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Cover Design by Jeffrey Homar
Title Page Design by Gail Mitchem
School of Fine Arts
University of Wisconsin-Milwaukee

rRjtNSilOTIONS OF THE
WISCONM ACADEMY
OF SCIENCES, ARTS
AND LETTERS

LV — 1966

Editor

WALTER F. PETERSON

EDITORIAL POLICY

The Transactions of the Wisconsin Academy of Sciences, Arts and Letters
is an annual publication devoted to the original, scholarly investigations of
Academy members. Sound manuscripts dealing with the state of Wisconsin
or its people are especially welcome, although papers by Academy members on
topics of general interest are occasionally published. Subject matter experts
will review each manuscript submitted.

Contributors are asked to forward two copies of their manuscript to the
Editor. The manuscript should be typed and double spaced on 8^/4 x 11" bond
paper. The title of the paper should be centered at the top of the first page
of the manuscript and should be typed in capital letters throughout. The
author^s name should appear in capital and lower case letters, and should
be underlined and centered directly below the title. A note identifying the
author by institution or background should be placed at the top of a fresh
page, immediately after the text of the article. Upper right hand page nota¬
tions from the second page on should read 2 — Brown, 3 — Brown, 4 — Brown, etc.

The cost of publishing the Transactions is great. Therefore, articles in excess
of twenty-five printed pages will not normally be accepted. In the rare instance
in which a longer paper is approved, the contributor may be asked to help
subsidize publication.

Documentary footnotes should appear at the end of the paper under the
heading “References Cited.'’ Supplementary or . explanatory notes of material
too specialized to appear in the text itself shoufd be typed on a separate sheet
entitled “Footnotes” and appended to the section entitled “References Cited.”
Contributors should avoid unnecessary documentation wherever possible. Other
matters of style should be in harmony with current practice in the subject
matter area.

Galley proofs and manuscript copy will be forwarded to the author for
proofreading prior to publication; both should be returned to the Editor within
two weeks. Papers received on or before July 15th will be considered for pub¬
lication in the current year. Papers received after that will be considered for
publication the following year.

Contributors will be given five offprints of their article free of charge.
Additional offprints in sets of 100, 200, etc. may be ordered at the time galleys
and copy are returned to the Editor. Price will vary according to quantity
desired and the length of the article.

Manuscripts should be sent to :

Professor Walter F. Peterson
Editor, Transactions of the Wisconsin Academy
Lawrence University
Appleton, Wisconsin 54911

TRiMCTMS OF THE
mSCOH ICitOEMY

Established 1870
Volume LV

INFLUENTIAL TEACHERS OF LITERATURE AT

THE UNIVERSITY OF WISCONSIN 1

Harry Hayden Clark

ANATOMY OF A DECIPHERMENT 11

Alan D. Corre

University of Wisconsin — Milwaukee

DEUSTER AS A DEMOCRATIC DISSENTER DURING THE CIVIL
WAR: A CASE STUDY OF A COPPERHEAD 21

Frank L. Klement

Marquette University, Milwaukee

GEORGE MADISON HINKLEY

SAWMILL ENGINEER FOR E. P. ALLIS 39

Walter F. Peterson
Lawrence University

WISCONSIN TERRITORIAL AND STATE CENSUSES 47

Walter H. Ebling

Department of Agricultural Economics
University of Wisconsin

DETOURING CALAMITY IN WATER RESOURCE DEVELOPMENT
A CASE IN POINT: SOUTHEASTERN WISCONSIN 59

Spenser W. Havlick

THE TRENTON METEORITES 77

W. F. Read and H. 0. Stockwell

FISHES OF SOUTHWESTERN WISCONSIN 87

George C. Becker

Department of Biology

Wisconsin State University, Stevens Point

THE SEASONAL DISTRIBUTION OF FISHES
IN VERMILION BAY, LOUISIANA

Carroll R. Norden

Department of Zoology

University of Wisconsin — Milwaukee

EVOLUTIONARY TRENDS OF THE MUCROSPIRIFER

LeRoy R. Peters

INTERACTION OF PHAGE T1 WITH STRAINS OF
ESCHERICHIA COLI

Marvin D. Whitehead and J. Roger Christensen

NOTES ON WISCONSIN PARASITIC FUNGI. XXXII

H. C. Greene
Department of Botany
University of Wisconsin, Madison

THE PRESETTLEMENT VEGETATION OF
IOWA COUNTY, WISCONSIN

Wayne J. Stroessner and James R. Habeck

REAPPRAISAL OF THE GROWTH POTENTIAL OF JACK PINE
AND RED PINES ON DIFFERENT SOILS OF WISCONSIN

S. A. Wilde, R. R. Maeglin, and Ch. Tanzer

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 55.
COMPOSITAE IV— COMPOSITE FAMILY IV (TRIBES
HELENIEAE AND ANTHEMIDEAE)

Carol J. Mickelson and Hugh H. litis
Herbarium

University of Wisconsin, Madison

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 56.
COMPOSITAE V— COMPOSITE FAMILY V TRIBE INULEAE
(ANTENNARIA, GNAPHALIUM, ANAPHALIS, AND INULA)

Edward W. Beals and Ralph F. Peters
Herbarium

University of Wisconsin

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 57.
FOLEMOMACEAE— PHLOX FAMILY 243

Dale M. Smith

Department of Biological Sciences
University of California, Santa Barbara

Donald A. Levin

Department of Botany

University of Illinois at Chicago Circle

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 58.
HYDROFHYLLACEAE— WATERLEAF FAMILY 255

Jack W. Shields
Dept, of Botany

University of Wisconsin — Milwaukee

i

i

I

HARRY HAYDEN CLARk|

t

iSth President of the |

WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS

INFLUENTIAL TEACHERS OF LITERATURE AT
THE UNIVERSITY OF WISCONSIN

Harry Hayden Clark

May I invite you at this evening’s banquet of the Academy to
share memories of some of the influential teachers of literature
of the generation before ours at the University of Wisconsin? Per¬
haps such memories will help to supplement what you have already
heard during the last two days about the great advantages our state
has in the way of environment, technology, recreational facilities,
technology, and other physical things. After all, an outsider in
moving to this state does not have to divorce himself or his chil¬
dren from opportunities to benefit from the non-physical cultural
resources of mankind in general, from a cultural heritage which
is both international and which stretches across the ages from
classical antiquity. Our forefathers were far-sighted and generous
in providing Wisconsin people with a state university which in its
literary departments was designed to act as a kind of “trans¬
former” to bring down to the level of individuals who live here the
great over-arching currents of whatever the ages have proved to
be conducive to ethical guidance, to human happiness, social justice,
and emotional enjoyment.

Since our Academy is distinctive in combining a concern with
the sciences and with the arts and literature (only two or three
other academies of the fifty states combine these), it may not be
amiss to mention a few of the common denominators of these
approaches. Both the sciences and the literary arts at their best
are fertilized by the imagination — the scientist such as Newton
usually proceeds by formulating an hypothesis, an imaginary sup¬
position to be tested inductively by the extent to which it explains
the action of particulars ; and the great writer such as Shakespeare
relates his particular characters to types arrived at by imagination.
Both the scientist and the literary man assume that the universe
is orderly, that given causes will produce predictable consequences.
Mixing certain chemical elements will insure producing repeatedly
certain compounds. Jefferson, the father of democracy, remarked
that one can learn more about the psychological ramifications of
filial ingratitude from King Lear than from mountains of abstract
sermons. And Macbeth clarifies the predictable outcome of murder¬
ous ambition. The psychiatrist and novelists such as Hawthorne,
the historian of the guilty conscience, centering on exploring the

1

2 Wisconsin Academy of Sciences ^ Arts and Letters [Vol. 55

individual’s inner life, have much in common. Both the scientist
and the literary man reason from particulars to universals, to
immutable scientific laws abstractly stated and to principles about
how man’s mind and heart react. It was no accident that most of
the founders of the Royal Society of Science in 1662 had been
trained in the literature of the ancient classics which taught one
to look for universals. Incidentally, Willard Gibbs first presented
his epoch-making “rule of phase” in the transactions of one of our
state Academies, and many theorists claim that investigations
hardly transcend pedestrianism or vocationalism unless they “fol¬
low through” and rise to the plane of universal laws true for all
times and places. In other words, both the scientist and the liter¬
ary man seek to penetrate beneath the surface appearances guided
by methods which try to be objective and austere. Individuals in
both disciplines may adopt sectarian religions, but in their distinc¬
tive professional concerns both tend to generalize not so much from
the supernatural or the miraculous as from verifiable human or
mundane experience. In short, the scientist and the literary man
in their higher reaches have much in common, and the founders
of our Wisconsin Academy were wise in uniting them in the same
organization, which has now endured for nearly a century.

I come now to our immediate topic, teachers of literature at the
University of Wisconsin from about 1915 to 1935. They may be
roughly divided into three categories according to what they chose
to emphasize. (In actual practice, in catering to classes of differ¬
ent ages, most teachers have used many approaches and methods;
but emphases may be illuminating in terms of individual tempera¬
ments and student interests.) First, reflecting the current vogue
of historiography and of concern with change sanctioned by the
vogue of evolutionism, teachers such as Karl Young, Alfred Holfeld,
and Arthur Beatty tended to emphasize a turn from Coleridgean
abstractions and judicial absolutes, and toward a concern in the
study of literature with historical continuity and sequence, with
historical influences, with explaining why a given book came to
take the shape it did as it grew out of specific backgrounds. Second,
continental theorists such as Brunetiere, having emphasized the
evolution of literary forms. University of Wisconsin scholars such
as Neil Dodge, Warner Taylor, J. F. A. “Sunny” Pyre and Ruth
Wallerstein tended to emphasize the place of literary techniques
and formal patterns in the resources of creative genius and in
enhancing the beauty of a given poem or novel. These people
stressed not so much background as foreground, a close and appre¬
ciative study of the actual text iso as to illuminate literary devices
which help to explain different nuances of aesthetic quality. Third,
continuing the classical concern with an appraisal of literary val-

1966]

Clark — Influential Teachers of Literature

o

o

ues, University of Wisconsin teacher-scholars such as Grant Sho;W-
erman, William Giese, Philo Buck, and F, W. Roe emphasized bring¬
ing a given drama or poem into relation with the norms provided
by the long tradition of values which had endured through the
centuries, trying to winnow and sift literary masterpieces which
in their wisdom and beauty might be of use to modern readers in
quest of self-realization and serenity, if not happiness. Such judi¬
cial critics thought of literature — often in Matthew Arnold’s sense
— as roughly allied with religion as a potential guide to fruitful liv¬
ing in their own day. They tended to try to avoid antiquarianism
and subjective appreciation, yet they approached literature in terms
of the authority of examples of past excellence and they exalted
beauty (as did Emerson) as the mark or by-product of virtuous
living.

Granting some overlapping of these broad classifications among
the teachers mentioned, I shall now proceed to more personalized
sketches. Karl Young, born in Iowa in 1879, had taken his B.A.
at the University of Michigan and his doctorate under G. L. Kit-
tredge at Harvard. His teaching apprenticeship at Annapolis before
coming to Wisconsin (where he taught for fifteen years, 1908-
1923) accentuated his tendency toward a military manner. (His
eminence is illustrated by the fact that Yale called him to head
its English Department, and he was elected national president of
the Modern Language Association.) Mr. Young was tall and slender
and handsome, with prematurely gray hair, the epitome of poise,
energy, refinement, and dynamic personal drive. He was the
scourge, especially during his chairmanship at the University of
Wisconsin, of anything fuzzy or nebulous or subjective or based
on guesswork. He began publishing in the Transactions of our own
Academy in 1910 on liturgical drama, an impressive early article
which was a link in a long chain of historical investigations which
led to his monumental two-volume book on The Drama of the
Mediaeval Church (Oxford, 1933). The current perverted slogan,
“Publish or Perish,” would have seemed to him a basic contradic¬
tion in terms. Himself a masterful teacher whose work benefited
from the student discussions he inspired, Mr. Young held that by
publishing one’s best insights he not only helped other overworked
teachers but kept his own teaching from perishing in the short
memory of one generation. He was a realist if not a pragmatist,
working inductively in rigorous relation to historical facts and
often newly unearthed texts which he used to explain why such
a master as Shakespeare should have been able to create hiis master¬
pieces.

Alfred Holfeld, also national president of the Modern Language
Association, led his well-organized German Department not only

4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

in teaching the language but in producing a host of historical mono¬
graphs illustrating in massive detail such matters as the actual
vogue of German literary ideas in our early American magazines
and the influence in America of writers such as Goethe. His oro¬
tund speeches in the Faculty of Letters and Science were master¬
pieces of persuasiveness and academic wisdom, and were very
influential.

Arthur Beatty, a native of Canada with a doctorate from Colum¬
bia, early illustrated the historical approach in studies (published
by our Academy) in the early drama in the roots that he found
in non-classical folk-festivals and the early mummers’ plays and
other works associated with the rebirth of the seasons. Later, Mr.
Beatty’s epoch-making book on Wordsworth (1922) stressed the
poet’s use of the Hartleyan psychology of associations to empha¬
size retrospect and the continuity of his own personal experience
as the subject-matter of The Prelude, a poem on the “growth of
a poet’s mind.” (This approach eventually merged into Words¬
worth’s later concern with the continuous tradition of his people
and their national religion in his later sonnet sequences and in his
anti-revolutionary admiration for Edmund Burke, spokesman for
the values of the continuity of tradition.) Mr. Beatty was a short,
stout, roly-poly kind of man, beloved by his students for his genial¬
ity. In later years he and his motherly wife enjoyed escorting
groups of American students through the places in England (such
as the Lake Country) which have historic associations with great
writers. He and his wife were active in various literary clubs and
he did much to interest students in literature through his gusto
and his ability to dramatize the man behind the book, to humanize
literature as growing out of actual three-dimensional individuals
who grew out of specific historical places and eras.

Since Merritt Hughes, a past president of this Academy is still
happily with is, he must be left to a future memorialist. Mention
will be made here of his long chairmanship of the English Depart¬
ment and of his editing of the works of Milton with voluminous
notes illustrating matchless erudition. His devotion culminated in
the publication of his many essays (Perspectives on Milton) by the
Yale University Press, with a eulogistic introduction by Harvard’s
Douglas Bush.

Turning now to the second of our categories of scholars, Neil
Dodge, born in the far west (Washington) in 1867, took his B.A.
and M.A. at Harvard, and in lieu of a doctorate studied intens-
sively for three years in the great libraries of Italy, France and
England. Beginning in 1898 he taught the rest of his long life at
the University of Wisconsin, serving as an austere chairman dur¬
ing his last years. His rearing in the family of an army colonel

1966]

Clark — Influential Teachers of Literature

5

was reflected in his ostensible sternness, which seldom hid his
kindly warmth of heart from those who knew him well His rich
knowledge of Italian writers such as Tasso and Ariosto led him to
serve as chairman of a committee on Comparative Literature which
eventually brought Philo Buck to the University of Wisconsin to
found a department in that field. Mr. Dodge’s concern was literary
in the strictest sense, emphasizing (especially in his teaching)
matters of versification and formal techniques. I recall his once
having his students, over a period of several weeks, demonstrate
that the first part of Milton’s '‘Lycidas” could be read in some
twenty ways in terms of prosodic emphasis involving matters such
as the substitution of an occasional anapest or dactyl for an iamb.
He was chosen as a major authority on Edmund Spenser to edit
his work for the renowned Cambridge Edition of the great poets.
Beyond mastering the texts themselves, Mr. Dodge was especially
concerned with getting his students to ferret out the distinctive
secrets of a poet’s verbal harmony and of how he used technical
devices to create beauty. Mr. Dodge was also famous for a course
in Advanced Creative Writing in which he coached mature students
in a highly personal way much as did Dean Briggs at Harvard;
he did much to get his students to avoid all affectations such as
what he called ''a highly rouged style” and any kind of extremism.
Literary art to Mr. Dodge was the organic by-product of the con¬
cept of the gentleman, a concept which he incarnated with impres¬
sive distinction and ethical elevation.

“Sunny” Pyre advanced the appreciation of literature for thou¬
sands of students in his large survey course. His concern for form
and technique are illustrated in his book on versification, The For¬
mation of Tennyson's Style, 1921, which Mr. Dodge helped to
inspire.

Warner Taylor took his B.A. and M.A. at Columbia University,
where he also taught for six years before beginning in 1911 his
36-year teaching career at the University of Wisconsin. Here, as
Director of Freshman English, Mr. Taylor had a tremendously
wide influence in molding the writing habits of tens of thousands
of students. His manner was quiet and gentle and persuasive. In
the early days his great sympathy and understanding and warm
hospitality helped to induct large numbers of young staff members
into the art of teaching. Mr. Taylor had a rare combination of
responsiveness to beauty along with an almost scientific interest
in trying to ferret out the mechanistic secrets of how and why a
given style produced its distinctive effect. (He was also a distin¬
guished ornithologist and won national prizes for his remarkably
beautiful photographs of birds.) Beside doing national surveys on
the teaching of Freshman composition, Mr. Taylor published highly

6 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

technical work such as his “Prose Style of Dr. Johnson’' (1918),
which illustrates his inductive concern with such matters as vari¬
ations of sentence length, the use of parallelisms and antithesis or
balance, imagery and metaphors, rhythm and cadence. Those of us
who were privileged to listen to Mr. Taylor’s annual talks to the
Department on the English Qualifying Exam for Freshmen, an
exam which included analyses of different kinds of style and liter¬
ary structure, will always remember his lucidity and orderly pro¬
cedure in elucidating the underlying design of the poems or pas¬
sages studied. Many middle-aged faculty members today who owe
part of their success to their power of expression have remarked
to me that they feel they got their start in a composition class
given by Warner Taylor, the man who stressed the importance of
how one says a thing.

Ruth Wallerstein came to the University of Wisconsin in 1920
after studying the ancient classics at Bryn Mawr and at the Uni¬
versity of Pennsylvania, where her doctoral dissertation was on one
of Shakespeare’s plays. She is aligned to those who emphasized
literary form by virtue of her posthumous Metrical Principles of
English Poetry (1961) and her Seventeenth Century Poetic (1950),
which had the rare distinction of winning a national prize from
Phi Beta Kappa. According to the Faculty Memorial of 1958, “She
knew how to communicate her enthusiasms to her students, and
she had a positive genius for discovering and awakening the gifted
student. . . . All who came into contact with Miss Wallerstein as
teacher and colleague and friend felt the charm and magnetism
of a very rare spirit.” (Incidentally, these faculty memorials of
individual teachers provide a most interesting supplement to the
history of the University.)

Since William Ellery Leonard, besides being a poet, taught lin¬
guistics rather than literature, his ,work is not quite relevant to
this talk on teaching literature. However, his foible of unceremoni¬
ously holding up younger colleagues to ask them precisely where
obscure passages appeared in the great writers, and his flamboyant
excoriation of anyone ignorant of such things, did much to get
young teachers to read with more precision. Trained by Kittredge,
in his younger days Mr. Leonard was a dynamic and inspiring
teacher and scholar.

Let us turn now to teachers who represent, broadly speaking,
judicial criticism roughly in the tradition of Matthew Arnold.
Grant Showerman and his Eternal Rome, along with his increas¬
ingly ethical familiar essays, paved the way for the blithe Ray
Agard, who in the Hellenistic spirit enjoyed illustrating in a
very broad and humane way the cross-fertilizations of ancient his¬
tory, literature, architecture, and jurisprudence as these drew upon

1966]

Clark — Influential Teachers of Literature

7

the vitality of the Great Tradition. Philo Buck, a native of New Jer¬
sey but reared in India by Missionary parents until he was sixteen,
came to the University of Wisconsin in 1925 to found the Depart¬
ment of Comparative Literature, having been educated at Ohio
Wesleyan and Harvard. With a famous course in the Great Books
and another entitled '‘History and Myth on the Stage,” Mr. Buck
reached thousands of Wisconsin listeners in his radio program. His
vitality, warmth, and genial individuality, along with his cultivated
manner of lecturing, somewhat in the British tradition, tended to
mellow the austerity which sometimes accompanies the judicial
approach. But the latter, involving classical standards, was readily
apparent in his critical guide entitled The Golden Thread, which
regarded as a norm what Plato (and Emerson) meant by the One,
the unity of mankind or the consensus gentium, which has had a
continuous endurance through the ages. Books, according to this
view, tend to live and to be applicable to the problems of each new
age in proportion as they focus particulars upon timeless universals
or the unity of human experience amid varying ephemeral distrac¬
tions. As Philo Buck’s memorial concludes, "He developed the new
department along his own specific lines as a medium for correlating
literature, art, philosophy and history and revealing man’s efforts
through the centuries to find meaning and value in life. If one might
find the 'golden thread’ of his thought, it was his optimistic faith in
man as evidenced by the best he has thought and felt.” Mr. Buck,
who made gracious living a fine art in itself, was able to lift the
spirit of his listeners to a glimpse of beauty and to awaken us to
a sense of kinship in mankind’s finest cultural heritage from all
the ages.

William Giese of the French Department, trained at Harvard
along with his friend Irving Babbitt, carried on something of the
latter’s tradition in his judicial criticism of such writers as Victor
Hugo and Sainte-Beuve, whose short-comings he did not minimize.
Contemplative and polished as a stylist, and not without asperity,
Mr, Giese has been likened to a light-house of good sense amidst
the more perilous shoals of nineteenth-century romanticism. Since
Helen White is happily still with us, and last year’s Academy’s
tribute (eloquently phrased by Mr. Welty) sums up her renowned
personality and achievements, I need not speak of her at length
here. Her main concern with devotional literature and the religious
writers of the seventeenth century tends to locate her among the
judicial ethical critics. Her unique sense of imaginative amplitude
has inspired a very large number of nationally known scholars such
as Mark Schorer. And her uncanny ability to appraise the poten¬
tialities of young candidates for University of Wisconsin positions

8 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

has done much to insure the recruitment of those who will carry
on her rich heritage.

Frederick W. Roe, who served as Junior Dean of Men, took his
doctorate at Columbia and came to the University of Wisconsin in
1905, serving as chairman of the English Department during his
later years. Of patrician tastes, living graciously in University
Heights near Dean Sellery and Mr. Buck, Mr. Roe taught and
published on the Romantic and Victorian writers as well as his
beloved Emerson. (In a nation-wide speech reminiscing about his
student days, the actor Frederick March once mentioned Mr. Roe’s
course in Emerson as one of his most memorable experiences.)
He had a special talent for getting students to visualize the sharp
individuality of authors such as Carlyle (to whose criticism he
devoted an early book), but he was mainly concerned with the social
criticism of authors such as Ruskin and Arnold. (Mr. Roe was
distinctive in his Ruskin-like love of painting, and like Emerson he
regarded beauty as a kind of symbol of the virtuous harmony and
paternalism of the good society.) He had a talent for friendship,
and did much in a personal way through correspondence to assure
his best students that someone at the University of Wisconsin was
interested in their continuing to do their best. Arnold’s sweetness
and light derived partly from the Hellenistic spirit represented his
norm as a judicial critic in quest of values which we might use
today to insure not only individual serenity but social justice.

^ As disciples of Arnold and Emerson, literary men such as Mr.
Roe approached our cultural heritage with certain assumptions.
Men do not live by bread alone, and all the material prosperity and
mechanical gadgets in the world will not alone insure the inward
emotional harmony essential to humane self-fulfillment. To this
end, one of our reliable aids is a cultural education which includes
a sense of the individual’s initiation into the great tradition of the
best that men have thought and that has already proved its power
to endure and to sustain successive generations. The great writers
such as Hawthorne have held that we need to balance the head
against the heart, rationalism against humane feeling and com¬
passion or the kind of empathy which underlies true brotherhood.
In an era celebrating the anti-hero, Mr. Roe liked to remind us
that Tennyson said that “we needs must love the highest when we
see it,” that Carlyle’s “Hero-Worship” and the desire to emulate
greatness and magnanimity are distinctively human traits still po¬
tentially among our resources. Amid all the bewildering changes
in modern life, there are still constants in human motivation and
reaction. As Joseph Campbell reminds us, the hero has a thousand
different faces, but most of the heroes in the great books which
have endured go upon somewhat the same journey through life.

1966]

Clark — Influential Teachers of Literature

9

which involves an alienation, initiation into life’s troubles, and
then our return as humbler but wiser men to the universal certi¬
tudes. One of the resources of literature as an ally of religion in
guiding us toward the good life or the good society is that it teaches
inductively and indirectly by example touched by beauty. As Emer¬
son remarks, ‘The beautiful is the highest, because it escapes the
dowdiness of the good [which sometimes appears merely negative]
and the heartlesisness of the true,” [when rationalism or science
are reduced to syllogisms or formulas]. He concludes, “Beauty is
the mark God sets upon virtue.” In other words, ethics as suggested
in the great books tends to benefit by the appeal of graciousness,
by making goodness seem desirable and attractive. As for the tri¬
partite concern of our Academy, it should be remembered that,
after Darwin especially, not to mention problems created by tech¬
nology, one of the great themes of literature has been ways and
means by which the individual can incorporate the findings of
science into hiis total thinking, religious and ethical, in such a way
as to maintain some degree of emotional balance and serenity. In
an era of fragmentation and specialization, as Emerson’s “Ameri¬
can Scholar” reminded us, literature’s primary concern is to help
the individual pick and choose so as to develop his personality
into an integrated and well-proportioned whole, rounding hiis com¬
plete life to the circle fair of orbed fulfillment.

Theodore Roosevelt once defined Americanism as practical ideal¬
ism. Our Academy can be justly proud of Wisconsin’s substantial
resources in land and factories and climate and city-planning and
the conservation of our physical resources. Let us not forget, how¬
ever, our non-physical resources in idealism as this has been made
available to our younger citizens by the “transformers” at our
state university who have enabled them to claim among their re¬
sources their birthright to a kinship in the great cultural traditions
which have sustained and guided other generations toward great¬
ness, toward self-fulfillment and emotional enrichment.

And literature, in books such as those by Dr. 0. W. Holmes and
Sinclair Lewis (Arrowsmith) , being concerned with suggesting
preferences, value-judgments, and envisaging goals for the good
life, has done much to motivate the better kind of scientists not
only in their quest of universal laws but in the practical relief of
suffering and in implementing humane compassion.

ANATOMY OF A DECIPHERMENT

Alan D, Corre
University of Wisconsin-Milwaukee

Outstanding research in the humanities too often goes unrecog¬
nized. For this reason in 1965 the Wisconsin Academy of Sciences,
Arts and Letters established a program of annual cash awards for
authors of meritorious papers in the humanities.

Since its founding in 1870, the Wisconsin Academy has sought
to encourage the diverse research interests of its members. Phil¬
ology, the broad field we now label language and literature, was
singled out from the start as worthy of support. Consequently the
Academy Committee for Recognition of Research in the Humanities
is pleased to make its first award in the field of linguistic
scholarship.

The First Place Academy Award in the Humanities goes to Dr.
Alan D. Corre for his fine essay ‘The Anatomy of a Decipherment.'’
Dr. Corre is Associate Professor and Chairman of the Department
of Hebrew Studies, University of Wisconsin-Milwaukee. As experts
and lay readers alike will discover, the following essay contains
much of interest and value.

Should readers of the Transactions wish to learn more about the
humanities prizes, they may write to the Chairman of the Academy
Awards Committee: Professor Goodwin F. Berquist, University
of Wisconsin-Milwaukee, Department of Speech, Milwaukee, Wis¬
consin 53201.

Introduction

‘‘Lecturer Learns Ugaritic’’

From our Correspondent
Johannesburg

Mrs. Leah Bronner, of Johannesburg, a lecturer in Hebrew at Wit-
watersrand University, has learned to speak Ugaritic, the language of
the Canaanites in 1400 B.C.

She learnt the language and Aramaic in order to write a thesis for a
doctorate. Mrs. Bronner, a mother of three children, will be the first
woman to receive a doctorate in Semitic literature at Pretoria University.

With all due respects to the distinguished newspaper ^ which
published this item, one might differ with it on two counts. First,
it is doubtful if anyone can learn to speak Ugaritic. The Ugari-

^ Jewish Chronicle (London), March 28, 1964.

11

12 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

tians, like certain other peoples in the Near East, unfortunately
did not indicate their vowels unequivocally, so that we cannot be
sure what the vowels were. Scholars reconstruct these vowels with
apparent certainty, but could we invent a time machine and chat
with the Ugaritians, we should doubtless be in for some shocks.^
Many factors of which we can have no knowledge may have been
in operation to make the vocalic structure of the language very
different from what we think it was. Second, it is rather surpris¬
ing that learning Ugaritic is any longer considered newsworthy.
Admittedly, Ugaritic shows no signs of becoming what political
jargon terms a “criticah’ language; yet Ugaritic is now well estab¬
lished as a member of the Semitic group of languages, having been
readmitted some 35 years ago when its sleep of 3,000 years was
first disturbed by a peasant on the Syrian coast right across from
the island of Cyprus, who found some small pieces of pottery at
Minet-el-Beida. The Archeological service in Beirut heard about it
and sent a man to investigate. He decided that the peasant had run
across a Mycenean tomb similar to ones found in Cyprus dating
from the thirteenth or twelfth pre-Christian centuries. Just half a
mile from this spot lies the mound of Ras Shamra, one of the many
heaps of earth in this part of the world that signal the existence of
a long dead city. Ras Shamra turned out to be the site of the an¬
cient city of Ugarit, already known from references in ancient
sources, whose location had previously been entirely lost. The de¬
cipherment of the tablets discovered there in a previously un¬
known cuneiform script presents a case history in decipherment of
lasting interest.

Who Deciphered Ugaritic?

It is generally agreed that the decipherment of Ugaritic was
“one of the shortest cases of decipherment on record.”^ The tablets
bearing the hitherto unknown cuneiform script were unearthed by
C. F-A. Schaeffer and G. Chenet about May 14, 1929. The first an¬
nouncement of their partial decipherment was published just a
year later, on June 4, 1930, by which time the tablets had been
exhibited locally, shipped to Paris, cleaned, transcribed and pub¬
lished. By 1931 the decipherment was virtually complete. This
stands in contrast to the decipherment of such languages as Egyp¬
tian and Akkadian which took long years of patient toil before they
yielded their secrets; but of course the difficulty of their scripts

2 One could not guess from written records that the vowels of merry/marry /'Mary
have fallen tog-ether in mid-western American English. We are’ in no better shape for
Ugaritic, which in general does not indicate vowels. While Ugaritic has archaic
features, it may Jiave been highly innovating in others. The difficulties of bringing
the distribution of the “three alefs’’ into any order is ah indication of how little we
really know about the vowel phonemes of this language.

3 I. J. Gelb. A Study of Writing (Chicago, 1963), p, 129,

1966] Corre — Anatomy of a Decipherment 13

was far greater than that of Ugaritic, with its small number of
signs. No bi“ or tri-lingual text was available for UgaritiCj unlike
Egyptian, which was deciphered only after the discovery of the
Rosetta stone with its parallel Greek and Egyptian inscriptions.
Akkadian too had to depend on multidingual texts for its decipher¬
ment (the Achaemenid inscriptions), although the other scripts
in the inscriptions were also previously undeciphered.

Some have reported that the decipherment of Ugaritic was
achieved independently and almost simultaneously by Hans Bauer,
E. Dhorme and Ch. Virolleaud.^ Others attribute priority to Bauer.
Thus the discoverer of the tablets writes:

It is to the credit of a German scholar, the late Professor Bauer of the
University of Halle, that he was the first to recognize that this language
was of Semitic origin, and that he tracked down certain words . . . work¬
ing on the same lines, two French scholars in their turn unravelled the
secret of the Has Shamra alphabet , .

More recently Johannes Friedrich has also given first place to
Bauer.® W. F, Albright, however, credits Bauer and Dhorme
jointly/ while A. M, Honeyman ascribes the decipherment to
H, Bauer, E. Dhorme “and other Semitists.''®

Who was really the first to decipher Ugaritic? As we shall see,
this question has no ready answer.

Preliminary Studies

On August 9, 1929, C. Schaeffer and G, Chenet brought before
the French Academic des Inscriptions et Belles Lettres, meeting
under President Rene Dussaud, the discoveries they had made
three months previously at Ras Shamra.® On September 20, 1929,
the French scholar Charles Virolleaud,^® to whom Schaeffer had
entrusted the tablets for study, presented to the Academy an as¬
sessment of the finds.

In his lecture he dealt briefly with the Akkadian tablets which
had been discovered, and went on to the tablets in the hitherto

^ Ibid. Cf. C. H. Gordon, Ugaritic Manual (Rome, 1955), p. 1. This seems to be
Gordon’s considered judgment. In his earlier Ugaritic Grammar (1940) he ascribes
priority to Bauer.

® C. F-A. Schaeffer. The Cuneiform Texts of Ras— Shamra Ugarit (London, 1939),
p. 37.

Johannes Friedrich, Extinct Danguages (New York, 1957), p. 83. (German edition:
Entzifferung Verschollener Schriften und Sprachen. Berlin, 1964, p. 70).

'^H. H. Rowley (ed.). The Old Testament and Modern Study (Oxford, 1951), p. 30:

. . its decipherment by Hans Bauer and Dhorme in 1931 (sic) . , . and its definitive
interpretation by Virolleaud. , .” Cf. also his statement in “The Old Testament and
Canaanite Language and Literature,” Catholic Biblical Quarterly VII (1945), pp. 9-10.

® Rowley, The Old Testament and Modern Study, p. 272; “Decipherment of this
system is due to the efforts of H. Bauer, E. Dhorme, and other Semitists.”

® C. F-A. Schaeffer “Les Pouilles de Minet el Beida”, Syria X (1929), pp. 285—303.
Charles Jean Gabriel Virolleaud was born July 2, 1879, at Barbezieux. He studied
at the Bcole des Langues Orientates in Paris.

14 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

unkno,wn script.^ Already he had taken some important steps
toward decipherment. He recognized only 26 or 27 signs/- which
meant without any possible doubt that the script was alphabetic.
He recognized too that the words were for the most part separated
by vertical word-dividers; that the vowels were not represented,
since the words were so short, rarely of more than five symbols;
that although some signs were formally identical with some Akka¬
dian signs, they would not have the same value, and in fact that the
Akkadian script would have no value for the decipherment; and
that there were different classes of texts. Virolleaud further ob¬
served that a number of adzes were inscribed with six signs, and
that these same six signs were preceded at the beginning of one
tablet by a seventh sign. He concluded that the six signs formed a
name of two parts (since they were elsewhere divided into two)
and that the seventh corresponded to the Akkadian ana denoting
possession, on the assumption that the tablet was addressed to the
owner of the adze. Another adze bore the same assumed name pre¬
ceded by four signs, two of them already occurring in the name.
He assumed that this was the word for adze. As it turned out,
Virolleaud was correct in all of these assumptions, with only
minor correction. However, his guess that the language of the tab¬
lets might be identical with the autochthonous language of Cyprus
written in the Cypriot syllabary was incorrect.

The news of the Ras Shamra excavations reached a far wider
public on October 12, 1929, when the French magazine L’ Illustra¬
tion published an article by Schaeffer and Chenet entitled “Des
Tombeaux Royaux et un Palais du 2® Millenaire avant J-C.”^^ The
article refers to the finding of tablets written in alphabetic cune¬
iform, but as yet undecipherable.

In the Revue Biblique for January 1930 Edouard Dhorme^® drew
attention in a brief note to the '‘sensational discoveries’" in Syria,
and looked forward to the publication of the texts. Publication
came in April 1930 as a supplement to Virolleaud’s address to the

C, Virolleaud. “Les Inscriptions Cuneiformes de Ras Shamra” in Syria X (1929),
pp. 304-340.

Thirty are recognized currently. Actually there are more, hut the additional signs
are variants of other signs, made by the addition of an extra wedge. Cf. C. H. Gordon,
Uyaritic Manual (Rome, 1955), pp. 11-12.

^ If one regards the West Semitic scripts as syllabaries (i.e. a consonant plus any
vowel), one might rephrase this in the sense that the script belonged to the West
Semitic rather than to the East Semitic syllabaries, despite appearances. Cf. Gelb,
op. cit., chapter 4.

“ Pp. 401 ff. Another popular article by Schaeffer, with excellent photographs,
appeared in the National Geographic Magazine, October 1930 (vol. 58), pp. 476-516,
under the title : “A New Alphabet of the Ancients is Unearthed.” Here Schaeffer
again refers to the ‘‘undeciphered” script, although by this time the script had in
fact already been deciphered.

Edouard Paul Dhorme was born at Armentieres on January 15, 1881. He studied
at the Ecole Biblique in Jerusalem and the Sorbonne.

1966]

Corre — Anatomy of a Decipherment

15

Academy on September 20, 1929, published in Syria.^^ The texts
were now available to the scholarly world in a clear and careful
transcription.

Bauer’s Decipherment

On April 22, 1930, Virolleaud’s transcription reached Hans
Bauer^^ in Halle. Bauer immediately began decipherment and com¬
pleted his tentative list five days later.^® The next day he communi¬
cated with Rene Dussaud of the French Academy, who passed on
the word to the Academy on May 23 and published an announce¬
ment in Syria, according to which Bauer had identified some 20
letters. In the meantime (on May 15) Bauer had notified the Ber¬
lin newspaper Die Vossische Zeitung of his discovery, and the
news was published in the supplement (das Unterhaltungshlatt)
to the issue of June 4, 1930. Here Bauer claims to have identified 20
characters with certainty and four tentatively (27 Buchstahen,
ivovon 20 sicker, U 'yait W ahrscheinlichkeit bestimmt sind). He re¬
fers specifically only to t and 2 He also claims to have read several
words, among them those for god, three, priests, and ax (which he
renders garzen)?^ Thereby he demonstrated the Semitic nature of
the language and refuted Virolleaud’s Cypriot hypothesis.

Further details were given in Forschungen und Fortschritte for
August 20, 1930. He had used as his starting point the assumption
that the language was Semitic, then the fact that west Semitic
has a limited number of consonants which are used as prefixes and
suffixes. He recognized that Virolleaud had already given the clue
to the prefix I denoting possesision. Bauer then sought common
words such as mlk (“king”)^^ and FI (“Baal”). He also interpreted
a number of phrases, and promised to publish shortly a full scale
work on the texts, which appeared in due course under the title
Die Entzifferung der Keilinschrifttafeln von Ras Schamra (Halle,
1930), incorporating his erroneous interpretation of half a dozen
characters. He even became the first in three millennia to write
something in the Ugaritic script, concluding this book with an

16 See note 11.

17 Hans Bauer was born at Grassmansdorf, Bavaria, January 15, 1878. He died
at Halle in 1937, where he had been professor ordinarius,

1® Cf. Hans Bauer. Entzifferung der Keilinschrifttafeln von Ras Schamra (Halle/
Saale 1930), p. 3.

Syria XI (1930), p. 200.

-6 In point of fact, Bauer had not read the word for priests. The word at issue is in
2.10 (Gordon, Ugaritic Manual, p. 129) and is to be transliterated mhnL By pure co¬
incidence Bauer’s errors made this read khnm (cf. Die Entzifferung, p. 13). The
rendering- garzen was also incorrect, the true form being- closer to the Akkadian cog¬
nate. It is doubtless a loanword, and it is entirely possible that the Hebrew qrdni,
which also means a tool, is a doublet of this word which came via a different route
and acquired a different meaning.

“1 Bauer erroneously read slm (tablet 1, line 8) as mlk and slmm (tablet 3, line 52)
as mlkk, thereby introducing a confusion from which he never recovered until he was
helped by Dhorme.

16 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

attempt to write in Ugaritic — or, more accurately, in Hebrew with
Ugaritic characters — ‘^Blessed art Thou, 0 Lord our God. Amen
and Amen.” It would probably be true to say that a Ugaritian scribe
would have had more difficulty in understanding what Bauer wrote
than vice versa.

Let us now examine Bauer's decipherment. In the Vossische
Zeitung he claimed to have interpreted 20 signs with certainty
and four tentatively. Two months later in Forschungen und Fort-
schritte, he published the values of eighteen signs, although he
again affirmed that 20 could be read with certainty. Of these, ten
have withstood the test of time fully (b, d, h, h, w, 1, n, r, t).
In the two alefs (now transcribed a and i) Baiier did not reach
the whole truth, although he came very close. Hence Friedrich's
statement^^ that Bauer had interpreted 17 characters correctly by
April 1930 needs this much emendation. Six signs were incorrectly
interpreted — g (which should be h) , another w (for k) , k (for m) ,

z (for s), m (for s) and s (for y. In the Entziffering he adds
two further correct interpretations — g, which he writes g because
he already had another incorrect g and y — and five further incor¬
rect interpretations — g (for q), q (for p), h (for u), p (for s)
and p (for s). In view of the fact that Bauer, between publication
of Forschungen und Fortschritte and the Entzifferung, had added
five new incorrect interpretations, compared with two new correct
ones, one may be permitted to wonder whether he would ever have
achieved a full decipherment, i.e, one permitting the reading of
connected texts, without the help he was to receive from Dhorme,
because several of his errors were in letters of high frequency,
and he had transcribed the entire corpus of texts then available
without sensing the basic errors in his proposed decipherment.
However, this help from Dhorme was forthcoming even before
the Entzifferung left the press.

Dhorme's Decipherment

Dhorme began his research about the same time as Bauer. When
Bauer's article appeared in the Vossische Zeitung, he had already
independently identified El, but had confused n and t, an error
which Bauer's article corrected for him. Since, however, Bauer
only hinted at his full decipherment, Dhorme continued his re¬
searches, fortunately, since he was not so advanced as Bauer by
June, and might possibly have accepted Bauer's erroneous decipher-

22 Not surprisingly, since the whole truth still eludes us. Cf. J. Reif, “The Loss of
consonantal aleph in Ugaritic,’’ Journal of Semitic Studies 4.1 (January 1959), pp.
16—20. There is no doubt that this problem must be solved by observing the actual
distribution of these alefs in Ugaritic, and not trying to fit them in to preconceived
notions as to the nature of proto-Semitic.

23 Johannes Friedrich. Extinct Languages (New York, 1957), p. 84.

1966]

Corre- — Anatomy of a Decipherment

17

ment if it had been published at that time.-^ As it turned out, he
achieved a much better result than Bauer. In the Revue Bihlique
for October 1930 he published an article in which he deciphered
correctly 18 characters (b, d, h, w, h [which he transcribes h],

t,2'^ y, k, 1, m, n, s, p, q, r, s, t) and six incorrectly (s [for the
correct u], ‘ [for g], g [for h], z [for s], is [for d] and s [for t]).
Additionally, like Bauer, he had come close to the truth in two
of the alefs. This decipherment enabled Dhorme to read the in¬
scription on the adzes as meaning ‘The chief of the priests’’ (which
Virolleaud had suggested must be a name) and much more besides.

This study was completed on August 15, 1930. A month later,
Dhorme, alerted by Rene Dussaud, read Bauer’s article in
Forschungen und Fortschritte, which he found to his surprise dif¬
fered basically from his decipherment. He thereupon added a post¬
script to his article in which he commented that “it will be inter¬
esting to see which of us is right,” and sent the proofs to Bauer.
At this time Bauer had just completed his Entzifferung ; Dhorme’s
communication obliged him to add a Wichtiger Nachtrag in which
he accepted Dhorme’s interpretation as more fruitful in explaining
enigmatic passages. On October 3 he wrote to Dhorme accepting
his findings, and on October 5 he communicated to Dhorme a re¬
vised decipherment representing the combined efforts of both
scholars which was published by Dhorme the next year.-® This
list was quite good enough for all practical purposes. Of the 27
letters they recognized at the time, 23 were correct — b, g, d, h, w,
h, h, t, y, k, 1, m, n, s, s, (which they wrote S2), p, s, q, r,
s, t. The interpretation of the two alefs remained the same; the
third alef was transcribed h, and d was transcribed s. Thus, about
a year after the publication of the texts, a decipherment was avail¬
able which was substantially correct.

ViROLLEAUD’S DECIPHERMENT

Virolleaud had also been working on the decipherment. About
the same time as Bauer was publishing his results in Forschungen
und Fortschritte, Virolleaud had received a new set of tablets
found by Schaeffer in 1930. These took about a month to clean, and

Cf. E. Dhorme. “Un nouvel alphabet semitique.” Revue Bihlique XXXIX (October
1930), p. 573; and “Le dechiffrement des tablettes de Ras Shainra,” Journal of the
Palestine Oriental Society, 1931, reprinted in E, Dhorme. Recueil Edouard Dhorme
(Paris, 1951), pp. 531-536.

^ There is a nondescript sign for this in his list, which is probably a transcriber’s
error, since Bauer had not succeeded in identifying- this sign, but it appears later in
the joint Bauer— Dhorme list.

E. Dhorme. “Premiere traduction des textes pheniciens de Ras Shamra,’’ Revue
Bihlique XL (January 1931), p. 33.

27 In Dhorme’s transcription the sign for z is omitted, doubtless an error.

18 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

a few days later Virolleaud had confirmed his previous supposi¬
tions with regard to the decipherment, which seemingly he was un¬
willing to publish until confirmation was forthcoming. On October
3, 1930 (the very day on which Bauer wrote to Dhorme accepting
his corrections), Virolleaud’s communication was read to the
French Academy, and three weeks later he himself presented his
results.^^ Like the others, Virolleaud used the 1 as his point of
departure. He then searched for the frequent words containing
the 1, mlk, b'L A set of signs in which the first and last were
identical and the middle was 1 furnished a word cognate with

Hebrew sis (three). These findings were confirmed by a text
containing several of the numerals.^® Virolleaud also recognized
that Ugaritic possessed three signs for the alef (only two had been
recognized previously) and that one of them contained the vowel a.
Virolleaud does not point out specifically the value of some com¬
mon signs (such as d, h, y [which he transliterates i], n, r) al¬
though he certainly had them, because he correctly translates words
containing them.’"^^ In addition to these five he had 17 other signs
correctly deciphered (a, i [written as e, which this sign may often
represent], b, g, z, h, h, t, k, 1, m, s [which he transliterates s],

p, s, q, t). His incorrect decipherments are u (which he trans¬
literates e), z (which he transliterates f), g (of which he is un¬
certain, but suggests may be another h) , s (which he transliterates
s), and t (which he transliterates s) . For some reason he fails to
mention w altogether, although he probably knew its value. Virol-
leaud’s treatment of the subject indicates that his purpose was first
to get to the meaning of the texts and not secure a decipherment
only.

This presentation was treated by the French newispapers^^ as the
first decipherment of this “mysterious alphabet’', whose enthusias¬
tic reports received a tart rebuttal from Dhorme. “Our readers will
know,” he declared, “what reliance can be placed on these state¬
ments.”^-

^ Cf. C. Virolleaud. “Le Dechiffrement des Tablettes Alphabetiques de Ras-Shamra,”
Syria XII (1931), pp. 15-23.

This text (which was not available to Bauer and Dhorme) was published in
Syria XV (1934), p. 249.

3* Virolleaud admits (La Legende Phenicienne de Dane!, Paris, 1936, p. 71) that
he obtained the value of the d from Bauer, presumably from the article in Forschungen
und Fortschritte.

31 For example, Le Figaro for October 25, 1930, reported (p. 3):

M. Virolleaud est parvenu a dechiffrer, par une methode qu’il a expose, a cette
Compagnie, les tablettes cuneiformes trouvees par MM. Schaeffer et Chenet, a
Ras-Shami-a . . . Da decouverte de M. Ch. Virolleaud . . . ne souffre d’ailleurs
. . . aucune incertitude, et le dechiffrement admirable fait par M. Virolleaud
en est deflnitif.

“Nos lecteurs savent a quoi s’en tenir sur la portee de ces affirmations.’’
E. Dhorme, “Premiere traduction des textes pheniciens de Ras Shamra,” Revue
Biblique XD (January 1931), p. 33.

1966]

Corre — Anatomy of a Decipherment

19

Nor was this remark by Dhorme an end to the dispute. In 1936
Virolleaud indicated that the information which was read to the
French Academy on October 3, 1930, had also been communicated
to Bauer, who used it to correct his work, and it was later published
under Dhorme's name, as we have seen,^^ This produced an indig¬
nant rebuttal from Bauer, who called Virolleaud’s assertion “eine
glatte Unwahrheit.’' He indicated that Virolleaud had indeed writ¬
ten to him, but did not communicate any usable information. What¬
ever may in fact have passed between the two men, it iseems fairly
clear that Bauer and Dhorme’s combined efforts were sufficient to
produce the ‘‘alphabet of 5 October'' without Virolleaud's help.

Conclusions

What then was the contribution of each of these scholars to the
decipherment? Unquestionably Bauer was the first to publish, on
June 4, 1930, the correct decipherment of some signs. Although no
cuneiform signs appear in the article in the Vossische Zeitung,
Bauer's comment

so bedeutet z.B, der einfache liegende Keil, der im Babylonischen asch
zu lesen ist, in unserer Schrift t . . .

leaves no doubt that he had deciphered the t. Bauer's incredibly
rapid progress in the decipherment calls forth admiration, and one
cannot doubt the brilliance of his initial efforts. But this admira¬
tion must be tempered by the fact that his later work was unsound,
and one cannot avoid the impression that his urge to publish in
haste entirely set aside the need to sift his findings.

Dhorme was the first to publish (in the Revue Biblique for Octo¬
ber 1930) an alphabet sufficiently accurate to permit the reading
of texts. Thus Dhorme was able to read the inscription of the adze,
which according to the decipherment of Bauer's third publication
on the subject (die Entzifferung) was to be read rb whnk — which is
meaningless. However, Dhorme had received isome early help from
Bauer, as we have seen.

It would seem therefore that Bauer and Dhorme should share the
honors, as Albright suggests. What of Virolleaud? It is entirely pos¬
sible that Virolleaud had achieved a partial, or perhaps almost com¬
plete, decipherment before the others ever started.^^ Virolleaud's ex¬
position of October 24, 1930, shows such detailed understanding of

^'3 C. Virolleaud. La Legends Phenicienne de Danel (Paris, 1936), p. 71. . . en

meme temps que j’informais I’Academie des inscriptions, j’avais cru bon de communi-
quer a M. Bauer les resultats complets de mes recherches personelles. M. Bauer a
immediatement adopte ces resultats . .

H. Bauer. “Zur Entzifferung- der Keilschrift von Ras Schamra,” Orientalistische
Literatur^seitung, XL (1937), col, 81-83.

®^As of February 14, Virolleaud was still orienting- his research to Cyprus, since he
communicated thus to the Societe Asiatique. Cf. H. Bauer, Das Alphabet von Ras
Schamra (Halle, 1934), p. 41.

20 Wisconsin Academy of Sciences^ Arts and Letters [Vol. 55

the contents of the tablets that it is clear that the decipherment was
far behind him. Particularly his discovery of the statement ‘‘He
pleads the case of the widow, he judges the suit of the orphan’' was
a strong, almost prophetic, hint of the importance that Ugaritic
was to assume in the elucidation of the Hebrew Bible, and points
to his grasp of the texts. He himself testifies that he was just about
ready to publish his decipherment when Bauer communicated his
finding to Dussaud. Should we therefore perhaps grant priority
in the discovery to Virolleaud in spite of Bauer’s publications?

The answer may safely be left to the historian of the decipher¬
ment of Akkadian, Robert W. Rogers, who, in granting “an unas¬
sailable superiority in translating” to Sir Henry Rawlinson over
Edward Hincks, remarks that in Hincks’ notes he shows great
skill as a translator, but for some reason he did not publish, Rogers
goes on :

The judgment must remain as it is, for the historian of the science
must base his decision on the published work of the pioneers and not
upon that which they left hidden in their notes,^

Similarly, Bauer, joined perhaps by Dhorme, must remain the first
decipherer, whatever may have lain on Virolleaud’s desk the day
before the issue of Syria reached Halle.

But one must admire Virolleaud’s part in this whole story, for
he displayed a remarkable scholarly altruism. He could easily have
delayed publication of the tablets until he was sure of a decipher¬
ment, or despaired of achieving one. As it was, he published them
immediately, knowing full well that another might thereby carry
off the prize of elucidating them first, as in fact happened. It
was no doubt the fact that Bauer had rushed into print — “un peu
prematurement, peut-etre” to quote Virolleaud — while Virolleaud
was still working on a really sound decipherment that brought
about the note to which Bauer objected so violently.

But if Bauer was the first decipherer, and Dhorme the first accu¬
rate decipherer, Virolleaud, by virtue of his great contributions
then and later, is the true father of Ugaritic studies. As Rogers
says: “To each man his own gifts and his own reward.”

.Robert W. Rogers. History of Babylonia and Assyria (New York, 1915), I, pp.
239-240.

DEUSTER AS A DEMOCRATIC DISSENTER DURING THE CIVIL
WAR: A CASE STUDY OF A COPPERHEAD*

Frank L. Klement
Marquette University, Milwaukee

Wisconsin made a notable contribution to the winning of the
Civil War, No Northern State, in proportion to population, had a
better record in furnishing soldiers. No state's soldiery received
more acclaim for courage and heroism. Three Wisconsin regiments
helped to give the famous Iron Brigade its enviable reputation, and
General William T. Sherman one time said that he always consid¬
ered a Wisconsin regiment “equal to an ordinary brigade,” Wis¬
consin's five Civil War governors, all Republicans, gave full support
to the war effort. Such newspapers as the Milwaukee Sentinel, the
Racine Advocate, and the (Madison) Wisconsin State Journal con¬
sistently endorsed all the war measures of the Lincoln administra¬
tion. Most of the state's citizens were patriots. Yet, on the other
hand, Wisconsin also furnished some well-known critics of Abra¬
ham Lincoln and Civil War policy. Marcus Mills (“Brick”)
Pomeroy of the La Crosse Democrat brazenly opposed most meas¬
ures of the Lincoln administration and had the gall to label Lincoln
“a flat-boat tyrant”-— even hoping for the president's assassination,
Edward G. Ryan, destined to become Wisconsin's most famous
jurist, wrote a scholarly critique of Civil War policy and instructed
the Democracy to oppose the changes which the conflict was impos¬
ing upon the country. Moses M. Strong of Mineral Point frequently
spoke against some of President Lincoln's war measures and
worked hard to put a Democratic president in the White House in
1864. George H. Paul of the Milwaukee News, Flavius J. Mills of
the Sheboygan Journal, and Stephen D. (“Pump”) Carpenter of
the (Madison) Wisconsin Patriot wrote editorials critical of
Lincoln and the war. No Wisconsin Democrat, however, developed
as solid an anti-war bloc as Peter V. Deuster, the prominent politi-

* Several years ago the author did a very brief and semi-popular article entitled
“Peter V, Deuster, the See-Bote, and the Civil War” for the Historical Messenger
(of the Milwaukee County Historical Society), XVI (December, 1960), pp. 2-6. That
cursory study has served as a springboard for this article.

21

22 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

cian who used the (Milwaukee) See-Bote as an outlet for his anti-
Lincoln and anti-war views.^

By 1860 Peter Victor Deuster served as chief spokesman for
the thousands of German Catholics who lived in southeastern Wis¬
consin. His rise to leadership was no accident — rather it was the
result of his ability, experience, and audacity. He had much in
common with the thousands of German Catholics who found their
way to Wisconsin in the 1840’s and 1850’s. He was born near Aix-
la-Chapelle (Aachen) in Westphalia on February 13, 1831. As a
lad of sixteen he accompanied his parents from the Rhineland to
Milwaukee, just as Wisconsin Territory was preparing for state¬
hood. After spending a year with his parents on a farm near Mil¬
waukee, young Deuster went to work for Moritz Schoeffler, pub¬
lisher of a German-language newspaper named the Wiskonsin-
Banner. Four years later he undertook the publication of his own
newspaper, a German-language family weekly named The Haus-
freund. Six months later he sold that paper and became business
manager of the (Milwaukee) See-Bote, a newspaper with great
influence among German Catholics of the area. In 1854, tired of
administrative chores, he moved to Port Washington to edit the
Zeitung, the fourth German-language newspaper with which he
was associated. Deuster became a community leader in Port Wash¬
ington, serving as notary public, clerk of the circuit court, and
postmaster as well as editor of the Democratic-oriented Zeitung.
In 1856 he had a chance to return to Milwaukee to team up with
August Greulich in publishing the See-Bote, and that paper flour¬
ished. Less than four years later, in January of 1860, the twenty-
eight year old immigrant American became sole proprietor of the

1 Wisconsin’s contribution to the war effort is reviewed and summarized in two dif¬
ferent papei’backs, Frank L. Klement, Wisconsin and the Civil War (Madison, 1963),
and Robert Wells, Wisconsin in the Civil War (Milwaukee, 1963). The role of the
Iron Brigade is superbly related in Alan T. Nolan, The Iron Brigade (New York,
1961) and the role of the governors is judiciously portrayed in Robert H. Jacobi,
“Wisconsin Civil War Governors’’ (M.S. thesis, typewritten, University of Wisconsin,
1948). The general subject of Democratic opposition to the Civil War is treated in
Frank L. Klement, Lincoln’s Critics in Wisconsin (Bulletin No. 14, Lincoln Fellow¬
ship of Wisconsin, Madison, 1956) and in Frank L. Klement, “Copperheads and Cop-
perheadism in Wisconsin ; Democratic Opposition to the Lincoln Administration,’’
Wisconsin Magazine of Histoo'y, XLII (Spring, 1959), pp. 182-188. The author has
discussed “Brick” Pomeroy as an anti-war man in three different articles — see Frank
L. Klement, “ ‘Brick’ Pomeroy, Copperhead and Curmudgeon,” Wisconsin Magazine
of History, XXXV (Winter, 1951), pp. 106-113; “A Small-Town Editor Criticizes Lin¬
coln: A Study in Editorial Abuse,” Lincoln Herald, LV (Summer, 1952), pp. 27-32,
60; and “‘Brick’ Pomeroy and the Democratic Processes: A Study of Civil War Poli¬
tics,” in Transactions of the Wisconsin Academy of Science, Arts and Letters, 1963
(Madison, 1963), pp. 159-169. The role of Edward G. Ryan as a critic of Lincoln and
the war is treated in scholarly fashion in A. J. Beitzinger, “The Father of Copper-
headism in Wisconsin,” Wisconsin Magazine of History, XLIX (Autumn, 1955), pp.
17-25. Both Ryan and Moses M. Strong have found competent biographers — see A. J.
Beitzinger, Edward G. Ryan: Lion of the Laio (Madison, 1960) and Kenneth W.
Duckett, Frontiersman of Fortune: Moses M. Strong of Mineral Point (Madison,
1955).

1966] Klement — Deuster as a Democratic Dissenter 23

See-Bote, and he rose rapidly in importance as a force in the
community.^

The See-Bote dated back to the early 1850's, when it was founded
by Bishop John Martin Henni to counteract the radicalism and
anti-Catholicism preached by several newspapers published in Mil¬
waukee. The Volksfreund, the Flugblaetter, and the Humanist
were anti-clerical, and their editors believed Catholicism the antith¬
esis of freedom and individualism. Alarmed at the anti-Catholic
tone of the press, Bishop Henni authorized the Reverend Dr.
Joseph Salzman to establish a German-language weekly to present
the Catholic point of view upon the issues of the day. Father Salz¬
man founded the See-Bote and boldly entered the conflict. “Radical¬
ism,” he wrote, “has egotism as its basic principle, which will turn
to destruction, if necessary, in order to fulfill the attainment of its
desires.”^ Although the See-Bote was never the “official” newspaper
of the diocese, it published a good deal of church news, and it neu¬
tralized the radicalism expressed by the other German-language
journals.

When Peter V. Deuster took over direction of the See-Bote early
in 1860, sectionalism and abolitionism were national issues, emo¬
tionalizing the country. Several months earlier, John Brown had
stoked the controversy at Harpers Ferry and Southern radicals
talked of secession and separation. Furthermore, 1860 was an elec¬
tion year and Republicans and Democrats argued party politics
heatedly and accused each other of bigotry. William H. Seward and
Salmon P. Chase seemed to be favored by most Wisconsin Republi¬
cans, while Democratic chieftains argued the merits of James
Buchanan and Stephen A. Douglas.

Editor Deuster worked hard to keep his readers from joining
the Republican party. Since prominent Torty-eighters like Carl
Schurz and Bernhard Domschcke were free-thinkers and abolition¬
ists as well as Republicans, it was easy for the editor of the See-
Bote to make a case against the radicalism of the Republican party.
Deuster could also point out that most of the Know-Nothings
(members of a nativist, anti-Catholic movement of the 1850’s) and
most temperance advocates were also Republicans. It was possible
for Deuster, therefore, to convince German Catholics that their
interests were tied to the Democratic party.^

“3 Fairly leng-thy obituaries of Peter V. Deuster appear in the Milwaukee News, De¬
cember 31, 1904, and the Milwaukee Journal, December 31, 1904. The Dictionary of
Wisconsin Biography (Madison, 1960), p. 100, contains a recital of his achievements.

® (Milwaukee) See-Bote, March 1, 1854, quoted in Peter Leo Johnson, Crosier on the
Frontier: A Life of John Martin Henni (Madison, 1959).

4 The rise of the Republican party in Wisconsin is treated in Andrew W. Crandall,
The Early History of the Repuhlican Party (Boston, 1930). Bernhard Domschcke’s
contribution to Wisconsin politics is well discussed in J. J. Schlicher, “Bernhard
Domschcke,” Wisconsin Magazine of History, XXIX (March-June, 1946), pp, 319-
332, 435-456, and Carl Schurz’s link to Badger State history is superbly summarized
in Chester V. Easum, The Americanization of Carl Schurz (Chicago, 1929).

24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55 j

After the northern wing of the Democracy nominated Stephen
A. Douglas for the presidency and the Republicans nominated
Abraham Lincoln, election fever swept the state. Although two
other contenders, John Bell and John C. Breckinridge, entered
the presidential race in 1860, most Milwaukeeans knew that the
race was between Lincoln and Douglas. Deuster criticized Lincoln
less than he did the party which “the Rail-Splitter” represented.
He defined the Republican party as “a conglomeration of isms” —
radicalism, abolitionism, prohibitionism, and Know-Nothingism. He
labeled Carl Schurz, who campaigned for Lincoln, “a political
mountebank” who would do anything for money, incite passions
and encourage fanaticism. He warned his readers not to be de¬
ceived by Schurz’s excursions in oratory nor his claim that Lincoln
would get a large percentage of the German-American vote.^

Although Deuster successfully convinced German Catholics to
repudiate Carl Schurz and vote for Stephen A. Douglas, he could
not prevent Lincoln from carrying Wisconsin by 20,000 votes. Wis¬
consin counties most heavily populated by German-Americans gave
Douglas a majority over Lincoln. The Democratic party schism, the
Lincoln image. President Buchanan’s bungling, and the homestead
plank— not the officious oratory of Carl Schurz — put Lincoln in
the White House in 1860.^^

Dramatic events followed each other rapidly. Southern states
seceded. Peace and compromise efforts failed. The Fort Sumter
affair inaugurated a civil war between the North and the South.
President Lincoln called for volunteers to suppress the “insurrec¬
tion” and issued a number of emergency or extraordinary procla¬
mations. A wave of patriotism swept over the countryside, and flags
flew on every hand, “till the whole Northern heavens seemed a
perfect aurora borealis of stars and stripes.”'

Deuster’s failure to bow to the surge of patriotic passion brought
attacks from several quarters. “Raise the Palmetto Flag at once,”
advised the editor of the Mihvaukee Journal, “and openly declare
that the Government, under which you live, is not to be supported.”
The Republican editor of the Sentinel added, “Those who are not
for their country are against it, and in times of war it is best that
all men should be known.” The next day the Sentinel again criti¬
cized Editor Deuster: “The See-Bote of yesterday, in reply to an
item in the Sentinel, endeavors to squirm out of the unenviable
position in which its secession predilections have placed it. We
again ask our German contemporary a plain question — on which

^See-Bote, October 31, November 7, 28, 1860.

« The Lincoln-Doug-las contest of 1860 in Wisconsin is well summarized in Lloyd
Spohnholtz, “Wisconsin and the National Election of 1860” (M.A. thesis, typewritten,
Marquette University, 1962),

" (Chicag-o) Prairie Farmer^ June 6, 1861.

1966] Klement — Deuster as a Democratic Dissenter 25

side are you? There can be no incivility in this. How many words
are wasted by men when they get in a corner, trying to convince
themselves and the public that facts are optical illusions. We affirm
that unless the See-Bote is in favor of supporting the government,
he [it] is an enemy of the government, and we shall wait with no
little anxiety to see how long an article its editor will write to-day
to mystify and abuse us, when the thing could be settled in half a
dozen words.”®

The Sentinel's comments stirred up some misguided patriots, who
threatened to burn down or destroy the See-Bote establishment.
When fire destroyed Deuster's home in the Fifth Ward several
months later, there was a tendency to suppose “patriots” . had
gained revenge because Deuster and the See-Bote had not given
unequivocal support to the war. The fire, however, was unrelated
to the patriotic passions of the hour, for it had started in a bakery
and spread to nearby houses.'*

The See-Bote's anti-war editorials were mild enough in the early
months of the war. Deuster blamed Republicans for defeating
compromise efforts, he blamed abolitionists for bringing on the
war, and he expressed the fear that the war might evolve into an
abolitionist crusade. He questioned the constitutionality of some
of President Lincoln’s emergency measures, and he predicted that
burdensome taxation and compulsory conscription would be im¬
posed upon the citizenry. Some Democrats, nevertheless, talked of
establishing a pro-war Democratic paper to counteract the See-
Bote's influence and “to neutralize its mischievous effects.” Self-
styled patriots again talked of using the torch. The Sentinel lashed
out at Deuster and the See-Bote for undermining support of the
war “among the German element.” “We will not call the See-Bote
a pestiferous sheet,” concluded one editorial writer; “its influence
iis that of a deleterious miasum [miasma] that mingles with the
purer air of our city, entering the shades of those who have no
safeguards, nor antidotes . . .

Actually, the See-Bote gave the Lincoln administration qualified
support in the first year of the war. Deuster, strongly opposed to
abolition, deplored the pressure exerted upon the president to add
emancipation as a second objective of the war. When Lincoln re¬
voked General John C. Fremont’s proclamation freeing the slaves
of rebels within his jurisdiction, Deuster gave the president a pat
on the back. For a time he envisioned Lincoln to be “a faithful,
conscientious, constitutional ruler” holding back the flood tide of
abolition. He did, however, continue to hope for peace and com-

^ Milwaukee Sentinel, April 18, 19, 1861.

^Ibid., April 19, 20, June 19, 1861; See-Bote, April 25, 1861.

Milwaukee Sentinel, August 9, 21-23, 28, 1861; See-Bote, August 21, 1861.

26 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

promise, wanting “spontaneous action by the people’’ to effect “the
calling of a national convention.” He also pointed out that taxes
were “ruinous” and arbitrary arrests “unnecessary.” Always he
criticized Yankees and the mailed fist of Puritanism. Always he
defended General George B. McClellan, a Democrat, against the
attack of radical Republicans. Yet he asked for popular support
of the “war policies” of the Lincoln Administration, arguing that
since the people had not adequately supported compromise before
the war, they were obligated to complete “the work which they had
endorsed.” A military draft, Deuster re-asserted, would be a
necessity.^i

Deuster’s aversion to abolition turned him against Lincoln as the
president retreated before abolition pressure and as the number
of arbitrary arrests multiplied. Lincoln’s message of March 6,
1862, endorsing compensated emancipation, made Deuster realize
that his fears had a solid basis. He referred to abolitionists as
“disunion demagogues” or “disunion devils,” and he warned that
emancipation would release “a flood of free Negroes and cheap
labor” to rob the immigrant Americans of the crumbs on their
tables. German-Americans, the See-Bote asserted, would lose their
jobs to the “contrabands” (the newly freed slaves). One issue of
the See-Bote carried a long and well-written article entitled “Aboli¬
tion the Worst Enemy of Free White Labor.” Another carried the
story of an employer offering eight contrabands a mere twenty-five
cents a day. Free Negroes, Deuster argued, would be shipped to
northern cities to replace white laborers — he even said that some
abolition-minded employers preferred Negro labor to the immi¬
grant Americans. Using words quite like those earlier enunciated
by Karl Marx, Deuster wrote, “Workmen! Be Careful! Organize
yourself against this element which threatens your impoverish¬
ment and annihilation.” He added, “Let us resist this evil from the
beginning! The North belongs to the free white man, not the Negro.
To him. Nature has provided other regions.

Spouting pessimism, Deuster saw the war as destroying the
ideals which had brought immigrants by the hundreds of thousands.
“It is strange,” wrote the editor of the See-Bote, “that so many
men emigrate to this country. Either the people in the old country
do not know that we have worse times before us, or they are having
worse times themselves. But the latter is not so. The motives which
induce immigration are hope of freedom and lighted taxation. But
neither freedom nor light taxation is to be found here. The taxes
in the future will be heavier and more oppressive than they were

See-Bote, January 3, 15, 21, February 19, 20, March 12, April 19, May 14,
28, July 16, 1862.

^Ibid., April 9, 16, 23, 30, June 4, 1862.

1966] Klement—Deuster as a Democratic Dissenter 27

in Germany^ and as for freedom^ we confess that we are living in
the century of the Bastile and a muzzled press„ The freedom which
we receive from Washington is gone forever, and under the name
of political necessity, the government takes away peu a peu the
constitutional rights of the people/'^'^

As 'Deuster became more critical of the Lincoln administration,
the editor of the Sentinel cracked the whip again. He accused the
See-Bote of substituting partyism for patriotism. He charged that
Deuster was guilty of “false statements'' regarding the state of
affairs in the country. He contended that the See-Bote neutralized
the efforts of the recruiting officers, “spawning its treacherous stuff
for the special delectation of this class [the German Americans]

One letter-writer who signed himself “Union" suggested extra-legal
measures to silence the See-Bote. “Such papers," he wrote, “should
be suppressed. ... If they will continue in their work of treason,
the .people should take the matter into their own hands. I do not
wish to encourage violence, but there is a time when forbearance
ceases, etc. Traitors at home should be dealt with as the common
enemy. Let them beware, 'A word to the wise' etc."^^ Other Mil¬
waukee newspapers added censure. The Daily Wisconsin repri¬
manded Deuster for preaching “a pro-Southern gospel" and for
trying to lead German Americans down the road of treason,^® Wil¬
liam W, Coleman of the German-language Herold offered “to dis¬
continue publication" of his paper and join the army if Deuster
would do the same.^^

Deuster, of course, ignored the criticisms which Republican
editors tossed in his direction. The Lincoln administration lost
popularity during 1862, after time tempered the patriotic passions
which swept the North at the start of the war. The agricultural
and financial depression which engulfed the upper Mississippi area
in 1861 continued to affect pocketbooks throughout 1862. The fre¬
quency of arbitrary arrests gave critics of Lincoln a chance to
chant that a despotism seemed to be enveloping the country. The
ascendancy of New England industry and eastern capital gave
western sectionalists a chance to say that their region was becoming
“slave and servant" of New England. Furthermore, the specious
spirit of Negrophobia was abroad in the land-~™it was widespread
among German- and Irish-American laborers. Deuster excelled
in appealing to the latent spirit of Negrophobia to keep German
Catholics in Democratic party ranks. He had referred to Negroes

^Ibid., June 11, 1862.

'^Milwaukee Sentinel^ June 16, July 22, 29, August 5, 1862.

Letter, “Union” to “Editors, Sentinel/’ August 7, 1862, published in Milwaukee
Sentinel, August 8, 1862,

(Milwaukee) Daily Wisconsin, August 13, 1862.

Milwaukee Herold, August 13, 1862; Milioaukee Sentinel, August 14, 1862.

28 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

as “black cattle’’ and warned his readers that abolitionists intended
to establish a “Negrocracy” in America. He gloated when he heard
that Wendell Phillips, “the orator of freedom,” had been stoned
and mobbed in Cincinnati, and he hoped that Milwaukeeans would
give him like treatment if he appeared in their city. When anti-
Negro riots erupted in Cincinnati and Toledo, Deuster blamed
abolitionists for “those realistic results.” And he referred to the
(Milwaukee) Herold, which endorsed emancipation, as belonging
to “the German Nigger Press.

When President Lincoln finally bowed to abolition pressure and
issued the preliminary proclamation of September 23, 1862, Deus¬
ter acted as if the world had come to an end. He labeled the procla¬
mation “ridiculous,” calling it “an unsavory piece of paper.” He
claimed it was “unconstitutional” and deplored its “consequences.”
It would incite Negroes in the South to insurrection, repeating the
scenes of horror which had been enacted by the Sioux in Minne¬
sota. He again recited his time-worn statement that free Negroes
would flood northern cities and take jobs and security away from
the German Americans. When the editor of the Milwaukee Sentinel
supposed that Negro labor was apt to be “non-competitive,” Deus¬
ter disagreed vigorously, and added, “. . . if free Negro laborers
came to take the place of white workers sent off to war to be
killed, the problem would be less acute.” When the editor of the
Cincinnati Gazette suggested that those who feared the competi¬
tion of the free Negroes could secure “employment” by presenting
themselves “at any of the army recruiting offices,” Deuster’s in¬
dignation knew no bounds. It was plainly evident that Peter V.
Deuster knew how to develop Negrophobia and weld readers of the
See-Bote into a solid Democratic bloc.’^

The election returns of November 4, 1862, heartened Deuster
and other Democrats. Deuster, a candidate for the State Assembly
in the Fifth District, carried his ward by an impressive margin —
726 votes to 364 for his Republican opponent.^® The anti-administra¬
tion tide of the fall of 1862 gave the Wisconsin Democrats three
of the six congressional seats — they were deprived of another seat
in Congress by the Republican-devised political stratagem called

See-Bote, January 28, April 2, June 18, 23, 1862. The sectional and economic
aspects of Democratic and midwestern opposition to the Lincoln administration are
analyzed in Frank L. Klement, “Economic Aspects of Midwestern Copperheadism,”
The Historian, XIV (Autumn, 1951), pp. 27-44, and “Middle Western Copperheadism
and the Genesis of the Granger Movement,” Mississippi Valley Historical Review,
XXXVIII (March, 1952 ), pp. 679-694. Midwestern Negrophobia is discussed and dis¬
sected in Jacque Voegli, “The Northwest and the Race Issue, 1861—1862,” Mississippi
Valley Historical Revieiv, L (September, 1963), pp. 235—251, and Frank L, Klement,
“Midwestern Opposition to Lincoln’s Emancipation Policy,” Journal of Negro History,
XLIX (July, 1964), pp. 169-183. Deuster is quoted in the latter article.

