two additional treatment aquaria

were set up using fluorescein dye alone and fluorescein dye with

15 ppb antimycin. The amount of fluorescein used was based on

calculations for 11 gallons of water per information from Mr.

William Selbig, Fisheries Biologist, Department of Natural Re¬

sources. The 27 C and 15 ppb antimycin were chosen as criteria

for the additional tracer dye tests because these conditions best

approximate the actual treatment conditions in the Rock River

reclamation project (personal observation, pre- and post-

1974] Antonioni — Antimycin — Beyond Teleocide 291

treatment survey of the East Branch, Rock River, summer, 1969 ;

and personal communication with Mr. William Selbig, Project

Director, Rock River Hearing, Juneau, Wisconsin, Spring, 1971).

Twelve individuals of L. siliquoidea and 12 individuals of

E. dilcitatus were exposed to the two additional treatments and

observed for dieoff during the 27-day period for Run III.

The dosage and temperature criteria outlined above for all runs

were chosen because they best approximate those conditions actu¬

ally being used in held treatments, and those used for similar

experiments with fish, from which a great deal of our available

information about the effects of antimycin has been derived.

An exposure limit of 27 days was chosen because it was a long

enough period of time and yet not so long as to jeopardize the

validity of the results by stressing the mussels through unneces¬

sarily long exposure to unnatural habitat conditions ; i.e., aquarium

life, lack of food, etc. The time of 27 days was also given as the

peak dieoff time for Elliptio individuals (Bratley and Mathiak,

1971).

The two main independent variables were dosage level and tem¬

perature. The pH, another important correlate to the lethality of

antimycin, was held at a constant 7.5. It has been observed that

pH does not fluctuate greatly for the types of warm- water stream

habitats which have been treated in the Rock River drainage

system, but remains between a maximum of 8.2 and a minimum

of 7.5 (personal communication with Mr. William Selbig). The

present experiment was carried out at pH of 7.5; it neither in¬

creased nor decreased throughout the experiment, and was un¬

affected by the presence or absence of the experimental organisms.

Analyses other than pH were not made of the water used in this

experiment. However, an extensive analysis of the Biotron tap

water was made by the Wisconsin Alumni Research Foundation in

March, 1972 (see Table 2), and the main parameter of concern for

the present study was pH.

The dependent variable was dieoff, which was recorded on a

daily basis for both species, throughout the three 27-day runs.

Antimycin Formulation Used

The liquid Fintrol Concentrate of antimycin (furnished by

Ayerst Laboratories) was used for the experiment. Liquid Fintrol

is the formulation for treatment of shallow running waters such

as the Rock River drainage system.

An initial stock solution of antimycin was prepared at the outset

of the experiment to ensure uniformity, and, in order to prevent

degradation and activation, it was mixed with an acetone retainer

292 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 2. ANALYSES OF WATER SAMPLES TAKEN IN BIOTRON,

UNIVERSITY OF WISCONSIN, MADISON, MARCH, 1972

solution (per instructions from Ayerst Laboratories) until ready

for addition to the treatment aquaria. The formula for determining

the amounts of antimycin stock solution to add to water in order

to arrive at desired concentrations in parts per billion, was as

follows :

[A] [B] [.35] = C

where

A = Number of gallons of water in aquarium

B = Desired concentration in parts per billion

C = Milliliters of stock solution to add to treatment aquaria

.35 = Conversion factor for gallons to milliliters

The stock solution was prepared by taking 5 milliliters of the ace-

tone/antimycin solution and diluting it in 1 gallon of water

(procedure suggested by Ayerst Laboratories). It was also sug¬

gested that at least 10 gallons of water be used in the treatment

vessels in order to ensure proper mixing of the solution; hence, 11

gallons of water was used for all calculations in the present study.

The Tracer Dye

Standard fluorescein dye (C2oHia05) was obtained from the

Aldrich Chemical Company in Milwaukee, Wisconsin, and added to

the appropriate acetone/ antimycin milliliter solution to obtain

15 ppb antimycin and fluorescein. The dye was also added to the

acetone solution alone to provide proper dissolution when added

to water for the fluorescein test without antimycin.

1974] Antonioni — Antimycin — Beyond Teleocide 293

RESULTS

Experimental Analyses

In a one-way analysis of variance procedure according to Hayes

(1963), a null hypothesis of no difference in dieoff due to treatment

with antimycin (across dosages, within temperatures) was tested

and rejected at the .05 level of significance (See raw data Tables

3, 4, and 5, and summary Table 6.) A separate one-way analysis of

variance was performed for the 27 C data, which included two addi¬

tional treatment blocks [i.e., antimycin at 15 ppb plus fluorescein

(D6) , and fluorescein dye alone (D7) ]. A null hypothesis of no

difference due to treatment with antimycin was tested for Di ( Con¬

trol), D5, D6, and D7, for both species, and rejected at the .05 level

(See summary Table 7).

Following the overall F-test results, individual pairwise compari¬

sons of interest were made according to a post-hoc test between

means (Scheffe, 1959).

A graph of the parameters (number of individuals dead versus

dieoff day) indicated that these two parameters were dependent

and thus either could be chosen as the parameter for the analysis

of variance tests. Since the observations made in this experiment

were carried out over a specified time of 27 days, there was some

question as to whether an analysis of variance procedure could

be used for the data. But the fact that 27 days is extremely short

compared to the life expectancy of clams, indicates that the termi¬

nation of observations after this time would not invalidate the

analysis of variance approach. It was further decided to look at

the average dieoff times at the various treatment criteria. Days

for which dieoff was observed were converted to logarithmic func¬

tions for a more accurate estimation of the average dieoff day and

for conciseness of graphing (See Tables 8 and 9).

Results

Following the overall F-tests performed on the data, several

post-hoc comparisons (using dieoff as the parameter) were made

and tested for significance at the .05 level.

For both species, increase in dieoff as temperature increased,

was significant for 12 ppb (D4) and 15 ppb (DG), but not for 5

ppb (D2) , or 10 ppb (D3) . There was an overall significant increase

in dieoff as temperature increased.

At Run III (27 C), a comparison between 15 ppb (Dr>), and 15

ppb antimycin plus fluorescein (D6), yielded no significant differ¬

ence in dieoff for either L. siliquoidea or E. dilatatus.

The increase in dieoff for L. siliquoidea was significantly greater

than the increase in dieoff for E. dilatatus for Di — D5. The total

294 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 3. RAW DATA FOR RUN I (17 C). NUMBER DEAD OUT

OF TOTAL EXPOSED (12 INDIVIDUALS AT EACH DOSAGE)

D i = 0 ppb (parts per billion antimy cin / water ; Control)

Do = 5 ppb

D 3 =10 ppb

D 4 = 1 2 ppb

D 5 =15 ppb

number of dead individuals was significantly greater for L. sili-

quoidea than for L\ dilatatus at 22 C and 27 C, but not at 17 C.

The following observations (using average dieoff day and num¬

ber of dead individuals for each day as parameters) were made,

though not tested for statistical significance:

1974] Antonioni — Antimycin — Beyond Teleocide 295

TABLE 4„ RAW DATA FOR RUN II (22 C). NUMBER DEAD OUT

OF TOTAL EXPOSED (12 INDIVIDUALS AT EACH DOSAGE)

D i = 0 ppb (parts per billion antimycin/ water; Control)

D 2 = 5 ppb

D 3 = 10 ppb

D 4 = 1 2 ppb

D( =15 ppb

The only remarkable change in the average dieoff day as tem¬

perature increased was for L. siliquoidea , for which a decrease

of 10.2 days is noted between 22 C and 27 C (See Table 9). In¬

crease in dosage did not greatly affect the average dieoff day for

either species. The average dieoff day for E. dilatatus was greater

296 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 5. RAW DATA FOR RUN III (27 C). NUMBER DEAD OUT

OF TOTAL EXPOSED (12 INDIVIDUALS AT EACH DOSAGE)

D i = ppb (parts per billion an timycin/ water; Control)

D 2 = 5 ppb

D 3 =10 ppb

D4 = 12 ppb

D 5 = 15 ppb

D c = 15 ppb Antimycin plus fluorescein tracer dye

D 7 = 0 Fluorescein tracer dye

than for L. siliquoidea at 27 C, but was essentially the same for

22 C and slightly lower at 17 C.

It was evident from the present study that antimycin is harmful

to mussels not only at relatively small dosages, but over a long

i

1974]

Antonioni — Antimycin — Beyond Teleocide

297

TABLE 6. SUMMARY TABLE FOR ONE-WAY ANOVA

OVER ALL DATA

^Significant at .05 level.

TABLE 8. AVERAGE DIEOFF DAYS FOR TOTAL INDIVIDUALS

AT ALL DOSAGES

period of time. Whether or not there is some physiological condi¬

tion unique to the clam which would make its utilization of anti¬

mycin different from that of other organisms is a question beyond

the scope of this study. Its intent was simply to eliminate as many

field variables as possible and concentrate on whether or not there

was dieoff as a direct effect of exposure to antimycin at differing

dosages and temperatures.

DISCUSSION

While there seem to be some advantages to the use of antiniycin

(especially when compared to what else is available), it is the

author’s opinion that there are many loose ends regarding the

298 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 9. AVERAGE DIEOFF DAYS AT EACH DOSAGE

Temperature 17C

Lampsilis siliquoidea Elliptio dilatatus

Average Average

Dosage No. Dead Day No. Dead Day

D i _ 0 _ 0 _

D2 _ 3 18.3 0 _

D3 _ 1 12.0 2 13.3

D4 _ 0 _ 1 12.0

D5 _ _ 1 13.0 0 _

Temperature 22C

Lampsilis siliquoidea Elliptio dilatatus

Average Average

Dosage No. Dead Day No. Dead Day

D i _ _ 1 12.0 0 _

D2 _ 1 17.0 1 17.0

D3 _ 3 11.9 0 _

D4 _ 2 23.2 0 _

Ds _ - _ 5 18.0 0 _

Temperature 27C

Lampsilis siliquoidea Elliptio dilatatus

Average Average

Dosage No. Dead Day No. Dead Day

D i _ 0 _ 0 _

D2 _ 3 4.2 1 18.0

D3 _ 4 4.1 0 _

D4 _ - _ 5 5.6 1 13.0

D5 _ 11 8.5 8 11.8

D6 _ 8 9.5 7 16.2

D 7 _ 0 _ 0 _

research that should be done before a toxicant such as antimycin

is used as a single management tool in the environment.

There have been few exhaustive studies on the effects of anti¬

mycin on non-aquatic and aquatic life other than fish. Antimycin

has also been used in conjunction with other fish toxicants (e.g.

rotenone) ; but the effects of such interactions have not been care¬

fully studied. Howland (1969) studied the interaction of rotenone

1974] Antonioni — Antimycin — Beyond Teleocide 299

and antimycin in fish bioassay experiments. It was concluded that

the toxicity of antimycin and rotenone together is greater than

either used alone; that is, the two chemicals when used in combi¬

nation appear to have an additive effect (though not concluded

to be synergistic).

A few studies have been done regarding effects of antimycin

on mammals, and one or two attempts have been made to determine

whether there are any harmful effects from antimycin residues in

tissues of fish consumed by humans. Herr, Greselin, and Chappel

(1968) studied the toxicity of antimycin and its degradation prod¬

ucts to mammals. Hematologic, organ, and tissue studies were

made on rats which were fed varying doses of antimycin for differ¬

ent periods of time. In oral administration, they concluded that

the toxicity of antimycin was quite low for mice, rabbits, and

quail, but that it was lethal for guinea pigs at comparable dosages.

When injected intraperitoneally, however, the toxicity was about

0.18 mg/kg body weight. Infusion in dogs at the rate of 10 micro-

grams/kg/min. was fatal. In a study of the oral administration

effects of antimycin on reproduction in rats, it was found that no

observable changes occurred during pregnancy but three weeks

after weaning, pups born to treated parents weighed 10% less than

those born to control parents. Skin and eye irritations were noted

in rats and rabbits, when antimycin was applied topically to these

areas (Herr, et al., Ibid).

Suggestions and Implications For Further Research

Baker (1928) has suggested that the nature of the water in

which mussels live profoundly influences them for good or ill. For

example, changes in micro-fauna, vegetation, siltation, or turbidity

might adversely affect a mussel population in short periods of time,

to say nothing of longer range effects by elimination of fish hosts

or interruption of fish-host specificity. Mussel glochidia “react to

certain species of fish and will not complete metamorphosis on

others. . . . When we consider that only a relatively small number

of species of fish are susceptible to infection by glochidia, it be¬

comes evident that there must be a definite ecological relationship

between the fish and parent mussel, such that both must frequent

the same kind of environment at the same time when the glochidia

are ready to be discharged.” Glochidial infection has also been

shown to give immunity against infection by more serious para¬

sites, thus benefiting the host fish (Baker, 1928).

It is also generally conceded that mussels, because of their sensi¬

tivity to pollution, are good indicator species for monitoring

degrees of degradation in streams or lakes. Since mussels are

300 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

generally confined to a particular portion of a lake or stream (a

mussel shoal), they have been used extensively to measure fallout

and pesticide content, as well as other forms of water pollution

(Van Der Schalie, 1970). Thus, clams can be used as indicators of

some of the changes brought about in a river system by human

activity.

It is the suggestion of the author that replications of the present

study (varying pH, etc.), followup studies and similar studies with

other organisms be done to determine the exact nature of the

detrimental effects of antimycin.

The present study has shown that mussels are killed by anti¬

mycin. The next logical step would be to find out why. Perhaps

there is something unique to the physiology of the clam which

makes it sensitive to antimycin in fairly low dosages; perhaps

the respiratory system of the clam is unique in its method of as¬

similation, storage, or utilization of antimycin; perhaps the fact

that clams are filter feeders has an effect on how much antimycin

is taken by the organism.

Hopefully, answers to questions like these will help provide us

with vitally needed information about the mode of action of anti¬

mycin in a variety of organisms.

Those agencies responsible for alterations in the environment

such as are brought about by use of toxicants should be aware of

the fact that they may be unable to “clean up their mess.” Losses

in the biota are irreversible and the alterations will leave gaps

in inter-relationships of flora and fauna, as well as in the oppor¬

tunities for studying unaltered ecosystems.

Stream ecosystem dynamics must be evaluated more carefully

and exhaustive attempts made to discover rare and unique species.

The calculation of correct dosages for streams or lakes with diverse

temperatures, fauna, flora, pH, etc., should be more accurately

gauged to ensure annihilation of target species only. In a manage¬

ment report for the Department of Biological Sciences, University

of Idaho, Moscow, Idaho, Rabe and Wissmar, 1969, cautioned that

more research is necessary to refine recommended concentrations

of antimycin under varied environmental conditions. Even 10 days

post-treatment at 5 ppb, trout fry exposed to the oligotrophic lake

(at 19 C and pH near neutrality), die within 12 hours. The fry

were observed to be in distress less than an hour after the distribu¬

tion of the poison and some were dead in less than 1.5 hours.

Thirteen days after the poisonings, no living crustaceans were

found in the lake samples.

The effects of removal of forage species and rates of growth

of introduced species should be determined over a long period of

1974] Antonioni — Antimycin — Beyond Teleocide 301

time in reclamation projects. A long-range cost/benefit analysis

should then be attempted to evaluate the success of such large-

scale management programs.

BIBLIOGRAPHY

BAKER, F. C. 1928. The fresh water Mollusa of Wisconsin. Pt. 2 Pelecypoda.

Bull. 1527 Univ. Wis. and Bull. 70 Wis. Geol. Nat. Hist. Survey. 495 pp

[p. 13].

BERGER, B. L., LENNON, R. E., and J. W. HOGAN. 1969. Laboratory

studies on antimycin as a fish toxicant. U. S. Bureau of Sport Fisheries

and Wildlife, Investigations in Fish Control: 26, 21 pp.

BURRESS, R. M., and C. W. LUHNING. 1969. Field trials of antimycin as

a selective toxicant in channel catfish ponds. U. S. Bureau of Sport Fish¬

eries and Wildlife, Investigations in Fish Control: 28, 12 pp.

DERSE, P. H., and F. M. STRONG. 1963. Toxicity of antimycin to fish.

Nature 200:600-601.

HAYS, W. L. 1963. Statistics, Holt, Rinehart and Winston, Inc. New York,

New York.

HERR, F., GRESELIN, E., and C. CHAPPEL. 1968. Toxicology studies of

antimycin, a fish eradicant. American Fisheries Soc., Trans. 96:320-326.

HOWLAND, R. M. 1969. Interaction of antimycin and rotenone in bioassays.

Prog. Fish Culturist 31:33-34.

LEBEN, C., and G. W. KEITT. 1948. An antibiotic substance active against

certain phytopathogens. Phytopath. 38:889.

LENNON, R. E. 1970. Control of fresh water fish with chemicals. Proceed¬

ings: Fourth Vertebrate Pest Conference, West Sacramento, California,

pp. 129-137.

LENNON, R. E., et al. 1971. Food and Agricultural Organization Technical

Paper 100.

RABE, F. W., and R. C. WISSMAR. 1969. Some effects of antimycin in an

oligotrophic lake. Prog. Fish Culturist 31 (3) :163.

RADONSKI, G. C. 1967. Antimycin: useful in perch control? Wis. Conserv.

Bull. 32 (2) : 15-16.

RADONSKI, G. C., and R. W. WENDT. 1966. The effect of low dosage appli¬

cation of Fintrol on the yellow perch. Wis. Conserv. Dept., Fish Manag.

Div., Manag. Rpt. 10.

SCHEFFE, H. 1959. The Analysis of Variance , John Wiley and Sons, New

York, New York.

VAN DER SCHALIE, H. 1970. Mussels in the Huron River above Ann Arbor

in 1969. Sterkiana, No. 39, p. 17.

ARBOVIRAL ANTIBODY SURVEY OF WILD MAMMALS

IN SOUTHEASTERN WISCONSIN

Omar M. Amin and Wayne H. Thompson

University Wisconsin — Parkside and

— Madison

ABSTRACT

A survey of host-ectoparasite-pathogen systems was undertaken

in a study of a partially forested area in southeastern Wisconsin

between April and October 1972. Sera of 17 species of small and

medium sized wild mammals were screened for antibodies with

antigens for Eastern, Western, St. Louis, Powassan, California,

and Bunyamwera group viruses. Results of tests with all sera were

negative, except for one from a gray squirrel which had a low titer

HI reaction with Western encephalitis virus. Antibodies to La

Crosse virus, associated with California encephalitis in south¬

western Wisconsin and commonly found in chipmunks and squirrels

there, were not found in this southeastern Wisconsin area.

INTRODUCTION

A definite relationship exists among disease pathogens, arthropod

vectors, and vertebrate host systems. The present study was de¬

signed to obtain information on the ecological role played by

southeastern Wisconsin arthropods and wild mammals in the main¬

tenance and dissemination of vector-borne diseases of man and

animals in nature. The primary stimulation came from the fact

that certain viral disease of man, i.e., California encephalitis,

La Crosse strain, has consistently been reported from southwestern

Wisconsin, Thompson and Inhorn (1967) and Moulton and Thomp¬

son (1971), while no cases have been implicated as being infected

in the southeastern corner of the state. It was not known whether

cases did not actually occur in the southeast, or whether they were

overlooked. The main known vertebrate hosts (gray squirrels and

chipmunks) and mosquito vector (Aedes triseriatus ) are, however,

common to both portions of the state (SE and SW).

The heavily populated southeastern portion was originally leveled

by glaciers and consists of cultivated farmlands and a number of

deciduous woodlots which often contain suburban or rural homes.

These offer many opportunities for exposure of children to forest

dwelling arthropods. The originally non-glacial southwestern por-

303

304 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

tion of the state is a well drained upland region with forests mostly

associated with hillsides and valleys. More information was needed

on the differences in arthropod and mammal populations in these

two areas as well as the pathogens that they harbor. The ecology

of collected arthropods, host, and mosquito populations will be

reported elsewhere.

MATERIALS AND METHODS

The study areas: The Parkside study area encompasses about

75 acres located in Kenosha County, north of the Pike River and

south of County Road A, and about 2.5 miles (4 kilometers) west

of the Lake Michigan shoreline. This ecological habitat consists

of natural undisturbed stream-fed deciduous forested areas inter¬

spaced with lowland and upland prairies (Fig. 1). More prairies

of similar composition and wood lots (wet, dry, and mesic) lie

directly south (across the Pike River) and east of the above de¬

scribed study area. County Road A runs east-west directly north

of the study area across which similar habitats as well as farm

lands are located. A golf course partially separates the western

boundaries of the study area from the larger deciduous forests

(southern mesic) of Petrifying Springs Park.

Animal capture and examination: Havahart mammal live traps

(sizes 1, 2, 3A) were set in 24 designated grids (Fig. 1) 8 per

station, and were regularly operated for 3 consecutive days and

nights every other week between April and October, 1972. A pilot

study was earlier made in the same area during fall, 1971 (Septem¬

ber, October) . Traps were checked twice daily, at dawn and shortly

before sunset. Captured animals were anesthetized with ether after

being shaken out of the traps into cloth laundry bags. They were

sexed, measured, weighed, tagged, and cleaned of ectoparasites.

Blood samples were taken primarily by heart puncture with 3 or 5

ml. syringes and 18, 23, or 25 gauge needles, depending on the size

of the animal, after which they were released. The samples were

collected in 10 ml. screw cap tubes which were then promptly stored

on wet ice until later centrifuged in the laboratory and serum and

clot were each frozen in separate vials (— 80°F.) for later anti¬

body and isolation tests. Temperature, precipitation, and relative

humidity in the study areas were regularly recorded.

Serology: Tests of sera for antibodies were conducted in the

Zoonoses Research laboratory in the State Laboratory of Hygiene,

Madison. Sera were screened in: (1) Hemagglutination-inhibition

(HI) tests with antigens for Eastern equine encephalitis (EEE),

Western equine encephalitis (WEE), St. Louis encephalitis (SLE),

Powassan (POW), and Bunyamwera (BUN) group viruses. (2)

TABLE 1. SEROLOGICAL DATA ON PARKSIDE MAMMALS TESTED FOR EEE, WEE, SLE, POW, BUN, LAC,

TVT, JC, SSH, 1971, 1972

1974]

Amin and Thompson — Survey of Wild Mammals

305

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306 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Neutralization tests in BHK2i tissue culture cells (TCNT) with

California encephalitis (CE) group; La Crosse (LAC), Trivittatus

(TVT), Jamestown Canyon (JC), and Snowshoe Hare (SSH) ;

and Bunyamwera (BUN) group (Wisconsin mosquito isolate

#523) . Sera which had been initially diluted upon sampling to 1 :2,

1:3, or 1:5, were further diluted in the laboratory to a final 1:10

for use in the TCNT tests. Initial dilutions of 1 :10 were not further

diluted. For HI tests, only undiluted sera or those diluted 1:2 or

1:3 in the field were tested. They were each further diluted 1:10

during the acetone treatment prior to testing.

RESULTS AND DISCUSSION

A total of 335 and 441 sera from 292 and 377 different mammals

of 17 species captured in the Parkside study area (September,

October, 1971 and April-October, 1972) were screened in hemag¬

glutination-inhibition and tissue culture neutralization tests,

respectively. The results are shown in Table 1. We found no evi¬

dence of detectable antibodies neutralizing CE or BUN group

viruses in sera of these mammals from the Parkside area, i.e.,

chipmunks and gray squirrels (Table 1), species which commonly

have antibodies when collected from forests in southwestern Wis¬

consin. Sera neutralizing La Crosse virus from chipmunks and

squirrels in southwestern Wisconsin were included as additional

controls for the sensitivity of these tests.

CALIFORNIA ENCEPHALITIS

Human cases of CE in Wisconsin occur mostly in the western

and southwestern parts of the state where forest dwelling mammals

(mainly chipmunks and squirrels) and mosquitoes (primarily

Aedes triseriatus ) are involved in the natural cycle of La Crosse

virus in nature (Thompson and Inhorn, 1967 ; Moulton and Thomp¬

son, 1971). That role has also been experimentally studied (Pantu-

watana et at., 1972). Isolates of La Crosse and other CE group

viruses (TVT, SSH, JC) have also been obtained from pools of

Aedes triseriatus, A. communis group, A. trivittatus, and Culex

pipiens by Thompson et al. and Defoliart et al., 1972). Isolates of

Bunyamwera group virus have also been reported in other parts of

Wisconsin (Anslow et al., 1969). Activity of La Crosse virus in

Wisconsin chipmunks was noted as far east as Dane County

(Thompson, pers. comm.), about 100 miles from the Parkside

study area.

Data on Parkside mammal population density and dynamics

is presented in an accompanying paper (Amin, p. 311 these Trans¬

actions) . The appropriate vectors of these viruses are also abundant

1974] Amin and Thompson — Survey of Wild Mammals

307

FIGURE 1. The Southeastern Wisconsin study area at Parkside.

in southeastern Wisconsin as well as in the southwestern portion of

the state where endemic virus activity has been detected. Table 2

summarizes data for six Aedes mosquito species collected during

July, 1972 in the Parkside study area as well as in nearby localities

of similar botanic composition. More detailed information on Park-

side mosquitoes will be published elsewhere. The absence of neutral¬

izing antibodies can thus only be accounted for by a comparative

lack of La Crosse virus activity in the area. The phytogeographic na¬

ture of southeastern Wisconsin might represent an important factor

supporting this hypothesis. The deciduous wooded areas where

TABLE 2. AEDES COLLECTIONS FROM PARKSIDE STUDY AREA AND NEARBY LOCATIONS, JULY 1972

308 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

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2Three CDC light traps and 3 dry ice (CO 2) traps were operated for four 24 hour periods.

1974] Amin and Thompson — Survey of Wild Mammals 309

A. triseriatus breeds (in basal tree holes) and chipmunks and

gray squirrels abound, in the southeastern corner of the state,

are usually represented by small isolated wood lots. This situation

could present a kind of natural barrier to the spread of these

viruses.

The absence of antibodies to BUN and other California group

(TVT and JC) viruses in the sera of these chipmunks and squirrels

is not as indicative of the lack of activity in the area, because these

animals have not been established as good hosts for replication of

these other viruses and for antibody production, as they have been

with La Crosse.

POWASSAN VIRUS

No reportable HI reactions with Powassan antigen were found

in the routinely collected samples in this study. One low titer reac¬

tion (1:10 only) was seen with the serum of a raccoon which had

been road-killed, but is not considered as significant because of the

likely presence of non-specific inhibitors. Serological evidence dem¬

onstrated antibody reaction against Powassan virus in raccoons,

foxes, and humans in the state of New York (Whitney, 1963).

Natural foci of infection appear to exist in Connecticut and South

Dakota in Ixodes spinipalpis taken from Peromyscus mice as well

as in Colorado in Dermacentor andersoni (Thomas et al., 1960 in

McLean and Larke, 1963).

WESTERN EQUINE ENCEPHALITIS

The one low level HI reaction with the antigen for WEE virus

with the serum from a gray squirrel (0.7 Kg. male) indicates pos¬

sible presence in other animals or birds in this area. Cases of WEE

have been occurring each year in Wisconsin horses. A few human

cases have also been diagnosed during the past five years ; it is likely

that other mild cases were missed (Thompson, 1971). Culex

tarsalis, a common vector of WEE collected only occasionally in

Wisconsin, was not obtained in this study area. Aedes vexans, the

most abundant mosquito in southeastern Wisconsin (Table 2) and

elsewhere in the state, has also been reported as a natural vector

of WEE virus (Miles, 1960). To date, no WEE virus isolates have

been obtained from mosquitoes in Wisconsin. Infections in humans

and horses are usually considered as dead-end infections because

of relatively low and transient viremias.

REFERENCES

ANSLOW, R. O., W. H. THOMPSON, P. H. THOMPSON, G. R. De-

FOLIART, O. PAPADOPOULOS, and R. P. HANSON. 1969. Isolation

of Bunyamwera-group viruses from Wisconsin mosquitoes. Amer. J.

Trop. Med. Hyg. 18:599-608.

310 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

DeFOLIART, G. R., R. 0. ANSLOW, W. H. THOMPSON, R. P. HANSON,

R. E. WRIGHT, and G. E. SATHER. 1972. Isolations of Trivittatus virus

from Wisconsin mosquitoes, 1964-1968. J. Med. Ent. 9(l):67-70.

McLEAN, D. M. and R. P. B. LARKE. 1963. Powassan and Silverwater

viruses: Ecology of two Ontario arboviruses. Can. Med. Assoc. J. 88:182-

185.

MILES, J. A. R. 1960. Epidemiology of arthropod-borne encephalitis. Bull.

Wld. Hlth. Org. 22:339-371.

MOULTON, D. W. and W. H. THOMPSON. 1971. California group virus

infections in small, forest-dwelling mammals of Wisconsin, some ecological

considerations. Amer. J. Trop. Med. Hyg. 20 (3) :474-482.

PANTUWATANA, S., W. H. THOMPSON, D. M. WATTS, and R. P. HAN¬

SON. 1972. Experimental infection of chipmunks and squirrels with

La Crosse and Trivittatus viruses and biological transmission of La Crosse

virus by Aedes triseriatus. Amer. J. Trop. Med. Hyg. 21(4) :476-481.

THOMPSON, W. H. 1971. Arbovirus surveillance in Wisconsin — 1971 Re¬

port, pp. 1-5.

THOMPSON, W. H., R. 0. ANSLOW, R. P. HANSON, and G. R. De¬

FOLIART. 1972. La Crosse virus isolations from mosquitoes in Wisconsin,

1964-1968. Amer. J. Trop. Med. Hyg. 21(l):90-96.

THOMPSON, W. H. and S. L. INHORN. 1967. Arthropod-borne California

group viral encephalitis in Wisconsin. Wis. Med. J. 66:250-253.

WHITNEY, E. 1963. Serologic evidence of group A and B arthropod-borne

virus activity in New York State. Amer. J. Trop. Med. Hyg. 12:417-424.

DISTRIBUTION AND ECOLOGICAL OBSERVATIONS OF WILD

MAMMALS IN SOUTHEASTERN WISCONSIN

Omar M. Amin

University Wisconsin — •

Parkside

ABSTRACT

Seventeen species of small and medium, wild mammals were

trapped during the 1972 season as well as during a pilot study (fall

1971) in two ecologically distinct habitats (deciduous woods and

prairies) in the Parkside study area, Kenosha County, southeastern

Wisconsin. Seasonal and spatial distribution, population dynamics

and other ecological information are discussed for the more com¬

monly encountered mammals; i.e., chipmunks, meadow jumping

mice, gray squirrels, opossums, white footed mice, and raccoons.

Activity usually increased from low in the spring to a peak in the

summer and declined by October. Activity of gray squirrels was

relatively higher in the spring with a peak in August. Chipmunks

experienced a lull in activity during August and the activity peak

occurred during September. Summer breeding activity in chip¬

munks was noted. Variations in population densities of the meadow

jumping mice from year to year were observed. The occurrence of

at least a second breeding season in these mice is suggested. Female

raccoons that failed to conceive during the regular breeding season

can apparently mate again and conceive during May or June. Addi¬

tional notes on breeding and activity periods, preferred habitats,

trap preferences, travelled distances, measurements and weights,

and recapture data are also included.

INTRODUCTION

A survey of host-ectoparasite-pathogen systems was undertaken

in a study area in southeastern Wisconsin between April and Oc¬

tober, 1972, after a pilot study in fall 1971. Studies of arboviral

antibodies and ectoparasitic arthropods (including mosquitoes) are

reported by Amin and Thompson (preceding paper p. 303). The

following data are concerned with the seasonal activity patterns,

distribution, population dynamics and other ecological observations

on some of the 17 mammal species captured.

311

312 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

MATERIALS AND METHODS

The Study Areas. The Parkside study area has been described

by Amin and Thompson. The most prevalent ground layer species

of plants in the southern mesic woods are Osmorthiza claytoni,

of plants in the southern mesic woods are Osmorhiza claytoni,

and Smilacina racemosa. The most prevalent ground layer species

of the southern dry mesic woods are S. racemosa, Amphicarpa

hracteata, O. claytoni, and Desmodium glutinosum.

Considerably more area of similar prairies and wood lots (wet

and dry mesic and mesic) lie directly south (across the Pike River)

and east of this area. County Road A runs east-west directly north

of it across from which similar habitats as well as farm lands are

located. A golf course partially separates the western boundaries

of the study area from the larger deciduous forest (southern mesic)

of the Petrifying Springs Park.

Animal capture and examination: The methods for capture of

the animals and the records made in classification of them have

been described in Amin and Thompson. Details as to the dates of

capture, recapture, and descriptions of the animals are given in

this paper. Temperature, precipitation, and relative humidity were

regularly recorded.

RESULTS AND DISCUSSION

Seventeen mammal species were captured, 2 of which were ob¬

tained only during the pilot study; i.e., Prairie vole (Microtus

ochrogaster) and short tail shrew (Blarina hrevicauda) . These data

are shown by season in Table 1. During the 1972 season, a total of

356 animals were initially captured of which 158 were recaptured

358 times (Table 3).

Eastern Chipmunk, Tamias striatus ohionensis Moulthrop

The Eastern Chipmunk was the most commonly encountered

mammal. Twenty two specimens (and 16 recaptures) were re¬

corded during the pilot study. During the 1972 season the total

number of trap entries exceeded half that of all other mammals

combined (Table 1). Of the 160 initial captures, 35 (22% ; 16 males

and 19 females) were juveniles which entered the traps between

June and October.

Animals weighing 50-74 g measured 200-260 mm (230) 1 in total

length and 70-100 mm (87) in tail length (N = 21). Animals weigh¬

ing 75-99 g measured 215-280 mm (249) in total length and 80-

110 mm (94) in tail length (N = 40). Animals weighing 100 to

1 Figures in parenthesis represent means.

1974]

Amin — Observations of Wild Mammals

313

TABLE 1. MAMMALS CAPTURED IN THE PARKSIDE STUDY AREA

(KENOSHA COUNTY) DURING 1972

*24 hrs.

2No. of initial captures, no. of recaptures (trap entries) of recaptured animals.

3 Total no. of trap entries.

130 g measured 230-300 mm (263) in total length and 75-115 mm

(96) in tail length (N = 82). Adult chipmunks of Wisconsin were

reported by Jackson (1961) to reach 115 g in weight and a maxi¬

mum of 265 mm in total length and 100 mm in tail length.

Sex ratio of all captures was equal. Chipmunks were almost

exclusively active during daytime hours (especially in the morning

and the late afternoon) in the deciduous woods. They entered trap

size 1 almost as often as trap size 2 but rarely entered trap size

3 A (Table 2). Chipmunks had the highest number of recaptures

with 2 specimens recaptured 9 times (bait bias might be involved)

and with a maximum distance recorded of 475 meters travelled by

a male during 1972 (Table 3). Burt (1969) mentioned a home

range of “seldom more than one hundred yards in greatest diame¬

ter,” Jackson (1961) indicated a “maximum radius of about two

hundred feet from homesite.”

The following information suggests that male chipmunks might

be relatively more active than females as judged by the frequency

of trap entries and of recapture away from homesite (footnote,

Table 3). Of the 160 initially captured chipmunks, 92 (46 males

and 46 females) were recaptured 1-9 times. The average number

of trap entries was 3.3 for males and 2.7 for females. The majority

of males and females (60%) were consistently recaptured within

their homesite (same trapping station). However, 25% of the

males and 35% of the females were recaptured within 75-100

meters from their homesite. The remainder, 15% and 5% of the

314 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 2. SEX RATIO, ACTIVITY HOURS, HABITAT PREFERENCE, AND

TRAP SIZE PREFERENCE OF PARKSIDE MAMMALS, 1972

'From last column, Table 1.

2Based on column (A).

3Based on column (B); the remainder were active at night (recovered at dawn) or was captured in prairie.

4The remaining 2% appear to have been active shortly before or during sunset and thus mixed with the night

catch.

5Three of the 6 chipmunks reported in the prairie were yearlings captured during June-August.

"Trap sizes: 1 = 45x13x13 cm 2 = 60x18x18 cm 3 = 105x28x32 cm.

males and females were recaptured as far as 475 and 425 meters

from the homesite, respectively. The average distance travelled

was, however, about 34 meters by both sexes.

Population density in Parkside deciduous woods was estimated

at 11 chipmunks per acre. This figure is relatively higher than the

average of 6-8 per acre, but lower than the maximum of 20 per

acre found by Jackson (1961) elsewhere in Wisconsin.

Chipmunks, as well as other mammals, were blood sampled by

heart puncture for the arboviral antibody survey. Data regarding

the effect of such sampling procedure on the weight changes in the

mammal species were available only for chipmunks because of the

adequate number of recaptures 1-2 days following initial blood

sampling during the first day of the biweekly trapping sequence.

Animals initially weighing 85-99 g (92.5) (N = ll) from which

0.1-1. 5 ml (0.83) of blood was withdrawn by heart puncture lost

an average of 1.9 g (—13 to +11) in body weight 1-2 days later.

Animals weighing 100-122 g (114) (N = 15) from which 0.2-1. 4

ml (0.59) of blood was withdrawn lost an average of 3.7 g (—13

to +8) in body weight. Weight loss in older chipmunks was about

twice that in younger animals despite the smaller average volume

of blood withdrawn from the older group. Apparently younger

chipmunks are less susceptible to weight loss, or more efficient in

recovery.