See-Bote, August 13, October 1, 8, 15, 22, November 3, 1862.

^Milwaukee Sentinel, November 14, 1862.

1966] Klement — Deuster as a Democratic Dissenter

29

“soldier-voting-in-the=field’’---and nearly captured control of the
state legislature. Democratic victories in northern states like Illi¬
nois, Indiana, Ohio, Pennsylvania, and New York gave Deuster’s
colleagues a chance to crow and a chance to sponsor “party jollifi¬
cations.’’ One of Deuster’s colleagues, pleased that Democracy was
again in style, composed the headline : “Fall Fashions — Democratic
Victories.

After Wisconsin Democrats finished celebrating their election
victories, they fixed a jaundiced eye upon the pending state draft,
which Governor Edward Salomon, at the request of Republican
politicos, had postponed from August until after the November
election. Early in the war Deuster had supposed that a draft would
be necessary and he restated that contention in mid- July, 1862.
Fully aware of anti-draft sentiment among most Milwaukee Ger¬
mans, Deuster did not hesitate to make political capital out of the
issue. He built up apprehension about the “coming draft” and he
gave publicity to Governor Salomon’s proclamation of August 13,
1862, that all foreign-born citizens who had exercised the fran¬
chise would be enrolled and subject to conscription, even if they
had not applied for their final naturalization papers. The See-Bote
also reported on the “draft disorders” in Pennsylvania, and Deus¬
ter built up a draft consciousness among his readers. He exposed
the postponement of the draft until after the November election as
“a political trick.” He also questioned the constitutionality of con¬
scription, pointing out that the Pennsylvania State Supreme Court
had declared the federal “limping” Draft Act of July 17, 1862 (a
federal act providing for states to conscript and recommending
procedures for a state-conducted draft) , as unconstitutional. Deus¬
ter added his own concern about military power becoming superior
to civil authority. Furthermore, he complained that excessive and
unfair quotas had been assigned to Democratic wards and he im¬
plied that the draft would be administered dishonestly because
abolitionists. Republicans, and Know-Nothings had been appointed
county draft commissioners. And he added that the draft of No¬
vember, 1862, would be but the first of many. “God Almighty only
knows,” concluded the doughty Democrat, “when the drafting will
stop.”^^2

^''-Sheboygan Journal, November 21, 1862. Historians generally interpret the fall
elections of 1862 as “a repudiation” of Lincolnian policy; see Winfred H. Harbison,
‘‘The Elections of 1862 as a Vote of Want of Confidence in President Lincoln,” in
Michigan Academy of Science, Arts, and Letters Papers, XIV (1930), pp. 499-513,
and Harry E. Pratt, ‘‘The Repudiation of Lincoln’s War Policy in 1862 — the Swett-
Stuart Congressional Campaign,” Journal of the Illinois Histoy'ical Society, XXIV
(April, 1931), pp. 129-140. The thesis that soldier-voting was little more than a po¬
litical stratagem is advanced in Prank L. Klement, ‘‘The Soldier Vote in Wisconsin
during the Civil War,” Wisconsin Magazine of History, XXVIII (September, 1944),
pp. 37—47. In the See-Bote of November 12, 1862, Deuster called the election returns
‘‘a popular revolution.”

--See-Bote, July 16, August 6, 20, November 5, 12, 1862.

30 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Governor Salomon and draft officials had every reason to be
apprehensive about the approaching lottery. During the enrolling
process, officers had received a hostile reception at many homes —
several German housewives had used broomsticks to chase out ‘‘the
intruders’" and one Milwaukee woman had thrown scalding water
upon an inquisitor. Enrolling officers expressed amazement at the
embarrassing number who claimed physical disability, set out for
Canada or “the woods,” or filed exemptions as “aliens.” Most men
with families and mortgages worried about the draft. Who would
pay the mortgage when it fell due if the breadwinner marched on
far-away battlefields or was killed fighting the Confederates? Op¬
position to the draft seemed to occur in those counties which held a
predominance of German-Americans. As immigrants arriving in
New York City, many had been met by state agents who described
Wisconsin as a land of milk and honey — a region of freedom and
opportunity. Compulsory conscription seemed to violate promises
made by the state agents. Furthermore, most German-Americans
did not understand what the war was all about, nor did they have
a chance to develop much loyalty to their newly adopted land.
Then too, newspapers like the See-Bote and the (Port Washington)
Ozaukee County Advertiser had dealt rather harshly with the
Lincoln administration, even challenging the propriety and con¬
stitutionality of the draft. The readers of the See-Bote and the
Ozaukee County Advertiser were apt to adopt the editorial views
of those newspapers as their own.

As “D-day” approached, uneasiness and apprehension increased.
Some Milwaukeeans — many from Deuster’s own ward — marched
up and down the streets shouting “No! No!” (“Nein! Nein!”) and
carrying “No Draft” (“Nein Militardienst”) signs. When the cup
of forbearance overflowed in nearby Ozaukee County and brought
forth the Port Washington draft riot of November 10, 1862, the
draft commissioner of Milwaukee County wisely postponed the
draft for a week, until troops from nearby camps could be brought
in to overawe the crowds and supervise the lottery.^^

Deuster received more than a fair share of the blame for the
unrest in Milwaukee and for the Port Washington draft riot. After

-^Milwaukee Banner & Volksfrennd, November 11, 1862; Milwaukee Sentinel,

November 10, 1862 ; See-Bote, November 19, 26, 1862. The story of the Port Washing¬
ton Draft Riot is summarized in Lawrence H. Larsen, “Draft Riots in Wisconsin,
1862,’’ Civil War History, AUI (December, 1961), pp. 421-427, although Larsen fails
to deal adequately with causation. Peter Leo Johnson, “Port Washington Draft Riot
of 1862,’’ Mid- America, I (January, 1930), pp. 212-220, develops the thesis that anti-
Catholic policy in the naming of an army chaplain brought on the draft disorder in
Ozaukee County. John W. Oliver, “Draft Riots in Wisconsin during- the Civil War,’’
Wisconsin Magazine of History, II (March, 1919), pp. 334-337, is most superficial.
Lynn I. Schoonover, “A History of the Civil War Draft in Wisconsin” (M.A. thesis,
typewritten. University of Wisconsin, 1915) is a third-rate study and badly outdated.
The best narrative account of the Port Washington riot can be found in History of
Washington and Ozaukee Counties (Chicago, 1881), pp. 360-366, 493-496.

1966] Klement — Deuster as a Democratic Dissenter 31

all, the See-Bote enjoyed a widespread circulation among German
Catholics in Milwaukee and Ozaukee counties and most of the
rioters were German Catholics. The Milwaukee Sentinel, of course,
blamed Deuster and the See-Bote for creating a climate of appre¬
hension and hostility which transformed citizens into rioters and
mobsters. The (Madison) Wisconsin State Journal derided the
editors of the See-Bote and the Democratic-oriented Milwaukee
News for teaching the German-Americans (this poor, deluded and
ignorant class of men”) that the war had been ''provoked'’ by
abolitionists and that it had evolved into a crusade to free the
slaves. According to the State Journal, Deuster's hands were blood¬
stained and he was clearly guilty of causing the riots. The
Republican-minded editor of the (Hartford) Home League also put
the blame for the riot at Deuster's door. The See-Bote was "a
baneful influence" among "a large portion of our citizens" —
Deuster’s words were "the law and gospel" to many German-
Americans and he could have soothed apprehension instead of agi¬
tating it. Frederick W. Orban of the (Milwaukee) Banner & Volks-
freund tried to blame Deuster and the See-Bote for the discontent,
yet at the same time he felt sorry for his fellow Germans.

The poor misguided ones — because we don’t believe they are anything
else — ^will now realize that their resistance against the law has reacted
adversely. But since their resistance was caused by love for their mostly
poor families — delivered to need without their supporting hands — the
authorities should use mildness as far as compatible with the law. It is
easier to pronounce a harsh judgment on poor, hard-working people when
sitting in a soft, upholstered easy-chair than to bear the miseries of
life without questioned reservation.^^

Questions as to who deserved the blame for the Port Washington
draft riot were also raised in the state legislature. In the state
senate, Herman L. Humphrey pointed his finger in Deuster’s direc¬
tion and asserted that the See-Bote deserved most of the blame.-^
Certainly the editor-publisher of the See-Bote was attacked from
many directions.

Emboldened by the election returns, Deuster refuted Republican-
made contentions that the rioters were poor, deluded, and ignorant.
He ridiculed Republicans for seeming to claim that they had a
monopoly upon virtue, literacy, and learning. He offered no apolo¬
gies for being a critic of conscription or emancipation. He refused
to bow before his critics. Instead he seemed to become bolder and
more aggressive, more critical of the Lincoln administration with

Banner & Volks freund, November 18, 1862; (Hartford) Home League, September
7, 1862, March 14, 1863; (Madison) Wisconsin State Journal, November 11, 1862;
(Madison) Wisconsin Patriot, November 17, 19, 1862 ; Milwaukee Sentinel, November
11-17, 1862.

restate of Wisconsin, Journal of the Senate . . . 1863 (Madison, 1863), pp. 458-459;
(Madison) Wisconsin Patriot, March 13-14, 1863.

32 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55 ;

each passing week. He interpreted the election returns of Novem- ■
her, 1862, as a mandate to cease his qualified support of the war i
and become an all-out critic. He bluntly blamed Republicans for the
deplorable state of affairs.^^ i

Republicans countered with editorial and oratorical blasts at i!
Deuster and the See~Bote. The federal marshal in Milwaukee called >!
Deuster and Christian Ott, who wrote most of the editorials for the i
See-Bote, into his office, trying to intimidate them and threatening !
arrest.“‘ The Union general commanding the Army of South-east ;
Missouri, through his provost marshal (Major Gustavus Heinrichs) |
forbade the circulation of the See-Bote in his sector and among his :
soldiers. He claimed that Deuster’s newspaper rendered “aid and ;
comfort to the enemy’’ and issued a boycott. “Public journals using I
as mean and disgraceful language as this paper,” concluded the !
military edict, “is [szc] injurious to military discipline, and is not 1
the literature to be tolerated in the army.”-® I

The edict suppressing the See-Bote in one military sector stirred |
up a controversy, and the issues of treasonable conduct and free !
press received a public airing. Republicans generally endorsed the j
army edict and Democrats claimed that a constitutional guarantee i
had been violated. The issue received a hearing in the State Assem- ;
bly, where Deuster sat as a member. Andrew J. Turner, of Portage, ;
introduced a resolution which gave “hearty approval” to Major ,
Heinrichs’ action in expelling the See-Bote from his department.
The resolution was referred to the Committee on Federal Relations, j
The majority report, dated March 31, 1863, endorsed Major Hein- j
richs’ edict. “We are of the opinion,” the report read, “that all '
such newspapers should be suppressed in and out of the army |
lines.” The report concluded : ;

In times like these there is no neutral ground. We are either for the j
government or against it — either patriots or traitors. We cannot be loyal ^
to the government and disloyal to the administration, ... We, therefore, 'j
regard the sentiments promulgated by the See-Bote ... as of the most jj
dangerous character; and that Major Heinrichs was fully warranted in '
prohibiting its circulation in the army under his command.^® |

Democratic members of the Committee on Federal Relations, :
quite naturally, disagreed with the majority report. Alden S. San- i
born, of Madison, presented a dissenting report which defended ;
Deuster and freedom of the press. Sanborn’s minority report de- ■!

- I

See-Bote, November 19, December 10, 17, 31, 1862, January 7, 14, 21, 1863. ;

^ Ibid., January 21, 1863. j

28 The edict, dated January 12, 1863, and signed by Major Gustavus Heinrichs as
“Provost Marshal General, Army of Southeast Missouri,’’ was published in the Mil¬
waukee Sentinel, January 27, 1863. For some unknown reason, the document does
not appear in Official Records of the Union and Confederate Armies (128 vols., 1880-
1901).

20 State of Wisconsin, Journal of the Assembly . . . 1863 (Madison, 1863), pp. 106, ,

123-124, 895, 956-957.

1966] Klement — Deuster as a Democratic Dissenter 33

scribed the See-Bote as ''the uncompromising friend of the people,
firmly attached to the principles of liberty, an unwavering advocate
of the restoration of the United States into the same fraternal
relations that existed before sectional parties menaced their
disruption/’^^ Democratic and Republican legislators, wearing par¬
tisan spectacles, saw the same act assuming different shapes.

Neither the Sentinel’s fulminations, the threats of a federal
marshal, nor a general’s edict checked the See-Bote's criticism of
the Lincoln administration. When Congress discussed the need for
federal conscription during February, 1863, Deuster printed his
anti-draft views. He claimed that federal conscription would de¬
stroy civil liberties of individuals as well as the sovereignty of the
states. It would keep the Republican party “permanently in
power,” wiping out the opposition party. The presidency would
evolve into a dictatorship and the republic turn into a despotism.
Negro troops, Deuster warned his wary readers, might even be
employed to enforce the draft and drag white men off to war, in¬
sulting them in the process. Yes, “compulsory conscription” and
“the excesses of the Administration” might even force the liberty-
loving people of the North “to the edge of the chasm,” bringing
civil war to them,^^

After the Conscription Act of March 3, 1863, became law, Deus¬
ter continued to play critic. He compared the federal measure to
“the Polish forcing act,” reminiscent of the drafting of the Poles
by the Russian government. German Americans, Deuster asserted,
would be sacrificed at the whim of New England Yankees. Deuster
also criticized “the $300. commutation clause,” a provision which
absolved a man of military service upon the payment of $300. Rich
Republicans, the See-Bote supposed, had incorporated that “iniqui¬
tous section” into the Conscription Act so that they might stay at
home while the poor immigrant Americans would then die upon
the battlefields.^^

Other issues besides federal conscription drew the wrath of
Peter V. Deuster and the See-Bote. When the Lincoln administra¬
tion carved West Virginia from the northwestern section of “the
Old Dominion,” in violation of a constitutional clause guaranteeing
the integrity of each state, Deuster printed his protest, labeling
such action unwarranted and unconstitutional. When successive
issues of greenbacks or legal tender notes were authorized by a
Republican-dominated Congress, Deuster cried “Foul !” and claimed
that property rights were sacrificed and inflation sanctified. When
Congress raised the tax on distilled spirits and considered doubling

30 Ibid., pp. 958-959.

See-Bote, February 11, 1863.

^^Ibid., March 25, April 20, July 29, 1863.

34 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

the levy on fermented liquors, the See-Bote again raised its voice.
Deuster believed beer “the healthiest and most innocent alcoholic
drink.” The proposed beer tax would fall heaviest upon the labor¬
ing classes and those immigrant Americans whose cultural pat¬
terns made them beer-drinkers. Deuster also criticized the removal
of General George B. McClellan from command of the Army of the
Potomac and decried the arrest of Clement L. Vallandigham, prom¬
inent Copperhead and critic of the Lincoln administration, early
in May of 1863. He called the trial of Vallandigham of Ohio by a
military commission in an area where the civil courts were open
“an outrage,” arguing that force and arbitrary measures had been
substituted for wisdom and justice. He applauded when Ohio Dem¬
ocrats retaliated by nominating Vallandigham as their party’s
gubernatorial nominee. Such bold action, Deuster argued, was a
proper protest against “usurpation and tyranny.” As far as Deus¬
ter was concerned, the wheel of revolution turned too fast and too
far. The “more radical measures of the Lincoln Administration”
could be compared with “the excessive measures of the French
Revolution.” Carl Schurz, a onetime Milwaukeean, seemed to be
one of Deuster’s favorite targets. The See-Bote seldom missed a
chance to throw mud at Schurz, who vainly sought military glory
upon Civil War battlefields. Deuster considered General Schurz
“incompetent” and “egotistical,” qualified only to carry a gun in
“Wide-Awake parades.

A peace movement gathered momentum during the first six
months of 1863 and Deuster jumped with alacrity upon that band¬
wagon. Continued war might crush out the last vestiges of civil
rights, for it continued to centralize the government. The weary
boatman at the river Styx ferried heavy loads, and the people on
the home front tired of the bloodshed and shuddered at the long,
long casualty lists. Defeatism became more and more widespread
as some became convinced that the South could not be conquered.
Then, too, ruinous taxes sapped the nation’s economy and robbed
men of their hard-earned dollars. Many Democrats were convinced
that the original objective of the war had been perverted. New
England capital seemed to have moved into the driver’s seat, using
Lincoln as a pawn in its game to make western interests servile to
eastern interests. Conciliation and compromise could stop the
bloodshed and the centralization of the government, giving mid-
westerners a chance to regain the balance of power they had held
politically before the war. “When will the hideous moloch who
holds the press and sword of this nation,” asked one of Deuster’s

^Ihid., December 24. 1862. May 7, 13, June 10, 17, July 2, October 14, 1863.

1966] Klement — Deuster as a Democratic Dissenter 35

friends, ‘‘call off his dogs of war, and suffer peace once more to
bless our bleeding country

Just when war weariness and defeatism seemed to be taking over
the northern heart, the fortunes of war changed. The tide turned
at Gettysburg and Vicksburg early in July, 1863. The peace move¬
ment then retreated, for it vacillated with the vicissitudes of war,
advancing with Union defeats and ebbing with Union victories.
Deuster, evidently convinced that a draft was necessary, quit criti¬
cizing the Conscription Act and turned, instead, to promoting “a
social plan” to help needy draftees. Deuster’s devastating criticism
of the administration also seemed to soften, giving way to mildness.
Early in November, 1863, the See-Bote even printed an advertise¬
ment from the general government. The editor of the Milwaukee
Sentinel, somewhat chagrined, protested, asserting that the Herold
was more deserving than the See-Bote, ‘‘Such being the case,” con¬
cluded the editor of the Sentinel, “We hope no further official pa¬
tronage will be bestowed upon a paper which is doing all it can
to embarrass the government. Let the proper authorities look to
the matter.”^^

Early in 1864, Republicans and Democrats began to talk of presi¬
dential candidates and to weigh the possibility of Lincoln’s re-
election. Union military victories, like those at Gettysburg and
Vicksburg, combined with Republican political victories at the polls
in October and November, 1863, gave Lincoln a claim to renomina¬
tion and re-election. Although some dissident Republicans favored
John C. Fremont as a candidate, the party’s national convention put
Lincoln’s name at the head of the ticket. From the first, it was
almost a foregone conclusion that Deuster’s party would name
George B. McClellan as its choice in the presidential contest of
1864. The presidential race stirred party ism and Deuster joined
other Democrats in denouncing Lincoln and praising McClellan.
The See-Bote seemed to delight in reporting critical comments
made by Deuster’s fellow Democrats. Edward G. Ryan, prominent
party mogul, had described Lincoln as “a weak, vain, amiable man”
characterized by “his utter imbecility and . . , moral incapacity”
— “a mere doll, worked by strings.”^® Mayor Abner Kirby, another
Milwaukee Democrat, labeled Lincoln “a weak and vacillating
president” and “a tool of fanatics” — “the weakest man on the whole
list of presidents.”^' Deuster, who had earlier judged President
Lincoln “the most incapable of statesmen and the most irresponsible
of butchers of men,” predicted that history would deal harshly with
the president. He claimed that Lincoln’s nomination was made at

Sheboygan Journal, April 9, 1863.

^See-Bote, November 5, 1863; Milwaukee Sentinel, November 6, 1862.

Quoted in the Milwaukee News, July 2, 1863; See-Bote, October 14, 1863.

See-Bote, April 24, 1864.

36 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

a convention dominated by rascals — “griffins, hypocrites, pharisees,
shoddy contractors, and hwo-legged cattle’/’ There was not “an
honest man in the whole convention.” These “hell-on-earth men”
nominated Lincoln, Deuster wrote, despite “the sighing of the
widows, the complaining of the children, and the moaning of the
wounded upon the battlefield.” Lincoln had no “conscience;” he
was guilty of telling smutty stories while soldiers were dying. The
See-Bote associated “Godlessness,” “perjury,” “irresponsibility”
and “dirty ditties” with Lincoln and radicalism. Evidently God
was punishing the nation for the sins of the Lincoln administration,
imposing suffering, taxes, and hardship upon the people because
radicals and fanatics directed affairs in Washington.'"^^

The See-Bote, of course, endorsed McClellan’s candidacy and
placed his picture on the front page. It reviewed McClellan’s qualifi¬
cations most favorably, praising the man and the soldier. McClellan
could lead the nation out of “the desert of troubles” and into the
promised land. Lincoln’s policy was failure; it was time for a
change. Lincoln’s re-election would mean more “troublous times”,
more drafts and more bloodshed.'^**

Party loyalties and the heat of political campaigns sometimes
prompts men to make irrational statements. Such was the case
when “Brick” Pomeroy of the LaCrosse Democrat hoped that “some
bold hand” would pierce Lincoln’s heart “with dagger point for the
public good.” Such was the case when the editor of the Beaver Dam
Argus wrote, “History shows several instances where the people
have only been saved by assassination of their rulers, and history
may repeat itself in this country. The time may come when it is
absolutely necessary that the people do away with their rulers in
the quickest way possible.” And such was the case when Deuster
wished Lincoln were dead. Upon hearing the story that a soldier
standing next to Lincoln (while the President was visiting the
front lines) was wounded by a bullet, Deuster wrote, “Oh, if a
fortunate coincident had caused that bullet to pierce the black,
inhuman heart of this great butcher of men, rather than lodge in
the leg of the poor soldier/’-^^

Despite the heat generated by newspaper editors and excited
orators, the election passed off with few incidents in Milwaukee
and Wisconsin. Although Deuster’s ward and the city of Milwaukee
gave McClellan a two-to-one margin (4,908 votes for McClellan,

^ Ibid., December 18, 1862, June 15, July 27, 1864.

Ibid., September 14, October 25, 1864.

^0 La Crosse Democrat , August 23, 1864 ; Beaver Dam Argus^ September 14, 1864 :
See-Bote, August 3, 1864. An account of the election of 1864 in Wisconsin can be
found in Frank L. Klement, “Wisconsin and the Re-election of Lincoln in 1864 :
A Chapter of Civil War History,” in Wisconsin in Three Wars [Historical Messenger,
XXII (March, 1966)] pp. 20-42.

1966] Klement — Deuster as a Democratic Dissenter 37

2,535 for Lincoln), Lincoln carried Wisconsin by 17,000 votes and
won re-election by a comfortable margin'd^

Deuster was not surprised by the election returns, but he ex¬
pressed his disappointment nevertheless. His editorials seemed to
say, “The Republicans started this war ; let them finish it/' “Every¬
one," he wrote, “views the future Avith apprehension and anxiety."
He seemed depressed and disgusted, and he tossed another taunt in
Lincoln's direction: “His watchword is war- — that's what the vote
meant. Disintegration of the country, with the end of civil order
and collapse of the government, will come. Then the people, de¬
ceived by Lincoln, will wake up and realize their plight/'^'^

In the closing months of the war Deuster remained a caustic
critic and an unredeemed Democrat. He referred to the country's
president as “a usurper,” and he moaned each time the president
called for more troops.^'^ Yet Deuster and most Democrats sym¬
pathized with President Lincoln when he feuded with the radicals
in his own party over Reconstruction policy. Lincoln favored a
rather mild Reconstruction policy, whereas the radicals wanted
vindictive measures and civil rights for the newly freed Negroes.
Democrats like Deuster and Edward G. Ryan openly supported the
president against most leaders of his party.

Lincoln's death at the hands of an assassin shocked Peter V.
Deuster. He feared that the president's death might give the radical
Republicans control of Reconstruction and that “retribution and
revenge” might become official policy. Deuster even claimed that
Democrats could mourn with a “pure conscience,” He rationalized
for his readers: “We have voted against Lincoln's election; written
against it; spoken against it. What we have said and written was
done with a clear conscience. We may say with an equally clear
conscience that there are no more sincere mourners today- — none
who deplore the death of President Lincoln more than the Democ¬
racy of the Northern States,”^^

Deuster's role as a Copperhead and critic of Lincolnian policy
did not adversely affect his business or political success. The See-
Bote became a prosperous business enterprise in the postwar years.
For two yeans, until the “Great Chicago Fire” of 1871, he also
published the Chicago Daily Union, another German-language
neAvspaper. During the postwar years he again sought public office,
serving one term in the state senate and three in Congress. In the

The soldier vote padded the rather scant majority Lincoln received of the home
vote in Wisconsin. The canvassers counted 68,906 Lincoln votes and 62,494 McClellan
votes cast in Wisconsin — they set aside the Kewanee County votes (157 for Lincoln,
753 for McClellan) because “no seal was attached.” Lincoln received 13,805 of the
16,789 votes cast by soldiers in the field,

See-Bote, November 23, December 14, 1864.

^ Ibid., February 11, 25, 1865; Milwaukee Sentinel, February 27, 1865.

^Milwaukee Sentinel, April 26, 1865.

38 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

postwar era he gained recognition as the most forthright and re¬
spected champion of the wants and rights of the German Catholics
of the Milwaukee area.

Peter V. Deuster can be classified as a conservative. He opposed
the changes which the Civil War imposed upon his adopted country.
He opposed the centralization of the government, for the war
helped to transform a federal union into a truly national state. He
opposed the triumph of industrialization and its ascendancy over
agriculture, objecting to the trend which caused the upper Midwest
to bow to the economic domination of the Northeast. He opposed
the extension of democratic rights to the former slaves ; he opposed
emancipation and the granting of civil rights to the newly free.
Yet he was a leader and spokesman for many German Americans
because he could put into words the hopes and the fears of his
countrymen, immigrant Americans adjusting to their American
environment.

GEORGE MADISON HINKLEY
SAWMILL ENGINEER FOR E. P. ALLIS

Walter F. Peterson
Lawrence University

In 1905 the Americcm Lumberman at the death of George M.
Hinkley, honored him with a special article. He was “among the
men who had done much to elaborate and perfect saw mill ma¬
chinery.’’ His contribution to sawmilling was widened when “for¬
tune cast his lot with one of the largest machinery manufacturing
houses in the country or the world/’ Edward P. Allis and Com¬
pany. The career of G. M. Hinkley, master sawmill designer and
builder, cannot be separated from that of E. P. Allis, whose Re¬
liance Works in Milwaukee, Wisconsin, manufactured the ma¬
chinery that made Hinkley famous among lumbermen.^

Edward Phelps Allis (1824-1889) was a New Yorker who turned
from the study of law to go West to seek his fortune as a business¬
man. By 1873 Allis had established himself as a leader in the
Wisconsin business community, had purchased and expanded the
Reliance Works founded in Milwaukee in 1847 by Decker and
Seville, and employed more than 300 men and apprentices. Mill¬
stones and mill supplies, along with castings and engines, were the
principal products. Although sawmill equipment had been listed in
the catalog for some years, it was no more than a minor line.^
Allis developed a technique of management that made him the
largest manufacturer in Wisconsin in the late nineteenth century.
“It has been Mr. Allis’ policy to secure the assistance of the best
specialists in the different lines of machinery manufacture, and
thus turn out the best machinery made, to which is due in a large
measure his great success,” reported an observer.^ Allis brought
together the engineering talent for the production of goods and
the financial support to secure the constant expansion of his works.
It was up to his engineers to provide the excellence of product and
efficiency in production that would yield profits.

In 1873 Allis invited George Madison Hinkley to become head of
the Reliance Works’ sawmill department. Hinkley was one of the
men who made up an engineering triumvirate which would lead

^American Lumberman, December 23, 1905, p. 1.

^Dictionary of American Biography (New York, 1928), pp. 219-220, Milwaukee
Sentinel, April 2, 1889.

^Sentinel, January 2, 1889.

39

40 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Allis and the Reliance Works to international fame and financial
success. The second major appointment was that of William Dixon
Gray to head the flour milling department. E. P, Allis rounded out
his staff of brilliant engineers by securing in 1877 the services of
Edwin Reynolds, who became the great steam engine builder of the
late nineteenth century. The American Society of Civil Engineers,
which had invited Allis to become a Fellow in 1883, published this
appraisal of his successful business technique : “Mr. Allis was not
an engineer, not an inventor, not a mechanic, but he had in full
measure that rare talent for bringing together the work of the
engineer, the inventor, the mechanic, that it might come to full
fruition, and the world at large be the gainer thereby.”^ The
achievement of Edward P. Allis was based on the success of Hink-
ley's sawmill equipment, Gray’s flour milling inventions, and Rey¬
nolds’ steam engines, which powered the sawmills and flour mills.
As it turned out, E. P. Allis could not have picked better men than
Hinkley, Gray and Reynolds.^

Allis and his engineers could hardly have lived at a better time.
After the wreckage of the depression of 1873 had been cleared
away, the United States very rapidly developed to maturity as an
industrial nation. From an economic point of view the period 1873
to 1893 was in some respects a golden age of American history.
During this period the public debt was rapidly reduced, even
though taxes were low. The federal government was usually more
concerned with a surplus than a deficit. Gradually, after the violent
shock of civil war, the spiritual unity of the nation was restored.
Manpower resources were unlimited as young and ambitious Euro¬
peans settled in cities and on farms. Inventions of all kinds added
greater comfort and convenience to daily life. But most of all,
there was a consciousness of progress, development and growth
which made possible an optimism in American life that has per¬
haps never been so great.

E. P, Allis was always alert to business possibilities. The lumber
industry, found in his own back yard, presented a remarkable op¬
portunity. Given the enormous stands of accessible timber, Allis
might almost have anticipated that between the Civil War and
1890 the principal center of the lumber industry would be the Great
Lakes region. In fact during that period Michigan and Wisconsin
accounted for nearly 30 per cent of the national production. During
the decade following Hinkley’s appointment as manager of the
sawmill department, the quantity of white pine sawed annually in
the Great Lakes area was to double, increasing from roughly four

^Proceedings of American Society of Civil Engineers, 1889. Louis Allis Scrapbook,
VoL 1. Courtesy of Mrs. Louis Allis, Milwaukee, Wisconsin.

® Walter P. Peterson. “E. P. Allis : A Study in Nineteenth Century Business Tech¬
nique,’"’ Marquette Business Review, Fall, 1962. pp. 44-48.

1966]

Peterson — George Madison Hinkley

41

billion to eight billion feet. Moreover, the industry was soon to de¬
velop in the West and in the South. During the decade of the
eighties the total value of the product was to increase from $210
million to $404 million. Supplying the rapidly expanding lumber
industry with equipment represented an enormous opportunity.®

The change that Allis must have noted was that sawmill methods
during the previous decade had been undergoing a rapid develop¬
ment. Introduction of the circular saw increased cutting capacity
more than ten times, although early circular saws were exceedingly
wasteful, sawing out at each cut a half inch of kerf. The movements
of the log carriage had been accelerated and the double edger and
later the gang edger had been introduced. At the close of the sixties
steam replaced manual labor in handling logs. These and many
other lesser improvements were accompanied by the increasing
efficiency and power of the driving engines. In short, the better
sawmill of 1870 bore little resemblance to the mill of 1860, and
was still improving.'^

Hinkley, born in Seneca, New York, May 24, 1830, was appointed
head of Allis' sawmill department in October, 1873. As a young man
he had recognized the great future in the lumber industry and in¬
creasingly aware of his growing taste for mechanical work, decided
to learn the millwright trade. His first effort in this new occupation
was in 1851 on a mill at Zilwaukee, Michigan. He then worked on
mills at East Saginaw and Thetford, Michigan, and one on the
Flint River.®

The Civil War broke out while Hinkley was operating a shingle
mill in Tuscola county. On September 11, 1862, he enlisted as a
corporal in Company 1, Sixth Michigan Calvary, and was mustered
into service on October 11, 1862. On May 6, 1864, Hinkley, now a
sergeant, crossed the Rapidan with General Grant and on June 11
he was captured by the Confederate forces during the battle of
Trevellian Station, when his horse was shot from under him. As
a prisoner he was confined in Confederate prison camps, including
Anderson ville, until he was paroled in late November, 1864.®

After the war Hinkley was kept busy building mills; first the
Farr mill at Muskegon; then a mill at Manistee; and in 1866 a
shingle mill in Milwaukee. After its completion, John Eldred, the
owner, engaged Hinkley as the operator. In 1870 Hinkley decided

6 Victor S. Clark. History of Manufactures in the United States (New York, 1929),
11, pp. 482-3.

"^Frederick Merk. Economic History of Wisconsin During the Civil War Decade
(Madison, 1916), pp. 69-71.

^Dictionary of Wisconsin Biography (Madison, 1960), p. 171. American Lumher-
man, December 23, 1905, p. 1.

^History of Milwaukee, Wisconsin (Chicag'o, 1881), p. 1288. AlKs-Chalniers Sales
B'lilletin, December, 1905, p. 1. Hinkley kept a diary during the Civil War. Although
the original has been lost by the family, portions of the diary in typescript are in the
files of the Allis-Chalmers Manufacturing Company, Milwaukee, Wisconsin.

42 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

to develop his ideas for improving sawmill machinery and to estab¬
lish his own business. He invented and sold a saw swage, a mill
lathe and other devices which Filer and Stowell, sawmill manufac¬
turers in Milwaukee, produced for him. His worth and his potential
as inventor and engineer moved E. P. Allis to hire him for the
Reliance Works.

Upon joining the Allis company, Hinkley contributed his sense
of organization, his drive, his inventiveness and his engineering
abilities. Actually his productivity in new sawmill devices had just
begun, for during the 32 years that he was head of the sawmill
department he patented 35 inventions. So that Allis might secure
not only the services of such inventive minds as Hinkley, and later
Gray and Reynolds, but also keep them in his organization, he
allowed departmental managers to hold all or part of their patents,
as well as those of their departmental co-workers. The company
then paid the managers for the use of their patented devices. More¬
over, the name plates on machines and company catalogs frequently
featured the name of the department head, thus giving him inter¬
national recognition.^-

When George Madison Hinkley came to the Reliance Works, the
annual sales of sawmilling equipment had not reached $1,000.
Hinkley poured all of his talent and energy into his job. At the
outset he did all the drafting, traveled, and carried on the corre¬
spondence. Most of the machinery turned out was under Hinkley’s
patents and his genius was such that some of the mill appliances
invented by him were manufactured and used in mills without
marked change for two to three decades afterward. As the reputa¬
tion of the Reliance Works and of Hinkley ’s inventions grew, so
did the sales of sawmill equipment.^^

Logging was a rough, tough business in the late nineteenth
century and the sawmill owners were a hard-bitten lot. It took a
particular type of person, besides the quality of the product, to sell
effectively to them. Hinkley was known for his commanding bear¬
ing, hiis forceful manner and pungent speech. A fine beard added
to his impressive appearance. His outbursts were considered classic.
One sawmill man vividly remembered his ‘‘highly scientific and
gifted knowledge of picturesque language.” Once when something

American Lumberman, December 23, 1905, p. 1.

^iW. H. Whiteside, President of Allis-Chalmers Company, Circular Letter No. 62,
December 20, 1905.

^ The Sentinel, February 29, 1888, notes that a patent was granted on a sawmill
carriage, one-half to George M. Hinkley and one-half to E. P. Allis and Company.
The contract, in Allis-Chalmers flies, between William W. Allis, President of Edward
P. Allis Company, and Edwin Reynolds, April 9, 1890, reaffirmed his previous con¬
tract, which gave him full right to his patents. Ernest C. Shaw, who knew G. M.
Hinkley well, understood that Hinkley held the same rights to his patents as did
Reynolds and also some patents of departmental co-workers. Edward P. Allis and
Company Catalog, 1885.

American Lximberman, December 23, 1905, pp. 1, 37.

1966]

Peterson — George Madison Hinkley

43

had gone wrong, Charles Allis, the third oldest of the Allis boys,
rushed out of his office to suggest less profanity, only to give up
when G. M. Hinkley furiously expanded on his original statement
with even greater force and added that he would “kow-tow to
nobody But he understood the loggers and sawmill owners and
could speak their language. Here was a man who knew what he
wanted and had the courage and ability to go after it. Hinkley
employed no tricks of salesmanship but sold the products of the
Reliance Works solely on their merits, “recommending them for the
value that was in them, and of that value and its most minute de¬
tails no man ever had more intimate and thorough knowledge.”^'"^

When George M. Hinkley assumed management of the sawmill
department, the company produced only a circular saw which was
described as a fast-running disc “with teeth on its periphery.’'
Only two years after Hinkley joined the Reliance Works, the catalog
of the sawmill department was increased to a fat 70 pages. Hink-
ley’s patents, together with his ingenuity and energy, had made the
difference.^^

In 1876, three years after joining the Allis Company, Hinkley
sent his first complete sawmill to Japan, and filled many larger
domestic orders as the reputation of the department continued to
grow. In the spring of 1878 ten carloads of sawmilling equipment
were sent to Texas, including two large double sawmills, setworks,
engines, boilers, and everything necessary for a complete outfit.
Later the same year the Sentinel reported that “in the matter of
sawmills the reputation of Messrs. Allis & Co. stands alone.

In the hard-fisted and free-wheeling sawmilling business a less
energetic man than Hinkley and a smaller concern than the Edward
P. Allis Company would have had difficulty maintaining the iden¬
tity and integrity of its patents. The mechanical “dog,” the device
to hold the log in place on the log carriage, was of critical impor¬
tance. In 1880 the Allis Company brought suit against Filer,
Stowell and Company for infringement of a patent dog used in
sawmills. Allis and Hinkley sought to recover royalties from all
firms that had manufactured or were using their patented device
to the extent of $600 to $800 for the use of the dog during past
years and recognition of rights in the future. When the Allis posi¬
tion was sustained by the courts, the lumbermen of Oshkosh, Wis¬
consin, formed the Northwestern Sawmill Protective Association
to defend themselves against an additional Allis claim of 25^ per

■•^Ernest C. Shaw to Alberta J. Price, August 23, 1954, Axel Soderling- to Alberta J.
Price, August 3, 1954. Interviews in Allis— Chalmers historical files.

American Lumberman, December 23, 1905, p. 37.

Edward P. Allis and Company Catalogs, 1871, 1875.

^'’Sentinel, October 9, 1876 ; March 19, 1878; May 15, 1878.

44 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

1000 feet of lumber cut by mills using its devices if not manufac¬
tured by the Reliance Works. The decision on this claim was in
favor of the Allis company and a referee was appointed to deter¬
mine the extent of the damages. Allis and Hinkley continued to
press their claims against a growing list of firms and lumbermen.
The first case, against Filer, Stowell and Company of Milwaukee,
was settled in 1883 when that company agreed to pay for past in¬
fringement and take out a license from E. P. Allis and Company
covering future use of the patent. This action provided the prin¬
ciple for settlement of the remaining cases.’ ^

At the fairs and exhibitions popular after the Civil War manu¬
facturers of all types entered their products in competition for
prizes and to widen their markets through the education of the
public. Hinkley supervised elaborate displays of Allis sawmill
equipment all over the country during the seventies and eighties.
The progress made by Hinkley in developing a first class sawmill
department can be seen in the impressive collection of prizes
awarded his sawmill equipment at the New Orleans World’s Fair
of 1885. For a circular sawmill in practical operation he was
awarded a medal of second class; headblocks in operation with
circular sawmill, medal of second class; collective display of saw¬
mill machinery, medal of second class; gang edger, medal of first
class; automatic lumber trimmer, medal of first class; two-saw
lumber trimmer, honorable mention; flooring machine, medal of
first class; for the Reliance mill dogs, operated with circular saw
mills, medal of first class. This record becomes more impressive
when it is compared with those of two other Milwaukee manufac¬
turers who also entered their equipment at the New Orleans fair.
Filer, Stowell and Company received honorable mention for its dis¬
play of mill machinery, and the T. H. Wilkin Company a medal
of first class for its saw stretcher. G. M. Hinkley’s sawmill depart¬
ment was obviously helping to establish the national and inter¬
national reputation of the Allis company.’^

Although Hinkley did not invent the band saw, he is given credit
for perfecting it.^’’ This was a machine carrying a saw made from
an endless steel band with teeth on one edge running over two
flat-faced wheels, one above and one below the level at which the
log was sawed. The great advantage was that the steel band was
one-half the thickness of the old circular saw and reduced the
waste from sawdust proportionately at every cut. When Hinkley

Sentinel, August 16, September 27, 1880; January 29, 1881; October 4, 1882;
March 22, September 2 1883.

Sentinel, May 23, 1885.

'■^American L^imber7nan, December 23, 1905, p. 1.

1966]

Peterson — George Madison Hinkley

45

was convinced that the band saw could work a great advantage,
he proceeded to perfect it,^^

With his characteristic skill and energy Hinkley pushed the de¬
velopment of the band mill. His first band mill was announced on
December 6, 1885, in a notice entitled ‘TO THE ATTENTION OF
LUMBERMEN.^^

We have just completed our new band saw mill, which is without ques¬
tion, the best machine of its kind ever offered to the market. One of these
mills is now set up at our works, corner of Florida and Clinton Streets
where it will remain on exhibition until December 15. It will then be re¬
moved to Dorchester, Wisconsin, and placed in active operation about
January 1 in the mill of the Jump River Lumber Company. We make
this announcement in order that parties interested in band saw mills may
have an opportunity to inspect our machines.^^

This was a nine-foot mill designed for saws ten inches wide. The
lower wheel had a cast-iron rim on the outside of which was
bolted a hardwood rim. The weight of this lower wheel was about
3,000 pounds. The top wheel was constructed almost entirely of
the best seasoned hardwood to make it as light as possible and at
the same time perfectly rigid:^^ Soon after the new band mill was
placed in operation at the Jump River Lumber Company, Prentice,
Wisconsin, the E. P. Allis Company received the following letter :

Your combined Band and Rotary Mill put in for us was started up
about the first of February last. It started off perfectly and our satisfac¬
tion has been constantly increasing. We are cutting from mixed logs,
knotty, frozen, shaky and sound, at the rate of 3,000 feet per hour, of
measured lumber, requiring no more care than a circular mill. We expect
with a little more familiarity with operating the mill, to saw 35,000 feet
per day. We have examined other mills in operation and unhesitatingly
say we have seen none that compare favorably with this one. We cordially
recommend anyone desiring a mill to examine this one in operation.

Jump River Lumber Company"*

Although the later development of the Hinkley Automatic Power
Swage and the Hinkley Power Guide, along with his other numer¬
ous inventions, rounded out his contributions to the sawmill indus¬
try, it was the perfected band saw that the American Lumberman
regarded as “the monument of his rare genius and mechanical
ability.^’^^

21 Ernest C. Shaw to Alberta J. Price, August 23, 1954. Allis-Chalmers historical
files. It was characteristic of all the sawmill developments of the sixties and seventies
that they were calculated to secure increased output or a saving- of labor. Little effort
was made toward achieving- a saving of timber which was both cheap and abundant.

^-Sentinel, December 6, 1885.

23 In the Southern huniherman, December 15, 1931, p. 82, E. A. Hall, then manager
of the milling machinery department of Allis-Chalmers, provided details on construc¬
tion of the mill.

2i Jump River Lumber Company, undated letter in Allis-Chalmers historical files.
American Lumberman^ December 23, 1905, p. 1.

46 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

It is significant that Hinkley lived and produced his equipment
during the period of greatest lumber expansion, when every manu¬
facturer of sawmilling equipment was pushed to the utmost to meet
both the great demand and the intense competition. At his death
in 1905 the American Lumberman paid him tribute:

Mr. Hinkley was as great a man in his line of business as Carnegie in
his. He has been as useful in his day and generation, in view of the cir¬
cumstances which surround him, as any great inventor whose name could
be mentioned. His relation to the improvement of saw mill machinery was
almost akin to that of Edison to electrical development or of Ericson to
the evolution of naval construction. Had he so elected his name would
have been as eligible to enrollment in a national hall of fame as any of
those cited. But he chose — if he gave that matter a thought — that his
works should be his monument.^®

Hinkley distinguished himself within the company well beyond
his ingenuity as an inventor and machinist. It was the business
ability of George Madison Hinkley that E. P. Allis prized equally
highly. With the management and the sales of the department
wholly in his charge, he raised the status of his department to the
first rank in the field and annual sales to nearly $400,000 by 1889
when Allis died. Hinkley had vindicated the business technique by
which E. P. Allis operated and whose fortunes, in part, were cre¬
ated by him.^^

^Ibid., p. 37.

After more than 32 years of service to the company as manager of the sawmill
department, G. M. Hinkley died on December 14, 1905.

WISCONSIN TERRITORIAL AND STATE CENSUSES

Walter H. Ebling

Department of Agricultural Economics
University of Wisconsin

State census work can be best understood against the background
of the important and excellent United States censuses. Although
the national census organization in the U.S. has become perhaps
the world’s best, the development from a simple beginning in 1790
to the present was slow, at least in the early decades. Art. A, sec.
2 of the U.S. Constitution provided for the U.S. census :

Representatives and direct taxes shall be apportioned among the several
states which may be included within this Union according to their respec¬
tive numbers, which shall be determined by adding to the whole number
of free persons, including those bound to service and excluding Indians
not taxed, three-fifths of all other persons. The actual enumeration shall
be made within three years after the first meeting of the Congress of the
United States and within every subsequent term of ten years in such
manner as they shall by law direct.

Although this was a landmark in census development, it limited
the work to an enumeration of the inhabitants of the country. Very
early there were demands for other information, such as data on
agriculture and industry. In the rapidly developing states and terri¬
tories the ten-year interval was sometimes longer than convenient
for state and local government, especially in frontier areas.

As demands came for more frequent or more detailed data on
population, manufacturing, industry or agriculture, the national
census organization was lacking in both experience and skills. For
nearly fifty years the question of the constitutionality of such addi¬
tional census work was a deterrent to progress. When people needed
more data on population or in new fields, they turned to the states
for them.

Actually State census work goes back into colonial times, census
enumerations being reported in Massachusetts as early as 1643;
Rhode Island, 1708; and New Hampshire, 1767.^ In 1854 the
Superintendent of the United States Census reported that 20 of the
31 states then in the Union had some kind of state census.^ Al¬
though the earliest work was concerned largely with population,
some later state enumerations included agriculture, manufacturing
and mining. These state censuses have now disappeared, except for
the mid-decade one in Massachusetts^ and a somewhat different one
which provides population data annually in Kansas.^

47

48 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Territorial Census Work by Wisconsin

Like other states, Wisconsin engaged in census-taking during
territorial days. Eleven state censuses were taken between 1836
and 1905. The first five came between 1836 and 1847. These terri¬
torial censuses were concerned only with population, first for the
organization of the territorial government and then for statehood.
A provision for state censuses at ten-year intervals was also writ¬
ten into the Wisconsin State Constitution in 1848.

The first territorial census of 1836 was described in a report of
the State Historical Society of Wisconsin in 1892.^ The editor
pointed out that the Act of Congress, April 20, 1836, establishing
the territorial government of Wisconsin provided that previous to

WISCONSIN TERRITORIAL CENSUS ■ 1836
SEX RATIO OF POPULATION

THREE COUNTIES- BROWN, IOWA,
MILWAUKEE

AND OVER

POPULATION
UNDER 21

In the census of 1836 the population was shown by age groups. The inhabitants
age 21 years and over were 77 per cent men and 23 per cent women for Brown,
Iowa, and Milwaukee Counties. Crawford County is not included in this chart
because the data were influenced by the military personnel stationed at Fort
Crawford.

1966] Ehling — Wisconsin Territorial and State Censuses 49

the first election the governor should order a census or enumeration
of inhabitants of the several counties to be made by the sheriffs
and reported to him.^ Upon the basis of this census the governor
was to apportion in the ratio of population the council members
and representatives, Indians excepted.

The first territorial census for Wisconsin was taken in July of
1836, No printed blanks were furnished for the enumeration.
Sheriffs were instructed simply to report in writing the names of
white families, with the number of persons in each family, divided
into four groups:

I. Males under 21 years

II. Females under 21 years

III. Males 21 years and over

IV, Females 21 years and over

In 1836 Wisconsin Territory was much larger than present Wis¬
consin because it included most of Iowa, Minnesota, and other land
west of the Mississippi River to about the present site of Bismark,
N. D. The enumeration, however, covered only the populated parts
of Wisconsin and some of the counties of Iowa west of the Missis¬
sippi River, an area temporarily attached to Wisconsin Territory

iN MID i9f^ CENTURY MOST STATES HAD THEIR OWN CENSUS •

SOURCE : 1850 US. CENSUS COMPENDIUM

In 1854 of the thirty states in the U.S., twenty-one had their own census. Wis¬
consin took censuses from 1836 to 1905.

50 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

pending further organization. Governor Dodge provided for repre¬
sentation in the Territorial Council (13 members) and House of
Representatives (26 members) on the basis of population, which
for the four counties — Brown, Crawford, Iowa, and Milwaukee —
amounted to 11,683 persons.

In 1838 the Territory took another census of population, again
by the county sheriffs, with the data recorded by towns and cities."^
No age divisions were required. The sheriffs recorded the names
of the master, mistress, steward, or overseer of each household, and
the township in which the family lived. They recorded the number
of white males, white females, free males of color, and free females
of color, with a column for totals and one for remarks. Each
sheriff was required to summarize the reports and submit them to
the Secretary of the Territory.

In 1842 another territorial census was authorized and taken. The
headings were the same as those in 1838, with the addition of a
column for errata.

As statehood approached, a further census enumeration was nec¬
essary. An Act in Relation to the Formation of the State Govern¬
ment, January 31, 1846, provided in sec. 1 that every white male
inhabitant above the age of 21 who resided in the territory six
months previous to the census and who was a citizen of the U. S,
or had filed his declaration of intention, according to U. S. natural¬
ization laws, was authorized to vote for or against the formation
of a state government in Wisconsin.® Sec. 3 provided for the gov¬
ernor to appoint in each of the counties some suitable person to
enumerate the number of inhabitants, omitting non-citizen Indians
and officers and soldiers of the U. S. Army. The census-takers were
empowered to choose as many assistants as necessary, assigning
to each one a portion of his county accurately defined either by
Congressional Township lines, the boundaries of towns organized
for town government, or distinctly bounded by water courses or
public roads.

The appointment of special persons to take the 1846 census was
a major departure from previous census work by the sheriffs or
their deputies in each county. The appointed persons were required
to take an oath that they would obtain an exact enumeration of all
residents within their county or division and make duplicate reports
for the Secretary of the Territory and the Register of Deeds. A
penalty was provided in sec. 6 of the Act for failure to perform
assignments properly. The enumeration was to begin on June 1,
1846, and be completed within 30 days.

Upon the basis of the population determined in this census the
governor was to issue a proclamation and appoint delegates appor¬
tioned to each county and territory according to population for the

1966] Ebling- — Wisconsin Territorial and State Censuses 51

first state constitutional convention. Thus this census differed from
previous ones, the details being carefully prescribed for the pur¬
pose of Statehood.

The first Constitution for Wisconsin, produced by a constitutional
convention in 1846, was rejected by the voters in a referendum in
April, 1847. To provide a basis in the territory for the apportion¬
ment of members for a second constitutional convention, a special
legislative session in October 1847, passed a new act for the forma¬
tion of a state government. Secs. 13-19 provided for another census
in December 1847, only 18 months after the previous one. This
census recorded 210,546 people, an almost unbelievable increase
of 36 per cent in a year and a half.

State Census Continued Under the
Wisconsin Constitution

The five censuses of population during territorial days in Wis¬
consin were largely for the apportionment of representatives to
the territorial legislature and the constitutional conventions, but
of course they also showed the rapid growth in population and the
advancement of the frontier.

After Wisconsin became a state, the Constitution provided for
the continuing of the state census enumerations at ten-year inter¬
vals for the mid-decade years ending in five.^ Six isuch censuses
were conducted for the mid-decade years from 1855 to 1905.

WISCONSIN POPULATION 1820-1960

52

Wisconsin Academy of Sciences, Arts and Letters [VoL 55

^ m

O Q

_J OT
3 3
Q. O
O X
a. H

o

1820

1966] Ebling- — Wisconsin Territorial and State Censuses 53

Obviously during a period of rapid settlement, population change,
and the frequent addition of new counties, apportionment of the
members of the legislature according to population needed to be
made frequently. The state Constitution provided that the mem¬
bership of the legislature be reapportioned after each census
enumeration, both federal and state, every five years. As the state
became more mature and population more stable, however, the
need for isuch frequent reapportionment was less pressing than
during the days of most rapid growth and geographic advance
of the population.

In November, 1910, a constitutional amendment relating to re¬
apportionment was adopted. It provided that apportionment of
members of the legislature according to population should be done
only at ten-year intervals in accordance with the U.S. census, thus
eliminating the need for a state census of population. The 1905
census, therefore, the sixth one under statehood, was the last of
the Wisconsin state censuses as provided under the Constitution.

Agricultural, Manufacturing and Mineral Data in the
Wisconsin State Census

The territorial and state census as in Wisconsin was developed
originally for the enumeration of the inhabitants. An explosive
increase in the work began with the 1885 census. The Revised
Statutes of 1878 had made substantial additions to state census
work. These included a long list of questions on agriculture (ani¬
mals, crop acreage, land tenure, equipment, product values), man¬
ufactured products and minerals produced in the state. The new
material was so extensive that much larger reports for the censuses
for 1885, 1895 and 1905 were required, with major portions de¬
voted to the new subjects.

The responsibility for carrying out this enlarged work was as¬
signed to the Secretary of State, who prepared the schedules and
sent them to county clerks for enumeration by town, city, and
village clerks. The county clerks filed the original reports with the
Registers of Deeds and sent copies to the Secretary of State, who
was responsible for tabulation and publication.

Filing original reports in the counties and making hand-written
copies for the Secretary of State had serious faults. There is no
evidence that counties had much use for the original documents,
many of which were lost, and the making of copies by cheap labor
in the counties resulted in errors and omissions which reduced the
accuracy of the tabulations.

Although the reasons that triggered the spectacular enlargement
of the Wisconsin state censuses beginning with 1885 are not en-

54 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

tirely clear, several are apparent. To begin with, the state economy
was largely agricultural and with the post-Civil War depression
of the 1870’s and 80’s, data on agricultural trends and changes
were of great interest. Another and perhaps major reason was that
the U.S. Congress in the census legislation for 1880 authorized
the Secretary of the Interior to pay states and territories half the
cost of a mid-decade census in 1885 if they met certain require¬
ments. Apparently Congress hoped that if all states could perform
a mid-decade census patterned after the U.S. census, mid-decade
data for the nation might be produced. As a result more elaborate
state censuses, including many of the U.S. census inquiries of 1880,
were taken in various states and territories, but the U.S. census of
1880 had been so enlarged that states could not duplicate it en¬
tirely. A few received federal payments but most of them, like
Wisconsin, did not. Although this federal legislation applied only
to 1885 and was not re-enacted,^® Wisconsin continued the enlarged
program through the 1905 census.

Another subject included in the state census of Wisconsin be¬
ginning in 1885 and continuing for the following two censuses —
1895 and 1905 — was the “enrollment of militia.” Wisconsin fur¬
nished 91,327 men in the Civil War. The 1885 state census recorded

NUMBER OF FARMS IN WISCONSIN , 1850-1960

U.S. Decennial Census Wisconsin State Census ^^U.S. Mid-decade Census

of Agriculture

Table 1. Some Characteristics of Wisconsin State Censuses.

1966] Ehling — Wisconsin Territorial and State Censuses

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56 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

the names and addresses of 29,686 veterans living in Wisconsin.
By 1895 the number of Civil War veterans in the state declined
to 26,367 and by 1905 to 9,521. Clearly by another decade few
Civil War veterans would remain and the need for this information
would disappear.

It is not surprising, therefore, that with the greater stability of
the state's population, thus reducing the need for frequent legisla¬
tive reapportionment, with agriculture more prosperous and be¬
coming a smaller segment of the state’s growing economy, and
with the number of Civil War veterans greatly reduced, the move
developed to amend the state constitution to eliminate the state
census. The amendment was passed by referendum in 1910, thus
closing seven decades of state census work.

These censuses, in addition to serving important purposes in
their time, provide a rich mine of historic information. Because
they were published by various divisions of the state, by counties,
towns, cities, and villages, they provide useful detail for historic
studies. Partly because a mid-decade census of population is useful
and because some present problems require more frequent censuses
than at ten-year intervals, administrators of new projects dealing
with apportionment of federal funds are now demanding a federal
mid-decade census. In recent years hearings have been held, with
many agencies urging Congress to provide a mid-decade census
of population. The latest of these hearings was in Washington on
May 4 and 5, 1965 ; a fifty million dollar appropriation was being
sought for this purpose.

Summary

State census taking developed in early American history when
population and frontiers were changing rapidly. The U.S. Census
at ten-year intervals was not frequent enough to provide informa¬
tion necessary under those conditions. State censuses, undertaking
to fill a part of the need, for a time served an important purpose.
However, they could not provide for the needs of an increasingly
complex society over the longer period and they have largely dis¬
appeared. Continuing needs for data at shorter intervals caused
the U.S. Census in 1925 to undertake a Census of Agriculture at
five-year intervals. A Census of Manufacturing is also taken at
shorter intervals. Because of other needs for population data at
five-year intervals one may expect that the U.S. Census will take
action within the next decade.

References

1, Ebling, Walter H. Evolution of Agricultural Data Systems, Agricultural
Estimates Division, U. S. Dept, of Agriculture and Dept, of Agricultural
Economics, University of Wisconsin, i960.

1966] Ebling — Wisconsin Territorial and State Censuses 57

2. U. S. Census Compendium 1850, by J. D. B. DeBow, Superintendent,

3. Statement by Raymond D. Lavalle, Census Director, State of Massachu¬

setts, May 4, 1962, before congressional subcommittee on census and gov¬
ernment statistics of the Committee on Post Office and Civil Service,
Hon. Harley O, Staggers, Chairman, pp. 653-660. Part 4, Mid-decade
census hearings.

4. Statement by Dr. Conrad Taeuber, Assistant Director, U, S, Census

Bureau.

5. Wisconsin Historical Collections, Vol. XIII, 1892, pp. 247-270.

6. Organic Act establishing the Territorial Government of Wisconsin of April

20, 1836, Sec. 4.

7. The legal basis for the 1838 Census is found on pages 239-244 Territorial

Laws of Wisconsin 1837, Act No. 53, providing for the taking of a sec¬
ond census or enumeration of the inhabitants of the Territory of Wis¬
consin, approved December 30, 1837.

8. Laws of Territory of Wisconsin, 1846, Act approved January 31, 1846.

9. Section 3 of Article 4 of the Wisconsin Constitution read as follows: ‘‘Sec¬

tion 3. The legislature shall provide by law for an enumeration of the
inhabitants of the state in the year 1855 and at the end of every ten
years thereafter: and at their first session after such an enumeration
and also after enumerations made by the authority of the United States
the legislature shall apportion and district anew the members of the
Senate and Assembly according to the number of inhabitants exclud¬
ing Indians not taxed and soldiers and officers of the U. S. Army and
Navy.

10. Wright and Hunt,, History and Growth of the U, S. Census 1790-1890,
U. S. Government Printing Office 1900, page 67 and footnote.

In addition to the above mentioned sources, the various laws and published
reports relating to this work in Wisconsin have been examined. Credit
must also be given to J. E. Boell, the state archivist, for encouraging a
study of which this paper is a part and to the staff of the State Histori¬
cal Society, especially Librarian Ruth Davis, who has been most helpful.
The Secretary of State’s office, especially Miss Kay Thompson, assisted
in making records available.

DETOURING CALAMITY IN WATER RESOURCE DEVELOPMENT
A CASE IN POINT: SOUTHEASTERN WISCONSIN

Spenser W. Havlick*

Water pollution control, inadequate water^based recreation facil¬
ities, and flood control loom as a trio of critical issues which the
American urban dweller must face with new urgency. The approach
in this discussion is first to present difficulties in water-resource
planning in general terms and second to analyze the southeastern
Wisconsin situation, using the Milwaukee River basin as an
example of a potential and relatively untapped water resource.
Implicit in the discussion is the assumption that the Milwaukee
River Valley could qualify as an experiment and model demonstra¬
tion of water planning and development in an urbanizing basin —
a matter of local as well as state and national concern.

The use of untreated surface and/or well water for metropolitan
centers began to be questioned in Milwaukee and across the nation
at the turn of the century, when surface water deteriorated. Sani¬
tary engineers had followed the European practice of combining
storm and sanitary wastes in a sewer network whose effluent was
discharged into the available water course. In arid regions where
stream flow was undependable and meagre, the population density
was at first both scattered and transient. Rotation of privies re¬
solved the waste problem until density became high. Technologies
in transportation opened new opportunities to expanding popula¬
tions and industries. In most areas, however, it appears that natu¬
rally available water resources for supply and waste assimilation
became inadequate. By the 1920’s, the need for better waste treat¬
ment and water purification was recognized in all of the large
American cities. The shortage was particularly critical in western
water supplies and in the eastern industrial centers with water
pollution.

Curiously, metropolitan cities on sizable lakes or rivers were
usually the last to be forced to take action in water development
schemes. By the 1950's and early 1960’s state and federal agencies
were authorized to take stronger measures in guiding water re-

* The author is afRliated with the Department of Conservation, School of Natural
Resources and the Department of Environmental Health, School of Public Health,
University of Michigan.

Manuscript received June, 1965.

59

60 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

sources, especially in flood control, pollution abatement, and in
water supply. The Water Quality Act of 1965 is a preview of more
comprehensive efforts by the Johnson administration to strengthen
the federal role in water quality management in 1966 and 1967.

With new and more leisure hours, increased income, interstate
highways, and the congestion of urban areas, water availability
has taken on values previously assumed to be costless (see Fig. 1).
Water-based recreation demand has increased sharply. Forty-four
percent of the United States’ population prefers water-based rec¬
reation activities over all others ( Outdoor Recreation for America,
1962). New premiums are attached to water resources because of
increased demand from waste disposal operations, water supply,
real estate developers, irrigation, power, and fish and game inter¬
ests. According to the Kerr Committee Report of 1960 the rate of
increase will be dependent upon the level of the population growth.
The improvement and application of technology to keep pace with
this increase, and the more intensive use of our land and water
will require more research and acceleration of programs for con¬
servation, development, and management of these resources (Water
Resources Activities in the United States, 1960).

Gilbert White, University of Chicago geographer, suggests that
there is a tremendous gap between what exists and what is tech¬
nically feasible. There seems little doubt that in every basin of
more than 2000 square miles drainage area and in many smaller
ones, there is the physical possibility of evening out flow by further
storage, of decreasing the pollution of waters, and of readjusting
upstream land use so as to reduce unnecessary soil loss and make
wise use of water. Of course the social feasibility of such water
and land management is a separate problem, according to White
(1957),

Without a crisis in a river basin, rational long-term planning
struggles along with the most modest budget. A severe drought or
flood catalyzes activity — often misdirected because of the urgency
of a recent catastrophe (Hart 1957). If protective legislation or
policy is not soon established, however, the justification for the
law fades with the memory of the crisis. With sustained public
interest and support, ultimate decisions about present and future
degrees of regulation and basin development are eventually cast
into the political arena — as they should be. Through the political
and institutional processes, objectives can be clarified and refined,
countervailing forces can be organized and operated, public infor¬
mation can be dispersed and the goals of the public can be brought
to fruition. As in so many other matters of rational land and water
planning for metropolitan growth, there is a general apathy on
the part of the uninformed and already overburdened taxpayer as

1966]

Ravlick — -Water Resource Development

61

well as a reluctance on the part of the politician to lead the way
for fear of controversy that might cost him a vote (Higbee 1960),

The engineer is able to anticipate the quality conditions of any
river passing through a basin. For example, assume that a stream
receives domestic or industrial wastes from a community. After a
period of recovery or '‘self-purification” under certain conditions,
the stream is restored to specific levels of quality in terms of dis¬
solved oxygen, bacteria, sludge deposition, biochemical oxygen de¬
mand (B.O.D.), or even temperature. As other communities load
the stream with effluents regardless of the level of recovery, the
conditions of the river at any point can be calculated with surpris¬
ing accuracy. Yet the growing pool of knowledge is still incomplete,
with several glaring examples such as the effect of algal photo¬
synthetic activity on oxygen levels in flowing water or the effect
of agricultural fertilizers accumulating by runoff in the river or
lake.

Conceivably a better understanding of the human ecology in a
river basin is an avenue which must be explored for better water-
resource planning in the future. Once the human interdependencies,
better deciphered, are superimposed upon the matrix of biophysi¬
cal interactions, the calamity of irrational planning may well be
avoided. The calamity, disaster, or sometimes merely the misfor¬
tune of the basin plan or design has been a national misallocation
of resources and perhaps more important — an undermining of self-
help by the region and population directly affected because of crisis
decisions which force federal jurisdictions upon the disaster area.
Hart (1957) emphasizes that a premium is placed on the unanimity
engendered by crisis, and a hindrance placed on mobilization of a
general interest of people in an interstate region in planning.

Numerous crises in basin development stem from an order of
events which should be reversed. Many times projections for popu¬
lation growth and economic development have been used as a fun¬
damental premise for water-resource development without the con¬
cept of a carrying capacity at present technology and prices. Would
it not seem advisable first to establish clearly the carrying capacity
of the basin in question and the array of alternatives available
under various costs and intensities of development? With these data
at hand, the political, economic, and administrative machinery at
local (basin) and national levels would function with the realization
that water resources are a major determining factor in economic
activity, population growth, and the stabilization of a basin.

The benefit-cost analysis of public water resources projects
assumes that prices in private markets generally register social
values. But relief measures are not initiated by most polluters un¬
less they are forced. Seldom can an inarticulate public (often unin-

62 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

formed) prevail against either the organized lobby of the heavily-
endowed polluter or the small unknown polluter.

Too many departures from the narrow scope of ideal market
conditions can occur for us to place great faith in benefit-cost anal¬
ysis when it is subjected to the pragmatic test. Often dumping
wastes into a river or a lake appears costless, on the assumption
that natural processes will do the job. When the self-purification
capacity is overloaded, however, the necessary job cannot be done.
Thus a fuller cost should be assessed, although perhaps no cost
whatever is calculated. The serious drawback is that distorted re¬
source allocations and social costs are frequently applied because
market or engineering plans create inefficient mixes of dilution,
water supply treatment and waste treatment. The polluter wonders
why he should incur a cost whose benefits are diffused downstream,
often unclaimed but available to all.

My contention is that when the data are more comprehensive,
a fee and bounty system could be arranged and subsidized through
public-private cost-sharing. Costs have been assessed and accepted
by users and nonusers of the national and state highways. The
‘Truckers” of our streams should certainly be charged, once the
data gap is closed. Once the computers can be fed the pertinent
information, the terms and costs can be assessed with considerable
certainty.

When computing the costs of alternative quality control devices,
we should consider competitive and complementary relationships
between water values and uses. For example, dilution through flow
augmentation would have to be evaluated in light of the fact that
this alternative is usually competitive to prime power production
and complementary perhaps to flood control, navigation, and irri¬
gation (depending on the season for the latter) .

Visualize a private basin with many manufacturing activities of
a single owner operating where the only allowable pollution would
be that for which he would be willing to bear the full costs of sew¬
age disposal and water treatment. Water treatment, to deal with
higher pollution levels from effluent and flow augmentation, theo¬
retically would substitute partially for sewage treatment.

Let us assume that the sales and purchases of goods and services
in a model basin-wide firm provide an adequate yardstick. With
this “market device”, pollution abatement can be measured in eco¬
nomic terms as the ratio of inputs/outputs. Public decisions about
pollution can be inferred from the actions of a firm which bears
the total costs. Two problems confront public policy when it recog¬
nizes an area (our basin) as an interdependent system that would
produce results varying in different beneficient ways from those
yielded by the operation of free markets in a basin with independ-

1966]

Havlick — Water Resource Development

63

ent pollution-producing and water-using units. One is the problem
of devising an optimum system for waste control and treatment
of water. The other is provision for an appropriate distribution of
costs among economic units and activities.

One shortcoming of the hypothetical firm is an inability to show
peoples' preferences in significant social values in the market of
goods and services. Although not a new problem, it is one which
demands a more thorough investigation. Another flaw is that the
^^economic efficiency" is much too narrow, coupled with a gross lack
of information on pollution interrelationships. If this sort of wel¬
fare maximization (or cost minimization) were to become national
policy, great care would have to be taken to prevent industries and
municipalities or districts from passing on excess costs. Public regu¬
lation could experience difficulty keeping in step with these cost
movements. No effort could be made until the necessary data is
available, political valuations filling the vacancy for the time being.
Overall, however, market criteria in a basin can establish guide-
posts and indicators of social value for the majority of goods and
services.

Some of the unique organizational and engineering constraints
which can be avoided only at very high cost are evident in the ina¬
bility to internalize many pollution-created externalities. Sometimes
a lake receiving 90% treated effluent over time shows an irrevers¬
ible eutrophication. Indeed, no one can fix a cost on lake-aging in
terms of littoral sludge buildup, a diminished hypolimnion (less
depth and more toxicity), or increased turbidity. It is equally
difficult to ^'charge" for high levels of electrolytes, alkyl-benzene-
sulphonates, and assorted inorganics which resist breakdown be¬
cause of infrequent detection and inadequate treatment. Inorganic
concentrations may increase as water use increases and even as
treatment continues. Political constraints have less rigorous con¬
ditions but add in with technical constraints to represent the extra
cost limitation put upon achieving an ultimate goal.

Attention is called to southeastern Wisconsin, which will illus¬
trate the merits and objections about the basin-wide firm. My case
in point is the Milwaukee River, which begins 80 miles northwest
of Milwaukee, Wisconsin, and meanders among morainal landscape
into a heavily industrialized urban area of more than one million
people. Dairy farms, heavy industry, tanneries, breweries and food
processing represent key economic activities in the 845-square-mile
drainage basin. As the stream approaches and parallels Lake Michi¬
gan, cities of progressively larger population pour untreated and
treated wastes into the Milwaukee River, The river enters Lake
Michigan at Milwaukee, spewing industrial and municipal wastes
into the lake, which is after considerable dilution the source of

64 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

drinking water for numerous cities, including Milwaukee, along the
Wiscbnsin-Illinois shore. Before 1900 and the subsequent indus¬
trial and population growths, the river was used for swimming,
boating, water supply, fishing, power, and navigation. Today pollu¬
tion curtails the first four uses; the last two have been halted for
other; reasons.

Three abbreviated models tailored from Kneese (1962) suggest
solutions for the pollution problem in southeastern Wisconsin. Since
the vast industrial and manufacturing complex at Milwaukee rep¬
resents the key economic growth and the greatest user of the basin,
benefits should be based mainly on industrial expansion and in¬
creased waterfront use envisioned after pollution abatement. Please
assurne that the present economic growth will continue, and that
factors of production (labor and capital) will be mobile. Also as-
sumejthat governmental agencies and political structures will main¬
tain present constraints in addition to the physical or technical
constraints of today’s pollution level, which has drawn the limit on
industrial growth and municipal use of the river. Let us now con¬
sider models X, Y, and Z.

Model X proposes specialization of the river. Certain tributaries
are zoned as clean-water streams, others and the main trunk as
legal carriers of waste. Regrettably, time and information avail¬
able do not permit a thorough presentation of pertinent data about
benefits to industry and to recreation, value added, least alternative
costs, and the benefit-cost ratio. For all alternatives, the data must
be complete and specific if courses of actions are to be qualified,
compared, and evaluated. The physical layout of Milwaukee sug¬
gests that specialization under Model X might be ideal : when the
river becomes loaded with pollution beyond the point of marginal
costs of treatment, industries and users turn to the clean streams
left in the basin, to ground water, or to Lake Michigan. But ground
waters are sinking out of economic sight and clean streams have
found competitive uses in recreation as well as in the complemen¬
tary use of diluting the main polluted stream. The lake is logically
next. Biologically and chemically, however, something is happening
to the quality of the lake water, making it progressively more ex¬
pensive to treat and use. With the relaxation in Model X of waste
treatment to substitute for greater water treatment, a problem has
been created far beyond the basin model.

Lake Michigan, despite its size, has begun to show significant
signs of eutrophication primarily from excessive siltation, agricul¬
tural runoff, and wastes from the eight million people along its
shores. Hard to calculate in the workings of the model is the water
quality level which Chicago would like to maintain for dilution

1966]

Havlick — Water Resource Development

65

purposes as it takes Lake Michigan water to augment flow in the
Chicago Sanitary Canal — Illinois River flowing to the Mississippi.

Model Y presents an alternative of flow augmentation which
strangely enough in one situation means recirculation. Again
necessary volumes of data, benefits, and costs are needed. Three
types of flow augmentation should be applicable. The first is a
system of storage reservoirs which would flood a highly developed
flood plain at very high social costs for translocation, etc.

A second possibility is flushing the river with Lake Michigan’s
water, an action which presupposes the need for river water qual¬
ity maintained by cold, oxygen-abundant lake water. However, the
intake for the flushing tunnels would include increasing amounts
of water polluted by the river, whose mouth is nearby. A combina¬
tion of the first two techniques could be a third type of augmenta¬
tion. Would this water quality permit the reopening of city beaches
now closed?

It might be of interest here to note that a proposal from the
mayor’s special water pollution committee suggested that the lake
pollution be abated by chlorinating the harbor basin (where the
river empties into Lake Michigan). Inadequate information is con¬
spicuous when investigators with or without cost analyses come
up with such a suggestion.

Model Z offers the possibility of widespread secondary treatment
(activated sludge, trickling filters, etc.) by users^ — individuals, in¬
dustries, and municipalities. Refined secondary treatment, stabili¬
zation ponds, tertiary treatment, separate storm and sanitary sys¬
tem, and better-than-nothing primary treatment could and should
be evaluated singly or as a composite activity for the basin plan. In
addition, as in the other methods and models, the downstream and
downlake effects (in an interdependent system) must somehow be
ascertained and the cost functions of alternatives including the pol¬
lution damages of models X and Y must be known. Only when
these are available can the over-all comparable costs of alternative
systems be explored.

These three models could serve as frameworks on which to hang
various data. A present normal is provided in the assumptions. A
budget analysis of opportunity costs, comparable values, and
benefit-cost ratios is implied before and after a particular model is
applied. A new allocation of resources should follow if suggested
by economic efficiency. In short, the goal is the most efficient com¬
bination of factors to minimize cost and maximize social welfare in
the basin. Sometimes, as on the Miami River, Ohio, a power plant
is forced out of an area because of demands of economic efficiency.
Loss of the tax base and the farther distance of power transmis¬
sion pass along a higher cost to the consumer. Yet the total ‘'eco-

66 Wisconsin Academy of Sciences^ Arts and Letters [Vol. 55

nomic ecosystem'’ must be taken into account before a final
judgment.

By drawing up comparable budgets of anticipated returns and
costs of certain alternatives, we can make significant strides to¬
ward water allocation in a water-dependent economy which may
make the fullest use of available resources. Alternative choices
need not, however, be judged totally on economic efficiency. Suc¬
cess is apparently forthcoming in Germany’s Ruhr Valley. The
Milwaukee situation illustrates, however, that a system has a con¬
straint in the dependence on larger watersheds and basins. Time,
distance, and natural processes make cost assessment difficult and
highly complex. As well as quantity, water quality must be treated
as a variable.

Pollution abatement facilities must be judged on more than just
technical-engineering estimates. When alternate economic terms
are combined with engineering solutions, the prospective benefi¬
ciaries are still confronted by institutional and administrative
problems. Even an economically and technically sound proposal can
be crushed in the institutional meshwork, whether it be in the val¬
ley of the Milwaukee or the Missouri or the Huron. The marginal
approach is not perfect because marginal data are often unavail¬
able and ideal market conditions do not exist in a river basin, even
if the economic service area is identical with the basin boundaries.
Nevertheless, these alternatives help to bring out the real problems,
which is a step toward finding answers for their solution.

Upon close scrutiny, even “non-consumptive” uses can, in fact,
be costly. Hirshleifer et aL (1960) lists the very significant values
of water that can be lost when the “non-consumptive use” is for
cooling, navigation (streamflow conflicts with values accruing to
pollution dilution and/or irrigation and/or hydropower peaking
pools), and water percolating underground, which is lost because
of extraction costs or minerals added. To avoid a calamity in water-
resource planning stemming from a crisis in the basin, economic
analysis alone will not suffice. In sectors of the western states,
water has become more scarce than the dollars needed to recapture
it. Folz (1957) warns that if additional water supplies are not
made available, water may become a limiting factor to economic
expansion — and such situations are increasing. Certain parts of
California are already facing an arrest of growth owing to water
shortage ; and the future growth of the populous industrial areas in
the East will largely depend on their ability to restrict those uses
of water the marginal utility of which is lower than that of urban
development uses.

It would appear unwise, therefore, in spite of temptations that
will be presented in the future, to base the expansion of the econ-

1966]

Havlick — Water Resource Development

67

omy on temporary increases in the supply of investment capital and
similar increases in the supply of water, since almost certainly the
future will bring renewed periods of drought. The wiser course
would seem to recognize the carrying capacity or minimum supplies
available in the long run — permitting adjustments from innovations
in technology — and development of the economy on those criteria.

The idea should not be conveyed that every basin crisis brings
a calamity because of irrational planning or even that every basin
is destined to experience a catastrophe. Commentators on the Dela-
ware River Basin insist that “there is time to plan and build to
provide for all the uses of as much water as engineering and eco¬
nomic techniques are capable of providing. There is no overpower¬
ing crisis (flood, famine, depression) today that is compelling
precipitate action toward ill-considered, unbalanced, and unwise
construction along the Delaware, Water development clearly is
not the key to economic growth in this humid eastern area, hence
other broad social and economic considerations will need to be
taken into account if the maximum economic potential of the basin
is to be realized. Serious shortages of good water may appear by
around 1980, for the supply is becoming progressively less generous.
Undoubtedly, development will be planned and construction begun
before any crisis appears” (Martin et al., 1960) .

Aside from some work being done by the Wisconsin Southeastern
Regional Planning Commission, other commendable efforts toward
basin-resource planning in the absence of critical crises are those
in the Huron River Basin in southeastern Michigan. After more
than seven years of citizen participation in organizational planning,
including a Huron River Watershed Intergovernmental Committee,
the 1964 Michigan legislature provided the basin and the state with
an enabling act (Act. No. 253, Approved May 28, 1964) which
authorizes units of local government to cooperate in planning and
carrying out a coordinated water management program in the
watershed which they share. If the planning process enables the
citizenry to foresee and prevent a crisis in the basin, the manage¬
ment and planning effort should serve the best public interests over
the long run as well as the short run. The recently formed Huron
River Watershed Council is a positive step.

For comparison of a less envious record of achievement in a river
basin development for the public good, attention is directed to an¬
other small watershed in southeastern Wisconsin, the Milwaukee
River Basin. During the early part of the twentieth century,
exemplary efforts were made by the Milwaukee County Park Com¬
mission in river parkway recreational development (See Fig. 2).
In recent decades, however, Milwaukee and its environs have grown
in a typical urban sprawl without concern for the river or the basin

68 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

FIG. 1 MILWAUKEE RIVER BASIN

1966] Havlick — Water Resource Development 69

Figure 2. View from Gordon Park across Milwaukee River one block down¬
stream from Locust Street bridge circa 1920. (Milwaukee Public Museum
photo.)

beyond the political boundary of the county (Fig*. 1). Perhaps the
most calamitous assumption in the minds of local planners was
that because the Milwaukee metropolis adjoined one of the largest
and deepest fresh-water lakes in the world, the city would never
face a water resource problem. Lake Michigan may not always be
the water planner's Elysium. Nevertheless, this visual and mental
association with “limitless” Lake Michigan has prevented the
intensification of public interest even in light of minor crises. Only
recently has some effort been made to call upon the Southeastern
Wisconsin Regional Planning Commission to suggest a plan for
the Milwaukee Valley, as was done in another smaller basin whose
representatives expressed concern. In summary, it is felt that cer¬
tain modest proposals should be offered in the form of hypothetical
recommendations which if implemented through proper and as yet
unestablished administrative and political channels might prevent
a calamity in the event of a crisis in basin resource allocation and
planning.

The five recommendations which follow offer the most feasible
possibilities for basin water development from the author's obser¬
vations of the predicted growth in the basin and his analysis of

70 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

the physical features of the basin landscape. It is suggested that the
proper authorities (still to be determined) propose a schedule of
priorities for these or other suggested projects in the hope that
one politically and economically practical may emerge. It is absurd
to contend that all the developments must begin at once, and it is
equally absurd that all must wait — especially in the light of present
demand for recreational facilities and even the most modest
population projections for the region.

(1) Fifteen existing reservoirs should be brought into greater
use. A number of new small reservoirs in the upper regions of the
Middle and North Branches of the Milwaukee River would provide
excellent pools for swimming, boating and fishing. Some should be
designed particularly as wildlife refuge. Long Lake, Kettle
Moraine Lake, Mauthe Lake, Lake Ellen, Wallace Lake, Silver
Lake, and Little Cedar Lake all typify the use and the congestion
in Milwaukee River Basin lakes. The topography southeast of Eden
in southern Fond du Lac County has abutments for artificial lake
impoundment. Much of the land along the North Branch of the
Milwaukee River is marginal pasture or sub-marginal cropland.
Most of the land suggested for the small catch basins on both
branches is presently marsh. With reforestation to improve the
terrain bordering the artificial lakes, attractive recreation areas
can be created now at low social costs. Proposed recreation sites
are about 45 minutes by auto from Milwaukee.

(2) Four and one-half miles north of the interchange of U.S.
Highway 141 and Wisconsin 100, the Milwaukee River loops within
about 5,400 feet of Lake Michigan. A recommendation is made that
a two-way channel or tube be constructed which would carry in
times of flood almost 10,000 c.f.s. from a small auxiliary reservoir
on the Milwaukee River to Lake Michigan. Perhaps a pump-siphon
arrangement is feasible (the river channel downstream has been
improved to carry 6,000 c.f.s. and the maximum flow on record is
15,100 c.f.s.) . In dry weather, pumps reversed from flood conditions
could lift Lake Michigan water to the ‘‘aqueduct” to augment flow
according to the needs of industry and waste assimilation. Lake
water would have to be lifted about 104 feet before gravity would
carry it down to the river channel ( See Fig. 1 ) .

(3) It appears that at least four stretches of “blue-green corri¬
dors” are or will be urgently needed to prevent flood plain buildup
as well as to provide critical recreation area along the Milwaukee
Waterway. The Milwaukee County Park Commission has success¬
fully used the parkway-river bank idea. Estabrook, Lincoln, and
Kletzch Parks are excellently-designed examples of what needs to
be done basin-wide.

1966]

Havlick — Water Resource Development

71

Taking into consideration present residential and commercial
development, the author urges the establishment of at least these
four blue-green corridors, which offer scenic river beauty and rec¬
reation besides a safeguard against future severe property loss
from flood inundation (all four corridors are shown shaded in Fig.
3).

(a) A four-mile river parkway just north of Kewaskum, Wis¬
consin, with easy access from U.S. Highway 45. It is assumed
that land would be obtained 0.5 mile in both directions from the
river bank, usually by easement, donation (as in most of Detroit’s
metropolitan parks) , or outright purchase.

(b) Another four-mile scenic corridor from Kewaskum south
along the river to almost the Barton-West Bend area. A small
corner of the Kettle Moraine State Forest reserves an additional
mile near the northern portion of this corridor (See Fig, 3).

(c) East of West Bend a scenic six-mile waterway meandering
toward Newburg. Easy entry to the sparsely settled river bank
would be possible from roads paralleling the corridor on the north
and south. Before the heavy picnicking and camping season, high
water canoeing should be popular in this and in the other corridors.

(d) Probably the first blue-green corridor which should be estab¬
lished, a five-mile stretch from the Waubeka-Fredonia vicinity
south to Saukville. Fortunately the Ozaukee County Parks of Wau-
bedonia and Ehler’s begin and end this proposed blue-green corri¬
dor. Again highways parallel the corridor conveniently on the east
and west. Parenthetically, the greatest expected flood plain resi¬
dential and commercial development will be from Waubeka-
Fredonia south along the river through Saukville, Grafton, and
Mequon to Milwaukee.

(4) Several Lake Michigan bathing beaches in Milwaukee are
closed 25-30% of the swimming season because of increased coli-
form counts after at least 0,1 of an inch of rain and excessive
aquatic plant growth from eutrophication caused by nutrients from
basin runoff and normal “efficient” sewage treatment. Even with
cool lake water most of the summer and deteriorating water qual¬
ity, excessive crowding on the beaches is a critical problem. In
providing recreation facilities for the 1970’s, the suggestion is
made that a new Milwaukee Metro-Basin Lake Michigan Beach
Park be created 36 miles north of Milwaukee, east of Lake Church.
Approximately 640 acres with 6,000 feet of sandy beach frontage,
in no danger of water pollution, appear to offer excellent potential
as a recreation area. The high bluffs of the Lake Michigan shore¬
line near Milwaukee are absent here. Land back from the lake in
the proposed park (held by private out-of-state owners) is wooded

72 Wisconsin Academy of Sciences^ Arts and Letters [Vol. 55

KETTLE MORAINE

Proposed
River Corridor

FIG. 3 MILWAUKEE RIVER BASIN WITH PRESENT AND PROPOSED RECREATIONAL AREAS

1966]

Havlick — Water Resource Development

73

and contains a small bedrock lake. See the triple circle along Lake
Michigan shore in Fig. 1 or Fig. 3.

(5) The final recommendation focuses on the revitalization of
mid-metropolitan boating. An extensive small craft marina is under
construction and a boat launch ramp is in operation in the outer
harbor area. With this development it would seem economically
rewarding to attract marine craft into the downtown area via the
lower river. Riverside docks, shops and promenades would be possi¬
ble if storm-sanitary sewer separation continues, along with in¬
creased waste treatment and reduced ground water infiltration into
the sanitary system.

Above North Avenue dam abandoned property already owned
by the Milwaukee County Park Commission and private firms
should be converted to river bank-parkland similar to upstream
Estabrook and Lincoln Parks. Kern, Riverside, Hubbard, and Gor¬
don Parks are fine beginnings along the two-mile double bank po¬
tential. Forty years ago, when pollution was probably an equally
or more severe problem (before separate sanitary sewers), great
activity occurred along the mid-city waterway (See Figures 4
and 5). If boating facilities were provided, following this recom¬
mendation, and riverside landscape improved, recreation of the
past could be restored, offering additional use and enhancement of
the present green corridor. Augmentations of stream flow sug¬
gested in (2) would be provided and balanced in a volume benefi¬
cial to industry, civic amenities, and river recreation but not to
jeopardize presently useful Lake Michigan beaches and water
intakes.

Conclusions

(1) Lake Michigan offers unlimited water supply for industrial
and population growth but at increasing cost of water treatment as
pumping increases and as intakes are extended. Part of the added
cost is caused by deteriorating lake water quality. No use of Mil¬
waukee River water is for potable supplies.

(2) Lake Michigan offers limited recreational facilities in the
Milwaukee vicinity. As an alternative, several possibilities are
spelled out in the form of upriver recreation corridors and small
artificial lakes zoned for specific uses. These reservoirs should not
be justified primarily by flood-control benefits as has been at¬
tempted previously. The dual purpose diversion and flow augmenta¬
tion facility east of Thiensville would provide both flood and low
flow protection. Minimum monthly average flows could most prob¬
ably be maintained in excess of 300 to 500 c.f.s., using the new
facility in combination with present reservoirs.

74 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Figure 4. Excursion boat on Milwaukee River at Wisconsin Avenue bridge.
Milwaukee City Hall in background. Marine National Exchange Bank has re¬
placed buildings on the right. Circa 1920. (Milwaukee Public Museum photo.)

(3) Instead of major waste lagooning, flow augmentation, or
both to furnish drastic pollution abatement, the proposal is made
to provide increased and alternative recreation areas within easy
driving distance of a growing metropolis, and hold the line on river
and lake water quality deterioration through treatment, separation
of sanitary and storm sewers, and elimination of ground water in¬
filtration into the Metropolitan Sewerage Commission facilities.

(4) A basic assumption is that a dilatory program in recrea¬
tional water development and enhancement of the basin environ¬
ment can act as the greatest constraint on growth in the basin,
which has ample water, transportation facilities, and other factors
to expedite growth and social welfare.

We can be certain that the demand and need for recreation facili¬
ties along the Milwaukee River Waterway will continue. It is hoped
that these wants can be met through appropriate water-resource
development and management. Crisis should be avoided in plan¬
ning to satisfy the multiplicity of wants of any basin. The water
potentials which move states, cities, and the nation to act hastily
are fleeting and capricious and incapable of being harnessed eco¬
nomically, save by measures which assume long-term human enter¬
prise. One alternative before us, therefore, is to continue to try to

1966]

Havlick — Water Resource Development

Figure 5. Mid-city Milwaukee River recreation activity at Riverside and Gor¬
don Parks about 1918. Revived river use is proposed in conjunction with vig¬
orous water pollution abatement. (Milwaukee Public Museum photo.)

rationalize our national government and to make uniform devolu¬
tions to the states — although consent born of crisis will continue
to thwart and misdirect those efforts (Hart 1957). Organizational
interrelationships of a basin are a fascinating phenomenon in hu¬
man ecology. When considered together with the basin carrying
capacity and with the physical and economic features which deter¬
mine substantive policies and practices, they should help maxi¬
mize present opportunities and permit prudent, rational future
alternatives in basin planning.

References Cited

Folz, William E. “The Economic Dynamics of River Basin Development,”
Law and Contemporary Problems, Vol. XXII, No. 2, Duke University
School of Law, 1957, p. 207.

Hart, Henry. “Crisis, Community, and Consent in Water Politics,” Law and
Contemporary Problems, Vol. 22, No. 3, Duke University School of Law,
1957, p. 537.

Higbee, Edward. The Squeeze — Cities Without Space, William Morrow & Co.,
New York, 1960, p. 270.

Hirshleifer, Jack, James C. DeHaven, and Jerome W. Milliman. Water
Supply: Economics, Technology and Policy, University of Chicago Press,
1960, p. 68.

Kneese, Allen V. Water Pollution: Economic Aspects and Research Needs,
Resources for the Future, Inc., Washington, D. C., 1962.

76 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Martin, Roscoe C., Guthrie S. Birkhead, Jesse Burkhead, and Frank J.
Hunger. River Basin Administration and the Delaware, Syracuse Uni¬
versity Press, 1960, p. 228.

Outdoor Recreation for America. A Report to the President and to the Con¬
gress by the Outdoor Recreation Resources Review Commission, Washing¬
ton, D. C., 1962, pp. 173-182.

Water Resources Activities in the United States, Senate Select Committee on
National Water Resources, Committee print No. 12, S. Res. 48, 86th Con¬
gress, 2nd Session, U. S. Government Printing Office, 1960, p. 3.

White, Gilbert F. “A Perspective of River Basin Development,” Laiv and
Contemporary Problems, Vol, XXII, No. 2, Duke University School of
Law, 1957, p. 159.

THE TRENTON METEORITES

W. F. Read and H. O. Stockwell

Although meteorites are commonly named for the town large
enough to have a post office nearest to their discovery location, the
Trenton meteorites are an exception. Trenton is not a town but a
36-square-mile township in Washington County, Wisconsin. The
center of the township is about four miles east of West Bend, or
roughly 30 miles north of Milwaukee,

First published notice concerning the discovery of iron meteor¬
ites in this township was a short article by J. Lawrence Smith^ in
the American Journal of Science for 1869, Smith reported that
four specimens had been found, weighing 62, 16, 10, and 8 lbs,, and
that all had been acquired by the German Natural History Society
of Wisconsin. F, Brenndecke reported to the Natural History So¬
ciety in 18692 that the 62 lb. mass was found in 1858 and pur¬
chased by I. A. Lapham. The three smaller specimens turned up
“in the years immediately following’’ and went into the Society’s
collection. A fifth piece was said to have been found but could not
be located. The 62, 16, 10, and 8 lb. specimens will be referred to as
Nos. 1, 2, 3, and 4.

In 1872, Lapham reported® the finding of two additional speci¬
mens: one of 16l^ lbs, in 1869 and another of 33 lbs. in 1871. He
purchased the first for his own collection. The second was sent to
M. Von Baumbach “to be taken to Europe.” The 16% and 33 lb.
specimens will be referred to as Nos. 5 and 6.

Mr. Carl Gauger has advised the authors that about 1880 a speci¬
men weighing approximately 10 lbs. was found on his property and
taken to the Milwaukee Public Museum. This specimen will be re¬
ferred to as No. 7,

H. 0. Stockwell of Hutchinson, Kansas, visited the area in Sep¬
tember, 1952, and went over considerable ground with a metal de¬
tector. Results were spectacular. On the second day he found one
mass of 413 lbs. a few feet away from another of 527 lbs. Later
he found a small specimen weighing 11/2 lbs., and purchased two
more specimens from local residents. One was a 6V2 lb. mass report¬
edly found before 1890. The other, weighing 3 lbs., was said to
have been found about 1933. The 61/2 lb. mass will be referred to
as No. 8; the 3 lb. mass as No. 9; and the 413, 527, and II/2 lb.
masses as Nos. 10, 11, and 12.

77

78 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Some notes concerning- the disposition of Stockwell’s five speci¬
mens are in order. About 80 lbs. were removed from the 527 lb.
mass and sold to Ward’s Natural Science Establishment. The re¬
mainder of this and the entire 413 lb. mass have been purchased
by the U. S. National Museum. The 6I/2 lt>. mass and half of the
11/2 lb, mass were sold to R. A. E. Morley of Salem, Oregon.

In August 1964, W. F. Read and his son discovered another
specimen of 9I/2 lbs. while working with a metal detector similar
to the one used by StockwelL^ This will be referred to as No. 13.

A summary of the finds to date is as follows :

Found Weight

1

1858

62

2

1858-68

16

3

10

4

8

5

1869

I6I/4

6

1871

33

7 c.

1880

10?

8 c.

1885

61/2

9

1933

3

10

1952

413

11

yy

527

12

yy

11/2

13

1964

91/2

Location of Finds

The only finds whose locations have been recorded with any pre¬
cision are those made by Stockwell and Read. Smith reported that
the first four specimens were found “within a space of ten or
twelve yards very near the north line of the 40 acre lot of Louis
Korb”. Lapham’s manuscript notes^ include a map which shows
that the Korb property was the SWi/4 of the NEl^, Sect. 33, T 11
N, R 20 E, and that the meteorites were found near the center of
the north line. Lapham’s 1872 report on the finding of Nos. 5 and
6 states only that they were found “in the same field”. His manu¬
script notes, however, say that No. 5, at least, came from “very
near” the place where Nos. 1-4 were found. The approximate dis¬
covery sites of Nos. 7 and 9 were pointed out to W. F. Read by
Mr. Carl Gauger, who now owns the property. According to infor¬
mation obtained locally by H. 0. Stockwell, No. 8 was discovered
on an old stone pile formerly about 500 ft. northwest of the Gaedke
barn.

Figure 1 shows with varying degrees of accuracy the discovery
sites of all specimens except No. 8. Coordinates of the main site

1966] Read and Stockwell—The Trenton Meteorites

79

Figure 1. Northeast quarter of Sect. 33, T 11 N, R 20 E. Meteorite discovery
sites: cross indicates precise location; circle, fairly precise location; dot,
approximate location. Dashed line shows limits of detector coverage by W. F.
Read; dotted line, approximate limits of detector coverage by H. O. StockwelL

(Nos. 10--13) are Lat, 43" 22^ 44"; Long. 88° 6" 30", (Smith
gives the latitude as 43° 22^ and the longitude as 88° 8^) The
nearest town is West Bend, about 4 miles to the northwest, for
which according to modern usage the meteorites should have been
named.

No, 5. External Form

The Greene collection at Milwaukee-Downer College included a
161/4 lb, uncut iron meteorite identified in the catalog as from
Washington County, Wisconsin, Presumably this is specimen No.

80 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

5, found in 1869 and acquired originally by Lapham. When and
how it came into the Greene collection is unknown. When
Milwaukee-Downer merged with Lawrence College in 1964, the
bulk of the Greene collection was purchased by the University of
Wisconsin-Milwaukee. This specimen was loaned to W. F. Read for
study. Its external form is shown in Fig. 2. The original shape
has doubtless been somewhat modified by oxidation. The bottom
side in the upper photograph (same as upper two thirds of lower
photograph) shows low knobs separated by shallow depressions
and may be an ablation surface from the exterior of the parent
mass. The other three surfaces are evidently the result of rupture,
with no apparent subsequent modification by ablation. The one to
the left of the label in the upper photograph is jagged and suggests
rupture by pulling apart. The bottom surface in the lower photo¬
graph is smoothly curved, as if by shearing. The (poorly shown)
top surface in the upper photograph is about two thirds smooth
and one third jagged, suggesting a combination of shearing and
pulling apart. Whether rupture took place on or before impact (or
both) is not clear.

No. 5. Structure and Composition

Fig. 3 shows the appearance of an etched section. Kamacite
bands are about .7 mm. wide, making this a medium octahedrite,
as noted in the Prior-Hey Catalogue.^ Since the Widmanstatten
pattern is continuous across the entire section, this is evidently a
fragment from a single large Ni-Fe crystal.

An interesting feature of the kamacite bands is their tendency
to show a certain amount of curvature. This can be seen by using
a straight edge on Fig. 3. Presumably the bending is from stress
encountered either (1) during the meteorite’s pre-terrestrial his¬
tory, (2) while passing through the earth’s atmosphere, or (3)
on impact. These alternatives are certainly not mutually exclusive.
Along the upper right edge of the section as shown in Fig. 3, the
Widmanstatten figure disappears in a jumble of irregular kamacite
grains. These are transected by a small “fault”, clearly traceable
for a distance of about 6 mm. The fault is quite tight, certainly
not an open fracture, and suggests shearing under high pressure,
presumably pre-terrestrial. The reason for the granular structure
and its genetic relation, if any, to the fault, remains a question.
The oxide-filled fracture visible along the lower edge of the section
in Fig. 3 clearly differs in origin from the fault. It appears to be
the result of incipient rupture under low confining pressure.
Another indication of stress (Uhlig’s interpretation”^) is seen in the
occurrence of Neumann lines in many of the kamacite bands.

1966] Read and Stockwell — The Trenton Meteorites

81

Figure 2. Two views of Trenton No. 5. The side shown in the lower photo¬
graph is at the bottom in the upper photograph. Short lines indicate the posi¬
tion of the sawcut for the etched surface shown in Fig. 3.

Perry® has called attention to the prevalence of ‘‘hatching” (re¬
garded by him as a gamma-alpha transformation structure) in
the kamacite of Trenton specimens at the U.S. National Museum.
This is conspicuous also in the kamacite of Trenton No. 5.

82 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Figure 3. Etched face of end piece cut from Trenton No. 5. The black “vein”
at the bottom is oxidized material following a fracture.

Plessite fields are numerous and of variable structure. Some —
usually the smaller ones — consist of “dense’', apparently homogene¬
ous material etching dark grey. Some contain abundant small gran¬
ules of kamacite in a dark grey matrix. And some show fine kama-
cite bands instead of the granules, the bands running in one or
more directions conforming to the surrounding Widmanstatten
pattern. When bands and granules occur in the same field, the
bands tend to be disposed around the borders with granules toward
the center.

Troilite occurs in Trenton No. 5 as nodules, thin plates, and small,
irregular grains (See Fig. 4). The nodules (Fig. 4 shows two) lack
a continuous envelope of swathing kamacite, but are surrounded by
irregular kamacite grains that stand out clearly from the adjacent
Widmanstatten pattern. It is well known that troilite undergoes
a considerable volume increase by inversion at 130° C. This may
explain the fact that some of the oxide-blackened fractures visible
in Fig. 3 seem to be roughly radial to the troilite nodules. Note
especially how the large fracture along the bottom edge turns
upward at its right-hand extremity and terminates against the
nodule in this vicinity. The thin plates of troilite may be straight or
distinctly curved. They grade into more or less lenticular bodies.
Some of the plates and small grains may have failed to show up

1966]

Read and Stockwell — The Trenton Meteorites

83

Figure 4. Distribution of troilite in the etched face shown in Fig. 3. The
smaller particles were located by means of a sulfur print.

on the sulfur print from which Fig. 4 was taken. For example, the
straight, black line extending toward the upper left from the left-
hand nodule in Fig. 3 appears to be a completely oxidized thin plate
of troilite.

Trenton No. 13

As noted above, Trenton No, 13 was discovered by W. F. Read
and his son in August 1964. It lay at a depth of about 1^ ft., where
the oxide crust was undisturbed by cultivation. The surface which
appears at the top of the upper photograph in Fig. 5 is smoothly
convex and was probably shaped by ablation. The opposite surface,
shown in the lower photograph, is extremely irregular. It is heavily
encrusted with limonite, locally forming short, finger-like protuber¬
ances, The surface of the metal underneath is apparently quite
jagged, probably indicating a rupture surface formed by pulling
apart. Trenton No. 13, which has not yet been sectioned, remains
for the present at Lawrence University.

Acknowledgment

The West Bend News was most helpful in paving the way for
StockwelFs collecting work. Reuben Gauger, who then occupied the
Gaedke farm, and Carl Gauger kindly permitted Stockwell to work
parts of their farms with his metal locator. Subsequently Robert
Gaedke and Carl Gauger extended similar hospitality to Read. For

84 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Figurf] 5. Two views of Trenton No. 13. The side shown in the lower photo-
g]'aph is the bottom of the specimen as shown in the upper photograph.

the loan of Trenton No. 5, described in this paper, Read is indebted
to Prof, R. A. Pauli of the Geology Department at the University
of Wisconsin-Milwaukee. Mr. R. A. E. Morley of Salem, Oregon,
furnished valuable information on the history of Trenton finds.
For data derived from Lapham’s manuscript notes, the writers
are indebted to Mr. Walter E. Scott of Madison.

1966] Read and Stockwell — The Trenton Meteorites

85

References

1, Smith, J. L. A nev/ meteoric iron — “the Wisconsin meteorites” — with some

remarks on the Widmanstatten figures, Amer, Jour. Sci., 2nd ser., Vol.
47, p. 271-272, 1869.

2, Brenndecke, F. On meteorites: extract from a discourse, February 7, 1869,

before the Society of Natural History of Wisconsin. An, Rept. Smiths.
Inst 1869, p. 417-419.

3, Lapham, I. A. The Wisconsin meteorite. Amer. Jour. Sci,, 3rd ser., Vol. 3,

p. 69, 1872.

3, Read, W. F. The Hedden-Stockwell meteorite detector. Meteoritics, Vol. 2,
No. 4, p, 369-370, 1965.

5, Lapham, I. A. Manuscript notes in possession of the Wisconsin State His¬

torical Society.

6, Prior, G. T. Catalogue of meteorites, 2nd ed. revised by M, H. Hey. London,

The British Museum, 1953.

7. Uhlig, H. H. Contribution of metallurgy to the study of meteorites, Part

II— -the significance of Neumann bands in meteorites. Geochim. et Cos-
mochim. Acta, Vol. 7, p. 34-42, 1955,

8. Perry, S. H. The metallography of meteoric iron. U. S. Nat. Mus. Bull. 184,

1955.

FISHES OF SOUTHWESTERN WISCONSIN

George C. Becker
Department of Biology
Wisconsin State University, Stevens Point

The last extensive sampling of the fish fauna of southwestern
Wisconsin was made by C. Willard Greene during the late 1920’s.
In 1935 he published The Distribution of Wisconsin Fishes. Al¬
though many studies on game fish have been made in southwestern
Wisconsin since Greene’s time, no inventory of fish species has ap¬
peared since. The present study includes inland and boundary wa¬
ters of the counties of Richland, Crawford, Grant, Iowa, and La¬
fayette. The stations sampled appear on Map 1.

From June 27 to June 30, 1960, eleven stations (L6-16) were
sampled on the Pecatonica River and its tributaries in the eastern
half of Lafayette County and one station (112) in Iowa County.
From June 20 to August 18, 1962, 135 stations were sampled along
the lower Wisconsin River, the inland waters of the counties of
Richland, Crawford, Grant, Iowa, and Lafayette counties, and one
station on the Mississippi River (M4). From July 15 to 18, 1963,
four samples (M2, M3, M5, M6) were taken from Pools 10 and 11
of the Mississippi River. On June 27, 1964, a sample (Ml) was
taken from Pool 9,

One hundred species and more than 90,000 individuals were seen
or handled. Readily identified species were returned to the water.
Those whose identification was questionable were preserved in 5%
formalin, sorted, and identified later. Examples of all species have
been transferred to 40% isopropanol and are stored at the Biology
Museum, Wisconsin State University, Stevens Point.

In medium-sized streams to large rivers the most useful collect¬
ing device was a nylon seine 25 feet long, 6 feet deep, with 3/16-
inch bar measurement. It was used almost exclusively on the Wis¬
consin and Mississippi Rivers along sand and mud bottoms in water
4 feet deep or less. Because over shallow rocky bottom the seine
was less effective in capturing darters, at stations W12 and W15
(see Map 1) we used an alternating current shocker with 100 feet
of cable. The darters were extracted from between the rocks with
scape nets. A boom shocker, powered by a direct current generator,
was used to capture the deep-water fish from the Wisconsin River.
The direct current drew fish momentarily to the electrodes, from
which they were removed with scape nets. We found the boom
shocker ineffective against fish less than 4 inches long.

87

88 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55 t

One to two hours were devoted to collecting* fishes at each collec¬
tion site. An attempt was made to sample from all possible habitats.
On the Wisconsin River, for instance, we collected from sloughs,
riffs over sand flats, island pools, isolated overflow pools, swift
water, slow-moving water, and seepage bayous. A 12-foot boat
powered by a 5-horsepower outboard motor carried us to a wide
variety of habitats at each station.

Geology of the Region

The counties of Richland, Crawford, Grant, Iowa, and Lafayette
(except for a small area in the southeastern corner) fall within
the unglaciated portion of Wisconsin. This region, spared the level-

1966]

Becker— Fishes of Southwestern Wisconsin

89

ing effect of glaciation, is uniquely beautiful, with craggy bluffs,
pillars, and natural bridges carved out by wind, rain, and other
forces. The high relief of the terrain has deep valleys or coulees
alternating with high knolls. Loose rocks, irregular and sharp-
edged, are of the same material as the bedrock of the region. Caves
and sink holes are common and frequently quite large.

Streams, largely devoid of falls or rapids, follow regular courses
and show a dendritic pattern. Marshes and lakes are scarce and
are found only in the valley bottoms. The region is well known for
its flash floods, particularly the Kickapoo River,

Most of the soil of the region is derived from the underlying bed¬
rock and is referred to as residual soil (Martin 1916). The residual
material in the limestone belts is chiefly a fine brown or reddish
clay, representing the more or less insoluble residue from the decay
of the limestone. Much of this fine soil is carried down the steep
slopes and into streams, frequently raising turbidity.

On the higher and more level areas of this section there is a
layer of light or buff-colored silt soil called loess, which was
brought there by the wind (Whitson 1927), Part of it came from
the far western plains, although some of it was probably derived
from rock flour exposed around the borders of the glacial area
where streams flowed out from under the ice. The loess forms a
blanket varying from a few inches to several feet thick.

Although most of this portion of the state was originally wooded
and the soils are of a comparatively light color, some portions, es¬
pecially belts along the tops of the ridges, were prairies and had
darker soil. The soils of this character were formed largely from
the loess.

Recent Changes in Distribution

Southwestern Wisconsin is a strategic crossroads in fish distribu¬
tion. With the Mississippi River as a distribution route, this part
of the state is frequently the first to show the movement north¬
ward of southern species and the movement eastward of the west¬
ern plains fishes.

Some species which have come into Wisconsin in the recent past
are the Ozark minnow, the pirateperch and the warmouth. Thus
far these species appear to be common nowhere, but there is evi¬
dence that they are spreading gradually into state waters from
which they had not previously been taken, Greene (1935) captured
the Ozark minnow in Iowa and Lafayette counties. Our survey has
disclosed several colonies in the Platte River and its tributaries in
Grant County.

Our four collections of the pirateperch from three stations on
the lower Wisconsin and one station on Bear Creek, one of its

90 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

tributaries, indicate a firm establishment of this species in the in¬
land waters of the state. Greene took this species at only five sta¬
tions in Wisconsin. One was Mill Creek, 6 miles southwest of Ste¬
vens Point, indicating” that at an early date it had migrated up the
Wisconsin River to the middle of the state.

The warmouth had been reported by Greene from southwestern
Wisconsin only from the Mississippi River. My collection from Mill
Creek, a tributary of the lower Wisconsin in Richland County, in¬
dicates the presence of this species in the Wisconsin River drain¬
age. Wisconsin Conservation Department personnel report the
warmouth as common in the region. Since it is a desirable pan¬
fish, a considerable amount of minnow pail stocking may have
taken place. Hence the problem of evaluating natural distribution
of this species is the same as for yellow bass, which in recent years
has been captured in many new waters (Helm 1958).

In comparing our findings with Greene’s we observe that the
following species at least have increased in numbers and have
extended their ranges during recent years in southwestern Wis¬
consin: silver redhorse, golden redhorse, longnose dace, brassy
minnow, Ozark minnow, grass pickerel, western sand darter,
orange-spotted sunfish, pumpkinseed, and rockbass. The intrusion
of the rockbass, considered a glacial lake species, into the driftless
area is an example of adaptation.

Although the slimy sculpin was expected in southwestern Wis¬
consin, this glacial relict was not until recently found in Citron
Creek, Crawford County. The starheaded topminnow, previously
collected infrequently from southeastern Wisconsin, has appeared
in a recent collection from an Iowa County lagoon of the Wiscon¬
sin River.

The skipjack and blue catfish seem to have become exceedingly
rare or may even be absent. Greene examined collections of these
species from the upper Mississippi River, but we have no recent
reports from there. A 1963 survey showed both species far down¬
stream in the vicinity of the Kentucky-Tennessee line (pers. comm.
— Nord, Jan. 10, 1964.) The ghost shiner, formerly common in the
Wisconsin and Minnesota portions of the Mississippi River, has
not been collected there since 1944.

Species with drastic reduction in numbers are the paddlefish
and the channel catfish, probably because of the rapidly deteriorat¬
ing conditions on the Mississippi. Many commercial fishermen re¬
port the catfish industry in jeopardy. On the same river the chan¬
nel mimic shiner appears to be decreasing and is rarely found
today.

Brook trout, even in the smaller colder streams, find little suit¬
able habitat. If the brown trout were not stocked on a put-and-

1966] Becker — Fishes of Southwestern Wisconsin 91

take basis, trout fishing- in southwestern Wisconsin would be an
activity of the past.

What is the opportunity of adding new species to our Wisconsin
hsh fauna? Over 30 species of fish known from Illinois have not
been found in Wisconsin (Forbes and Richardson 1920). Some of
these may find conditions suitable here. The recent shift northward
of some of our Wisconsin species indicates a trend that may apply
to Illinois species.

I anticipate that the red shiner, Notropis lutrensis (Baird &
Girard), will soon be listed from Wisconsin. It has been reported
from a small stream in Dubuque County, Iowa, opposite Grant
County, Wisconsin (Harlan and Speaker 1956), and again from
the Mississippi River a few miles below the Wisconsin line (pers.
comm. — Nord Jan. 10, 1964).

Species of Fishes

Southwestern Wisconsin, because of the Mississippi and Wis¬
consin Rivers, is rich in species number. Our collections contained
82 species from the Wisconsin River; 60 species from the Mis-
sippi River; 64 from the inland waters of Richland County; 53
from Iowa County; 49 from Grant County; 44 from Crawford
County; and 36 from Lafayette County. A report from Nord (Sep¬
tember 24, 1962) lists 74 species of fish collected from or seen in
Pool 10 of the Mississippi River during Upper Mississippi River
Conservation Committee surveys. Pool 10 extends from Gutenberg
(Grant County) lock and dam to Lynxville (Crawford County)
lock and dam (U.S. Army Engineer Division, 1963).

The present survey captured 29 species of minnows (Cyprini-
dae) ; 15 species of suckers (Catostomidae) ; 15 species of perch¬
like fishes and darters (Percidae) and 10 species of sunfishes and
allies (Centrarchidae) . These four families of fishes were the best
represented in these waters, and in numbers of individuals cap¬
tured. In addition to my collections and observations I have in¬
cluded in the following list reliable reports from the literature and
from informants.

Map 1 indicates those waters in which collections were made.
The sampling stations are designated by a letter followed by a
number. The letters have the following values :

W — Wisconsin River
M — Mississippi River
R — Richland County
C^ — Crawford County
G — Grant County
I — Iowa County
L — Lafayette County

92 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

The Mississippi River, Wisconsin River and the counties have
separate numbering systems, each beginning with the number 1.
Station numbers are assigned from west to east in the county;
i.e., in Crawford County the westernmost stream sampled was
Gran Grae Creek, which received a station designation of Cl, fol¬
lowed by the Little Kickapoo with C2. On a stream with a system
of tributaries, such as the Kickapoo River, numbers were as¬
signed as follows: first the stations on the tributaries of the west
side of the Kickapoo; then the Kickapoo itself with upstream sta¬
tions being followed by downstream stations; and then the tribu¬
taries on the east side of the Kickapoo River.

After the species names which follow, I have indicated by key
letters and station numbers all of the stations where those species
were captured. If one species appeared at several consecutive sta¬
tions, the first and last station number are separated by a hyphen ;
e.g., M2-5 (after the longnose gar) means that it was captured at
stations 2, 3, 4, and 5 on the Mississippi River. Numbers within
parentheses, such as W (2-8), refer to a capture or captures some¬
where between station 2 and station 8 on the Wisconsin River. This
was a boom shocking float trip in which no attempt was made to
pinpoint the site of capture of a particular species. Other boom
shocking collections are represented by the following designations :
W(12-13), W(18-19), W(20-22).

Silver lamprey — Ichthyomyzon unicuspis Hubbs and Trautman.
W16. A single adult was collected from the Wisconsin River at
Boscobel by Mr. Larry Bolchen on June 28, 1962. It has been pre¬
served and placed in the State University collections. Several rec¬
ords from the Iowa side of the Mississippi River are reported by
Harlan and Speaker (1956). This species of lamprey as well as
those that follow are probably more abundant than the collections
records seem to indicate. Electric currents drive them out of the
mud and sand. The ammocetes seldom are taken with the seine
because of their burrowing habits.

Chestnut iRmprey— -Ichthyomyzon castaneus Girard. Nord (pers.
comm. — Sept. 24, 1962) lists this species as uncommon on the
Mississippi. Greene (1935) believes that this species may be com¬
mon in the larger rivers of the Mississippi system. Harlan and
Speaker (1956) report it from the Mississippi River opposite
Allamakee County, Iowa.

American brook lamprey — Lampreta lamottei (LeSueur). R4,
Rll, R14-16, R18, R20, R23, G12, G34, G36, 12, 15 (13 collections;
157 + individuals). This species seems to be distributed commonly
in the smaller, dear-water streams in the region.

1966]

Becker- — Fishes of Southwestern Wisconsin

93

Bsiddlefish—Polyodon spathula (Walbaum). The paddle fish is
reported from the Wisconsin River upstream to the Prairie du Sac
dam. John Truog, Wisconsin Conservation Department, reports
that in recent years schools of paddlefish have been observed in
spring below the dam. The Wisconsin Conservation Bulletin of
March 1937, carries the following item: “Prairie du Sac— A 80
pound ispoonbill catfish was imprisoned by the swift current at the
power dam here. The fish was four feet long.’’ Troug examined a
57-inch paddle fish found dead in the Wisconsin River at the mouth
of Blue River (Grant County) on Oct. 28, 1962. Robert Searles,
Biology Department, Wisconsin State University, Stevens Point,
found a partially decomposed paddlefish under the bridge crossing
the Wisconsin River at Muscoda (Grant County) in August 1960.

The paddlefish is today greatly reduced in the Mississippi River,
from which it was taken in great numbers during the early 1900’s
(Coker 1930). Nord (pers. comm. — Feb. 27, 1964) reports that
44,857 pounds of this species were taken commercially in 1961.
Only 801 pounds came from Pool 10 and none from Pool 11. Harlan
and Speaker (1956) write that the paddlefish in the boundary
waters of Iowa is not considered an important angling fish. Only
occasional specimens are caught, whereas most of them are illegally
hooked by snagging, largely below dams or obstructions on the
Mississippi River.

Lake sturgeon — Acipenser fulvescens Rafinesque. This species is
rare to uncommon on the lower Wisconsin River and the Mississippi
River opposite Crawford and Grant Counties. Greene (1935) re¬
cords a report from the Wisconsin River at Prairie du Sac. Nord
(pers. comm, — ^Sept. 24, 1962) reports it as rare in the Mississippi
River in Pool 10. Harlan and Speaker (1956) list several collections
from the Mississippi River opposite Crawford and Grant Counties,
and report that it is rarely, if ever, caught on hook and line,

Shovelnose sturgeon — Scaphirhynchus platorynchus (Rafines¬
que). W(2-8), W(12-13), W16, W19 (4 collections; 5 individuals).
The shovelnose sturgeon is commonly taken on hook and line in
the lower Wisconsin and the upper Mississippi Rivers. I have ex¬
amined several specimens between two and three feet long. A
specimen caught at Boscobel (W16) weighed just two pounds and
was 27% inches in total length (part of caudal filament was missing
for which no allowance was made in measurement) . The specimens
captured during the summer of 1962 were taken in deep water
either by hook and line or with boom shocker.

Longnose gar — Lepisosteus osseus (Linnaeus). W4, W8, W16,
W17, W19, W(20-22), W21, M3-6, C8 (13 collections; 35 individ¬
uals) . The longnose gar is common in the Wisconsin and Mississippi

94 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Rivers and the lower reaches of their larger tributaries. With the
seine we took these frequently in very shallow, sand-bottomed bays
on downstream sides of islands in both the Wisconsin and Missis¬
sippi Rivers. A longnose gar captured at W17 with total length of
26% inches weighed 22.1 ounces.

Shortnose gar — Lepisosteus platostomus Rafinasque. W14,
W (20-22), W21, R9, Cll (5 collections; 8 individuals). In south¬
ern Wisconsin the shortnose gar is found in the same habitat as
the longnose gar. Nord (pers. comm. — Sept. 24, 1962) reports the
shortnose gar as abundant in the Mississippi River at the conflu¬
ence of the Wisconsin River and the longnose as common. The short¬
nose is heavier-bodied than the longnose gar. A shortnose captured
at W14 with a total length of 22% inches weighed 23.7 ounces.

Bowfin — Amia calva Linnaeus. W19, M3 (2 collections; 3 indi¬
viduals). The bowfin is present in both the Wisconsin and Missis¬
sippi Rivers. Greene (1935) examined specimens from Knapp
Creek, Richland County, and Pine River, one mile west of Gotham,
Richland County. My specimens were taken from sloughs. Nord
(pers. comm. — Sept. 24, 1962) reports this species as abundant
from the Mississippi at the mouth of the Wisconsin River.

Mooneye — Hiodon tergisus LeSuer. W(2-8), W(12-13), W18,
W (20-22), M5, C9, CIO (7 collections; 11+ individuals). The
mooneye appears to be common in the Wisconsin River and occa¬
sionally found in the lower reaches of its larger tributaries. Nord
(pers. comm.- — Sept. 24, 1962) reports this species as common in
the Mississippi River.

Goldeneye — Hiodon alosoides (Rafinesque). Greene (1935) ex¬
amined collections from Lake Pepin of the Mississippi River, some¬
what north of the area under consideration. Nord (pers. comm. —
Sept. 24, 1962) reports the goldeneye as uncommon in Pool 10 of
the Mississippi River.

Gizzard shad — Dorosoma cepedianum (LeSueur). W16, W19,
W20-23, Ml-6, R9 (13 collections; 521+ individuals). This species
is abundant on the Wisconsin River from Boscobel down to its
juncture with the Mississippi River. Large numbers of young were
taken in the quiet and shallow waters of both the Wisconsin and
Mississippi Rivers. We found a number of shad in a landlocked
pool (25 feet wide, 30 feet long and 4 feet in greatest depth) on
an island on the Mississippi River about % mile below the Lynx-
ville Dam, Crawford County.

Skipjack herring — Alosa chrysochloris (Rafinesque) Greene
(1935) writes: “Since the construction of the Keokuk Dam, the
skipjack is said by fishermen to have become very much less com-

1966]

Becker — Fishes of Southwestern Wisconsin

95

mon if not extinct in Wisconsin waters.” Nord (pers. comm. — Sept.
24, 1962) writes that the skipjack herring has not been seen or
collected from the Mississippi River in recent years by survey
parties. His knowledge of this species in Wisconsin is confined to
citations in the literature. Coker (1929) gives evidence that the
construction of the hydroelectric dam across the Mississippi River
at Keokuk, Iowa, may have been responsible for the marked reduc¬
tion of the river herring in the Upper Mississippi River. Of the
Mississippi River opposite Iowa, Harlan and Speaker (1956) write:
“The fish has not been taken in the last twenty years and is now
thought to be rare or absent.”

Brown tmut—Salmo trutta Linnaeus. Rl, R6, Rll, R18, R20,
C3, C5, C13, G2~4, G6, G8, G9, G27, G34, G36, 14, LI (19 collec¬
tions; 95+ individuals). The head- waters of the many streams
in this region are frequently suited to hold this species. Reproduc¬
tion is to a large degree limited. The trout fishery in southwestern
Wisconsin is largely dependent on a continued stocking program.
Truog (in conversation) reported that early in the spring trout
have been taken from the Wisconsin River at Spring Green.

Rainbow trout—Salmo gairdneri Richardson. Rl, R20, C3 (3
collections ; 3+ individuals) . The rainbow has been stocked in these
waters on a put-and-take basis. Because of its migratory habit,
this species, unless caught during the season when stocked, may
be altogether lost to the trout fisherman. Natural reproduction is
limited.

Brook trout — Salvelinus fontinalis (Mitchill). C5 (1 collection;
7 individuals). Truog has records of small wild populations of
brook trout from the headwaters of the following streams : Fancy
Creek (6/5/58), Malancthon Creek (6/19/58), and Hawkins
Creek (7/22/60) . All three streams are tributary to the Pine River
in Richland County. He also found small wild populations of brook
trout in Crooked Creek, Grant County (4/13/61) . The trout ranged
from 3.9 to 10.3 inches.

The brook trout does poorly in southwestern Wisconsin. Water
from springs cold enough to support brook trout quickly warms
up, precipitating conditions unfit for this species. Although south¬
western Wisconsin was originally brook trout range (Brasch, et ah,
1958), today trout species tolerant of higher temperatures must
fill the niche vacated by them.

Quillback — Carpiodes cyprinus (LeSueur). W(2-8), W2-5,
W7-10, W12-13, W15-23, Ml-3, M5, G14, 110 (28 collections;
512+ individuals). The quillback is common on both the Wisconsin
and Mississippi Rivers. It may also be taken in their larger, heavily

96 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

silted tributaries. Reproduction is especially high in the Wisconsin
and Mississippi Rivers, where the young were taken in dense
schools from quiet water, often no more than six inches deep, over
silty bottom.

River carpsucker — Carpiodes carpio (Rafinesque) . W(2-8), W2,
W8, W9, W (12-13), W15-18, W (20-22), Ml, R3, R17, G7, 12, 15
(17 collections; 69+ individuals). The range and habitat of this
species appear to be similar to those of the quillback. Nord (pers.
comm. — Sept. 24, 1962) lists it as abundant in the Mississippi
River. Although adults are often mistaken for the quillback, the
river carpsucker has a distinct tubercle in the middle of the lower
lip which is absent in the quillback. Adult river carpsuckers reach
large size. One specimen from the Wisconsin River weighed four
pounds 151+ ounces and was 201+ inches in total length.

Highfin carpsucker — Carpiodes velifer (Rafinesque). W(2-8),
W (12-13), W16, W17, M5 (6 collections; 57+ individuals). The
highfin carpsucker is confined to the Wisconsin and Mississippi
Rivers and usually in moderate to swift currents. Its distribution
along these waterways is probably more extensive than the col¬
lections indicate. Many carpsucker young were taken which are
impossible to distinguish between C. velifer and C. carpio. Not
until they reach a length of 75 to 100 mm. can these species be told
apart.

Bigmouth buffalo — Ictiobus cyprinellus (Valenciennes), W18,
W20, W21, W22, M4, R9, G14, 17, 110, L8 (10 collections; 18 +
individuals). This species is common locally in medium to large¬
sized rivers in large holes where the current is sluggish.

Smallmouth buffalo — Ictiobus bubalus (Rafinesque). W(2-8),
W8, Wll, W16, W( 20-22), M3, M5, C14 (8 collections; 20 individ¬
uals). The smallmouth buffalo is a large-water species, although
occasionally taken from the mouths of small streams. A single
specimen, 44 mm. long, was captured near the mouth of Richland
Creek (C14) where it was only 10 to 15 feet wide. Most of this
species captured during the summer of 1962 were young-of-the-
year.

Black buffalo — Ictiobus niger (Rafinesque). W(2-8). The single
specimen Truog and I captured with boom shocker from the Wis¬
consin River was 22 inches in total length and weighed six pounds,
1/2 ounce. Nord (pers. comm. Sept. 24, 1962) reports this species as
uncommon in the Mississippi River.

Blue sucker — Cycleptus elongatus (LeSueur), W(2-8), W13,
W (20-22). A limited population of this interesting sucker is
present in the lower Wisconsin River. On a boom shocking trip

1966]

Becker — Fishes of Southwestern Wisconsin

97

W(2-8) between Spring Green (W2) and Lone Rock (W8), six
individuals were brought to net and, three more were seen at the
electrodes. The smallest fish captured measured 22.8 inches in total
length and the largest 29.0 inches. The latter weighed just eight
pounds. These fish were all taken from deep water adjacent to
islands where the banks were badly eroded and a great number of
trees had toppled into the water. At W13 with seine I captured a
single young-of-the-year 34 mm. long. Harlan and Speaker (1956)
report it as uncommon to rare in the Mississippi.

Northern redhorse—Moxostoma macrolepidotom (LeSueur).
W(2-8), W2, W3, W6, W8, WIO, W ( 12-13 ), W14-16, W18, W(20-
22), W21, M3, R9, R16, R17, R20, R21, C7-10, C14, G13, G14,
G16, G28, G30, G31, G41, G43, 11-3, 17-11, LI, L2, L4, L8, L14,
LI 6 (49 collections; 323+ individuals). Harlan and Speaker
(1956) write that the northern redhorse is the most common spe¬
cies of sucker in the Mississippi River. It is abundant in the Wiscon¬
sin River and in medium to large-sized tributaries to -the Wisconsin
and Mississippi Rivers. Occasionally this species is taken in the
lower reaches of small streams opening into these rivers. A speci¬
men from the Wisconsin River, 17% inches in total length, weighed
two pounds five ounces.

Golden redhorse — Moxostoma erythrurum (Rafinesque) ,
W(2-8), W2, W7, Wll, W(12-13), W14-18, Ml-2, M6, R9, C7,
G16, 12, 17-12, L2, L7, L16 (26 collections; 184 individuals). The
golden redhorse is frequently taken with the northern redhorse,
although its distribution is more spotty. Where encountered on the
Wisconsin and Pecatonica Rivers, it appears abundant.

Silver redhorse — Moxostoma anisurum (Rafinesque). Wl, WIO,
W14, W16, W18, W (20-22), W23, R9, R13, R16, C8, 18-12, L3,
L4, L6, L16 (20 collections; 66+ individuals). The silver redhorse
was encountered less frequently on the Wisconsin River than the
northern and golden redhorses. Where found, only one or two
specimens were captured per station. On the Mississippi River it
is considered uncommon (pens. comm. — Nord, Sept. 24, 1962). It
is common on the Pecatonica River and its medium-sized
tributaries.

Greater redhorse — Moxostoma valenciennesi Jordan. W(2-8).
A single specimen 51 mm. long was captured from the Wisconsin
River somewhere between Spring Green (W2) and Lone Rock
(W8). This species is rare in southwestern Wisconsin. There is no
record of it from the Mississippi River opposite Crawford and
Grant Counties.

Black redhorse— Moxostoma duquesnei (LeSueur). Although
ascribed to southern Wisconsin (Hubbs and Lagler 1958), this

98 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

ispecies has never been recorded from counties in the present study.
Greene (1935) captured a single specimen from Black Earth Creek,
Dane County, close to the Iowa County line (Black Earth Creek
flows through the extreme northeastern corner of Iowa County,
where it joins the Wisconsin River). This sucker is extremely rare
on the western edge of its range, which includes southern Minne¬
sota and northeastern Iowa. Its presence in southwestern Wiscon¬
sin is probable.

Spotted sucker — Minytrema melanops (Rafinesque). Wll, W18,
W19, M4, R17 (6 collections; 20 individuals). Nord (pers. comm.
— Sept. 24, 1962) lists this species as common for the Mississippi
River in Pool 10. We encountered the spotted sucker in sloughs and
backwaters of the Wisconsin River. It prefers little or no current.
Ten individuals were collected from station R17 on the lower Pine
River (Richland County), by far the largest collection made.

Lake chubsucker — Erimyzon sucetta (Lacepede). W19, RIO.
This species was taken only from the Wisconsin River (one individ¬
ual) and the lower extremity of Indian Creek (one individual).
Greene (1935) captured it from the Wisconsin River at Boscobel
and at Blue River. A young-of-the-year captured from a slough at
station W19 was 33 mm. long.

Northern hog sucker — Hypentelium nigricans (LeSueur).
W(2-8), W(12-13), W14, W(20-22), Rll, R14, R16, R18, R20,
Cl, C3, C4, C6-10, C14, Gl, G5, G15, G31, II, 12, 17-11, L2, L4,
L8 (32 collections; 201+ individuals). This species is abundant
locally in medium to large streams, especially in swift water. It is
uncommon on the Mississippi River (pers. comm. — Nord, Sept. 24,
1962).

White sucker — Catostomus commersoni (LacepMe). W13, W16,
W19, W20, M4, Rl-23, Cl, C3-8, C12-14, Gl-9, Gll-12, G14-47,
Il_5, I7_i2, Ll-2, L4, L6, L8-15 (107 collections; 4,679 individ¬
uals) . The white sucker is the most ubiquitous sucker in Wisconsin.
In southwestern Wisconsin its distribution is general. Although
abundant in small to medium-sized streams, it is less common in a
large river like the Wisconsin and extremely rare in the Mississippi.

Carp — Cyprinus carpio Linnaeus. W (2-8), W8, W (12-13), W18,
W (20-22), W21-22, M2, M5-6, R7-9, Rll, R16, R20, R23, C9,
Cll, G14-15, G28, G31, G41, 12, 17, L2-4, L16 (31 collections;
170 individuals). The carp is commonly taken in most medium¬
sized streams or large rivers. It is abundant in the Mississippi
River (pers. comm. — Nord, Sept. 24, 1962).

Central stoneroller — Campostoma anomalum pullum (Agassiz).
W19, M3, Rl-2, R4-14, R16, R18-19, Cl, C3-8, CIO, C12-14, Gl-4,

1966]

Becker — Fishes of Southwestern Wisconsin

99

G6-7, G9, Gll-26, G28-30, G32-38, G40-42, G44-47, II, 14, 17-12,
LI, L4, L6-16 (90 collections; 8,198+ individuals). The central
stoneroller was the most abundant fish found in smaller streams :
several hundred individuals were taken at many stations. It is
generally found in large schools in riffle areas, although in a few
instances I have encountered considerable numbers in quiet pools.

Largescale stoneroller— Campos^oma anomalum oligolepis Hubbs
and Greene. M5-6, 19, Ill, L2, L6, L13 (7 collections; 16+ indi¬
viduals). The largescale stoneroller, although extremely common
in central and northern Wisconsin, is seldom taken in southern
Wisconsin, where it is supplanted by the central stoneroller.

I have noted both Campo stoma a. pullum and Campos toma a.
oligolepis from the collections at stations 19 and Ill, with single
specimens from each station showing intergrade characters.
Nybakken (1961) has found intergrades from my collections at
112, L6 and L13. He has recently found both forms of Campostoma
and intergrades in the Dodge Branch of the W. Pecatonica River
from southeastern Iowa County.

Campostoma prefers swift waters in medium to small-sized
streams. In southern Wisconsin the greatest numbers were taken
in streams only a few feet wide.

Longnose dace — Rhinichthys cataractae (Valenciennes). R19,
R20, Cl, C3, C5, C7, C8, C13, Gl-5, G7, G12, G13, G15, G16, G20,
G29, G30, G32, G33, G35, G37, G44, G45, II (28 collections; 274
individuals). The longnose dace is common in the small and
medium-sized streams of Crawford, Richland and Grant Counties.
Greene (1935), although sampling heavily in the same counties
during the late 1920’s, failed at that time to find this species. This
suggests that an expansion of the range may have occurred in the
interim. The longnose dace from the northeastern corner of Iowa
may have assisted in this extension of range. Our record from
Menominee Creek (G45) is so close to Illinois that we may expect
to find this species in the northwestern corner of that state. Forbes
and Richardson (1920) report this minnow as rare in Illinois, with
a single individual captured near Waukegan and three individuals
from Big Creek in the extreme southern part of the state.

Blacknose dsice—Rhinichthys atratulus (Hermann). Rl, R4,
R6-8, RlO-14, R18-20, Cl, C3, C5, C7, CIO, C13, G2-8, Gll, G12,
G29, G33, G41, G42, II (33 collections; 1,365+ individuals). The
blacknose dace is a common inhabitant of small, spring-fed streams
throughout the region. It is frequently taken with the preceding
species, although it is less tolerant of high water temperatures.

Hornyhead chub — Hyhopsis biguttata (Kirtland). W15, C14,
G12, G15-47, II, 17-12, LI, L2, L4, L6-9, L16 (50 collections;

100 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

997+ individuals). For some peculiar reason this species, which
appears to adapt itself to a variety of conditions, is rare to occa¬
sional in the streams tributary to the Wisconsin River, but it is
abundant and widely distributed in the streams of Grant, Iowa,
and Lafayette Counties, which flow southward into the Mississippi
River. Greene (1935) speculates that the hornyhead chub finds
unsuitable this large central portion of the driftless area because
it is underlain with potsdam sandstone. The absence of pebbles,
required for spawning, may be the limiting factor.

Silver chub — Hybopsis st07-eriana (Kirtland). W(2-8), W(12-
13), W16, W (20-22), W23, Ml-3, M5-6, R16, 12, 13, 15 (15 col¬
lections; 50+ individuals). The silver chub is a large-water form.
We captured it only from the Wisconsin and Mississippi Rivers
and the lower extremities of their large tributaries. The large
adults of the silver chub are taken generally in deep water, can
effectively be collected with boom shocker, and will readily take
earthworms on a hook.

Speckled chub — Hybopsis aestivalis (Girard). Wl-6, W(2-8),
W( 12-13), W22, M5, R17, 12 (12 collections; 574+ individuals).
The speckled chub is locally abundant in large rivers. We cap¬
tured it in considerable numbers from shallow riffs over sand on
the Wisconsin River between Arena (Wl) and the mouth of Sneed
Creek (W6). Elsewhere on the river it was uncommon.

Creek chub — Semotilus atromaculatus (Mitchill). W14-16, W19,
Ml, Rl-16, R18-20, R22, R23, Cl, C3-8, CIO, C12-14, Gl-27, G29,
G32-47, 11-4, 17, 18, 112, LI, L2, L4, L8-9, Lll-16 (98 collections;
3,190+ individuals). The creek chub is abundant in small to
medium-sized streams but uncommon in the large rivers. It is a
minnow adapted to a wide variety of habitats and a wide range
in water temperatures. Next to the bluntnose minnow it is the
most widely distributed minnow in the region.

Southern redbelly dace — Chrosomus erythrogaster (Rafinesque) .
RIO, G4, Gll, G12, G15-18, G21-27, G29-30, G32, G33, G35-36,
G42, G45-46, II, 14, 112, LI, L7, L9, Lll, L13, L15 (33 collections;
1,542+ individuals). The southern redbelly dace is abundant in
small clear streams up to ten feet wide and common in some
medium-sized streams. We failed to capture this species in many
likely-looking streams of Crawford and Richland counties.

Redside dSiCe—rClinostomiis elongatus (Kirtland). Rll, R12, R14,
R18 (5 collections; 117+ individuals). The redside dace is abun¬
dant locally in the tributaries of the Pine River (Richland County),
in small streams, most of which have trout populations. We en¬
countered it nowhere else in the region. Some hybridization with
the northern common shiner was noted.

1966] Becker — Fishes of Southwestern Wisconsin 101

Golden shiner — Notemigonus crysoleucas (Mitchill). Wll, W14,
W16, W19-20, W22, Ml-5, R9, R17, G14 (15 collections; 467 -f
individuals). The golden shiner was taken primarily in the sloughs
and backwaters of the Wisconsin and Mississippi Rivers and in the
lower extremities of the larger tributaries of the Wisconsin River,
Seldom was it taken from water with current. Several collections
came from landlocked pools adjacent to the large rivers.

Bullhead minnow — Pimephales vigilax (Baird and Girard). Wi¬
ll, W13-17, W19-23, Ml-3, M5~6, R9, R17, R23, G14, 12, 13 (37
collections; 4,2894- individuals). The bullhead minnow is found
commonly in the Wisconsin and Mississippi Rivers, in large tribu¬
taries to these rivers, and in the lower extremities of small streams
flowing into them. It is more abundant than any other fish, with
the exception of the spotfin shiner, in the larger river. It has been
taken from sloughs and from water with moderate current.

Bluntnose minnow — Pimephales notatus (Rafinesque). Wl-2,
W6, W9-22, R2-3, R5-21, R23, Cl, C4-8, CIO, C14, Gl, G3, G5,
G7, G9, Gll-47, 11-9, Ill, 112, L2, L6-16 (113 collections; 6,770+
individuals) . The bluntnose minnow is probably the most successful
fish in its distribution. It is adapted to great variations in water
size, temperature and quality. Although most abundant in small
streams, large numbers were taken from several stations on the
Wisconsin River, where it occurred with the bullhead minnow.
Although I did not find the bluntnose in the Mississippi River,
Harlan and Speaker (1956) report several records from the river
opposite Crawford and Grant counties.

Fathead minnow—Pimephales promelas Rafinesque. W4, W6,
W16, W19, M5, R2, R6-9, R13, R15, R17, C4-6, C8, C12-14, G3-4,
G12, G16-17, G23-26, G33, G36, G40, 12, L15 (35 collections; 245
individuals). The fathead is most commonly taken in moderate¬
sized streams which are silty. It seems to be generally distributed
in the region, occurring in small streams to large rivers but seldom
abundant. The largest collection (114 individuals) was taken from
Rattlesnake Creek (G17). In numbers this species is far less suc¬
cessful than the other species in this genus.

Pugnose minnow — Opsopoeodus emiliae Hay. Wll, W18-21,
M3-4, R9 (9 collections; 120 individuals). This small minnow was
taken from sloughs of the lower Wisconsin River where the bottoms
were covered with dead leaves and other organic debris. Seventy-
two young-of-the-year were captured near the west end of Newton
Island (W20). A more intensive survey of quiet waters in the
lower reaches of large tributaries to the Wisconsin and Mississippi
Rivers would undoubtedly disclose a greater distribution than the

102 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

present survey indicates. In another paper I have reported the
presence of this species on the western end of Lake Poygan in
eastcentral Wisconsin (Becker 1964). The pugnose minnow, a
southwestern form, appears to have extended its range in Wiscon¬
sin since Greene made his survey.

Suckermouth minnow — Phenacohius mirabilis (Girard). M5-6,
R3, RlO-12, R16-17, R19-20, C7, G12, G14~16, G18-20, G21, G23,
G26, G28-35, G37, G39-42, G44-45, G47, 11-2, 17, 110, 112, L2,
L6, L8 (45 collections; 389 individuals). The suckermouth minnow,
a southern form, has established itself well in the driftless area of
the state. It is a common minnow in rivers of all sizes, but the
largest collections were made in medium-sized streams. It prefers
swift-running water over a gravel bottom, although we have taken
it from a wide variety of habitats.

Brassy minnow — Hyhognathus hankinsoni Hubbs. W2-3, W6-9,
R2-4, R7-11, R15, R17, R19-22, Cl, C4, C7~8, 12, 14 (28 collec¬
tions; 334 individuals). The distribution of the brassy minnow in
these counties is spotty. It is generally taken in moderate-sized
streams or small rivers. The collections of this species in the upper
section of the Wisconsin River were unexpected. Where it ,was
taken, we generally did not find the silvery minnow (Hyhognathus
nuchalis). Greene (1935) has pointed out that these closely-related
species are complementary; i.e., their ranges are separate and
adjacent. Fair numbers were found in Knapp Creek, Mill Creek,
Willow Creek (Richland County streams), in the Kickapoo River
(Crawford County) and in Otter Creek (Iowa County). Greene
(1935) has only two records of this species in the area covered by
this survey. It appears that in recent years this minnow has become
more successful in the driftless area.

Silvery minnow — Hyhognathus nuchalis Agassiz. W(2-8), W2,
WIO, W13-17, W19-22, Ml, R5, RIO, R16, R20, Cl, C6, C9, C14,
G3, G5, G7, G9, G12-14 (31 collections; 626 individuals). The sil¬
very minnow inhabits medium to large rivers and the lower extrem¬
ities of small streams opening into such waters. In the last, many
adults were captured. The smaller waters may serve as spawning
areas for this minnow. Young-of-the-year numbering 230 were cap¬
tured from the Blue River near its mouth. (G14).

Ozark minnow — Dionda nuhila (Forbes). G32, G34, G37 (3 col¬
lections; 37 individuals). The Ozark minnow has been reported
from streams in Iowa and Lafayette counties (Greene, 1935). The
present survey adds the upper end of the Platte River and its tribu¬
taries, Grant County. The Ozark minnow was taken in clear small
to medium-sized streams where it travels in fairly dense schools

1966]

Becker — Fishes of Southwestern Wisconsin

103

near the surface of the water. Because of the schooling habit, this
species would be easy to miss.

Common ishiner — Notropis cornutus (Mitchill) . W7, W16, W(18-
19), W19, W23, M6, R6-9, Rll, R12, R14, R18, Cl, Gl, G5, G7,
G12-47, II, 12, 17-10, 112, Ll-4, L6-10, L12, L14-16 (76 collec¬
tions; 4,621 -f individuals). The common shiner prefers streams
of small to medium size. It is captured over a wide variety of
bottom types and found in greatest numbers in clear water, al¬
though it can tolerate considerable turbidity. Recently it has become
extremely rare in the Mississippi River.

Emerald shiner- — Notropis atherinoides Rafinesque. Wl-10,
W12-23, Ml-3, M5-6, R3, R9, R17, R20, R21, R23, C9~10, C14,
G5, G14, G16, G31, G43, 12, 15 (50 collections; 1,636-|- individuals).
The emerald shiner occurs commonly in large streams and rivers
and at the lower extremities of small streams opening into rivers.
Although abundant on the Wisconsin River, it is outstripped in
numbers by the spotfin ishiner, another member of the same genus.
Our records indicate that on the Mississippi River the situation is
just reversed, with the emerald shiner more abundant than the
spotfin shiner.

Rosyface shiner- — Notropis rubellus (Agassiz). G19-22, G25,
G27-30, G36-40, G43-44, G47, II, 17, 19-10, 112, L2-3, L5-8, L14,
L16 (30 collections; 658+ individuals). The rosyface shiner gen¬
erally inhabits medium-sized streams, although several populations
were located in small streams. This shiner travels in schools near
the surface of the water. Its apparent absence from the streams of
Crawford and Richland Counties is difficult to explain.

Spotfin ^hmev—N otropis spilopterus (Cope), Wl-23, Ml-6, R3,
R5, R7-9, R13, R16-17, R19-21, R23, Cl, C4, C7-10, C14, G5,
G14-15, G43, 12-3, 15, 112, L2, L5~8, LIO, L12, L14, L16 (74 col¬
lections; 27,238+ individuals). The spotfin shiner is encountered
in moderate-sized streams and rapidly increases in abundance with
the increase in stream size. It was by far the most abundant fish
captured from the Wisconsin River and one of the most common
in the Mississippi. The spotfin prefers a moderate current and is
generally taken in shallow water over sand bottom. It is far less
common in the quiet waters of sloughs.

Spottail shiner— Notropis hudsonius (Clinton). W(2-8), W2,
Wll, W14-16, W19~23, Ml-6 (19 collections; 2,103+ individuals).
We collected the spottail only from the main channel and back¬
waters of the Wisconsin and Mississippi Rivers. It is uncommon
to common in the Wisconsin River. For most stations it was re¬
corded with the collection of a single specimen. At Boscobel (W16)

104 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

we collected 95 individuals in a single collection. It is one of the
most abundant fish species in the Mississippi River, where hundreds
were seined at each station. It was common in sloughs and abun¬
dant in moderate currents.

Weed shiner— Wo tropis texanus (Girard). W2, Wll, W16, W19,
W (20-22), W21-22, M4, Cl, Gl, 16 (14 collections; 87 individuals).
The weed shiner is an inhabitant of the quiet or sluggish sections
of medium-sized streams to large rivers. Occasionally it may be
taken from the lower reaches of small streams emptying into a
large river. The distribution of this southern minnow is spotty. It
appears to be rare to uncommon on the Wisconsin River, where
collections were small; i.e., from one to seven individuals per col¬
lection. Fifty-six individuals were taken in a single collection (Cl)
from near the mouth of Gran Grae Creek.

River shiner — Notropis hlennius (Girard). Wl-4, W6-7, W9-10,
W(12-13), W14, W16-23, Ml-3, M4-5, G5 (27 collections; 1,606 +
individuals). The river shiner is found commonly in the Wisconsin
and Mississippi Rivers and occasionally in the lower extremities
of their tributaries. It occurs in fair numbers on the Wisconsin
River and is abundant on the Mississippi, where it comprises a
large percentage of the catch. Our records show hundreds taken
from the Mississippi River where the waters sampled had a moder¬
ate to fast current. Next to the spottail shiner, it was the fish most
commonly captured from the Mississippi.

Blacknose shiner — Notropis heterolepis Eigenmann and Eigen-
mann. This minnow was not collected during the survey, although
Greene (1935) reports it from Bear Creek in the southeastern
corner of Richland County.

Sand shiner — Notropis stramineus (Cope), Wl, W5-10, W(12~
13), W13-19, W21, W23, M3, M5-6, G14~16, G22, G28, G37-41,
G43-45, G47, 17-9, 112, L2, L6-8, L12, L14, L16 (46 collections;
1,631+ individuals). The sand shiner is rare in small streams,
fairly common in medium-sized streams and small rivers, and ap¬
pears to diminish in numbers in large rivers except locally where
large concentrations may be found. This species prefers running
water and is most frequently taken over a sand bottom.

Northern mimic shiner — Notropis volucellus volucellus (Cope).
Harlan and Speaker (1956) record several collections of this sub¬
species from the Iowa County side of the Mississippi River opposite
Allamakee and Clayton counties. Its occurrence on the Wisconsin
side opposite Grant County is probable. Christenson and Smith
(1965) report this form from a backwater of the Mississippi River
west of Fountain City, Buffalo County, Wisconsin.

1966]

Becker— Fishes of Southwestern Wisconsin

105

Channel mimic shiner — Notropis volucellus wickliffi Trautman,
M2-3 (2 collections; 4 individuals). Three collections of the chan¬
nel mimic shiner taken July 1944, by John D. Black, from the
Mississippi River in Pool 9 opposite Crawford County are in the
Museum of Zoology, University of Wisconsin, Madison. This sub¬
species is present only in the Mississippi River, where it is
uncommon.

Ghost shiner — JSlotropis huchanani Meek. Three collections of
this species taken in July 1944, from Crawford County and Alla¬
makee County (Iowa) by John Greenbank and Melvin Monson are
in the Museum of Zoology, University of Wisconsin, Madison. Nord
(pers. comm.— -Jan. 10, 1964) reports that during the 1963 Missis¬
sippi River small fishes survey this species was not encountered in
the Mississippi River opposite Wisconsin, but it appeared down¬
river in the collections between Pools 13 (Bellvue, Iowa) and 26
(Beechridge, Alexander County, Illinois).

Bigmouth shiner — Notropis dorsalis (Agassiz). Wl-3, W6, W9,
W13, W17, W19, W22-23, Ml-3, R2-4, R6-8, RlO-14, R16, R19-
21, Cl, C3-8, C14, G4-5, G7-9, Gll-15, G17-20, G22~23, G25-27,
G29~30, G32-35, G39~42, G45-46, 11-2, L6 (71 collections; 1,330 +
individuals) . The bigmouth shiner occurs commonly over sand
bottoms in streams of small to medium size. In large rivers this
species is rare to uncommon. On the Wisconsin and Mississippi
rivers its presence was based on the capture of single individuals
at stations where found.