Only one specimen was captured during April (Table 1). Chip¬

munks were, however, observed sporadically in the study area and

1974]

Amin — Observations of Wild Mammals

315

in nearby localities at least during most of March. Their activity

index increased sharply between April and May following the

termination of the last freezing temperatures in the latter part of

April. Activity continued to increase reaching two peaks in July

and September. Dunford (1972) observed the same phenomenon

in New York and was unable to relate it to unfavorable high tem¬

peratures, breeding activity, or food shortage. A dramatic decline

FIGURE 1. Activity (total trap entries) of 6 mammal species captured in

the Parkside study area correlated with seasonal temperatures, 1972.

Percentages in the lower figure represent the proportion of recaptures in

the total number of trap entries of each mammal species during the cor¬

responding month.

316 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

in activity occurred in October following near and below freezing

temperatures in that month (Fig. 1). Chipmunks were seldom

seen after October. The percentage of recaptures on the activity

curve increased progressively from 13% in May to 86% in October.

Between November and March the eastern chipmunk was reported

to be inactive-dormant but not a regular hibernator (Jackson,

1961).

Highest activity in terms of total number of trap entries was

noted near streams bordered by hill slopes; i.e., in W 1, W 2, and

W 12 (Fig. 1 of Amin and Thompson). This might have resulted

partially from proximity of nesting sites. Activity was second

highest in W 6-W 11, W 13, but minimal in the center of the

wooded plateau farther away from the stream (W 3-5) as well as

in the relatively isolated W 14-W 16.

Data on seasonal weight changes show that spring mating ac¬

tivity appeared to have occurred at least during the latter half of

March (Fig. 2C). Supporting data include the capture of pregnant

female no. 99 on 5 May which was lactating on 27 May and the

capture of the earliest and youngest female no. 64 on 9 June. Ac¬

cording to Jackson (1961), the young do not leave the den before

40 days after birth following a gestation period of about 31 days.

By using data from Allen (1938), male no. 104 which weighed

80 g on 25 June (Fig. 2C) was estimated to have been about 2

months old ; i.e., born during the last week of April. Similarly, male

no. 208 which weighed 78 g on 19 August was estimated to be about

2 months old. The above data show that births resulting from

spring breeding activity continued for at least 2 months between

late April and late June, 1972. The same data (Fig. 2C) also show

that summer breeding activity does take place. Jackson (1961) in¬

dicated that summer mating in Wisconsin occurs between the end

of June and the end of July. My data for pregnant female no. 28

captured on 21 July and lactating females nos. 28, 227, and 247

captured on 5 August and 2 and 16 September, respectively, suggest

that summer mating started during the second half of June and

continued until early August. Similar observations were made by

Allen (1938) in New York.

Fig. 2C also indicates a relatively fast rate of growth in juveniles

between June and early August. Weight increase slowed down con¬

siderably between early and mid-August and mid-September. This

slow down might be related to the late summer lull in activity

referred to above. During the remainder of September and Oc¬

tober faster weight increases in the young resumed. Chipmunks

over one year of age did not seem to experience such dramatic

weight changes. Weights of breeding females, nos. 99 and 28 (Fig.

2C), represented the expected changes resulting from pregnancy

1974]

Amin — Observations of Wild Mammals

317

Figure 2. Seasonal weight changes of raccoons, opossums, and chipmunks.

and birth. Weights of breeding males, e.g., no. 94, show a moderate

fall increase. The increase in weight in all above categories might

be partially related to anticipated overwintering. Forbes (1966)

noted that fall weight increases in some Minnesota chipmunks

corresponded to accumulation of fat in relation to hibernation.

318 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

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1974]

Amin — Observations of Wild Mammals

319

Meadow Jumping* Mouse, Zapus hudsonius intermedins Krutzsch

The jumping mouse had the second highest initial capture count

(Table 1). The fact that none was captured during the pilot study

(Sept., Oct. 1971), in contrast to 25 trap entries during September

and October, 1972 in the same area might be explained by variation

in population density from year to year. Jackson (1961) indicated

that peaks of abundance of Z. hudsonius “do not seem to occur at

the same time as peaks in other species.” Supporting evidence from

this study includes the fact that 15 meadow voles (Microtus

pennsylvanicus) and no jumping mice were captured during the

pilot study, whereas only 2 meadow voles and 73 jumping mice

were captured during the 1972 trapping season. Similar associa¬

tions between the above two mammals were reported in Michigan

by Blair (1940) .

Animals weighing 10 to 19 g measured 180-235 mm (206) in

total length and 105-140 mm (124) in tail length (N = 50). Ani¬

mals weighing 20-25 g measured 200-240 mm (217) in total length

and 125-145 mm (131 mm) in tail length (N=25). Adult jumping

mice of Wisconsin were reported by Jackson (1961) to reach a

maximum of only 227 mm in total length and 140 mm in tail length.

More females than males entered the traps (1.3:1) as shown in

Table 2. However, Townsend (1935) and Blair (1940) reported

61% and 53% of 163 and 64 field collected mice, respectively, as

males. The mice were usually active at night in open prairie situa¬

tions. About 2/3 of the total captures occurred in traps of size 1

and the remaining in traps of size 2 (Table 2) . No more than 2 re¬

captures were recorded of the same individual with a maximum

distance recorded of 100 meters travelled by a female (Table 3).

Although only 14 jumping mice were recaptured, females ap¬

peared to be more active than males. None of 8 recaptured females

was trapped in the same site twice with an average and maximum

distance travelled of 85 and 100 meters, respectively. On the other

hand, all but 2 of the 6 recaptured males were recaptured in the

same site with an average and a maximum distance travelled of

25 and 75 meters, respectively.

No mice were captured in April and only one on 27 May. They

were reported to emerge from hibernation in Wisconsin in late

April to early May (Jackson, 1961). After the end of May, the

activity index climbed rather rapidly to a peak in July. The steady

decrease in activity during August and September continued to a

minimum in October with 75% recaptures (Fig. 1). The above

activity curve corresponded with the overall temperature changes

during 1972. Entrance into hibernation presumably followed the

first heavy frosts early in the fall.

320 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

The main breeding season apparently occurs in late May. Authors

do not seem to agree whether Z. hudsonius has 1 or 2 other breed¬

ing seasons per year; see Jackson (1961) and Burt (1969) for

lack of supporting information. Hamilton (1935) observed only

one litter per year in New York. The following data indicate a

second and possibly a third breeding season during July-August

and during September, respectively, at least in southeastern Wis¬

consin. One pregnant and two laetating females were captured on

22 July; one pregnant and one laetating female were captured on

19 August; one laetating female was captured on 29 September.

Blair (1940) indicated 2 peaks of breeding activity in Michigan,

in the spring and in late summer.

Eastern Gray Squirrel, Sciurus carolinensis hypophaeus Merriam

The gray squirrel was the second most commonly encountered

animal in terms of total number of trap entries (Table 1). Only

one juvenile, weighing 0.162 Kg and measuring 370 mm in total

length and 180 mm in tail length, was captured on 7 July. A con¬

siderably greater proportion of immatures was reported by Chap¬

man (1938) in Ohio. Animals weighing 0.40 to 0.59 Kg measured

410-520 mm (480) in total length and 160-240 mm (215) in tail

length (N = 18). Animals weighing 0.60 to 0.80 Kg measured 460-

560 mm (501) in total length and 190-260 mm (221) in tail length

(N = 29). Adult gray squirrels of Wisconsin were reported by

Jackson (1961) to reach a maximum of only 540 mm in total length

and 252 mm in tail length.

More males than females entered the traps (1:0.8). Sex ratio

of captured squirrels appears to vary from year to year. Chapman

(1938) reported 49, 60, and 39% males among 144, 55 and 89 gray

squirrels trapped during 1935, 1936, and 1937, respectively, in

Ohio. They were almost exclusively active during day hours and

only in wooded situations. Most preferred entry in size 3A traps

but 17% and 18% entered traps size 1 and 2, respectively (Ta¬

ble 2). Animals captured more than once did not show any pro¬

nounced consistency in trap size preference. Up to 9 recaptures

of a female and a maximum distance of 425 meters travelled by

a male (Table 3) were recorded. Jackson (1961) mentioned a usual

home range of the gray squirrel “within one thousand feet of its

nest”, although it might travel distances of 4 miles or more in

search of food.

Population density in Parkside deciduous woods was estimated

at 2.4 gray squirrels per acre. Jackson (1961) found that one pair

per acre is a common number in Wisconsin and that “populations

of 20 or more to an acre of woodland may occur.”

1974] Amin — Observations of Wild Mammals 321

None of the 33 recaptured animals was exclusively trapped in

the same site. Males appeared to be relatively more active than

females. The 18 males travelled an average and a maximum dis¬

tance of 112 and 425 meters. Corresponding figures in females

were 70 and 325 (N = 15). Similar observations were reported by

Hungerford and Wilder (1941) in Connecticut. The average num¬

ber of trap entries was 3 for recaptured males and females.

The activity curve started relatively high in April (Fig. 1) indi¬

cating a moderate activity prior to the initiation of trapping in that

month. The slight decrease of activity in May corresponded with

the spring canopy coverage and the possible relative obscurity of

traps. Activity peaked in July and August but declined considerably

in September. The slight increase during October indicates that

activity did not cease after trapping was discontinued but probably

continued at least during November. Personal observations sub¬

stantiate the above information and also indicate that gray squir¬

rels are also intermittently active throughout the winter, at least

in southeastern Wisconsin.

Opossum Didelphis marsupialis virginiana Kerr

Three opossums were captured during the pilot study. During

1972, a total of 24 animals were captured (Table 1), of which 11

(45%; 4 males and 7 females) were juveniles, which entered the

traps between July and October. Holmes and Sanderson (1965)

reported that about 2/3 of over 400 trapped Illinois opossums were

juveniles. Animals weighing 0.4 to 0.99 Kg measured 420-630 mm

(506) in total length and 170-250 mm (206) in tail length

(N = ll). Animals weighing 1 to 1.9 Kg measured 600-700 mm

(655) in total length and 210-300 mm (260) in tail length (N = 4) .

Animals weighing 2-3 Kg measured 650-800 mm (738) in total

length and 230-320 mm (280) in tail length (N = 8). One pregnant

female weighing 3.4 Kg which was captured on 9 June, measured

830 mm in total length and 300 mm in tail length.

More females than males entered the traps (1.7:1), as shown

in Table 2. Jackson (1961), however, indicated that wild popula¬

tions in Wisconsin have “normally about 20 percent more males

than females.” Similarly, Lay (1942) and Reynolds (1945) ob¬

served sex ratios of 1.33 males : 1 female in Texas and 1.22 males :

1 female in Missouri, respectively. Holmes and Sanderson (1965),

however, reported an even sex ratio in Illinois with males being

caught less frequently than females.

Opossums were active strictly at night in wooded habitats.

McManus (1971) indicated that spring and summer nocturnal

322 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

activity of captive opossums in New York peaked during midnight

hours and markedly decreased toward dusk and dawn. However,

in fall and winter, they were active at approximately the same

level throughout the night. This hourly activity pattern was shown

to be related to seasonal variation in ambient temperature. Over

half of the captures were recorded in traps size 3A and most of

the remainder in traps size 2 (Table 2). Up to 8 recaptures were

recorded for one female with a maximum distance of 550 meters

travelled by a female (Table 3). Jackson (1961) indicated that

individuals “may wander a mile or more from the home site” in

search of food, with a normal home range of 20-40 acres.

Females appeared to have been relatively more active than males

with an average number of trap entries in recaptures of 2.4 and

1.3 respectively. The four females travelled an average and a maxi¬

mum distance of 215 and 450 meters. Corresponding figures in

males were 175 and 200 (N = 3) . The sample size of 7 recaptured

animals was, however, too small to validate the above statement.

Holmes and Sanderson (1965) indicated that adult males are much

more mobile than adult females and juveniles.

Opossum activity gradually increased from low in April to a

peak in July (corresponding with increased temperature), declined

in August but retained a moderate level during September and

October. The per cent recaptured in the latter 2 months was low

because of the more frequent trap entries of juveniles. After Oc¬

tober and during winter, opossums were occasionally seen wander¬

ing at night in and around the study area.

The recovery of pregnant females nos. 51, 73, and 24, during

May and June and of juveniles between early July and September

(Fig. 2B) indicated a breeding season between March and end of

May and possibly early June.

White-footed Mouse Peromyscus leucopus noveboracensis (Fischer)

White-footed mice appear to have been more abundant in 1971

than in 1972. During September, October, 1971 17 specimens were

trapped (with 2 recaptures), while during the whole 1972 season

a total of 20 animals entered the traps (with 6 recaptures) (Ta¬

ble 1) . Animals weighing 10 to 19.9 g measured 135-190 mm (158)

in total length and 60-80 mm (70) in tail length (N = 9). Animals

weighing 20 to 26 g measured 145-190 mm (165) in total length

and 65-80 mm (75) in tail length (N = 10).

More males than females entered the traps (1:0.7) ; see Table 2.

In Michigan and Connecticut, Nicholson (1941) and Hirth (1959)

reported similar observations (68 and 58% males, respectively).

1974]

Amin — Observations of Wild Mammals

323

Nicholson (1941), however, reported more even sex ratios among

litters and immatores. They .were exclusively active at night, mostly

in wooded areas (Table 2). Activity was reported to peak at and

shortly after sunset in Ontario (Mann, 1954) ; near Madison (Em-

len et ah , 1957) ; and in Connecticut (Hirth 1959), as well as

shortly before sunrise near Madison (Emlen et ah, 1957). They

were only captured in size 1 traps (Table 2). No more than 6 re¬

captures were recorded with a maximum distance of 175 meters

travelled by a male (Table 3). The 2 recaptured females were each

trapped twice in the same site. Nicholson (1941) and Hirth (1959)

indicated greater travelling distances for males than for females

in southern Michigan and in Connecticut, respectively. Jackson

(1961) indicated a normal home range of about 0.25 acre.

After April, activity increased progressively during the trapping

season to a peak in October. There was a slight decline only during

August (Fig. 1). Burt (1969) mentioned a resting period (from

breeding) in July or August. The consistent increase in activity

after August indicated a marked activity at least during late fall

and early winter in southeastern Wisconsin. Hirth (1959) sug¬

gested an inverse correlation between rainfall (when it occurred

before midnight) and activity (judged by frequency of trapping

success) .

Breeding was reported to occur between March and October

(Nicholson, 1941, Hirth, 1959, Jackson, 1961, and Burt, 1969)

with 4 broods being raised between April and November (Jackson,

1961, and Burt, 1969). The recovery of pregnant females on 9 July,

19 August, 15 September, and 13 October suggest the presence

of the 3rd and the 4th breeding seasons in southeastern Wisconsin.

Raccoon Procyon lotor hirtus Nelson and Goldman

Fourteen raccoons were captured of which 6 (43% ; 2 males and

4 females) were juveniles, which entered the traps between July

and September. Mech et ah (1968) indicated an even ratio of young

to adults in Minnesota. Stuewer (1943) observed 65% juveniles in

Michigan and Sonenshine and Winslow (1972) reported about

20 and 23% juveniles in two Virginia localities. Animals weighing

1.5 to 3.45 Kg measured 590-700 mm (630) in total length and

150-220 mm (180) in tail length (N = 6). Animals weighing 3.5

to 5.49 Kg measured 750-880 mm (828) in total length and 250-

290 mm (266) in tail length (N=5). Two animals weighing 5.5

and 7.2 Kg measured 830 and 970 mm in total length and 230 in

tail length. One pregnant female weighing 8.5 Kg, which was cap¬

tured on 23 June, measured 970 mm in total length and 330 mm

in tail length. Adult raccoons of Wisconsin were reported by Jack-

324 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

son (1961) to reach a maximum of only 960 mm in total length

and 275 mm in tail length.

More females than males entered the traps (2.2 :1) . Other “field”

sex ratios showed variable bias toward males, e.g. Stuewer (1943)

(1.08:1) and Sonenshine and Winslow (1972) (1.64:1). However,

sex ratio in litters was observed to be 1 male :1. 36 females in

Michigan (Stuewer, 1943). Mech et al. (1968) observed an even

sex ratio in yearlings and adults in Minnesota. Raccoons were

exclusively active at night and mostly in wooded areas. All entered

size 3A traps (Table 2). No more than 3 recaptures were recorded

with a maximum distance recorded of 300 meters travelled by a

male. Stuewer (1943) observed that males travel more extensively

than females. Jackson (1961) indicated a normal home range of

about 2 miles.

The activity of raccoons increased from low in April to a peak

in August, declined considerably in September, and was not back

to peak in October; a pattern corresponding closely with that of

the prevailing temperatures.

Jackson (1961) and Burt (1969) referred to one mating season

during late January and February with young being born in April

and May after a gestation period of about 64 days. Juveniles nos.

6, 18, 11, and 22 (Fig. 2A) appear to be the products of such a

breeding season. Juveniles nos. 6 and 18 were estimated to have

been 13 and 11 weeks old (using Stuewer’s, 1943 weight/age data)

and to have been born in mid- or late April, respectively. The preg¬

nant female (no. 95) which was captured on 23 June, however, did

not fit this pattern. She must be considered as a female that did

not conceive during the regular breeding season and that mated

again sometime during May or June. Stuewer (1943) and Jackson

(1961) also reported such cases. Births as late as August and

September were reported in New England States and Indiana by

Whitney (1931) and Lehman (1968), respectively. Dorney (1953)

took 2 males weighing less than 4 pounds from Horicon Marsh,

Wisconsin in November.

Cottontail Rabbit Sylvilagus fioridanus mearnsii (J. A. Allen)

Seven cottontails weighing 0.9-1. 2 Kg and measuring 420-470

mm (443) in total length and 40-70 mm (54) in tail length were

captured.

Striped ground Squirrel Citellus tridecemlineatus (Mitchill)

Five squirrels weighing 120-155 g and measuring 260-290 mm

(270) in total length and 90-100 mm (95) in tail length were

captured.

1974] Amin — Observations of Wild Mammals 325

Long-tailed Weasel Mustella frenata noveboracensis (Emmons)

Three weasals weighing 184-215 g and measuring 390-440 mm

(407) in total length and 130-180 mm (157) in tail length were

captured.

Eastern Fox Squirrel Sciurus niger rufiv enter

Geoff roy-Saint-Hilaire

Two animals weighing 0.55 and 0.80 Kg and measuring 460 and

580 mm in total length and 180 and 260 mm in tail length were

captured. Adult Fox squirrels of Wisconsin were reported by Jack-

son (1961) to reach a maximum of only 565 mm in total length.

Meadow Vole Microtus pennsylvanicus (Ord)

Two specimens weighing 40 g each and measuring 145 and 160

mm in total length and 40 and 42 mm in tail length were captured

during the 1972 season. This species was apparently more abundant

during 1971; 15 specimens were captured during the pilot study

(with one recapture).

Franklin’s Ground Squirrel Citellus franklinii (Sabine)

One female weighing 300 g and measuring 360 mm in total length

and 135 mm in tail length was captured.

Norway Rat Rattus norvegicus (Berkenhout)

One pregnant female weighing 230 g and measuring 370 mm

in total length and 160 mm in tail length was captured on 30

September.

Striped Skunk Mephitis mephitis hudsonica Richardson

One skunk was captured on 21 July, 1972. Another one was

trapped during the pilot study on 10 October, 1971 which was

recaptured the following day.

Southern Woodchuck Mar mot a monax monax (Linnaeus)

One specimen weighing 4.1 Kg and measuring 710 mm in total

length and 150 mm in tail length was captured on 28 May, 1972.

Adult woodchucks of Wisconsin were reported by Jackson (1961)

to reach a maximum of only 640 mm in total length.

Additional information on the above less abundant mammals is

included in Tables 1-3.

ACKNOWLEDGEMENT

Supported in part by a Summer 1972 WARF grant awarded by

the University of Wisconsin — Madison, Graduate School.

326 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

REFERENCES

ALLEN, E. G. 1938. The habits and life history of the eastern chipmunk,

Tamias striatus Lysteri. New York State Mus. Bull. 314. 122 pp.

BLAIR, W. F. 1940. Home ranges and populations of the jumping mouse.

Amer. Midland Natur. 23:244-250.

BURT, W. H. 1969. Mammals of the Great Lakes region. Univ. Michigan

Press, Ann Arbor. 246 pp.

CHAPMAN, F. B. 1938. Summary of the Ohio gray squirrel investigation.

3rd N. Amer. Wildlife Conf. Trans. : 677-684.

DORNEY, R. S. 1953. Some unusual juvenile raccoon weights. J. Mamm.

34:122-123.

DUNFORD, L. 1972. Summer activity of eastern chipmunks. J. Mamm. 53:

176-180.

EMLEN, J. T., R. L. HINE, W. A. FULLER, and P. ALFONSO. 1957.

Dropping boards for population studies of small mammals. J. Wildlife

Mgt. 21:300-314.

FORBES, R. B. 1966. Fall accumulation of fat in chipmunks. J. Mamm. 47:

715-716.

HAMILTON, W. J. JR. 1935. Habits of jumping mice. Amer. Midland Natur.

16:187-200.

HIRTH, H. F. 1959. Small mammals in old field succession. Ecology 40:417-

425.

HOLMES, A. C. V. and G. C. SANDERSON. 1965. Populations and move¬

ments of opossums in east-central Illinois. J. Wildlife Mgt. 29:287-295.

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SAMPLING AND ANALYSIS OF BOTTOM

SEDIMENTS OF SOME WISCONSIN LAKES

Laverne C. Strieker

and

Robert N. Cheetham, Jr.

U.S.D.A. Soil Conservation Service —

Madison

ABSTRACT

From 1971 to 1973, bottom sediment core samples were taken in

nine Wisconsin lakes. The samples were first described in the field

and then analyzed for physical characteristics, organic content,

and phosphorus.

The watershed and lake characteristics of Big Spring Pond in

Adams County and East Silver Lake, Monroe County, are discussed

in detail.

INTRODUCTION

Bottom sediment samples were collected through the ice and

from a boat with a core sampler developed at the University of

Wisconsin Water Chemistry Laboratory. Nine lakes were sampled:

Big Spring Pond — Adams County

Little Green Lake — Green Lake

County

Lilly Lake — Kenosha County

Angelo Pond — Monroe County

Perch Lake — Monroe County

East Silver Lake — Monroe County

Lake Leota — Rock County

Spauldings Pond — Rock County

Wautoma Pond — Waushara County

The samples were first described in the field and then analyzed

for physical characteristics, organic content and inorganic phos¬

phorus. Analyses were done according to Schulte and Olsen, 1970,

procedures at the Soil and Plant Analysis Laboratory, Department

of Soil Science, University of Wisconsin— -Madison.

The studies consisted of evaluating watershed and lake charac¬

teristics. Based on these findings, recommendations are made for

pond or lake improvement.

Two of the nine lakes were selected to emphasize the extreme

variation of watershed and lake characteristics — Big Spring Pond,

Adams County and East Silver Lake, Monroe County.

BIG SPRING POND

Watershed Information

Big Spring watershed is in New Haven Township, southeastern

Adams County, Wisconsin, about eight miles NE of Wisconsin

327

328 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Dells. The watershed has a drainage area of 1,677 acres, or 2.62

square miles. Big Spring Pond has a surface area of 7.3 acres.

The Big Spring area is topographically mapped on the 15-minute

Briggsville, Wisconsin, Quadrangle, 1958 edition, published by the

United States Geological Survey. The contour interval is 20 feet.

Agricultural Stabilization and Conservation Service (ASCS) con¬

tact aerial photographs, scale 1 inch equals 1,667 feet, AJA-1FF-96

of 7/15/65 and AJA-1FF-96 of 7/15/65 provide stereographic

coverage.

Adams County has a humid continental climate with wide

extremes of temperature. The coldest month is January with an

average temperature of 17 F. July, the warmest month, has an

average temperature of 73 F. The average rainfall is 31 inches

and occurs mainly during the growing season of 130 to 140 days.

The 1968 map Soils of Wisconsin by Hole, et al., indicates a gen¬

eral I 17 association of loams, clay loams, and sandy loams. Typical

soil series are Manawa, Oshkosh, and Poygan. Adams County is a

part of the Wisconsin Central Plain. Big Spring watershed is in

that portion of the county invaded by the Pleistocene Green Bay

ice lobe. Nearly horizontal quartz sandstones of Upper Cambrian

Age are almost completely obscured by soils, alluvium and Pleisto¬

cene pitted outwash and till. To the east of Big Springs are glacial

lacustrine deposits and to the west is the Johnstown Moraine of

Wisconsin Age.

Land use was determined by field reconnaissance and aerial

photographs.

Acres

Cropland - 965

Pasture _ 274

Woodland and Wildlife _ 356

Roads, Buildings, and Other - 82

Big Spring Pond - 7

Total _ 1,684

The Big Spring area was first settled in the fall of 1849 after

treaty was made with the Menomonee Indians. By 1870 the area

was in farmland and culturally accelerated erosion commenced.

With a modified Musgrave equation, a 635-acre sample was used

to determine annual soil loss of cropland.

E = KCR

(%S) 1.35 (LS)

°5

.o

(10)

(72.6)

ACPM

Where :

E is erosion in tons per acre

K is a function of soil type

R is rainfall erosiveness

1974] Strieker and Cheetham — Sampling of Sediments 329

S is slope

LS is length of slope

A is area in acres

C is rotation — crops

P is practice — as contour farming

M is management factor

Eight cooperators with the Adams County Soil and Water Con¬

servation District own 1,231 watershed acres and six non-coopera¬

tors own the remaining 446 watershed acres. This is exclusive of

the 7.3 acre lake.

Present average annual soil loss from cropland, as shown in

Table 1, was estimated at 1.99 tons per acre; 0.12 tons per acre of

soil loss from pasture; 0.12 tons of soil loss from woods and wild¬

life, and 4 tons per acre from roads and from dwelling and barn¬

yard areas. Common cropland rotations are corn, oats, and two,

three, or four years of hay. Streambank erosion and gully erosion,

which are minor, were estimated after field inspection to cause

10% of the delivered upland sheet erosion to Big Spring Pond.

Converting to tons per year based on 100 lbs per cu ft yields an

annual input of 1698.4 tons of sediment from erosion. The aver¬

age soil loss from the drainage area is 1.01 tons per acre per year.

The difference in the current estimated soil loss of 0.48 tons per

acre per year and the determination of 1.01 tons per acre per year

based on loss of pond capacity may be attributed to continuing and

improved land treatment measures during the past 20 years.

TABLE 1. AVERAGE ANNUAL SEDIMENT YIELD BY SOURCES

IN BIG SPRING WATERSHED (PRESENT CONDITIONS)

GULLY EROSION

STREAMBANK EROSION l Est. at 10% of delivered sheet erosion 73

STREAMBED EROSION \

TOTAL 806 Tons

330 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

Pond Information

Big Spring Pond is described by Klick and Threinen, 1966, as a

hard-water drainage lake with 7.3 surface acres. The water is light

brown, alkaline, and has a low transparency. The fishery is large-

mouth bass, bluegills, pumpkinseed, and brook and brown trout.

Game includes muskrat and ducks. Access, without parking facil¬

ities, is near the 11-foot head dam on a town road.

Based on interviews with several residents of Big Springs, it was

determined that in 1951 maximum water depth of the lake was

approximately 6 ft and average water depth 4 ft. Surface area then

was 7.3 acres.

In 1971 surface area was unchanged, but maximum water depth

(save at outlet) was 3 ft and average depth was 2 ft to the water-

solids interface. The lake capacity or volume in 1951 was an esti¬

mated 29.2 acre feet and in 1971 only 14.6 acre feet. Therefore, in

20 years the loss of lake capacity averaged 0.78 acres feet per year.

In order to determine sediment accumulation in Big Spring Pond,

visits were made to the area in January and June 1971. Water qual¬

ity analyses and temperatures were taken in January. In June a

core sample was obtained from a small pier west of the dam. Two

attempts were made for core samples at the north end of the pond

and at the outlet-dam area, with poor recovery because of rock

and gravel fill near the dam. They were discarded.

The recovered sample was extruded from the plastic core barrel

and representative upper, middle, and lower core segments were

collected for analysis with results as given in Table 2.

The visual description of these sediments is as follows:

Sample

0.8' Watery collapsed — brown — black silt — no noticeable odor — modern

plant fibres-roots.

1.0' Uniform brown-black silty sand. No noticeable odor — some organic

content.

0.2' Silt, black, organic — fibres — shells.

0.5' Silt, black, numerous fibres and shell fragments. Some fine sand.

0.2' Silt, black, uniform, tr. sand.

0.1' Silt, black, organic.

1.0' Silt, black, fairly uniform with trace of clay and some find sand. Firm-

poorly stratified. Some organic content.

Interpretation

The sediments in Big Spring Pond are mostly silt and organic

as judged from the core samples take near the center of the lake.

1974] Strieker * and Cheetham — Sampling of Sediments 331

The surface of the sediments is mostly silt and has a very high

inorganic phosphorus level. It is thought this layer of sediments

was transported from the watershed through sheet erosion. A con¬

siderable amount of fertilizer is applied to the cropland in the

watershed each year which may account for the high phosphorus

level. Organic matter derived from plants and algae was not high

at this location.

Below the top layer (2'-2.5') there is a mostly sandy layer. The

sand may have been transported simultaneously with the top silt

but the sand, because of volume weight, settled out more rapidly.

The sand probably originated from streambanks and road ditches

and is not as fertile as the silt. The bottom layer is higher in or¬

ganic matter and inorganic phosphorus. This was probably the

original ground level before construction of the dam. The area was

originally marsh and was inundated at time of dam construction.

EAST SILVER LAKE

Watershed Information

East Silver Lake is in Adrian Township on the Camp McCoy

military reservation, Monroe County, Wisconsin. The lake is 7.5

miles SSW of Tomah. The watershed has a drainage area of

2,080 acres, or 3.25 square miles. The lake has a surface area of

8.5 acres. The East Silver Lake Area is topographically mapped on

the 15-minute Tomah, Wisconsin Quadrangle, 1947 edition, pub¬

lished by the United States Geological Survey. The contour interval

is 20 feet. Agricultural Stabilization and Conservation Service

(ASCS) contact aerial photographs, scale 1 inch equals 1,667 feet,

55081 of 372-18 and 55081 of 372-19 of 10-7-72 provide stereo¬

graphic coverage.

Monroe County has a humid continental climate with wide ex¬

tremes of temperature. The coldest month is January with an

average temperature of 14 F. July, the warmest month, has an

average temperature of 73 F. The average rainfall is 30 inches and

occurs mainly during the growing season of 135 to 145 days.

332 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

The 1968 map of Soils of Wisconsin by Hole, et ah, indicates

a general C-6 association of loamy sand and sand soil series such

as Plainfield, Sparta, Gotham, and poorly drained undifferentiated

alluvium; and, D1 steep rock land, loams, and silt loams whose

typical soil series are Gale, Norden, Hixton, Fayette and Seaton.

Monroe County is a part of the Western Uplands. East Silver Lake

watershed is in the “Driftless” area of the county. East Silver

Lake, fed by springs and seeps, drains westerly to the La Crosse

River, a tributary to the Mississippi River.

Cambrian sandstones, shales, and siltstones crop out in the

watershed but are somewhat obscured by loess, soils, alluvium,

and colluvium. The sandstones outcrop near the upper end of the

lake and are occasionally exposed by road-cuts. The Paleozoic

formations appear almost horizontal except where slumped or

settled. The higher bluffs have a thin Ordovician dolomite cap

rock. Windrow Bluff, about three miles northeast of Silver Lake,

is the type locality of the Windrow high level gravels of Cretaceous

age.

Land use was determined by field reconnaissance and aerial

photographs.

Acres

Cropland _ 2.0

Pasture _ —

Woodland and Wildlife _ 1967.5

Wetland _ 21.0

Roads, Ditches, Buildings, misc. _ 83.0

East Silver Lake _ 6.5

Total _ 2080

There are two landowners (Department of Defense and one

farmer) who are cooperators with the Monroe County Soil and

Water Conservation District. Several non-cooperators have parts

of their farms in the upper reaches of the watershed.

Monroe County was created in 1854. Fruit and berry culture,

dairying, and general farming were practiced. After 1890 the trend

was toward dairying and general farming. Between 1908 and 1910

land was acquired by the Army for a military reservation known

as Camp Robinson. After a varied history it was renamed Camp

McCoy in 1926. From 1933-35 part of the camp was a CCC supply

base of the U. S. Department of Agriculture.

Upland sheet erosion computations were made for two acres of

hayland — on Boone Loamy sand. The land, classed as VIIs2, is on

a 12% slope.

As shown in Table 3, the present average annual soil loss from

cropped land is estimated at 1.6 tons per acre. Soil loss from wood-

1974] Strieker and Cheetham — Sampling of Sediments 333

TABLE 3. EAST SILVER LAKE: AVERAGE ANNUAL SEDIMENT

LOAD BY SOURCES (PRESENT CONDITIONS)

TOTAL 442 tons

land and wildlife is 0.16 tons per acre, and soil loss from roadside

and ditch erosion is 10 tons per acre. Predicted current sediment

yield from gross erosion delivered to East Silver Lake is 0.22 tons

per acre per year. The most conspicuous sediment producing areas

are the numerous roadbanks and ditches adjacent to and above the

lake. Streambank and gully erosion are negligible to severe above

the lake but were not studied separately in detail. Two gullies have

fanned deltas into the lake. An estimate of 10% of the delivered

upland sheet erosion was made for this type of sediment yield.

TABLE 4. ANALYSES OF THE 4 CORES FROM EAST SILVER LAKE

4

334 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

Lake Information

East Silver Lake is described by Klick, Gebken, and Threinen,

1969, as a hard-water drainage impoundment of 6.5 surface acres.

The water is clear, medium hard, alkaline, and has a high trans¬

parency. Located entirely within Camp McCoy, the lake is managed

by the federal government. Largemouth bass, northern pike, crap-

pies, pumpkinseed, brook, brown, and rainbow trout compose the

fishery. Access is by permit.

Sampling and Analyses

Four holes were made through the ice and cores collected as

follows :

Upstream Thickness Bottom

Core From Dam Of Ice Depth Profile Sediment

1 _ 125' 9" 7' 0 — 1' 10" Sand

1' 10"— 2' 6" Silty sand

2' 6"— 3' 8" Sand

2 _ 450' 11" 8. -5" 0 — 7" Sandy silt

7" — 10" Silt and sand

3 450' 9" 8' 0 — 6" Sandy silt

6"— 3' 10" Sand

4 670' 8" 6' 0 —1' 4" Sand, silty

1' 4"— 3' 0" Sand

Interpretation

As shown in Table 4, the sediments in East Silver Lake are

mostly sand and organic with some silt. Along the steep shoreline

the sediments are mostly sandy on top. This is because of recent

sliding of sand from the steep banks and delta deposits that have

fanned out on the lake bottom. Further out, as in cores 2 and 3,

the sediments are silty with higher organic content. At one time

this layer was on the surface of the sediments but now has been

covered by the sliding sand.

The sandy layer has encroached to a depth of around 6 ft in

some areas. The deepest part of the lake appears to be around 9 ft.

The silty organic layer is on the surface of the sediments toward

the center of the lake ; see core 2, Table 4. The silt in this layer

probably originally eroded from agricultural land in the watershed.

Most of this sedimentation occurred immediately after the lake

was formed. Present-day erosion is carrying mostly sand particles

and some silt.

The organic material in this layer is derived from decayed plant

and animal remains within the lake. At times, aquatic plants and

algae are abundant. After severe storms, branches and leaves may

be transported to the lake.

1974] Strieker and Cheetham — Sampling of Sediments 335

The sandy layer is infertile; the silty organic layer is very fer¬

tile. Phosphorus levels mentioned here include only inorganic

phosphorus. If organic phosphorus was included, much higher

readings would be obtained for the organic material.

When the silty organic layer is in contact with water, the phos¬

phorus may be continually recycled. This leads to an excessive

nutrient supply in the waters of the lake and resultant aquatic

plant problems.

PROPOSED IMPROVEMENT

To prevent aquatic plant and other problems, the sediment layer

furnishing nutrients should be either covered or removed.

Big Spring Pond has a very fertile top sediment layer that

should be removed. Improved oxygen condition in the pond would

delay future buildup of this organic layer. Mechanical aeration

is recommended. Releasing the outlet water from near the bottom

of the pond would also delay organic sediment buildup. A new type

of structure is recommended here. Nutrient contribution to the

pond could be lessened through improved land treatment measures.

The sliding sand is covering the edges of East Silver Lake now.

In time, this deposit could cover most of the lake bottom. Problems

of lake shallowness could then occur. It would be best to attempt

to stop the sliding sand where it is now and remove the silty or¬

ganic layer from the center of the lake. This could be done by lake

drawdown and allowing the layer to dry out, or by physical re¬

moval with a dredge or dragline. After removal of the silt organic

layer, better nutrient discharge could be obtained by installing a

bottom release outlet. The Soil Conservation Service has standard

drawings for this type of outlet.

Land treatment measures to control erosion and sedimentation

are needed above both bodies of water. Gully control structures

and diversions should be planned for reducing the amount of sedi¬

ment. The critical areas of roadside erosion should be resloped

where possible, hand-mulched, limed, and seeded to vetch and

grass.

Critical reaches of streambank erosion should be delineated and

erosion control measures applied. The control may be simple or

complex, depending on bank height, channel width, soil materials

in bank profile, and presence or absence of bedrock in the channel.