Pallid shiner— Notropis amnis Hubbs and Greene. M3, W21. I
captured a single specimen from the Mississippi River at
Wyalusing and another from the Wisconsin River between the
mouths of Gran Grae Creek and Little Kickapoo Creek. Three speci¬
mens were collected by John Kennedy from the mouth of Big Green
River at its juncture with the Wisconsin River (Grant Co.) on
Aug. 15, 1960. Seven collections from the Mississippi River opposite
Crawford County were made by Greenbank, Monson and Black in
July to August 1944. These and additional collections from the
Upper Mississippi River are in the Museum of Zoology, University
of Wisconsin, Madison. Nord (pers. comm. — Jan. 10, 1964) reports
that this species was captured from Pools 9 and 13 during the 1963
Mississippi River small fishes survey.

Flathead catfish — Pylodictis olivaris (Rafinesque) , James Kin-
cannon of Blue River, Wisconsin, caught 12 flathead catfish from
the Wisconsin River between the Blue River bridge and a point
about a mile upstream (vicinity of W15). These fish were caught
on set lines using bullheads and catfish as bait. The average size

106 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

of the fish caught was 14.6 pounds, the smallest weighing five
pounds and the largest 35 pounds. He writes:

They were all taken along- steep, grassy banks which displayed a pre¬
dominance of hard clay or dirt rather than sand. Old tree roots and logs
were also in evidence at each site. A large majority of the fish contained
spawn. Although I made sets at areas of unstable (sand) banks no fish
were taken in this type of place. Once again I noticed that unseasonably
cool weather has an adverse effect on the number of fish taken. Also as in
the past the catch tapered off near the middle of June.

Two flathead catfish were seen by Truog on August 18, 1962, on
a boom shocking run near Bridgeport (W22). According to com¬
mercial fishermen on the Mississippi River, this species in the vicin¬
ity of Prairie du Chien has been on the decline in recent years.

Blue catfish — Ictalurus fnrcatus (LaSueur). Greene (1935) ex¬
amined a collection from the Mississippi River near Lansing, Iowa,
opposite Crawford County. Eddy and Surber (1947) write that it
formerly occurred in the Mississippi River and larger tributaries
from Minneapolis southward but that it is now very rare in Minne¬
sota waters with no specimens taken in recent years. Nord (pers.
comm.^Sept. 24, 1962) does not list this species from Pool 10 of
the Mississippi River.

Channel catfish — Ictalurus punctatus (Rafinesque). Wl, W(2-
8), W15-16, W18-19, W (20-22), W22, R9, R17, CIO, 13, L14 (12
collections; 18 individuals). This species is common in medium to
large-sized rivers and is occasionally taken in smaller tributaries
to such streams. The channel cat is perhaps the most important
game fish on the Wisconsin and Mississippi Rivers. From the
Mississippi River Nord (pers. comm. — Feb. 27, 1964) reports that
in the commercial catch this species ranked third in numbers after
bluegills and crappies in 1962-63, although it has dropped in recent
years. The channel catfish recently has been the object of consider¬
able research on the lower Wisconsin and the Mississippi Rivers.

Yellow bullhead — Ictalurus natalis (LeSueur). W12, G15-16 (3
collections ; 4 individuals) . The yellow bullhead is uncommon in this
region. Nord (pers. comm. — Sept. 24, 1962) reports this species
from the Mississippi River in Pool 10. Greene (1935) captured it
from several stations on the Mississippi opposite Crawford County.
Harlan and Speaker (1956) report that this species is taken occa¬
sionally in the Mississippi River.

Brown bullhead — Ictalurus nehulosus (LeSueur). M3. Greene
(1935) captured this species from the Mississippi River in the
vicinity of Lynxville (Crawford County). Several collections in the
University of Wisconsin Museum of Zoology were made by Green-
bank and Monson from the Mississippi River opposite Allamakee

1966]

Becker — Fishes of Southwestern Wisconsin

107

County, Iowa, in July and August 1944. Other records in the same
vicinity are recorded by Harlan and Speaker (1956), who write
that it inhabits the sloughs and river lakes.

Black bullhead — Ictalurus melas (Rafinesque) . W14, W16, W19-
20, R17, G26 (6 collections; 139 individuals). This species is found
principally in the backwaters and sloughs of the Mississippi and
Wisconsin Rivers, and in large quiet pools in their tributaries. In a
sand-bottomed pool about 100 feet from the Wisconsin River we
captured hundreds of black bullheads ( W20) . The pool was approx¬
imately 60 feet long, 15 feet wide and had a maximum depth of
two and one-half feet. These fish must have been trapped in this
pool during high water.

Stonecat — No turns flavus Rafinesque. W12, W15, W18, G14,
G18--21, G30, G32, G34, G36-37, G40, G47, 13, 17-11 (21 collections;
68 4- individuals). This species is locally abundant in swift waters
over stony bottoms. It has been taken from the Mississippi and
Wisconsin Rivers and their tributaries. Nord (pers. comm. — Sept.
24, 1962) considers it uncommon in Pool 10 of the Mississippi
River. I have frequently captured the stonecat while electrofishing
among the boulders and rocks under highway bridges.

Tadpole madtom — Noturus gyrinus (Mitchill). M2-5, G31
(5 collections; 9 individuals). Nord (pers. comm. — Sept. 24, 1962)
lists the tadpole madtom as common on the Mississippi River. Else¬
where this species appears rare. Greene (1935) captured this
species from the Wisconsin River in the vicinity of Boscobel
(W16).

American eel — Anguilla rostrata (LeSueur). Greene (1935) ex¬
amined collections of the American eel from the Mississippi River
at Lynxville, Crawford County, and near Lansing, Iowa, opposite
Crawford County. Harlan and Speaker (1956) report only a single
collection since 1945 in the vicinity of Lansing. Nord (pers. comm.

, — Sept. 24, 1962) lists the eel as uncommon. Greene (1935) ascribes

the decrease of the eel from Wisconsin waters to dam construction
on the Mississippi River.

Central mudminnow — Umbra limi (Kirtland). Wll, W16, W19,
R4, R17, R22, Cl-2, C6, G6, GIO, 12-3, 15-6, 112 (16 collections;
64 individuals) . I have found the mudminnow under a wide variety
of conditions : small streams to large rivers, clear to turbid water,
cold, spring-fed to warm waters.

Grass pickerel — Esox americanus vermiculatus LeSueur. Wll,
W14, W16, W18-19, W (20-22), W21, M4, RIO, R17, Cl, C13, 15-6
(16 collections; 43 individuals). The grass pickerel is found in the
quiet backwaters of the Wisconsin and Mississippi Rivers, where

108 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

it is frequently taken with the northern pike. It is also found in the
lower reaches of tributaries of the Wisconsin River. Nord (pens,
comm. — Sept. 24, 1962) considers it uncommon in Pool 10 of the
Mississippi River. Contrary to its name, in many of the backwaters
from which we have taken the grass pickerel, there was practically
no vegetation. Several specimens were captured from sand-
bottomed pools which were entirely devoid of vegetation. It is my
opinion that this species has extended its range considerably in
southwestern Wisconsin since Greene made his survey in the late
1920’s, when he reports a single collection from the Mississippi
River in the vicinity of Ferryville.

Northern pike — Esox Indus Linnaeus. W8, Wll, W16, W18-22,
M3-4, R9-10, Cl, C14, G9 (18 collections; 59 individuals). The
northern pike is found in the sloughs and backwaters of the Wis¬
consin and Mississippi Rivers and in the lower reaches of their
tributaries. Nord considers it common in the Mississippi River and
writes (pers. comm. — Feb. 27, 1964) that a 1956 to 1958 study
showed lamprey scars on 68 fish, 25 of which were northern pike.

Muskellunge — Esox masquinongy Mitchill. Harlan and Speaker
(1956) consider the muskellunge as rare in the Mississippi. Nord
(pers. comm. — Feb. 27, 1964) reports that he knows of none taken
from the Mississippi below Minneapolis-St. Paul in recent years. I
have no authentic records of this species from the lower Wisconsin
River, although fishermen I have spoken to maintain that there
are a few large muskellunge in the vicinity of Blue River (Grant
County). Possibly a few may escape from Lake Wisconsin (Sauk
County), which gets an annual fingerling stocking program (Poff
and Threinen 1965).

Blackstripe topminnow — Fundulus notatus (Rafinesque) . W19.
The three specimens which I took from a debris-filled lagoon were
the first of this species reported from the unglaciated portion of
Wisconsin and point up the possibility that this species did cross
over into the Great Lakes watershed of Wisconsin via the Fox-
Wisconsin waterway at Portage, Wisconsin. Greene (1935) be¬
lieved this species found the unglaciated area ecologically unsuit¬
able and was unable to explain its presence on the upper Fox River.

Starhead topminnow — Fundulus notti (Agassiz). About 18 speci¬
mens were collected by Marlin Johnson, University of Wisconsin-
Madison, from a lagoon of the Wisconsin River, T8N R5E Sec. 9
N%, Iowa County, on June 31, 1965. Five specimens from this
collection have been placed in the Museum at Wisconsin State Uni¬
versity, Stevens Point.

Burbot — Lota lota (Linnaeus). Gl, G9. Three small specimens
were taken at station Gl and one at station G9. Truog records this

1966] Becker — Fishes of Southwestern Wisconsin 109

species from Rush Creek, Crawford County, TllN R6W Sec. 27,
April 27, 1963, and also from Blue River, Grant County, T7N
RIW Sec. 4, July 12, 1963. Nord (pers. comm. — Sept. 24, 1962)
considers the burbot as rare in the Mississippi.

Trout-perch — Fercopsis omiscomaycus (Walbaum). W21, Ml (2
collections; 2 individuals). Nord (pers. comm. — -Feb. 27, 1964) lists
this species as uncommon for the Mississippi River but says that
‘‘fairly large numbers” have been captured in the vicinity of La
Crosse. I would consider it rare on the lower Wisconsin River.

Pirate perch — Aphredoderus sayanus (Gilliams). W16, W19,
R23 (4 collections; 5 individuals). The pirate perch is rare to un¬
common in the Mississippi and Wisconsin Rivers and the lower
extremities of some of their tributaries. The specimens from the
Wisconsin River were taken from quiet backwaters. Two individ¬
uals, 83 mm. and 88 mm, long, were taken from Bear Creek (Rich¬
land County) in water with moderate current.

White bass— chrysops (Rafinesque) . W(2-8), W2, W4~5,
W8-11, W16-23, Ml-6, R9 (25 collections; 557 -f individuals). The
white bass is common to abundant on the lower Wisconsin and on
the Mississippi Rivers, occasionally taken in the lower extremities
of their larger tributaries.

Yellow bass— mississippiensis (Jordan and Eigenmann).
W2, W6, M2-5 (6 collections; 40 -|- individuals). The yellow bass
is uncommon on the lower Wisconsin River and uncommon to com¬
mon on the Mississippi River opposite Crawford and Grant Coun¬
ties. We encountered this species in small numbers at five of the
six stations which we sampled on the Mississippi River. Greene
(1935) took this species only twice on the Mississippi River oppo¬
site Crawford County and listed it as rare in Wisconsin. Today the
yellow bass is found in many lakes and larger rivers of southern
and eastcentral Wisconsin (Helm 1964). Although this extension
of range may result from the Fox-Wisconsin canal at Portage, it
is more likely that stocking programs are responsible for the rapid
range extension.

Yellow perch — Berea flavescens (Mitchill). W(2-8), W2, W6,
W8, WlO-11, W13-16, W19-21, W23, Ml-4, R23 (22 collections;
302 -f individuals). The yellow perch is common locally in the
Mississippi and common along the lower Wisconsin River. Two
specimens were captured from the lower end of Bear Creek (R23) .

Sauger — Stizostedion canadense (Smith). W(2-8), W(12-13),
W18, W(20-22), M2--3, C9, Cll, 12, 16 (9 collections; 34 individ¬
uals). The sauger was readily taken with the boom shocker from
the Wisconsin River, where it appears to be common. It is also

110 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

found in the lower extremities of its larger tributaries. Nord (pers.
comm. — Feb. 27, 1964) reports it as abundant in the Mississippi
and more frequently taken than the walleye. He reports that in a
recent creel census 4,299 saugers were observed as compared to
1,406 walleyes. With the seine we failed to pick up any young-of-
the-year. By contrast walleye fry were captured in numbers from
both the Wisconsin and Mississippi Rivers.

Walleye — Stizostedion vitreum vitreum (Mitchill). W(2-8), W6,
W9~ll, W (12-13), W13-17, W19, W21-22, M2-4, R17, Cl, C9,
C14, 13, L14 (25 collections; 219+ individuals). Nord (pers. comm.
— Sept. 24, 1962) lists the walleye as common in the upper Missis¬
sippi. The present survey captured many young-of-the-year on the
Wisconsin River; e.g., 76 at W15, 27 at W16, 19 at W13. Walleyes
are also found in the large, deep tributaries to the Wisconsin and
Mississippi Rivers.

Western sand darter — Ammocrypta clara Jordan and Meek.
Wl-5, W7, W9~10, W13-14, W16-17, W19, W20-23, Ml-3, M5-6
(25 collections; 246+ individuals). This darter is common to abun¬
dant on the Wisconsin and Mississippi Rivers. It is captured in
moderate to swift currents over fine sand where water depth is
from a few inches to 18 inches. It prefers extensive sand flats where
frequently it is the only fish taken.

Crystal darter — Ammocrypta asprella (Jordan). W(12-13). Six
specimens of this rare darter were collected from a rock shelf bot¬
tom in water depth of a foot or less. The collecting site was 2.5 miles
east of Orion on the Richland County side of the river. This is
the first time this species has been collected within state waters.
Length of these specimens ranged between 4%" and 6+2". Greene
(1935) captured the crystal darter from the Mississippi River at
Cassville, the only other record of this species from the region.

River darter — Percina shumardi (Girard). W(2-8), W (12-13),
W12, W15, W (20-22), M2-3, M5-6 (9 collections; 95 individuals).
This is a common darter of large rivers such as the Wisconsin and
Mississippi. It is generally captured over gravel and rock bottoms.
The river darter is probably more common on the Wisconsin River
than our collections indicate. Twenty-three specimens were cap¬
tured at station W12 and 43 at station M6.

Gilt darter — Percina evides (Jordan and Copeland). Although
this species has not been recorded from the waters within or bor¬
dering the area studied, the area lies within the range for this i
species (Trautman 1956). The gilt darter has been taken from |
the Rock River of Illinois (Forbes and Richardson 1920) and from

1966] Becker — Fishes of Southwestern Wisconsin 111

the Black and St. Croix Rivers of Wisconsin (Greene 1935).
Gerking (1945) suggested that in recent years this darter has de¬
creased greatly in Indiana,

Blackside darter — Percina maculata (Girard). W2, W12, W14-
16, M6, R5, R13, R16, R20-21, R23, C4, C7, 17-9, Ill, L16 (20
collections; 53 individuals). The blackside darter occurs within
streams and rivers of all sizes in clear to turbid water. Although
it appears frequently, it is nowhere abundant. I have taken this
darter over soft bottoms covered with organic debris, but it favors
a gravel bottom.

Slenderhead dsLYter—Percina phoxocephala (Nelson), Wl, W(2~
8), W(12-13), W12, W15, C8, 12, L7-8, L16 (11 collections; 58
individuals). On the Wisconsin River this darter ,was generally
taken in company with the river darter. They appear to be
equally common. On the Mississippi River the slenderhead is rarely
taken. This species is occasionally found in moderate to large-sized
streams but because it selects rubble and large gravel for habitat,
capture with seine is difficult. In such habitats it is easily collected
with electrofishing equipment.

Logperch — Percina caprodes (Rafinesque) . Wl, W(2-8), W2,
W4, W6, WIO, W (12-13), W12, W14-18, W20-23, Ml-6, R20, G15,
G31, G41, G43 (33 collections; 209 individuals). The logperch in¬
habits moderate to large-sized streams and rivers. It is adapted to
a wide variety of bottom types, although it prefers a hard bottom
of gravel. I have captured it most frequently in moderate currents,
although I have taken it from swift currents and from quiet
sloughs.

Bluntnose darter — Etheostoma chlorosomum (Hay). This south¬
ern darter has been collected as far north as the Root River,
Houston County, Minnesota (Eddy and Surber, 1947). Records
from the Mississippi River come from small isolated ponds between
New Albin and Minnesota slough on the lowa-Minnesota border
just across from Victory, Vernon County (Harlan & Speaker,
1956). In the University of Wisconsin-Madison Museum of Zoo-
ology are two specimens from this locale collected on August 21
and 23, 1944,

Johnny darter— nigrum Rafinesque. Wl-6, W8-17,
W19-23, Ml-6, Rl-4, R6-14, R16-20, R22-23, Cl, C3-8, CIO, C14,
G2-3, G5, G7, G9, Gll, G27, G29-47, 11-12, Ll-2, L6-16 (126 col¬
lections; 2,833 -f individuals). The Johnny darter is the most suc¬
cessful member of the family Percidae. It is found in the smallest
stream and in the largest river over a wide variety of bottom types.
In a few stations where it was not captured, it would undoubtedly
have been found with more intensive sampling.

112 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Banded darter — Etheostoma zonale (Cope). W12, W15, W(20-
22), R20, G13-14, 12, 15, 17, L2 (10 collections; 64 individuals).
The banded darter is a common darter in some waters where the
bottom is strewn with light gravel. It is generally found in dear-
water streams of medium to large size. Over a rock shelf in the
Wisconsin River 22 were captured at W15 and 14 at W12, the
largest collections made of this species during the survey. Harlan
and Speaker (1956) record this darter from the Mississippi River
north of Dubuque, Iowa.

Iowa darter — Etheostoma exile (Girard). R15. The Iowa darter
is uncommon in southwestern Wisconsin. We captured a single
individual from the Pine River at station R15. Greene (1935) re¬
cords two collections from the Wisconsin River (Richland County),
one collection from the Mississippi River at Lynxville and several
collections from the Pecatonica River in the vicinity of Argyle.

Rainbow darter — Etheostoma caeruleum Storer. W12, W14~16,
RIO, 16 (8 collections; 16 individuals). The rainbow darter, nor¬
mally an inhabitant of moderate-sized streams, is like the banded
darter, generally taken over gravel. It appears in several small
populations in the Wisconsin River, but it is rare to uncommon in
this part of the state.

Mud darter — Etheostoma asprigene (Forbes). W15, M4, Cl, 13.
Only one specimen was captured at each of the four stations. This
small species must be considered rare in the sloughs of the Wis¬
consin and Mississippi Rivers and in the lower extremities of their
tributaries. It prefers turbid water over a soft bottom.

Least darter — Etheostoma microperca Jordan and Gilbert.
Greene (1935) captured the least darter from the Pine River at
Richland Center (Richland Co.). It was not encountered in the
present survey.

Fantail darter — Etheostoma flabellare Rafinesque. W12, W15,
Rl, Rll-12, R14, R16, R18, G13-14, 12, 15, 17, L2 (47 collections;
1,016 individuals). The fantail darter is generally taken over rock
or gravel. It occurs in small to moderate-sized streams and is
especially abundant in the streams of the Pine River watershed in
Richland County. On the Wisconsin River it is common locally over
rock shelves in shallow water, where it is easily collected by electro¬
fishing. Greene (1935) has reported it from several stations on
the Mississippi below the mouth of the Wisconsin River in Grant
County.

Smallmouth bass — Micropterus dolomieui Lacepede. Wl, W (2-
8), W2, W4-6, W8-10, W12-16, W18-22, M2-3, G16, G25~26, G35,
12, 15, 17-11, LI, L4, L6, L8, L16 (52 collections; 314+ individ-

1966]

Becker — Fishes of Southwestern Wisconsin

113

uals). Fair numbers of smallmouth bass, ten inches and larger, are
found in moderate to swift current along the rocky banks of the
lower Wisconsin River. Many young-of-the-year were captured
from eddies along sand banks. On the Mississippi River the small¬
mouth is uncommon (Nord, pers. comm. — Feb. 27, 1964). Nord
writes : ‘'Much of the favored habitat . . . has been altered or
destroyed since the inception of the 9-foot navigational channel. The
distribution of this species now appears to be quite spotty.”

An interesting phenomenon was called to my attention by Truog.
Grant County is traversed from east to west by a ridge. North of
this ridge, the streams draining into the Wisconsin River contain
no ismallmouth bass, although there appears to be ample stream
gradient, rubble bottom and clear water. South of the ridge, the
streams, even small ones less than ten feet wide, contain good popu¬
lations of smallmouth bass.

Largemouth hs^^^—Micropterus salmoides (Lacepede). W4-7,
WlO-11, W13-14, W16, W18-23, Ml-6. R9, R17, R20, R23, C14,
Gl, G16, G25-26, G35, 12, 15, LIO L14, L16 (40 collections; 694 +
individuals). The largemouth bass is common in the sloughs, back¬
waters, and occasionally is trapped in the landlocked pools of the
Wisconsin and Mississippi Rivers. Hundreds of- young-of-the-year
were taken at some stations. This species occurs in moderate to
large tributaries of the large rivers and is occasionally found in
the lower extremities of small streams opening into them.

Warmouth — Chaenohryttus gulosus (Cuvier). Cl, R9. I cap¬
tured a single young-of-the-year from Gran Grae Creek on Septem¬
ber 23, 1966, The only specimen captured in the survey came from
a quiet widespread of Mill Creek below the millpond. According
to Harlan and Speaker (1956) and Nord (pers. comm. — ^Sept. 24,
1962) it is rare to uncommon in the Mississippi River. Because of
its preference for weed-filled ponds or lakes with mud bottom, the
warmouth in southwestern Wisconsin should occur more frequently
in artificial lakes and their backwaters than in streams. Truog
reports the warmouth as common.

Green sunfish — Lepomis cyanellus Rafinesque. W7, R2-3, R9,
Rll, R17, R20, R23, C2-3, C6, C8, G16-17, G20, G25-26, G31,
G37~38, G40, G43„ 12, 15, 17, 112, L14 (26 collections; 129+ indi¬
viduals). The green sunfish is a common species in moderate-sized
waters with soft bottom and sluggish current, rare to uncommon
in the Wisconsin and Mississippi Rivers.

Pumpkinseed — Lepomis gibbosus (Linnaeus). W4, Wll, W14,
W16, W20~21, W(20-22), M2-3, R9, R16-17, 16 (13 collections;
51+ individuals). The pumpkinseed is nowhere abundant in the

114 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

unglaciated region. It occurs occasionally in the Wisconsin and
Mississippi Rivers and their tributaries.

Bluegill — Lepomis macrochirus Rafinesque. Wl-6, W8, Wll,
W14-16, W18-20, W (20-22), W22-23, Ml-6, R17, R23, CIO, G36-
38, 15-6, L9-10, L12, L14, L16 (40 collections; 790+ individuals).
The bluegill is abundant in the Wisconsin and Mississippi Rivers
and is occasional in their medium and large-sized tributaries. Large
numbers were captured from the Pecatonica River and its tribu¬
taries in Lafayette County.

Orange-spotted sunfish-+^cpom2s humilis (Girard). W16, W19,
Ml-3, M5, R9, R16, 17-8, L6-8 (15 collections; 56+ individuals).
This small sunfish is found in quiet waters of moderate-sized
streams to large rivers. Nord (pers. comm. — Sept. 24, 1962) re¬
ports this species as uncommon in the Mississippi River. It appears
to be extending its range into the inland waters of these counties.
Thirty years ago Greene (1935) captured this species only from
the Mississippi River and the Galena River (Lafayette Co.) near
the Illinois line.

Rock bass — Amhloplites rupestris (Rafinesque). M3, 17, 19-11,
L6 (6 collections; 10+ individuals). The rock bass is seldom en¬
countered in the driftless region. Nord (pers. comm. — Sept. 24,
1962) considers it as uncommon in the Upper Mississippi River.
Truog reports this species as quite numerous in Pool 9 of the
Mississippi and frequently captured around old tree roots and
stumps. Interestingly enough Greene (1935), although sampling
heavily, had no capture from these counties. The records of the
present study indicate a recent adaptation to streams in the un¬
glaciated area.

Black crappie — Pomoxls nigromaculatus (LeSueur). W(2-8),
W4-5, W8, WlO-12, W14, W16-23, M2-6, R9, R17 (27 collections;
1,272+ individuals). The black crappie is abundant in the sloughs
and backwaters of the Wisconsin and Mississippi Rivers. It is
occasionally captured in the lower extremities of the larger tribu¬
taries of the Wisconsin River.

White crappie — Pomoxis annularis Rafinesque. W14, W16, W18-
22, M2-4, R9, C8, G38 (17 collections; 63+ individuals). This
species is common in the lowermost parts of the Wisconsin River
and in the Mississippi River. It is occasionally captured in their
larger tributaries. The white crappie is now found in the larger
rivers and lakes of eastcentral Wisconsin, which are parts of the
Great Lakes drainage basin (Becker, 1964). Since Greene (1935)
encountered this species only in the Mississippi drainage, it is
possible that the Fox-Wisconsin canal at Portage may in part be
responsible for this extension in range.

1966]

Becker — Fishes of Southwestern Wisconsin

115

Brook silverside — Labidesthes sicculus (Cope), Wl-6, Wll,
W14-16, W19-21, W23, M2~6. R9 (21 collections; 361+ individ¬
uals). The brook silverside is common in the Wisconsin and Missis¬
sippi Rivers. A single specimen .was captured from Mill Creek
(R9). This species is found primarily in quiet water and over a
variety of bottoms.

Freshwater drum — Aplodinotus grunniens Rafinesque. Wl,
W(2-8), W12, W16, W18--23, M2-3, M4-5, R9 (16 collections;
89+ individuals). The freshwater drum or sheepshead is abundant
in the lower Wisconsin and in the Mississippi Rivers. It is fre¬
quently caught on hook and line.

Mottled sculpin- — Cottus hairdi Girard. R6, R18, R20, R22, C3,
Gll, 14, L13, L15 (10 collections; 154+ individuals). The mottled
sculpin is common in the headwaters of many small streams of the
region, frequently the most common fish in the sample. It prefers
cool waters over heavy gravel with vegetation, often in the same
locale as trout.

Slimy sculpin — Cottus cognatus Richardson. C5. Marlin Johnson,
University of Wisconsin-Madison, collected this species from
Citron Creek at the Hwy E bridge (T9N R5W Sec. 36 NE14,
Crawford County) on October 24, 1964. Specimens were sent to
Dr. Reeve Bailey at the University of Michigan Museum to verify
identification. Although this species is common in springs and
spring runs of northeastern Iowa, this is the first record of this
species from southwestern Wisconsin. I sampled the same station
; on Sept. 24, 1966, and captured over 40 specimens. The sculpin was
the most common fish, followed by the brook trout.

Brook stickleback — Culaea inconstans (Kirtland). R2, R13, R15,
R22, C5-7, G42, 14, 112, Lll-16 (15 collections; 68+ individuals).
The brook stickleback is uncommon to common in small to
! moderate-sized streams. I have taken it from clear water but more
i frequently from turbid water that had been roiled by cattle.

: Acknowledgment

I I wish to thank Kenneth Derr, John R. Truog, James Kincannon
i and C. W. Threinen, all of the Wisconsin Conservation Department,
i who provided me with working space and materials, and fish collec-
f tions and records. Two species were recorded from the collections
: of Marlin Johnson, University of Wisconsin-Madison. My gratitude
also to Robert C. Nord, Survey Director of the Upper Mississippi
I River Conservation Committee, Bureau of Sport Eisheries and
^ Wildlife, who supplied me with recent unpublished records from
the Mississippi River. I am indebted to my sons Kenneth and Dale,

116 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

who provided yeoman service behind electrodes and seine for many
hours beyond the normal work day. The paper was read critically
by Threinen, Truog and Nord. I have used their suggestions when
feasible. If I have not followed their suggestions and the paper
suffers error, I assume full responsibility. Dr. Reeve M. Bailey,
Curator, University Museums, University of Michigan, was kind
enough to help me unravel some of the knottier problems in identifi¬
cation. Funds for carrying on the survey were supplied in part
through a research grant from the Board of Regents, Wisconsin
State Universities.

(Corrections and additions to be made in Becker, George C. 1959. Distribu¬
tion of Central Wisconsin Fishes. Trans. Wis. Acad. Sci. Arts & Letters 48:
65-102.

p. 84, 1. 2 — Insert ^‘Campostoma anom. oligolepis’* in lieu of “Campostoma
anom. pull.”

p. 89, 1. 18 & 19— Crss out ‘‘T 12; P 13, 14”

p. 90, 1. 20 — Insert “Largescale Stoneroller — Campostoma anomalum
oligolepis Hubbs & Greene” in lieu of “Central Stoneroller — Campostoma
anomalum pullum (Agassiz)”

p. 96, 1. 14 — Add “T 7” at end of line

p. 96, 1. 20— Cross out “7” in “T 6, 7”

p. 102 — Add to the list of species: ‘'Moxostoma erythrurum (Rafinesque)
T12; P 13, 14. Chrosomus neogaeus (Cope) T 1; LW 1; Eske Creek outlet
(T24N RlOE Sec. 19) Portage Co., IV:23:60. Notropis heterodon (Cope) Eske
Creek outlet (T24N RlOE Sec. 19) Portage Co., IV:23:60.”)

References

Becker, George C. 1964. The fishes of Lakes Poygan and Winnebago. Trans.

Wis, Acad. Sci. Arts & Letters. 53:29-52.

Brasch, John, James McFadden and Stanley Kmiotek. 1958. The eastern
brook trout, its life history, ecology and management. Wisconsin Conser¬
vation Dept. Publ. 226:11pp.

Christenson, Lyle M. and Lloyd L. Smith. 1965. Characteristics of fish
populations in upper Mississippi River backwater areas. U.S. Dept. In¬
terior. Fish & Wildlife Service. Circular 212. 53pp.

Coker, Robert E. 1929. Studies of common fishes of the Mississippi River at
Keokuk, Bull, Bureau of Fisheries, Document 1072:141-224.

Eddy, Samuel and Thaddeus Surber, 1947, Northern fishes, Univ. of Minne¬
sota Press, 276pp.

Forbes, Stephen A., and Robert E. Richardson, 1920. The fishes of Illinois.

Illinois Nat. Hist. Survey Bull. 3:357pp,

Gerking, Shelby D. 1945. The distribution of the fishes of Indiana. Invest.
Ind. Lakes and Streams 3(1) :137pp,

Greene, C. Willard. 1935. The distribution of Wisconsin fishes. Wisconsin
Conservation Commission. 235pp.

Harlan, James R., and Everett B. Speaker. 1956. Iowa fish and fishing. State
Conservation Commission, Des Moines, Iowa, 377pp.

Helm, Wm. T. Yellow bass in Wisconsin. Trans. Wis. Acad. Sci. Arts & Let¬
ters. 53:109-125.

1966]

Becker — Fishes of Southwestern Wisconsin

117

Martin, Lawrence, 1916. The physical geography of Wisconsin. Wisconsin
Geological and Nat. Hist. Survey Bull. 36:549pp,

Nybakken, James W, 1961, Analysis of the sympatric occurrence of two
subspecies of the cyprinid fish Campostoma anomalum (Rafinesque) in
Wisconsin. M.S. Thesis. Univ, of Wisconsin, Madison. 35pp.

PoFF, Ronald J., and C. W. Threinen. 1965. Surface water resources of
Columbia County. Lake & Stream Classification Project. Wisconsin Con¬
servation Dept, 55pp,

TrautMan, Milton B. 1957. The fishes of Ohio. Ohio State Univ. Press. 683pp.

U.S. Army Engineer Division. 1963. Navigation charts middle and upper
Mississippi River. North Central Corps of Engineers, Chicago, Ill. 61
charts.

Whitson, A. R. 1927. Soils of Wisconsin. Wisconsin Geological and Nat. Hist.
Survey Bull, 68:270pp.

■r

. X

t

f

1

THE SEASONAL DISTRIBUTION OF FISHES
IN VERMILION BAY, LOUISIANA

Carroll R. Norden
Department of Zoology
University of Wisconsin-Milwaukee

Vermilion Bay is one of several shallow estuaries along the
north shore of the Gulf of Mexico. The fish fauna of a number of
these bays, from Texas to Florida, have been investigated by other
workers (Gunter 1938a, 1938b, 1945; Suttkus et al 1953-54;
Reid 1955a, 1955b; Simmons and Hoese 1959; Arnold et al. 1960).
Except for a study of nearby White and Grand Lakes by Gunter
and Shell (1958), Vermilion Bay has received little attention from
ichthyologists and fishery biologists.

For a three-year period, monthly trips were made to Vermilion
Bay in order to assess the fish fauna inhabiting the area. A primary
objective of the study was to obtain an inventory of the fishes in
the bay and to determine their seasonal movements. A second
objective was to interpret the seasonal changes of the fish fauna
in relation to hydrological conditions.

Materials and Methods

Fish collections were made each month from October, 1960, to
August, 1963, Samples were collected during 32 of the 35 months
and all months were sampled at least twice during the study
period.

A variety of collecting gear was used in order to sample as wide
a size range as possible of the fish population. The gear consisted
of three 125-foot gill nets with II/2- and 2-inch mesh, a 300-foot
trammel net with 3-inch mesh, a 16-foot shrimp otter trawl, 10-,
25-, and 50-foot nylon minnow seines, plankton nets with number
6 and 12 meshes, a 6-foot beam trawl, developed by the Galveston
Laboratory of the U. S, Bureau of Commercial Fisheries to sample
post-larval shrimp, dip nets, trot lines, and hook and line.

Most of the collecting was done in three general areas, around
Southwest Point, Redfish Point, and Cypremort Point. On Redfish
Point the University of Southwestern Louisiana has a small labo¬
ratory which was used as a base for operations, and the greatest
amount of sampling was done in that area.

119

120 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Generally, the fishes were collected over a two-day period,
with one or more water samples taken and salinity determined
by silver nitrate titration (Marvin et at. 1960), reported as parts
per thousand of salinity. From the same water samples, values for
pH were obtained by using a Beckman Model G pH meter. Tem¬
peratures of air and water were procured by standard centigrade
thermometer. The Secchi disc was used as an index of turbidity.

The large specimens taken with gill and trammel nets were
weighed, measured, sexed, and discarded. Total length measure¬
ments were made with a measuring board for the larger specimens,
dividers and a millimeter ruler for the smaller specimens. Total
length is to the nearest millimeter. The common names given follow
Bailey cf (x/. (1960).

In some instances, particularly in collections made with the
trawl or seines, certain species were so abundant that not all
specimens could be preserved and returned to the laboratory.
In such cases, random samples of the abundant species were taken
along with the rare specimens specifically selected from the catch,
A large number of the fishes were preserved in ten percent formalin
and are stored at the University of Southwestern Louisiana or at
the University of Wisconsin-Milwaukee.

Certain limitations are inherent in the study, primarily because
of weather conditions and a shortage of personnel. It was not
always possible to sample all areas of Vermilion Bay with equal
frequency. Over the three-year period, however, each month was
sampled quite consistently with gear that was selective for the
different size-age groups.

Acknowledgments

Special thanks are extended to Dr. Lewis T. Graham, head of
the Department of Biology at the University of Southwestern
Louisiana, and to the following students who assisted in making
the collections: Donald Burney, Lewis T. Graham, Jr., William
Mason, Anthony Romano, Samuel Riche, David Williams, and
especially to Semmes Lynch, who made available some of the most
recent data. Mr. Kenneth Lantz of the Louisiana Wild Life and
Fisheries Commission also contributed data and assistance. Dr.
Reeve M. Bailey, Curator of Fishes, University of Michigan, veri¬
fied the identification of several species, Drs. Rezneat M, Darnell,
Marquette University, and Gordon Gunter, Gulf Coast Research
Laboratory, read the manuscript and offered many helpful sug¬
gestions. The help of all these people is greatly appreciated. The
work during the 1963 season was subsidized in part by Contract
No. 14-17-0002-48 from the U. S. Bureau of Commercial Fisheries.

1966] Norden — Seasonal Distribution of Fishes 121

Description of Vermilion Bay

Vermilion Bay (Fig. 1) is located at about 92 °W., 29° 40' N.
It is a shallow body of water with a surface area of approximately
208 square miles and an average depth of about five feet. Except
for Southwest Pass, its greatest depth is ten feet.

It is surrounded by marshland on three sides, to the north, east
and west. The marsh consists of extensive areas of typical salt-
marsh vegetation, such as white cord grass, Spartina patens, big
cord grass, S, cynosuroides, black rush, Juncus roemerianus, and
three-cornered grass, Scirpus olneyi. Remiains of this marsh vege¬
tation have resulted in the deposition of humic materials along the
shores and at the bottom of Vermilion Bay.

Numerous small bayous empty into the bay. The largest is the
Vermilion River, entering in the northwest corner. The Intracoastal
Waterway borders the north and eastern margins of the bay. To
the south. Vermilion Bay is connected to the Gulf of Mexico at
two points. The western channel is narrow, over 90 feet deep, and
lies between Southwest Point and Marsh Island. During tidal ex¬
change, a strong current of water flows through Southwest Pass.
The eastern channel. East and West Cote Blanche Bay, is wider
and shallower and lies between Marsh Island and Cypremort Point.
Thus Marsh Island, with an area of approximately 125 square miles
(Orton 1959), partially isolates Vermilion Bay from the Gulf of
Mexico.

Hydrography

The water temperatures of Vermilion Bay tend to fluctuate
rather closely with atmospheric temperatures. The monthly range
of temperatures at the time fish collections were made is indicated
in Fig, 2. It will be noted (Fig. 2) that for five months (May
through September) the waters of Vermilion Bay were always
above 20 °C. and that the cold months, when water temperatures
were less than 10 °C., were December, January, and February. The
minimum temperature recorded was 6°C. on December 17, 1960;
the maximum was 35° on August 6, 1963. At Redfish Point (Table
1) the monthly averages of surface water temperatures for the
three-year period varied between a low of 9°C, in January to a
high of 32.7 °C. in August. These water temperatures are similar
to those reported for Lake Ponchartrain, Louisiana (Darnell 1958) ,
and East Lagoon, Galveston, Texas (Arnold et al, 1960).

Vermilion Bay waters generally exhibit lower salinities than
those reported from the Texas estuaries (Gunter 1945; Reid 1955a;
Simmons and Hoese 1959) or from Tampa Bay, Florida (Springer
and Woodburn 1960). This is in part because of the heavy annual

122 Wisconsin Academy of Sciences, Arts and Letters

[Vol. 55

Figure 1. Vermilion Bay, Louisiana.

1966]

Norden — Seasonal Distribution of Fishes

123

35

30
d 25

o

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Figure 2. Monthly maximum-minimum range of temperatures at which fishes
were collected in Vermilion Bay, Louisiana.

rainfall of this section of Louisiana, the greater amounts of fresh
water draining into Vermilion Bay, and the influence of the
Mississippi-Atchafalaya waters moving from east to west along the
Louisiana coast and blocking out the more saline waters from the
Gulf of Mexico. The monthly averages measured at Redfiish Point
ranged from a low of 2. in June to a high of 9.%° in July (Table
1). This is not much different from Lake Ponchartrain, in which
Darnell (1962a) reports salinity varying from 3, to 8.%°,

The range of salinities recorded at the time fish collections were
made is shown in Fig. 3. The minimum salinity was 0.8%° on
April 30, 1963, whereas the maximum during the entire study was
32.8%° on July 13, 1962. This high salinity occurred during a pro¬
longed period of isouthwest wind from July 10, to July 23, 1962,
which blew the more saline waters from the Gulf of Mexico into
Vermilion Bay.

Little variation occurs in the pH of Vermilion Bay waters. The
monthly average at Redfish Point (Table 1) varied between a
pH of 7.2 in December to 8.2 in October. The pH probably exerts
little influence on the seasonal movements of fishes.

124

Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Figure 3. Monthly maximum-minimum range of salinities at which fishes were
collected in Vermilion Bay, Louisiana.

The bottom deposits of Vermilion Bay consist mostly of fine silt
and humic materials, with some sand and an occasional shell reef.
Like those of Lake Pontchartrain (Darnell 1958, 1961), these bot¬
tom deposits are continually being disturbed by wind action. The
waters of Vermilion Bay are highly turbid and the average monthly
Secchi disc readings at Redfish Point (Table 1) ranged from a low
of 19 centimeters in October to a high of 89 centimeters in
September.

The Fishes of Vermilion Bay

Eighty species of fish with representatives in 41 families (Table
2) were collected during this investigation. In addition. Megalops
atlantica was seen in the bay but not collected. Later, in the fall of
1963, Eleotris pisonis and Porichthys porosissimus were collected
by Mr, Semmes Lynch. Ictiohus bubalus was collected by Mr. Ken¬
neth Lantz (Lantz 1963) in 1962. Thus 84 species are reported
from Vermilion Bay.

As has been pointed out by Gunter (1945, 1956a, 1956b) and
others, the fish fauna of an estuary is essentially a marine fauna.
Vermilion Bay is typical, because few freshwater fishes invade wa¬
ters which are even moderately saline. Eight species of freshwater
fishes (Lepisosteus oculatus, L. platostomous, Ictiobus bubalus,
Ictalurus furcatus, Roccus mississippiensis, Lepomis macrochirus,
L. punctatus, and Aplodinotus grunniens) were collected. Seven of
the eight species would be considered rare in the bay. The single
important exception is Ictalurus furcatus, which is particularly
abundant during the winter months. This is not surprising as it has
been reported repeatedly from estuarine waters (Gunter 1945;
Darnell 1958; Gunter and Shell 1958), and Darnell (1962b) has
classified this species as a facultative invader of brackish waters.

1966]

Norden — Seasonal Distribution of Fishes

125

Table 1. Monthly Averages of Hydrologic Data at Redfish Point,
Vermilion Bay, Louisiana. (Oct. 1960-Aug. 1963)

Month

Water

Temperature

°C

Salinity

PPT

Secchi Disc
Centimeters

pH

January .

9.0

6.47

26.2

7.3

February .

17.3

6.73

57.8

7.7

March .

18. 1

5.51

44.5

7.8

April .

22.0

3.73

24.0

7.9

May .

24.9

3.71

25.9

7.9

June .

29.1

2.22

57.6

7.6

July .

30. 1

9. 15

64.9

7.3

August .

32.7

7.25

75.0

7.4

September .

25.2

6.55

89.0

7.6

October .

19.5

6.74

19.0

8.2

November .

19. 1

7.42

37.0

7.4

December .

11.4

4.84

26.5

7.2

Salinity may influence the age groups of a species which enter
an estuary (Gunter 1945, 1956a), and low salinity gradients may
keep out certain species altogether. Thus the threadfin, Polydac-
tylus octonemus, reported as being very abundant by Gunter
(1938b, 1945) in Barataria Bay, Louisiana and Aransas Bay,
Texas, was not collected within Vermilion Bay. The same may be
said for the butterfish, Poronotus triacanthus, the moonfish. Vomer
setapinnis, and the star drum, Stellifer lanceolatus. Gunter and
Shell (1958) reported that they collected three specimens of Poly-
dactylus octonemus in Little Bay, which is an extension of Ver¬
milion Bay. Therefore, it probably should be added to the check¬
list (Table 2) as an occasional straggler in Vermilion Bay,

Only six species, Prishls pectinatus, Synodus foetens, Chloro-
scombrus chrysurus, Selene vomer, Trichiurus lepturus, and Pepri-
lus paru (Table 3), gave evidence of preferring highly saline wa¬
ters. All six species were collected during the period of highest
salinities (26.9 to 32.8%°) from July 10 to July 23, 1962,

As Table 3 indicates, a greater variety of marine and brackish
water species is present during the summer than during the winter
months. The fish fauna of Vermilion Bay was most diversified dur¬
ing July, when 60 species were collected, whereas only 13 species
were collected during January and 18 species in February.

Adults of several marine species are common in Vermilion Bay
in the summer (Table 3). Among the more conspicuous of the
summer fish fauna are Carcharhinus leucas, Dasyatis sahina,
Bagre marinus, and Galeichthys fells. These species are less com-

Table 2. Relative Abundance of Fishes Collected in Vermilion Bay, Louisiana. (Oct. 1960-Aug. 1963)

126 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

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Table 2. Relative Abundance of Fishes Collected in Vermilion Bay, Louisiana.

(Oct. 1960- Aug. 1963) — Continued

1966]

Nor den — Seasonal Distribution of Fishes

127

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Table 2. Relative Abundance of Fishes Collected in Vermilion Bay, Louisiana.

(Oct, 1960-Aug. 1963) — Continued

128 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

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Seasonal Distribution of Fishes in Vermilion Bay, Louisiana. (Oct. 1960-Aug. 1963)

1966]

Norden — Seasonal Distribution of Fishes

129

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Table 3. Seasonal Distribution of Fishes in Vermilion Bay, Louisiana. (Oct. 1960-Aug. 1963) — Continued

130 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Dec.

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Table 3. Seasonal Distribution of Fishes in Vermilion Bay, Louisiana. (Oct. 1960-Aug. 1963) — Continued

1966]

Norden — Seasonal Distribution of Fishes

Dec.

1

i ! ! !

1

1

1

!

Nov.

!

— GO

i 1 r !

” 1

1

1

1

1

^ vrN t\ t\ ' — '

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1 1

1

Oct.

!

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c/)

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1

114

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1 1

1

GO GO sD 0 — — ^

GO GO GO ^ ”11

1

1

July

GC

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1 . ^ 1

1

1

"O'vDO' r^iirNC^OX'

O' <^i 1 u". — — ' 1

— 1 1

1 1

i

!

1

June

2*

1

7

I

188

1!

1

1

O' sD 0 itn

0 O) GO 1 GO 1 1

^ 1 11

1 1 1

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May

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Apr.

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^ 1 III

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Mar.

sO

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1 *

ro ^

^ 0 —

^1 ! ! !

1

Feb.

Mll‘1

1

^ ir\

1

1

Jan.

MM*!

1

1 1 1 ^ ^ 1 1

1

Species

c

S

C

i

c

i

1

c

u

LO piCiUriL^ .

Gobioides broussonneti .

Gobionellus hastatus. . .

Gobionellus shufeldti .

Gobionellus stigmaticus .

Gobiosoma bosci . .

- 1..-,

• !

. !

• 3 ;

i'2 J
n -o ,

<0 'C '
2 <

^ li

i

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yus u .

Mugil cephalus .

Membras martinica .

Menidia beryllina .

Citharichthvs spilopterus .

Paralichthys lethostigma .

Trinectes maculatus .

Symphurus plagiusa .

Gobiesox strumosus .

Sphaeroides nephelus .

* :3

! ^

• -2
. ^

. 0

3 -Q.

^s.

3 J

5 .2

:)Q,

1

t Observed.

() Collected by K. Lantz (1963).

* Collected by Semmes Lynch (1963).

132 Wisconsin Academy of Science, Arts and Letters [Vol. 55

mon in cooler waters and tend to migrate out of the bay as the
water gets colder (Gunter 1945; Simmons and Hoese 1959; Dar¬
nell 1961). The adults of only three species, Dorosoma cepedianum,
Ictalurus furcatus, and Mugil cephalus could be considered very
abundant at cold temperatures.

Adults of a number of species which are fished in Vermilion
Bay are rather sparsely represented in the collections, particularly
Cynoscion nehulosus, Pogonias cromis, Sciaenops ocellata, and
Paralichthys lethostigma. It has been suggested (Gunter 1945;
Simmons and Hoese 1959) that these fishes tend to elude the collect¬
ing gear rather effectively.

Of the 84 species recorded from Vermilion Bay, 26 species or
nearly 31 percent were collected at all seasons of the year. Only
eight species or 9.5 percent were prsent every month of the year.
In contrast to these data, 26 species of marine fishes and 6 species
of freshwater fishes (Table 3), about 38 percent of the species
complex, are represented as occasional stragglers and comprise only
a small portion of the fish population of the bay.

The young of marine species tend to be more abundant in an
estuarine environment, and a number of investigators (Gunter
1957, 1961) have noted that an estuary serves as a nursery for
many marine species. The larvae and young of one or more marine
species may be collected in Vermilion Bay during every month of
the year ; a greater variety, however, is present during the spring
and summer (Table 4).

Larval menhaden begin appearing in November (Table 4) and
continue into April, which is consistent with the findings of Suttkus
and Sundararaj (1961). These larvae have been assigned to
Brevoortia patronus because only juveniles and adults of that
species have been collected from Vermilion Bay. The finescale
menhaden, B. gunteri, however, is present in the nearby Gulf
(Suttkus 1958 ; Christmas and Gunter 1960) , and some of the larvae
may belong to that species. The same may be said for the larval
anchovies (Table 4), which appear from April to August. Some
of the smaller individuals may be Anchoa hepsetus, although this
species is rare in Vermilion Bay, whereas A. mitchilli is very
abundant.

Many of the larvae were cleared and stained for identification
purposes and the larval sciaenids were identified by using the cri¬
teria of Hildebrand and Cable (1930, 1934). These data for
Bairdiella chrysura, Leiostomus xanthurus, Menticirrhus ameri-
canus, and Micropogon undulatus are consistent with the findings
of Hildebrand and Cable (1930, 1934) and Suttkus (1954). The
appearance of Cynoscion nehulosus in June, July, and August is
not surprising in view of the work of Sundararaj and Suttkus

Table 4, Seasonal Appearance of Larval Marine Fishes in Vermilion Bay, Lousiana.

1966]

Norden — Seasonal Distribution of Fishes

lx.

1 1 7 1 11

1 1 i 1 T 1

1 7 1 1 1 1 1 1 1

1 1 1 1 1

1 1 1 1 r-a 1

1 00 1 1 1 1 1 1 1 1

I I

I I

rx. — <

X I

1 lx.

A

July

200-2 7 ^
25

10-38

37

35-37

X

6-46

17-29

10-13

12-25

8-10

Mil

June

75-78

10-29

III

6-32

12-23

11-12

18-19

• Jn 1

15-21

May

13-25

59-63

15-25

15-44

10-12

5-11

Ml!

Apr.

30-40

12- 57

13- 33

43-61

14-34

IX

13-51

Mar.

38-43

17-49

42-65

13- 45

14- 18

15- 17
14-37

13-20

11- 30

12- 17

CXD

00

vrs

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1 1 1 1

1 °° 1

1 1 1 1 1 ^ 1

1 1 1

1 1 1 1

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c u
3

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Q 3

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^ ^ 3

CO 2

?5 9

•<5.5

3dt^

Total lengths in millimeters.

X Males carrying eggs and young.

134 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

(1962). The appearance of larvae in November and December
(Table 4) may indicate two spawning peaks, as suggested by
Gunter (1945).

Only a few larvae corresponding to the descriptions of Gobio-
nellus and Microgobius (Hildebrand and Cable 1938) were col¬
lected; larvae with characteristics of Gobiosoma bosci, however,
were abundant during May, June and July.

Many of the young, metamorphosed Myrophis punctatus were
captured with a dip-net and nightlight as they swam near the
surface. Springer and Woodburn (1960) captured young worm eels
in a similar manner in Tampa Bay during April. As pointed out
by Gehringer (1959), the leptocephalus of Flops saurus shorten
before they metamorphose. Thus, the specimen collected in July
was more advanced than the earlier specimens. Except for this
single exception, however, their seasonal appearance agrees with
that of Arnold et al. (1960) for Galveston, Texas.

Males of Galeichthys felis and Syngnathus scovelli were collected
carrying eggs and young, the former in July and the latter in both
July and August. During both June and July pregnant Dasyatis
sabina were caught in trawl hauls and the young born alive in the
boat.

Evidence from this work as well as from work of others (Hilde¬
brand and Cable 1930, 1934, 1938; Gunter 1938b, 1945) indicates
that few species actually spend their entire life cycles within an
estuary. Examination of sexually mature adults, observations of
breeding colors and behavior, and the fact that larvae, young and
adults were collected at all seasons of the year suggest that five
species (Cyprinodon variegatus, Fundulus grandis, Gobiosoma
bosci, Menidia beryllina, and Gobiesox strumosus) remain to spawn
and complete their life cycles within the confines of Vermilion Bay.
In addition, small populations of Adinia xenica, Lucania parva,
Gambusia affinis, Mollienesia latipinna, and Gobionellus shufeldti
may be self-sustaining in scattered areas along the edge of the bay.

In comparing the total number of individuals collected, it was
found that three species (Table 2) contributed over 75 percent of
the total. A large portion of these consisted of small individuals,
less than 100 millimeters in total length, and therefore this is an
estimate of numbers, not of weight.

Gunter (1945) in his work in Capano Bay and Aransas Bay,
estimated that Micropogon undulatus, Anchoa mitchilli, and
Menidia beryllina comprised the largest species mass in that area
and Suttkus et al. (1953-54) in his work on Lake Ponchartrain,
reported that Micropogon undulatus and Anchoa mitchilli com¬
prised 80 percent of the trawl catches and that Micropogon un¬
dulatus and Brevoortia patronus accounted for 47 percent of the

1966] Norden — Seasonal Distribution of Fishes 135

seine catches. Further estimates from Louisiana waters, Barataria
Bay, Grand Lake, and White Lake (Gunter 1938b, Gunter and
Shell 1958), indicate that the Atlantic croaker, bay anchovy, and
largescale menhaden were the most abundant species in those areas.
These three, plus an additional eight species (Table 2), Fundulus
grandis, Cynoscion arenarius, Leiostomus xanthurus, Gobiosoma
bosci, Mugil cephalus, Membras martinica, Menidia beryllina and
Trinectes maculatus, made up more than one percent of the total
catch by number. Four of these eight species, Cynoscion arenarius,
Leiostomus xanthurus, Menidia beryllina, and Trinectes maculatus,
comprised between one and ten percent of the catch in Lake
Ponchartrain (Suttkus et al. 1953-54). The eleven species named
contributed nearly 95 percent of the total individuals collected from
Vermilion Bay during the three-year period. The remaining 73
species taken made up only five percent of the total catch.

Summary

Some 70,000 specimens of fishes were collected during the inves¬
tigation. Eighty-four species with representatives in forty-one
families comprised the fish fauna of Vermilion Bay, Louisiana.

Three species, Anchoa mitchilli, Micropogon undulatus, and
Brevoortia patronus, comprised over 75 percent of the total indi¬
viduals collected. These three, plus an additional eight species,
account for nearly 95 percent of the total number collected. The
remaining 73 species contributed about five percent of the total.

Vermilion Bay is a nursery ground for marine fishes, with the
larvae of 22 (Table 4) and the young of 16 other species (Table 3)
appearing at various seasons of the year.

Only eight species of freshwater fishes were taken in Vermilion
Bay and of these eight only Ictalurus furcatus appeared in any
numbers.

Salinity in Vermilion Bay is rather low, less than 10%° during
most of the year, which may tend to reduce the numbers of some
species and keep others out of the bay altogether. Six species indi¬
cated a preference for the highly saline waters recorded in July
1962, although subsequently (in October, 1963) the harvest fish and
the lizardfish were collected by Mr. Semmes Lynch in waters of
7.1%° salinity.

Water temperature, rather than salinity, appears to exert a
greater influence on the seasonal movements of fishes in and out
of Vermilion Bay. Twenty-six species were present at all seasons
of the year, but only eight were collected every month of the year.
The adults of three species were common at cold temperatures,
whereas the adults of 16 species were common at high tempera¬
tures.

136 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

This study and evidence from other investigations, indicates that
five and probably not more than ten species complete their entire
life cycles within the confines of Vermilion Bay.

Literature Cited

Arnold, E. L., Jr., R. S. Wheeler, and K. N. Baxter. 1960. Observations on
fishes and other biota of East Lagoon, Galveston Island. U. S. Fish and
Wildl. Serv., Spec. Sci. Kept., 344:1-30.

Bailey, R. M., E. A. Lachner, C. C. Lindsey, C. R. Robins, P. M. Rodel, W. B,
Scott, and L. P. Woods. 1960. A list of common and scientific names of
fishes from the United States and Canada. Am. Fish. Soc., Spec. Pub. No.
2:1-102.

Christmas, J. Y., and G. Gunter. 1960. Distribution of menhaden, genus Bre-
voortia, in the Gulf of Mexico. Trans. Am. Fish. Soc., 89(4) :338-343.

Darnell, R. M, 1958. Food habits of fishes and larger invertebrates of Lake
Pontchartrain, Louisiana, an estuarine community. Pub. Inst. Mar. Sci.,
5:353-416.

- . 1981. Trophic spectrum of an estuarine community, based on studies

of Lake Pontchartrain, Louisiana. FcoL, 42(3) :553-568.

- . 1962a. Ecological history of Lake Pontchartrain, an estuarine commu¬
nity. Am. Mid. Nat., 68(2) : 434-444.

- . 1962b. Fishes of the Rio Tamesi and related coastal lagoons in east-

central Mexico. Pub. Inst. Mar. Sci., 8:299-365.

Gehringer, J. W. 1959. Early development and metamorphosis of the ten-
pounder Flops saurus Linnaeus. U. S. Fish and Wildl. Serv., Fish Bull.
155., 59:619-647.

Gunter, Gordon. 1938a. The relative numbers of species of marine fish on the
Louisiana coast. Am. Nat., 72:77-83.

- . 1938b. Seasonal variations in abundance of certain estuarine and

marine fishes in Louisiana, with particular reference to life histories. Ecol.
Monographs, 8 (3) : 313-346.

- . 1945. Studies on marine fishes of Texas. Pub. Inst. Mar. Sci., 1(1) :1-

190.

- , 1956a. A revised list of euryhalin fishes of North and Middle America.

Am. Mid. Nat., 56(2) :345-354.

- . 1956b. Some relations of faunal distributions to salinity in estuarine

waters. EcoL, 37(3) :616-619.

- . 1957. Predominance of the young among marine fishes found in fresh

water. Copeia, 1:13-16.

- . 1961. Some relations of estuarine organisms to salinity. Limnol.

Oceanog., 6(2) : 182-190.

- and W. E. Shell, Jr. 1958, A study of an estuarine area with water-

level control in the Louisiana marsh Proc. La. Acad, Sci., 21:5-34.

Hildebrand, S. F, and L. E. Cable. 1930, Development and life history of
fourteen teleostean fishes at Beaufort, N. C. Bull. U. S. Bur. Fish., Fish.

Doc. No. 1093. 46:383-488.

- and - . 1934. Reproduction and development of whitings or king-

fishes, drums, spot, croaker, and weakfishes or sea trouts, family Sciaeni-
dae, of the Atlantic Coast of the United States. Bull. U. S. Bur. Fish., Bull.
No. 16. 48:41-117.

- and - . 1938. Further notes on the development and life history

of some teleosts at Beaufort, N. C. Bull. U. S. Bur. Fish., Bull. No. 24.
48:505-642.

1966]

Nor den — Seasonal Distribution of Fishes

137

Lantz, K. E. 1963. Cypremort Cove, Iberia Parish. La. Wild Life and Fish.
Comm., 10th Biennial Kept., 1962-1963:127-128.

Marvin, K. T., Z. P. Zein-Eldin, B. Z. May, and L. M. Lansford. 1960. Chem¬
ical analyses of marine and estuarine waters used by the Galveston Bio¬
logical Laboratory. U. S. Fish and Wildl. Serv., Spec. Sci. Kept., 349:1-14.

Orton, E. W. 1959. A geological study of Marsh Island, Iberia Parish, Louisi¬
ana. La. Wild Life and Fish Comm., Tech, Bull., 1-28.

Reid, G. K., Jr. 1955a. A summer study of the biology and ecology of East Bay,
Texas. Pt. I. Texas J. Sci., 7(3) :316-343.

- . 1955b. A summer study of the biology and ecology of East Bay, Texas.

Pt. II. Texas J. Sci., 7(4) :430-453.

Simmons, E. G.- and H. D. Hoese. 1959. Studies on the hydrography and fish
migrations of Cedar Bayou, a natural tidal inlet on the central Texas
Coast. Pub. Inst. Mar. Sci., 6:56-80.

Springer, V. G. and K. D. Woodburn. 1960. An ecological study of the fishes
of the Tampa Bay area. Fla. State Board Conserv., Prof, Pap,, Series No,
1:1-104.

SuNDARARAJ, B. I. and R. D. Suttkus, 1962. Fecundity of the spotted seatrout,
Cynoscion nebulosus (Cuvier), from Lake Borgne area, Louisiana. Trans.
Am. Fish. Soc., 91(1) :84-88.

Suttkus, R. D. 1954, Seasonal movements and growth of the Atlantic croaker
(Micropogon undulatus) along the east Louisiana Coast. Proc. Gulf and
Carib. Fish. Inst., 7th Ann. Sess., 1-7.

- . 1958. Distribution of menhaden in the Gulf of Mexico. Trans. 23rd No.

Am. Wildl. Conf., 401-410.

- , R, M. Darnell, and J. H. Darnell, 1953-1954. Biological study of

Lake Pontchartrain. Zool. Dept., Tulane Univ., Ann. Rept., 1953-1954,
1-59.

- and B. I. Sundararaj. 1961. Fecundity and reproduction in the large-

scale menhaden, Brevoortia pair onus Goode. Tulane Stud. ZooL, 8(6) :177-

182.

/

EVOLUTIONARY TRENDS OF THE MUCROSPIRIFER

LeRoy R. Peters'^

Abstract

The Brachiopods Mucrospirifer mucronatus and Mucrospirifer
thedfordensiis from the Silica formation of the Devonian of Ohio
were studied and were shown to be in the process of evolving from
a compact form to a long narrow form and to two flattened vari¬
eties of the original form. Through this study of the Mucrospirifer
of the Devonian of Ohio, the Devonian of Wisconsin may be more
closely correlated with the Devonian of Ohio after a similar study
is made of the Devonian of Wisconsin.

In 1964 John R. Tilman of Ohio Wesleyan University studied the
Mucrospirifer of Ohio and on the basis of considerable overlap in
variation, reclassified the Mucrospirifer into two species and de¬
scribed a third new species. Over 500 specimens were studied, all
of them collected from the North Quarry of the Medusa Portland
Cement Company near Sylvania in Lucas County, Ohio, SE Sec.
7, T. 9 S., R. 6 E. The graphs in this paper were plotted with only
a few points in order to show trends more clearly.

The present study is based on the external features because . .
the shape and the general proportions of the whole shell seem to
be characters of the greatest value in distinguishing between
species” (Tillman 1964). The characteristics studied here are the
width of the fold, length, thickness, and the width of the shell.

In the past, ratios based on the width of the shell have been of
doubtful value since both cardinal extremities are rarely preserved.
In this study an effort was made to collect specimens that had re¬
tained at least one cardinal extremity. Because Brachiopods are
bilaterally symmetrical, on specimens that had only one cardinal
extremity the measurement was taken from the center of the pedi~
cal opening to the end of the existing cardinal extremity and
doubled without loss of accuracy.

* The author at the time this paper was written was a senior majoring- in geology
at the University of Wisconsin-Milwaukee. At the present time he is a second lieu¬
tenant in the United States Army.

139

140 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Figure 1. Features of the Mucrospirifer used in this study.

Since the size of each specimen varied, it was necessary to find
a way to compare them. This was accomplished by setting the shell
width equal to one and comparing it with the other features in a
simple ratio similar to the method used in crystallography to com¬
pare lengths of crystal axes.

Example :

Length X
Width ~ 1

X = Ratio of Length to Width

li _ A

55 ~ 1

X = 0.25

In figures 2 and 3, three main divisions are present, each extend¬
ing from a central area. This area represents the ancestral form
Mucrospirifer mucronatus variety I found in the lower portions
this formation. Extending from this area are Mucrospirifer mucro¬
natus variety II, Mucrospirifer thedfordensis variety I, and Mucro-

1966] Peters — Trends of the Mucrospirifer 141

Figure 2.

spirifer thedfordensis variety II, which are found in the upper
portions of the formation. The farther a specimen lies from the
central area the higher in the formation it is found,

Mucrospirifer mucronatus variety I is a compact form which
widens while retaining its thickness/length ratio to form M. muro-
natus variety II, M, mucronatus variety I reduces the thickness/
length ratio by about one half , while retaining approximately the
width/length ratio to form M. thedfordensis variety I, M. mucro¬
natus variety I reduces the thickness/length ratio by about one half
while increasing the width/length ratio to form M, thedfordensis
variety II,

These evolutionary trends continue until the thickness/width
ratio reaches about 20% in the three advanced forms. This appears
to be the most effective shape for this environment.

A later study will be made of the relationship of lithology to the
evolutionary trends, because there was a change of environment.
After these studies are finished, similar studies will be made on the
Mucrospirifer of Wisconsin with the hope of more closely correlat¬
ing the Devonian of Ohio with that of Wisconsin,

142 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Figure 3.

1966]

Peters — Trends of the Mucrospirifer

143

Figuee 4. The changes in the shape of the Mucrospirifer as indicated by this
study.

References Cited

Tillman, John R. “Variation in Species of Mucrospirifer from Middle Devo¬
nian Rocks of Michigan, Ontario, and Ohio,’’ Journal of Paleontology, V,
38, No. 5, (1964) pps. 952-964.

7.

INTERACTION OF PHAGE T1 WITH STRAINS OF
ESCHERICHIA COLI

Marvin D. Whitehead and J , Roger Christensen^

In an effort to find strains of Escherichia coli for which page T1
exhibits the phenomenon of host-controlled variation, we treated
for susceptibility to the phage 290 cultures of this organism iso¬
lated by the Bacteriological Laboratories of Strong Memorial Hos¬
pital, Rochester, N. Y.

Each culture was streaked on a nutrient agar plate and the streak
spotted with a loopful of T1 lysate having a titer of 10^^ per mb
The lysate designated as Tl-B had been made on E. coli B. Three
of the cultures designated as Wh24, Wh57 and Wh96 showed lysis,
and when further tested, each of these strains showed different
responses to Tl.

Phage Tl had a plating efficiency of 10“^ to 10“^ on strain Wh2Ji.
Phage was isolated from these plaques, plated again on Wh24 and
reisolated.

This reisolated phage, designated as Tl-24, gave about one-tenth
as many plaques on Wh24 on E. coli B. The phage isolated from the
plaques of T1--24 when plated on B had reverted to possess prop¬
erties of Tl-B. This is considered to be a typical case of host-con¬
trolled variation.

Phage Tl produced only tiny, dim plaques on strain Wh24 by
the two-layer technique. More distinct plaques were obtained by
spreading the phage and bacteria on the surface of nutient agar
plates, but the plaques formed were still smaller than those obtained
by the same technique on strain B.

Strain Wh24 fails to absorb sufficient amounts of Tl from a
liquid suspension. Judging from the size of the plaques, conditions
in soft agar are probably less conducive for attachment than those
on the surface of a solid medium. Variations in the ionic compo¬
sition of the medium, age of the cells, or the presence of nutrient
broth were all without effect on phage absorption.

* This research was supported in part by the Woodward Fund for Medical Research
and the Georgia Southern College Research Fund. The Senior author was the recipient
of an N.S.F. Summer Research Participation Fellowship at the Dept, of Microbiology,
School of Medicine and Dentistry, University of Rochester. Dr. Whitehead is Pro¬
fessor of Mycology, Georgia Southern College, Statesboro, Georgia, and Dr. Christensen
is Professor of Microbiology, Univ. of Rochester, Rochester, N. Y.

145

146 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Strain Wh24, because of failure of efficient attachment, was con¬
sidered of little value on the investigation of host-controlled vari¬
ation.

Because strain Wh57 showed a plating efficiency approximately
equal to that of B with phage Tl-B, phage Tl-24, and Tl-57, it was
not investigated further.

A loopful of Tl-B at 10^*^ per ml. gave a clear spot on strain
Wh96, but a loopful of Tl-B at 10® had no visible effect. With inter¬
mediate concentrations, there was a gradual transition in the clar¬
ity of the infection spot, but distinct plaques were never observed.

Phage Tl-B attached well to Wh96, but the infected cells failed
to produce plaques, either on B or on Wh96. A plot of the surviv¬
ing cells versus the multiplicity of phage irreversibly attached,
shows that up to a multiplicity of approximately 10 phage per cell,
about one page in five is effective as a cell killer. At higher multi¬
plicities, the apparent killing efficiency decreases, but by using high
multiplicities more than 99.9% of the cells are killed. It is apparent
that some cells are killed even by a single infection, some resist
moderate multiplicities, nearly all are killed by high multiplicities,
but none make recognizable phage.

The fact that phage susceptibility is of frequent occurrence in
E, coil populations of varying sources makes it probable that in a
continued search a strain having desired properties for studying
host-controlled variation can be obtained.

Summary

Lysing strains of Escherichia coli showed differing responses to
phage Tl. Typical host-controlled variation was demonstrated.
Phage Tl lysate of E. coli B plated on isolated strain Wh24 and
the phage reisolated from developing plaques showed reactions dif¬
fering from that originally obtained on E. coli B.

NOTES ON WISCONSIN PARASITIC FUNGI. XXXII.

Ho C. Greene

Department of Botany, University of Wisconsin, Madison

The season of 1965 in southern Wisconsin was not very favorable
for the development of parasitic fungi, owing to the continuation
of drought conditions up to midsummer. Unless otherwise speci¬
fied, all collections referred to in the following notes were made
in 1965.

General Observations

Venturia sp. on Chamaedaphne calyculata (L.) Moench., col¬
lected in small amount near Trout Lake, Vilas Co., July 17, by J.
Medler, does not seem identical with any of the members of this
genus reported on Ericaceae in Wisconsin. The perithecia are
hypophyllous on dead diistal areas, gregarious but not crowded,
developing well within the host tissue, but still erumpent, black,
globose, thick-walled, up to about 125 p diam., with stiff black
setae approx. 50-60 x 4. fx. The asci are 50-55 x 15-16 p. broadly
clavate or subova te, the ascospores about 20 x 7.5-8 p, greenish
hyaline with septum almost median, but with one cell slightly larger
than the other.

Phyllachora sp., collected on Phalaris arundinacea L. at Madi¬
son, October 30, 1964, unfortunately does not have mature asco¬
spores. Orton, in his monographic treatment of North American
graminicolous species of Phyllachora (Mycologia 36: 39. 1944),
described Phyllachora phalaridis as a new species, known to him
at that time only from the type locality in Massachusetts. The
U. S. D. A. Index mentions no other species of Phyllachora on this
host.

Mycosphaerella sp. occurs on spots primarily due to Ramularia
plantaginis on a leaf of Plantago rugelii Dene, collected August
10 near Leland, Sauk Co., and does not correspond to other species
of Mycosphaerella reported on Plantago. Possibly it is connected
with the Ramularia. The inconspicuous perithecia are grayish-
brown, subglobose, about 75-85 p diam., the slender-clavate asci
36--38 X 6.5-7. 5 p, the hyaline, subfusoid ascospores ca. 8 x 3 p.

147

148 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

Mycosphaerella sp. is associated with Ascochyta compositarum
J. J. Davis on dead areas of leaves of Heliopsis helianthoides (L.)
Sweet collected at Madison, September 3. The black, thick-walled
subglobose perithecia are about 125-150 y diam., the asci clavate,
short-pedicellate, approx. 35-40 x 6-7 y, the hyaline ascospores
isubfusoid, about 12 x 3 ju, with median septum.

PucciNiA sp. (or Uromyces?), represented only by an amphi-
sporic stage, has been noted on a specimen of Carex comosa Boott
collected by H. H. litis near Hope Lake, Jefferson Co., July 28, 1956.
The amphiispores range from oblong, subellipsoid or subfusoid to
more or less broadly ovate, tend to be truncate at base and taper
more or less above to a subacuminate apex, are ca. (30-) 40-55 x
(13-) 14-16 (-18.5) /A, the wall golden-yellow, .8-1.2 y thick at base
and sides, 3-5 (-7) /x above, finely verruculose, the pores 2-3 (-4),
equatorial or superequatorial. A few of the spores have fragments
of pedicels istill attached, but in most they have fallen away. Fig.
1 showts some of the amphispores and was provided by G. B. Cum¬
mins of the Arthur Herbarium at Purdue University, to whom
the specimen was submitted for examination. It seems possible
this may be connected with one of the varieties of Puccinia caricina
DC.

Aleurodiscus oakesii (B. & C.) Cooke is the name usually
applied to the thelephoraceous fungus associated with, and pre¬
sumably causing, “patchy bark” of white oak, Quercus alba L.,
and less commonly bur oak, Q. macrocarpa Michx., in Wisconsin.
This condition is very prominent on the large, open-grown white
oaks in the woods on the University of Wisconsin Observatory

Figure 1. Amphispores on Carex comosa. X800

1966]

Greene — Wisconsin Parasitic Fungi

149

property near Pine Bluff, Dane Co, Some of the trees have lost
all, or practically all, the original bark from ground level to 20 feet
or more up the trunk. Such trees are noticeably whiter and
smoother than uninfected specimens and usually show many of the
tiny, cup-like fruiting structures of the organism on the surface of
the trunk. Large trees do not appear to be seriously damaged by
the fungus, but it gives evidence of being at least mildly parasitic
on struggling small oaks in the partial shade of the bigger trees.
Some of these small trees are completely covered with the fungus
and have died. It seems likely, from examination of cuts made into
the trunks, that the cambium layer of the smaller trees has been
invaded, thus in effect girdling them.

Phyllosticta nebulosa Sacc. was reported in my Notes 31 as
occurring on Lychnis viscaria L. in Wisconsin. Reliance was
placed on named plants in a botanical garden, but examination of
authentic specimens of L. viscaria indicates that though the plants
so named are some species of Lychnis, they cannot be L. viscaria.

Phyllosticta minima Ell. & Ev. has subglobose conidia about
7-8 X 5-6 /X. In a collection of this species on Acer ruhrum L.,
made near Denzer, Sauk Co., July 31, a few of the pycnidia have
conidia which are cylindric and biguttulate, about (4-) 5-6 (-6.5)
X 1.7-2 /X, The spots are very sharply defined and the infection
does not appear to be a mixture of species. Phyllosticta minutis-
sima Ell. & Ev., which occurs commonly on maple, has much smaller
conidia of a micro-type.

Phyllosticta diervillae J. J. Davis on Diervilla lonicera Mill,
was found in the Madison School Forest near Verona, Dane Co.,
July 25, All previous collections were made by Davis in extreme
northern Wisconsin, the latest in 1923.