The majority of eroded material is derived from unstable banks,

cut-banks, bank freeze and thaw, and bank-full waters. Riprap,

deflectors, and bank sloping and seeding are most generally used.

Occasionally debris basins or drop structures are needed in con¬

junction with other measures for streambank protection. While the

336 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

acreage of cropland is very small, some additional land treatment

measures are needed on the rather steep Class VII land, such as

contouring. The most satisfactory treatment would be a changed

land use to woodland and wildlife. Soil losses from wooded areas

are being reduced by complete forest land management.

BIBLIOGRAPHY

Crop and Livestock Reporting Service, Wisconsin State Department of

Agriculture, 1956, Wisconsin Rural Resources , Monroe County, 56 pp.

Crop and Livestock Reporting Service, Wisconsin State Department of

Agriculture, 1957, Wisconsin Rural Resources , Adams County, 58 pp.

HOLE, F. D., M. T. BEATTY, C. J. MILFRED, G. B. LEE, and A. J.

KLINGELHOETS. 1968. Soils of Wisconsin : Univ. Wisconsin, Geo.

Nat. His. Survey, Dept. Soil Science, U.S. Soil Conserv. Serv., and U.S.

Forest Serv. Map 1:710,000.

KLICK, T. A., D. F. GEBKEN, and C. W. THREINEN. 1969. Surface Water

Resources of Monroe County, 119 pp.

KLICK, T. A., and C. W. THREINEN. 1966. Surface Water Resources of

Adams County, 71 pp.

MARTIN, L. 1932. Physical Geography of Wisconsin, Wis. Geo, Survey Bull.

36. (2nd ed.).

Railroad Commission of Wisconsin, 1915, Report of the Railroad Commission

of Wisconsin to the Legislature on Water Powers, Part II, Gazetteer of

Streams, pp. 489-540.

SCHULTE, E. E., and C. C. OLSEN. 1970. Wisconsin Soil Testing and Plant

Analysis Procedures . Dept. Soil Science, Univ. Wisconsin, 45 pp.

Wisconsin Legislative Reference Bureau, 1970, The Blue Book of Wisconsin.

STUDIES ON AQUATIC OLIGOCHAETA IN

INLAND WATERS OF WISCONSIN

Richard P. Howmiller

University California — •

Santa Barbara

INTRODUCTION

Aquatic oligochaete worms, especially the Tubificidae, are com¬

mon in the benthos of many lakes and rivers. In some aquatic

environments, for example in polluted stretches of rivers and the

profundal zone of many lakes, these organisms are often the most

abundant of benthic macroinvertebrates.

Many of the aquatic oligochaetes occupy a niche similar to that

of their terrestrial counterparts, ingesting and burrowing in the

bottom sediments. Through these activities they may have pro¬

found effects upon the functioning of aquatic ecosystems. Under

experimental conditions tubificids have been shown to promote

the release of plant nutrients (Howmiller, unpublished) and pol¬

lutants (Jernelov 1970) from sediments, and to increase the rate

of biochemical oxidation of sediments (Zvetlova 1972).

The worms are preyed upon by numerous invertebrate predators

and by bottom-feeding fishes. Their importance as a food item of

fishes is often underestimated because the worms are so rapidly

digested that only unrecognizable fragments remain in stomach

samples handled in the usual fashion. However, the fact that some

tubificids serve as intermediate hosts of fish parasites (Calentine

and Delong 1966, Calentine and Mackiewicz 1966) attests to their

use as food by several fish species. The value of the worms as fish

food is recognized by tropical fish fanciers who support a small

industry supplying them with “tubifex worms” — the only direct

economic importance of aquatic oligochaetes.

Pollution biologists have long associated an abundance of

oligochaetes, coupled with a scarcity of other benthic invertebrates,

with severe organic pollution. Thus one often sees the worms

collectively referred to as “sludgeworms”. In fact, only a few

species of tubificids are tolerant of gross pollution. Thus, in recent

years, attention has been focused on the possibility of a more sensi¬

tive index of environmental quality based on the relative abun¬

dance of species found to be intolerant, moderately tolerant, and

very tolerant of organic enrichment. This interest has been a pri¬

mary consideration in several studies of oligochaete species’

337

338 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

distributions in waters of the Great Lakes. The worm fauna of

the Great Lakes is now quite well known and it has become clear

that polluted bays and harbors, and moderately enriched areas

such as southern Lake Michigan, have characteristic species as¬

semblages which are distinct from those of the open unpolluted

areas of the lakes. The worm fauna of Great Lakes’ waters border¬

ing on Wisconsin has been studied by Hiltunen (1969a) in the

Apostle Islands region of Lake Superior, Howmiller and Beeton

(1970) in Green Bay, and Hiltunen (1967) and Howmiller (1972)

in Lake Michigan proper. The fauna of these areas includes forty-

one species of Oligochaeta (Howmiller and Beeton 1970).

Despite the importance of aquatic oligochaetes, and the fre¬

quency with which they are encountered by aquatic biologists, the

fauna of most regions is poorly known. Though the aquatic biota

of Wisconsin has probably been recorded more thoroughly than

that of any other state, the literature contains only occasional

records of oligochaetes at the species level. For many years, the

records of Sparganophilus eiseni (= S. tamesis) , Lumbriculus

limosa ( = L. varigatus) , Chaetogaster, Nais, Pristina, Limno-

drilus claparedeianus, and Tubifex tubifex from Lake Mendota

(Muttkowski 1918) have constituted the most complete account

of the aquatic Oligochaeta of Wisconsin. The purpose of the present

communication is to record additional species from Wisconsin and

to give an account of some ecological observations.

RECENT WISCONSIN RECORDS

Oligochaete species identified from my recent collections are

listed in Table 1, in which localities are given as code letters.

Explanation of the code letters, and locations of these lakes and

rivers, are as follows:

YL

LK

LM

SP

TM

GL

LGL

GP

KR

MR

RR

CL

WR

LP

LMa

Code

Yellow Lake

Lake Kegonsa

Lake Mendota

Salmo Pond

Theresa Marsh

Green Lake

Little Green Lake

Grand Portage (Tank) Lake

Kinnickinnic River

Milwaukee River

Root River

Clear Lake

Wisconsin River

Lake Pepin

Lake Mallalieu

Lake or River

County

Burnett

Dane

Dodge

Green Lake

Iron

Milwaukee

Oneida

Pepin

St. Croix

1974]

Howmiller — Studies on Aquatic Oligochaeta

339

Nomenclature used in Table 1 follows that in the recent mono¬

graph of Brinkhurst and Jamieson (1971) except that Limnodrilus

spiralis (Eisen) is here retained as a distinct species. Limnodrilus

spiralis is separated from L. hoffmeisteri and other Limnodrilus

species on the basis of differences in penis sheath morphology.

The sheath of L. spiralis is, on the average, longer than that of

L. hoffmeisteri and has its opening at the tip, surrounded by a flat

plate with one edge upturned (Fig. la). The shorter sheath of

L. hoffmeisteri is usually somewhat curved and is hooded with the

opening at 90° to the shaft (Fig. lb). Brinkhurst (Brinkhurst and

Jamieson 1971) considers L. spiralis a synonym of L. hoffmeisteri

and feels that a separate species should not be erected on the basis

of the differences mentioned above unless the worms can be shown

to be different in some other way. However, these differences are

pronounced and, in my experience, intergrades are rare or non¬

existent. It is difficult to be very positive on this last point since

penis sheathes are often somewhat deformed when the animal is

placed under a coverslip, especially in specimens where the sheath

is just forming. Thus it is possible that a specimen with a sheath

deformed in slide preparation could be mistaken for an intergrade,

or vice versa. Differences in distribution of mature specimens of

the two forms, in a lake in which both occur (Table 2), suggest

that they are ecologically different in addition to being morpho¬

logically distinct.

Hiltunen (1967) found L. spiralis in Lake Michigan, but con¬

sidered it a variant of L. hoffmeisteri. He has since recorded

elsewhere1 that L. spiralis is discontinuously widespread in the

Great Lakes region, in mesotrophic and eutrophic habitats, and

that specimens intermediate between it and typical L. hoffmeisteri

are infrequently observed.

The question may merit further taxonomic investigation, but

at present it seems appropriate to retain L. spiralis as a distinct

species.

1 Hiltunen, J. K. 1973. A Laboratory Guide; keys to the tubificid and naidid

Oligochaeta of the Great Lakes region. Unpubl. ms. 24 pp.

340 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

TABLE 1. AQUATIC OLIGOCHAETA IDENTIFIED FROM RECENT

COLLECTIONS AND LOCALITIES AT WHICH THEY WERE FOUND

(FOR EXPLANATION OF CODE LETTERS SEE TEXT). SPECIES

REPORTED BY MUTTKOWSKI (1918) FROM LAKE MENDOTA

ARE MARKED WITH A SINGLE ASTERISK. ALL OTHERS

ARE NEW RECORDS FOR INLAND WATERS OF WISCONSIN

Species

Lumbriculidae

Lumbriculus variegatus (Verrill)*

Localities

SP, CrL

Naididae

Arcteonais lomondi (Martin)

Aulophorus vagus Leidy

Dero digitata (Muller)

Haemonais waldvogeli Bretscher

Nais elinguis (Muller)

Nais simplex Piguet

Ophidonais serpentina (Muller)

Stylaria fossularis Leidy

Stylaria lacustris (Linnaeus)

LMa, DL

TM

TM, LMa, DL, LN, FR

TM

LN

TM, LMa, DL, IFR

LGL, DL

TM, CL, LMa

Tubificidae

Aulodrilus americanus Brinkhurst & Cook

Aulodrilus limnobius Bretscher

Aulodrilus pigueti Kowalewski

Aulodrilus pluriseta (Piguet)

Branchiura sowerbyi Beddard

Ilyodrilus templetoni (Southern)

Limnodrilus cervix Brinkhurst

Limnodrilus claparedeianus Ratzel*

Limnodrilus hoffmeisteri Claparede

Limnodrilus prof undicola (Verrill)

Limnodrilus spiralis ( E isen ) * *

Limnodrilus udekemianus Claparede

Peloscolex multisetosus multisetosus (Smith)

Peloscolex multisetosus longidentus Brinkhurst

& Cook

Potamothrix hammoniensis (Michaelsen)

Potamotlvrix moldaviensis (Vejdovsky &

Mrazek)

Tubifex kessleri americanus Brinkhurst &

Cook

Tubifex tubifex (Muller)*

YL

YL, LMa, DL

YL, LMa, DL, LW

DL

LP, LG

YL, LM, TM, LGL, GP,

LMa, DL, CrL, TL, FR

LD, FR

LGL

YL, LK, LM, TM, GL, LGL,

GP, KR, MR, RR, WR, LP,

LMa, DL, CrL, TL, LD, LG,

PL, IFR, LW, FR

GL

LG

RR, LMa, LG, IFR, FR

LMa, LG, FR

KR

LG

GLY, LMa, DL

GL

GL, KR, MR, WR, LMa, LG,

FR

** see text

1974]

Hoivmiller — Studies on Aquatic Oligochaeta

341

FIGURE 1. Illustrating differences in form of the penis sheath in (a)

Limnodrilus spiralis and (b) L. hoffmeisteri. Drawn from Lake Geneva

specimens.

The purpose of listing localities in Table 1 is not to suggest the

existence of discrete species’ distributions within the state but

to indicate the degree of commonness or rarity of each of the

species. In this regard it must be pointed out that there was some

bias in sampling as most samples were taken in deeper waters

free of macroscopic vegetation. The tubificids are thus more ade¬

quately represented than the naidids which prefer weedy littoral

situations. However, many readers will recognize that the waters

sampled represent a considerable variety of environmental condi¬

tions ; from nearly oligotrophic Crystal Lake to the very eutrophic

conditions of Lakes Winnebago and Delavan, and from the very

shallow Theresa Marsh to Green Lake, the deepest lake in the

state.

OLIGOCHAETA AND LAKE TYPES

Introduction

There is, of course, always an academic interest in defining the

sorts of environments in which particular organisms are found.

In recent decades there has been an increasing interest in asking

342 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 2. ABUNDANCE OF TUBIFICID WORMS, AS INDIVIDUALS

PER SQUARE METER, AT FIVE DEPTHS ALONG A TRANSECT

IN LAKE GENEVA. ESTIMATES ARE BASED ON A SINGLE

(15 X 15 CM) EKMAN GRAB SAMPLE AT EACH DEPTH

EXCEPT FOR THE 27.5 M STATION WHERE FOUR

SUCH SAMPLES WERE TAKEN. SEVERAL SAM¬

PLES TAKEN FROM DEPTHS OF 36.5 AND

43.5 M CONTAINED NO MACROSCOPIC

INVERTEBRATES

Depth, meters

this question in a slightly different way, viz; Which species of

organisms are typically found in a particular type of environment?

Adequate knowledge of this sort may allow us to use organisms

as indicators of environmental quality. For example, changes in

composition of the biota have provided useful indices for docu¬

menting the eutrophication of lakes (Brinkhurst 1969, Hooper

1969).

The bottom fauna of littoral areas has only limited value as an

index of the trophic state of a lake, because littoral areas are

strongly influenced by edaphic factors and often receive consider¬

able inputs of allochthonous detritus. The profundal region, on

the other hand, offers relative uniformity of environmental con¬

ditions. In many respects the conditions are related to the produc¬

tivity of the overlying water. Chief among these are the quantity

and quality of organic matter and the dissolved oxygen concen¬

tration (Jonasson 1972). Consequently, most attempts to correlate

lake trophy and bottom fauna have been concerned with the pro¬

fundal benthos.

Earlier work of this sort focused upon the larvae of chironomid

midges (Thienemann 1922, Deevey 1941, Brundin 1958). Diifer-

1974] Howmiller — Studies on Aquatic Oligochaeta 343

ences found, at the generic level, resulted in the formation of a

“bottom faunistic lake type system” which seemed to have wide¬

spread applicability (Brundin 1958), although the relationship

between productivity and bottom fauna was often complicated by

morphometric influences (Brundin 1949).

Brinkhurst (1964) investigated the oligochaete fauna of a num¬

ber of lakes in the English Lake District with this same question

in mind. His results are a bit difficult for the reader to evaluate as

species abundances are indicated merely as absent, present or

abundant. Also, he presents no data to indicate how one might

rank the lakes investigated using other criteria, saying only that

they “have been held to represent a series according to the classical

taxonomy of lakes”. He concluded that his findings indicated

several species to be common in a wide variety of lakes. Thus,

while Peloscolex ferox seemed to be characteristic of less produc¬

tive lakes and Euilyodrilus (Potamothrix) hammoniensis was in¬

creasingly common in more productive lakes, the data offered no

more quantitative relationship between lake type and worm fauna

composition.

In view of the previously mentioned studies on the 'Great Lakes,

which have found species assemblages differing greatly under

different environmental conditions, Brinkhurst’s (1964) results

seemed surprising. There seemed a possibility that relationships

between worm faunal composition and lake type might be found

elsewhere, where perhaps, there was a richer fauna to offer more

ecologically specialized worm species. Thus the present study was

undertaken.

Procedure

The twenty-six lakes chosen for investigation included all those

studied by Lueschow and co-workers (1970). They measured a

number of parameters indicative of trophic status of the lakes

and calculated a “composite rating” which allowed a ranking of

the lakes from most oligotrophic to most eutrophic. The other

lakes included in the present investigation, with the exception of

Devils Lake, were among those studied by Hilsenhoff and Narf

(1968). Hilsenhoff and Narf measured various environmental

factors thought to be important in regulating chironomid popula¬

tions but did not rate the lakes as oligotrophic or eutrophic.

Environmental parameters measured during the investigations

reported here included depth profiles of temperature and dissolved

oxygen, transparency, color, conductivity, alkalinity, seston, and

organic and carbonate content of sediments.2 The data, particularly

3 A detailed account of methods and limnological data comprises an August 1972

progress report to the Wisconsin Department of Natural Resources.

344 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

transparency, color, organic seston and total phosphorus values,

were used to arrange the lakes in a trophic series (Fig. 2). Those

lakes studied by Lueschow and co-workers (1970), are listed here

in the same order as ranked by them on the basis of their “com¬

posite rating”. Thus the data collected during the present investi¬

gation have essentially been used only to place the other lakes in

their series. The order in Fig. 2 is thus meant to indicate trophic

status, with the more oligotrophic lakes at the top of the list, and

the most eutrophic at the bottom.

Samples for study of the oligochaete fauna were taken at some

distance from shore, usually from a depth near maximum depth.

In one instance (Lake Geneva, See Table 2) samples from a shal¬

lower depth, still well within the profundal zone, were substituted

when deeper samples were found devoid of macroscopic organisms.

Four replicate samples were taken at the single sampling site

in each lake using a 15 x 15 cm tall form Ekman grab. Samples

were sieved immediately with a U.S. Std No. 30 screen (0.565 mm)

and the residue from the screen preserved in 10% formalin. All

organisms were picked from the preserved residue by hand, with

the aid of low power magnification. Oligochaete worms were

mounted whole on microscope slides with Turtox CMC or Amman’s

LAKE

TRANSPARENCY

0.5 1

I - T

5 10

“I - 1

CRYSTAL

DEVILS

GREEN

GENEVA

TROUT (N)

TROUT (S)

ROUND

PINE (WAUKESHA)

MIDDLE

OCONOMOWOC

PLEASANT

RHEINHARDT

BOOTH

PINE (ONEIDA)

LIME

EAST HORSEHEAD

ONEONTA

GRAND PORTAGE

MENDOTA

PEWAUKEE

PEPIN

YELLOW

LITTLE GREEN

KEGONSA

DELAVAN

WINNEBAGO

mm

Em

E3ESEEZ S3

mmm.

MM

33

m

COLOR

V X XV XX

l - r-

m3

■vV^&!lS3

mmm mm

mssmmm

mmxmm

isillili

mmmsmw

ORGANIC SESTON

0.5 1 2 5 15

0

33

33P3]

mmm

mm

mm®

wmmimmm

mmm&immmsm

PHOSPHORUS

10 50

200

mm

B!1

m

mmm

>;i T.'V'

mmmmsm

FIGURE 2. Illustrating- values of some limnological parameters for twenty-

six Wisconsin lakes. Transparency is in meters and was measured with a

20 cm all-white Secchi disc. Color was determined by comparison with a

Forel-Ule scale. Organic seston data are given in mg/liter and were ob¬

tained by measuring loss on ignition of material on Whatman GF/C filters.

Total phosphorus values are given in Mg/liter; the samples were taken from

a depth of 1.0 m and determinations made by the method of Schmid and

Ambuhl (1965).

1974] Howmiller — Studies on Aquatic Oligochaeta 345

Lactophenol or a mixture of the two. They were examined after

waiting several weeks for clearing of soft tissues.

Results and Discussion

Samples from the profundal of nine of the lakes (Round, Pine

in Waukesha County, Middle, Oconomowoc, Booth, Lime, Oneonta,

Pewaukee, and Delavan) contained no oligochaete worms. Note

that while these lakes without worms fall in a range which may

be described as mesotrophic to very eutrophic (Fig. 2), the six

most oligotrophic lakes all contained oligochaetes in the profundal

sediments.

All the oligochaetes found In these profundal samples belonged

to the family Tubificidae. Table 3 lists the abundance of individual

taxa for each of the seventeen lakes from which worms were col¬

lected. Absolute abundance is given as individuals per square

meter, based on the mean of counts from the four Ekman grab

samples. The standard error of the mean is also given in the table.

Since in all cases the number of samples was four, the standard

deviation is twice the standard error. The relative abundance of

each taxon, as a percentage of the total number of tubificids, is

also given In Table 3.

Before further reviewing Table 3, it should be pointed out that

many common tubificids can be positively identified only when

they are sexually mature and bearing sexual structures (e.g.

genital chaetae or penis sheathes). Thus, in almost every case, it

was impossible to identify many of the worms in the samples.

These are listed as undetermined immatures and placed into two

groups ; those which possessed hair chaetae and those lacking hair

chaetae. Knowing which species are present in the samples, from

positive identifications of mature specimens, allows one to make

a reasonable guess at the identity of immature specimens. Thus, in

Table 3, I have listed probable abundance of Limnodrilus hoff-

meisteri; this number including L. hoffmeisteri and a portion or

all of the immature worms without hair chaetae. In cases where a

portion of the immatures without hair chaetae could have belonged

to another species (e.g. L. prof undicola in Green Lake) the im¬

matures were assigned to the two species on the basis of relative

abundance of positively identified mature specimens. In the same

manner, undetermined immature specimens without hair chaetae

were included among probable numbers of Ilyodrilus templetoni or

Tuhifex tubifex depending on which was found as mature speci¬

mens in the particular lake. In one case (Lake Geneva) only a

portion of the immatures with hairs was included among probable

Tubifex tubifex because these could also have included immature

Potamothrix hammoniensis .

346 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

This manipulation of data no doubt involves some error since

it assumes that all species populations included some mature in¬

dividuals. This is obviously not true, since many lakes contained

immatures with or without hairs and no mature specimens which

fit the category. Also, assignment of immatures to two species

was done with the assumption that mature specimens constituted

the same proportion of both populations. This assumption, too,

is certainly in error to some extent. Unfortunately, there is no

way to assess the magnitude of error introduced by these assump¬

tions. As long as there is no way to reliably identify immature

specimens and we lack life history information which might indi¬

cate the normal proportion of mature specimens under given

environmental conditions, calculations of the sort described above

will provide the only means of estimating the number of species

in a lake and the relative abundance of each.

In these seventeen lakes, mean abundance of tubificids ranged

from 11 individuals / m2 in South Trout and Pine (Oneida County)

to over 27000/m2 in Green Lake (Table 3). The standard errors

associated with these means are in many cases quite large, but

small enough to leave no doubt that there are differences of

several orders of magnitude in the abundance of tubificids in these

lakes. It is clear that the four most oligotrophic lakes were the

four in which oligochaetes were most abundant, but there was

no consistent relationship between worm abundance and lake tro¬

phy when all seventeen lakes were considered (compare Fig. 2,

Table 3) .

It is apparent from Table 3 that Limnodrilus hoffmeisteri was,

by far, the most common worm in the lakes. Of the seventeen lakes

having worms in the profundal benthos, this species occurred in

twelve (in thirteen if one includes L. spiralis in L. hoffmeisteri) .

It was clearly numerically dominant in most of these lakes (Ta¬

ble 3). This was not unexpected. L. hoffmeisteri is cosmopolitan

and is generally the most common worm in any region (Brinkhurst

and Jamieson 1971). It has been found to be the most abundant

worm in studies of many types of environments, though in extreme

cases this has often been associated with organic pollution (Brink¬

hurst 1969).

Ilyodrilus templetoni was the second most common and abundant

species. It was represented by mature specimens in six lakes and

probably occurred among the immature specimens of five other

lakes (Table 3). In Crystal Lake and Lake Mendota, I. templetoni

accounted for a considerable proportion of the worm fauna and

this may have been true of several other lakes (South Trout,

Rheinhardt, Pine in Oneida County, East Horsehead; Table 3).

1974]

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1974]

H ow miller — Studies on Aquatic Oligochaeta

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350 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

These findings contrast sharply with Brinkhurst’s (1964) data

from British Lakes which indicate that I. templetoni (as Tubifex

templetoni) occurred in few lakes and was never abundant. How-

miller and Beeton (1970) found I. templetoni common in a portion

of Green Bay but not so above the latitude of Sturgeon Bay or

below Long Tail Point near the mouth of the Fox River. The

species thus seems to be favored by moderate organic enrichment

but intolerant of gross pollution.

Tubifex tubifex, third most common worm in these profundal

samples, occurred in only two lakes but there comprised most of

the worm fauna (Table 3). It may be of significance that the two

lakes in which it occurred, Green and Geneva, are at the oligo-

trophic end of the series and are the deepest of these lakes. Tubifex

tubifex, in puzzling contrast with its well-deserved reputation for

tolerance of severe organic pollution in rivers and harbors, has

frequently been reported as a relatively abundant species in the

profundal of oligotrophic lakes. Brinkhurst (1964) reported that

it alone occurred in material from high alpine lakes of Austria

and this seems to be true as well of some oligotrophic sub-alpine

lakes in the Sierra Nevada (Howmiller, unpublished). Tubifex

tubifex occurs in the profundal benthos of the oligotrophic upper

Great Lakes (Hiltunen 1967, Howmiller 1972) and in the less pro¬

ductive regions of the lower lakes (Brinkhurst and Jamieson

1971).

The three species just discussed, L. hoffmeisteri f /. templetoni,

and T. tubifex, are the only common oligochaetes in the profundal

region of these Wisconsin lakes. While six other species were

found in the present series of samples, each occurred in only a

single lake (Table 3). These lakes are thus at least as poor in

species as those of the English Lake District investigated by

Brinkhurst (1964). Of the twenty lakes which he sampled, four

yielded no worms, seven contained only one species, six contained

two species, and three had three species. Of the five species re¬

corded from these English lakes ; Peloscolex ferox, Aulodrilus

pluriseta, Tubifex tubifex, Ilyodrilus templetoni, and Limnodrilus

hoffmeisteri, only the latter occurred in more than half the lakes.

Aulodrilus pluriseta and T. tubifex were found in only two lakes

each.

The paucity of species in the profundal of inland lakes contrasts

strongly with findings in bays, harbors, and littoral areas of the

Great Lakes. For example, thirty species of Oligochaeta, including

nineteen tubificids, have been reported from Green Bay. As pointed

out by Dahl (1970), there are many possibilities for overland dis¬

persal of oligochaetes. The occurrence of only a few species in

1974] Howmiller — Studies on Aquatic Oligochaeta 351

the profundal of these Wisconsin lakes, considering their close

proximity to the rich fauna of the Great Lakes, thus indicates that

in terms of the requirements of oligochaete species, the profundal

regions of inland lakes constitute a set of very similar environ¬

ments. The corollary of this is that composition of the profundal

oligochaete fauna has no index value for distinguishing between

lakes within the range offered by the series investigated. There

does seem some tendency for more oligotrophic lakes to be less

likely to lack worms in profundal regions, and perhaps T. tubifex

occurs in abundance in the profundal zone only in more oligo¬

trophic lakes. However, it would seem to require an investigation

of much wider scope to determine whether these are valid

generalizations.

VERTICAL DISTRIBUTION OF OLIGOCHAETA IN LAKES

While the investigation just described was concerned primarily

with worms of profundal regions, samples were taken at several

shallower depths in two of the lakes. These samples were examined

for evidence of depth differences in composition of the worm fauna,

as has been reported for several British and European lakes

(Brinkhurst 1964).

Benthic samples were taken from seven depths in Lake Geneva

on a single date (Table 2). The samples from the two deepest

stations (36.5 and 43.5 m) contained no macroscopic invertebrates.

This is no doubt a result of prolonged anoxia in the deepest part

of the basin. Vertical zonation of tubificid species was suggested

by composition of the samples from the five lesser depths.

Two species, Branchiura sowerbyi and Limnodrilus udekemianus

occurred only in the shallowest water, from 12.5 m in depth. Two

others, L. hoffmeisteri and Peloscolex multisetosus, reached their

maximum abundance in this sample. In addition to these four

species, the sample contained immature specimens of at least one

other. The most probable number of species was considered to be

five, more than at any greater depth.

A sample from 17.5 m depth contained typical L. hoffmeisteri as

well as L. spiralis. Of these, only L. spiralis was represented among

mature specimens at greater depths. The 17.5 m sample also con¬

tained mature Potamothrix hammoniensis and T. tubifex and was

dominated by unidentifiable immature specimens representing

mostly or entirely T. tubifex.

At 21.0 meters the sample was dominated by immatures without

hairs. The mature specimens in the sample can be assigned only

to L. spiralis. Potamothrix hammoniensis, Peloscolex multisetosus

352 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

and a relatively small number of immatures resembling T. tubifex

were also found at this depth.

The sample from 24.5 m depth contained still fewer immatures

with hairs and was dominated by mature L. spiralis.

Samples from 27.5 m indicated a fauna dominated by T. tubifex;

the immatures with hairs are believed to represent mostly this

species. Small numbers of L. spiralis and P. hammoniensis were

also found (Table 2).

This series, while inadequate in that it represents only a single

transect with single samples from most depths, provides evidence

of vertical differences in composition of the worm fauna in a single

lake. It is also clear that the paucity of species in profundal sam¬

ples is due to the special conditions there and does not apply to

the lake as a whole. While this series from Lake 'Geneva included

at least seven species of tubificids only 3 were found at 27.5 m,

the depth taken to represent the profundal (Table 3). Lastly, we

may note that the proportion of mature individuals in some popu¬

lations varies with depth ; with numbers of mature L. hoffmeisteri

exceeding immature specimens at 12.5 m, mature Limnodrilus spp.

much less numerous than immatures at 17.5 and 21.0 m, and ma¬

ture L. spiralis considerably more abundant than immatures at

24.5 m. A careful analysis of a situation of this sort through the

course of a year could indicate more precisely the proportion of

immatures attributable to each taxon and indicate conditions

conducive to maturation and breeding of the species.

Samples taken in Devils Lake at three depths on several dates

further illustrate vertical zonation of oligochaetes as well as sea¬

sonal variations in the abundance of some species. Samples taken

on two dates at two nearshore stations (Table 4) included variable

numbers of three species of naidids. Doubtless, a series of samples

taken throughout the summer and winter would have reflected

greater fluctuations in abundance. Muttkowski (1918) noted strong

seasonal variation in abundance of naidids in Lake Mendota, with

higher population densities associated with seasonal development

of the littoral macrophytes.

Aulodrilus pluriseta in Devils Lake also seemed to undergo a

seasonal change in abundance (Table 4). This is, in the extremes

indicated here, unusual for a tubificid but apparently not unusual

for A. pluriseta. In a series of samples taken monthly in two

British lakes, Bala and Windermere, Brinkhurst (1964) found

strong seasonal fluctuations in abundance of A. pluriseta, with

maxima in mid- to late summer. He reported that very few sexu¬

ally mature specimens were found and suggested that asexual

1974]

Howmiller — Studies on Aquatic Oligochaeta

353

TABLE 4. ESTIMATES OF ABUNDANCE OF AQUATIC

OLIGOCHAETES, AS INDIVIDUALS PER SQUARE

METER, AT TWO NEARSHORE STATIONS ON

DEVILS LAKE

reproduction was occurring. In Bala Lake more breeding specimens

were found, especially in late summer and this may be the time of

a single annual period of sexual activity (Brinkhurst 1964). No

breeding specimens were observed in the Devils Lake samples and

the seasonal fluctuation is thus believed to be the result of rapid

asexual reproduction.

On several dates naidids appeared in samples taken at a mid-lake

station with a depth over 13 m. Maximum numbers were observed

here in fall (Table 5), and may reflect active or passive movement,

or both, of the worms from littoral areas as macrophytic vegeta¬

tion died down. Most, if not all, naidids are capable of swimming.

Of the three Aulodrilus species found at the nearshore station,

only A. pigueti was recorded from mid-lake (Table 5). To the best

of my knowledge, swimming has not been recorded for Aulodrilus

and dispersal may thus not occur as readily as with naidids.

Again, the full richness of oligochaete fauna of the lake was not

reflected in profundal samples. While samples from Devils Lake

have revealed six species of tubificids (Tables 4, 5) only three have

been found at the mid-lake station (Table 5).

354 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 5. ESTIMATES OF ABUNDANCE OF AQUATIC

OLIGOCHAETES, AS INDIVIDUALS PER SQUARE

METER, AT A MID-LAKE STATION ON

DEVILS LAKE

Date

SUMMARY

This paper presents records of twenty-eight taxa of aquatic

Oligochaeta from the inland waters of Wisconsin. Most of these

were not previously reported from the state.

An investigation of the profundal oligochaete fauna in a series

of lakes revealed no consistent correlation of abundance or species

composition with lake type. In general, the profundal worm fauna

is species poor; species other than Limnodrilus hoffmeisteri, Ilyo-

drilus templetoni, and Tubifex tubifex are seldom found.

Samples taken from shallower depths in two lakes provided evi¬

dence of vertical differences in composition of the worm fauna,

with littoral samples containing species not found in the profundal

benthos.

ACKNOWLEDGEMENTS

Thanks are due to Dr. Robert Calentine for allowing me to ex¬

amine his specimens from Lake Mallalieu, and to Ted Ringger who

provided samples from Theresa Marsh.

1974]

Howmiller — Studies on Aquatic Oligochaeta

355

Professors A. M. Beeton and A. D. Hasler assisted greatly by

making available facilities of the Center for Great Lakes Studies

and the Trout Lake Biological Station, respectively.

These investigations were given financial support through a

grant from the Wisconsin Department of Natural Resources.

LITERATURE CITED

BRINKHURST, R„ 0. 1964. Observations on the biology of lake-dwelling

Tubificidae. Arch, HydrobioL 60:385-418.

— - - . 1969. Changes in the benthos of Lakes Erie and Ontario. Bull.

Buffalo Soc. Natur. Sci. 25:45-65.

- — and B. G. Jamieson. 1971. The Aquatic Oligochaeta of the World.

Univ. Toronto Press, Toronto and Buffalo, xi -f- 860 pp.

BRUNDIN, L. 1949. Chironomiden und andere Bodentiere der Sudschwe-

dischen Urgebirgseen. Inst. Freshw. Res. Drottningholm Kept. No. 30:1-

914.

— — - . 1958. The bottom faunistical lake type system and its application

to the southern hemisphere. Verb. Internal Verein. Limnoh 13:288-297.

CALENTINE, R. L., and B. L. DeLONG. 1966. Archigetes sieboldi (Cestoda:

Caryophyllaeidae) in North America. J. Parasitol. 52:428-431.

- - and J. S. MACKIEWXCZ, 1966. Monobothrium ulmeri n. sp. (Ces¬

toda: Caryophyllaeidae) from North American Catostomidae. Trans.

Amer. Microsc, Soc. 85:516-520.

DAHL, I. 1970. Borsteorme (Oligochaeta) fra indvande i Thy. Flora og

Fauna 76:49-65.

DEEVEY, E. S. 1941. Limnological studies in Connecticut VI. The quantity

and composition of the bottom fauna of thirty-six Connecticut and New

York lakes. EeoL Monogr. 11:413-455.

HXLSENHOFF, W. L., and K P. NARF. 1968. Ecology of Chironomidae,

Chaoboridae, and other benthos in fourteen Wisconsin lakes. Ann.

Entomol. Soc. Amer. 61:1173-1181.

HILT UN EN. J. K. 1967. Some oligochaetes from Lake Michigan. Trans. Amer.

Microsc. Soc. 86:433-454.

- . 1969a. Invertebrate macrobenthos of western Lake Superior. Michi¬

gan Acad. 1:123-133.

- - - — •. 1969b. Distribution of oligochaetes in western Lake Erie, 1961.

Limnoh Oceanogr. 14:260-264.

HOOPER, F. F. 1969. Eutrophication indices and their relation to other

indices of ecosystem change, p. 225-235 In Eutrophication: Causes, Con¬

sequences, Correctives. Nat. Acad. Sci., Washington, D.C.

HOWMILLER, R. P. 1972. The oligochaete fauna of central Lake Michigan,

p. 58-62 In R. P. Howmiller and A. M. Beeton. Report on a Cruise of

the R/V Neeskay in Central Lake Michigan and Green Bay, 8-14 July

1971. Wisconsin- — Milwaukee, Center for Great Lakes Studies Spec. Rept.

No. 13.

• - and A. M. BEETON. 1970. The oligochaete fauna of Green Bay,

Lake Michigan. Proc. 13th Conf. Great Lakes Res. pp. 15-46.

JERNELOV, A. 1970. Release of methyl mercury from sediments with layers

containing inorganic mercury at different depths. Limnoh Oceanogr. 15:

958-960.

JONASSON, P. M. 1972. Ecology and production of the profundal benthos

in relation to phytoplankton in Lake Esrom. Oikos Suppl. 14:1-148.

356 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

LUESCHOW, L. A., J. H. HELM, D. R. WINTER and G. W. KARL. 1970.

Trophic nature of selected Wisconsin lakes. Trans. Wisconsin Acad. Sci.

Arts Lett. 58:237-264.

MUTTKOWSKX, R. A. 1918. The fauna of Lake Mendota: a qualitative and

quantitative survey with special reference to the insects. Trans. Wisconsin

Acad. Sci. Arts Lett. 19:374-482.

SCHMID, M., and H. AMBUHL. 1965. Die Bestimmung geringster Mengen

von Gesamtphosphor im Wasser von Binnenseen. Schweiz. Z. Hydrol.

27:184-192.

THIENEMANN, A. 1922. Biologische Seetypen und die Grundig einer hydro-

biologischen Anstalt am Bodensee. Arch. Hydrobiol. 13:347-370.

ZVETLOVA, L. I. 1972. Oligochaeta in the oxygen balance of reservoirs. Proc.

Symp. Aquatic Oligochaeta in Tartu, 1967. pp. 118-125. (Russian, English

Summary) .

A HISTORY OF THE VEGETATION OF

EAU CLAIRE COUNTY, WISCONSIN

William J. Barnes

University Wisconsin —

Eau Claire

ABSTRACT

The General Land Survey records of ca. 1850 were used to

reconstruct the vegetation of presettlement Eau Claire County.