Phyllosticta wisconsinensis H. C. Greene described occurring
on Helianthus occidentalis Ridd. (Trans. Wis. Acad. Sci. Arts Lett.
53: 211. 1964) has long-cylindric conidia (8.5-) 10-13 (-16) x
2. 5-3. 5 /X and large pycnidia, often 200 /x or more in diam. An
additional specimen on the same host, collected at Madison in 1965,
is practically identical in type of lesion and in microscopic char¬
acters. Two specimens on the closely related Helianthus rigidus
(Cass.) Desf., one collected in 1961 near Cassville, Grant Co., and
the other in 1965 near Albany, Green Co., have very similar
rounded to fusoid lesions and large pycnidia like those of Ph. wis¬
consinensis, but the conidia are shorter, not more than 8 /x, and
somewhat wider, similar to the conidia of Phyllosticta favillensis
Greene (Amer. Midi. Nat. 48: 50. 1952), described from a speci¬
men on Silphium integrifolium Michx. and currently represented

150 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

in the Wisconsin Herbarium by four specimens on this host. The
Albany and Cassville specimens on H. rigidus are being filed
temporarily with the Phyllostictae indet. They appear, however,
to be related to Ph. wisconsinensis rather than to Ph. favillensis.

Phyllostictae, appearing parasitic, but so far undetermined as
to species, continue to be found on diverse hosts, as indicated in
the following descriptive notes: 1) On Pteridium aquilinum (L.)
Kuhn var. latiusculum (Desv.) Underw. collected near Leland,
Sauk Co., August 31, 1964. On indeterminate, dull reddish-brown
areas; pycnidia epiphyllous, black, subglobose, widely ostiolate,
pseudoparenchymatous, small, about 60-75 /x diam., tending to be
in lines following the venation ; conidia hyaline of the micro-type,
about 4. 5-6. 5 x .7-1 /x. 2) On Quercus ellipsoidalis. Collected at
Madison September 14. Spots very sharply defined, rounded, with
rather wide reddish-brown borders and very light brown centers,
4-6 mm. diam. ; pycnidia epiphyllous, loosely to closely gregarious,
shiny black, deeply seated in tissue, globose or subglobose, approx.
100-150 fx diam. ; conidia subglobose to ovoid, 6.5-8 x 9.5-
10.5 (-12) /X. Phyllosticta globulosa Thum., which also occurs on
oak, is described as having subglobose or ovate-globose conidia 6-9
/X diam., but plainly differs in other characters. 3) On Oxyhaphus
nyctagineus (Michx.) Sweet collected in Dane Co., near Arena,
July 8, 1964. Spots dull brown, small and marginal, usually bearing
only one or two pycnidia, but occasionally more; pycnidia amphi-
genous, mostly epiphyllous, black, subglobose, about 125-175 /x
diam., the ostiole delimited by a dense ring of black cells; conidia
hyaline, narrow-f usoid, approx. 8-11 x 2. 4-2. 7 (-3) /x. The
conidial shape and the rather large black pycnidia suggest that
this may prove to be a species of Phomopsis, but no scolecospores
were seen in the mounts studied. 4) On three specimens of
Caulophyllum thalictroides (L.) Michx., the first collected July 6
at Gov. Dodge State Park, Iowa Co. The conspicuous spots are
ashen with a very narrow yellowish-brown border, orbicular to
oblong, .3-7 cm. in short diam. ; pycnidia epiphyllous, scattered,
from somewhat flattened to subglobose, thin-walled, pallid
yellowish-brown, small, about 50-75 /x diam.; conidia hyaline, sub-
cylindric to subfusoid or broadly subfusoid, straight or slightly
curved, about 4.5-8 (-10) x 2. 4-3. 2 /x. The second specimen was
taken a few days later, July 14, at the same station. Here the
lesions are large, effuse, sordid greenish-brown areas involving
the distal portions of leaflets; pycnidia many, flesh-colored, about
100-150 /X diam.; conidia similar in shape to, but slightly larger
than, the July 6 specimen. The third specimen was gathered
August 28 at Wildcat Mt. State Park, Vernon Co. Here the con¬
spicuous lesions are wedgeshaped, distal in situation, up to 5 cm.

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long by 3 cm. at widest point, subzonate, tan, with narrow darker
margin ; pycnidia loosely gregarious, epiphyllous, rather dark
brown and thick-walled, subglobose, about 125-200 p. diam. ;
conidia similar to those in the other two specimens. Perhaps all
represent progressive stages in the development of the same thing.
I have found no report on any Phyllosticta on Caulophyllum. 5)
On Potentilla recta L. collected at Tower Hill State Park, Iowa
Co., October 13. Very much like a Phyllosticta which occurred on
Fragaria virginiana Dene., as reported in my Notes 26 (Trans.
Wis. Acad, Sci. Arts Lett. 49: 89. 1960). In both specimens the
zonate banding of the spots is similar, the conidiophores well-
developed, the conidia correspond in size and shape, and the
pycnidia are erumpent, but lighter in color and less markedly
rostrate on Potentilla. 6) On Staphylea trifolia L. collected at
Nelson Dewey State Park, Grant Co., September 19, 1961. The
spots are ashen-brown, immarginate, irregular, approx. 1 cm. diam.,
pycnidia hypophyllous, gregarious, dark brown, subglobose, appar¬
ently without ostioles, about 75-90 p diam.; conidia hyaline, short
rod-shaped, 3-3.5 x .8-1 ix. 7) On Menyanthes trifoliata L. col¬
lected June 12 in Hope Lake Bog near Cambridge, Jefferson Co.
The spots are tan with narrow darker border, rounded, about 4-6
mm. diam.; pycnidia epiphyllous, gregarious, light brown,
pseudoparenchymatous, subglobose with prominent ostiole, about
80-110 jtt diam.; conidia hyaline, rod-shaped, about 2.5-3 x .7-1
fji, very numerous. 8) On Scrophularia marilandica L., two speci¬
mens, from Gov. Dodge State Park, Iowa Co., August 23, and from
near Leland, Sauk Co., August 24. Spots sordid brown, sometimes
purple-bordered, ranging from rounded and only 2-3 mm. diam. to
large irregular blotches; pycnidia epiphyllous, scattered to gre¬
garious, pallid brownish, thin-walled, subglobose, ca. 90-140 p
diam. ; conidia hyaline, ellipsoid to short-cylindric, quite variable
in size, seeming to run somewhat smaller in the Sauk Co. specimen,
but intergrading, approx. (3.5-) 5-7 (-8.5) x 1.5-2 p. Similar to
but better developed than specimens on this host reported on in
my Notes 30. European species are described on Scrophularia, but
none correspond in conidial size with the Wisconsin specimens.
9) On Pentstemon gracilis Nutt. var. wisconsinensis (Penn.)
Fassett collected near Lodi, Columbia Co., June 7, 1960. Spots
narrow, elongate, subtramslucent, ashen with brownish borders;
pycnidia seriately arranged, black, subglobose, approx. 135-175 p
diam., the ostiole delimited by a very thick ring of black, heavy-
walled cells; conidia very numerous, hyaline, straight or slightly
curved, narrowly cylindric, 4-6 x 1-1.3 p. 10) On Aureolaria
(Gerardia) pedicularia (L.) Raf. collected near Leland, Sauk Co.,
August 14. Spots small, about 2-3 mm. diam., rounded, subzonate.

152 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

dark brown but often with a paler center; pycnidia epiphyllous,
mostly closely crowded in the central portion of the spot, pallid
sooty=brown, isubglobose, approx. 125-150 p. diam. ; conidia hyaline,
ellipsoid to broadly ellipsoid, short-cylindric or occasionally sub-
fusoid, biguttulate in some pycnidia, variable in size, (3.5-) 5-
7 (-11) X 2-3 (-4) IX, 11) On Triosteum perfoliatum L. Two
specimens, the first from near Pine Bluff, Dane Co., September
5, 1964. Spots ranging from tiny, angled and ashen, about 1 mm,
diam. to larger, indefinite, light reddish-brown areas; conidia
hyaline, very small, 2.5-3 x .5-.7 p,. Similar material has been
collected in August or later in several localities and twice leaves
have been held out-of-doors over winter without any further de¬
velopment. The second specimen was taken September 12, 1964,
at Gov. Dodge State Park, Iowa Co. Here the lesions are large,
about 2-2.5 cm. diam., ovate and brownish, with a cinereous center;
pycnidia epiphyllous, blackish, subglobose, approx. 125-150 p
diam.; conidia hyaline, ellipsoid, small, 3-3,5 (-4.5) x 1-1.3 p.
12) On Viburnum cassinoides L. (cult.) collected at Madison,
October 5. Spots small, rounded, dark, elevated; pycnidia epiphyl¬
lous, black, subglobose, about 125-150 p diam.; conidia hyaline,
subcylindric or subfusoid, approx. 5-8 x 2.5-3 p. 13) On Solidago
canadensis L. from the Flambeau State Forest near Oxbow, Sawyer
Co., July 23, 1964. Spots rounded and dark-bordered, with cinere¬
ous centers, small, about 1-2 mm. diam. ; pycnidia epiphyllous, one
or two per spot, pallid sooty-brown, thin-walled, globose, about
150-175 p diam., the ostiole delimited by a ring of dark, thick-
walled cells ; conidia hyaline, narrow-cylindric, straight or slightly
curved, often guttulate, approx. 6-10 x 1.8-2. 3 p. 14) A Phyllos-
ticta very similar microscopically to the preceding occurs on Aster
prenanthoides Muhl. collected near Leland, Sauk Co., August 19,
1964. The fungus is hypophyllous on orbicular lesions 1-2 cm.
diam., which are purplish above and dull yellowish below. 15) On
Silphium perfoliatum L. collected near Leland, Sauk Co., September
19, 1964. Spots rather indefinite, mostly small and somewhat
rounded, but becoming confluent over considerable areas, mottled
dark gray and ashen; pycnidia epiphyllous, mostly rather closely
clustered on, but not confined to, the lighter portions of the spots,
small, black, globose, about 50-90 p diam., without true ostioles,
although some pycnidia have rounded, thin areas in the walls;
conidiahyaline, short rod-shaped, 3-5 x .6-8 p. 16) On Helian-
thus giganteus L. collected near Leland, Sauk Co., August 12, 1964.
Spots small, angled and ashen on larger indefinite brown areas;
pycnidia epiphyllous, usually only one to a spot, black, subglobose,
about 150-175 p diam, ; conidia hyaline, cylindric, 6-7 x 1.5-2 p.
17) On Arctium minus Bernh. from Gov. Dodge State Park, Iowa

1966]

Greene— Wisconsin Parasitic Fungi

153

Co., October 1. Spots mottled, cinereous through blackish-brown,
irregular in shape and size; pycnidia epiphyllous, scattered, dark
brown, subglobose, about 150-175 /x diam. ; conidia hyaline, cylin-
dric, 4-5.5 x 1.5-1. 8 /x, sometimes biguttulate.

CONIOTHYRIUM spp. indet, and possibly parasitic have been
noted. 1) On Salix discolor MuhL collected near Leland, Sauk Co.,
August 4, 1964. Spots small, fuscous, marginal ; pycnidia epiphyl-
lous, scattered, black, erumpent, subglobose, approx. 115-125
diam, ; conidia greenish-gray, oblong or broadly ellipsoid, 5-6.5 x
2.5-3 2) On Ulmus americana from near Leland, Sauk Co.,

June 18. Spots sharply defined, usually only one or two to a leaf,
pallid- to reddish-brown, with narrow darker borders, rounded,
about 2-4 mm. diam. ; pycnidia epiphyllous, more or less closely
gregarious, black, rather thick-walled, subglobose, mostly about
100-150 fi diam. ; conidia smooth, clear olivaceous-gray, broadly
elliptic, 4-7 x 2. 7-3. 5 ,/x. Because many of the leaves in this large
collection bear, on rather similar spots, the Phyllosticta which has
in Wisconsin lists been doubtfully referred to P. ulmicola Sacc., it
seems possible that the Phyllosticta was primary, especially since
a few spots show a mixture of Coniothyrium and Phyllosticta.
The form in hand does not in any way correspond to Tharp's
Coniothyrium ulmi (Mycologia 9: 116. 1917). 3) On Cotoneaster
“melanocarpa'' (cult.) collected at Madison, September 11, 1964.
The spots are rounded or angular, about 2-5 mm. diam., essentially
sordid- or rufous-brown, but the centers often appear cinereous,
due primarily to a loosened and somewhat uplifted cuticle ; pycnidia
epiphyllous, erumpent and gregarious, appearing intraepidermal in
origin, black, thick-walled, subglobose, approx. 100-160 diam.;
conidia light grayish-olivaceous, oblong to broadly ellipsoid, or
sometimes almost globoid, about 2.7-3. 8 x 5-5.5 ,/x. 4) On Acer
negundo L. from near Leland, Sauk Co., August 10. Spots orbicu¬
lar, ashen and translucent, with narrow yellow-brown border, about
.3-1 cm. diam, ; pycnidia mostly epiphyllous, scattered, globose or
subglobose, approx. 100-150 diam. ; conidia very numerous,
smooth, clear light gray, 4-5.5 x 2.3-3. 5 ./x. Evidently not Coniothyr¬
ium negundinis Tehon & Daniels, which occurred at the bases of
twigs, had pycnidia twice as large, and smaller, olivaceous, spheri¬
cal to ovoid conidia,

Phomopsis sp, was present in profusion on still attached over¬
wintered fruit of a cultivated species of Rosa collected June 22 at
Madison. The large, crowded, black, globose pycnidia are approx.
175-250 .p. diam,, the hyaline scolecospores ca, 18-22 x 1.2-1. 6 /x,

154 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

from almost straight to sinuously curved, enlarged at one end, the
other conidia subfusoid, 6-9 x 2-2.5 y. Both types of spores are
abundant. Parasitic in origin?

Phomopsis sp. ? occurs on cinereous areas of leaflets of Caragana
arhorescens Lam. (cult.) collected at Madison, July 20. Pycnidia
black, thick-walled, prominently ostiolate, gregarious to crowded,
mostly epiphyllous, subglobose, variable in size, about 100-250 /x
diam. Deflnite scolecospores were not seen, but conidia range
from rather broadly fusoid to moderately slender in one group
which run 7-10.5 x 3-3.8 /x, to a second group where they are
about 12-13 X 2 /X at one end and tapering to 1 /x at the other, thus
verging on a scolecosporous condition, with an aspect strongly sug¬
gestive of Phomopsis. Phomopsis caraganae Bond, on stems has
fusoid conidia of similar size,

Ascochyta spp., ranging from well-developed to more or less
presumptive, have been found on 1) Apios tuberosa Moench. col¬
lected at Gov. Dodge State Park, Iowa Co., July 6. Well character¬
ized and appearing mature. Spots conspicuous, greenish to pallid
brownish with narrow dark brown margin, translucent, orbicular,
about .5-1.5 cm. diam. ; pycnidia carneous, thin-walled, gregarious
to crowded, subglobose, approx. (90-) 125-175 /x diam.; conidia
hyaline, subcylindric or subfusoid, 7-10 (-11) x (2.6-) 3-3.5 (-4)
/X, regularly uniseptate, occasionally slightly constricted at septum.
In essentials this seems very similar to an undetermined Ascochyta
reported in my Notes 29 (Trans. Wis. Acad. Sci. Arts Lett. 52:
236. 1963) and it seems likely that other specimens on both Apios
and Amphicarpa with this type of lesion and pycnidia, referred
doubtfully to Phyllosticta phaseolina Sacc., in reality belong here.
More collections on both hosts would be desirable. 2) on Convolvu¬
lus sepium L., Madison, July 9, 1964. The blackish subglobose
pycnidia are about 125 /x diam., the hyaline, guttulate, uniseptate
conidia about 10-12 x 2.5 /x. The pycnidia are on the same type of
reddish-brown, zonate spots characteristic of Stagonospora con¬
volvuli Dearn. & House, so it seems possible thisi is merely a some¬
what depauperate development of that species. 3) On Polemonium
reptans L. collected near Leland, Sauk Co., July 15. Spots diapha¬
nous, ashen-brown, orbicular to ovate, about 1 cm. diam. ; pycnidia
scattered to gregarious or even crowded, pallid brownish and thin-
walled, with a well-defined ostiole delimited by a narrow ring of
dark cells, subglobose, mostly 125-200 /x diam., even more in a few
cases; conidia hyaline, short-cylindric, broadly ellipsoid, or sub¬
fusoid, a small number with a median septum, about 5-7.5 x
2.4-3 ,/x. The specimen appears well-matured, but perhaps more
conidia would have developed septa in time. There seem to be no

1966]

Greene — Wisconsin Parasitic Fungi

155

reports of Ascochyta or Phyllosticta on Polemonium in North
America. Ascochyta polemonii Cav. on cultivated P. caeruleum L.
in Europe has conidia 12-14 x 3 /x. 4) On Glecoma (Nepeta)

hederacea L. collected near Albany, Green Co., October 1, 1964.
Spots rounded, sordid brownish with darker border, about 5 mm.
diam. ; pycnidia epiphyllous, gregarious, thin-walled, subglobose,
about 100-125 ,/x diam.; conidia hyaline, cylindric, uniformly and
markedly biguttulate, 5-7 x 1.8-2. 2 /x. The conidia are much
smaller than those of Ascochyta nepetae Davis which occurs on
Nepeta cataria L. 5) On a leaf of V eronicastrum virginicum (L).
Farw. collected July 15 near Leland, Sauk Co. Spot blackened,
strongly zonate, about 4 mm. diam. ; pycnidia flesh-colored, sub-
globose, about 100-125 ^ diam. ; conidia uniformly uniseptate, hya¬
line with granular contents, cylindric and obtuse, about 7.5-10 x
3-3.5 /X. Directly centered on the reverse of the spot is a sorus of
the microcyclic rust Puccinia veronicarum DC. In the same gen¬
eral area on September 11, on the same host, a possible Ascochyta
was found on orbicular, blackish, zonate areas, about 1-2.5 cm.
diam. ; pycnidia sooty-brownish, thin-walled, subglobose, approx.
100-125 /X diam., scattered and very inconspicuous ; conidia hyaline,
cylindric, sparingly uniseptate, about 9-10 x 2-2.3 /x, mostly non-
septate and smaller, about 4. 5-7. 5 x 1.3-1. 7 /x. 6) On Antennaria
parlinii Fern, collected August 31, 1964, near Leland, Sauk Co.
Spots sordid brown with narrow darker margin, irregular in shape
and involving the distal portions of the leaves; pycnidia epiphyl¬
lous, gregarious, black, subglobose, thick-walled, erumpent, approx.
125-150 /X diam. ; conidia hyaline, cylindric, uniformly uniseptate,
not constricted at septum, (11-) 12-13 (-14) x 2. 5-2. 8 /x. The ra¬
tio of length to width of the conidia suggests that this might ulti¬
mately prove to be a Stagonospora.

Aster prenanthoides Muhl., collected near Leland, Sauk Co.,
August 12, 1964, bears on the living leaves a possibly parasitic and
peculiar sphaeropsidaceous fungus which seems to fall between
Diplodina West, and Chaetodiplodina Speg. There are no spots.
The pycnidia are sooty brown, relatively thin-walled and pseudo-
parenchymatous, globose, about 110-225 /x diam., the ostiole small
but sharply outlined by a ring of dark cells, hypophyllous, scat¬
tered and superficial on a small, loosely organized subiculoid net¬
work, the component hyphae of which are brownish and appear to
originate as strands from wall cells at various points on the pyc-
nidium. Conidia are hyaline, straight or slightly curved, cylindric,
long-cylindric, or subfusoid, many appearing continuous, but many
uniseptate and a few obscurely 3-septate, approx. (15-) 17-30 (-33)
X 3.5 — 5 ,/x.

156 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Septoria sp. is strictly confined to the spermogonial surface of
aecial sori of Puccinia dioicae P. Magn. on leaves of Solidago
patula Muhl. collected July 1 near Leland, Sauk Co. The sori have
a conspicuous blackish-purple margin, quite unlike sori on adjacent
leaves which bore the rust alone. The tiny black pycnidia are only
about 35-50 /x diam., the hyaline acicular spores approx. 15-20
X 1 jit.

Hendersonia sp. — a very large-spored form — occurs on conspic¬
uous spots on leaves of Spartina pectinata Link collected at Madi¬
son, September 3, 1965. This spotting of Spartina has been noted
over the years at various stations, and specimens have been pre¬
served. In my Notes 20 (Trans. Wis. Acad. Sci. Arts Lett. 43:
173. 1954) I described the spots of a specimen collected in 1952
near Mazomanie, Dane Co., as “remarkably conspicuous, large,
orbicular . . . with grayish centers and wide, purplish-brown
borders on the upper surface of the leaves. On the lower surface
and coinciding with the spots are wefts of sordid-whitish, largely
superficial, yet closely appressed mycelium. Microscopically this
mycelium is thin-walled, septate, and somewhat verrucose. . .

At that time I overlooked the very scanty and inconspicuous devel¬
opment of the Hendersonia, as I did in subsequent specimens, until
the 1965 Madison collection, where there is a relatively profuse de¬
velopment of pycnidia. Taking all the specimens into account, one
finds that the pycnidia are from scattered and very few in some to
gregarious and fairly numerous in others. They are subglobose,
about 160-200 /x diam., sooty-olivaceous, with a small ostiole de¬
limited by a ring of dark, thick-walled cells. The large conidia are
from almost straight to variously curved, widest at or near the
middle, ends obtuse, clear yellowish-olivaceous, 6-9, but mostly
7-septate, approx. (60-) 75-100 (-120) x (11-) 12-13 (-15) /x. The
relation of the superficial hypophyllous mycelium to the Hen¬
dersonia is not clear, although it occurs in greater or less develop¬
ment on the reverse of all the spots and is confined to them.

Cylindrosporella ( ?) sp. occurs on mottled green and brownish
areas of leaves of Podophyllum peltatum L. collected May 29 near
Albany, Green Co. The subcuticular acervuli are most inconspicu¬
ous and even in section discernible with difficulty. They are very
slightly concave to almost plane and the conidiophores on them
very short, almost obsolete. The conidia, which are produced in con¬
siderable numbers, are hyaline, cylindric or subcylindric, straight
or slightly curved, (6-) 8-10 (-12) x 1.2-2. 6 /x. This plainly has
no connection with Septotinia podophyllina (Ell. & Ev.) Whetzel,
which has much larger septate conidia and lesions of a different
aspect.

1966]

Greene — Wisconsin Parasitic Fungi

157

COLLETOTRICHUM spp. indet, have been noted on several hosts as
possible parasites. 1) On leaf midribs of Corylus americana Walt,
collected near Pine Bluff, Dane Co., September 5, 1964. The leaf
adjacent to the midrib is brownish and discolored, suggesting
parasitism. The fructifications are elongate and deep-seated in the
tissue, the stiff setae black below, somewhat paler toward the tip,
1-2 septate, approx. 65-150 x 5-6 p. (somewhat wider at the very
base) , a few shorter and narrower, mostly in pairs or small groups.
The conidia hyaline, falcate, 20-23 x 2. 5-3. 2 p. 2) On Asarum
canadense L. collected in Wyalusing State Park, Grant Go., June
24. Spots blackened, orbicular, about .5-1 cm. diam. ; acervuli
epiphyllous, scattered to gregarious; setae rather coarse, fascicled
and prominent, clear purplish-brown below to slightly paler above,
tapering gradually to the subacuminate tips, slightly to moderately
curved, approx. 90-140 x 5-7 p, 2-5 septate; conidia hyaline,
falcate to almost lunate, about 17-23 x 2.5-3 p. The affected leaves
were growing among and surrounded by healthy leaves on a high,
deeply shaded bluff where frost damage could scarcely have been a
factor. Possibly parasitic, but Asarum canadense is notable for
lack of parasites, with the only so far determined fungus reported
on it from Wisconsin or elsewhere being Synchytrium asari Arth.
& Holw. 3) On dead upper portions of stems of Desmodium brac-
teosum (Michx.) DC, var. longifolium (T. & G.) Rob. from the
New Glarus Woods Roadside Park, Green Co., October 1, 1964. The
plants were, in the main, still living and it seems probable that
the fungus caused the death of the stem tips. No setae were ob¬
served, but the organism is perhaps referable to Colletotrichum
in the usage of von Arx. The acervuli occur in profusion, are dark,
subepidermal, small, mostly only about .2 x .1 mm. ; conidiophores
closely compacted and olivaceous below, but paler above in the free
measurable portion where they are from about 8-10 x 2,5-3 p. ap¬
pearing to be confined mostly to the margin of the acervulus;
conidia subhyaline, cylindric, about 11-15 x 3. 5-4. 5 p. 4) On
Asclepias exaltata (L.) Muhl. collected at Madison, September 11,
1964, Lesions elongate, pallid greenish with wavy black borders,
approx. .5 cm. wide by 2-4 cm. long; acervuli epiphyllous and gre¬
garious; setae peripheral, slender, flexuous, uniform clear dark
brown, little if any paler at the subacuminate tips, sparingly sep¬
tate, approx. 60-85 x 3. 5-4. 5 p; conidia hyaline, straight, cylindric
I and obtuse, occasionally subfusoid, about 14-17 x 3. 5-5. 5 p.

I Marssonina (?) sp. occurs on Quercus velutina Lam. collected
j at Madison, October 3, 1964. The flesh-colored acervuli are hy-
I pophyllus on elongate brownish areas along the principal veins,
j In my Notes 26 (Trans. Wis. Acad. Sci. Arts Lett. 49: 95. 1960) I

158 Wisconsin Academy of Sciences, Arts- and Letters [Vol. 55

mentioned a very similar fungus on Quercus alha L., where the
conidia “vary from rarely obclavate to cylindric, broadly cylindric
or ellipsoid, or curved Marssonina-like, continuous so far as ob¬
served, 18-36 X 6.5-9 fx.” One would suspect that the great irregu¬
larity in conidial size may be the result of late season development,
with accompanying wide temperature extremes, but even so this
organism does not seem close to any fungus currently reported
from this area on either white or black oak.

Botrytis (B. vulgaris Fr.?) appears strongly parasitic on elon¬
gate brownish lesions which encircle the stems of greenhouse-
grown tobacco, Nicotiana tabacum L., at Madison in September
1965. The fungus is fruiting profusely, with whorls of subhyaline,
short-clavate branches produced near the tips of comparatively
long, septate, brownish-olivaceous conidiophores. The conidia are
grayish, smooth, broadly elliptic, oval or oblong, 8-10 (-13) x 6-9 ju.

Cladosporium sp., appearing parasitic, is epiphyllous on Andro¬
meda glaucophylla Link collected in Hope Lake Bog near Cam¬
bridge, Jefferson Co., June 12. The spots are rounded, reddish-
purple, small, 1-2 mm. diam., or confluent and larger, mostly mar¬
ginal on the narrow leaves ; conidiophores loosely clustered from a
isubstromatic base to fascicled from a well-developed stroma, multi-
septate, from almost straight to slightly curved or sinuous, a few
subgeniculate, simple, subdenticulate, clear brown below, becoming
pallid above, 35-60 x 3-4 jx ; conidia ellipsoid to subfusoid, continu¬
ous or 1-septate, catenulate, roughened, grayish-olivaceous, about
10-12 (-15) X 3-4 jx, I have not found any reports of Cladosporium
on Andromeda or related plants.

Cercospora sp. in small amount, has been observed on Gaultheria
procumbens L. collected by J. A, Curtis in the Bittersweet Lake
Scientific Area near Eagle River, Vilas Co., July 25, 1963. This
bears no resemblance to Cercospora gaultheriae Ell. & Ev. nor to
other species listed on Ericaceae in Chupp’s monograph. The fun¬
gus is hypophyllous and quite diffuse, the conidiophores clear
brown, markedly geniculate and tortuous, often rather intricately
branched and intermingled, but not compacted into a stroma, about
4.5-6 fx diam. at the base, several-septate and up to 150 /x long. The
conidia are dilute olivaceous, narrowly obclavate to almost acicular,
straight to moderately curved, 5-11 or more septate, base subtrun¬
cate, 55-105 X 3.5-4.5 /x.

Alternaria sp. in an apparently parasitic condition on Euphor¬
bia esula L. was first noted at Madison in 1949 and commented upon
briefly in my Notes 14. The material was rather old, however, and
not suitable for close study. In 1965, in the same general location.

1966]

Greene- — Wisconsin Parasitic Fungi

159

much fresher and quite plainly parasitic Alternaria was collected
and studied on this host. The spots are sordid brown with a narrow
darker border, rounded or angular in basic outline, usually extend¬
ing from about the midrib up to, and involving, the margins of the
narrow leaves, subzonate and approx. 2-5 mm. diam. The conidi-
ophores are amphigenous, but most prominent on the upper leaf
surface, and appear to be produced from stomates in a few cases,
in broadly diverging groups of half a dozen or more. They are clear
olivaceous-brown, 3-4 septate, up to 60 x 4 ju and from almost
straight to curved and mildly tortuous-geniculate. The main spore
bodies are olivaceous-gray or olivaceous, broadly ovate to clavate,
about 28-50 x 12.5-14 (-15) /x, the narrow beaks subhyaline to dilute
olivaceous, approx. 15-30 (-35) x 2-3 /x. The spores with beaks
mostly run about (50-) 60-70 (-80) x 12-15 /x with 3-9 transverse
septa and about 2-5 vertical septa. Alternaria brassicae (B.) Sacc,
is reported in Seymour's Index as occurring on Euphorbia esula, but
according to Neergaard in his authoritative “Danish Species of
Alternaria and StemphyliuriF’ , A. brassicae is a much coarser spe¬
cies, The Wisconsin specimen likewise does not correspond with
Macrosporium (Alternaria) euphorbiae BarthoL, which has wider
spores and much longer beaks.

Graphiothecium vinosum J. J. Davis was described (Trans,
Wis. Acad. Sci. Arts Lett. 18(1) : 90. 1915) as occurring on Ribes
americanum Mill, at Madison, with the observation that the fungus
reached full maturity only after overwintering. The last previous
collection on this host was made nearly 50 years ago, but in Sep¬
tember 1964 at Tower Hill State Park, Iowa Co., leaves of Ribes
americanum, infected with what appeared to be possibly an imma¬
ture Ascomycete, were gathered and overwintered out-of-doors at
Madison, In May 1965 these were found to bear numerous vinous-
purplish synemmata characteristic of Graphiothecium vinosum.

Graphiothecium sp. developed on leaves of Lonicera tatarica L.
collected at Madison, October 6, 1964, with the fungus in immature
condition, and held out-of-doors until May 1965. The closely gre¬
garious synemmata are amphigenous and deeply seated in the leaf
tissue, with a black, bulbous base about 125-150 /x diam. The synem¬
mata proper are approx. 175-200 /x long by 25 /x or more thick, black
and compact below, but becoming hyaline and more loosely organ¬
ized above. The conidia are hyaline, fusoid to subcylindric, about
10-17 X 2,5-4 /X, often produced at right angles to the stalk.
Although a late season collection, the fungus appeared to have ini¬
tiated its development as a parasite. So long as unblighted by frost,
Tatarian honeysuckle and similar exotics remain green and active
much later in the fall than most native species. At the time of col-

160 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

lection the fruiting structures resembled immature pycnidia or
perithecia, with no indication of the ultimate synemmatal develop¬
ment which evidently occurred in the spring of 1965.

Lonicera morrowi Gray observed near Ridgeway, Iowa Co.,
August 5, 1964, bore on the under surface of the leaves a curious,
possibly parasitic, presumed fungus which is snowy white and had
developed in narrow lines in the form of closed rings, partial rings,
and other less regular serpentine patterns. All leaves on any one
branch consistently bore the fungus and, except for it, were very
clean-appearing with no evidence whatsoever of the debris one
would ordinarily expect had the growth accompanied insect infes¬
tation. Although superficial in aspect, the organism is very closely
appressed and not readily removed. Microscopically it consists of
masses of closely interwoven, non-fruiting, hyaline, very slender,
hyphae-like threads, about 1 jx wide, which are only obscurely sep¬
tate, or perhaps even non-septate. These threads quite regularly
ascend the trichomes but appear to be superficial on them.

Additional Hosts

The following hosts have not been previously recorded as bearing
the fungi mentioned in Wisconsin.

Plasmopara viburni Peck on Viburnum opulus L. (cult.). Mil¬
waukee Co., Fox Point, September 24, 1962. Coll. & det. J. W.
Baxter.

Syzygites megalocarpus Ehr. ex Fr. on Entoloma (grayanum
Peck?). Dane Co., Madison, September 27. Coll. & det. R. J. Boles.

Microsphaera alni (Wallr.) Wint. on Lonicera morrowi Gray.
Iowa Co., near Ridgeway, August 5, 1964.

Phyllachora gram inis (Pers.) Fckl. on Elymus iviegandii
Fern. Grant Co., Wyalusing State Park, August 28, 1957. Coll. H. H.
litis.

Ophiodothis haydeni (B. & C.) Sacc. on Aster sagittif olius
Willd. Sauk Co., near Leland, September 11.

Metasphaeria leersiae (Pass.) Sacc. on Leersia virginica
Willd. Jackson Co., Gullickson’s Glen near Disco, August 21, 1963.

Leptosphaeria elymi Atk. on Elymus canadensis L. Sauk Co.,
near Leland, July 15.

Ceratostomella ulmi Buisman on slippery elm, TJlmus rubra
Muhl. E. B. Smalley, Dutch elm disease specialist at the University
of Wisconsin, informs me that the disease is now general upon slip-

1966]

Greene — Wisconsin Parasitic Fungi

161

pery as well as upon American elm in southern Wisconsin, and that
he has noted its natural occurrence on Ulmus pumila L. in the
state, although this species is comparatively resistant.

Cronartium ribicola Filsch. II, II on Ribes diacantha Pall,
(cult.). Dane Co., Madison, September 22.

Melampsora abieti-caprearum Tub. II, III on Salix adeno-
phylla Hook. (cult.). Dane Co., Madison, October 5.

PucciNiA CARICINA DC. II on Carex comosa Boott. Oneida Co.,
near Woodruff, July 5, 1958. Coll. H. H. litis.

PucciNiA DIOICAE P. Magn. I on Solidago uliginosa Nutt. Sawyer
Co., Flambeau State Forest near Oxbow, July 24, 1964.

PUCCINIA DIOICAE P. Magn, II, III on Carex annectens Bickn.
var. xanthocarpa (Bickn.) Wieg. LaCrosse Co., Town of Farming-
ton, June 29, 1959. Coll. A. M. Peterson. This is the punctate form,
Puccinia vulpinoidis Diet. & Holw., now regarded as a synonym.

PUCCINIA KARELICA Tranz. II, III on Carex crinita Lam. Florence
Co., Lost Lake, July 14, 1939. Coll, A. L. Throne. Det. J. W. Baxter.

Puccinia asteris Duby on Aster puniceus L, Iowa Co., Gov.
Dodge State Park, July 14.

Uromyces SILPHII (Burr.) Arth. I on Helianthus tuherosus L.
Sauk Co., near Leland, June 16.

Uromyces sporoboli Ell. & Ev, III on Sporobolus heterolepis
Gray. Kenosha Co., near Woodworth, August 10, 1954. Coll. P. B.
Whitford. Det. J. W. Baxter.

CiNTRACTiA CARicis (Pers.) Magn. on Carex meadii Dewey. Iowa
Co,, near Ridgeway, June 14.

Phyllosticta hispida Ell. & Dearn. on Smilax ecirrhata
(Engelm.) Wats. Green Co., New Glarus Woods Roadside Park,
September 8, 1951. Although an adequate specimen was placed in
the herbarium at the time of collection, it was overlooked and not
recorded in these notes.

Phyllosticta monardae Ell. & Barth, on Monarda punctata L.
Sauk Co., near Leland, September 11. P. monardae is said to be syn¬
onymous with Phyllosticta decidua Ell. & Kell., but because the
description does not fit P. decidua as I understand it, on an interim
basis I have applied the name P. monardae to a species of Phyllos¬
ticta with non- translucent spots which occurs on Monarda, Ble-
philia, Lycopus, Mentha and Py cnanthemum in Wisconsin and
which corresponds well with the description.

162 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Phoma polygramma (Fr.) Sacc. var. plantaginis Sacc. on
Plantago rugelii Dene. Racine Co., Racine, October 30, 1894. Coll.
J. J. Davis. This is labeled Phyllachora plantaginis Ell. & Ev. and
is presumably a portion of the specimen from which Ellis and Ever¬
hart described that species. Except for the fact that the fusoid
conidia are slightly longer than in specimens on Plantago lance-
olata L., discussed by me in my Notes 27 (Trans. Wis. Acad. Sci.
Arts Lett. 50: 159. 1961), the overall aspect is very similar indeed,
although in P. rugelii the fungus is on the leaves, whereas in P.
lonceolata it is normally confined to the flowering scapes.

Rhizosphaeara kalkhoffi Bub. on Picea pungens Engelm.
(cult.). Grant Co., Wauzeka, October 1964. Coll. G. L. Worf.

Neottiospora arenaria Syd. on Carex lanuginosa Michx. Adams
Co., Springville Twp., September 13, 1958. Coll. T. G. Hartley
(6273).

CONIOTHYRIUM FUCKELii Sacc. on Rubus strigosus Michx. Sauk
Co., near Leland, August 31, 1964.

Ascochyta violae Sacc. & Speg. on Viola sororia Willd. Sauk
Co., near Leland, August 31, 1964.

Ascochyta compositarum J. J. Davis on Ambrosia trifida L.
Iowa Co,, Gov. Dodge State Park, July 6. The smaller-spored
variety.

Stagonospora albescens j. j. Davis on Carex hystricina Muhl.
Florence Co., near Lost Lake, June 14, 1959. Coll. H. H. litis. On
C. rostrata Stokes. Sawyer Co., Flambeau State Forest, June 24,
1959. Coll. E. Beals.

Septoria nematospora j. j. Davis on Carex intumescens Rudge.
Sawyer Co., near Loretta, June 12, 1959. Coll. L. Hathaway.

Septoria caricinella Sacc. & Roum. on Carex lactust7is Willd.
Price Co., Chequamegon National Forest near Park Falls, June 20,
1959. Coll. E. Beals.

Septoria caricina Brun. on Carex pedunculata Muhl. Sauk Co.,
Parfrey's Glen, June 8, 1964.

Septoria glycines Hemmi on Amphicarpa bracteata (L.) Fern.
Dane Co., Madison, August 6, 1964. Hemmi gives the pycnidial
diameter as 44-100 jx and the length of spores at 21-52 y. On Amphi¬
carpa they are about 50-65 fx and 20-40 y, respectively and the
lesions are dull brownish, cuneate, about .5-3 cm. long by not more

1966]

Greene — Wisconsin Parasitic Fungi

163

than 1 cm. wide, usually involving the tips of leaflets. The pycnidia
are epiphyllous and gregarious, mostly concentrated along the prin¬
cipal veins,

Septoria lobeliae Peck on Lobelia kalmii L, Door Co., Egg Har¬
bor, August 27, 1945. Coll. R. A. McCabe. On basal leaves of a pha¬
nerogamic specimen,

Septoria matricariae Hollos on Anthemis cotula L. Dane Co.,
Madison, June 3.

Sphaceloma rosarum (Pass.) Jenkins on Rosa rugosa Thunb.
(cult.). Dane Co., Madison, October 5.

Hainesia lythri (Desm.) Hoehn. on Ruhus deliciosus Torr.
(cult.). Dane Co., Madison, September 22.

COLLETOTRICHUM MADISONENSIS H. C. Greene on Carex comosa
Boott. Jefferson Co., Hope Lake Bog near Cambridge, July 28, 1956,
Coll. H. H. litis. On C. rostrata Stokes. Sawyer Co., Flambeau State
Forest, June 24, 1959. Coll. E. Beals. On C. vulpinoidea Michx.
Waupaca Co., Clintonville, July 28, 1959. Coll. K. D. Rill.

Colletotrichum helianthi J. J. Davis on Helianthus tuberosus
L. Sauk Co., near Leland, June 16.

Cylindrosporium rubi Ell. & Morg. on Ruhus odoratus L.
(cult.). Dane Co., Madison, October 5.

Monochaetia discosioides (Ell. & Ev.) Sacc. on Rosa rugosa
Thunb. (cult.). Dane Co., Madison, October 5. Considerable uncer¬
tainty attaches to the nomenclature of these forms, but this is the
same entity reported as M. discosioides on native roses in Wis¬
consin.

Myrioconium comitatum j. j, Davis var. salicarium Davis on
Salix petiolaris Smith. Ozaukee Co., Cedarburg and Waukesha Co.,
Big Bend. Both specimens collected by Davis in June 1930, but not
reported and overlooked until recently.

Monilinia fructicola (Wint.) Honey. Monilia stage on fruit of
Prunus nigra Ait. Sauk Co., near Leland, August 24.

Ramularia canadensis Ell. and Ev. on Carex normalis Mack,
Sauk Co., near Denzer, July 31.

Cercospora caricis Oud. on Carex alopecoidea Tuckerm. Port-
i age Co., near Junction City, August 21, 1953. Coll. G. Ware.

Cercospora oxalidiphila Chupp & Muller on Oxalis europea
i Jord. Sauk Co., near Leland, August 12, 1964.

164 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Cercospora umbrata Ell. & Holw. on Bidens coronata (L.)
Britt. Columbia Co., Gibraltar Rock County Park, August 13, 1964.
In small amount, associated with Cercospora hidentis Tharp.

Tuberculina persicina (Ditm.) Sacc. on Gymnosporangium
juniperi-virginianae Schw. Ill on Juniperus virginiana L, Dane
Co., Madison, Picnic Point, May 21. Coll. & det. J. L. Cunningham.
The spore horns have been aborted and replaced by the sporodochia
of Tuberculina, The first Wisconsin record on telia of a heteroecius
rust.

Additional Species

The fungi mentioned have not been previously reported as occur¬
ring in Wisconsin.

Gibberidea abundans (Dobr.) Shear on Prunella vulgaris L.
Sauk Co., near Leland, August 18, 1962. In my Notes 29 this was
erroneously reported as Linospora brunellae Ell. & Ev., a species
which, so far as I am now aware, has not yet been found in Wis¬
consin or elsewhere in the Midwest.

USTILAGO TREBOUXii H. & P. Sydow on Panicum virgatum L.
Dane Co., Madison, July 7. A large clump of this grass in a garden
on the University of Wisconsin Campus was heavily infected. The
plant was moved several years ago from a spot in Sauk Co., near
Lone Rock, Wis. According to Fischer, U. trebouxii is fairly wide¬
spread on various grasses in the western United States, but he does
not list P. virgatum as a host. The U. S. D. A. Index of Plant Dis¬
eases does mention it (as U. underwoodii Zundel) as occurring on
P. virgatum in New York State.

Phyllosticta cystopteridis sp. nov.

Maculis obscuro-brunnei'S, immarginatis, plerumque in pinnulis
totis; pycnidiis sparsis, pallido-brunneis, muris tenuibus translu-
cidisque, subglobosis, indistincte ostiolatis, mensuriis variis, ca,
(100-) 125-150 (-225) y, diam. ; conidiis hyalinis, angusto-cylindra-
ceis vel subfusoideis, plerumque biguttulatis, (6-) 8-10 (-12) x
(1.5-)1.7-2.5(-3) y.

Lesions dull brown, immarginate, usually involving entire pin¬
nules ; pycnidia scattered, pallid brownish, thin- walled and translu¬
cent, subglobose, obscurely ostiolate, variable in size, approx.
(100-) 125-150 (-225) y diam.; conidia hyaline, narrowly cylindric
or subfusoid, mostly biguttulate, (6-) 8-10 (-12) x (1.5) 1.7-2. 5
(-3) y.

1966]

Greene — Wisconsin Parasitic Fungi

165

On living leaves of Cystopteris fragilis (L.) Bernh. Gov, Dodge
State Park near Dodgeville, Iowa County, Wisconsin, U. S. A., July
14, 1965.

A very sizable type specimen was obtained and in a number of
mounts made from it no conidia with septa were seen, despite the
ratio on length to width. Although few sphaeropsidaceous fungi
have been reported on ferns, the writer’s experience would indicate
that they are not so very uncommon on these hosts.

Phyllosticta argillacea Bres. on Ruhus strigosus Michx. Sauk
Co., “Hemlock Draw” near Leland, August 14. Since 1958 nineteen
specimens of this fungus have been collected .Twelve are on R. alle~
gheniensis Porter, all collected in the Madison School Forest near
Verona, Dane Co., and 4 on R. occidentalis L., one from Madison,
one from Abraham’s Woods near Albany, Green Co., and two from
Gov. Dodge State Park, Iowa Co. An interesting example of a
fungus, first described in 1894 on the cultivated European rasp¬
berry, R. idaeus L. and of which a number of European exsiccati
have been distributed, now apparently reported for the first time
from North America, yet widespread in southern Wisconsin for
almost a decade on native Ruhus, {R. strigosus, it should be noted,
is closely related to R. idaeus and by some is considered to be but a
variety of it) . J. J. Davis in his long collecting career seems not
to have found this fungus, nor did the writer prior to 1958.
Although the pycnidia are flesh-colored and difficult to discern ex¬
cept by transmitted light, the host lesions are very noticeable and
it seems unlikely that the organism could have escaped attention
over the years had it been present in any considerable amount.

Macrophoma farlowiana (Viala & Sauv.) Tassi on Vitis aes¬
tivalis Michx. Dane Co., near Verona, September 14, 1964.

Ascochyta leonuri EIL & Dearn. on Leonurus cardiaca L. It
appears that various Wisconsin collections on this host which were
referred to Ascochyta nepetae J. J. Davis are better placed in A,
leonuri because the conidial dimensions at their upper limit corre¬
spond to those of the latter species and are out of the range of A.
nepetae.

Stagonospora trifidae sp. nov.

Maculis nigris, irregularibus et indefinitis, saepe magnis; pyc-
nidiis hypophylliis, spar sis vel gregariis, vel confertis in venis
primis; flavido-brunneis, muris tenuibus, subglobosis, ca. (125-)
150-175 fx diam., conidiis hyalinis, obtusis, cylindaceis vel subcy-
lindraceis. 3-4-septatis, saepe guttulatis, (20) 23-33 (-37) x 7-8

(-10) IX.

166 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Spots black, irregular and indefinite, often large; pycnidia
hypophyllous, scattered to gregarious, or crowded on the principal
veins, yellowish-brown, thin- walled, subglobose, approx. (125~)150
-175 ft diam. ; conidia hyaline, obtuse, cylindric or subcylindric, 3-4-
septate, often guttulate, (20-) 23-33 (-37) x 7-8 (-10) ft.

On living leaves of Ambrosia triflda L. collected in the East
Marsh of the University of Wisconsin Arboretum at Madison, Dane
County, Wisconsin, U. S. A., September 3, 1965.

Stagonospora ambrosiae Savile (Mycologia 38: 453. 1946) was
on lesions primarily produced by Entyloma compositarum and has
narrow conidia 10-33 x 2. 5-3. 5 ft.

Septoria liquidambaris Cooke & Ell. on Hamamelis virginiana
L. Sauk Co., “Hemlock Draw” near Leland, August 31, 1964. On
Liquidambar in the specimens that I examined, including N. Amer.
Fungi 530, the fungus is hypophyllous, contrary to the statement
by Cooke and Ellis that it is epiphyllous. On Hamamelis, however,
the fruiting structures are definitely epiphyllous and most, but not
all, are Cylindrosporium-like and compressed by the cells of the
palisade layer. In Liquidambar the more loosely organized meso-
phyll tissue allows better pycnidial development. The spores are
quite characteristic and very similar on both hosts. Instead of
scolecospores, a few of the rather imperfect pycnidia on Hamemelis
contain microconidia, about 3 x 1 ft. On Hamamelis the lesions are
very striking, with the dark brown spots surrounded by a brilliant
salmon-colored halo.

Botrytis byssoidea J. C. Walker on Allium cepa (cult.). Racine
Co., Racine, 1918. Coll. Walker. This should have been included in
the earlier Wisconsin lists since, as described by Walker (Phyto¬
path, 15: 708-713. 1925), it is definitely parasitic on onion bulbs.

THE PRESETTLEMENT VEGETATION OF IOWA COUNTY,
WISCONSIN

Wayne J. Stroessner and James R. Haheck^'

Introduction

A istudy of the presettlement vegetation of Iowa County, Wis¬
consin (Fig, 1), was initiated for the purpose of determining the
variety and distribution of vegetation types at the time of .white
settlement in the 1830’s, The original land survey records were
employed in this study in the same manner as other workers have
in other investigations of presettlement Wisconsin vegetation
(Ellarson 1949, Coder 1956, Neuensch wander 1956, and Finley
1951),

Iowa County (Fig, 1) lies in the unglaciated, southwestern
corner of Wisconsin (90°00" W, 43°00' N). The county is bordered
on the north by the Wisconsin River, which flows westward and
joins with the Mississippi River. One of the most conspicuous
physiographic features of the county is the occurrence of the Mili¬
tary Ridge, which is an elevated ridge of Galena limestone extend¬
ing across Iowa County in an east-west direction. The Military
Ridge dissects the county into two nearly equal halves, a north
and south section, which are markedly differentiated from one an¬
other in climate, geology, soils, and vegetation. Detailed descrip¬
tions of this interesting feature will be discussed in a later section
of this paper.

Early settlers were first attracted to the Iowa County area by
the presence of lead and zinc deposits occurring throughout much
of southwestern Wisconsin. Miners and other white settlers were
well established in this region by the late 1820’s. Mining remained
the major occupation of the first settlers for many years, although
in the later decades, during the 1840’s and 1850’s and up to the
present time, agriculture became increasingly more important in
the county’s economy.

The vegetation in Iowa County, as well as elsewhere in Wiscon¬
sin, has no doubt been markedly influenced by man’s activities for
many hundreds of years, beginning with the first primitive Indian
tribes in southern Wisconsin. Indians are known to have exerted

* The senior author is currently a biolog'y instructor at Monroe Senior High School,
Monroe, Wisconsin ; the junior author is Associate Professor of Botany at Montana
State University.

167

168 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

a significant influence on the native vegetation through their regu-
lar use of fire. Through the purposeful and accidental use of fire, |
the Indian is thought to have been an effective tool in the establish- f
ment and maintenance of grassland and oak opening vegetation in -
many areas in southern Wisconsin. Curtis (1959) discusses the f
role of early Indians in Wisconsin in affecting the vegetation of ffi
this region. S

1966] Stroessner and Habeck — Vegetation of loiva County 169

Miners and early agriculturalists in Iowa County also initiated
their own influence on the vegetation as soon as they arrived. Min¬
ing activities no doubt created a need for timber for mine shaft
construction and other uses. Oak forests in southwestern Wiscon¬
sin supplied timber for the mining industry. The prairies and oak
openings prevalent in the southern half of Iowa County were the
first attraction to farmers settling in this area, since these areas
did not require clearing the land.

Since the mid-nineteenth century, when the area was well settled,
Iowa County vegetation has been subjected to a wide variety of
uses and misuses. Some forests have been cut to various degrees,
some of the most select hardwoods have been used for construction
of railroads, some have been burned, and others have been used as
pastures. Very few areas of vegetation have remained undisturbed
since the time of settlement. An analysis of changes in the Iowa
County vegetation during the past 130 years will be the subject of
future investigation. This present report will confine itself to a
detailed description of the vegetation before its severe alteration
by white man.

The Original Land Survey Records

One of the most reliable sources for determination of the type
of vegetation present during the presettlement period is the invalu¬
able records written by the surveyors during the original land
survey of the Northwest Territory made during the 1830’s.

Most of the surveying was done between 1831 and 1834. It is
assumed that most of this work was not very enjoyable, since the
survey crew had to walk approximately 100 miles through un¬
inhabited country for each township surveyed. Not only did they
have to contend with the natural elements, but they also had to
be on the lookout for Indians, In May and June of 1832, there were
several fierce encounters with several of the Indian tribes during
which many people were killed, scalped, and sometimes decapitated.

Even though the surveyors’ records were not intended to describe
the vegetation in great detail, much quantitative and qualitative
material can be extrapolated from these field notes. The data which
have been most valuable for determining the type of vegetation
present during the presettlement period include not only the de¬
scription of the vegetation, but also the information given for the
witness trees. From these data one is able to establish fairly accur¬
ately the type, distribution, density, and the basal area of a stand
of trees ; and, in turn, one can determine the relative frequency, the
relative density, and the relative dominance of either a single forest,
a township, or the entire county. Importan'ce values can easily be
obtained from these known quantities.

170 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

The relative frequency is described as the number of occurrences
at section and quarter section corners of one species as a percentage
of the total number of occurrences of all species at the corners.
The relative density is the number of individuals of the species tab¬
ulated as a percentage of the number of individuals of all species
tabulated within a prescribed area. The relative dominance is the
basal area of the species in comparison to the total basal area
of all species expressed in per cent. The importance value is only
relative and is determined by summing the relative frequency, the
relative density, and the relative dominance of each species. This
technique is described by Ward (1956) .

Discussion of Field Notes

A summary of the data derived from the field notes is provided
in Tables 1 and 2. Table 1 includes only those trees that were used
as witness trees, either at quarter section or corner section posts.
Table 2 is a tabulation of trees that were found directly on the
survey line. Since the trees were directly on the line, it was not pos¬
sible to obtain any absolute values. Absolute values as well as some
relative values of these trees could serve no significant function,
but their presence was of value in determining whether an area
was or was not heavily populated with trees.

From these two tables it is easy to note that most of the trees
in Iowa County are species of oak (Qiiercus). White and bur oak
(“Burr Oak” as recorded in the original field notes) are by far
the most dominant trees, comprising approximately three-quarters
of all trees listed. All of the oaks combined occupy 95.8% of all
trees used as witness trees. The common names of the trees are
listed here as found in the field notes, along with their probable
current scientific names.

Because most of the surveying was done during the winter
months, the exact identification of some of the trees may be ques¬
tioned. Six different oaks were listed in the surveyors’ records. It
is possible that yellow oak and black oak are the same species,
Quercus velutina (Ellarson 1949), although there are several in¬
stances in which the surveyors used both common names even at
the same site, indicating that the surveyors recognized differences
between the two. The following entry is one of many by Sylvester
Sibley, implying that yellow oak and black oak are two different
species :

North between sect 22 and 23

SO'.OO ' Y. oak 8 S 87 W 49

B. oak 7 S 37X E 31

1966] Stroessner and Haheck — Vegetation of Iowa County 171

Table 1.

Trees Used as Witness Trees in Iowa
County During the 1830's

No.

of

Trees

Tot.

No. of
Quart.

Rel.

Freq.

%

Rel.

Dens.

%

Rel.

Dom.

%

Imp.

Value

White Oak

Quercus alba L .

1, 524

1, 004

37.7

39.8

52.2

129.7

Burr Oak

Quercus macrocarpa Michx .

1 253

820

30.8

32.9

23.5

87. 1

Yellow Oak

Quercus muhlenbergii Engelm .

576

427

16.0

15.1

13.5

44.6

24.3

.4

Blaek Oak

Quercus velutina Lam .

305

245

9.2

8.0

7.1

. 1

Finn Oak

Quercus ellipsoidalis E. J. Hill .

5

4

.2

. 1

Red Oak

Quercus borealis Michx. f .

2

2

. 1

. 1

. 1

.3

Hickory

Cary a spp .

39

39

1.5

1 .0

.5

3.0

Elm

Ulmus spp .

18

18

.7

. 5

.8

2.0

Aspen

Populus spp .

25

25

.9

.7

.3

1 .9

Sugar (Maple)

Acer saccharum Marsh .

13

13

.5

.3

.7

1.5

Lynn (Basswood)

Tilia americana L .

10

10

.4

.3

.2

.9

Willow

Salix spp .

8

7

.3

.2

. 1

.6

Cherry

Prunus spp .

3

3

. 1

. 1

. 1

.3

Yellow Pine

Pinus resinosa Ait .

9

9

.3

.2

.2

.7

White Birch

Betula papyri/era Marsh .

7

7

.3

.2

. 1

.6

Yellow Birch

Betula lutea Michx. / .

10

10

.4

.3

.2

.9

I ronwood

Ostrya spp. Scop .

2

2

. 1

. 1

.0

.2

White Ash

Fraxinus americana L .

11

11

.4

.3

.2

.9

Black Ash

Fraxinus nigra Marsh .

3

3

. 1

. 1

.1

.3

Plum

Prunus spp .

1

1

.0

.0

.0

.0

Hackberry

Celtis occidentalis L .

1

1

.0

.0

.0

.0

Totals .

3, 825

2, 661

100.0

100.2

100.0

300.2

172 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Table 2.

Species

Trees Found Directly on Survey Lines

IN Iowa County During the 1830's

No. of
Trees

Total Basal
Area

Rel. Dens.

%

Rel. Dom.

%

White Oak

Quercus alba L .

234

30, 050

52.9

54.2

Burr Oak

(Quercus macrocar (ya Michx .

5 3

5, 231

12.0

9.4

Yellow Oak

(Quercus muhlenbergii Engelm .

108

14,365

24.4

25.9

Black Oak

Quercus velutina Lam .

26

3, 327

5.9

6.0

Pinn Oak

Quercus ellipsoidalis E. J . Hill .

1

64

.2

. 1

Red Oak

Quercus borealis Michx. f .

1

201

.2

.4

Hickory

6

283

1 .4

.5

Elm

Ulmus spp .

4

917

.9

1 .7

Aspen

Populus spp .

3

208

.7

.4

Sugar (Maple)

Acer saccharum Marsh .

2

355

. 5

.6

Lynn (Basswood)

'TiJin nmprirnnn I .

—

Willow

Salix spp .

—

—

Cherry

PrunuM ^pp

—

Yellow Pine

Pinus resinosa Ait .

2

100

.5

.2

White Birch

Pell tin papyri fern Marsh

—

Yellow Birch

Betula lutea Michx. f .

1

254

.2

.1

I ronwood

O.’itr'Da spp Sr.ap

—

White Ash

Fraxinus americana L .

1

79

.2

. 1

Black Ash

nl(frn A/fnrxh

—

—

Plum

Prunus spp .

—

—

Hackberry

C'.pJtidi nrriripntnl ix f

—

—

Totals .

442

55, 443

100.0

100.0

1966] Stroessner and Habeck — Vegetation of Iowa County 173

The ‘‘B. oak” is a black oak, even though it may be misinterpreted
by some to be a bur oak. Sibley was consistent in his entries and
used the '‘B,” abbreviation only to represent black oak; if the “B.”
had referred to bur oak, he would not have used the method of
recording shown below:

North bet sect 4 and 5
4.00 Stream
19.20 Stream
40.00 B oak 7

Burr oak 8

S A'jA W 56

N 82 E 2.95

also :

West bet sect 16 and 21

39.92 Burr oak 4 S

B. oak 6 N

3

11

W .9

W 62

In the first example the B. oak is listed first. If it had been a bur
oak, he would not have written the entries in two different ways
and he probably would have used ‘‘Do” (ditto) to represent the
other witness tree. In the second example he again would have
used “Do” to represent the second witness tree.

In this study “Yellow Oak” is considered to be Quercus muhlen-
bergii, since this species is commonly called either Chinquapin Oak
or Yellow Oak (Grimm 1957). A few of these trees can still be
found in Iowa County, generally along the edges of bottomland
forests.

Scrub oak was not listed as a witness tree even though it was
often recorded as the main type of vegetation in the undergrowth.
In a few areas pine trees were found. These relict pine stands are
still found today and have been described by McIntosh (1950). At
the edges of some clearings and near some mine diggings and
along some of the streams, a few aspens were recorded. Hickory
trees were sparsely scattered throughout the entire county. Along
the bottoms of the Wisconsin River the composition of the wooded
areas changed somewhat; yellow oak became more abundant, and
white oak became less prominent. A greater variety of species was
found in the wooded areas along the river bottoms; namely elm,
lynn (Tilia), and maple occurred frequently, with white ash, black
ash, yellow birch, plum, and hackberry also present.

North of the center of the county in Township 7 North, Range 3
East (Fig. 2), some stands were quite dense in contrast to those in
most of the other townships ; however, this township did not have
as large a number of trees used as witness trees or as great a total
basal area as did several other townships.

174 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Of approximately 2,400 points that were used as quarter section
or corner section markers in the entire Iowa County

367 points were in the open prairie. No trees were listed, but
mounds were built to mark the locations of the posts;

104 points had only one tree listed, usually because the crew was
in a prairie, and as a result no other trees were available;
or because the trees were at the north boundary of the
county along the Wisconsin River, in which case only one
tree’s diameter was usually given;

31 points were not listed properly or were not listed at all. On
some occasions only the diameters were given without the
distances, or the distances and not the diameters ;

15 points had trees located exactly at the spot where the stake
was to be driven ; hence only one tree was listed, giving the
name and diameter; and

3 points were in streams or in a lake, and again no trees were
listed.

This indicates that the remaining 1,880 of the 2,400 points were
used to determine the types of forest cover present in Iowa County.

Method for Construction of a Map from Survey Records

To determine the nature of the vegetation present in Iowa County
at the time of settlement, it was believed useful to construct an
accurate map for that period. It seemed evident after preliminary
investigation that the most suitable and accurate source of infor¬
mation is the record of the original land survey. Since the original
surveyors’ field notebooks could not be removed from the Public
Land Office, it was necessary to transcribe all needed information
directly from the field notes. Descriptions of vegetation vary con¬
siderably from one surveyor to the next. It became apparent that
there were very few '‘dense” stands of trees anywhere in the entire
county, and it was difficult to determine precisely the density of
these wooded areas. All surveyors apparently agreed on the descrip¬
tions of the prairies, since no trees were available as witness trees
within a grassland area, as shown in the following entry :

North bet sections 1 1 and 12

40.00 when raised a mound for X section corner

56.50 Stream 9 S.E.

57.40 Spring brook 4 S.W.

80.00 Where raised a mound for cor sections 1,2, 11, 12
Land 1st rate rolling prairie

It is considerably more difficult to determine the density of cer¬
tain forests, since the descriptive terminology is general and vari¬
able. For example: “Very thinly timbered with . . “very thinly
scattered with . . and “broken prairie with some timber . .

1966] Stroessner and Habeck — Vegetation of loiva County 175

usually meant that this area was prairie or opening. When the
terminology is ‘'thinly timbered . . “tolerably well timbered with
. . or “barrens'", it is very difficult to ascertain exactly what
was meant by the description; hence it seemed that some method
was necessary to convert the available data in the records to quan¬
titative values to determine the densities of various stands in the
county. In order to accomplish this, the diameters and basal areas,
as recorded by the surveyors, were transferred to specially designed
tally sheets with a color code for identifying the density for the
various stands of trees. Usually two trees were selected as witness
trees at each site. Occasionally a surveyor would list four trees at
a section corner. Two of the four trees were insufficiently described
for use in this study, since only their diameters and the sections
in which they were located are listed.

Because of minor discrepancies in comparisons of surveyor de¬
scriptions and quantitative values determined from the spacing
of the witness trees as recorded on the tally sheets, a differentiation
was made concerning terminology used by the surveyors in describ¬
ing “prairie”, “oak opening” and “oak forest”. To distinguish be¬
tween these three classifications, it was necessary to convert the
quantitative values from the field notes to some usable values. The
arbitrary values ,were established in order to have consistency. The
basic values were derived from descriptions of communities already
used by other authorities in the field. In the study made by Curtis
and others (Curtis 1959), a minimum of one tree per acre separates
savanna from prairie. Brown (1950) considered areas with 2 to 8
trees per acre with an average distance of about 100 feet between
pine trees to be pine savannas. The “oak openings” used in this
project are equivalent to the “savannas” mentioned above. Habeck
(1961) differentiates between “oak opening” with trees 50 feet
or more apart and “oak forest” with trees less than 50 feet apart.
The problems associated with the definition and recognition of
savanna vegetation are discussed in detail by Dyksterhuis (1957).
To establish values similar to those used in other forest community
studies, and for the convenience of converting the survey data into
usable form on the tally sheets for the construction of the vegeta¬
tion map, the following vegetational divisions were established to
differentiate between various tree densities.

Prairie: Prairies are defined as having less than one tree per
acre. It is necessary to have trees separated by a minimum distance
of 209 feet in order to have less than one tree per acre. This value
can easily be obtained from the field notes by determining the
distance from the point to the two witness trees. It has been found
that the average link distance from post to tree gives a reasonably

176 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

close value for the average distance in feet between the survey
trees (Cottam 1953) . Therefore an average distance that is greater
than or equal to 209 links from the point to the witness tree indi¬
cates that the area is prairie. In this study, if either witness tree
is more than 300 links from the point, the area is also considered
to be prairie. This serves as a correction factor for any situation
in which, by chance, one of the trees happens to be very near to
the post and the other one extremely far away.

Oak Opening: The oak opening is characterized by having from
50 to 209 feet between trees or an average of from 50 to 209 links
from the point to the two witness trees. These values were selected
to correspond with those selected by other authorities. Simple
mathematics shows that such an area has from one tree per acre
to 17.4 trees per acre. If any of the witness trees are located at a
distance greater than 80 links from the point, then this area might
also be considered as oak opening.

Oak Forest: An oak forest has a density of 17.4 or more trees
per acre. This means that there are fewer than 50 feet between trees
or an average distance of less than 50 links from the point to the
two witness trees.

Bottomland Forest: Bottomland forests are found along river
bottoms and near marshy areas. The trees found in this area are
the same as those mentioned earlier in this paper.

On the map (Fig. 2), differentiation is made between '‘upland
prairie’’ and “wet prairie and marsh”. This division of the two
types of prairies is necessary in order to distinguish between the
various types of plant communities generally associated with each
habitat.

In the preparation of the map, three preliminary maps were
constructed by various methods. One map was made from the quan¬
titative data which was organized on the survey record tally sheets.
A second map was made by using the surveyors’ qualitative descrip¬
tions of the vegetation. At the end of each mile covered by the sur¬
vey team, the surveyor entered a brief written description of the
topography and vegetation of the land similar to the following
notation :

North bet sect 33 & 34
7.50 Stream 8 east

8.30 Road E &z W about 8 chs. E from this line is a recently evacuated
log house

40.00 set mile post

B. Oak 12 S 59 E 4.16

Do 12 Marked XS S 3 W 3.46

44.00 enter prairie

80.00 Where raised a mound in prairie cor to sect 27, 28, 33, 34
Land 1st mile 2nd rate hilly & thinly timbered. Oak and
2nd mile 1st rate rolling prairie

1966] Stroessner and Habeck — Vegetation of loiva County 177

(^UPLAND PRAIRIE ^ WET PRAIRIE a MARSH

Figure 2. Map of presettlement vegetation in Iowa County, Wisconsin. The dis¬
tribution of the five major vegetation types in the county as well as the loca¬
tion of the Military Ridge are illustrated. The Range and Township lines en¬
close areas which are each 36 square miles.

For the construction of this latter map, a color key similar to the
one used for the first map was used. The divisions for the vegeta¬
tion types were grouped according to the descriptions used by the
surveyors :

Prairie — Described as “prairie”.

Oak Opening — Described as “scattered”, “thinly timbered”, “barren”,
“well timbered”, “reasonably well timbered”, and “forest” (only when
witness trees were distantly located).

Oak Forest — Described as “forest” (providing witness trees were quite
close together), “heavy” or “dense forest”, and “dense thicket”.

178 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Other terms which are descriptive of the areas surveyed are:
“bottoms'’ — referring to the lowland regions along a river or
stream bed, and “marsh” — which is self-explanatory.

According to Mr, Tester Bakken of the Public Land Office, Madi¬
son, Wisconsin, governmental personnel in Washington, D. C., drew
up maps of each township after the surveyors’ notebooks were
completed and sent to Washington. Each map was constructed from
the original field notes, which usually included a page containing a
very brief and simplified sketch of the township made by the sur¬
veyor while he was collecting data in the field. On some occasions
the map was quite different from the surveyor’s sketch, but in the
final analysis, the resulting maps corresponded quite closely to
present maps of the same area. A third map showing the results
of their findings was prepared by photographing and reproducing
their maps so that a clearer outline for this study could be made.
By combining the observable features of the above three maps, a
fourth and final map was constructed, which contains a compila¬
tion of all of the material and original information available con¬
cerning the presettlement period.

Discussion and Conclusions

The northern half of Iowa County is by far the most heavily
forested portion. It is separated from the southern half by
stretches of true prairie where the old “Military Ridge” road was
once located. Military Ridge is the trail which army troops and
supplies once traversed in the late 1800’s when traveling between
Madison and various military points in the western part of the
state, Iowa, and Minnesota. The terrain is quite level, the land is
high, and streams are not numerous ; therefore east-west traveling
was not difficult across this stretch of land.

All sizable streams and rivers recorded by the surveyors are
shown on the map. The location and size of these streams compare
quite favorably with present maps of the same areas. Therefore it
can be assumed that not only the listings of the rivers and streams
were accurate, but the data concerning witness trees and the de¬
scriptions of the land may be assumed to be accurate also.

Most forested areas are near a river or stream. The trellis drain¬
age system, characteristic of both the northern and southern halves
of the county, apparently does not permit a suitably moist environ¬
ment for forest development in the central portion. The southern
half of the county has more acreage in marsh than does the north¬
ern half. Some oak forests are found in the southern half, but the
stands are few and small in comparison to the forests of the north¬
ern half. The surveyors generally described the southern portion
as rolling prairie with only a few hilly areas, whereas the northern

1966] Stroessner and Habeck — Vegetation of lotva County 179

portion was generally described as rather uneven or rolling and
hilly. Some rather deep ravines were found along some of the
rivers and streams in the northern part of the county, but most of
the river banks in the southern portion apparently were not as
prominent. In the extreme northern portions of the county the
lowlands or river bottoms were generally described as flat, with
‘‘first rate soil” where there was prairie, or “second rate soil”
where trees were thinly scattered.

Throughout the county, oaks are by far the most prevalent spe¬
cies except in the lowland regions along the Wisconsin River on
the northern border. In these lowland areas a larger variety of
other hardwood species appear, as has been described earlier. Up¬
land prairies, which occupy a major portion of the southern half
of the county, are nearly absent from the northern half.

Some of the first settlers to inhabit Iowa County were miners,
the first mine operations being located in the southern part. Shortly
after the miners moved into the county in the latter part of the
1820’s, some of the first homesteads became established. Because
of the occurrence of upland prairie, farming in this southern por¬
tion of the county was perhaps much easier to initiate than in the
northern half, since much of the land did not necessitate forest
clearing before crops could be planted.

It is concluded that Iowa County during the 1830’s consisted of
large areas of prairie, oak opening, and oak forests. Very few other
species of trees or types of vegetation were present in the county
during the presettlement period. What has happened to these areas
since presettlement will be a subject of future investigation.

References Cited

Anon. 1881. History of Iowa County, Wisconsin. Western Historical Company
— Chicago 970 pp.

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.
Brown, R. T. 1950. Forests of the Central Wisconsin Sand Plains. Bull. Ecol.
Soc. Am. 31:56.

CoTTAM, G. and J. T. Curtis. 1956. The Use of Distance Measures in Phytoso-
ciological Sampling. Ecology. 37(3) :451-460.

- , - , and B. W. Hale. 1953. Some Sampling Characteristics of a

Population of Randomly Dispersed Individuals. Ecology. 34:741-757.
Curtis, J. T, 1950. Original Forest Structure. Paper at AAAS Symposium on
structure on Plant Communities. Cleveland. 30 Dec.

- , 1959. The Vegetation of Wisconsin. University of Wisconsin Press.

657 pp.

Dyksterhuis, E. J, 1957. The Savannah Concept and Its Use. Ecology. 38:435-
442.

Ellarson, R. S. 1949. The Vegetation of Dane County, Wisconsin in 1835.
Trans. Wis. Acad. Sci., Arts and Letters. 39:21-45.

180 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

Finley, R. W. 1951. The Original Vegetation Cover of Wisconsin, An unpub¬
lished doctoral dissertation.

Gleason, H. A., and A. Cronquist. 1963. Manual of Vascular Plants of North¬
eastern United States and Adjacent Canada. D. Van Nostrand Company,
Inc. Princeton, New Jersey,

Gorder, H. a. 1956. Pre-settlement Vegetation of Racine County. Trans. Wis.
Acad. ScL, Arts and Letters. 45:169-176.

Grimm, W. C. 1957, The Book of Trees. The Stackpole Company, Harrisburg,
Pennsylvania.

Habeck, J. R. 1959. A Phytosociological Study of the Upland Forest Commu¬
nities in the Central Wisconsin Sand Plain Area. Trans. Wis. Acad, of
Sci., Arts and Letters. 48:31-48.

- . 1961. The Original Vegetation of the Mid- Willamette Valley, Oregon.

Northwest Science. 35(2) :65-77.

- . 1962. Forest Succession in Monmouth Township, Polk County, Oregon

Since 1850. Proceedings of the Montana Academy of Sciences. 21:7-17.

McIntosh, R. P. 1950. Pine Stands in Southwestern Wisconsin. Trans. Wis.
Acad. Sci., Arts and Letters. 40:243-257.

Neuenschwander, H. E. 1956. The Vegetation of Dodge County, Wisconsin
1833-1837. Trans. Wis. Acad. Sci., Arts and Letters. 46:233-254.

POTZGER, J. E., M. E. POTZGER, and J. McCormick. 1956. The Forest Primeval
of Indiana as Recorded in the Original U. S, Land Surveys and an Evalu¬
ation of Previous Interpretations of Indiana Vegetation. Butler Univer¬
sity Botanical Studies. Butler University, Indianapolis 7, Ind. 13(1) :95-
111.

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.

REAPPRAISAL OF THE GROWTH POTENTIAL OF JACK PINE AND
RED PINES ON DIFFERENT SOILS OF WISCONSIN*

S. A, Wilde, R. R. Maeglin, and Ch. Tanzer'

Under the influence of MayUs monograph (1890) and other re¬
ports dealing with natural forest distribution, foresters of the
Lake States adopted a credo that jack pine is better suited for re¬
forestation of coarse sandy soils than is red pine. The advanced
growth of Wisconsin plantations of the two tree species provides a
constantly increasing evidence that this thesis has notable excep¬
tions, a disregard of which leads to large losses in the volume of
produced timber. As revealed by soil and mensuration analyses,
the relative performance of jack and red pine does not depend on
soil texture alone, but is strongly influenced by the mineralogical
composition and root permeability of soils.

Years ago, Mr. F, G. Wilson, a member of the Wisconsin Con¬
servation Department, had shown the senior writer windbreaks
established simultaneously on coarse sandy soils in which red pine
produced a much faster growth than did jack pine. A study by
Voigt (1951) explained this ''deviation from the assumed norms of
behavior’' by a higher capacity of the roots of red pine to penetrate
previously farmed soils in which the root channels have undergone
a deterioration. Similar observations were later reported by Wilde
(1961).

The recent survey of Wisconsin plantations has indicated another
digression in the soil-growth relationship of jack and red pine.
On coarse-textured soils enriched in silicate minerals, such as feld¬
spar, mica, and hornblende, red pine shows an appreciably better
performance than does jack pine (Fig. 1).

The survey encountered five pairs of adjacent or closely-located
plantations of the two tree species, supported by feldspathic sandy
soils of nearly identical state of fertility (Plainfield and Omega
sands). The results of the growing stock analyses show that red
pine attained a significantly higher increment than did jack pine,
in some instances yielding nearly a 40% higher volume of wood
(Table 1).

Information of broader significance on the growth potential of
the two species is provided by the average data derived from ran-

1 Contribution from Soil Science Dept., Univ. of Wisconsin, and the U. S. Forest
Products Laboratory, in cooperation and supported in part by the Wisconsin Con¬
servation Department. Publication approved by the director of the Wis. Agric. Expt.
Station.

“ Professor of Forestry, Wood Technologist, and Pi'oject Assistant in Soil Science.

181

182 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Figure 1. Trees of average height and DBH illustrating the
relative performance of 17-year-old plantations of jack pine
and red pine established the same day on Plainfield sand
(Nepco Industrial Forest, Adams County, Wisconsin).

domly surveyed jack pine and red pine plantations (Wilde et al,
1964a and 1964b). The results, given in Table 2, show that red
pine plantations of all three site classes delivered a significantly
higher volume than did jack pine plantations; according to the
weighted average of all plantations, red pine produced 23 per cent
more wood per year in comparison with jack pine.

An examination of the average fertility levels of soils supporting
plantations of different site quality, given in Table 3, permits addi¬
tional inferences.

Table 1. The Growth of Simultaneously Established Jack Pine and Red Pine Plantations on Non-podzolic
Sandy Soils Enriched in Silicate Minerals (Results on Per Acre Basis).

1966] Wilde, Mmglin and Tanzer — Growth Potential

183

Red Pine

Vol.

Cu. Ft.

351

1,365

734

463

2,960

Basal

Area

Sq. Ft.

O oJ 00 o

t\ r-1 00 t\

No. of
Stems

1,580

1,060

800

1,600

940

DBH

Ins.

2.9

4.7

4.5

3.0

5.9

Ht. Ft.

16.9

32.0

27.0

19.8

44.7

Jack Pine

Vol.

Cu. Ft.

307

1,270

461

294

1,909

Basal
Area
Sq. Ft.

CO O O xD

^ sD ^

No. of
Stems

1 ,600

990

830

1,600

960

DBH

Ins.

2.6

4.5

3.6

2.6

5.2

Ht. Ft.

16.5

33.0

24.0

17.5

38.5

Age

Yrs.

16

21

20

17

34

Plantation No.*

12, 13 .

173, 172 .

177, 176 .

181, 189 .

314, 315 .

184 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55
Table 2. Relative Performance of Jack Pine and Red Pine Plantations

ON Non-podzolic and Mildly Podzolized Coarse Sandy Soils Not
Influenced By Ground Water (Results on Per Acre Basis).

Tree Species

Age

Yrs.

Height

Ft.

D3H

In.

No. OF
Trees

Basal
Area
Sq. Ft.

Volume
Cu. Ft.

Ave.
Annual
Growth
Cu. Ft.

Low Site Quality

Jack pine .

Red pine .

22

23

22.0

21.3

3.4

4.0

1,241

1,282

70

101

473

688

21.5

29.9

Med

lUM SiTF

1 Quality

Jack pine .

Red pine .

24

21

30.5

27.5

4. 1
4.8

1,177

978

107

117

1,106

1,300

46. 1
61.9

Hic

:h Site

Quality

Jack pine .

Red pine .

23

20

38.6

32.9

4.6

5.1

1 ,029
1,054

117

136

1,514

1 ,960

65.9

98.0

Weighted

Averag

E OF All

Sites

Jack pine .

Red pine .

23.3

21.7

31.6

27.8

3.9

4.6

1,113

1,052

100

112

1,133

1,377

48.6

63.5

The fertility of soils supporting red pine plantations of low site
quality is definitely higher than that of jack pine plantations. This
is undoubtedly because of the prevailing tendency to plant jack
pine on soils depleted by wind erosion and previous farming. There¬
fore, the increase of the annual increment by 8 cu. feet per acre,
shown by red pine, cannot be attributed to its inherent growth
potential.

On the other hand, the fertility levels of soils supporting planta¬
tions of medium site quality reveal information of practical im¬
portance. The production of extra 16 cu. feet per acre per year,
constituting nearly 35 per cent increase in the annual increment,
was achieved by red pine on soils of a nearly similar average level
of fertility in comparison with soils supporting jack pine planta¬
tions. These results indicate that misoriented tree planting may
deprive the landowner of one-third of the volume of young timber;
and it may inflict a still greater loss at the end of rotation.

Examination of the fertility of soils supporting plantations of
high site quality further emphasizes the critical importance of the
soil productive capacity in reforestation aiming at maximum re-

1966]

Wilde, Maeglin and Tanzer — Growth Potential

185

w Q

Ph <X>
Oi
Q tH

<

a ^

^ H

"I

g g

5 ^

a ■<

O 'W^
Ph

Ph m
P a
w 2

M a

o !=>
w O'

a a
o H

a w
a

a

>

a

<

D

a

DJ

H

U)

Exch.

Mg

S\ l\

vD

r<i ro

bb

o

o

o o

O O

d d

Exch.

Ca

s

o

sD bN.

00

^ —
fS ^

d d

>-< O

" —

J

>

<

w

o

CO

\

^ X)

vtn
xO tx.

^ —
t\ rvi

j

CO ■

J

O' 00

tx

r-i

— H IN.
rN

<

Total

KT

H

-u

CL

.037

.048

.074

.059

O sD

O' o

O O

O

Z £ H
< hU

o o

00 (Nl

O

00

^ s:

— O'

o < .

62“^

d —

ro

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Clay

P. Ct.

6.9

7.3

9.4

8.8

ZTI

001

OQ

u X <

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z ou.

Z Di .

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U

21.5

29.9

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61.9

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Tree

Species

Jack Pine
Red Pine

Jack Pine
Red Pine

Jack Pine
Red Pine

186 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

turns. On soils of slightly higher fertility, red pine produced 32
cubic feet per year or 50% more than did jack pine. Moreover, the
high rate of growth, yielding about one cord per year per acre,
was obtained on soils whose fertility is not greatly superior to that
of soils supporting jack pine plantations of medium site quality.
The difference in the output of the two trees is enormous, consti¬
tuting a debit of 52 cu. feet or a more than 100 per cent loss of
the increment.

The observed superior growth of the red pine on non-podzolic
coarse sandy soils is not necessarily valid on sandy loam soils,
particularly those modified by a podzolization process or derived
from quartzitic parent materials. Occasionally, jack pine on such
soils was found to produce more than a 50% higher increment than
the red pine. Planting jack pine on infertile soils could be justified
only for the purpose of erosion control, soil amelioration, or isola¬
tion of parasites attacking other tree species.

Abstract

On coarse-textured soils enriched in silicate minerals, such as
feldspar, mica and hornblende, red pine, Pinus resinosa, usually
produces appreciably higher yield of timber than does jack pine,
Pinus hanksiana. This is especially true of soils in which prolonged
cultivation or grazing has caused deterioration of root channels.
The superior growth of red pine on non-podzolic coarse sandy soils
is not necessarily valid on sandy loam soils, particularly those
modified by podzolization process, or derived from quartzitic parent
material. As shown by a survey of soils and growing stock, a dis¬
regard of the productive potential of soils and subsequent mis-
oriented planting may lead to the loss of more than 50 per cent
of the yield by either tree species.

References

Mayr, H, 1890. Die Waldungen von Nordamerika. Paul Parey, Berlin,
Voigt, G. K. 1951. Causes of injury to conifers during the winter of 1947-1948
in Wisconsin. Wis. Acad. Sci., Arts and Lett., 49:241-243.

Wilde, S. A. 1961. The soil-ameliorating effect of jack pine (Pinus hanksiana)
and red pine (Pinus resinosa) plantations. In “Recent Advances in Botany:
1631-1635. University of Toronto Press.

Wilde, S. A., J. G. Iyer, Ch. Tanzer, W. L. Trautmann and K, G. Wat-
TERSTON. 1964a, The growth of jack pine {Pinus hanksiana, Lamb.) plan¬
tations in relation to fertility of non-phreatic sandy soils. Soil Sci. 98:
162-169.

- . 1964b. The growth of red pine {Pinus resinosa, Ait.) plantations in

relation to fertility of non-phreatic sandy soils. For. Science, 10:463-470.

- . 1965. Location and ownership of Wisconsin coniferous plantations.

Tech. Notes No. 90. Coll, of Agr., U.W,, Madison, Wis.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 55
COMPOSITAE IV— COMPOSITE FAMILY IV

(Tribes Helenieae and Anthemideae)

Carol J. Mickelson and Hugh H. litis
Herbarium, University of Wisconsin, Madison

The Helenieae and Anthemideae are two of the ten tribes of
the Compositae, the largest family both in the plant kingdom and
in the flora of Wisconsin. The Helenieae, an artificial group segre¬
gated on the basis of the loss of the receptacular chaff, is repre¬
sented by two native North American genera. The Anthemideae,
a rather uniform tribe with most of the species native to the Old
World, is represented in Wisconsin not only by many species that
are introduced cultigens such as daisies and absinthe or weeds
such as dogfennel and pineapple weed, but also by several impor¬
tant North American natives.

Previous portions of the family that have been treated in a pre¬
liminary fashion are the genera Aster (Shinners 1941), Senecio
(Barkley 1963), Solidago (Salamun 1963), and the tribes Helian-
theae (Melchert 1960), Eupatorieae, Veronieae, Cynareae, and
Chicorieae (Johnson & litis 1963), and Inuleae (Beals & Peters
1966).

The present treatment of the Helenieae and Anthemideae of
Wisconsin is based on specimens in the herbaria of the University
of Wisconsin (WIS), University of Wisconsin-Milwaukee (UWM),
Milwaukee Public Museum (MIL), University of Minnesota
(MIN), Chicago Natural History Museum (F), Platteville State
University, and the University of California (UC). Grateful
acknowledgment is due to the curators of the above herbaria for the
loan of their specimens.

Dots on the maps represent exact locations, triangles represent
county records. Some records have been added from Thomas Hart¬
ley’s manuscript '‘Flora of the Driftless Area” (1962), from Paul
Sorensen’s 1965 studies on the plants of Glacial Lake Wisconsin,
and Duluth records from Lakela’s Flora of Northeastern Minnesota
(1965). The numbers in the map insets record flowering and fruit-

Field work and preparation of manuscript supported in part by the Research Com¬
mittee of the University of Wisconsin, on funds from the Wisconsin Alumni Research
Foundation, the printing costs of illustrations in part by the Norman C. Fassett Me¬
morial Fund.

187

188 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

ing dates in Wisconsin. Plants with vegetative growth only, in bud,
or with dispersed fruits are not included. Nomenclature and order
of tribes and genera follows that of Cronquist (1952). A key to
the Wisconsin tribes and genera has been published (Johnson &
litis 1963, pp. 257-262), the sequence and numbering of which is
followed in the present study.

Special thanks are due to the first author’s teacher, Dr. Florence
Neely of Augustana College, Rock Island, Illinois, for an excellent
introduction to botany, to Mr. Brian Marcks for much aid in field
and herbarium, to Dr. Donald Ugent for advice in the preparation
of diagrams, to Dr. John Moore, University of Minnesota, for his
kindness in sending additional locations for map records, and to
Mrs. Katharine Snell and Dr. John W. Thomson for various aids.

TRIBE II. HELENIEAE Benth. & Hook.

An artificial group (cf. Solbrig 1963) segregated on the basis of
the naked receptacles. Bracts herbaceous, usually in one series.
Styles truncate, branched as in the Helianthieae, the stigmatic
lines running to the apex. Leaves alternate or opposite. Pappus of
scales or chaff.

KEY TO GENERA

A. Leaves alternate; bracts free at base; pappus of long-awned
scales; heads globose, conspicuously radiate, the rays yellow

_ 18. HELENIUM.

AA. Leaves opposite; bracts united at base; pappus scales with
numerous, long bristles; heads ishort-radiate ; very rare ad-
ventive _ _ _ _ _ 18a. DYSSODIA.

18. HELENIUM L. Sneezeweed

Heads many-flowered, radiate, the ray flowers yellow, rather
large and showy, pistillate (fertile) or neutral (sterile), cuneate,
3- to 5-cleft, pubescent without. Disks subspherical, the flowers
perfect, 4- to 5-lobed, yellow or dark brown at maturity. Involucral
bracts reflexed at maturity, in 2 or 3 series, the outer longer than
the inner. Receptacle naked, globose. Pappus of 5-8, thin, l-nerved,
awned scales. Achenes 4- to 5-ribbed, the angles pubescent. Annual
or perennial herbs with alternate, usually basally decurrent leaves
covered with bitter, aromatic, resinous dots. Inflorescences few- to
many-headed corymbs.

A Western Hemisphere genus of about 40 species, resembling
sunflowers and blackeyed Susans (Helianthus and Rudbeckia) but
differing in the globose disks and globose, naked receptacles. The

1966]

Mickelson and litis — Flora of Wisconsin

189

resinous granules of Helenium give the plants a bitter taste, the
milk from cows which have eaten it a bitter flavor, appear to
poison livestock which have eaten quantities of it, and cause the
sneezing reaction which gives it its common name (Rock 1957).

Key to Species

A. Leaves lanceolate; stems winged by the decurrent leaf bases.

B. Disk flowers yellow, Sdobed; ray flowers pistillate; cauline

leaves 1-3.5 cm wide; widespread throughout _ _ _

_ _ _ _1. H. AUTUMNALE.

BB. Disk flowers dark brown, 4-lobed; ray flowers neutral;
cauline leaves to 1 cm wide; very rare, in central Wiscon¬
sin _ _ _ 2. H. FLEXUOSUM.

AA. Leaves filiform, less than 2 mm wide ; stems not winged ; disk

yellow; very rare introduced weed _ 3. H. AM ARUM.

1. Helenium autumnale L. Sneezeweed. Map 1.

Perennial, subglabrous or puberulent herbs with basal offshoots
from a stout rootstock. Flowering stems erect, 4-15 dm tall, con¬
spicuously winged by the decurrent leaf bases. Rosette leaves 4-6
cm long, 5-8 mm wide, oblanceolate, entire to shallowly toothed;
cauline leaves 3.5-15 cm long, 1-2 (-4) cm wide, sessile-decurrent,
lanceolate to elliptic, acuminate at apex and base, distantly serru¬
late, or rarely entire, sparsely pubescent and resinous-punctate.
Heads few to many in paniculate corymbs ; involucral bracts
clasping the young head, reflexed at maturity; receptacles naked,
conic-globose. Ray flowers 10-15, pistillate, bright yellow, 8-22
mm long; disk flowers perfect, yellow, 5-lohed, campanulate and
narrowed to a tube just above the achene, 2-3 mm long. Achenes
1-1.5 mm long with 7-8 membranaceous, awned pappus scales.
2n=34 (Darlington 1955).

Abundant in all but* northernmost Wisconsin, in sunny or shady
moist areas such as river bottom floodplain forests, low open woods
with alder, willow, elm, ash, red dogwood, silver maple, and yellow
birch, on sand and gravel bars of rivers and lakeshores, meadows of
Carex, Scirpus, and Juncus, and in low prairies, swales, or marshes
with Eupatorium perfoliatum, Bidens cernua, Aster, Solidago, and
Prunella, according to Curtis (1959) peaking in the wet prairies.
Flowering from the second week in July to mid-October; fruiting
from August through October.

Helenium autumnale var. canaliculatum (Lam.) T. & G., sup¬
posedly distinguished by its smaller size, narrow entire leaves, and
strongly narrowed, often basally channeled ligules, if it occurs in
Wisconsin at all, cannot be clearly segregated. The only linear-
elliptic, entire-leaved extremes that do compare well with some

190 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

St. Lawrence estuary plants of var. canaliculatum are J. J. Davis
SM., Spring Green, Sauk Co., and two collections in 1945 and 1955
by H. C. Greene s.n. from the calcareous Kenosha Prairie, Kenosha
Co. Both of these may well be but xeromorphic forms, perhaps
related to growth on calcareous substrates. Although Fernald lists
two collections from La Crosse and St. Croix Counties (not seen),
all our many collections from that region are clearly typical H.
autumnale.

The Menomini Indians used the dried, nearly mature, pulverized
heads of H. autumnale as a snuff to loosen head colds. The roots
were used by the Meskwaki Indians for medicine, and a tea was
made from the florets for catarrh of the stomach. In Indian lan¬
guages the name for this plant means sneezing spasmodically
(aiatcianitikun) or inhalant (tcatcamosikan, pitcikomate) (Smith
1923).

2. Helenium flexuosum Raf. Purplehead Sneezeweed. Map 2.

Helenium nudiflorum Nutt.

Perennial, puberulent or scaberulous herbs with basal offshoots
from a compact rootstock. Stem erect, solitary, 4-8 dm tall, striate-
sulcate, very prominently winged from the decurrent leaf bases.
Rosette leaves oblanceolate, 4-5 cm long, and 1 cm wide, the cauline
leaves linear-lanceolate to oblanceolate, 3-9 cm long, 0.5-1 cm wide,
sessile-decurrent, entire to irregularly toothed, sparingly pilose,
resin-dotted. Heads 7-12 or more in open paniculate corymbs;
involucral bracts narrowly lanceolate, reflexed at maturity; recep¬
tacle naked, conic. Ray flowers 8-13, neuter, yellow, 1-2 cm long;
disk flowers dark reddish-brown, Jf-lohed (rarely 5), perfect,
campanulate, 2-2.5 mm long. Achenes 1 mm long, with 5 mem¬
branaceous, awned pappus scales.

Centering in the southeastern United States (Rock 1957 :139,
map 3), abundant as far north as southern Illinois and Missouri
in upland pine-oak woods, mesic-moist grass-sedge prairies and
open places, spreading northward as a weed (Rock 1957), in Wis¬
consin known from but four collections, all since 1958, one from
an open flood-plain forest, and three from open sandy or moist
areas in former glacial lake beds, areas well-known for disjunct
stations of Coastal Plain species. Adams Co. : Highway 21 about
3 mi east of highway 13, a sandy, peaty, low, sedge-shrub prairie
with Liatris pycnostachya, Aronia melanocarpa. Lycopodium in-
nundatum, Cladium mariscoides. Spiraea tomentosa, Spiraea alba,
July 1962, litis & Sorensen 3638, (WIS). Jackson Co.: T22N ;
R3W ; Sec. 4, rather moist sandy area bordering highway 54, July
24, 1958, Hartley U^U7 (WIS). Trempealeau Co.: Low moist bot¬
tomland woods on flood-plain terraces between Tank Creek and

1966]

Mickelson and litis— Flora of Wisconsin

191

Black River (T18N ; R8W; sec. 33), with Salix, Solidago, Aster,
Eupatorium, Sept. 5, 1958, litis & Koeppen 11,947 (WIS) . Waupaca
Co.: Open sun, field with clay soil, Clintonville airport. Sept. 16,
1965, Rill 1496 (WIS). Flowering from the end of July to mid-
September; fruiting in September.

3. Helenium am arum (Raf.) Rock Map 2.

Helenium tenui folium Nutt.

Distinguished easily from H. autumnale and H. flexuosum by its
annual habit, linear-filiform leaves not exceeding 2 mm in width,
and small yellow heads. A weedy southern species collected twice
in Wisconsin, clearly adventive and non-persistent. Racine Co. :
C. & N. W. right of way, sec. 14-15, Sept. 11, 1901, Wadmond
13631/2 (MIN). Sheboygan Co.: Sheboygan, waste place. Sept. 11,
1921, Goessl s.n, (WIS).

18a. DYSSODIA Cav.

4. Dyssodia papposa (Vent.) Hitchc. Fetid Marigold. Map 2.

Ill-smelling, glabrous annual with slender taproot; stem 1-5 dm
tall, single or freely branching. Leaves opposite, 1-4 cm long, 1-3
cm wide, pinnatifid into 1 mm wide, linear, toothed lobes, with few
large, pellucid orange glands (these also on involucre). Heads few-
flowered, paniculate ; involucre 5-9 mm high, the outermost bracts
green, free, linear, the inner oblanceolate, reddish brown, and
united at the base. Ray flowers pistillate, 4 mm long, the very short
(1 mm), green ligules scarcely exserted; disk flowers perfect, cam-
panulate, 3 mm long. Achenes pubescent 3-4 mm long; pappus of
numerous, 2-2.5 mm long bristles.

Native of more arid, western regions, adventive east to New
England in disturbed, sandy or open habitats, collected once in
Wisconsin in the sandy flats of the Wisconsin River, an area where
other southern or western species occur (e, g. Diodia teres,
Leptoloma cognatum, Croton glandulosus and C. monanthogynus) .
Iowa Co. : Triangle at junction of U. S. highway 14 and Wise. 23,
Sept. 18, 1965, Murmanis s.n. (WIS).

TRIBE III. ANTHEMIDEAE Cass.

Mostly aromatic herbs or subshrubs; bracts commonly dry-
scarious, imbricate in several series. Receptacle naked, pubescent,
or chaffy. Flowers white, yellow, or green, the outer pistillate or
neuter. Style branches truncate, the anthers not tailed. Leaves
alternate, often finely dissected. Pappus short or absent.

192 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

KEY TO GENERA

A. Receptacle chaffy, the heads radiate,

B. Heads rather large, 1-4 cm wide, solitary and terminal on

long peduncles; receptacle conic at maturity; achenes
terete or angled _ _ ^19. ANTHEMIS.

BB. Heads small, 5 mm or less, densely corymbose; receptacle

flat; achenes compressed _ 20. ACHILLEA.

AA. Receptacle naked or villous, the heads radiate or discoid.

C. Inflorescence corymbose or heads terminal on long
peduncles; ray flowers showy, sometimes obsolete, yellow
or white.

D. Receptacle flat or low-convex.

E. Heads radiate (rarely discoid) ; pappus absent;

achenes 5-10 ribbed _ 21. CHRYSANTHEMUM

EE. Heads discoid or short-radiate ; pappus short-

membranaceous; achenes 3-5 ribbed _

_ ___22. TANACETUM

DD. Receptacle conic at maturity; leaves pinnatisect _

_ _ _ 23. MATRICARIA

CC. Inflorescence paniculate, racemose, or spike-like with
inconspicuous (2-8 mm high), discoid heads; flowers green
_ 24. ARTEMISIA.

19. ANTHEMIS L. Chamomile, Dogfennel.

Heads few to many, solitary on long peduncles, radiate and re¬
sembling a small daisy. Rays ligulate, pistillate or neuter, white or
yellow; disk flowers campanulate, perfect, yellow. Receptacle with
slender, scarious, prominent chaff. Pappus a minute crown or lack¬
ing ; achenes 4- to 5-ribbed, truncate. Aromatic, annual or perennial
herbs with alternate, finely pinnatisect leaves and terminal heads.
About 60 species, native to Europe, Asia, and Africa, ours weedy
introductions.

Key to Species

A. Ray flowers white ; receptacle conic ; involucre 2.5-5 mm high ;
achenes tuberculate (lOx) ; very common, especially southern

Wisconsin _ 1. A, COTULA,

AA. Ray flowers yellow; receptacle flat to shallowly convex; in¬
volucre 5-8 mm high ; achenes smooth ; rare adventive _

_ 2. A, TINCTORIA.

1. ANTHEMIS COTULA L. Dogfennel, Mayweed, Stinking Cotula.

Map 3.

Ill-smelling or aromatic annual, 2-6 dm tall, simple or branching
from a taproot, slightly tomentose when young. Leaves sessile, 2-6

1966]

Mickelson and litis— Flora of Wisconsin

193

i.^S'HELENIUM

FLEXUOSUM
OH. AMARUM
oDYSSODI A
■ PA.PPOSAl

194 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

cm long, 1-4 cm wide, delicately twice-pinnatifid into 0.3-0. 5 mm
wide segments. Heads few to many, radiate; involucre 2.5-5 mm
high; bracts glabrate to tomentose. Receptacle conic at maturity,
with linear-pointed chaff at the tip or covering most of the recep¬
tacle but not subtending the outermost disk flowers. Rays 11-15,
ivhite, neuter or pistillate and sterile, 8-11 mm long; disk flowers
yellow, 3 mm long; achenes prominently tubercled, 1.5-2 mm long;
pappus lacking. 2n=18 (Love & Love 1961).

Native of Eurasia, an abundant weed throughout S. Wis¬
consin, less so northward, in disturbed areas such as cultivated
fields, hog pastures, railroad yards, roadsides, farmyards, lake
shores, and bluffs. Flowering and fruiting from late May through
mid-October.

The quite similar Eurasian Anthemis arvensis L., weedy in E.
United States, but not yet known from Wisconsin, is distinguished
by pleasant odor, fertile ray flowers, chaff subtending all disk
flowers, and ribbed but smooth achenes. The similar, but rather
rare Matricaria chamomilla L. (q.v.) has a smooth, chaffless recep¬
tacle and ribbed but etuberculate achenes.

2. Anthemis tinctoria L. Golden Marguerite, Yellow Cotula.

Map 4.

Aromatic, rhizomatous perennials, with 1-several stems 3-7 dm
tall. Leaves gray-green tomentose, 4-6 cm long, 1-3 cm wide, twice-
pinnatifid into 0.5-1 mm wide lobes. Heads solitary, radiate ; involu¬
cre 5-8 mm high, the bracts tomentose. Receptacle flat to shallowly
convex; chaff linear-lanceolate, persistent and very prominent (ca.
5 mm long), covering the receptacle. Rays 16-51, bright yellow,
pistillate, 9-14 mm long; disk flowers dull-yellow, perfect, 3-4 mm
long ; achenes 2-2.5 mm long, ribbed ; pappus rudimentary. 2n=18
(Darlington 1955).

Native of Europe and western Asia, often cultivated for its
showy, all-yellow flowers, in Wisconsin a rare and sporadic escape
in railroad yards, driveways, roadsides, lake shores and rubbish
piles. The flowers yield a yellow dye (Clapham, et al. 1952) . Flower¬
ing from July through early September; fruiting from July through
September.

20. ACHILLEA L. Yarrow, Milfoil.

Heads many, small, densely corymbose, radiate, the 3-13 rays
short, white or rarely pink, pistillate and fertile ; disk flowers per¬
fect, 10-75. Involucral bracts with dry, scarious margins, imbricate
in 3-4 series. Receptacle chaffy, flat or conic. Achenes compressed,
callous-margined ; pappus absent. Aromatic perennial herbs with

1966]

Mickelson and litis — Flora of Wisconsin

195

alternate, subentire to finely dissected leaves; about 75 species of
the Northern Hemisphere, mostly in the Old World.

Key to Species

A. Leaves finely dissected into linear segments ; plant tomentose ;

ubiquitous throughout _ _ „1. A. MILLEFOLIUM.

AA. Leaves undissected, serrulate; plant glabrate to subglabrous;
rare advent! ve _ _ 2. A. PTARMICA.

la. Achillea millefolium L. ssp. lanulosa (Nutt.) Piper

Common Yarrow, Milfoil. Map 5.

Achillea lanulosa Nutt.

Achillea lanulosa f. Peroutkyi F. S. Seymour, FI. Lincoln
Co. Wis. 331. 1960 (Type: Rays pink, dry field, Rock Falls
Township, Lincoln Co., Wise. July 13, 1954, Seymour
IBySlJp WIS.). Synonymous with f. ruhicunda (Farw.)
Farw.

Villous, strongly aromatic, rhizomatous perennial herbs, 2-8 dm
tall. Leaves 1-2 times pinnatisect into slender, 0.2-1 mm wide, vil¬
lous to glabrescent segments, the basal 13-34 cm long, 2-5 cm
wide, long-petioled, the cauline 6-20 cm long, 1-3 cm wide, sessile.
Corymbs fiat-topped or rounded with many 3.5-6 mm high heads;
bracts linear-lanceolate, with scarious margins; receptacle chaffy,
conic at maturity. Flowers 18-28; rays 4-6, white (or pale pink in
f. RUBICUNDA (Farw.) Farw.), 2-3 mm long; disk flowers
14-22, with a green tube and 5 white lobes, perfect, 2-2.5 mm
long; achenes 2 mm long; pappus lacking or a small collar. 2n=36
(Clausen. Keck, & Hiesey 1940; Ehrle 1958).

Ubiquitous throughout Wisconsin in a variety of sunny habitats,
from disturbed areas such as sand bars of lakes or rivers, railroad
yards, abandoned fields, roadsides, pastures, and juniper glades to
prairies and open woods, an indicator of mesic prairies (68-69%
presence, cf. Curtis 1955). Flowering from June through mid-
October, the peak from mid-June through July; fruiting from July
to mid-October.

The Achillea millefolium polyploid complex consists of a series
of intergrading and morphologically often indistinguishable forms
(cf. Cronquist 1955) separable mostly by chromosome number and
pollen size correlated with distinctive geographic ranges. The wide¬
spread, evidently native North American Achillea is a tetraploid,
with 2n=36 and with smaller pollen (26-31 /x). Various studies
suggest that nearly all Wisconsin specimens belong to this taxon,
A. m. ssp. lanulosa.

lb. Achillea millefolium L. ssp. millefolium, the European
hexaploid (2n=54) with less dissected leaves, wider segments,

196 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

smaller heads, less pubescence and larger pollen (31-33 p) , has
been found by Mulligan and Bassett (cf. maps p. 76-77, 1959) and
Lawrence (1947) to be sparingly introduced along the northeastern
American coast but absent elsewhere. It is occasionally cultivated,
especially in its deep pink form, and is represented by a few Wis¬
consin collections : Dane Co. : in garden, June 2, 1955, E. J. Wil¬
liams s.n. (WIS). Rusk Co.: Ladysmith, July 30, 1915, /. J, Davis
s.n. (WIS). It may be that some of the wild plants belong to this
taxon, but this cannot be determined at this time.

2. Achillea ptarmica L. Sneezeweed, Map 6.

Glabrate, rhizomatous perennial herbs, 2-10 dm tall. Leaves un-
lobed, linear-lanceolate, serrulate, 3-7 cm long, 3-5 mm wide.
Panicles corymbose, 2-40 headed, the heads 3-4 mm high; bracts
lanceolate, keeled, the margin brown-scarious ; receptacle slightly
convex, chaffy. Rays usually 8-10, white, pistillate, 4-6 mm long,
the disk flowers perfect, 2-3 mm long; achenes with 2-3 light col¬
ored, longitudinal ribs, 1.5-2 mm long; pappus lacking. 2n=18
(Love & Love 1961) .

A Eurasian species, occasionally escaped from old gardens along
roadsides, railroads, vacant lots, and near abandoned homes. Flow¬
ering and fruiting from July to September.

Most of the Wisconsin collections are of a horticultural variant
with all flowers ligulate (f. MULTIPLEX (Reynier) Heimerl=f.
LIGULOSA hort.=var. “The Pearl”).

21. CHRYSANTHEMUM L. Chrysanthemum,
Ox-Eye Daisy.

Heads radiate (rarely discoid in C. halsamita) , singly at the
ends of long branches or corymbose. Rays numerous (rarely want¬
ing), white (ours), pistillate and fertile; disk flowers perfect, 4- or
5-lobed, yellow, campanulate. Involucral bracts imbricate with dry
scarious margins ; receptacle naked, flat or convex. Pappus a short
border or absent ; achenes angular or subterete with 5-10 ribs. An¬
nual or perennial herbs with alternate, entire, toothed, or dissected
leaves. About 150 species, mainly in the Northern Hemisphere,
chiefly in the Old World, many cultivated for their showy flowers.

Key to Species

A. Heads with conspicuous white rays.

B. Heads few, large, 4-6 cm in diam. ; leaves toothed to lobed.

C. Heads solitary on long, slender, naked peduncles ; upper
leaves strongly reduced or lacking; stems slender, 4-6

1966]

Mickelson and litis — Flora of Wisconsin

197

dm tall ; abundant throughout _ _ _

_ _ _ 1. C. LEUCANTHEMUM.

CC. Heads few to many at end of robust^ leafy, 1~2 m tall

stems; peduncles 5-10 mm long; rare escape -

_ _ _ _ C. ULIGINOSUM.

BB. Heads several to many, small, 12-22 mm in diam., corym¬
bose; leaves pinnatisect; rare escape _ _ _

_ _ _ _ 3. C. PARTHENIUM.

AA. Heads discoid or with minute, white rays, ca. 5 mm in diam. ;

leaves serrate; rare adventive _ _ _ 4. C. BALSAMITA.

1. Chrysanthemum leucanthemum L. Ox-Eye Daisy,

Common Daisy, Marguerite. Map 7.

Rhizomatous, glabrous, perennial herbs, with one to several, 2-8
dm tall, erect stems. Leaves irregularly toothed to lobed, the basal
spatulate-obovate, long-petioled, the cauline sessile, strongly re¬
duced above. Heads solitary, 3-6 cm in diam., terminal on long
peduncles; bracts imbricate, dark brown-margined. Rays (45) 20-
40 per head, white and showy, l-2( + ) cm long, pistillate; disks
1-2 cm broad, the flowers perfect, yellow, 2-3 mm long; achenes
10-ribbed; pappus absent. 2n=36, 54 (Darlington 1955).

Native of Europe and western Asia, widely naturalized in cool
temperate North America along roadsides, meadows, pastures, in
abandoned flelds, and other disturbed areas, in Wisconsin a com¬
mon weed throughout but particularly abundant in north central
Wisconsin (cf. map, Lindsey 1953), our plants generally referred
to var. FINN ATIFIDJJM Lecoq. & Lamotte (with pinnatifid basal
leaves, smaller heads), a variety of doubtful validity not differen¬
tiated here. Flowering from June to August (October) ; fruiting
from July to October.

2. Chrysanthemum uliginosum Pers. Giant Daisy, High Daisy.

Map 8.

Rhizomatous, glabrous, perennial herbs with erect, robust, 1-2 m
tall stems, leafy to the top and in dense clones. Leaves sessile,
lanceolate-serrate, 7-11 cm long, 1-2 cm wide. Heads few to many,
large and showy. Involucral bracts lanceolate, the margin brown-
hyaline, Rays about 20-22, white, pistillate; disks 15-20 mm broad,
yellow, the lobes dark-tipped with age; achenes with short, mem¬
branaceous pappus, 2n=18 (Darlington 1955),

A native of southeastern Europe (Hungary, Balkans), often
cultivated for its showy flowers, rarely escaped in moist, disturbed
areas, collected but 4 times in Wisconsin : Dane Co. : Roadside south

198 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

of Lake Waubesa, T6N, RlOE, sec. 19, Sept. 1938, Skinners s,n.
(WIS). Fond du Lac Co.: T15N, R17E, sec. 2, Sept. 24, 1960, Wol-
litz s.n. (WIS). Lincoln Co.: open pasture, T31N, R8E, sec. 12,
Aug. 27, 1950, Seymour & Peroutky 12,170 (WIS) . Winnebago Co. :
Neenah, filled marsh near Lake Winnebago shore. Sept. 23, 1965,
Barker s.n, (WIS), Flowering and fruiting from August to
October,

3. Chrysanthemum parthenium (L.) Bernh. Feverfew.

Matricaria parthenium L. Map 9.

Perennial, subglabrous to lightly pubescent herbs, one-to-several
stemmed, 3-10 dm tall. Leaves bipinnatisect into broadly obtuse
segments, the long-petioled lower 8-12 cm long, to 8 cm wide, the
sessile upper smaller. Corymbs open, the many heads 12-22 mm in
diam., whitish, radiate ; involucre 3-4 mm high. Achenes 10-ribbed ;
pappus short-membranaceous. 2n=18 (Darlington 1955).

A European introduction, commonly grown in old gardens as an
ornamental. An early, apparently oriental, introduction to Europe,
known to Theophrastus and other ancients, and valued as a medi¬
cinal plant for its abortive properties, for promoting menstrual dis¬
charge, as well as for other ailments such as dropsy, fever, nerves
and worms (Hegi 1929). A ruderal, occasionally escaped along
roadsides and lake shores. Flowering and fruiting from June to
August.

4. Chrysanthemum balsamita L. forma tanacetoides Hayek

Costmary, Mint-Geranium. Map 10.

Rhizomatous, aromatic, perennial herbs, 3-12 dm tall. Leaves
undissected, obovate-oblong, serrate, the juvenile silvery-pubescent,
glabrous with age, the lower with 12-25 cm long blade and petiole
of equal length, the upper oblong, smaller, sub-sessile. Heads many,
small, 5 mm broad, discoid, clustered near branch tips. Involucre
3-5 mm high; rays lacking (minute and white in f. BALSAM¬
ITA, this not in Wisconsin) ; disk flowers yellow; achenes 10-
nerved; pappus short, 2n=18, 54 (Darlington 1955).

Native of Asia Minor, Armenia and Persia, once often cultivated
as a medicinal and for the sweet odor of its foliage, in Wisconsin
an occasional garden escape along roadsides, lakeshores, and vacant
lots. Flowering and fruiting from August to October.

Chrysanthemum ooronarium L., Map 9, the Garland or Crown
Daisy, an annual with bipinnatisect leaves and yellow to white
flowers, was once collected (from cultivation?) in Rock Co.: Aug.
28, 1889, Skavlem s.n. (WIS).

1966]

Mickelson and litis — Flora of Wisconsin

199

200 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

22. TANACETUM L. Tansy.

Similar to Chrysanthemum (and often included in that genus),
but differing in the often rather tight, fiat corymbs, discoid or very
short-radiate heads, short-membranaceous pappus, and 3-5 ribbed
achenes. About 25 Northern Hemisphere species of mostly strongly
aromatic, perennial herbs.

Key to Species

A. Heads 25-100 or more in dense corymbs, 7-10 mm in diam,,
the ray flowers without ligules; leaves glabrate; plants in

dense, many stemmed clumps; common introduced weed _

_ l.T. YU LG ARE.

AA. Heads 3-17 in loose corymbs, 10-20 mm in diam., with 2-4
mm long, yellow ligules; leaves tomentose, stems solitary;

rare on inner beaches of L. Michigan, Door Co. _

_ 2. T. HURONENSE.

1. Tanacetum vulgare L. Common Tansy, Golden Buttons.

Chrysanthemum vulgare (L.) Bernh. Map 11.

Rhizomatous, glabrous, aromatic, perennial herbs with many
stems in dense clumps, 4-10 (-14) dm tall. Leaves twice-pinnati-
sect, the ultimate segments sharply toothed (curled in the occa¬
sional f. CRISPUM (L.) Hayek), 8-18 cm long, 3-11 cm wide,
sessile and clasping the stem. Corymbs dense, 5-12 cm wide, with
25-100 or more discoid, golden-yellow heads, 7-10 mm in diam.
Involucre 4-5 mm high; bracts imbricate with scarious margins;
receptacle flat. Heads heterogamous ; flowers fertile, resinous-
granular; rays inconspicuous, pistillate, 3-lobed; disk flowers per¬
fect, 5-lobed; achenes 1.5 mm long, 5-ribbed; pappus lobes small,
membranaceous. 2n=18 (Love & Love 1961) .

A favorite European (?) ornamental and ancient medicinal
plant (oil poisonous!), commonly cultivated, escaped in open or
disturbed areas along roads, railroads, fences, pastures, sandy
beaches of lake shores, and floodplains and banks of rivers. Flower¬
ing and fruiting from July to October.

2. Tanacetum huronense Nutt. Map 12.

Chrysanthemum huronense (Nutt.) Hulten
Tanacetum bipinnatum ssp. huronense (Nutt.) Breitung,
Amer. Midi. Nat. 58 : 66. 1957.

Perennial herbs, from long, slender rhizomes; stems 1-3, erect,
3-5 dm tall. Leaves deeply twice or thrice pinnatisect, ± tomentose,
the ultimate segments mucronulate; basal rosette leaves very large,
23-36 cm long, 3-9 cm wide; cauline leaves 10-23 cm long, 3-8 cm

1966]

Mickelson and litis— Flora of Wisconsin

201

wide. Inflorescence open, heads few, 3-12 (-22), 1~2 cm in diam.;
involucral bracts scarious-margined ; receptacle hemispheric. Rays
short, 2.5-4 mm long, yellow, pistillate ; disk flowers yellow, perfect,
5-lobed, 2-3 mm long; achenes 2-3.5 mm long, ribbed, truncate;
pappus lobes membranaceous.

Central Alaska to Hudson Bay and the northeastern United
States (Maine), in Wisconsin rare but locally abundant on Lake
Michigan shores in Door County, in ecologically open habitats isuch
as calcareous stony beaches, dry peaty and turfy limestone barrens,
interdunal swales, and on dunes, especially the outer Great Lakes
dunes with Ammophila breviligulata, Agropyron psammophilum,
Solidago ( bought oniif ) , Cirsium pitcherii, Prunus pumila, Salix
syrticola, and other Great Lakes specialities. In Wisconsin very
rare: Door Co.: Baileys Harbor, July 21, 1940, Goessl s.n. (WIS).
Jacksonport, Aug. 3, 4, 1929, J. J. Davis s.n. (WIS) ; Jacksonport,
Lake Michigan beach, July 15, 1940, Shinners & Sieker 2208
(MIL). Newport, July 3, 1905, Milwaukee Public Museum Expedi¬
tion s.n. (MIL). Flowering and fruiting from July to September.

Tanacetum huronense var. huronense has recently been mapped
by Guire and Voss (1963) , who accepted its endemism and distinct¬
ness from the three other varieties described by Fernald (1935).
However, in the Flora of Alaska, Hulten (1950) comments that
the Alaskan populations contain all of Fernald’iS varieties ! Although
in general Great Lakes collections are more robust and have more
heads than those from the East, one of Fernald’s main characters,
that of head number [high (6-30) in var. huronense, and low (1-6)
in the other varieties] , does not hold. Several Michigan collections
have as few as 3 or 4 heads [e.g. Ehlers 1061 (WIS), Gates 13,995
(UC), litis 15,853 & 15,850 (WIS)], yet are simply few-headed
extremes of var. huronense populations (cf. Fig. 1), and are nearly
indistinguishable from isuch collections as Fernald 69 (Aroostoock
Co., Maine, UC), Collins & Williams s.n. (St. John River, Maine,
UC) , or Fowler s.n. [Restigouche, Bass River, N.B. ( WIS) ] , of var.
johanense Fern, On the other hand, the northernmost Hudson Bay
collections and those from Newfoundland, with 1-3 heads and but
1-2 dm tall, appear to be depauperate plants strongly resembling
Alaskan collections of T. bipinnatum. These findings agree well
with those of Fernald (1923) which are more realistic than his
later conclusions (1935, 1950).

It seems likely that Alaska is the area of glacial survival and
the center of dispersal of this species. Here it is most abundant as
well as morphologically most diverse, perhaps due to hybridization
with T. bipinnatum. Furthermore, unless one accepts Fernald’s
“Nunataks'', this is the only region where the species occurs in un-

202 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

A

•

UPPER GREAT LAKES

( var .

huronense )

OO

o

MAINE, E. QUEBEC

( var .

johanense )

o

o

A

HUDSON BAY, ANTICOSTI

( var .

b j f ar i urn )

o

o

o

o

NEWFOUNDLAND

( var .

terrae-novae )

1 i 5

4 5 6 7

i 9

10

11 12 13 14 15 16

17 11

19 20 21

NUMBER OF HEADS PER STEM

Figure 1. Comparison of number of heads per stem with height of specimens
of Tanacetum huronense from four geographical areas in North America.

1966]

Mickelson and litis — Flora of Wisconsin

203

Fig.

TANACETUM

• T. HURONENSE
o T. DOUGLAS I I
© T. B I P I

* T. CAMPHORATUM
“ GLAC I AL MAX IMUM

glaciated territory (cf. Fig. 2). One can postulate that the modern
Eastern North American local populations are either the result of
sporadic post-glacial long-range dispersal to “open’’, often calcare¬
ous habitats and/or the scattered residues of once broader distri¬
butions which now survive only in special open habitats such as
dunes or gravel flats which are low in competition. Since only a
few plants became established at any one place, to subsequently
give rise to larger populations (either sexually or asexually), only
a fraction of the diversity of the original Alaskan population would
thus be represented. If to this lack of genetic variability one adds
effects of inbreeding, as well as local selection and direct phenotypic
responses to climatic factors, it is not surprising to find local popu¬
lations differing in minutiae. A similar situation has been discussed
in Gentianopsis (litis 1965).

204 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

23. MATRICARIA L. Wild Chamomile.

Heads radiate with pistillate, fertile and white rays, or heads dis¬
coid and rays absent (M. matricarioides) ; disk flowers perfect, fer¬
tile, campanulate, yellow or greenish, 4-5 lobed. Involucral bracts
2-3 seriate, slightly imbricate, scarious-margined. Receptacle
naked, hemispheric or conic. Pappus a membranaceous crown, or
absent. Achenes generally ribbed or wing-margined. Annual or
perennial herbs with alternate, finely pinnately-dissected leaves,
and few to many terminal heads. About 40 Northern Hemisphere
and South African species, ours introduced weeds.

Key to Species

A. Ray flowers none ; disk flowers greenish, 4-lobed ; heads short-
stalked; achenes marked by elongated red-brown oil glands;

very common throughout _ 1. M. MATRICARIOIDES.

AA. Ray flowers white; disk flowers yellow, 5-lobed; heads long-
stalked.

B. Receptacle conic at maturity; achenes ribbed, smooth, un¬
marked ; involucre 2-3 mm high ; rare, but see also Anthe-
mis cotula _ 2. M. CHAMOMILE A.

BB. Receptacle hemispheric at maturity; achenes prominently
ribbed, transversely rugulose or tuberculate with apical
oil glands; involucre 3.5-6 mm high; rare on Lake Supe¬
rior shores _ 3. M. MARITIMA.

1. Matricaria matricarioides (Less.) Porter Pineapple Weed.
Matricaria suaveolens L. Map 14.

Glabrate annual with strong aromatic pineapple odor ; stems one
to several, erect, 4-50 cm tall. Leaves finely twice-pinnatifid into
0.5 mm wide linear segments, sessile, 2.5-9 cm long, 1-2.5 cm wide.
Heads short-stalked, discoid and conical. Involucre 3-5.5 mm high ;
bracts 24-32, with red-brown midrib and broad scarious margins;
receptacle naked, hollow. Ray flowers lacking. Disk flowers ^-lobed,
yellow-green, 1-1.5 mm long; achenes 1-1.5 mm long,^ ribbed with
2 longitudinal, slender red-brown oil glands; pappus rudimentary.
2n=18 (Love & Love 1961) .

Native probably of NE Asia (Clapham, et al. 1952; Baker
1962), recently (1850’s) occurring as a weed in Europe and North
America, in Wisconsin the earliest collections from the northern¬
most counties (1915), not collected in Milwaukee until 1939 or in
Madison until 1942, ubiquitous in farmyards, fields, along road¬
sides, industrial areas, and in cities along walks, streets, public
grounds, and waste areas. Flowering from May through mid-
September; fruiting from June through September,

205

1966] Mickelson and litis — Flora of Wisconsin

2, Matricaria chamomilla L. Chamomile. Map 13,

Aromatic, glabrous annual ; stems erect, slender, 2™-4 dm tall.
Leaves sessile, 1--2 cm wide, 2-6 cm long, twice-pinnatifid into
linear-filiform segments 0.5 mm wide. Heads few, 5-12, radiate,
resembling a small daisy; receptacle naked^ sharply conic at ma¬
turity, hollow ; involucre 2-3 mm high ; bracts with brown midrib,
light scarious margins. Ray flowers pistillate, white, 13-20, 5-7
mm long; disk flowers yellow, S-lobed, 1-1,5 mm long; achenes
smooth, ribs not prominate, 1 mm long, 2n=18 (Love & Love 1961).

Native of southern and eastern Europe to west Asia, infrequent
in Wisconsin in disturbed habitats, waste places in towns and water
fronts. Flowering and fruiting from May to July. It has been known
since 1588 that a blue oil can be distilled from the plant (Karsten
1946),

The Wisconsin plants are all erect, dz corymbose, few-headed,
and much more uniform than European collections. The species
strongly resembles Anthemis cotula (q,v.).

3. Matricaria maritima L, ssp. inodora (L.) Clapham. Map 15,

Matricaria inodora L,

Matricaria maritima var, agrestis (Knaf.) Wilmott,

Annual, 3-7 dm tall, erect, glabrous-glabrate. Leaves sessile, 2-5
cm long, 1-3 cm wide, twice pinnatifid into 0.5 mm wide, linear-
filiform segments. Heads radiate, few to many; involucre 3-6 mm
high; bracts with brown, scarious margins. Ray flowers white,
12-20, pistillate, 1-2 cm long; disk flowers yellow, 1-2 mm long;
achenes with .2-3 wide ribs, 2 red-brown oil glands near the top,
transversely rugulose, 1-2.5 mm long; pappus rudimentary. Flower¬
ing from July through October; fruiting in September and October.

A Eurasian species, collected only near Herbster, Bayfield County
on Lake Superior. Bark Point, lake shore and open fields, Sept, 6,
1959, litis 15,515 (WIS). Bark Point, roadside, July 10, 1938,
Fassett 20fi'30 (WIS).

A plant cited by the collector as the above (Lakela 1965: 387),
but more resembling MATRICARIA MARITIMA var. MARITIMA
with short height (3 dm) and fewer, larger heads borne on long,
sturdy peduncles, was collected once near Duluth, Minnesota: in
bare soil of roadcut, Duluth Heights, Duluth, Oct. 18, 1947, Lakela
7m (WIS).

24. ARTEMISIA L. Wormwood, Sage, Sagebrush.

Annual, biennial, or perennial aromatic herbs or small subshrubs
with erect to decumbent stems. Leaves alternate, entire to deeply
pinnatifid or pinnatisect, pubescent to glabrous. Inflorescence a

206 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

spike-like, racemose, or profusely branched panicle of small and
inconspicuous, discoid heads. Receptacle naked or covered by hairs.
Bracts imbricate in several series, the inner with scarious margins.
Flowers resinous-granular, 1-3 mm long ; rays not ligulate, 5-lobed,
pistillate with bifid styles ; disk flowers perfect, sterile or fertile,
5-lobed with bifid, truncate-erose styles (simple in A. caudata) .
Seeds minute, ca. 1 mm long; pappus lacking.

About 200 species, over half in the USSR, distributed mainly in
cool or arid regions of the Northern Hemisphere, several species
dominant shrubs in prairies, plains, and deserts, cultivated for
foliage plants and, because of their aromatic, volatile oils and bitter
substances, for use in tonics, antihelminthics, and liqueurs.

Key to Species

A. Receptacle hairy; leaves white-silky canescent; plants peren¬
nial and somewhat woody at base.

B. Leaves short, 1-2 cm, the segments filiform, 0.5-1 mm
wide ; flowering stems ascending, to 5 dm tall, the vegeta¬
tive stems forming mats; inflorescence a narrow panicle;
very local on Mississippi River bluffs from Pierce Co. to
Trempealeau Co., rarely weedy elsewhere _1. A. FRIGID A.

BB. Leaves 5-15 cm long, the segments 2-3 mm wide ; flowering
stems erect, to 9 dm tall; inflorescence a leafy panicle;

sporadic adventive _ 2. A. ABSINTHIUM.

AA. Receptacle naked ; leaves tomentose to glabrous ; plants annual,
biennial, or perennial.

C. Disk flowers sterile, the achenes abortive; adult plants
usually glabrous.

D. First year lower leaves forming a basal rosette ; leaves
tomentose-glabrate ; involucre 2-3.5 mm high; tap-

rooted biennial; common in sandy areas _

_ 3. A. CAUDATA.

DD. Lower leaves not in a rosette; involucre 2 mm high;
robust glabrous herbs from a rootstock; very rare and

sporadic _ 4. A. DRACUNCULOIDES.

CC. Disk flowers fertile.

E. Leaves glabrous-glabrate, 2-3 times pinnatisect or
pinnatifid.

F. Annual or biennial herbs; involucre 1-2 mm high;
bracts glabrous.

G. Inflorescence a dense racemose panicle with
many spike-like branches from the leaf axils;
heads erect; common weed _ 5. A. BIENNIS.

1966]

Mickelson and litis— Flora of Wisconsin

207

GG. Inflorescence a broad terminal panicle with

nodding heads; rare annual weed _

_ _ 6. A. ANNUA.

FF. Perennial shrub; involucre 2-2.5 mm high; bracts
canescent or tomentose; rarely escaped cultigen --

_ 7. A. ABROTANUM.

EE. Leaves tomentose at least on one surface, simple or
dissected.

H, Leaves unlobed and linear-lanceolate, the margins
regularly serrate to entire in the inflorescence,
densely white-tomentose beneath, bright green-

glabrous above; moist deep-soil prairies _

_ 8. A. SERRATA.

HH. Leaves deeply lobed or cut, or entire with the mar¬
gins irregularly toothed.

I. Leaves delicately divided, the segments filiform,
gray-green pubescent; rarely escaped cultigen __

_ 9. PONTIC A.

11. Leaf segments broader or leaves entire.

J. Leaves green-glabrate above, white-tomentose
beneath, coarsely lobed; rare weed, eastern

Wisconsin _ 10. A. VULGARIS.

JJ. Leaves pubescent on both surfaces.

K. Involucre 2-4 mm high; leaves entire or
irregularly toothed, densely white-
tomentose beneath, tomentose to glabrate

above; common prairie species _

_ 11. A. LUDOVICIANA.

KK. Involucre 5-8 mm high; leaves obtusely
lobed, densely creamy white wooly; rarely
escaped on L. Michigan or L. Superior __
_ 12. STELLERIANA.

1. Artemisia frigida Willd. Prairie Sagewort. Map 16.

Mat-forming, gray-green tomentose perennial, decumbently
branched and ± woody at base, the several flowering stems 3-5
dm tall, erect-ascending. Leaves 1-2 cm long, finely dissected into
linear segments. Panicles 17-28 cm long, narrow with ascending
branches; heads many, 2-3 mm high; receptacle long -villous ;
bracts ca. 15. Flowers 26-34, fertile; rays 7-9; disk flowers 19-25;
achenes 1 mm long. 2n=18 (Kawatani & Ohno 1964).

A western Great Plains and Rocky Mountains grassland species,
ranging from central Alaska to New Mexico, in Wisconsin only
along the Mississippi River in Pierce, Pepin and Trempealeau

208 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

1966]

Mickelson and litis — Flora of Wisconsin

209

counties on exposed limestone (dolomite) bluffs, talus slopes, and
Mississippi River sand terraces (rarely adventive elsewhere in dis¬
turbed areas such as on railroads and road cuts). Flowering in
August and September ; fruiting in October.

These Mississippi and St. Croix River bluffs and adjoining sand
terraces appear to be among the most xeric habitats in Wisconsin.
Sloping steeply southwestward, they receive the maximum amount
of sunlight and accumulate little soil because of high winds and
extreme water run-off. In this environment water and nutrients are
limited (Curtis 1959), and only a few plants can survive. Some
fairly widespread dry-prairie species that are otherwise rare or
restricted in Wisconsin occur here, including Besseya bullii (Sala-
mun 1951), Castilleja sessiliflora (Salamum 1951), Psoralea escu-
lenta (Fassett 1939), Anemone patens var. wolfgangiana (Fassett
1947), Liatris cylindracea (Johnson & litis 1963), Bouteloua hir-
suta, B. curtipendula (Fassett 1951), Artemisia caudata, Aster
oblongifolius, and A. ptarmicoides.

In addition a number of otherwise widespread Great Plains spe¬
cies have highly localized Wisconsin distributions very similar to
that of A. frigida, and like that species reach their easternmost
limits here. These include such Wisconsin rarities as Psoralea ar-
gophylla, Astragalus caryocarpus, and Petalostemon villosum (Fas¬
sett 1939), Anemone caroliniana (Almon 1930), Liatris punctata
(Johnson & litis 1963), Bouteloua gracilis, and Muhlenbergia cus-
pidata (Fassett 1951), and Erigeron glabellus.

2. Artemisia absinthium L. Absinthe, Sagewort. Map 17.

Aromatic, rhizomatous, basally suffrutescent, perennial herbs,
6-9 dm tall. Leaves white-silky canescent, esp. when young, 6-15
cm long, 2-8 cm wide, long-petioled (4-13 cm), the upper sessile,
divided to entire, the lower 2-3 times pinnatifid-pinnatisect, the
lobes 2-4 mm wide. Panicles leafy, 17-50 cm long, to 30 cm wide,
the branches rather strict-ascending. Heads many, rather large
(3-4 mm), nodding; involucre 2-3 mm high; bracts 13-18; recep¬
tacle covered with villous hairs. Flowers 45-80, fertile; rays 10-21,
the disk flowers 35-58; achenes 1 mm long. 2n=18 (Love & Love
1961).

Native of temperate, dry regions from central Asia to south¬
western Europe, an ancient, prehistoric medicinal and magical
herb, at present still grown for use as a tonic, antihelminthic (hence
'Vormwood”), local anaesthetic, and in the preparation of the
liqueurs Vermouth and Absinthe, in Wisconsin in disturbed, esp.
calcareous areas along roads, in pastured fields, and waste places
in cities. Flowering from end of July to September; fruiting in
August and September.

210 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

3. Artemisia caudata Michx. Field Sagewort, Wormwood.

Map 18.

Biennial or occasionally short-lived perennials with a ± woody
taproot ; pubescence very variable, often gray-tomentoise when
young, usually glabrate throughout when mature. Stems 1-5, erect-
ascending, 6-10 (-16) dm tall, reddish-violet tinged. First year
rosette leaves 10-20 cm long, 6-12 cm wide, long petioled (4-8 cm),
2-3 times pinnatifid into l(--2) mm wide segments; cauline leaves
sessile, with axillary branchlet “fascicles’’, the lower 7-10 cm long,
2-6 cm wide, smaller, less divided above. Panicles leafy, 23-60 cm
long, 2.5-22 cm wide, ascending-branched, with many greenish
heads. Involucre 2-4 mm high; bracts 12-16, glabrous, green with
a red-brown midrib, scarious margins; receptacle naked. Flowers
21-41 ; rays fertile, 11-21, ca. 1 mm long disk flowers sterile, the
style simple (not bifid) 10-20, 1.5-2 mm long; achenes 1 mm long.
2n=18 (Kawatani & Ohno 1964).

A variable North American native of wide distribution, most
common in the eastern and central United States and Canada, rang¬
ing westward to the Pacific coast, in Wisconsin very common in
dry and sandy prairies (81-87% presence, Curtis 1959), sensitive
to competition and often found in ecologically open habitats such as
dry high-lime prairies, dry sand prairies (with Stipa spartea.
Delphinium virescens, Arahis lyrata, Oenothera rhomhipetala) ,
sandstone and limestone cliffs, inner beaches and dunes on Lake
Michigan, sandbars of rivers, and in jack oak-jack pine barrens,
often weedy in disturbed areas along roadsides or sandy railroads,
overgrazed or abandoned sandy fields, and waste areas of cities.
Flowering from mid-August through September; fruiting to early
October.

The smaller, somewhat similar, subarctic Artemisia canadensis
Michx. (=A. borealis Pallas), is cited for Wisconsin by Fernald
(1950), evidently on the basis of some depauperate specimens of
A. caudata,

4. Artemisia dracunculoides Pursh. Dragon Sagewort,

Estragon, Tarragon. Map 19.

Artemisia dracunculus L. of authors, not the Eurasian taxon.

Artemisia glauca Pallas, ex. Willd.

Glabrous, rhizomatous, robust perennials, 5-14 dm tall from a
woody caudex. Leaves dark green, often turning black, sessile,
(2) 4-7 cm long, simple or 3~parted into narrowly-linear, 1-2 mm
wide, unbranched segments, the upper simple. Panicles leafy,
diffuse, the slender branches often drooping, 3-6 dm long, to 2.5

1966]

Mickelson and litis — Flora of Wisconsin

211

dm wide; involucres 2-2.5 mm high; bracts 11-16, glabrous; recep¬
tacle naked. Flowers 19-32 ; rays fertile, 10-16 ; disk flowers sterile,

9- 16, 1.5 mm long; achenes 1 mm long.

In Wisconsin sporadic, in sandy soils, on dry, sandy prairie bluffs,
and along roads and railroads, similar to A. caudata (q.v.) but
quite uncommon. Flowering and fruiting from August to October.

A complex group native to the western and midwestern U. S. in
sandy or grassy habitats, probably consisting of several taxa poorly
understood, apparently divisible into two major types: one, rang¬
ing from Oklahoma-Colorado to Montana-Alberta, and west to
California, characterized by fewer, broader leaves, and erectly
branched panicles; and the other, ranging from Nebraska-Missouri
to Minnesota-Iowa and Wisconsin, characterized by numerous,
slender leaves, and diffuse panicles with drooping branches. With
both nomenclature and taxonomy confused, it is not possible to
determine here proper species limits or names.

5. Artemisia biennis Willd. Biennial Sagewort. Map 20.

Biennial (or annual?) glabrate-glabrous herbs, 3-7 dm tall, un¬
branched above, or with short, dense, lateral inflorescence branches
below, these often to the base. Leaves sessile, the lower 5-15 cm
long, 2-7 cm wide, deeply once- less often twice-pinnatifid, the few
sharp-toothed lobes 20-40 mm long, 1-5 mm wide, the reduced
upper leaves once-pinnatifid. Panicles dense, spike-like or racemose,
often axillary, usually exceeded by leaves. Heads small, erect,
short-peduncled or sessile, 1-2 mm high; bracts 11-16; receptacle
naked. Flowers 37-45, fertile; rays 17-22, the disk flowers 20-23;
achenes 1 mm long.

Presumably native to western North America, widely distributed
as a weed eastward, in Wisconsin on gravelly, sometimes muddy
lake shores and river banks, often weedy in waste areas in cities,
farmyards, gardens, railroads, and especially roadsides, one of the
relatively few native American weeds in this area. Flowering and
fruiting from very late August into November, peaking in mid-
September,

6. Artemisia annua L. Annual Sagewort. Map 21.

Glabrous, 3-7 dm tall annuals. Leaves flnely 1-3 times pinnati-
sect into 1 mm wide segments, the lower 3-10 cm long, 3-9 cm
wide, petioled, the upper sessile, smaller. Panicles leafy, 10-50 cm
long, 3-24 cm broad, very diffuse, with many minute heads. In¬
volucre 1.5-2 mm high; bracts 8-18; receptacle naked. Flowers

10- 29; rays 5-9, the disk flowers 5-20; achenes 1 mm long. 2n=18
(Love & Love 1961).

212 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

ARTEM ISIA
DRACUNCULOIDES

1 L L t ^]o^ s

I lL 1 No!\ s

ARTEMISIA

ANNUA

A. ABROTANUM

! . 1 r"

ARTEMISIA

SERRATA

NOi\ S

moT

ARTEMISIA
PONTIC A

ARTEMISIA

VULGARIS

LL. I N0!\

'• ARTEMISIA

BIENNIS

1966]

Mickelson and litis — Flora of Wisconsin

213

A temperate Asiatic and southeastern European (Balkans)
weed, sparingly naturalized in eastern North America in waste
places and old fields, known from only 3 Wisconsin collections.
Milwaukee Co.: Milwaukee, Sept. 2, 1940, Skinners 3S58 (WIS).
Racine Co.: Racine, Aug. 6, 1898, Wadmond s.n. (MIN). Sheboy¬
gan Co. : Sheboygan, waste place in city, Aug. 1911, Goessl s.n.
(WIS). Flowering and fruiting in August and September.

7, Artemisia abrotanum L. Garden Sagebrush, Southernwood.

Map 21.

Strongly aromatic, 5-10 dm tall sub-shrub, with the leaves finely
divided into linear-filiform segments, these glabrous above, hairy
below. Panicles leafy, 15-40 cm long, the many, small heads droop¬
ing, 2-2.5 mm high; bracts 8-18; receptacle naked. Flowers 15-35,
all fertile. 2n=18 (Love & Love 1961).

Native of southeastern Europe and temperate Asia (apparently
a race of A. paniculata Lam.) known only in cultivation or as a
rare escape (cf. Hegi, 1929, 6^L 635), in Wisconsin uncollected for
over 60 years, Racine Co. : Racine, garden, dz 1860, Hale s.n.
(WIS). Sheboygan Co.: Sheboygan, roadside. Sept. 9, 1903, Goessl
s.n. (WIS). Lakela (1965) cites a collection from Duluth.

8, Artemisia serrata Nutt. Genera of North American Plants 2 :

142, 1818. (Lectotype: “Near the Prairie du Chien, on the banks

of the Mississippi, in open alluvial soils” Nuttall s.n., in PH.)

Saw-leaf Mugwort. Map 22.

Erect, rhizomatous, robust, perennial herbs, 6-15 dm or more
tall. Leaves linear-elliptic, white pubescent beneath, dark green and
glabrous above, the lower 7-16 cm long, 1-2.5 cm wide, regularly
serrate, the upper smaller, less regularly serrate to entire. Panicles
leafy, 14-50 cm long, 2.5-18 cm wide; heads nodding, short-
peduncled, 3 mm high; bracts 9-15; receptacle naked. Flowers
14-25, fertile, the rays 5-10, the disks 9-15 ; achenes 1 mm long.
2n=36 (Keck 1946).

A localized prairie species of the upper Mississippi River Valley
(cf. fig. 3) occurring in rich moist soils along rivers and streams,
wet prairies and ditches, low wet meadows, and moist sandy al¬
luvial soils. Flowering and fruiting in August and September.

Although NuttalFs original description of A. serrata gives its
location as “near the Prairie du Chien, on the banks of the Missis¬
sippi, .... also on the banks of the Missouri”, the range of the
species does not extend to the Missouri River; therefore, the Wis¬
consin collection is here designated as the lectotype.

214 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

The Phytogeography of Artemisia Serrata
AND ITS Significance.

Because the flora of glaciated lands (except for Arctic or conifer¬
ous forest species that might have lived on top of the ice) must
be derived from unglaciated areas, because species or floras do not
migrate readily except into “open” habitats without much compe¬
tition, and because glaciation, especially that of the Wisconsin ice
advance, occurred only very recently, we can profitably use the mod¬
ern distribution of a species as shown on a detailed range map to
pin-point the probable area of its glacial survival and possible
routes of subsequent migration into once-glaciated territory. This
approach has great utility for understanding the histories of indi¬
vidual species and whole floras, as well as for pinpointing glacial
“refugia” (Hulten, 1937) or more appropriately “survivia” (litis,
1965). The resultant composite geographic patterns tend to center
at one end of the survival areas, from whence they radiate, depend¬
ing on the tolerance of the species, in ever-increasing “progressive
equiformal areas” (Hulten, 1937), these especially beautifully
shown by the Appalachian species in the upper Middle West.

It is a fact of the very greatest interest that, with the rarest
exceptions, all species present in glaciated territory of Eastern or
Central North America have at least one small part of their range
outside the glacial maximum, this then the area where we may
assume they survived glaciation.

Exceptions to this rule include first of all species which have
evolved since the last glacial period, such as the many dune and
strand taxa of the glacial lakes (Guire and Voss, 1963 ; Johnson and
litis, 1963) a few of which are rather distinct (e.g. Cirsium pitcheri
(Torr.) T. & G. ; cf. Johnson and litis, 1963) but most of which
are better treated as microspecies or geographic subspecies (e.g.
Tanacetum huronensis Nutt.; cf. p. 200-203; Iris cristata Ait. ssp.
lacustris (Nutt.) litis; cf. Mason and litis, 1965). Several Midwest¬
ern bog or cliff species with western affinities, if distinct at all,
belong here as well (litis, 1965), such as Dodecatheon amethysti-
num (Fassett) Fassett and Aconitum noveboracense A. Gray (=A.
columbianum Nutt.).

Secondly, there are a number of species of hybrid derivation,
such as Cyperus houghtonii Torr. (Marcks MS.), Penstemon ivis-
consinensis Pennell (Crosswhite MS.) and Quercus ellipsoidalis
E. J. Hill, these evidently also being post-glacial in origin.

Thirdly, there are a number of species or varieties in such large
and actively evolving genera as Bidens or Solidago, or in apomictic
genera as Rubus or Crataegus, which again we must assume to be
post-glacial in origin. These three groups listed so far are particu-

1966]

Mickelson and litis — Flora of Wisconsin

215

larly interesting for the evolutionist since in many cases they rep¬
resent recent and often rapid morphological evolution that can be
dated rather accurately by the retreat of the Wisconsin ice sheet.

Fourthly, there are a small number of highly distinct species
without North American relatives, which we may assume that
because of their microspermous seed were able to migrate here
by stratospheric transport (jet streams?) . Such rarities, with their
known or probable region of origin in parentheses, include the fab¬
ulous and now extinct Thismia americana Pfeiffer (Burmaniaceae
— ^Tasmania?) once collected in a wet prairie near Chicago, Cypri-
pedium arietinum R, Br, (Orchidaceae — China), Pedicularis fur-
bishiae S. Wats, of northern New England (Scrophulariaceae —
Himalayas?) and a fern of E. Asiatic affinity (W. H. Wagner,
personal communication).

Fifth and last are a small number of distinct to very distinct
taxa whose geographic origins and histories are harder to explain.
All these taxa, significantly we think, are associated with prairie
habitats, and are mapped here (Fig. 3) .

Of these, the most distinct is the monotypic genus Napaea, which
does grow in unglaciated parts of Ohio, but there apparently as a
recent weedy introduction (litis, 1963). It is a polyploid (litis and
Kawano, 1964) with a relationship, if very distant, to Sidalcea
malachroides (Hook & Arn.) A. Gray of California.

Artemisia serrata Nutt., a very distinct polyploid species (n =
18), appears closest to the diploid A. longifolia Nutt, (n = 9) of
the northern Great Plains (Keck, 1946).

Besseya bullii has its rather close congeneric relatives in the
Western mountains. Pennell (1935) suggests it may be of post-
Wisconsin origin since all stations south of the Wisconsin maxi¬
mum are on recent outwash plains.

Cirsium hillii (Canby) Fern. (C. pumilum (Nutt.) Spreng. ssp.
hum (Canby) Moore & Frankton), a perennial, appears to be de¬
rived from the biennial C. pumilum (Nutt.) Spreng. of the Eastern
United States, this or both in turn from C. drummondii Torr. &
Gray of the arid western Great Plains.

Excepting the enigmatic Erythronium propullans A. Gray from
near Minneapolis, the only good ''Linnean'' species that occur wholly
within glaciated territory are the feiv prairie species listed above.
litis (1963) originally suggested that Napaea dioica survived
the Pleistocene glaciations together with the tall grass prairie in
the Oklahoma-Kansas-Ozark region from where, poist-glacially,
the species vanished to migrate north into glaciated lands with the
tall grass prairie. However, in view of the fact that the other taxa
of diverse geographic relationships listed above all share this re¬
striction to glaciated prairie lands, as well as their occurrence in

216 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Artem i s i a serrata
Very distinct sp.jClosest
relative A. longifolia ?
of Northern Great Plains

After Keck I 946 , p . p

Napaea d i o i ca

Monotypic endemic genus
Very distantly related to .
Sidalcea malachroides from
;Ca I i f orn i a .Low prairies.^

!G I ac i a I max i ma

After litis 1963

C i rs i urn pum i I urn ssp. h i I I i 1

PERENNIAL OP DRY PRAIRIES, DERIVED PROM (?) SSP.
SANDY OPEN PLACES. CLOSEST RELATIVE IS CIRSIUM
WESTERN GREAT PLAINS.

ssp . pum i I urn

After Moore &
Frankton 1966

pro parte

1966]

Mickelson and litis — Flora of Wisconsin

217

Besseva bu I I i i

Dry, sandy, gravelly bluffs and ^
prairies. The 8 other spp. of the
genus all in the Western U.S.

nr: i \ L

Sti'q

1 1

L [ 1 .

Vb

i ill]

i

i [

Lespedeza I eptostachva

'Rare prarrie legume, extinct
^ in Wisconsin. Related to Lespedeza
■^^angust i f 0 I i a of the Atlantic Coastal Plain.

Figure 3. Five species whose ranges are restricted to deepsoil prairies in
glaciated regions of the Middle West. The Ohio records of Napaea dioica in
unglaciated territory appear to be introductions since they come from dis<
turbed areas.

lands both glaciated and unglaciated by the Wisconsin glaciation,
we may suggest a more specific hypothesis; namely that these

species

a) originally became established in the prairie in the open habi¬
tats following the retreat of the Illinois glacier, a time sufficiently
distant for considerable speciation, b) that subsequently they
evolved into new taxa there, and c) that they survived the Wiscon¬
sin glaciation in the Illinois-Iowa prairies as well, which must have
existed there then as they do now. The very distinct Napaea, of
course, probably had an older history as previously outlined.

218 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

It is suggestive that one other, but not as sharply distinct a taxon,
the rare Lespedeza leptostachya Engelm., is restricted to glaciated
lands but only of the Wisconsin glaciation (as already pointed out
by Fassett (1939)). Yet this restriction and its close relationship
to the southern Atlantic Coastal plain L. angustifolia (Pursh.)
EIL (cf. Clewell, 1966) suggests that its introduction into the
prairie vegetation was recent ( post- Wisconsin ) , hence its morpho¬
logical evolution is very slight (For a contrary opinion using the
“Driftless Area’' as a refugium, see Fernald, 1925). The signifi¬
cance of these speculations to the phytogeographic history of the
region is obvious. They suggest, contrary to believers in marked
periglacial climatic depressions and broad coniferous belts south
of the ice, that the prairie in fact did occur and survive in the Illi¬
nois area at the height of the last ice advance and that the climate
then ,was not too different from what it is now, views that agree
well with those in the many publications of E. L. Braun and one
of us (cf. litis, 1965; Mason and litis 1965), based on other and
varied evidence.*

Artemisia s errata, like Napaea dioica with which it may grow
(e.g. Dane Co, trunk PB, along a railroad near Verona; cf. photo
in litis 1963), is thus phytogeographically clearly a most unusual
species, one of a select group of prairie plants which originated in
the area of Illinoian glaciation and which increased their ranges
into the area of Wisconsin glaciation sometime between the retreat
of that ice and the present. Its present survival in a few wet rail¬
road rights-of-way, prairies which are now in ever-increasing dan¬
ger of needless and thoughtless herbicide spraying for “weeds”, is
precarious. Its habitats, the remnants of low prairies, are in des¬
perate need of protection and preservation.