About 80% of the county was prairie, oak opening or barrens,

while only 20% was forest. The prairies and oak openings occurred

primarily south and west of the tension zone, while the barrens

occurred on the sandy terraces of the Eau Claire River. The

pineries and hardwood forests occurred for the most part at the

eastern end of the county. Field samples and aerial photographs

were used to note changes which have occurred over the last 120

years. Farmland and oak forest now occur where previously oak

opening and prairie existed. This change is due to the relative rich¬

ness of the soil and to the cessation of periodic fires. The forests

have changed from being dominated by pine to having red oak and

aspen as the most important species. The composition of the bar¬

rens has not changed greatly over the years, although the character

of this vegetation type has changed from that of an opening of pine

and oak to a closed forest.

INTRODUCTION

The first permanent settlers in the county came to what is now

the city of Eau Claire in 1845. Eau Claire County was established

in 1856, with a population of several hundred people. The majority

of the first settlers were associated with the lumber mills at the

confluence of the Eau Claire and Chippewa Rivers. The population

grew rapidly during the early logging days, so that there were

about 22,000 people in the county at the climax of the lumbering

era, about 1900. Most of the early farms were located in the south¬

ern and western parts of the county, and agriculture changed from

wheat, barley and rye to primarily dairying at about the turn of

the century.

The character of the vegetation was described in early local

newspaper accounts as, “The county is finely diversified with

streams, timber, prairie and oak openings”, and,

357

358 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

The lands are mostly prairie, slightly rolling, and interspersed with an

abundance of fine timber. We (City of Eau Claire) are located within 15

miles of the inexhaustible pineries, one lying on the Chippewa River, the

other on the Eau Claire River.

One article written by a visitor to the city of Eau Claire includes

the statement, “Your great want seems to be a lack of timber in

the immediate vicinity”, while other accounts indicate that some

of the southern and western parts of the county were covered by

“brush”.

DESCRIPTION OF PRESENT EAU CLAIRE COUNTY

Eau Claire County is located in the west central part of the state

at about 91° W, 45° N. It is rectangular in shape, being 18 miles

from north to south (T25N to T27N) and 36 miles from east to

west (R5W to R10W), a total of 648 square miles. The south¬

western part of the county is quite broken and is part of the

western upland, while the rest of the county is rolling and is part

of the central plain (Fig. 1). The extreme southwestern portion

of the county lies in the driftless area, while all of the remainder

is covered by Illinoian drift (Martin, 1932). Elevations within

the county range from 1000 to 1100 feet above sea level in most

places, except in the lower terraces of the Chippewa and Eau

Claire Rivers where they are about 800 to 900 feet above sea level.

Eau Claire County is covered by a deep layer of Mt. Simon and

Eau Claire sandstone of Cambrian age. Glacial deposits cover all

FIGURE 1. Geological provinces and glaciation in Eau Claire County.

Vertical lines indicate the Driftless Area, the remainder is older Drift.

Horizontal lines indicate the Western Upland, the remainder is the Central

Plain.

1974] Barnes — History of Vegetation , Eau Claire County 359

but the driftless area and river valleys, and a layer of loess of

varying thickness occurs throughout the county. About 85 soil

types are represented in the county, most of which are sands,

loamy sands, or silty sands.

Eau Claire County is of particular interest from a vegetational

viewpoint, as the tension zone lies across the county from the

northwest to the southeast (Fig. 2). The southern and western

parts of the county belong to the prairie-oak forest province of

southern Wisconsin, while the extreme northeastern part of the

county is a part of the mixed conifer-hardwood province of north¬

ern Wisconsin. A major part of the county lies within the transi¬

tional zone between these two floristic provinces.

Climatic differences within the county are not great, although

there is a tendency for the northeastern part to be cooler and more

moist than the southwestern part for much of the year (Collins,

1968). Mean annual precipitation is 32 inches, the average date of

the last killing frost is May 4, and the mean length of the growing

season is 150 days (Barland, 1965).

METHODS

The species composition of the several vegetation types present

at the time of the land survey (1847-1853) was determined by

compiling data from photocopies of the original field notebooks

of the surveyors, on file at the County Clerk's Office. Details of the

type of record made by the surveyors can be found in Cottam

(1949) and Bourdo (1956). The methods used for making quali¬

tative and quantitative analyses of the surveyors' records and for

360 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

preparing a vegetation map are described by Cottam (1949),

Bourdo (1956) and Stroessner and Habeck (1966). In some cases

the common names used for the species of trees were different

from the names today. These are listed below as interpreted in this

study.

Pitch pine — Finns banksiana Lamb, (jack pine)

Black pine — Finns banksiana Lamb, (jack pine)

Spruce pine — Pinus banksiana Lamb, (jack pine)

Yellow pine — Finns resinosa Ait. (red pine)

Black oak — Qnercns velutina Lam. (black oak)

or

Qnercns ellipsoidalis Hill (Hill’s oak)

or

Qnercns borealis Michx. (red oak)

Jack oak — Qnercns ellipsoidalis Hill (Hill’s oak)

Water beech — Carpinus caroliniana Walt, (blue beech)

White maple — Acer rub rum L. (red maple)

Sugar maple — Acer saccharnm Marsh, (sugar maple)

Lind — Tilia americana L. (basswood)

Ironwood — Ostrya virginiana K. Koch, (hop hornbeam)

White walnut — Juglans cinerea L. (butternut)

Seven deputy surveyors were involved in surveying Eau Claire

County over a 6 year interval. Thus a species may have been given

more than one common name. Several cases of two or more species

given the same common name occur in the surveyors’ records.

None of the surveyors differentiated between Popnlns tremuloides

and P. grandidentata, both were simply called “aspen”. However,

this does not represent a serious limitation to interpretation of the

vegetation, as the ecological behavior of these two species is very

similar. However, the name “black oak” was apparently applied

to Qnercns velutina, Q. ellipsoidalis and Q. borealis. Since these

three species of oak have quite different ecological behaviors, the

failure of the surveyors to differentiate does present a problem

in the interpretation of their records.

Field samples of the tree composition were taken during the

summer of 1973, using the quarter method (Cottam and Curtis,

1956). From 150 to 200 points were sampled in the forested areas,

while 40 points were used in sampling an oak forest in an area

which was mapped as being an oak opening in ca. 1850. Sampling

stations were located on a map of the public properties of the

county, and only those stands which showed no signs of recent

disturbance were used. Only trees with a dbh of 4 inches or more

were recorded, although field notes of the character of the under¬

story and sapling frequencies were also kept.

1974] Barnes — History of Vegetation, Eau Claire County 361

VEGETATION TYPES

Mapping

When the surveyors established a section or quarter section

point, they measured the distance and compass bearing to the

nearest tree in each of 4 quadrants in the case of corner points

on the exterior township lines, or to the nearest tree on each side

of the line at all other points in the township. These data, along

with the common names of the trees and their diameters at breast

height (dbh) were then recorded in their field books. If there were

no trees nearby, they constructed a mound of earth and sod, and

so stated. Every major change in vegetation, trees which occurred

directly on the line, rivers, streams, ponds, lakes, caves, abrupt

topographical changes, windfalls, etc., were noted in their records.

Also, at the end of each section line they wrote a brief description

of the mile they had just traversed. From these data, a map of the

vegetation of the county, as it appeared in ca. 1850, was con¬

structed (Fig. 3).

Differentiation between prairies, oak openings, barrens and for¬

ests is made on the map. Wetlands are not extensive in the county,

occurring for the most part at the heads of smaller streams.

Prairies are defined as areas with less than one tree per acre. Thus,

all corners at which mounds had been constructed, or at which the

distance between trees was a minimum of 209 feet, were mapped

as prairies, providing this was consistent with section line descrip¬

tions. All areas in which the trees were less than 209 feet apart

but greater than 50 feet apart, or if the section line description

was appropriate, were mapped as oak openings if the trees were

predominantly oak, or barrens if the trees were predominantly

pine. When the trees were found to be less than 50 feet apart, the

area was mapped as forest.

Prairies

The area mapped as prairie represents the single most extensive

type of vegetation in the county at the time of settlement. Of the

2,053 points established in the county by the surveyors, 888 (44%)

were mapped as prairie. The greatest expanse of prairie occurred

in the central part of the county, south and west of the alluvial

terraces of the Eau Claire River. Here several whole townships

were nearly devoid of trees. The prairies, and the oak openings,

occur primarily south of the tension zone, on the western uplands.

Unfortunately, the surveyors rarely included any information

on the herbaceous species present in their field notes. Most section

line descriptions simply state “nothing but grass on it”, “rolling

prairie”, “level prairie”, or “first rate prairie”. A substantial

362 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

W,->V

K

I

fe

◄

a

ci

£

7i

O

Vff/’f.t

M\Y>

i> r:ur\

n

O

Indian trail • • • •

FIGURE 3. The vegetation of Eau Claire County in ca. 1850

1974] Barnes — History of Vegetation, Eau Claire County 363

number of section line descriptions indicated that much of the

area mapped as prairie had various amounts of oak scrubs present.

Included were descriptions such as “some oak brush”, “scattering

oak bushes”, “oak bushes”, and sometimes, “considerable oak

brush”, or “covered with oak brush”.

Some insight as to the species which occurred on these prairies

may be inferred from Buss’ study (1956) of plant succession on

the Meridean Prairie in adjacent southeastern Dunn County. He

found that the most abundant plants occurring on sites undisturbed

for at least 35 years were Andropogon gerardi Vitm., Tradescantia

ohiensis Raf., and Euphorbia corollata L. Species listed as being

common or numerous include Stipa spartea Trim, Phlox pilosa L.,

Artemisia ludoviciana Nutt., Solidago rigida L., Liatris scariosa

Willd. and L. pycnostachya Michx. Andropogon scoparius Michx.,

Sorghastrum nutans Nash., Petalostemum purpureum Rydb., and

P. candidum Michx. are species Buss listed as being scattered.

Of the trees which were recorded on the prairie, 41.1% were bur

oak, 36.8% were black oak, 9.7% were white oak, while 8.1% were

aspen.

The presence of large tracts of prairie in the Middle West has

been attributed to periodic burning (Gleason, 1913), and Curtis

(1959) discusses in some detail the role of fire in maintaining

the prairies of southern Wisconsin. Fire undoubtedly was respon¬

sible for the prairies in Eau Claire County. H. A. Towne, (Bar-

land, 1965) in discussing an elk hunt near Eau Claire in 1857,

provides the following description. “The low ground was covered

at the time by dense poplar thickets, through which the prairie

fire would run every few years killing them but leaving the dead

young trees still standing until a new fire should sweep the coun¬

try, and find the dry timber and consume them”. E. T. Sweet

(1875), a geologist, gives the following description of prairies

located northwest of Eau Claire near St. Croix, Wisconsin.

Analogous to the barrens are areas known as brush prairies and simply

prairie. In some of these prairies, young trees are springing up, and bid

fair, if undisturbed, to attain the usual size. These are appealed to as

examples of prairies returning to forest, since annual fires are no longer

permitted to ravage the region. So far as these areas are concerned, the

appeal seems to be well taken, save that we might, perhaps, justly dissent

from the use of the word prairies as applied to them; for there seems to

be no evidence that these ever were prairies in the sense of being com¬

pletely and compactly covered by prairie grass, to the exclusion of all

shrubs and stubs of arboreous plants, as in the case of true prairies. They

rather appear to have originally been open forest areas, which, on account

of the character of the soil, were especially subject to dryness, thus to

destructive action of annual fires; while moister adjoining areas escaped.

On the cessation of the destructive agent, they appear to be returning to

their normal condition.

364 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

The fact that much of the area, which was prairie at the time of

settlement, now bears tree growth supports the contention that fire

was responsible for the prairies of Eau Claire County.

No attempt was made in this study to take a contemporary

sample of the vegetation of the area mapped as prairie. However,

inspection of aerial photographs shows that most of the area is

presently under cultivation. Oak forests now occur throughout the

area, usually in the shapes of squares, rectangles, or other geo¬

metric forms, or are elongated and irregular and occur on the

steeper slopes or along streambeds.

Oak Openings

The oak openings occurred primarily in the southern and west¬

ern parts of the county in association with the prairies. Differenti¬

ation between prairie and oak opening was often difficult, and the

resulting boundaries are somewhat arbitrary. A total of 384 of

the corner points in the county (19%) are mapped as oak opening.

Section line descriptions of the oak openings again gave no

information concerning the herbaceous species, although it is pre¬

sumed they were not much different from those of the prairies.

“Some scattering oak trees”, “some oak timber”, “scattering oak”,

“timber scarce”, or, “oak barrens”, are typical section line

descriptions.

The average distance between trees was 90 feet, which is about

5 trees per acre. Of the 12 species of trees recorded by the sur¬

veyors, the most common species by far were black oak, with a

relative density of 38.5% ; bur oak, 35.0% ; and white oak, 16.4%.

The majority of these oaks had a dbh of from 8 to 12 inches.

Like the prairies, the oak openings were probably maintained

by fire. Muir (1913) described the oak openings in southern Wis¬

consin as open and sunny. He indicates that the oak forest devel¬

oped from the oak openings upon settlement by white man and

the subsequent cessation of fires. Cottam (1949) studied the

history of an oak forest in Dane County, Wisconsin, which was

once an oak opening. He came to the conclusion that it was fire,

rather than a change in climate, disease, or some other factor,

that was responsible for the oak openings at the time of settlement.

The major tracts of oak openings occurred along streams, espe¬

cially Lowe’s Creek and Otter Creek in the southwestern part of

the county, and Bridge Creek in the southeastern part of the

county. Similar patterns of oak opening distributions are apparent

in the studies of the vegetational histories of Beloit Township

(Rock County) by Ward (1956), of Dodge County by Neuen-

schwander (1957), and of Racine County by Goder (1956). The

patterns of distribution of the oak openings are probably related

1974] Barnes — History of Vegetation , Eau Claire County 365

to the reduced frequency and/or severity of fires in these moister

regions compared to the surrounding uplands, where almost no ma¬

ture trees were present.

A sample of 40 quarter points were taken in what is presently

an oak woods adjacent to Lowe's Creek in Section 4, T25N, R9W.

Although this sample is too small to place much statistical confi¬

dence, it is probably sufficient to provide some insight into the

major changes in the vegetation of this region, which have oc¬

curred since settlement.

The most obvious change in the oak openings is the change in

absolute density of the trees, from 5.4 trees per acre in ca. 1850

to 235 trees per acre in 1973. The major change in composition

is the increase in relative density of black oak, and the correspond¬

ing decrease in relative density of bur oak (Table 1). Curtis

(1959) states that bur oak is more resistant to fire than black oak,

which would seem to provide a reasonable explanation for the

opposite change in relative density of these two species, since the

time of settlement and the cessation of periodic fires. Other species

did not occur in sufficient numbers, either in the surveyor's records

or in the 1973 sample, to permit interpretation in any detail.

The oak woods present in Section 4 today is completely domi¬

nated by black oak, in terms of frequency of occurrence and rela¬

tive size, as well as relative density. Distribution of the size classes

of black oak is presented in Fig. 4. The rather large number of

trees in the lower size classes and the absence of many saplings of

any other species indicate that no major change in the composition

of this woods is likely to occur in the near future.

DIAMETER CLASSES (INCHES)

FIGURE 4. Size class distribution of black oak ( Quercus velutina Lam.)

366 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

TABLE 1. COMPARISON OF RELATIVE DENSITY AND DIAMETER

AT BREAST HEIGHT OF TREES IN THE OAK-OPENINGS IN

CA. 1850 AND 1973

The Oak-Pine Barrens

Most of the sandy stream benches along the western two-thirds

of the Eau Claire River were apparently occupied by a mixture

of oak and pine in ca. 1850. This is mapped as the oak-pine barrens.

The area includes 249 corner points, about 12% of the total for

the county. These barrens were very heterogeneous, with the oak

and pine generally forming a mosaic of separate stands of various

sizes. The mean distance between trees was calculated to be 86

feet, or 5.9 trees per acre, which is similar to the values obtained

for the oak openings. However, the variability of the distances in

the oak-pine barrens (Coefficient of Variation of 102%) is much

greater than that of the oak openings (c.v. of 67.5%). This indi¬

cates that the barrens were open areas that contained few trees

interspersed with rather dense stands.

E. T. Sweet (1875) wrote the following description of the bar¬

rens of northwestern Wisconsin.

The trees are either scrub-pine or black-jack oak, averaging in diameter

about three or four inches and in height not over fifteen feet. In some

places, as in the sand hills of the barrens, the trees are at considerable

distances from each other, and in other places the little scrub pines, not

over two inches in diameter, are so close together as to constitute a nearly

impenetrable thicket. On the sides of the barrens, and in low places, quite

large groves of norway pine are frequently found.

Moses Strong (1876), state geologist, described the barrens near

St. Croix, Wisconsin as follows:

The timber occupying these tracts is peculiar and does not justify the

application of the term barrens. Some portions are covered with scrub pine

1974] Barnes — History of Vegetation , Eau Claire County 367

to the exclusion of all else save underbrush. Most nearly similar are the

patches of norway pine. Other areas are covered with burr, black and

even white oak bushes, with occasional trees of these species. . . . There are

also areas where white pine occurs.

Section line descriptions of this area varied considerably and

included such entries as “scattering spruce pine”, “scattering oak”,

“scrub oak”, “nearly barren”, “timber small pine, quite thick”,

“some small pine on east one half, west one half oak barrens”,

and, “sparse pine timber, undergrowth of jack oak”.

Of the total of 157 jack pine trees recorded, 91, or 58%, were

in the 9 to 10 inch diameter class. Moreover, the diameters of the

trees at any given point rarely deviated by more than an inch

or two, although the diameters of this species ranged from 5 to

15 inches in these barrens. Red pine also apparently occurred in

even-age stands, while white pine was recorded primarily from

corner points near the Eau Claire River or in other low places.

These even-age stands of jack and red pine and the extensive areas

of oak shrubs attest to the fact that these barrens, like the prairies

and oak openings, were subject to periodic burning.

The 1973 composition of the oak-pine barrens was estimated

using 200 quarter points in 20 stands scattered throughout this

region. The results are presented in Table 1, along with compara¬

tive values obtained from the surveyors’ records. The major dif¬

ference in composition is the large decrease in red and white pine,

and the corresponding increase in the relative density of jack pine.

The decrease in red and white pine is due to logging in the latter

half of the 19th century. Jack pine and Hill’s oak saplings pres¬

ently occur in most of these stands, while white pine saplings are

infrequent, and saplings of red pine are rare. Another major

difference in composition between these two eras is the increase

in relative numbers of Hill’s oak and decrease in the numbers of

white oak. Apparently the openings created by logging, and the

fires that burned through the slashings, have favored the spread

of jack pine and Hill’s oak, at the expense of red and white pine

and white oak.