9. Artemisia pontica L. Roman Wormwood. Map 23.

Gray-green tomentose, rhizomatous perennials, 3-10 dm tall.
Leaves twice-pinnatisect into fine, linear, rather short segments, the
lower 1-3 cm long, petioled. Panicles 10-25 cm long, 2-4 cm wide,
the heads nodding, 2-3 mm high; bracts 12-18; receptacle naked.

* Substantiation for prairies in the “Wisconsin”-glaciated Midwest immediately
following- g-lacial retreat comes from pollen analyses of W. S. Benning-hof (The Prairie
Peninsula as a filter barrier to Postglacial plant migration. Proc. Indiana Acad. Sci.
1936. 72: 116-124. 1964). Abundant grass pollen in the earliest Late-glacial Spruce-
Fir zone deposits of Indiana and Ohio, and the relatively very late appearance of
Hemlock and beech pollen is taken as evidence in support of earliest Late-glacial
invasion by grassland elements ; i.e. that the “Prairie Peninsula” became established
in wake of glacial retreat, early enough to prevent its crossing by Beech and Hemlock.
In view of the several endemic animals (K. P. Schmidt) and the endemic plants
listed above, we may trace the Prairie Peninsula or part of it just one step further
back in time ; namely that prairie, is a pre-ice feature, which, growing during the
Wisconsin maximum next to the ice in Illinois and Iowa, simply moved eastward
following ice retreat. We may then also assume that the climatic pattern then was
not too different from what it is now.

1966]

Mickelson and litis — Flora of Wisconsin

219

Flowers 35-60, fertile, the rays 10-15 (-18), the disk flowers 25-45.
2ri=18 (Suzuka 1952; Kawatani and Ohno 1964).

Native mostly of southeastern Europe and western Asia in arid
grasslands, in America cultivated as a border plant for the grayish-
white foliage, in Wisconsin escaped but not established in waste
places in cities and roadsides : Douglas Co. : Brule River, sandy
roadside T45N, RllW, S4, July 2, 1943. Thomson, 5270 (WIS).
Green Co. : Juda, roadside, Sept. 29, 1957, Fell 57-1357 (WIS). She¬
boygan Co.: roadside in Sheboygan, July 1912, Goessl s.m (WIS).

10. Artemisia vulgaris L. Sagewort, Mugwort. Map 24.

Rhizomatous, 9-13 dm tall perennial herbs. Leaves white-
tomentose beneath, dark-green glabrous above, the lower 2-3 times
pinnatifld-pinnatisect, 5-11 cm long, 2-7 cm wide, the lobes rather
broad, acute. Panicles leafy, 25-60 cm long, 3-22 cm wide, the
heads short-peduncled, nodding, 3-4 mm high; bracts 12-16; re¬
ceptacle naked. Flowers 22-26, fertile; achenes 1 mm long. 2n=16
(Love & Love 1961).

Native of Eurasia formerly cultivated as a medicinal, as a substi¬
tute for hops in making beer, as tea in England, and for its foliage,
introduced in America and occasionally adventive in eastern Wis¬
consin in sandy or disturbed areas such as railroad yards or waste
grounds in cities.

11. Artemisia ludoviciana Nutt. ssp. ludoviciana Western

Mugwort, White Sage. May 25.

Artemisia gnaphalodes Nutt. Genera North American Plants
2: 143, 1818. (Type: ‘‘On dry savannahs about Green Bay,
Lake Michigan, and on the banks of Fox River, and the
Missouri’’ Nuttall s.n. (in PH?) ; cf. Keck 1946:441.)

Artemisia ludoviciana Nutt. var. gnaphalodes (Nutt.) Torr.
et Gray.

Densely white-pubescent, rhizomatous, perennial herbs, 3-7 (-12)
dm tall, the stems unbranched (rarely branched above), often in
extensive loose clones. Leaves densely white-tomentose beneath,
tomentose to glabrate above, oblanceolate-ellipitic, 4-12 cm long,
0.5-2. 5 cm wide, the lower often sharp-toothed, the upper reduced,
linear-lanceolate, entire. Panicles rather leafy, 7-43 cm long, 1-13
cm wide, the heads many, 2.5-4 mm high; bracts 10-15; receptacle
naked. Flowers 12-22, fertile ; rays 6-9, the disk flowers 6-12 ;
achenes 1 mm long. 2n— 36 (Keck 1946).

Characteristic of Wisconsin prairies, especially dry-meisic
(Curtis, 1959) and dry sand prairies, as well as deep-soil mesic,
and even moist prairies, often somewhat weedy on railroads and

220 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

roadsides. Flowering and fruiting from (late July) mid-August
through mid-October.

Widespread from Oregon to S. California, east to Indiana,
with seven, often intergrading subspecies recognized by Keck
(1946), in Wisconsin represented only by ssp. ludoviciana, here
fairly uniform (although certain collections, especially in Green
and Dane counties, with larger, more toothed leaves and tendencies
for the upper surface to become glabrate, suggest introgression
from A. serrata, a rarer species of moist prairies; cf. Keck l.c.).

12. Artemisia stelleriana Besser Beach Sagewort, Dusty

Miller Map 26.

Densely white-woolly , rhizomatous perennials with decumbent
to ascending, 3-7 dm tall stems. Leaves sessile, 1-2 times pinnatifid,
3-9 cm long, 2-5 cm broad, the segments broad or rounded.
Panicles narrow, spike-like, 9-30 cm long; heads it erect, large,
5-8 mm high; receptacle naked. Flowers 28-39, fertile; achenes 2
mm long. 2n=18 (Love & Love 1961).

Sand dune species native to N. Japan and N.E. Asia, widely
cultivated for its attractive foliage, in Wisconsin rarely escaped
but not established in sandy areas on Lake Michigan. Brown Co. :
Green Bay, cult., June 26, 1884, Schuette s.n. (F). Door Co.:
E. end of Sturgeon Bay Canal, in sand near lighthouse, June
1913, Davis s.n. (WIS). Sheboygan Co.: Elkhart, August 5,
1892, Schuette s.n. (F). The Duluth station is mapped from
Lakela (1965). Flowering and fruiting in July and August.

1966]

Mickelson and litis— Flora of Wisconsin

221

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Scrophulariaceae. Trans. Wis. Acad. Sci. Arts and Letters 40: 111-
138.

Smith, Huron H. 1923. Ethnobotany of the Menomini Indians. Bull. Public
Museum, Milwaukee 4: 30-31; 215.

SoLBRiG, Otto T. 1963. The tribes of Compositae in the Southeastern United
States. Journ. Arnold Arboretum. 44: 436-461.

Suzuke, Osamu. 1952. Chromosome numbers in Artemisia. 21st Annual Meet¬
ing of the Genetic Society of Japan.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN
NO, 56, COMPOSITAE V— COMPOSITE FAMILY V
TRIBE INULEAE

(Antennaria, Gnaphalium, Anaphalis, and Inula)

Edward W. Beals and Ralph F, Peters
Herbarium, University of Wisconsin

The following notes and maps of these, among the taxonomically
most difficult plants in Wisconsin, are based on collections in the
Herbaria of the University of Wisconsin — Madison (WIS), Uni¬
versity of Wisconsin — ^^Milwaukee (UWM), University of Minne¬
sota (MIN), and the Milwaukee Public Museum (MIL). We wish
to thank the curators of these herbaria for loan of specimens, Olive
Weber, Stephen Gilson, Frank Crosswhite, Harriet Peters, Katha¬
rine Snell and Carol Mickelson for aid in preparing the maps, and
Dr. H. H. litis for his cooperation and encouragement during this
study. Dr. Carroll Wood of the Gray Herbarium, Harvard Univer¬
sity, has been kind enough to check one of the Antennaria types.
Much of the herbarium work was supported by grants from the
Wisconsin Alumni Research Foundation, as were many field trips
on which Antennaria species have been collected. The field work
of one of us (E. Beals) was supported by Fellowship No.
GF-11,914, Division of General Medical Sciences, U. S. Public
Health Service.

On the maps, dots indicate the specific location where a specimen
was collected, triangles represent county records when the specific
location is not known. The months of flowering and fruiting, as re¬
corded on all the specimens observed, are shown in the lower left-
hand corner of each map. The distribution of dots reflects not only
the distribution of species but also the intensity of collecting.
Southern Wisconsin, especially Dane and Sauk Counties, is much
better represented than northern Wisconsin.

COMPOSITAE— TRIBE VI INULEAE

Anthers tailed at base; style-branches rounded or truncate, ex-
appendiculate ; leaves alternate; receptable naked or chaffy; heads
small, the corollas all tubular, or, in the large yellow heads of
Inula, the outer ligulate; plants more or less white-woolly; leaves
alternate.

223

224 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

KEY TO GENERA

A, Plants slender 1-5 (-10) dm tall, Heads 1 cm or less in diam.,
white or stramineous.

B. Cauline leaves few, strongly ascending, much smaller than
those of the persistent basal rosette; stolons creeping or
ascending; all plants dioecious _ 38. ANTENN ARIA.

BB. Cauline leaves many, about the same size as the basal
leaves which soon wither; stolons absent.

C. Phyllaries pure white, with conspicuous, longitudinal
creases creating the appearance of wrinkled tissue pa¬
per; plants dioecious, the fertile head often with a few
perfect but sterile flowers in center ; dry plants without
a strong odor _ 39. ANAPHALIS.

CC. Phyllaries grayish white, yellow, or brown, scarious,
with very small longitudinal ridges but no conspicuous
creases ; heads perfect ; dry plants with strong tobacco¬
like odor _ 40. GNAPHALIUM.

AA. Plants very large and robust, 1-2 m tall. Heads very showy,
the disk 3-5 cm in diam., yellow. Infrequently adventive with
large woolly leaves _ 41. INULA.

38. ANTENNARIA L, Pussy Toes, Everlasting, Ladies Tobacco
(By Edward W. Beals)

Perennial woolly herbs with alternate and basal leaves, char¬
acterized by many-flowered heads of all tubular dioecious flowers.
Heads often in congested inflorescences ; involucral bracts imbricate
in several series and often colored; receptacle flat or convex, not
chaffy. Pistillate flowers with filiform corollas, bifid styles, and a
row of capillary bristles united at the base. Achenes terete or
flattened. Staminate flowers with broader, lobed corollas, undivided
styles, caudate anthers, and thickened clavate or barbellate bristles.
A common genus occurring throughout the north temperate and
arctic zones and in southern South America, in Wisconsin among
the earliest of all spring flo,wers.

The genus has been subject to many taxonomic interpretations.
Linnaeus included two species of what we call Antennaria in the ^
genus Gnaphalium: G. dioica of the Old World and G. plantagini-
folia of Virginia, Brown (1818) transferred these to the genus An-
tennaria. Gray (1886) considered all Antennaria in eastern North
America as one variable species, A. plantag ini folia, while Barton
(1818) and Darlington (1853) segregated the narrower-leaved
plants as A. dioica, which is also the species in Europe. Greene
(1897) transferred the New World A. dioica to a new species, A.

1966]

Beals and Peters — Flora of Wisconsin

225

neglecta. As workers continued to study Antennaria, it was divided
into more and more species. The variability within the genus also
led to the recognition of numerous varieties and forms. Identifica¬
tion has become a very difficult procedure. On the other hand,
several workers (notably Cronquist, 1945, 1946) have tried to
simplify this confusion by combining taxa. Today the two extreme
views of this species complex are represented by Cronquist (in
Gleason 1952) and Fernald (1950) ; the former recognizes two
species whose range includes Wisconsin, and the latter, ten. Many
individual plants in Wisconsin exhibit such great leaf variation
that, using Fernald’s treatment, different mature leaves of the
same rosette may key out to different species!

The extreme variability and lack of well-defined taxonomic
groups in Antennaria can be correlated with widespread polyploidy,
apomixis, and very likely hybridization (Stebbins 1932a, 1932b).
However, unlike other confused, apomictic taxa, such as Rubus
and Crataegus, the original number of species contributing to the
present Antennaria complex in Wisconsin is small — perhaps only
two. Because of apomixis (the production of parthenogenetic seed),
the traditional concept of a species — a population which has shared
and will share genes freely— breaks down. To put species names on
these plants becomes a matter of convenience — they are simply
categories of plants with similar morphology and ecology. Such
naming cannot imply a future mixing of genes within the named
species — otherwise every apomictic plant would be called a new
species. Nor does it imply a sharing of genes in the immediate past.
It is quite probable that some apomictic ‘‘species,’’ similar mor¬
phologically and ecologically, are derived from several independent
occurrences of hybrdization and polyploidy.

The genus Antennaria in Wisconsin consists of a series of
morphological gradients. It would be helpful to taxonomic analysis
if the measurement of plants showed clustering along these gradi¬
ents, but in most cases there is no such clustering. Even the uni¬
versally accepted separation into narrow- and broad-leaved plants
puts many herbarium specimens with intermediate leaves on a
questionable border line. Morley (1958), in Minnesota, has made
the most determined attempt to segregate these two groups.

It is important, therefore, to remember that species names in
the genus Antennaria are convenient labels, and that there is
nothing sacred or inviolable about where the lines are drawn be¬
tween groups. For some work, finer lines may be drawn than for
other work. In the following discussion, we have divided the genus
about as finely as we dared for the purposes of describing the
geographic distribution in Wisconsin. The categories we have
chosen are usually distinguishable with adequate herbarium speci-

226 Wisconsin Academy of Sciences, Arts and Letters [VoL 55

mens (which should include mature rosette leaves, stalks with
flowers or fruits, and well-developed stolons) or with live plants
in the spring. These categories coincide somewhat with the tradi¬
tional categories (Fernald 1950), but they include several modifica¬
tions, There will always be borderline cases; however, with the
realization that species names are a matter of convenience and not
a matter of inviolable natural discretions, one can take less seri¬
ously the problem of labeling these borderline cases. This problem
could be more intelligently approached if we discarded species
names altogether and used some numerical system for identifying
Antennaria plants along one or more taxonomic gradients.

Chromosomes of a number of plants were observed. The basal
meristem of very young leaves provided the largest meristematic
cells for observation. The leaves were treated with a solution of
8-hydroxyquinoline (.29 g/1) for about four hours. They were
hydrolyzed without fixing with normal HCl at 60° for 10 minutes
and then washed. The mitotic region was smeared in aceto-carmine.
In diploid plants the chromosome count was 28. Because the
chromosomes in polyploids were too aggregated to count, it was
possible to distinguish only diploidy and polyploidy. Stebbins
(1932a, 1932b) has previously counted chromosomes of various
Antennaria species. Most of his polyploids, except for the tetraploid
A. neodioica, were hexaploid (chromosome number 84).

The Antennaria complex in Wisconsin includes two basic diploid
species, one broad-leaved, the other narrow-leaved, which are
usually quite easily distinguishable from each other. In addition
to these, there are various polyploids that cause confusion in two
directions. First, some of these are hard to distinguish from their
diploid ancestors. Second, among themselves there is every degree
of gradation between the two diploid progenitors. The polyploids
are all potentially apomictic, but several can apparently produce
seed sexually as well.

Figure 1 shows 108 flowering specimens with corolla length
(partially correlated with polyploidy) plotted against the width/
length ratio of the rosette leaves (indicator of hybridization) , Each
species is represented in proportion to the total number of female
specimens with flowers available, and specimens were selected at ^
random from the herbarium folders to make these measurements.

It should be noted that the two diploid groups, A, neglecta^ and
A, plantaginifolia, shown in Fig. 1 by solid black triangles and
circles respectively, remain distinct from each other, but that both
grade into the polyploid complex, especially A. neglecta. In Fig.

1, part of the apparent confusion in the polyploids is the result of
using only one character for the abscissa. Different polyploid
groups of Antennaria carry different characters which delineate

1966]

Beals and Peters— Flora of Wisconsin

227

A

0

☆

A. neglecta (22)
A. petaloidea (5)
A. canadensis (3)
A. neodioica (24)

M A. munda (7)

® A. plantaginifolia (5)

O A. fallax (21)

□ A. parlinii (21)

Figure 1. Distribution of two morphological characteristics of 198 randomly

selected female Antennaria specimens.

“species” arbitrarily chosen on the practical bases of recognizability
on herbarium specimens and of their relatively distinct and/or
qualitative nature. They are described in the key below.

The detailed study of the ecology and life history of the genus
in southern Wisconsin is reported elsewhere (Beals 1961), There
is some differentiation of species on the basis of their ecology, but
each species has rather broad ecological adaptability. Antennaria
is found in many plant communities, most prominently in oak
barrens, cedar glades, and dry-mesic prairies, but commonly also
in dry prairies, mesic oak forests, pine barrens and oak openings,
and in sand barrens, oak forests, pine forests, wet-mesic prairies,
and lake sand dunes. Glabrous forms are more common in woodland
than in the open, while tomentose forms are more common in open
vegetation. Antennaria increases under grazing pressure and, to a
lesser extent, with mowing. Factors which enable members of this

228 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

genus to compete successfully in various plant communities include
genetic diversity and polyploidy, drought resistance, wintergreen
leaves, which can photosynthesize while other iplants are dormant
if temperature and sunlight are sufficient, and the secretion of an
antibiotic into the soil which inhibits the growth of other species
of plants.

Key To Species

A. Rosette leaves with 1-3 prominent veins, the lateral veins if
present rarely prominent beyond broadest part of leaf.

B. Stolons prostrate, lash-like.

C. Cauline leaves with scarious appendages _

_ 1. A. neglecta.

CC. Cauline leaves without scarious appendages _

_ 2. A. petaloidea.

BB. Stolons short, ascending stiff.

D. Upper cauline leaves with scarious appendages; leaves

glabrous above _ 3. A. canadensis.

DD. Cauline leaves without scarious appendages; leaves

glabrous to pubescent above _ 4. A. neodioica.

AA. Rosette leaves with 3-7 prominent veins, the two main lateral
veins converging toward and nearly reaching the tip.

E. Stolons prostrate, lash-like _ 5. A, munda.

EE. Stolons short, ascending, stiff.

F. Involucre 4-7 mm high, pistillate corolla 4-6 mm long ;
staminate corolla 3-41/2 mm long; nodes on flowering

stem 3-5 _ 6. A. plantaginifolia.

FF. Involucre 6-8 mm high, pistillate corolla 5-8 mm long;
staminate corolla 4-5% mm long; nodes on flowering
stem 5-12.

G. Rosette leaves tomentose above _ 7. A, fallax.

GG. Rosette leaves glabrous above; stem often purple-
glandular _ 8, A. parlinii.

1. Antennaria neglecta Greene Maps 1, 2.

Female: Rosette leaves narrow, usually 1-veined, cuneate, lance¬
olate, or spatulate, often subpetiolar, 15-50 mm long, 5-15 mm
wide, sometimes becoming glabrous at maturity. Stolons spreading,
long and flexuous, the terminal rosettes opening tardily; stolon
leaves linear, 3-10 mm long. Flowering stems 4-30 cm high;
cauline leaves usually 3-4, linear, 12-30 mm long, 1-2 mm wide, the
upper with scarious appendage. Heads 1-7, in crowded inflores¬
cences at anthesis, developing (in typical form) into racemes as
flowers mature. Involucre 5-7 mm high; bracts white to purple,

1966]

Beals and Peters — Flora of Wisconsin

229

often with scarious tips. Corolla 4-6 mm long, regular or irregular ;
pappus 4-6 mm, achenes about 1 mm long.

Male: Similar, but heads remain crowded; involucre 4-6 mm
long, the bracts white and petaloid, without scarious tips ; flowering
stems 2-13 cm tall.

This diploid species is much more common in southern than in
northern Wisconsin, in pastures, sand and oak barrens, cedar
glades, oak openings, pine barrens, dry pine woods, dry oak woods,
and in all but the wet prairie types. The apparent absence of male
plants in NE Wisconsin raises interesting questions about repro¬
duction of this species.

Var. campestris may be differentiated by pistillate heads remain¬
ing crowded even when fruiting and by flowering stems generally
much shorter. Corresponding male plants are probably indistin¬
guishable from typical A. neglecta. The following may be of this
Great Plains variety: Douglas Co.: Superior, May 16, 1927, L. R.
Wilson .2104. (WIS) ; Boylston, May 29, 1927, L. R. Wilson 2087
(WIS). Jackson Co.: Black River Falls, June 11, 1947, D. F.
Gr ether 5 HO (WIS).

2. Antennaria petaloidea Fern. Map 3.

Female: Rosette leaves and stolons very similar to those of A,
neglecta. Flowering stems 8-35 cm high ; cauline leaves 5-8, linear,
8-25 mm long, 1-4 mm wide, without scarious tips. Head 1-10,
crowded to loosely corymbose or racemose. Involucre 6-9 mm high ;
bracts green, brown, or purple, tips not scarious. Corolla 5-7.5
mm long, regular or irregular; pappus 5.5-7 mm, achenes 0.8-1. 5
mm long.

This polyploid species, closely related to A. neglecta, though
nowhere very common, is widely distributed over Wisconsin in
prairies, pastures, and especially pine woodlands. Only female
plants are known from Wisconsin.

3. Antennaria canadensis Greene Map 4.

Female: Rosette leaves somewhat narrow, usually 1-veined, ob-
lanceolate to broadly obovate, short-^petioled to sessile, 20-40 mm
long, 5-20 mm broad, glabrous above. Stolons erect, short and stiff,
the large terminal rosettes developing early; stolon leaves linear,
5-20 mm long, 1-3 mm broad. Flowering stems 5-30 cm high;
cauline leaves 5-7, linear, 10-30 mm long, 1-2 mm broad, the
upper with scarious appendages. Heads 3-12, in crowded to loose
corymbs. Involucres 7,5-9 mm- long; bracts brownish or greenish,
sometimes with scarious tips. Corollas regular or irregular, 5-7.5
mm long; pappus 5-7 mm, achene 1-1.3 mm long.

230 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

1966]

Beals and Peters — Flora of Wisconsin

231

Male: Similar, except flowering stems shorter and involucral
bracts broader, with white petaloid tips.

This rather infrequent polyploid, also similar to A. neglecta, is
found predominantly in N. Wisconsin in pine barrens and wood¬
lands, oak woodlands, and occasionally in dry prairies. The only
male plant came from westernmost Wisconsin: St. Croix Co. : Pine
Lake, open isandy shores. May 31, 1960, litis 16,775 (WIS).

4, Antennaria neodioica Greene Map 5.

Female: Rosette leaves 1- to 3-veined (the lateral veins when
present never approaching the blade tip), oblanceolate to obovate,
15-40 mm long, 5-20 mm broad, tomentose to glabrous above, with
considerable variation in the same clone; petiole 4.5-8 mm long.
Stolons erect, short, and stiff, with early-developing terminal
rosettes; stolon leaves sessile, lanceolate, 6-25 mm long, 2-10 mm
broad. Flowering stems 15-40 cm high; cauline leaves 7-10, linear
to lanceolate, 12-30 mm long, 2-5 mm broad, without scarious ap¬
pendages. Heads 3-15 in crowded to loose corymbs. Involucres 5.5-
7.5 mm high; bracts green, purple, or brown, with scarious or
white-petaloid tips. Corollas 4.5-7 mm, regular or irregular; pap¬
pus 4-7 mm, achenes 0.7-1.3 mm long.

A polyploid species, intermediate between the narrow- and broad¬
leaved species, common throughout Wisconsin, especially in the
north, in dry to mesic prairies, oak and pine barrens or woodlands,
cleared-woodland pastures and cedar glades. No male plants have
been found in Wisconsin.

Stebbins (1935) postulated A. neodioica as derived from A. vir-
ginica, a diploid species of western Pennsylvania to western Vir¬
ginia. This would imply either that A. virginica became extinct
over a large area which would probably include Wisconsin, or that
A. neodioica spread from a relatively small region of the Appa¬
lachians, where it originated, to southeastern Canada, all of the
northeastern United States, and to the Midwest. Either way, a
drastic difference in success between diploids and polyploids must
be postulated. Wisconsin representatives of A. neodioica are as
easily explained as allopolyploid derivatives of A. neglecta and
A. plantaginifolia, since A. neodioica is intermediate in many char¬
acters between these two.

5. Antennaria munda Fern. Map 6.

Antennaria occidentalis Greene, of authors including Stebbins
who named many of our specimens.

Female: Larger rosette leaves 3- to 5-veined, the lateral veins ap¬
proaching the blade tip, broadly cuneate, spatulate, or obovate,
30-60 cm long, 10-30 cm wide, sparsely to densely tomentose

232 Wisconsin Academy of Sciences, Arts and Letters [Vol, 55

above ; tip of blade rounded or in forma FARWELLII subtruneate,
sparsely to densely tomentose above; petiole 5-15 mm. Stolons de¬
cumbent and elongated, sometimes flexuous, the terminal rosettes
developing early or late ; stolon leaves linear, sessile to subpetiolate,
5-20 mm long. Flowering stems 10-45 cm high; cauline leaves
5-11, linear to oblanceolate, 15-30 cm long, 1-5 mm broad, with¬
out scarious tip. Heads 4-15, in a dense corymb. Involucre 6.5-9
mm high; bracts green, brown, purple, with white, subpetaloid or
(occasionally) scarious tips. Corolla 5-7 mm long, regular or irreg¬
ular; pappus 6-8 mm, achene 1-1.5 mm long.

This polyploid, most common in south-central Wisconsin and ab¬
sent from the west-central part, is found mostly in prairies and
pastures. Only female plants have been collected. Although its
leaves would put it in the broad-leaved category, its stolons are
more or less like those of A. neglecta. Fernald (1950) included un¬
der the name A. munda plants with either “short and assurgent
or prolonged and decumbent” stolons, using the tapered base of
the rosette leaf blade to characterize the species. Those with short
and assurgent branches are hardly distinguishable from A. fallax
in Wisconsin, while the distinction between short- and long-
stoloned specimens is relatively clear. In light of this, I have
placed part of Fernald’s A. munda with my A. fallax and kept the
rest as a separate species A. munda, according to Fernald’s de¬
scription (1936) of the type specimen.

Two specimens, both from northwestern Wisconsin, which key
out to this species in the above key, would be called A. farwellii
Greene according to Fernald (1950). They are distinguished from
typical A. munda by the subtruncate leaf blades, but were tenta¬
tively called a form of A. munda (called A. farivellii on
map 6) because of the tapering of the leaf blade and the spreading
stolons. Since Greene’s description of A. farwellii (1898) described
the stolons as short and assurgent (which with reduction of rank
would make it a form of A. fallax), it is difficult to know where
these two specimens belong : Bayfield Co. : Siskowitt Lake, dry soil,
10 July 1938, Fassett 20015 (WIS). Sawyer Co.: Hayward, Lake
Windigo, June-July 1943, E. M. Gilbert s.n. (WIS).

6. Antennaria PLANTAGINIFOLIA (L.) Richards Maps 7, 8.

Female: Rosette leaves oblong, obovate to orbicular, 3- 7-veined,
25-60 mm long, 12-30 mm wide, canescent to glabrate above;
petiole 10-30 mm long. Stolons short, ascending, the terminal
rosettes opening early; stolon leaves linear to lanceolate, 15-25 mm
long. Flowering stalks 10-30 cm high; cauline leaves 3-5, linear
to lanceolate, 15-30 mm long, 3-10 mm wide, without scarious

1966]

Beals and Peters — Flora of Wisconsin

233

234 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

appendages. Heads 4-20, in dense to loose corymbs. Involucres 4-7
mm high, often purple at base with white tips. Corollas 4-6 mm,
usually irregular; pappus 5-5.5 mm, achenes 0.7-1. 5 mm long.

Male: Similar to female but flowering stalks 3-20 cm long; in-
volucral bracts broader and subpetaloid, and corollas 3-4.5 mm
long.

This species, predominantly diploid and sexual with male and
female plants occurring in about equal numbers, is most common
in S. Wisconsin, following the Mississippi and St, Croix Rivers into
the NW part of the state, and occurs in dry places such as oak
openings, sand barrens, pastures, and dry prairies. Some herbarium
specimens are on the borderlines between this and the next two
species.

7. Antennaria fallax Greene Maps 9, 10.

Female: Rosette leaves obovate to orbicular or often broadly
quadrangular, 3- to 7-veined, 40-70 mm long, 15-40 mm wide,
slightly to heavily tomentose above ; petioles 9-35 mm long. Stolons
short and ascending, the terminal rosettes developing early ; stolon
leaves linear to broadly lanceolate, 10-30 mm long. Flowering stalks
15-45 cm high; cauline leaves 5-12, linear to lanceolate, often
crowded below, 18-45 mm long, 3-10 mm broad, without scarious
appendages. Heads 5-22, in dense to loose corymbs. Involucres
6-8 mm high; bracts whitish, greenish, or purplish, sometimes
scarious ; corollas 5-7 mm, usually irregular but sometimes regular ;
pappus 6-8 mm, achenes 0.8-1. 5 mm long.

Male: Similar but flowering stalks 5-25 cm high, involucral
bracts broader, and corollas 4-5.5 mm long.

A polyploid species common throughout Wisconsin, except in the
north-central region, the male plants rather uncommon and only
near the rivers of the Mississippi Valley, especially the Wisconsin,
in most dry to mesic prairies, oak openings, and in almost all pre¬
viously grazed habitats, less common in ungrazed oak woods, in¬
frequent in pine barrens, and rare in sand and oak barrens.
Subglabrous specimens included here approach in character the
following species.

8. Antennaria parlinii Fern. Maps 11, 12.

Female: Rosette leaves obovate to orbicular or often broadly
quadrangular, 3- to 7-veined, 45-80 mm long, 20-55 mm wide, gla¬
brous and bright green above ; petiole 12-35 mm. Stolons short and
ascending ; terminal rosettes developing early ; stolon leaves linear to
lanceolate, 15-25 mm long. Flowering stalks 15-45 cm high, some¬
times purple-glandular; cauline leaves 5-10, linear to lanceolate,
often crowded below, 25-50 mm long, 5-12 mm broad, without scar-

1966]

Beals and Peters — -Flora of Wisconsin

235

ious appendanges. Heads 5-18, usually in dense corymbs. Involu¬
cres 6^8 mm high; bracts whitish, greenish, or purplish. Corollas
5-8 mm, irregular or regular; pappus 6.5-8 mm, achenes 0.8-15
mm long.

Male: Similar but flowering stalks 3-30 cm high, involucral
bracts broader, and corollas 4.5-5.5 mm long.

Common in the south, sparingly in the north, with male plants
very common in parts of the “Driftless Area'' and adjacent locali¬
ties in SW Wisconsin but absent elsewhere, predominantly in both
grazed and ungrazed oak woods, grazed dry prairies and other
pastures, but rare in ungrazed prairies, sand and pine barrens.
Although A. parlinii is closely related to A. fallax, the very charac¬
teristic glabrous upper leaf surface and occasional purple glands
on the stem suggest that introgression may have occurred in the
past with some unknown, now extinct species, (The glabrous- but
narrow-leaved, asexual, northern A. canadensis is an unlikely
candidate.)

39. ANAPHALIS DC. Everlasting

White-woolly herbs resembling Gnaphalium with many-flowered
corymbose dioecious heads or with a few sterile hermaphrodite
flowers in center of pistillate heads ; flowers all tubular ; receptacle
flat or convex, not chaffy. Leaves alternate, entire, sessile. A small

N. American and Asiatic genus, represented by a single species in
Wisconsin, which is highly polymorphic and widespread from N.
Asia to Eastern North America.

1. Anaphalis margaritacea (L.) C. B. Clarke

Pearly Everlasting. Map 13.

Erect unbranched perennial herb, 2-9 dm tall Leaves very vari¬
able, linear to linear-lanceolate, 5-11 cm long and 3-15 mm wide,
tomentose below, glabrous to tomentose above. Stem tomentum
gray-brown and spreading just below the heads, becoming
appressed and usually white lower on the stem. Heads 4-7 mm
high; phyllaries papery, pure pearly white with conspicuous longi¬
tudinal creases, imbricated in several rows. Pistillate flowers Ali¬
form with bifid style ; corolla brown at the base, yellow higher up,
with yellow-green, ascending lobes. Staminate flowers broader;
corolla yellow-green with both base and apex brown, the brown
corolla lobes recurved from the exserted anthers. Pistillate heads
with a few perfect, but sterile flowers in center. Achenes brown,

O, 4-0.8 mm long, conspicuously papillate under low power, terete
or somewhat ridged.

Common throughout northern Wisconsin in open, often sandy
places, most frequently on roadsides and lake shores, occasionally

236 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

in open forests. Flowering in July and August; fruiting in August
and September.

Fernald (1950) recognized four varieties, reflecting the tremen¬
dous variability the species shows throughout its range (and in
Wisconsin) in leaf number, width, and indument. Without careful
morpho-geographic analyses, however, .which though at present
lacking would be of great phytogeographic interest, these varieties
cannot clearly be defined, and therefore recognized, in Wisconsin.

40. GNAPHALIUM L. Cudweed, Everlasting,
Rabbits Tobacco

(By Ralph F. Peters)

Woolly slender herbs with many-flowered heads in corymbose or
capitate inflorescences. All flowers tubular, mostly pistillate and
very slender, a few central ones perfect and broader, the style bifid.
Pistillate corollas white or yellow ; perfect corollas green or yellow-
brown with brown base and brown or yellow lobes. Pappus a single
row of filiform white bristles. Phyllaries scarious, white or stramin¬
eous, in one to several rows. Receptacle flat, not chaffy. Achenes
0.4-0. 8 mm long, light to dark brown, wrinkled and flattened at
first, becoming terete and almost smooth as they mature. Leaves
alternate, entire, sessile or decurrent. A widely distributed, large
genus, especially in Andean South America and South Africa.

Key to Species

A. Heads 2-3 mm high, in capitate leafy-bracted clusters; upper
stems very densely white-floccose-tomentose, obvious to the

naked eye ; stems usually much branched, 1-2 dm tall _

_ 1. G. uliginosum.

A. Heads 4-6 mm high, capitate or corymbose ; upper stems with
appressed, or nearly microscopic loose-spreading tomentum;
stems erect, seldom branching except within a corymbose inflo¬
rescence, 1-10 dm tall.

B. Leaf bases decurrent; middle or lower stem with glandular-
hirsute pubescence 0.2-0. 5 (-1) mm long; leaves usually
10-15 times as long as wide, tapering gradually to an acute
tip; achenes distinctly papillate under high magnification __

_ 2. G. Macounii,

B. Leaf bases not decurrent; middle or lower stem with glan¬
dular-hirsute pubescence less than 0.25 mm long or lacking;
leaves usually only 7-10 times as long as wide, tapering more
abruptly to the acute tip; achenes ridged but glabrous and
not papillate _ 4, G. ohtusifolium.

237

1966] BeaU and Peters — Flora of Wisconsin

1. Gnaphalium uliginosum L. Low Cudweed. Map 14.

Loiv, usually much-branching or prostrate annuals, 4-20 (-30)
cm high. Leaves narrowly obianceolate, tomentose on both upper
and lower surfaces, 6-35 cm long. Upper stems with a very dense,
long, white-floccose tomentum, becoming appressed-lanate below.
Inflorescences densely capitate, usually several to a plant, each
including 2-16 small heads, these 2-3 mm high and greatly over¬
topped by their subtending leafy bracts ; phyllaries in 1 or 2 rows,
barely imbricated, narrowly acuminate, 1-2 long, stramineous, pale
brown or yellowish, often with green midribs; achenes papillate
under high magnification.

Introduced from Europe, frequent in disturbed or sandy, usually
damp and open places such as roadsides, lake shores, and pastures,
sometimes on ledges of sandstone cliffs, mainly in northern Wis¬
consin and along the major rivers and Lake Michigan. Flowering
from mid-June through September; fruiting from mid-July into
November.

Gnaphalium purpureum L. (Map 17), a common European
weed resembling G. uliginosum but with erect unbranched stems,
oblanceolate-ispathulate leaves, and “spicate"’ inflorescences is rep¬
resented by one undated Goessl collection from Sheboygan, very
likely from his garden.

2. Gnaphalinm macounii Greene Western Cudweed.

Gnaphalium decurrens Ives Map 15.

Erect biennial, 2-7 dm tall, unbranched except at the top. Leaves
usually ascending rather sharply, linear lanceolate to linear obian¬
ceolate, (7-) 10-15 (-20) times as long as wide, with relatively
broad bases, decurrent onto the stem as wings, tapering gradually
to an acute or acuminate tip; upper surface glandular, lower with
an appressed tomentum. Upper stem with a short, loosely spread¬
ing to appressed-lanate tomentum, grayish brown to white; lower
stem granular-hirsute. Inflorescence corymbose, generally not as
widely branching as in G. ohtusifolium; heads 20-100, 4-6 mm
high; phyllaries light gray to yellow, 1-4 mm long, imbricated in
2-4 rows, the outer ones broader than the inner. Achene papillate
under high magnification.

Common in northern Wisconsin in open places, and usually in
dry, sandy soil such as beaches, fields, sand pits, roadsides, gravel
banks, and edges of bogs and marshes. Flowering from late July
to early September ; fruiting from September to early October.

This western, Cordilleran species has apparently spread across
the northern United States and southern (Canada to New England

238 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

^ ANAPHALIS

MARGARITACEA

GNAPHALIUM
MACOUNII

I#. . • GNAPHALIUM O.
VAR SAXICOLA

o GNAPHALIUM

PURPUREUM
'JjGl

1966] Beals and Peters— Flora of Wisconsin 239

and Nova Scotia since the Pleistocene. It tends to be weedy and
most collections are recent, out of the past 30 years. The Racine
Co. station is based on two old (ca. 1860) sheets of Hale (WIS).

3. Gnaphalium obtusifolium L. X G. MACOUNII Greene

hollow circle, Map 15.

A single collection, intermediate between the parents in that the
leaves are but slightly decurrent and the achenes papillate suggest
it to be a hybrid : Lincoln Co. : open field, T 35N, R 7E, Sect. 17,
King, Sept. 11, 1951, F. C. Seymour 13229 (WIS).

4. Gnaphalium obtusifolium L. Catfoot, RabbiPs Tobacco

Maps 16, 17.

A very variable taxon with one abundant and two very rare vari¬
eties in Wisconsin.

Key to Varieties

A. Stem with extensive lanate or loose-spreading, non-glandular
tomentum.

B. Phyllaries in 3-5 rows ; stem closely appressed-lanate, occa¬
sionally with scattered, twisted flocks; 15 cm-1 m tall; com¬
mon throughout all but northernmost Wisconsin _

_ 4a. var. obtusifolium.

B. Phyllaries in 1-3 rows ; stem with loosely-spreading tomen¬
tum; weak-stemmed delicate herb 3-15 cm tall, very rare in

Wisconsin Dells region _ 4b. var. saxicola.

A. Stem glandular-hirsute with little or no tomentum _

_ 4c. var. micradenium.

4a. Gnaphalium obtusifolium var. obtusifolium Map 16.

Erect biennial, 15 cm-1 m tall. Leaves lanceolate, ascending or
spreading, (5-) 7-10 (-13) times as long as wide, glandular or
scabrous above, appressed-tomentose below, tapering to a sessile
but not decurrent base and rather abruptly to the acute tip. Stem
with a white tomentum, usually closely appressed-lanate, but occa¬
sionally in scattered, twisted flocks reminiscent of the Greek origin
of the generic name, meaning ‘docks of wool.’' Inflorescence corym¬
bose, of 20-150 heads, these 4-6 mm high ; phyllaries light gray or
tan to yellow, 1-5 mm long, imbricated in 3-5 rows, the outer
broader than the inner ones. Achenes wrinkled to ridged but
glabrous.

Throughout Wisconsin except in the extreme north, very common
in sandy soil of fields, roadsides, prairies, and occasionally in open,
dry woods. Flowering from August to early October; fruiting in
September and October.

240 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

4b. Gnaphalium obtusifolium var. saxicola (Fassett) Crong.

Gnaphalium saxicola Fassett, Rhodora 33 : 75. 1931.

[Type : Adams Co. : Sandstone ledges, Coldwater Canyon, Dells
of the Wisconsin River, Fassett, Uhler, & McLaughlin 9590 ( WIS) ]

Map 17.

Similar to var. obtusifolium but with very slender stems only
3-15 cm tall. Leaves usually oblanceolate, spreading, (3-) 4-9 times
as long as wide, and frequently but sparsely tomentose beneath.
Stems with a white, loosely spreading tomentum. Inflorescence with
1-15 heads; phyllaries much less imbricated, in 1-3 rows, usually
all narrow.

Very rare on mesic to dry sandstone ledges in the Wisconsin
Dells area. Flowering from late July to mid-September; fruiting
in September and October.

At the type locality in Coldwater Canyon, Peters recently found
it only on a single, south-facing ledge 5 m above the canyon floor.
At this point the canyon is about 50 m wide and rather marshy.
A black-top public path of the Dells Boat Company passes directly
below this ledge about 150 m before it enters the very narrow Cold-
water Canyon proper. The dry ledge is about 6 dm wide and 3 m
long and is overhung by another ledge 1 m above it. In 1963, less
than a dozen plants of var. saxicola, none taller than 6 cm were
scattered along this sandy, barren ledge. It is possible that there
are other colonies on similar, inaccessible ledges of the many sheer
cliffs which in this region commonly rise from the nearby Wiscon¬
sin River.

The earliest collection of var. saxicola was made in 1866 [by an
unknown collector (WIS)] at Congress Hall, now known as “Lost
Canyon’'. Here in 1963, Peters found only two plants, a little taller
than those of Coldwater Canyon, growing on a narrow ledge above
a smooth, 15-foot rock wall located in the west branch of the can¬
yon, at the turn-around for the horse-drawn wagons which carry
visitors through the rocky gorge, currently a popular tourist attrac¬
tion. Above the ledge, the cliff rises almost vertically for some dis¬
tance, with no overhang. Here, as in Coldwater Canyon, there is no
competition from other plants. This site is rather sunny and the
only known habitat that is not heavily shaded. The fact that var.
saxicola is just as depauperate here makes it extremely unlikely
that it is merely a form of var. obtusifolium stunted by the absence
of light, but rather that it is a cliff ecotype recently derived from
var. obtusifolium.

A few specimens (all in WIS) have been collected which are
intermediate to var. obtusifolium, especially in tomentum and/or
height : Sauk Co. : “crags,” Mirror Lake, shady coulee near Delton,

1966]

Beals and Peters — Flora of Wisconsin

241

1891, True s.n.. Iowa Co. : north facing wooded bluff along Wiscon¬
sin River opposite Lone Rock, 1930, Fassett 12549. Sauk Co. : end
of Lower Dells, Delton, 1949, Fassett 28120. Richland Co. : Boaz,
1959, Wasuwat s.n. These intermediates, coupled with the scarcity
of specimens, make it advisable not to follow Fassett (1931), who
considered it a separate species, but rather Cronquist (1946) who,
on a 1944 annotation slip on the 1866 sheet (WIS), wrote, ‘Tn my
opinion, G. saxicola Fassett constitutes a local variety of G. obtusi-
folium, with the True specimen being somewhat intermediate to
the typical variety.”

4c. Gnaphalium obtusifolium var. micradenium Weath.

Map 16.

Very similar to var, obtusifolium, but somewhat smaller, with
leaves only 1-5 mm wide. Stems with little or no tomentum, instead
covered with a very short (less than 0.25 mm) long glandular-
hirsute pubescence.

An eastern variety, represented in west-central Wisconsin by
only four specimens (all in WIS) : Adams Co. : oak-jack pine woods
10 mi NNE of Friendship, 1948 Brown 334. sandstone bluff, oak-
pine woods, 7 miles SSW of Adams, 1948, Brown 343. Jackson Co. :
black oak-sugar maple woods, north base of Bear Bluff, sandy soil,
1958, Witt. Vernon Co. : grazed bluff, Newman Valley, 1958, Melch-
ert & Witt s.n. (together with a plant of the typical variety).

41. INULA

A large Old World genus of small to robust yellow-flowered herbs,
with but one species escaped in Wisconsin.

1. Inula helenium L. Elecampane. Map 18.

Robust herbs 1-2 m tall from a stout mucilaginous rootstock;
basal leaves very large, on long petioles, the blades 4-5 dm long,
ca. 2 dm wide, woolly beneath, scabrous above, the upper similar
but reduced, sessile and cordate-clasping, ovate or ovate-lanceolate
and long acuminate. Heads few, solitary at tip of 2-15 cm long stout
peduncles, like a sunflower, massive, 2-3 cm tall, 3-5 cm in diam.,
the involucral bracts imbricate in several series, the outer folia-
ceous ; rays pistillate, yellow, showy, 2-4 cm long, the disk fls. per¬
fect. Pappus 7-10 mm long of brown-stramineous capillary bristles,
the achenes slender, 3-6 mm long glabrous, 4-angled. Occasional
escaped from cultivation and then often established in large col¬
onies in old fields, pastures, fencerows, roadsides and near old gar¬
dens, rarely in open woods, flowering from mid-July to late August;
fruiting into September.

242 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Bibliography

Barton, W. P. C. 1818. Compendium Florae Philadelphicae. voL 2. Corey and
Sons, Philadelphia.

Beals, E. W. 1961. Ecological life history of the genus Antennaria in south¬
ern Wisconsin. Ph.D. thesis. Univ. of Wisconsin, Madison,

Brown, R. 1818. Observations on the natural family of plants called Compos-
itae. Transactions of the Linnean Society of London 12:76-142.

Cronquist, a. 1945. Notes on the Compositae of the northeastern United
States. I. Inuleae. Rhodora 45:182-184.

- . 1946. Notes of the Compositae of the northeastern United States. III.

Inuleae and Senecioneae. Rhodora 48:116-125.

Darlington, W. 1853, Flora Cestrica. Kimber and Sharpless, Philadelphia.

Fassett, N. C. 1931. Notes from the Herbarium of the University of Wiscon¬
sin — VI. Rhodora

Fernald, M. L. 1936. Contributions from the Gray Herbarium of Harvard Uni¬
versity CXIII. VII. Memoranda of Antennaria. Rhodora 38:229-231.

- , 1945. Contributions from the Gray Herbarium of Harvard University

CLVII. I. Key to Antennaria of the “manual range”, Rhodora 47:221-
235; 239-247.

- . 1950. Gray^s Manual of Botany, ed. 8. Am. Book Co,, New York.

Gleason, H. A, 1952. The New Britton and Brown Illustrated Flora. Lancas¬
ter Press, Lancaster, Pa.

Gray, A, 1886. Synoptical Flora of North America. Smithsonian Institution,
Washington, D. C.

Greene, E. L. 1897. Studies in the Compositae V. Pittonia 3:172-185.

- . 1898. New or noteworthy species XXIII. Pittonia 3:343-349.

- . 1902. Some new northwestern Compositae. The Ottawa Naturalist 15:

278-279.

- . 1911. Antennaria in the Middle West. Amer. Midland Nat. 2:73-90.

Ives. 1819. American Journal of Science. 1:380-382,

Linnaeus, C, 1753, Species Plantarum. ed. 1. vol. 2. Holmiae.

Morley, T. 1958. Note on the distinction between the broad- and narrow-leaved
Antennarias of Minnesota. Rhodora 60:306.

Nelson, A. and Macbride, J. F. 1916. Western Plant Studies. III. Botanical
Gazette 61:46.

Stebbins, G. L., Jr. 1932a. Cytology of Antennaria. I. Normal species. Bot.
Gaz. 94:134-151.

- . 1932b. Cytology of Antennaria. II. Parthenogenetic species. Bot. Gaz.

94:322-345.

- . 1935. A new species of Antennaria from the Appalachian region. Rho¬
dora 37:229-237.

Weatherby, C. a. 1923. Some critical plants of Atlantic North America. Rho¬
dora 25:22-23.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 57
POLEMONIACEAE— PHLOX FAMILY

Dale M. Smith
Department of Biological Sciences
University of California, Santa Barbma

and

Donald A. Levin
Department of Botany
University of Illinois at Chicago Circle

Spring and early summer in Wisconsin are the principal seasons
of flowering of the Polemoniaceae. Although the plants are often
abundant, only three genera and seven species are found in Wiscon¬
sin, while the whole family comprises 18 genera and approximately
316 species (Grant 1959). Most of the genera are found in western
North America, but a few are amphitropical with both North and
South American representatives, and a few more are Eurasian,
with none endemic to eastern North America.

The members of the family found in Wisconsin show conspicuous
adaptive radiation. Some occur in riparian forests, others in upland
forests, and still others in prairies and sand-hills or even in weed
patches. Most species native to Wisconsin may occasionally be cul¬
tivated, with Phlox paniculata the common isummer-flowering gar¬
den phlox. Additional aspects of the natural history of the family
are well treated by Grant (1959), and Grant and Grant (1965).

POLEMONIACEAE A. L. De Jussieu Phlox Family

Perennial or annual herbs; leaves opposite or alternate, simple
or pinnately compound; flowers in terminal or axillary cymose
clusters, diffuse or nearly head-like, perfect. Calyx of five wholly or
partially united sepals, usually regular; sinuses between the lobes
often membranous ; corolla of five regular wholly or partially united
petals, broadly campanulate or salverform, contorted in bud ;
stamens 5, epipetalous, alternate with corolla lobes ; ovary superior,
3-celled, styles 1, or three partially united; stigmas 3, or 1, 3-lobed;
fruit a loculicidally dehiscent capsule; seeds 1-several per locule,
tending to become mucilaginous when wet.

1 Field work and preparation of manuscript supported in part by the Research Com¬
mittee of the University of Wisconsin, on funds from the Wisconsin Alumni Research
Foundation, by the Department of Botany, University of Wisconsin, Madison, and
by National Science Foundation grants GB-1227 and GB-2739 to the authors. The
aid of Hugh H. litis, Doris Bruch and Eunice Roe, University of Wisconsin Herbarium,
Madison, in preparation of text and distribution maps is gratefully acknowledged.

243

244 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

KEY TO GENERA

A. Leaves pinnately compound, leaflets 7--17 ; calyx becoming
enlarged, chartaceous in fruit; corolla broadly campanulate,
blue, _ 1, POLEMONIUM.

A, Leaves simple ; calyx not conspicuously enlarged in fruit, fre¬
quently breaking through intercostal membranes; corolla sal-
verform or narrowly trumpet-shaped.

B. Annual; calyx not breaking in fruit; corolla narrowly
trumpet-shaped; inflorescence of tightly crowded incon¬
spicuous flowers _ 2. COLLOMIA.

B. Perennial; calyx usually splitting along intercostal mem¬
branes as fruit matures; corolla salverform, conspicuous;
inflorescence large, open. _ 3. PHLOX.

1. POLEMONIUM L, Jacob's-Ladder

Twenty-three species, mostly perennial herbs, primarily of west¬
ern North America.

1. POLEMONIUM REPTANS L. Jacob’s-Ladder, Greek Valerian.

Map 1.

Perennial herb, 1=5 dm tall ; leaves pinnately compound ; leaflets
7=17, glabrous, villous or rarely glandular; inflorescence a loose
few-flowered corymbose cyme ; pedicels 4=8 mm long ; calyx broadly
herbaceous, accrescent in fruit, the lobes triangular, about equaling
the tube; corolla broadly campanulate, deep blue to near- white,
8=16 mm long, lobed to mid-point; stamens equally inserted, slightly
exceeded by length of corolla lobes ; style barely exceeding stamens.
2N=18.

Widespread in eastern United States deciduous forest but lacking
on the coastal plain, common in southern Wisconsin south of the
floristic Tension Zone, in rich often moist hardwoods of sugar
maple, basswood, elm and oak, northward in white pine-red maple,
frequently in low meadows, marshes and even sphagnum bogs, on
wooded bluffs of sandstone, rarely limestone, rarely in deep soil
mesic prairies (e.g. at Juda, Green Co. cf. Bray, R, ‘'Climax forest
herbs in prairie,'' Am. Midi. Nat. 58: 437, 1957), sometimes along
roadsides and railroad rights-of-way. Flowering from (late April)
earliest May to the second week of June, fruiting from late May to
August.

Wisconsin material of this taxon shows great variations in pubes¬
cence, e.g. density, trichome length, glandularity, which ranges from
totally glabrous to densely pubescent with long-glandular trichomes
on upper leaves, stems and inflorescences. Davidson (4950) con-

1966]

Smith and Levin- — Flora of Wisconsin

245

ducted progeny tests on variable material presumably from Penn¬
sylvania. At Berkeley, California, all of his plants were glabrous,
which led him to conclude that such variation was unworthy of tax¬
onomic recognition at any rank. Braun (1956) presented data dem¬
onstrating that a Polemonium localized in a few southern Ohio and
northern Kentucky populations did indeed have sufficient integrity
to warrant recognition as variety villosum E, L. Braun, which dif-
ered slightly in habit, leaf, and flower morphology. Braun demon¬
strated that the two varieties hybridize today where conditions are
appropriate and postulated that variation such as that noted in Wis¬
consin could have resulted from post-Pleistocene contacts between
var, villosum and var. reptans. Herbarium material from Wisconsin
and field observations on Illinois plants suggest a high degree of re¬
combination of the factors responsible for the determination of
pubescence traits, and their extremes as well as their intermediates
are evenly scattered throughout the range of the species in Wis¬
consin. Grant (1965) considers P. reptans an obligate outbreeder,
the condition being enforced by protandry and self-incompatibility,
with bumble-bees being the principal pollinators. This would tend
to maintain the genetic diversity of a mongrelized population pro¬
vided selective factors did not operate against recombinants.

The variation in Wisconsin seems not to include the extreme var.
villosum types described by Braun, which show an association of
several morphological and physiological characters with densely
glandular pubescence. Accordingly, all specimens examined fall
within F. reptans var. reptans.

2. COLLOMIA Nutt. Collomia.

Fourteen species, four perennials, 10 annuals, primarily of west¬
ern North America.

1. Collomia linearis Nutt. Collomia. Map 2,

Annual herb 1™6 dm tall, subglabrous below, strongly glandular
pubescent in inflorescence, unbranched or in vigorous specimens
branched above, each branch terminating in a leafy-bracted head¬
like cluster of sessile flowers; leaves mostly alternate, simple,
lanceolate to linear, narrowed to sessile base; calyx somewhat ac¬
crescent, to about 4 mm in fruit; corolla about 1 cm long, pale
pink, bluish or white; tube very slender; lobes very shorty ca. 1-3
mm long; stamens subequal, included; plants autogamous; seeds
mucilaginous when wet, 1-3 per capsule. 2n=16.

A Great Plains and western N. Am. mountain element, apparently
native but rare in sands and gravels in Wisconsin ; but this is diffi¬
cult to prove, since it occurs mostly in ''open habitats'' along rail-

246 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

roads, river and lake shores, sometimes in clearings in forest, and
docks and waste places in cities. Flowering from late June through
July (Oct.), fruiting from, early July through August.

3. PHLOX L. Phlox, Wild Sweet-William.

Erect or spreading perennial herbs (subshrubs or annuals not in
Wisconsin) ; leaves opposite, simple, entire, sessile, linear-lanceolate
to ovate; flowers prominent, pinkish or rarely blue or white, in
terminal, variously-shaped cymose clusters; calyx five-lobed, the
lobes acute or aristate, herbaceous, the sinuses membranous, often
rupturing as the 3-valved capsule enlarges; corolla showy, salver-
form, the lobes variously shaped, often with prominent eyemark-
ings; stamens unequally inserted in corolla tube; anthers usually
included in tube; style elongate and exceeding the calyx lobes
(rarely exserted from corolla) or shorter than calyx lobes; seeds
not mucilaginous when wet. Sixty-six species North American, one
Asiatic, all but three perennial.

Isolating Mechanisms in Phlox.

Geographical. The distribution of phlox in North America is bi¬
partite, with all Wisconsin taxa belonging to the eastern 25% of
the species. Within this group, certain taxa have greater affinity
with the Appalachians, others with the Ozarks, while in some geo¬
graphical affinity is obscure. The only Wisconsin phlox of appar¬
ent Appalachian origin is P. glaherrima subsp. interior, the west¬
ernmost lowland derivative of the Appalachian P. glaherrima,
Ozarkian elements are P. bifida, P. divaricata subsp. laphamii and
P. pilosa subsp. fulgida. The latter seems to have diverged directly
from P. pilosa subsp. ozarkana, while P. divaricata subsp. laphamii
is probably a hybrid derivative of P. divaricata subsp. divaricata
and P. pilosa subsp. ozarkana. Of the remaining Wisconsin phloxes,
the ancestry may not be readily traced. The few sporadic records for
P. paniculata are probably escapes from cultivation. Phlox divari¬
cata subsp. laphamii and P. pilosa subsp. fulgida, the only sympa-
tric species, are ecologically isolated and no interspecific hybridi¬
zation in Wisconsin has been seen.

Ecological. Ecological isolation is moderate to strong in all the
phlox species in Wisconsin. In the Kenosha Prairie where P.
glaherrima subsp. interior co-exists with P, pilosa subsp. fulgida,
their requirements differ sufficiently so that populations of the two
are usually contiguous rather than mixed, with wetter prairies oc¬
cupied by P. glaherrima. Apparently P. pilosa subspecies are not
isolated where they contact, but their generally separate ranges

1966]

Smith and Levin — Flora of Wisconsin

247

suggest that even they are ecologically differentiated. Phlox
paniculata typically occurs in moist woodland (as an escape).
Strongly sympatric in Wisconsin are P. divaricata laphamii of
woodlands and P. pilosa fulgida of prairies, these virtually totally
isolated ecologically.

Seasonal, The flowering periods of the respective species are
shown on each map. Throughout the seasonal sequence overlapping
occurs between species, but the flowering peaks are amply distinct.
Even with slight overlap, ecological isolation or flower constancy
of pollinators is usually sufficient to provide effective reinforce¬
ment of seasonal barriers. An interesting exception to typical
seasonal isolation occurs in Kenosha County, where a late-flowering
race of P. pilosa subsp. pilosa {P. argillacea Clute) overlaps for
nearly three weeks the flowering period of P, glaberrima subsp.
interior. In a similar population in northern Illinois where cross¬
pollination was documented, the pollen load carried interspecifically
was very low and no hybrids were found. However, artificial hy¬
brids have been produced, so that even the limited pollen transfer
may have abiding significance.

Incompatibility, Controlled pollinations involving all Wisconsin
Phlox taxa have demonstrated isolation by cross-incompatability.
Seed-set failed whenever P, bifida was involved, and unilateral in-
compatability was encountered whenever pollen from short-styled
species (P, divaricata and P, pilosa) was applied to long-styled
species (P. glaberrima and P, paniculata) , Even when seed-set oc¬
curred, reduced numbers of seeds indicated varying degrees of
incompatability. The weakest barriers were between the members
of section Protophlox, P, divaricata and P, pilosa, while inter¬
sectional crosses always presented a strong or absolute barrier.
Within Section Phlox, represented in Wisconsin by distantly re¬
lated species, barriers were also strong or absolute. (Fig. 1, cf. p.
252).

Key to Species

A. Style elongate, equaling or exceeding calyx lobes (Section

Phlox) _ B.

B. Plants erect, unbranched from base, leaves linear-

lanceolate to ovate, corolla lobes rounded or apiculate _ C.

C. Leaves elliptic-lanceolate to ovate, veins very prominent
and areolate on abaxial surface, corolla-tube pubescent,
anthers and pollen cream- white _ 1. P. paniculata.

CC. Leaves linear-lanceolate or slightly broader, veins ob¬
scure, corolla tube glabrous, anthers and pollen orange
_ 2. P. glaberrima subsp. interior.

248 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

1966]

Smith and Levin — Flora of Wisconsin

249

BB. Plants spreading, branched throughout, corolla lobes con¬
spicuously bifid, notched 3--5 mm _3. P. bifida suhsp. bifida.

AA. Style short, included deeply within calyx (Section Protophlox
Wherry)

D, Sterile basal branches conspicuous, their leaves broadly
elliptic, subevergreen, leaves of fertile branches herbace¬
ous, acutish with somewhat cordate bases ; flowers mostly
tending toward blue, corolla-tube glabrous, sepals acute.
_ 4. P. divaricata subsp. laphamii.

DD. Sterile basal branches absent or inconspicuous, all leaves
linear to linear-lanceolate, flowers usually some shade of
pink, usually with conspicuous eye-markings, corolla-tube

pubescent, sepals with aristate tips _ 5. P. pilosa.

1. Phlox paniculata l. Summer Phlox. Map 3.

Perennial herb, 10-20 dm tall, from short rhizome ; stem glabrous
to pubescent, green or streaked with red; leaves sessile or short-
petioled, lanceolate to ovate or elliptic, strongly areolate-veiny on
abaxial surface, glabrous or pubescent ; inflorescence a large cluster
of smaller cymes; calyx lobes short-aristate, intercostal mem¬
branes often folded ; corolla tube elongate, pubescent, petals mostly
dark-pink; style elongate, usually exserted along with one or two
cream-white anthers. 2n=14.2

Widespread in the eastern U. S. north of the Coastal Plain to
Chicago and northern Missouri, escaped in southern Wisconsin on
wooded roadsides and ditches, grazed woods, railroad tracks and
old cemeteries, in the East Wilderness Scientific Area of Wyal-
using State Park appearing native in the midst of flood plain
forest, but this close to the ‘Immigrant TraiF' and hence, too, ad-
ventive (sub litis 20,575). Flowering from late July into October.
This is probably the most extensively cultivated of all phloxes, and
often the unaltered wild plants are grown in perennial summer
gardens. Escapes from cultivation are common within and without
its range, the precise area of native distribution therefore not likely
to be determined.

2. Phlox glaberrima l. subsp. interior Wherry Smooth Phlox.

Map 4.

Perennial glabrous erect herb, 3-8 dm tall, from short rhizome ;
stems green; leaves linear-lanceolate to lanceolate, 5-10 cm long,
broadening upward; inflorescence a nearly flat-topped cluster of

2 Chromosome morphology of eastern North American Phlox has been reported in
detail by Smith and Levin (1967).

250 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

cymes ; calyx lobes subulate, the intercostal membranes flat ; corolla
glabrous, brilliant rose-pink ; style elongate ; anthers orange.
2n=14.

A subspecies of the interior low plateaus from Arkansas to north¬
western Ohio, in Wisconsin only from the low prairies along Lake
Michigan in Kenosha (Chiwaukee Prairie and Carroll Beach south
of Kenosha) and Racine County [wet meadows, 4 mi. E of
Rochester, July 21, 1843, /. A. Lapham s.n. (WIS)] and adjoining
Beach, Illinois, together with Allium cernuum, Thalictrum revolu-
turn, Liatris spicata, Fimbristylis drummondii, and other species
restricted in Wisconsin to this area, once common on low prairies
in Illinois and Indiana, the typical subspecies southern Appalachian.
Flowering from latest June through July.

3. Phlox bifida Beck subsp. bifida Ten-point Phlox. Map 4.

Perennial, low, mat-forming herb, 1-2 dm tall, the base persist¬
ent; leaves 6-10 per shoot, linear to linear-lanceolate, 1-2 cm long,
glabrous to slightly glandular near the few-flowered inflorescence ;
pedicels 15-30 mm long, glandular; calyx lobes subulate, the mem¬
branes mostly flat; corolla lobes deeply bifld (notched) for 4 mm,
light blue or white, the eye-markings prominent; style 1 cm long,
slightly exceeding calyx lobes. 2n=14.

An Ozarkian (there somewhat calcophilic) species, in southern
Wisconsin at very edge of range and very rare [Ozaukee Co. :
Cedar Creek, no date, C. T. Tracy s.n. (WIS). Rock Co.: sandy
woods, SWl^, Sec. 33, T. IN, R. HE, ca. 6 mi. W of Beloit, May
12, 1946 (full flower) ,E.W.& G. B. Fell Jf6253 (WIS) ] , the second
collection probably native, the species much cultivated together
with the deep purple-red P. suhulata, the Moss Phlox, which it
resembles and to which it is closely related, and their hybrids.

4. Phlox divaricata l. subsp. laphamh (Wood) Wherry

Phlox divaricata y?? laphamii Wood, Class Book of Botany ed. 2:
439. 1848 (Lectotype: Milwaukee, Wisconsin, Lapham s.n., WIS!).

Blue Phlox, Wild Sweet-William. Map 5.

Perennial herb, 2-5 dm tall; decumbent sterile stems subever¬
green, often rooting-at the nodes ; erect stems bearing a moderately
compact cymose inflorescence; leaves on sterile shoots somewhat
elliptic, those of the fertile branches cordate-based, lanceolate, ses¬
sile, pubescence glandular, 25-50 x 10-20 mm ; calyx lobes subulate,
the membranes mostly flat ; corolla usually bluish-purple, violet, or
white (in forma albiflora Farw.), the corolla lobes truncate,
rounded, or apiculate; style 1. 5-3.0 mm long, included. 2n=14.

Widespread in the central U. S. to southern Georgia and eastern
Texas and centering on the Ozarks (hence somewhat calcophilic).

1966]

Smith and Levin — Flora of Wisconsin

251

in Wisconsin locally very abundant south of the Tension Zone, es¬
pecially in damp flood plain forests (e.g. Avon bottoms, Green Co.,
where in early spring the whole forest floor is tinged blue), rich
mesophytic hillsides of sugar maple, beech, and basswood, some¬
times on sandstone bluffs, much cultivated in wild flower gardens.
Flowering from earliest May to early June.

This Phlox is well-defined in Wisconsin. In Illinois, Indiana and
Kentucky, variation in stature, leaf size and shape, development
of sterile basal branches, and most conspicuously, degree of petal
notching, are prominent. Wherry (1955) hypothesized that post-
glacially the two subspecies migrated, P. d. ssp. divaricata from
the Appalachians, and P. d. ssp. laphamii from the Ozarks, to over¬
lap and hybridize in an area roughly corresponding to the Illinois-
Indiana state line, where maximum variability occurs. Levin
(1966) postulates that P. divaricata subsp. laphamii arose in
Ozarkia through hybridization between P. divaricata and P. pilosa
ssp. ozarkana Wherry. The P. pilosa genes gave P. divaricata the
morphological attributes of P. d. ssp. laphamii and physiological
attributes which assured success in the forest-prairie border region.

5. Phlox pilosa l. Downy Phlox, Prairie Phlox.

Perennial herb, 2-6 dm tall, from a subligneous base; stem usu¬
ally pubescent; sterile shoots sparse, not rooting at nodes, not
evergreen; leaves linear to linear-lanceolate, generally pubescent;
inflorescence a rather compact cluster of cymes ; calyx lobes
aristate, membranes flat ; corolla typically pink, usually with promi¬
nent eye-markings; tube pubescent; styles short, 1. 5-3.0 mm, in¬
cluded within calyx.

One of the most variable species of Phlox, rather close to P.
divaricata, with two subspecies in Wisconsin.

Key to Subspecies

A. Plants glandular pubescent, at least in inflorescence ; rare _

- 5a. P. pilosa ssp. pilosa

AA. Plants with lustrous non-glandular pubescence ; very common
- 5b. P. pilosa ssp. fulgida.

5a. Phlox pilosa L. ssp, pilosa Map 6.

Throughout the E. United States, occurring sympatrically
with all subspecies except P. p. ssp. riparia in western Texas, the
subspecies ecologically and morphologically defined, but often
hybridizing in areas of sympatry, in general in dry forest, wood¬
land borders, prairies, or areas of slight disturbance, in Wiscon¬
sin rare, reported from deciduous woods, wooded shady slopes.

252 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

open sands with Lupinus perennis, limestone cliffs, or roadsides;
differentiated from P. pilosa ssp. fulgida by glandular pubescence
and more reflexed calyx lobes. 2n=14.

5b. Phlox pilosa L. ssp. fulgida Wherry Map 7.

Strictly a prairie subspecies, widespread from Kansas to Mani¬
toba and Indiana, not repopulating disturbed areas where prairie
has been eliminated as does the preceding subspecies, in Wisconsin
south of the Tension Zone (except in some sand areas), in a wide
variety of prairie habitats from low damp to dry and calcareous
‘‘Goat Prairies’", in open sandy oak-savannah with prairie flora,
limestone bluffs, open oak woods, railroad right-of-way relic
prairies, in recently burned jack pine stands; one of the showiest
spring wild flowers, sometimes pollinated by day-flying dear-wing
Sphingids. Flowering from mid-May to early (mid-) July, fruiting
from late June to late July.

The fine, lustrous, non-glandular pubescence, which is the only
feature differentiating this subspecies from ssp. pilosa, seems to
be too insignificant on which to base a subspecies; furthermore,
occasional non-glandular plants are found in other subspecies.
However, because of eco-geographical differences associated with
non-glandularity, recognition as a subspecies seems justified. The
two Wisconsin subspecies of P. pilosa intergrade in a broad diago¬
nal band across Illinois and some of this influence may be noted in
southern Wisconsin. 2n=14.

1966]

Smith and Levin — Flora of Wisconsin

253

GILIA R. & P. is not included as part of the flora of Wisconsin,
although one species, G. multicaulis Benth. {—G. achilleae folia
Benth,) was collected once in Two Rivers [Manitowoc Co. : no date.
Coll. E. Dapprich s,n, (MIL), det. L. Constance 1963]. The species
is native to the South Coast Range of California (Grant 1954) and
has previously been reported as a garden escape in the eastern
United States (Wherry 1936). Ipomopsis rubra of the southeastern
United States was once collected [Adams Co. : Aug. 19, 1937. Coll.
J, W. Thomson s.n. (WIS)], no doubt an adventive from a garden,
for the species has a long record of escapes from cultivation in the
midwestern states.

Literature Cited

Braun, E. Lucy, 1956. Variation in Polemonium reptans, Rhodora 58:103-116.
Davidson, J. F. 1950, The genus Polemonium (Tournefort) L. Univ. Calif.
Publ. Bot. 23:209-282.

Grant, V. 1954. Genetic and taxonomic studies in Gilia, IV. El Aliso 3:1-18.

- , 1959. Natural History of the Phlox Family. Vol. 1, Martinius Nyhoff,

The Hague, Netherlands.

- . and Karen A. Grant. 1965. Flower Pollination in the Phlox Family.

Columbia Univ. Press. New York.

Levin, D, A. 1963. Natural hybridization between Phlox maculata and P.
glaherrima and its evolutionary significance. Amer. Jour. Bot. 50:714-720.

- . 1966. Variation in Phlox divaricata. Evolution 19: (in press).

Smith, D. M. and D. A. Levin, 1967, Karyotypes of eastern North American
Phlox. Amer. Jour. Bot. 54: (in press).

Wherry, E, T. 1936. Miscellaneous eastern Polemoniaceae. Bartonia 18:52-59.

- ^ — . 1955. The Genus Phlox. Morris Arbor. Monogr. Ill, Philadelphia

174 pp.

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN NO. 58
HYDROPHYLLACEAE—WAIERIEA? FAMILY

Jack W. Shields
Dept, of Botany
University of Wisconsin— Milwaukee'^

The distribution maps of species in Wisconsin were compiled
from collections in the herbaria of the University of Wisconsin-
Milwaukee (UWM), the University of Wisconsin (WIS), the Uni¬
versity of Minnesota (MIN), and Milwaukee Public Museum
(MIL), as well as some records in southwestern Wisconsin and ad¬
joining states from the unpublished thesis of Hartley (1962) . Num¬
bers within the enclosures in the lower left-hand corner of each
map represent the number of specimens that were flowering or
fruiting in the respective months. Those which were flowering and
fruiting simultaneously are included in both enclosures for that
month. The habitats in Wisconsin are from data on the herbarium
specimen labels.

Grateful acknowledgment is made to Professors Alvin L. Throne,
Hugh H. litis, Gerald B. Ownbey, and Albert E. Fuller for the use
of their herbarium facilities for this study; and to Professors
Throne, Hugh H. litis and Peter J. Salamun for their assist¬
ance and advice in the preparation of this report. Assistance of
Doris Bruch and Eunice Roe, the Research Committee of the Uni¬
versity of Wisconsin on funds from the Wisconsin Alumni Research
Foundation, and of the Federal Work Study Program are grate¬
fully acknowledged.

HYDROPHYLLACEAE—WATERLEAF FAMILY

Annual, biennial, or perennial, more or less pubescent, succulent
herbs. Leaves alternate, lobed or pinnate; flowers solitary or in
scorpioid cymes, small to medium-sized, white to blue or purple,
regular, perfect, hypogynous. Sepals 5, distinct or connate at base,
persistent; corolla tubular to rotate, 5-lobed about to middle;
stamens 5, inserted on corolla-tube, alternate, included or exserted ;
ovary entire, 1-locular, with 2 parietal, 1-4-ovuled placentae; style
1, cleft at summit. Capsules with 1-4 seeds.

KEY TO GENERA

A, Flowers in terminal cymes or axillary scorpioid cymes ; leaves
alternate.

B. Corolla lobes convolute in the bud; calyx sometimes with
auriculate sinuses ; blades of median stem leaves generally

* Present address: Dept, of Botany, University of Minnesota, Minneapolis 55455.

After Sept. 1: Dept, of Biology, Macalester College, St. Paul 55101.

255

256 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

more than 8 cm long*, deeply pinnate or palmate, basal

leaves long-petioled ; perennials or biennials _

_ 1. HYDROPHYLLUM.

BB. Corolla lobes imbricated in the bud; calyx sinuses without
auricles ; blades of median stem leaves generally less than
8 cm long, pinnate-pinnatifid, basal leaves not distinctive ;

very rare adventive _ 3. PHACELIA.

AA. Flowers solitary; leaves opposite below _ 2. ELLISIA.

1. HYDROPHYLLUM L. Waterleaf

Herbs with large, lobed or divided leaves and succulent stems;
cymes terminal, one-sided, several- to many-flowered, repeatedly
forked. Corolla campanulate; lobes erect or somewhat spreading;
stamens slender, usually villous; ovary 1-locular, with 2 large
dilated placentae; ovules few; style 1, slightly bifid. Capsule glo¬
bose, pubescent. Small genus of 8 North American species.