Some of this change in composition, especially with respect to

the oaks, may be due to another factor. Although much of this

area is presently county owned, some of it is private property

under cultivation. These private properties, which were not in¬

cluded in the sample, represent some of the better soils, and their

inclusion would in all probability, have yielded higher values for

white oak and lower values for Hill’s oak. Recalculation of the

composition of the oak-pine barrens in ca. 1850, using data only

from corner points in areas which are presently county owned,

368 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

did in fact give somewhat lower values for white oak and higher

values for Hill’s oak.

The average diameter of all trees in ca. 1850 was 11.1 inches,

while in 1973 it is 8.6 inches. However, since the surveyors had

a decided preference for large trees (Bourdo, 1956), the mean

diameter of the trees has probably not changed very much over

this time interval. Relative density of the species present today

and their diameter class values suggest little future change in the

composition of the vegetation of this area.

The Pine Barrens

The area mapped as pine barrens is continuous with the oak-

pine barrens and occurs along the Eau Claire River in the eastern

part of the county. The major difference between these two vege¬

tation types is the increase in relative proportions of pine over

oak. The three species of pine constituted 80.1% of all the trees

present, while the oaks accounted for only 10.6%. The variability

of the tree distances (c.v. of 101%), and section line descriptions

by the surveyors, indicate that, like the oak-pine barrens, this area

contained numerous, relatively dense stands of pine interspersed

over an otherwise rather open landscape. A total of 116 corner

points (5%) are mapped as pine barrens in the county.

Section line descriptions by the surveyors included such entries

as “scattering pine and pine barrens”, “timber black and yellow

pine”, “timber black, yellow and white pine”, and, “white and

yellow pine, first rate”. Other descriptive notations included “Enter

black pine barrens”, “Leave barrens, enter creek bottom”, “Leave

timber, enter barrens”, and, “Leave bottom of Eau Claire River

and enter black pine barrens”.

White and red pine were apparently far more abundant in the

pine barrens than in the oak-pine barrens. However, there seems

to have been some bias by the surveyors in choosing white and red

pine as witness trees. The mean distance of any given species from

the corner points should be the same as for any other species,

regardless of its density or frequency of occurrence. Yet, the mean

distance from corner points of 41 white pine trees was calculated

as 103.5 links (link = 0.67 feet) ; for 89 red pine trees it ,was 90.5

links, while for 62 jack pine trees it was found to be only 72.9 links.

The average distance for all trees was computed as 87.6 feet. The

probability of obtaining such deviations by chance alone is less

than 10%. Also, the surveyors recorded all large trees which oc¬

curred directly on the section lines. These represent a small, but

unbiased, sample of the relative density of each of the species. In

comparing these values to the corner point data, it was found that

1974] Barnes — History of Vegetation, Eau Claire County 369

the relative density of white pine on section lines was 28.6%, while

at the corner points it was 37.0%. The comparable values for red

pine were 42.9% and 49.3%, and for jack pine they were 7.1%

and 4.1%. Thus, this apparent bias by the surveyors results in

somewhat of an overestimate of white pine, and to a lesser degree,

of red pine, and thus underestimate of the relative density of jack

pine in the pine barrens.

Comparative values of the composition of the pine barrens are

given in Table 2. Like the oak-pine barrens, the major difference

in composition of the pine barrens from ca. 1850 to 1973 is the re¬

duction in numbers of white and red pine, and the corresponding

increase in the relative density of jack pine. There has been a

noticeable increase in oak, as the relative density of the species

listed as black oak by the surveyors has changed from only 3.0%

in ca. 1850 to 19.1% in 1973. The “black” oak present in this area

today is always Quercus borealis. When one compares tree diame¬

ters for these two time periods it is apparent that many large indi¬

viduals of white and red pine have been replaced by many smaller

individuals of jack pine and red oak.

Jack pine stands in this area are presently composed of indi¬

viduals of rather small size, and do not contain many saplings of

other species. The red oaks are present in all size classes and no

one or two species of saplings occur consistently in the understory.

TABLE 2. COMPARISON OF RELATIVE DENSITY AND DIAMETER

AT BREAST HEIGHT OF TREES IN THE OAK-PINE BARRENS IN

CA. 1850 AND 1973

370 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

Thus, no clear future successional trend is apparent in this vege-

tational type.

Forests

With the exception of the sandy alluvial terraces along the Eau

Claire River, the far eastern part of the county was occupied by

forests. It seems probable that the location at the eastern end of

the county, the heavier soils, and the more numerous wetlands in

this area greatly reduced the frequency of fire, thus permitting a

forest to develop. Because of differences in composition, this for¬

ested region is separated into a south forest, south of the Eau

Claire River, and a north forest.

South Forest

A total of 159 corner points (8%) were recorded in the south

forest. The mean distance between trees was found to be 30.9 feet,

which is only about 46 trees per acre. This low density value is

due, to a large degree, to the failure of the surveyors to include

many trees in the 4 to 8 inch diameter class as witness trees, as

it was difficult to inscribe the required corner identification infor¬

mation on them. Since these smaller trees usually represent a sub¬

stantial portion of the trees present in a forest, their inclusion

would have resulted in a smaller mean distance, and subsequently

a higher absolute density.

The character of the south forest was that of a closed forest,

with the exception of the rather numerous windfalls. Twenty-two

TABLE 3. COMPARISON OF RELATIVE DENSITY AND DIAMETER

AT BREAST HEIGHT OF TREES IN THE PINE BARRENS IN

CA. 1850 AND 1973

1974] Barnes — History of Vegetation , Eau Claire County 371

of the 159 corner points, 13.8%, were located in windfalls. A

characteristic descriptive entry in the field notes is:

16:00 Leave swamp and enter windfall, bears N and S

40:00 Set post for 14 sec. corner

W. Pine 16" S89E 592 links

No other tree near

55:00 Leave windfall, bears N and S

Some characteristic section line descriptions of these windfalls

included “Timber dead pine”, “Timber mostly dead and fallen”,

and, “Undergrowth in windfall brush and briars”. White pine was

the most frequently used witness tree in these windfalls, and the

soil types were mostly wet alluvial soils, Elm Lake loamy sand and

Adrian mucky peat, which are poorly drained soils. The signifi¬

cance of the role of windfalls in perpetuating the diversity of the

mixed conifer-hardwood forests of northeastern Wisconsin has

been discussed by Stearns (1949), and windfalls appear to have

been of equal importance in the forests of Eau Claire County. The

site of some of these windfalls was examined in the summer of

1973, and tip-up mounds, left by the uprooted trees, are still in

evidence in some places.

Table 4 indicates that major compositional changes have oc¬

curred in this vegetation type since settlement. The relative density

of white pine has dropped from 43.6% to 6.7%, and its average

diameter has changed from 18.7 inches to 9.3 inches. The relative

TABLE 4. COMPARISON OF RELATIVE DENSITY AND DIAMETER

AT BREAST HEIGHT OF TREES IN THE SOUTH FOREST IN

CA. 1850 AND 1973

372 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

density of red pine has decreased from 9.0% to 0.5% and, as in

the barrens, there has been a corresponding increase in the relative

number of jack pine. However, unlike the barrens, the increase

in relative density of jack pine has not been great, from 0.7 to

6.0%. The two species which exhibit the greatest increase in rela¬

tive density are red oak and aspen, with gains of 6.9 to 30.7 % and

3.8 to 22.7% respectively.

The average diameter of all trees recorded in ca. 1850 was 15.1

inches, compared to a mean of 7.9 inches of 600 trees measured in

1973. Part of this difference is undoubtedly due to the surveyors’

avoidance of very small trees. Nevertheless, the difference is great

enough to indicate a major change in the structure of this forest

between these two time periods.

Possibly indicative of a future successional trend are the many

small individuals of red maple and their numerous saplings which

are common in the south forest. However, many small individuals

of red and white oak are also present, thus no great change in the

composition of this forest seems likely in the near future.

North Forest

A total of 255 corner points (12%) were mapped in the north

forest. This community type is the only one in the county lying

north of the tension zone. It differs from the south forest primarily

in the presence of sugar maple and basswood in the canopy and a

higher frequency of witch-hazel ( Hamamelis virginiana L.) , horn¬

beam and blue beech in the understory. Also, the dominance by

white and red pine was not nearly as great here (combined rela¬

tive density of 28.6%) as in the south forest (52.6%).

The mean distance between trees in the north forest, 26.7 feet,

was somewhat less than that of the south forest. The mean diame¬

ter of the trees in the south forest was somewhat greater (15.1

inches) than that of the north forest (13.2 inches) due to the

greater number of white pine trees in the south. Some of these

white pine were 48 inches dbh at the time of the survey, and many

were 24 to 30 inches. Large fire scarred stumps of pine are still

in evidence in both of these forests today. Basal area was nearly

identical in the two forests, about 58 square feet per acre in the

south and 59 in the north.

Windfalls were also extensive in the north forest, occurring at

24 (9.4%) of the 255 corner points. Like the south forest, most

of these windfalls were apparently downed pine which had devel¬

oped on poorly drained soils.

The comparative composition of the north forest is given in

Table 5. Again, a drastic reduction in the relative number of pine

has occurred, with a corresponding increase in red oak trees. Other

1974] Barnes — History of Vegetation, Eau Claire County 373

TABLE 5. COMPARISON OF RELATIVE DENSITY AND DIAMETER

AT BREAST HEIGHT OF TREES IN THE NORTH FOREST IN

CA. 1850 AND 1973

species which have increased in number appreciably are aspen and

white birch. The relative density of sugar maple has decreased

from 14.0% in ca. 1850 to 1.1% in 1973. However, the reason for

this reduction is probably a parallel of the situation of white oak

in the oak-pine barrens. That is, sugar maple occurred primarily

on richer soils (especially Withee sandy loam and Loyal loam),

and these are the soils now under cultivation, i.e. no longer

forested.

Red maple is presently the most abundant small tree in the north

forest, with individuals occurring in 54% of the quarter points,

and saplings occurring in every stand sampled in 1973. This would

seem to indicate a successional trend toward red maple stands in

the future. However, the relative density of large individuals of

this species has not changed since ca. 1850 (12%), making such

a prediction somewhat tenuous.

SUMMARY

Records of the surveyors (ca. 1850) indicate that approximately

80% of Eau Claire County was prairie, oak opening and barrens,

while only about 20% was forest. The prairies and oak openings

occurred in the south and west, the oak-pine and pine barrens

occurred for the most part on the terraces of the Eau Claire River,

374 Wisconsin Academy of Sciences, Arts and Letters [Vol. 62

while the forests occurred in the extreme eastern part of the

county. This open nature of most of the vegetation was almost

surely due to fire.

The forests, although escaping destruction by fire, were sub¬

jected to another catastrophe, windstorms. About 11% of the

corner points in the forests were recorded as being in windfalls at

the time of settlement. These occurred primarily in the white pine

stands which had developed on poorly drained soils.

Some of the patterns of the vegetation as it existed in ca. 1850

are related to the tension zone. Most of the area south of the

tension zone was prairie or oak opening, while the area north of

the zone was forested. The area within the tension zone was very

heterogeneous and included all of the vegetation types discussed

in this study. However, the largest part of the tension zone occurs

on sandy outwash, which was dominated by the pines, especially

jack pine.

The relationship between the vegetation as it existed in ca. 1850

and contemporary land use patterns is quite apparent. Regions

previously prairie and oak openings are now predominantly farm¬

land. The oak-pine and pine barrens, as well as some of the forests,

are presently county-owned forest.

The major portion of the vegetation of Eau Claire County at

the time of settlement was open, or dominated by species of trees

of low shade tolerance. This indicates the importance of fire, and

to a lesser extent, windstorms, on the development of the vege¬

tation prior to settlement by white man. With settlement came

the cessation of prairie fires and the cutting of the pineries. This

caused forests to spring up on what was once prairie and oak

opening, and the forests to change in composition from those domi¬

nated by white and red pine to dominance by oak and aspen. The

vegetation of the barrens has shown the least amount of change

in composition, although the character of this vegetation type has

changed to a more closed form.

REFERENCES

BAILY, JUDGE WM. F. 1914. History of Eau Claire County, Wisconsin.

C. F. Cooper & Co., Chicago.

BARLAND, L. 1965. The River Flows On. A Record of Eau Claire County

from 1910-1960. Worzella Publ. Co. Stevens Point, Wisconsin.

BOURDO, E. A. JR. 1956. A review of the general land office survey and of

its use in quantitative studies of former forests. Ecology 37:754-768.

BUSS, I. 0. 1956. Plant succession on a sand plain, northwest Wisconsin.

Trans. Wis. Acad. Sci., Arts, Lett. 45:11-19.

COLLINS, C. W. 1968. An Atlas of Wisconsin. College Printing and Publ.,

Inc. Madison, Wisconsin.

COTTAM, G. 1949. The phytosociology of an oak woods in southwestern

Wisconsin. Ecology 30:271-287.

1974] Barnes — History of Vegetation, Eau Claire County 375

COTTAM, G. and J. T. CURTIS. 1956. The use of distance measures in

phytosociological sampling. Ecology 37 : 451-460.

CURTIS, J. T. 1959. The Vegetation of Wisconsin. The Univ. of Wise. Press.

Madison.

GLEASON, H. A. 1913. The relation of forest distribution and prairie fires

in the Middle West. Torreya 13:173-181.

GODER, H. A. 1956. Pre-settlement vegetation of Racine County. Wis. Acad.

Sci., Arts, Lett. 45:169-176.

Inventory of Forest Resources. Eau Claire County Forests. 1959. Wisconsin

Conservation Department. Madison, Wisconsin.

MARTIN, L. 1932. The Physical Geography of Wisconsin. The Univ. Wis.

Press, Madison.

MUIR, J. 1913. The Story of My Boyhood and Youth . Houghton Mifflin

Co., N.Y.

NEUENSCH WANDER, H. E. 1957. The vegetation of Dodge County, Wis¬

consin. 1833-1837. Wis. Acad. Sci., Arts, Lett. 46:233-253.

Soil Survey Report. Eau Claire County, Wisconsin. 1969. Soil Conservation

Service. Wis. Dept. Agr.

STEARNS, F. W. 1949. Ninety years change in a northern hardwood forest

in Wisconsin. Ecology 30:350-358

STROESSNER, W. J., and J. R. HABECK. 1966. The presettlement vegeta¬

tion of Iowa County, Wisconsin. Wis. Acad. Sci., Arts, Lett. 55:167-179.

STRONG, M. 1880. Geology of Wisconsin. Survey of 1873-1879, Volume III.

Edited by T. C. Chamberlin, p. 375.

SWEET, E. T. 1880. Geology of Wisconsin. Survey of 1873-1879, Volume III.

Edited by T. C. Chamberlin, p. 326.

WARD, R. T. 1956. Vegetational change in a southern Wisconsin township.

Iowa Academy of Science 63:321-326.

ADDRESSES OF THE AUTHORS

LOUIS W. BUSSE, School of Pharmacy, Univ. Wisconsin — Madison 53706

THOMAS FOSTER GRITTINGER, Univ. Wisconsin, Sheybogan Center, P.O.

Box 719, Sheboygan, Wis. 53081

WARREN R. WADE, Univ. Wisconsin — Stout, Menomonie, Wis. 54751

WILLIAM STEUBER, 705 Schiller Court, Madison, Wis. 53704

PAUL C. BAUMANN, JAMES F. KITCHELL, JOHN J. MAGNUSON and

TERRENCE B. KAYES, Dept. Zoology, Zoology Research Bldg. (P.C.B.),

and Limnology Lab., Univ. Wisconsin — Madison 53706

ROBERT T. BALMER, College Engineering and Applied Science, Univ.

Wisconsin — Milwaukee 53201

EDWARD E. POPP, 543 N. Harrison St., Port Washington, Wis. 53074

J. W. MERTINS and H. C. COPPEL, Dept. Entomology, Univ. Wisconsin — •

Madison 53706

WILLIAM C. SONZOGNI and G. FRED LEE, Laboratory Water Chemistry,

Univ. Wisconsin — Madison 53706 and Institute Environmental Sciences,

Univ. Texas — Dallas, P.O. Box 30365, Dallas, Tex. 75230

JON C. COOPER, G. FRED LEE and ANDREW P. DIZON, Limnology

Laboratory, Univ. Wisconsin — Madison 53706 and Univ. Texas — Dallas,

P.O. Box 30365, Dallas, Tex. 75230

G. FRED LEE and WILLIAM WILSON, Univ. Texas— Dallas, P.O. Box

30365, Dallas, Tex. 75230

G. E. HENDRICKSON and R. L. KNUTILLA, U.S. Geological Survey, Madi¬

son, Wis. and Okemos, Mich, offices respectively

THOMAS D. BROCK and JAMES HOFFMANN, Dept. Bacteriology, Univ.

Wisconsin — Madison 53706

WILLIAM DILWORTH, 23 Minnie Ave., Beloit, Wis. 53511

377

378 Wisconsin Academy of Sciences , Arts and Letters [Vol. 62

JAMES LA MALFA, Univ. Wisconsin Center, Marinette County, Marinette,

Wis. 54143

RICHARD A. KARSTEN, HAROLD McCONNELL and THOMAS D. PAT¬

TERSON, Dept. Community Affairs, Univ. Wisconsin Extension, 9722

W. Watertown Plank Rd., Milwaukee 53226 and Dept. Geography, Florida

State Univ., Tallahassee 32306 and Dept. Geography, Northern Illinois

Univ., DeKalb, Ill. 60115

ARTHUR MARSHALL, 125 East Gilman St., Madison, Wis. 53703

THEODORE S. COCHRANE and PETER J. SALAMUN, Dept. Botany-

Herbarium, Univ. Wisconsin — Madison 53706 and Dept. Botany-

Herbarium, Univ. Wisconsin — Milwaukee 53201

PETER J. SALAMUN and THEODORE S. COCHRANE, Ibid

BRIAN G. MARCKS, Dept. Environmental Quality and Conservation, Hor-

vard, Needles, Tammen and Bergendorff, Alexandria, Va.

MARY ELLEN ANTONIONI, 6318 Piping Rock Rd., Madison, Wis. 53711

OMAR M. AMIN and WAYNE H. THOMPSON, Univ. Wisconsin— Parkside,

Kenosha, Wis. 53140 and Univ. Wisconsin — Madison and State Labora¬

tory of Hygiene, Madison 53706

OMAR M. AMIN, Ibid, senior author above

LAVERNE C. STRICKER and ROBERT N. CHEETHAM, JR., USDA Soil

Conservation Service State Office, 4601 Hammersley Rd., Madison, Wis.

53711

RICHARD P. HOWMILLER, Dept. Biological Science, Univ. California —

Santa Barbara, Santa Barbara, Calif. 93106

WILLIAM J. BARNES, Univ. Wisconsin — Eau Claire, Eau Claire, Wis. 54701

WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS

OFFICERS

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54481

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5370 6

1973-74

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The ACADEMY COUNCIL includes the above officers, officials

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