Key to Species

A. Perennial forming dense clones; principal cauline leaves
deeply pinnately divided; calyx sinus naked; peduncle and
pedicels subglabrous; fruiting inflorescences rather dense, to

8 cm across _ 1. H. VIRGINIANUM.

AA. Biennial, with stems solitary; principal cauline leaves pal-
mately divided to shallowly palmately lobed; calyx with re¬
flexed appendages in sinuses ; peduncles and pedicels pu¬
bescent, fruiting inflorescence very open, 10-20 cm across _

_ 2. H. APPENDICULATUM.

1, HYDROPHYLLUM VIRGINIANUM L. Virginia Waterleaf; John’s

Cabbage. Map 1.

Perennial from horizontal rhizome; stem 2-6 dm tall; stem,
cymes, pedicels and sepals pubescent with appressed to ascending
hairs up to 2 mm long ; leaves pinnate almost to midvein, 5-7 lobed,
broadly ovate or triangular, the terminal lobes and basal pair often
with 2-3 additional lobes, acute or acuminate apices; cymes very
dense. Sepals separate below the middle, narrowly linear, sparsely
hirsute-ciliate ; corolla white to purple, 7-10 mm long; stamens
long-exserted. Seeds (l-)2. 2n=18 (Wilson 1960).

Widespread from the Appalachians and Ozarks to Canada, in
Wisconsin common throughout except the northern tier of counties
in mesic or damp, shady rich deciduous woods of maple-elm or
beech, oak-basswood, or aspen-birch-red maple, in wooded pastures
and bluffs, thickets, and on stream banks and river bottoms. Flow¬
ering from mid-May to early July; fruiting from late June to
August.

1966] Shields — Reports on Flora of Wisconsin 257

2. Hydrophyllum appendiculatum Michx. Maple-leaved

Waterleaf. Map 2.

Biennial from a thin taproot, with lateral fibrous roots ; stem 2-6
dm tall, densely pubescent with both short, slender and spreading
pilose hairs 2-3 mm long ; upper cauline leaves shallowly palmately
5-lobed, truncate or cordate, the basal leaves pinnately 5-7 lobed
(the lowest with additional 2 remote lobes) ; cymes loosely flowered.
Sepals free nearly to base, lanceolate, densely hirsute, alternating
with small reflexed appendages; corolla lavender to pink-purple,
9-14 mm long; stamens equalling or longer than corolla. Seeds 1.
2n==18 (Wilson 1960).

An Ozarkian element widespread in the central US, in southern
Wisconsin in rich deciduous woods of beech-maple, on wooded oak
hillsides, and in riverbottom forest. Flowering from late May to
August, and fruiting from July through August.

2. ELLISIA L., Ellisia

Ellisia nyctelea L. Ellisia. Map 3.

Slender somewhat succulent annual, 1.5-3 dm tall, usually
branched from base; stems weak, glabrous or very sparsely hispi-
dulous; leaf blade ovate-oblong, 4-6 cm long, pubescent, pinnately
divided nearly to the mid-rib into 7-11 widely spreading, oblong
lobes; flowers solitary; sepals triangular-lanceolate; corolla white,
narrowly campanulate, 5-8 mm long, the lobes and the glabrous
stamens shorter than the tube ; ovary sub-globose, 1-locular but the
2 parietal placentae expanding and completely surrounding the 4
ovules. Capsule globose, much exceeeding the rotate accrescent
calyx, about 1 cm in diam., 4-seeded; fruiting pedicels drooping,
2-3 cm long. 2n=20 (Wilson 1960).

A monotypic genus of the prairies and plains from the Rocky
Mountains to the Ozarks and Indiana, with an isolated population
on the northern Atlantic slope, in Wisconsin somewhat weedy in
moist locally disturbed habitats such as sandy areas along stream
banks, eroding slopes to moist maple-basswood woods, chiefly in
the main drainage areas of the Mississippi, Wisconsin, Sugar, Rock
and Fox Rivers and the shore of Lake Michigan. Flowering from
mid-May to early June; fruiting from late May through June.

3. PHACELIA Juss. Scorpion Weed, Phacelia

Annuals or biennials with alternate, lobed to pinnatifid leaves,
flowers in dense strongly scorpoid cymes, of perhaps 100 species,
none native of Wisconsin.

258 Wisconsin Academy of Sciences, Arts and Letters [Vol. 55

Phacelia distans Benth. (det. L. Constance 1963), a slender
hispid annual ,with small pinnate-pinnatifid leaves, dense, pro¬
nouncedly scorpoid, paired cymes, 1 cm long flowers with exerted
stamens, and an endemic of California, Nevada, and Arizona, has
been collected once “along shore of Lake Superior near Herbster’’
[Bayfield Co.: July 15, 1951. H. A. Davis 9571 (MIL)], no doubt
an accidental introduction,

Phacelia franklinii Gray, a 1-6 dm tall, usually unbranched
sparsely pubescent biennial or annual with once pinnate or pinnati-
fid leaves, several scorpoid racemes, small blue to white flowers
with hairy filaments, is a northwestern North American species

1966]

Shields — Reports on Flora of Wisconsin

259

from Alaska to Wyoming. Occurs commonly from Duluth eastward
on the north shore of Lake Superior (cf. Lakela 1965, Gillett 1960,
Fig. 4) and can be expected in Wisconsin on gravelly lake shores.

Phacella purshii Buck!., listed by both Fernald (1950) and
Gleason (1952) for Wisconsin, ranges only as far north as central
Illinois (Constance 1949).

Literature Cited

Constance, L. 1940. The genus Ellisia. Rhodora 42:33-39.

- . 1942. The genus Hydrophyllum L. Amer. Midi. Nat. 27:710-731.

- . 1949. Revision of Phacelia, Subgenus Cosmanthus (Hydrophyllaceae) .

Contr. Gray Herb. 168:1-48.

Deam, Charles C. 1940. Flora of Indiana. Department of Conservation,
Indianapolis.

Fernald, M. L. 1950. Gray's Manual of Botany, Ed. 8. American Book Co.,
New York.

Gillett, George W. 1960. A systematic treatment of the Phacelia Franklinii
group. Rhodora 62:205-222.

Gleason, H. A. 1952. The New Britton and Brown Illustrated Flora, 3:102-
108. Lancaster Press, Lancaster, Pa.

Hartley, Thomas G. 1962. The Flora of the “Driftless Area.’’ State University
of Iowa, Botany Ph.D. Thesis (MS).

Lakela, Olga. 1965. A Flora of Northeastern Minnesota. University of Minne¬
sota Press, Minneapolis.

Wilson, K. A. 1960. The genera of Hydrophyllaceae and Polemoniaceae in the
Southeastern United States. Jour. Arnold Arh. XLI: 197-212.

r

■;

1

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ij

\

OFFICERS OF THE WISCONSIN ACADEMY OF
SCIENCES, ARTS AND LETTERS

President

David J. Behling
NML Insurance Co.
Milwaukee, Wisconsin

Vice-President (Sciences)

Jacob Shapiro
Depai-tment of Biology
Wisconsin State University,
Oshkosh

Vice-President (Arts)

James A. Schinneller
Department of Art
University of Wisconsin
Extension — Milwaukee

V ice-P resident ( Letters )

Frank L. Klement
Department of History
Marquette University,
Milwaukee

President-Elect

John W. Thomson
Department of Botany
University of Wisconsin —
Madison

Secretary

Eunice R. Bonow
Department of Pharmacy
University of Wisconsin —
Milwaukee

T reasurer

Norman C. Olson
NML Insurance Co.

Milwaukee, Wisconsin

Librarian

Jack A. Clarke

Department of Library Science
University of Wisconsin —
Madison

APPOINTED OFFICIALS OF THE ACADEMY

Editor — Transactions
Walter F. Peterson
Department of History
Lawrence University, Appleton

Editor — Wisconsin Academy Review
Ruth L. Hine

Wisconsin Conservation Department,
Madison

Chairman — Junior Academy of Science
Jack R. Arndt
Extension Division
University of Wisconsin — Madison

THE ACADEMY COUNCIL

The Academy Council includes the above named officers and officials and
the following past presidents of the Academy.

A. W. Schorger
H. A. Schuette
L. E. Noland
Otto L. Kowalke
Katherine G. Nelson

Ralph W. Buckstaff
Joseph G. Baier
Stephen F. Darling
Robert J. Dicke
Henry Meyer
Merrit Y. Hughes

Carl Welty
J. Martin Klotsche
Aaron J. Ihde
Walter E. Scott
Harry Hayden Clark

Cover Design by Jeffrey Homar
Title Page Design by Gail Mitchem
School of Fine Arts
University of Wisconsin-Milwaukee

TRANSACTIONS OF THE
WISCONSIN ACADEMY
OF SCIENCES, ARTS
AND LETTERS

LVI — 1967-1968

The current TRANSACTIONS is listed as LVI to maintain continuity and as
1967”68 to conform with the year of publication.

Editor

WALTER F. PETERSON

TRimCTIOl OF THE

mmmu iCiiDEMY

Established 1870
Volume LVI

THE SEARCH FOR SECURITY 1

David J. Behling

THE GREEK REVIVAL IN RACINE 9

Mary Ellen Pagel

RATTLESNAKES IN EARLY WISCONSIN 29

A. W. Schorger

THE WILD HONEYBEE IN EARLY WISCONSIN 49

A. W. Schorger

WISCONSIN PINELAND AND LOGGING MANAGEMENT 65

George W. Sieber

MARY MORTIMER: CONTINUITY AND CHANGE

AT MILWAUKEE FEMALE COLLEGE 73

Walter F. Peterson

A PROPOSITIONAL INVENTORY OF EXECUTIVE-

LEGISLATIVE CONFLICT 81

A. Clarke Hagensick

THE HANDWRITING ON THE LAND 93

Robert A. McCabe

RADIOCARBON DATES OF WISCONSIN 99

Robert F. Black and Meyer Rubin

GEOMORPHOLOGY OF DEVILS LAKE AREA, WISCONSIN 117

Robert F. Black

EVIDENCE FOR FAULT ZONES IN THE BEDROCK

OF MILWAUKEE COUNTY 149

Carl A. R. Distelhorst and A. G, Milnes

THE DISTRIBUTION OF IRON IN LAKE SEDIMENTS 153

Jerome 0. Nriagu

EFFECT OF FLOODING, DRAINAGE AND pH ON TRANSFOR¬
MATIONS OF Mn AND Fe IN 19 WISCONSIN SOILS 165

E. H. Graven and 0. J. Attoe

THE FRAGIPAN IN SOILS OF NORTHEASTERN WISCONSIN 173

Gerald W. Olson and Francis D. Hole

A COMPARISON OF RED CLAY GLACIO-LACUSTRINE
SEDIMENTS IN NORTHERN AND

EASTERN WISCONSIN 185

Gary W. Petersen

LIGHT PENETRATION STUDIES IN THE MILWAUKEE

HARBOR AREA OF LAKE MICHIGAN 197

Carroll R. Norden

THE MOVEMENT, RATE OF EXPLOITATION AND HOMING
BEHAVIOR OF WALLEYES IN LAKE WINNEBAGO AND
CONNECTING WATERS, WISCONSIN, AS

DETERMINED BY TAGGING 207

Gordon R. Priegel

THE TAXONOMY AND ECOLOGY OF LEECHES (HIRUDINEA)

OF LAKE MENDOTA, WISCONSIN 225

J. A. Sapkarev

RABIES AND RABIES CONTROL IN WISCONSIN 255

D. O. Trainer

NOTES ON WISCONSIN PARASITIC FUNGI. XXXIII 263

H. C. Greene

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN

NO. 59. PLANTAGINACEAE— PLANTAIN FAMILY 281

Melvern F. Tessene

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iOth President of the

WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS

THE SEARCH FOR SECURITY

David J. Behling

What is security? Perhaps the greatest insight into security is in
the Latin definition of the word securus, meaning ‘'without care.’’
Viewed positively, security is a good and productive force — peace
of mind, freedom from anxiety, freedom from uncertainty, freedom
from fear. With a sense of security, we are able to concentrate on
the productive aspects of life and living. Viewed negatively, in¬
security is a bad and unproductive force characterized by doubt,
apprehension, worry, fear, anxiety and other destructive and de¬
bilitating feelings.

All people want security. But as society has advanced the nature
of man’s search for it has changed. Primitive man’s chief concern
was the elemental protection of his life from hunger, from weather,
from beasts and his other enemies. Advancing civilization brought
a lessening of some sources of insecurity, but an increase in others,
including pestilence, famine and despotic rulers. With the indus¬
trial revolution, man made further progress.

It’s a fact^ — perhaps not too well known, but still a fact — that
any great degree of economic security, or any great development
of the insurance business that does much to provide economic se¬
curity, is nearly impossible except at times and in places charac¬
terized by a considerable economic and industrial development, a
general respect for law and order, a basically sound currency, and
reasonable stability of government. Favorable concurrences of
these conditions have obviously not existed for too long, and the
Roman historian Livy, may not have been thinking of insurance
when he wrote some 2,000 years ago words that have been trans¬
lated to say, “Nothing stings more deeply than the loss of money —
and security.”

On the other hand, maybe Livy did have insurance in mind when
he expressed in those words man’s need for economic security, for
he may have known that almost 2,000 years before his time the
Code of Hammurabi had shown the essentials of insurance to be
known to Babylonian traders. Hammurabi had also provided that
if a man were robbed and the criminal not apprehended, the gov¬
ernment would “render back to him whatsoever of his that was
lost,” — a sort of very early Social Security Act.

1

2 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Also, Livy may have known that some 900 years before his time
the merchants of the island of Rhodes had added important refine¬
ments to marine insurance when they devised the Rhodian Sea
Law. Storms and pirates were taking their toll of trading vessels,
not to mention imagined losses to ship-gulping sea monsters or to
sailing off the edge of the world into the surrounding void. The
Rhodians designed a system whereby when a ship failed to return,
each merchant absorbed a portion of the loss rather than allowing
the unlucky individual owner to be ruined.

This early insurance, whatever economic security it created in
the specific situations it covered, did not perceptibly increase the
security of the great majority of people. Yet we owe a great debt
to the merchant chiefs of the Mediterranean, for they formalized
the voluntary mutual assistance and risk-sharing principles on
which all insurance is based.

The Greeks, whose reverence for human life exceeded that of any
people who preceded them, were the first to apply these principles
to men’s lives. Their burial societies, not only met the burial ex¬
penses of deceased members, but also provided for at least some
of the temporary financial needs of their widows and orphans.

The business-like Romans left evidences that they had developed
rather complex forms of commercial insurance, and that they gave
continuity to the concept of life insurance through their payments
to the survivors of soldiers. As a matter of fact, some 2,000 years
ago the Roman, Ulpianus, for that purpose provided a table of life
expectancy so accurate that only slight changes have been made
since then.

Although the Greeks and Romans did make great strides in the
discovery of insurance principles and in the wider application of
these principles to more people, they brought increased security to
only a very small fraction of the population. In the time of ad¬
versity, like the death of the family’s breadwinner, the great ma¬
jority still had to depend for their economic necessities on the un¬
certain good will and generosity of their relatives and friends, who
themselves might or might not have resources to share.

The snail’s pace development of man’s cooperative efforts at
achieving economic security halted altogether with the fall of Rome
to the barbarians and the advent of the Dark Ages. The feudalism
of those days has been described as a compulsory form of security :
in return for his loyalty and labor for his ruler a man hoped to
obtain protection and the necessities of life for himself and his
family. This was not an ennobling form of security — and it existed
only at the whim of the lord of the manor and only so long as the
latter remained as strong or stronger than his rivals. This rather
insecure form of security may still compare favorably with the

1967-68]

Behling—The Search for Security

3

present situation of a sig*nificant proportion of mankind living in
countries not yet showing any considerable economic and industrial
development, respect for law and order, soundness of currency, or
stability of government.

The Renaissance marked the rebirth of mutual assistance efforts
on the part of western man. The merchant and citizen guilds, later
the friendly societies of England, other groups in other places, all
used insurance concepts to protect the security of their members.

The great fire of London in 1666^ while destroying five-sixths of
the city, had two beneficial side effects upon man's security. One, it
destroyed that section where the plague that periodically swept the
city was concentrated, and which has been credited with controlling
the future outbreaks of the disease. Two, fire insurance sprang into
being from the city's embers, enabling men to protect their homes
from the financial consequences of the disaster of fire. This is an~
another example of the fact that out of mankind's greatest dis¬
asters have often arisen humanity's means of salvation.

It was then, too, that life insurance policies and annuities entered
the scene, usually being offered to the public by companies under¬
writing both fire and life risks. So pertinacious were the agents
who solicited for these companies that an outraged poet of the day
complained :

By fire and life insurers next
I^m intercepted, pestered, vexed
Almost beyond endurance.

And though the schemes appear unsound,

Their advocates are seldom found
Deficient in assurance.

Among the numerous complaints were some whose titles seem
absurd even to our speculative generation of the 1960's :

Assurance of Female Chastity
Assurance From Lying
Insurance from Death by Drinking Gin
Insurance Against Going to Hell

But out of this great misconception of the true purpose of insur¬
ance— and it wasn't humorous at that time- — came the clearing of
the way for legitimate life insurance underwriting.

It is quite true that often in those early days of insurance, fire
and particularly life policies were either woefully underpriced or
overpriced by reason of misconceptions as to the principles in¬
volved. Efforts to clear up these misconceptions led to actuarial
science, dealing with the mathematics of life contingencies — ^that is,
the probabilities of life and death which were long greatly mis¬
understood, As a matter of fact, they are still surprisingly myste-

4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

rious, despite the fact that all the really basic principles of actu¬
arial science had been developed and presented in text books by the
time I studied the mathematics of life insurance at the University
exactly forty years ago.

This reference to my early insurance student days reminds me of
what probably ought to have been the frightening story I read at
the time about a life insurance man who was such a complete in¬
surance man that he filled many notebooks with statistical observa¬
tions of phenomena concommitant with life and death, attempting
to analyze each and thus extend the frontiers of his understanding
and knowledge of insurance.

In today's terminology we would very likely say that this had a
psychosomatic effect on him, for at a really not advanced age he
discovered one day while analyzing his notebooks that he had been
sleeping longer each night than the night before — fifteen minutes
longer, to be precise. This continued. The time arrived when he
slept for 23 hours and 45 minutes. On awakening he hastily called
his wife, children and grandchildren about him, gave them such
advice as the wisdom of a lifetime suggested, and bade them an
affectionate farewell. At the end of fifteen waking minutes, he
promptly fell asleep again, slept for precisely 24 hours — and then
quietly expired.

To my youthful mind that was a highly admirable example of
complete absorption in one's chosen vocation. Now that I am older,
I confess that I feel happy to observe that my sleep habits show
no such disquieting regularity to which my wife will attest.

Actuarial science, as I have indicated, was a mature science
when I came to it, and 1 promptly became convinced that the life
insurance problems of the future could be solved by the experience
and wisdom of the past. These have supplied the basic and im¬
mutable principles ; but the actual developments and innovations in
my lifetime were to be so extraordinary that no one would then
have conceived them to be possible of accomplishment in a short
forty years.

No one forty years ago would have guessed to what extent new
policies could be developed to provide new comprehensive programs
of life insurance and annuity benefits, “new" even if the elements
of each program were as old as actuarial science. No one would
have foreseen the development of new uses for life insurance in
the business world, such as to protect businesses in the event of
death of key executives, or to provide necessary additional security
for loans, or to assure orderly continuance of partnership busi¬
nesses after the death of one partner, or to assure Uncle Sam his
estate tax, with a sufficient amount left for the deceased's family.

1967-68]

Behling — The Search for Security

5

During the past forty years there have been extraordinary
changes in the methods by which life insurance is presented to the
purchaser, enabling him to analyze his financial situation and buy
the particular policies that fit into a logical program of protection
for the insurance needs of his family. This valuable programming
approach, together with other improved procedures and strength¬
ened standards of competence, have happily changed the public
image of life insurance agents from that of rather ineffectual but
annoyingly high-pressure salesmen, failures in other lines of en¬
deavor (all too generally true in fact forty years ago) to now, in
many cases, trusted confidential advisors of quasi-professional or
professional stature, with their own designation — that of Char¬
tered Life Underwriter, which is equivalent to the CPA in
accounting.

The life insurance business is usually counted among the very
conservative institutions in our economic and social life. It must
be conservative, because it is a trust sort of operation in which,
above all else, policyowner’s reserves must be safeguarded and ade¬
quate funds maintained to assure claim payments to all those who
will suffer the losses insured against. While the insurance man’s
need to be conservative may somewhat too often condition him to
oppose desirable, even inevitable change, it is extremely difficult
in the modern business world for any insurance or other business
organization to maintain the status quo for any length of time. In
fact, an obsession with security through the maintenance of the
status quo is the enemy of long term growth and even of existence.
It must be replaced by an intense desire to respond to new situa¬
tions arising in a changing world. Insurance history provides many
happy examples of such response.

During the early days of life insurance in this country, that is in
the 1800’s, policies became null and void if the insured traveled
too far from home, into the then unhealthy or dangerous regions
of the southern and western states, or into less settled parts of the
world or if he engaged in a duel, or even if he left the earth in a
hot air balloon! In the days when horses and wagons were the
usual means of land transportation, railroad engineers, firemen and
conductors had to pay extra for life insurance protection, and
anybody with nerve enough to serve as brakeman on a freight
train just couldn’t get insurance at any price. Similarly, it was
years after Kitty Hawk before aviators and their passengers could
get life insurance covering them in flight.

In contrast, one week before Major Gordon Cooper blasted off
on his twenty-two orbit space flight in 1963, the Aetna Life Insur¬
ance Company issued a $100,000 life insurance policy to Cooper
and to each of the six other original astronauts. Their life insur-

6 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

ance protection for their families was good anywhere on earth or
in outerspace. Yes, even on the moon, if and when some of those
men do get there.

A great many other details of the life insurance business and its
history might interest you. I must, however, move into the final
phase of my discussion of the part that life insurance plays in
man’s search for security — in helping him achieve economic
security.

Does this economic security mean protection against change in
man’s economic condition, to enable him to maintain the status
quo? Not by any means. We must not forget the paradox that an
obsession with security through the maintenance of the status quo
is the enemy not only of all progress, but also ultimately of security
itself.

Even if it seems to raise another paradox, let me try to explain
how insurance, by protecting the status quo against certain risks,
can enable a prudent man to incur other risks in order to progress
far beyond the status quo. A few examples may indicate what I
mean.

Life insurance has provided many a man’s widow and orphans
with their main or only means of self-respecting subsistence. Often
it alone has enabled the members of the bereaved family to remain
in what may be called their own world, something near to the kind
of life they have been used to, with some of the comforts and
amenities of our civilization in addition to the necessities of life,
and with the right kind of opportunities for the bringing up and
the education of the children. That is one side of the life insurance
coin; the other is that the ownership of an adequate amount of
life insurance enables a prudent man to incur larger financial ob¬
ligations and to take greater financial risks for the furtherance of
his career and, if he is successful, for the ultimate benefit and
satisfaction of his family. He can do so in reliance on life insurance
to pick up his financial responsibilities to his family if death inter¬
rupts his attainment of his business or professional objectives.

Then, too, the tendency of recent decades toward early marriages
and having children young gains with life insurance a measure of
economic prudence. The parents of a young girl can even prepare
for this by buying insurance on her life with the provision that if
and when she marries she can transfer it on to the life of her
bridegroom.

Insurance provides a bulwark against these hazards to economic
security, bulwarks it would be unthinkable to be without, for eco¬
nomic misfortune rarely if ever affects only the few persons whom
it directly strikes. If there is no method of relieving the financial
consequences of individual catastrophe, society as a whole suffers

1967-68] Behling — The Search for Security 7

both from the nonpayment of the liabilities of the insolvent and
from the interruption of the productive activities of all concerned.
And the other side of the coin is that in the absence of the security
that insurance can promise, man would not dare to invest either
his money or his efforts in the business and personal activities and
operations which make the modern world what it is and lead to the
great developments which the passing decades observe.

I should like to conclude with a short commentary on the life
insurance business’ own quest for security. How can the insurance
business insure itself? I do not at this point have in mind the tech¬
nical, but nonetheless important, matter of protection through re¬
insurance against too large claims or catastrophic aggregations of
claims. What I am thinking about is the long term security of the
insurance business. This will come from its adaptability to change,
from the new protections it provides against the financial conse¬
quences of the new hazards and perils which our country’s develop¬
ing economic and personal life incurs; from the extent to which
the security provided by insurance to business and industry, and
individuals, makes it prudent for business and industry, and indi¬
viduals to exercise the boldness and adventurousness which a good
pace of economic and personal progress will always require.

For all business — in fact for man himself — there can be no se¬
curity that is not grounded on courageous and wise adaptation to
the new situations that our changing world will bring. Man’s
search for security will continue to be a major personal and gov¬
ernmental preoccupation, but its pursuit ought not to obscure all
other values, and especially those on which security itself depends.

The words of Somerset Maugham as he watched the fall of
France in the first year of World War II are arresting —

‘‘Those who value security above freedom will lose their freedom,
and having lost their freedom, they will lose their security also.’’

THE GREEK REVIVAL IN RACINE

Mary Ellen PageV^

The Greek Revival style in architecture made its American debut
in 1798 v^ith Benjamin Henry Latrobe’s design for the Bank of
Pennsylvania at Philadelphia. Latrobe’s use of forms and details
derived from ancient Greek architecture was not without European
precedent; indeed, in his native England buildings in the Grecian
mode had been constructed as early as the 1750's, But in no Euro¬
pean nation was the style to prove more popular and enduring than
in the young United States, Its aesthetic merits, its ready adapta¬
bility to various functions and building materials, its evocation
of the ideals of Greek democracy — all these endeared it to Ameri¬
cans. And so it happened that within a decade of the completion
of Latrobe’s Philadelphia bank, the Greek Revival gained wide cur¬
rency in eastern architectural circles and, in most sections of the
new nation during the first half of the 19th century, was a pre¬
dominant style in which both professional architects and amateur
designers clothed public and residential buildings and, to a lesser
extent, commercial and religious structures as well. As Hugh Mor¬
rison has observed : “From 1820 on, the Greek temple became the
highest architectural ideal for a generation of Americans.’’^

A sketch of Racine, Wisconsin made in 1841 (just seven years
after pioneer settler Gilbert Knapp had erected Racine’s first build¬
ing) reveals that the tastes of the city’s early residents were not
at odds with this pattern. (Fig. 1). Their fondness for classical
architectural forms is evident in the modestly Grecian houses at
left and right and in the more monumentally treated county court¬
house near the center of the drawing.

* Department of Art and Office of Planning- and Construction, University of
Wisconsin Center System.

^ Early American Architecture from the First Colonial Settlement to the National
Period (New York, 1952), 575.

9

10 Wisconsin Academy of Sciences, Arts and Letters [VoL 56

Figure 1. Corner of Seventh and Main Streets, Eacine, in 1841 (drawing re¬
produced courtesy the Racine County Historical Museum).

Work on Racine’s first courthouse (Figs. 1, 2) began in 1839
under the supervision of William H. Waterman and Roswell Mor¬
ris, local contractors who were responsible, it appears, not only
for construction but also for design.^ Completed in 1840. their
courthouse was a chaste white frame building with a symmetrical,
temple-like fagade of four Doric columns. A domed octagonal cupola
crowned the building’s low-pitched gabled roof-line. In plan and in
front elevation the now-destroyed courthouse conformed to a type
held in high regard across America during the three decades pie-
ceding the Civil War. Among its relatives number such eastern
courthouses as those at Hillsboro, North Carolina (1845) and
Waynesburg, Pennsylvania (1850)^ and such midwestern examples
as the Third LaSalle County Court House at Ottawa, Illinois (1842 ;
razed), ^ the Milwaukee County Court House (1836; razed),® and

3 Fanny S. Stone (ed. ), Racine, Belle City of the Lakes and Racine County, Wis¬
consin (2 vols. : Chicago, 1916), I, 96. For the courthouse see also: The History of
Racine and Kenosha Counties, Wisconsin (Chicago, 1879), 367 ; Eugene W. Leach,
History of the First Methodist Episcopal Church, Racine, Wisconsin, with a Prelim¬
inary Chapter Devoted to the 'City of Racine 1836 to 1912 (Racine, 1912), 19, 31;
Rexford Newcomb, Architecture of the Old Northxvest Tex'Htory : A Stxidy of Early
Architecture in Ohio, Indiana, Illinois, Michigan, Wisconsin and Part of Minnesota
(Chicago, 1950), 133.

Illustrated in Talbot Hamlin, Creek Revival Architecture in America: Being an
Account of Important Trends in American Architecture and American Life prior to
the War Between the States (London, New York, and Toronto, 1944), Plates L and
LXXVII, respectively.

^Newcomb, 101 and Plate XLI.

® Harry H. Anderson, “The First County Court House,” Historical Messenger of the
Milwaukee County Historical Society, XXI (December, 1965), 100-109.

Figure 2. The Eacine public square c. 1860. First Baptist Church is at left,
the first Racine County Courthouse at right (drawing reproduced courtesy the
Racine County Historical Museum).

the well-known Iowa County Court House at Hodgeville (1859),
Wisconsin’s last important remaining example of the type.®

Enjoying concurrent and equally widespread popularity were
residential designs of the types represented at far left and far right
in the sketch of 1841. Rectangular, two-story structures with gently
sloping gabled roofs, these unpretentious homes lacked the colon¬
naded entrance porticoes of more elaborate Greek Revival buildings
but retained the compact silhouette, geometrical clarity, and, in
greatly simplified form, the classical details of the style. Familiar
Wisconsin examples include the York house near Zenda and the
diminutive Christophel house in Milwaukee — both mentioned by
Richard W. E. Perrin,'^ the brick residence at 922 North Cass Street
in Milwaukee, .and the so-called Old English House on Third Avenue
in Kenosha. The two early specimens in Racine, which, according
to local historian Eugene W. Leach, may have belonged to H, J.

Robert M. Neal, “The History of the Court House,” Centennial Story: The Iowa
County Courthouse 1859—1959 ( Dodge ville, Wisconsin, 1959) and Richard W. E. Perrin,
Historic Wisconsin Buildings: A Survey of Pioneer Architecture (Milwaukee Public
Museum Publications in History No, 4; Milwaukee, 1962), 30, 33.

Historic Wisconsin Buildings, 30, 35.

12 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 3. 1108 Douglas Avenue before recent renovation (photograph by Todd
Dahlen and Peter Vallone).

Smith (house at left) and to Paul Kingston (right) have not sur¬
vived, but similar residences still stand in the city.

At 1108 Douglas Avenue (Fig. 3), for example, is a home closely
resembling the lost Smith house. Designed and built in 1855 by

® Leach, G.

1967-68]

Pagel — The Greek Revival in Racine

13

Charles Fountain, it served briefly (1862-63) as the first Domini¬
can convent in Racine.^ Passing again to private ownership in 1863,
it has remained in use as a residence since then and had undergone
only minor remodelling and modernization on the interior until
1967, when the exterior was renovatedd^ Here, just as in the Smith
dwelling, one finds a two-story plan, a facade with three regularly
spaced windows on the upper floor, and, below, the entrance placed
to the left. The recessed entry way, with pilasters and narrow side¬
lights flanking the door, small transoms and an entablature above
— in the Fountain house the lone decorative element in an other¬
wise plain fagade — occurred in Neo-classical structures both hum¬
ble and sumptuous, both private and public. Variations on the
common theme appeared in Racine’s first prominent hotel (the
Racine House, 1837),^^ the city’s first brick residence (the Ives
house, c. 1840), the mid-19th century Fratt mansion, and the
Cooley-Kuehneman house of c. 1851-54 (Fig. 10) — ^^among numer¬
ous examples. This distinctive doorway treatment seems to have
been popularized by Asher Benjamin, whose widely circulated
architectural handbooks served as design sources and practical
guides for 19th century American builders.

Returning to the drawing of 1841, one finds that the Kingston
home at far right, while similar to those just considered, presents
a more ornamental fagade, with simplified Doric pilasters at the
corners and a fully-defined pediment above the second story win¬
dows. Like structures in present-day Racine include the attractive
gray and white frame house at 1201 College Avenue (Fig, 4).
Erected c. 1852-60, the home has been enlarged by additional con¬
struction at south and east but retains a sturdy, straightforward
ante-bellum flavor.^^

A late 19th century photograph of the one-time residence (now
the local American Red Cross headquarters) at 745 Wisconsin
Avenue suggests that it, also, was originally of this pedimented

® Sister Mary Hortense Kohler, The Life and Work of Mother Benedicta Bauer
(Milwaukee, 1937), 204—205,

Mary Ellen Pag’el (ed. ), Pagans and Goths in Nineteenth Ceutury Racine: Archi¬
tecture of the Classical and Gothic Revivals (mimeographed; Racine, 1964), 7—8 and
an unpublished report in the author’s collection written in 1964 by Carol Haberman
and Karen Nielsen of the University of Wisconsin— Racine Center.

11 Leach, 18.

^Ibid., 27,

10 The Historic American Building's Survey, Wisconsin Architecture: A Catalog of
Buildings Represented in the Library of 'Congress, with Illustrations from Measured
Drawings (Washington, D. C., 1965), 68.

11 Newcomb, 79 and Hamlin, Plate XCIV.

n* Pag'd, 8 and an unpublished report in the author’s collection prepared in 1964 by
Judy Sorensen and Barbara Monefeldt of the UW-Racine Center.

14 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 4. 1201 College Avenue (photograph by Todd Dahlen and Peter Val-
lone) .

type (Fig. 5). The dainty Victorian veranda which decorated the
home when it was photographed c. 1872-92 was, in all probability,
a post-Civil War addition, but the wing at right with the stout
Doric columns may have been part of the original design. Recent
remodelling has removed both the veranda and the sturdy colonnade

1967-68] Pagel — The Greek Revival in Racine 15

Figure 5. 745 Wisconsin Avenue in 1872-92 (photograph from the Racine
County Historical Museum).

and given the building its present facade (Fig, 6), Although there
is evidence that a structure, possibly a residence, stood on this site
as early as 1851, the precise date of the existing building, like its
original appearance and the name of its designer, is still to be
established.^®

Uncertain, too, are the identities of the architects of Racine's
best-known Greek Revival houses — the charming residence at 1247
South Main Street (Fig. 7) and its celebrated neighbor at 1135
South Main (Figs. 9, 10), both of which have been recorded by
the Historic American Buildings Survey of the U. S. Department
of the Interior.^' Earlier of the pair, 1247 South Main (first the
Hunt house; later called Westbourne; now the Harold C. Jensen
residence) dates from c. 1842-48 and is among the city's oldest

i'® An unpublished report in the author’s collection written in 1964 by Ruth Jensen
and Elizabeth Maroda, then at the UW-Racine Center, states that the property in
question was sold in 1851 by Alexander Bishop to Jacop Soatwall for $2,000, a sum
considerably in excess of amounts then asked for unimproved lots in the neighborhood.
The property increased in value in subsequent sales, having been reacquired by Bishop
in 1852 for $2,500, sold by Bishop to Walter Cooley for $3,714 in 1855, and by Cooley
to Hamilton Utley for $6,500 in 1870. See also Pagel, 6-7.

u Wisconsin Architecture, 68, 69.

16

Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 6. 745 Wisconsin Avenue in 1964 (photograph by Todd Dahlen and
Peter Vallone).

Figure 7. 1247 South Main Street (photograph by Todd Dahlen and Peter
Vallone) .

1967-68]

Pagel — The Greek Revival in Racine

17

extant homes. Tradition holds that it was built by William Hunt
as a gift for his wife and that the designer-builder was a local
carpenter.

In this connection, it is interesting to take note of a house men¬
tioned and illustrated some five decades ago by Racine historian
E. W, Leach (Fig. 8). In his day the house was standing at 416
Lake Avenue. He noted that it had been built ''about 1840” by a
carpenter named Chadwick and that it was still called the Chadwick
housed^ Writing a few years later, Mrs. David H. Flett repeated
this information and added that the home originally stood on Main
Street.^^ Early city directories reveal that one Reuben Chadwick,
cabinet maker, was residing at 141 Main Street by 1850,-*’ but
available evidence allows no more than speculation as to his identity
with the carpenter named by Leach and Mrs. Flett. The question
of the attribution and dating of the now-destroyed Chadwick house
assumes particular interest for students of the Hunt-Jensen home

18 Leach, 29.

^ “Notable Pioneer Homes” in Stone, I, 402.

The Racine Register, Business Directory, and Advertiser (Racine, 1850), 25. The
first Chadwick specifically described as a carpenter in city directories was Ellis Chad¬
wick, living with William Chadwick, woodturner, on Main Street, in 1868 ; see Richard
Edwards, Edwards’ Annual Directory of . . . the City of Racine for lS{>8-69 (Racine,
1868), 61, 62.

Figure 8. The Chadwick house c. 1912 (photograph from the Racine County
Historical Museum).

18 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

because of the obvious and striking similarities between the two
structures. So close are they in proportions and details that one is
tempted to suppose that the same hand drew both plans or that, at
the very least, a single source — to be discovered, perhaps, in a 19th
century builders’ guide — inspired them.

The Hunt-Jensen house has been moved several times during its
long history, but despite the transfers, it survives in good condition
and preserves a substantial portion of its original design. It re¬
mains an excellent example of the Greek Revival temple-house, with
the characteristic front portico of columns, here of the decorative
Ionic order. Typical, too, are the near symmetry of plan and facade,
the wood construction and siding, the uniformly white, smoothly
surfaced exterior, and the air of tranquility, dignity, and comfort:^^
The house presents many noteworthy details — among them the
pedimental ornament on the facade. Architectural historian Talbot
Hamlin has stated that pierced grilles of this type, executed in wood
(as in this case) or in cast iron, were one of several distinctly
American contributions to the Greek Revival decorative vocabulary
and were “common in frieze and attic windows all over the coun¬
try.”®^ The Hunt-Jensen grille, the sole surviving instance in Ra¬
cine, is not unlike the window grilles in Johnathan Goldsmith’s cot¬
tage at Painesville, Ohio (1841), which, Hamlin found, had been
borrowed from a plate in Minard Lafever’s The Modern Builders'
GuideP As we know, Lafever’s books were quite as popular among
19th century craftsmen as those of Asher Benjamin,

Called “perhaps the best remaining example of the Greek Revival
in Wisconsin” by the writers of the Historic American Buildings
Survey,-^ the William F. Kuehneman house (Figs. 9, 10) was built
for Eli R. Cooley, hardware merchant and third Mayor of Racine.
It can be dated between 1851, when Cooley acquired the property,
and 1854 when he sold it to Elias Jennings at a marked increase
in price.'^® The simple, beautifully proportioned home consists of a
two-story central block with a projecting porch of four slender
Doric columns and symmetrically disposed one-and-one-half-story

21 For the Hunt-Jensen house see also : Alexander C, Guth, “Historic American
Building’S Survey,’’ Wisconsin Magazine of History, XXII (September, 1938), 31-32;
Newcomb, 130 ; Writers’ Program of the Work Projects Administration in the State of
Wisconsin, Wisconsin: A Guide to the Badger State (revised ed. ; New York, 1954),
280. Pagel, 5-6; “Racine,’’ Wisconsin Architect, XXXIV (April, 1966), 16; and an
unpublished report in the author’s collection prepared in 1964 by Frank Chud and
Thomas Fuhrer, UW-Racine Center.

23 Hamlin, 354.

23/?>id. and Plate XCIII.

31 Wisconsin Architecture, 69.

=®Lucy Colbert, “Century-old Home Cited for Its Beauty, History,’’ Racine Journal-
Times Sunday Bulletin, October 14, 1956, sec. 2, 1 and an unpublished report in the
author’s collection written in 1964 by Lynn Meier and Marilyn Francis, UW-Racine
Center.

1967-68]

Pagel — The Greek Revival in Racine

19

Figure 9. 1135 South Main Street (photograph by Todd Dahlen and Peter Val-
lone) .

wings. Both exterior and interior have been carefully restored and
maintained by the present owner.'^^'

The gifted designer has not been identified with certainty, but
critics have suggested that Lucas Bradley (1809-89), Racine's first
architect, drew the plans. Born and educated in Cayuga County,
New York, Bradley worked as carpenter-architect for a brief period
in St. Louis, visited Racine in 1843, and settled there permanently
the following year. His two documented buildings in the Greek
Revival style— Second Presbyterian Church in St. Louis (1839-40;
razed and First Presbyterian Church of 1851-52 in Racine (Fig.
13) give evidence that he was a master of the first rank and, fur-

Much has been written about the Cooley— Kuehneman house. Additional sources
include: Newcomb, 130; Perrin, Historic Wisconsin Buildings^ 28-29; Pagel, 3-4; TVis-
consin Architect, XXXIV, 15. Perrin also discusses the home in his Historic Wisconsin
Architecture (Milwaukee, 1960), 11 and his “Greek Revival Moves Westward: The
Classical Mold, in Wisconsin,” Wisconsin Magazine of History, XLV (Spring, 1962), 201.

2T John A. Bryan, Missouri’s Contribution to American Architecture (St. Louis, 1928),
11, 27 ; by the same author, “Outstanding Architects in St. Louis between 1804 and
1904,” Missotiri Historical Review, XXVIII (1933-34), 85 ; Hamlin, 252 ; Newcomb, 135.

20 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 10. 1135 South Main Street, detail, entrance (photograph by Todd
Dahlen and Peter Vallone.

1967-68]

Pagel — The Greek Revival in Racine

21

ther, offer stylistic parallels with the Cooley-Kuehneman houser*^
This home, in turn, resembles a second residence in the area, as
Perrin has pointed out:

“A few miles northwest of Racine on the Nicholson Road, in the Town
of Caledonia, Racine County, is another Temple house which might be
called a country cousin of the Kuehneman house. The central Doric tetra-
prostyle portion resembles the Kuehneman house so very much that it
could be concluded that either the same architect or the same architec¬
tural handbook played a part in its design. This house is believed to have
been built by John Collins of New York State in about 1853.’’®®

Less appealing to church architects than was the contemporary
Gothic Revival style, the Greek Revival was, nonetheless, employed
for religious buildings. And Racine’s Neo-classical churches, like
its Grecian homes, range from the modest to the majestic. Repre¬
senting the former extreme is the tiny building at 806 Superior
Street (Fig. 11), Erected for the First Scandanavian Baptist
Church c. 1859, the structure is remotely Grecian in the pediments
and pilasters of its facade and in its boxy, compact shape. The now-
anonymous designer apparently felt constrained to modify the
pagan implications of the Greek Revival style and punctuated the
side elevations with Gothic lancet windows. One finds this curious
combination of classical and Gothic motifs in several other early
Wisconsin churches, including St. Peter’s Church (1839), formerly
in Milwaukee and now on the grounds of St. Francis Seminary, St.
Augustine Church at New Diggings (1844), and the Moravian
Church at Green Bay (1851).'^^ Even closer in form and spirit to
the little Racine church, though lacking Gothic aisle v^indows, are
the Painesville Chapel in Franklin (1832)'^^ and the Congrega¬
tional Meetinghouse at Cato (1857).'^- Racine’s Scandinavian
Baptists occupied their church until 1903. In 1887 they had built a

For Lucas Bradley see also: Racine city directories 1850-88; The History of
Racine and Kenosha Counties, Wisconsin, 375, 568 ; obituaries in the Racine Daily
Times, January 10, 1889 and in the Racine Journal, January 16, 1889 ; Stone, I, 401 ;
Alexander C. Guth, “Early Day Architects in Wisconsin,” Wisconshi Magazine of
History, XVIII (December, 1934 ), 143; Henry Steketee, “Architect Given Praise for
Planning Racine Church,” Racine Journal-Times Sunday Bulletin, February 19, 1939,
5 ; the Rev. Sydney H. Croft, “A Hundred Years of Racine College and DeKoven
Foundation,” Wisconsin Magazine of History , KJCKV (Summer, 1952), 251, 253; Henry
F. Withey and Elsie R. Withey, Dictionary of American Architects (Deceased) (Los
Angeles, 1956), 73; Dictionary of Wisconsin Biography (Madison, Wisconsin, 1960),
45—46 ; George Miller, “Cite Architecture of Six County Buildings,” Racine Journal-
Times Sunday Bulletin, October 23, 1960, sec, 1, 3 ; Pagel, 2, 12 ; sources cited for
First Presbyterian Church (below). There also exist a number of unpublished papers
dealing with Bradley and his work in the collections of Beloit College, the Racine
County Historical Museum, the Racine Public Library, and the author.

^Historic Wisconsin Buildings, 29.

These three churches are discussed and illustrated in Wisconsin Architecture, 71,
62, and 46, respectively.

^^lUd., 44.

33 Perrin, Historic Wisconsin Buildings, 34, 40.

22 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 11. 806 Superior Street ( photog-raph from the Racine County Histori¬
cal Museum).

parsonage nearby, and during this century the two small buildings
were joined and put to residential use.‘^'^

More architecturally pretentious was the city’s First Methodist
Church of 1844-45 (Fig. 12), Pilasters defined and divided its
fagade and acted as visual supports for the heavy pediment above.
The rectangular forms of the centralized entry echoed the building’s
shape, its geometrical ornamentation, and its squat, squared
belfry.^"^^ In several of these features First Methodist calls to mind
the church at Streetsboro, Ohio, illustrated by Hamlin, and,
among Wisconsin specimens, the First Baptist Church at Merton
(1845)'"^® and the Muskego Meetinghouse (formerly the Free-Will
Baptist Church) at Prospect (1859). Mid-19th century Racine
boasted at least two more houses of worship of this type— First
Baptist Church completed in 1848 (Fig. 2)^® and the Universalist

Pagel, 9 and an unpublished report in the author’s collection written in 1964 by
William Adams, Don LaFave, and Dennis Zwaga, UW-Racine Center.

The history of First Methodist is discussed in The History of Racine and Kenosha
Counties, Wisconsin, 382-383 and in Leach, 82-83,

35 Plate DXXXII.

Wisconsin Architecture, 55.

Perrin, Historic Wisconsin Buildings, 34, 40.

38 The History of Racine and Kenosha Counties, Wisconsin, 384-386 ; Leach, 8 ;
Stone, I, 359-360.

1967-68] Pagel — -The Greek Revival in Racine 23

Figure 12. First Methodist Church (photograph from the Racine County His¬
torical Museum).

Church of 1851-52.^^ Regrettably, neither these buildings nor First
Methodist come down to us.

Greek Revival church design in Racine culminated in the greatly-
admired First Presbyterian Church at Seventh Street and College
Avenue (Fig. 13), praised by Rexford Newcomb for its “sincere
and highly refined design”^® and described by Perrin as “perhaps
the finest example of brick church architecture in the Greek Re¬
vival Style. First Presbyterian’s members had built their first
church in 1842 and, to accommodate a growing congregation, en¬
larged this simple wood-framed structure the following year. Five
years later they passed a resolution calling for a new church and
appointed church member and architect Lucas Bradley to the build¬
ing committee. Fund-raising continued through 1850, with the lot
purchased in December of that year. In 1851 church historian
Stephen Peet wrote : “Measures have been taken and a subscription

^ The History of Racine and Kenosha Counties, Wisconsin, 387-388, 391 ; Leach,
20, 30 ; Stone, I, 376.

^0 Newcomb, 135.

Historic Wisconsin Buildings, 55-56.

24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 13. First Presbyterian Church (photograph by Todd Dahlen and Peter
Vallone) .

raised, amounting to near $8000, towards a more commodious
house, to be erected the coming season.’'^'^ By March Bradley had
been awarded the contract, and on May 6 the cornerstone was laid.

^ History of the Preshyterian and Congregatioival Ministers in Wisconsin (Milwau¬
kee, 1851), 152.

1967-68]

Pagel — The Greek Revival in Racine

25

Specifications called for a building of the Grecian-Doric style, and
this Bradley provided in a design stongly reminiscent of his earlier
Presbyterian church in St. Louis. Once again Doric columns dom¬
inated the monumental facade and were surmounted by an entabla¬
ture of the same order. Crowning both compositions were spires
with engaged columns of the Ionic order — spires less indebted to
Greek precedent, of course, than to those of British architects Sir
Christopher Wren (1632-1723) and James Gibbs (1682-1754).^*^
That Bradley's design is both derivative and eclectic detracts in no
way from its success.^^

First Presbyterian was dedicated on June 10, 1852, and within a
few months, work on the closely related First Congregational
Church (now St. George Serbian Orthodox Church) at 826 State
Street was underway (Figs. 14, 15). Two years earlier a dissident
portion of First Presbyterian’s membership had broken away to
found First Congregational, and in February, 1851 they had dedi¬
cated their original church — ^^according to Peet, an example of “the
Swiss Cottage and Gothic Style . . . with 5 pointed arch windows
on each side and one in front between two large porches, which
terminate in 4 pointed buttresses. This church perished in a fire
later the same year. Completion of the Congregationalists’ second
church was signalled by dedication services on November 17, 1854.
The building clearly owed a great deal to Bradley’s design for
the pilasters adorning the fagade and dividing the side elevations
the pilasters adorning the facade and dividing the side elevations
into bays, the Ionic order decorating the octagonal spire — all looked
back to the older church.

Good fortune has marked First Presbyterian’s subsequent his¬
tory : the church has seen few major alterations, and, by and large,
modifications have been carried out in the spirit of the original
fabric. First Congregational has been less fortunate: lightning
destroyed the spire in 1912; fire forced abandonment and sale of
the building in 1948; and for the next nine years it served as a

For the combination of Wren-Gibbs and Greek Revival elements in American
church design see Hamlin, 344-345.

Like the Cooley-Kuehneman house and architect Bradley, First Presbyterian
Church has received considerable attention from writers. Sources, in addition to those
already cited, include: The History of Racine and Kenosha bounties, Wisco7isin, 384;
Stone, I, 375-376; “Church to Observe 75th Year; Presbyterians Here to Hold Anni¬
versary Week,” Racine Journal-News, October 12, 1927, 1, 11 ; Guth, Wisconsin Maga¬
zine of History, XXII, 21-22. Henry Steketee, “Church to Mark Its 100th Year,” Racine
Journal-Times Sunday Bulletin, January 29, 1939, 1, 5 ; “To Dedicate Presbyterian
Parish Hall,” Racine Journal— Times , June 1, 1942, 9 ; Perrin, Historic Wisconsin
Architecture, 11 ; Perrin, Wisconsin Magazine of History, XLV, 201 ; Pagel, 1-2 ; Wis¬
consin Architecture, 67 ; Wisconsin Architect, XXXIV, 17.

^Peet, 153.

26 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Figure 14. First Congregational Church (now St. George Serbian Orthodox
Church) before 1912 (photograph from the Racine County Historical Museum).

1967-68]

Pagel — The Greek Revival in Racine

27

Figure 15. St, George Serbian Orthodox Church in 1964 (photograph by Todd
Dahlen and Peter Vallone).

dance hall. In 1957 it was acquired by the present owners and has
since undergone extensive remodelling and restoration.^®

The Greek Revival chapter in Racine’s architectural history came
to a close within a decade after First Presbyterian and First Con¬
gregational were dedicated. Here, as elsewhere in the United States
during the 1860’s, long-prevailing classical tastes surrendered to
the rising picturesque current — expressed in the rich, complex,
decorative forms of the Italian Villa, Gothic Revival, and Second
Empire styles.

A sequel to the ante-bellum Greek Revival story was written by
the Academic Reaction in architecture of the late 19th and 20th
centuries, when Grecian forms and details once again found favor
among American designers and their patrons. In Racine this re¬
surgent classicism was heralded by the home at 820 Lake Avenue
(Fig. 16), designed c. 1885-87 by James Gilbert Chandler of Ra¬
cine for the McClurg family (and, since 1938, home of the local

4G For First Cong-regational-St. George Serbian Orthodox see also : The History of
Racine and Kenosha Counties, Wisconsin, 394-396 ; First Congregational Church, Ra¬
cine, Wisconsin, 1851-1911, Celehratmg Sixty Years of Church Life (Racine, 1911),
1-6 : Stone, I, 365-366 ; Lucy Colbert, “Historic Landmark Becomes New St. George
Serbian Church,” Racine Journal-Times Sunday Bulletin, October 5, 1958, sec. 2, 1,
18; Pagel, 2-3; and an unpublished report in the author’s collection prepared in 196 1
by Peter Charnon and James Gilmore, UW-Racine Center.

28 Wisconsin Academy of Sciences, Arts and Letters [VoL 56

Figure 16. 820 Lake Avenue (photograph by the author).

chapter of the Veterans of Foreign Wars).^" Its decorative details
remind one of Minard Lafever’s conceptions,^® but its grand scale
reflects the tastes of this new classical era. Popularized by the
structures at the World’s Columbian Exposition of 1893 in Chicago,
the Academic Reaction enjoyed a long lifespan in this country,
flourishing for some forty years. Typically, Racine’s last prominent
buildings in the classical vein — City Hall and the Main Post Office
— were erected in 1930-31.

In chronology, in many aspects of design and technology, in the
amateur-craftsman status of most of its designers, the Greek Re¬
vival in pre-Civil War Racine had also conformed to typical mid-
western patterns. At the same time, Racine’s case takes on more
than ordinary interest, for alongside the city’s characteristic Neo¬
classical structures had been built a number of the outstanding
Greek Revival buildings in the Old Northwest. Fortunately, several
important examples have survived the years in estimable states of
preservation: the Hunt- Jensen house, the Cooley-Kuehneman
house, and First Presbyterian Church are cases in point and rank
among the great treasures of Wisconsin’s architectural past.

stone, I, 391-392; “V. F. W. Opens Veterans Club for 25th Anniversary,” Racine
Journal-T imes Sunday Bulletin, October 18, 1950, 4 ; and unpublished papers in the
authoT-’s collection written by Marilyn Francis in 1964 and by Daniel J. iMoriarity in
1966.

48 Compare Hamlin, Plates XCII-XCIV.

RATTLESNAKES IN EARLY WISCONSIN

A. W. Schorger

The first mention of rattlesnakes (serpens sonnettes) in Wis^
consin was by Hennepin (1903:222) during his voyage on the
upper Mississippi in 1680. Le Sueur (1902:184) in 1700 reported
it was dangerous to enter the caverns near Lake Pepin because of
rattlesnakes. He saw some which were six feet long, but generally
they did not exceed four feet.* According to Owen (1852:57) they
inhabited the bluffs below Lake Pepin.

While descending the lower Wisconsin River in 1814, Anderson
(1882:192) allowed his men to stop at sand banks to collect turtle
eggs and kill rattlesnakes. These he thought beautiful with their
bright golden color crossed with black markings. In descending the
same stream, Marryat (1839:105) considered it dangerous to wan¬
der far from the bank because of the rattlesnakes. He believed that
there was probably no place in America where the two species of
rattlesnakes were larger and more numerous than in Wisconsin.
Brunson (1872, 11:172) in 1843 made an overland trip from Prairie
du Chien to La Pointe, his route running to Cashton, Tomah, Black
River Falls, and Chippewa Falls (McManus, 1919). Before reach¬
ing the Black River his party saw both species of rattlesnakes, and
between the Black and Chippewa Rivers, a few ''massasaugers'’
only. They saw none beyond the Chippewa.

Species

Wisconsin has only two species of rattlesnakes, the timber rattle¬
snake (Crotalus horridus horridus) and the massasauga (Sistrurus
catenatus catenatus) , The timber rattlesnake, also known as the
banded, yellow, mountain, and rock rattlesnake, is rarely found
far from rock outcrops, and in Wisconsin rock rather than timber
would be a more appropriate name. Although Pope (1930:282)
reports western diamondbacks (Crotalus atrox atrox) in Vernon
County in 1928, these were probably timber rattlesnakes with aber¬
rant markings, or the progeny of an escape.

* Evidently the lengths are estimates. The French foot was 12.789 inches.

29

30 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

The massasauga* was also known as the prairie, and spotted
rattlesnake. Its habitat is marsh, low prairie, and the low banks
of streams. Bunnell (1897 :323) records that in the region of the
upper Mississippi the massasauga was quite local in distribution,
being found in the swampy meadows along creeks ; it occupied the
bottoms of the Mississippi River only above flood level. Less
resistant to the ecological changes produced by man than the tim¬
ber rattlesnake, it is now the rarer of the two species. The Sistrurus
catenatus kirtlandi Holb., a dark form said to have occurred in
Walworth County (Higley, 1889:161), is a synonym.

Size

The maximum length of the timber rattlesnake is six feet two
inches (Klauber, 1956 :149) . George Knudsen has informed me that
he captured a specimen near Gotham in the spring of 1965 which
was four feet five inches. Breckenridge (1944:159) mentions a
“very large rattler,” taken in southeastern Pierce County in 1929,
which was four and one-half feet long, with fifteen rattles and a
button. A supposed diamondback, killed near Viroqua in 1928, was
four feet, ten and one-half inches (Pope, 1930:282). The largest
rattler ever captured by Elmer Keitel in Sauk County was close to
five feet. Even a snake four feet long is considered large (Mac-
Quarrie, 1941:83). The longest rattler found by Messeling (1953:
23) was four feet three inches, and the greatest number of rattles
23. A rattler five feet long with 26 rattles was once reported from
Alma (Alma, 1878.2). The number of rattles is indicative only of
age. A new rattle is grown each time that the skin is shed, which
may be two or three times a year.

The massasauga is much smaller, the maximum length being
37.5 inches (Klauber, 1956:144). The usual length is about 24
inches. A female captured near Nelson, Buffalo County, was 23
inches (Breckenridge, 1944:152). Two specimens from Portage
were 22 and 26.5 inches (Pope, 1926).

Abundance

The data available give only a faint idea of the abundance of
rattlesnakes in the last century. At 'Dodgeville, 48 timber rattle¬
snakes, all but one being young, were once found under a large
rock and killed (Dodgeville, 1878). Two parties killed 42 at Devil’s
Lake (Reedsburg, 1872). On the ridge near Ash Creek, town of
Orion, Crawford County, 38 were killed at a den (Richland, 1869).

* Derived from a branch of the Chippewa living- on a stream of this name on the
north shore of Lake Huron. There are many variants in the spelling-. According- to
P. W. Hodge (Handbook of Indians north of Mexico) the proper spelling is missas-
sauga.

1967-68] Schorger — Rattlesnakes in Early Wisconsin

31

Three men killed 66 rattlesnakes in a meadow in the town of Har¬
mony, Vernon County (Viroqua, 1882). During a rattlesnake and
spermophile hunt at Gilmanton, 99 snakes were killed (Alma,
1962). The Cooke family, which settled near Gilmanton in 1856,
kept a careful record of the number: 150 rattlesnakes killed during
the first year (Cooke, 1940:286). Messeling (1953:23) stated that
he collects annually for the bounty about 1000 rattlesnakes, count¬
ing old, young, and unborn.

Massasaugas, in 1835, were abundant in the marshes which then
existed on the site of the city of Milwaukee. Of that time Olin
(1930 :214) wrote : “The first day we mowed we killed any quantity
of rattlesnakes. I will not say a thousand for fear some one will
think it a snake story.” In 1845, they “swarmed” on the prairie of
northeastern Walworth County (Burlington, 1882). When Rodolph
(1900 :354) settled in the town of Gratiot, Lafayette County, snakes
were more abundant than he had ever seen them elsewhere. He
killed hundreds of rattlesnakes. Conrad Colipp when he came to
Portage in 1849 killed “thousands in the spring and summer, often
averaging a few hundred a day” (West. Hist. Co., 1880:885), in
which case he must have done little else than kill snakes. While
breaking prairie near River Falls, two men killed 39 rattlesnakes
in one day (River Falls, 1873).

The following data on the number of snakes bountied in Craw¬
ford County, furnished by Milo C. Cooper, County Clerk, show that
the timber rattlesnake is still by no means rare :

Year 1964: 4,382 mature snakes

6,086 young or unborn
Year 1965: 4,086 mature snakes

7,952 young or unborn.

Young

The young are born in early fall from eggs held within the fe¬
male. A timber rattlesnake, four feet in length, killed on the lower
Wisconsin River on August 14, 1820, was opened by the Indians,
who removed eleven young (Schoolcraft, 1821 :363). A massasauga
found at Portage on July 12, 1926, contained ten embryonic eggs
(Pope, 1926). A female taken near Nelson, Wisconsin, gave birth
on August 6, 1933, to eight young which were slightly under eight
inches in length. A second female taken in the same locality on
July 22 contained five young about 6.75 inches long (Beckenridge,
1944:152). The young when born have a button on the end of the
tail. Rattles develop later. For September 9, 1875, there is a report
of nine young massasaugas on display in Watertown (Watertown,
1875). Four young were said to have entered the mother’s mouth

32 WiscoTisin Academy of Sciences, Arts and Letters [Vol. 56

when closely pursued, and to have been killed simultaneously with
the mother. Carver (1838:297). who was in Wisconsin in 1766,
affirms that he once killed a female containing seventy fully formed
young which entered her mouth when pursued. That the female
swallows her young when in danger is an old and persistent myth.

Habits

In denning, the timber rattler is not exclusive in its associations.
It is recorded for Licking County, Ohio: “Dens were found con¬
taining very discordant materials, twenty or thirty rattle-snakes,
black-snakes and copper-heads, all coiled up together” (Howe,
1847:297). At the mouth of a den in Richland County in May,
1874, rattlesnakes and bull snakes (Pituophis) were living together
(Richland Center, 1874). Messeling (1953) found in southwestern
Wisconsin the same den occupied by seven or more species of
snakes, along with skunks and raccoons. In a den in Sauk County,
opened by blasting, Elmer Keitel found 35 snakes, rattlers, bull
snakes, garter snakes, and other species, well intermingled (Mac-
Quarrie, 1941:83).

George Knudsen has informed me that in Wisconsin the massa-
sauga winters in decayed stumps, foundations of deserted buildings,
mammal burrows, and piles of old slabs. In Pennsylvania it is said
to hibernate in fissures in the earth, burrows of mammals, beneath
heavy moss, and under overturned trees (Miller, 1938:17).

Rattlesnakes disperse in summer. According to Klauber (1956:
402), they sometimes wander two miles from the den, but usually
less than a mile. Experts in Wisconsin think that the timber rattle¬
snake usually travels less than 1000 feet from the winter den.

Neither species always gives warning by rattling, nor is it neces¬
sary to be within two or three feet of the reptile to produce it.
Messeling (1953:22) reports that about half the time the rattle¬
snake gives no warning before striking, and he has known them
to rattle when distant twenty feet. The rattle of the massasauga
is weak. When McKenney (1868:181) was at Portage, he likened
the sound to the ticking of a watch. The rattle is more like the buzz
of an insect.

Rattlesnakes are excellent swimmers. When Pond (1908:335)
descended the Wisconsin in September, 1740, he wrote: “As we
Descended it we saw Maney Rattel Snakes Swimming across it and
Kild them.” At the large den on Mount Trempealeau, the yellow
rattlesnakes swam from it in spring and returned by the same
method in fall (Brunson, 1855:114).

Rattlesnakes can climb well. They have frequently entered build¬
ings in Wisconsin, even reaching the second floor. Audubon started
considerable controversy when he painted a rattlesnake in a tree

1967-68] Schorger— Rattlesnakes in Early Wisconsin

33

containing the nest of a mockingbird. Examples of these snakes in
trees and shrubs are not rare. Many times Keitel has found timber
rattlesnakes in trees where presumably they had gone for birds
(MacQuarrie^ 1941). George Knudsen, who has caught many hun¬
dreds of timber rattlesnakes, has never found one in a tree.

A peculiar habit which does not appear in the scientific litera¬
ture is the rattlesnake's tendency to go over an obstruction rather
than around it. Pope (1923:25) kept some timber rattlesnakes in
a cage two feet high. When the lid was removed and a snake could
place its head over the edge, it could draw up its body. Garland
(1917:33), lived on a farm near Onalaska, La Crosse County,
where timber rattlesnakes were plentiful. One of the largest ever
seen on the farm was killed in the act of climbing over a barrel
in the farmyard. He wrote : “I cannot now understand why it tried
to cross the barrel, but I distinctly visualize the brown and yellow
band made as it lay an instant just before the bludgeon fell upon
it, crushing it and the barrel together.” Thomas Harry, who came
to Racine County in 1849, saw massasaugas crawl over his men
resting on the ground while breaking the prairie (Lake City Publ.
Co., 1892:264). Two Germans hired to dig and curb a well on the
old Frost farm near the outlet of Lake Monona, at Madison, re¬
ported a rattlesnake approaching from behind, had crawled up the
back and over the shoulder of one of the men, presumably reclining,
to disappear in the tall grass (Brown, 1934:8). There are several
instances of rattlers crawling to the tops of beds in log cabins.

Food

Rattlesnakes feed principally on small mammals and birds. Little
specific information exists on the food in Wisconsin. A large timber
rattler captured in Pierce County had swallowed a fully grown
gray squirrel (Sciurus carolinensis) (Breckenridge, 1944:159). The
white-footed mouse ( Per omy sens ) , cottontail (Sylvilagus) , and
young woodchuck (Marmota) (obtained by entering the burrow) ?
are mentioned by Jackson (1961:117, 129, 219). Messling (1953:
21) lists gophers (Citellus), mice, small birds, frogs, and blackber¬
ries. The inclusion of blackberries is inexplicable unless present in
the prey. The very young feed on flies. According to Hoy (1883)
the massasauga subsisted .almost exclusively on meadow voles
(Microtm) . Other writers think frogs the common food. George
Knudsen has known them to eat frogs, voles, short-tailed shrews
(Blarina), and small snakes.

Enemies

Rattlesnakes have few natural enemies. There is an old tradition
of enmity between the white-tailed deer and the rattlesnake, al-

34 Wisconsin Academy of Sciences, Arts and Letters [VoL 56

though few encounters have been observed. This may be because
of the largely nocturnal feeding habit of the rattlesnake, especially
in hot weather. Seton (1929:288) mentions a hunter seeing in
Coahuila, Mexico, a deer cut a rattlesnake to ribbons by jumping
upon it several times with all four feet. A doe attacked a rattle¬
snake in Pennsylvania in the same way (Aldous, 1938). McDowell
(1950:46) would not commit himself on the question of whether
or not deer would kill snakes, but he did affirm that deer in pens
showed the greatest terror towards snakes of all kinds. A piece of
rope manipulated to simulate a snake would prevent a buck from
charging when a club would not. Bunnell (1887 :329) mentions that
a deer would leap high into the air and, with its four feet bunched,
come down on the rattlesnake. Keitel (MacQuarrie, 1941:83) felt
certain that deer attack rattlers, although he never witnessed the
act. He had, however, found many snakes with gouges in the backs
which could have resulted only from the hoofs of a deer.

Badgers in South Dakota, according to Jackley (1938), will
attack and eat rattlesnakes, especially during hibernation. A simi¬
lar observation has not been made in Wisconsin, where badgers
were once plentiful and are still not rare.

It is probable that birds are minor enemies. Bunnell (1897 :326,
329) states that while rattlesnakes of all sizes were being killed
at a den at Homer, Minnesota, ‘‘falcons or swift hawks of the
Mississippi bluffs” would swoop down and bear off writhing snakes.
The peregrine falcon (Falco peregrinus) is not known to capture
snakes. Raptors, however, are greatly attracted to sick or injured
animals. Bunnell also mentions eagles and hawks as enemies.

In 1873, a man hauling stone from a bluff at Trempealeau ob¬
served a domestic turkey gobbler battling four rattlesnakes, two
old and two young ones. He killed the young snakes, but the old
ones escaped. The turkey was completely exhausted (Trempealeau,
1873). Several accounts in the literature report wild turkeys at¬
tacking, if not killing, rattlesnakes.

Man has been the greatest enemy of the rattlesnake since the
first European set foot in Wisconsin. He also imported an able
assistant, the hog. Keitel has said that although he has never seen
a pig killed by a rattler, he has often seen a hog kill and eat one
(MacQuarrie, 1941:83). James Allen Reed, when he settled at
Trempealeau in 1840, found the place so infested with rattlesnakes
that it was called “The Rattle Snake Hills.” The Winnebago called
it Wa-kon-ne-shan-ah-ga, meaning “the place of the sacred snakes
on the river.” Bunnell (1897:184, 327) informed Reed of a breed
of hogs noted for their skill in hunting snakes, some of which Reed
brought from Prairie du Chien. In a short time the number of
rattlesnakes was greatly reduced. Bunnell mentions that a hog, lean

1967-68] Schorger — Rattlesnakes in Early Wisconsin

35

from a scanty winter diet, rushed among the numerous snakes at
a den. After killing several, the hog instead of eating them stag¬
gered away and took refuge in a mud hole. On recovery, she showed
no further interest in rattlesnakes. The hog’s lack of fat had en¬
abled the snakes to inject their venom into the blood vessels, al¬
though it is generally assumed that hogs are immune to the venom
since the normal layer of fat prevents the fangs from reaching
the circulatory system.

It was not uncommon in Grant County at one time to find a
rattler under an unbound bundle of wheat, or for a man loading
the wheat to find that a snake had been pitched to him along with
the bundle. When hogs became numerous, the snakes were largely
destroyed (Holford, 1900:49). Green River, in northern Grant
County, was once a good trout stream where the timid were warned
not to frequent its banks until the hogs had exterminated the snakes
( Platte ville, 1854).

The Norwegian settlements in Dane, Jefferson, and Waukesha
counties were visited by Lovenskjold (1924:88) in 1847. He wrote:
'Tn some places, especially where there are large sloughs, there
are poisonous snakes, but they are reduced in number year by year,
as the land is being cultivated. Their worst enemy is the hog, and
as the settlers keep large numbers of hogs because it costs but little
to feed them in the summer, they devour the snakes wherever
found.”

Killing and eating rattlesnakes is not confined to the semi-feral
animals which comprised the stock of the first settlers.

Lethal Effects of the Venom

Many writers on Wisconsin have expressed surprise, in view of
the abundance of rattlesnakes, that so few people have been bitten
and that only a very small number have died. Of 70 Wisconsin cases
which I have found in the literature before 1880, only 12 people
were reported to have died. Nearly all the deaths occurred in areas
occupied by the timber rattlesnake. The massasauga is so small
that the amount of venom injected was rarely fatal. Some of the
fatal cases are mentioned under the counties. Of the people bitten
30 were men, 29 children, and 11 women. The fatal cases comprised
5 men, 4 children, and 3 women. Six people were hospitalized for
snake bites in Wisconsin in 1958 and 1959, with no deaths (Parrish,
1965). No fatalities occurred in Wisconsin during the ten-year
period 1950-59, although the estimated number of snakebites was
15 annually.

Probably few large domestic animals fall victim to rattlesnakes.
If the venom rarely kills a human being, the chances of horses and
cattle dying are slender. Fonda (1868:281) relates that during the

36 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

removal of the Winnebago, just before making camp on the main
Baraboo ridge on May 15, 1848, his horse was bitten on the nose
by a rattlesnake. He thought that the horse, its head swelled to
twice normal size, would certainly die. An old Frenchman offered
to cure it. The next morning the horse was well, but he learned that
all the Frenchman had done was to look at the horse and talk to it.

Information on the circumstances under which an animal died
is meager, no mention being made of a snake having been seen or
killed. In four cases where cattle were found dead, the deaths were
attributed to snake bite. A colt 18 months old was found dead in
the road soon after being bitten by a snake (Alma, 1877.2). A
mule recovered from a bite, supposedly as a result of treatment
with snakeroot (Augusta, 1878.1). One horse nearly died from a
bite (Baraboo, 1871), and another succumbed twelve hours after
being bitten (Prescott, 1866). A horse, bitten beside the Platte
River in Grant County, swelled to an enormous size, but was cured
with sage tea and milk (West. Hist. Co., 1881). Cooke (1940:286)
says that when a fine horse was bitten on the nose, his father made
it drink a quart of whiskey and it recovered.

Antidotes

The early remedies were based on folklore. Most of the physicians
of the period were on the same medical level as the country people,
their treatments doubtfully efficacious. Often it is surprising that
the patient survived the treatment rather than the snake’s venom.
By far the most popular treatment was the internal use of alcohol.
Its general use must have been intensified by the report of Dr.
Burnett (1854) , who declared that because the venom was a depres¬
sant, the best antidote was alcohol, a powerful stimulant. His find¬
ings were widely copied. Many statements testify to the fact that
regardless of the amount of alcohol taken, intoxication did not
follow.

Some of the numerous external antidotes used in Wisconsin
were: salt and onions; a mixture of gunpowder, salt and egg
yolk; gall of any species of snake; black mud and tobacco; clay;
tobacco applied to the wound and also eaten ; freshly killed chicken ;
tincture of iodine; ammonia; whiskey, saleratus (sodium bicarbon¬
ate), and cornmeal; and alum taken internally. Dr. Ward’s treat¬
ment for a child bitten at Madison was a poultice of wood ashes
and copious draughts of whiskey punch. Since the child recovered,
the treatment was recommended highly (Madison, 1855). The vari¬
ous snakeroots, of which Polygala senega was so popular elsewhere,
were rarely used. Sometimes a slit was made in the wound, or a
large piece of flesh cut from it, and suction applied by mouth. While
at Portage, MoKenney (1868:188) was told that the Indians ob-

1967”68] Schorger — Rattlesnakes in Early Wisconsin

37

tained immunity by rubbing over their bodies the dried, powdered
flesh from the neck of the turkey vulture.

A man at Fennimore, bitten by a massasauga while binding
grain, underwent heroic treatment. When questioned by Bishop
Kemper, he replied that after reaching the house he drank half a
pint of alcohol and camphor, then a quart of whiskey, followed by
a quart of pure alcohol, and all this with no symptoms of intoxica¬
tion, The following morning he drank a pint of alcohol and swal¬
lowed a quarter pound of finely cut tobacco boiled in milk (Lan¬
caster, 1866) . In a way, it is disappointing that he did not die.

The use of a tourniquet is of no value. If incision and suction are
employed immediately, about 40 percent of the venom can be re¬
moved, but they are useless if more than one-half hour has passed
since the snake bite. The only really effective treatment is with
antivenin (Hyde, 1964).

Range

The formal papers on the reptiles of Wisconsin give only occa¬
sional places where rattlesnakes have been found. Most of my data
on distribution has come from newspapers. Unfortunately the in¬
formation is often insufficient to determine the species. Usually it
is possible to determine species from the dimensions given for the
snake, or from the habitat. Because the timber rattlesnake never
occurred east of the longitude of Madison, any rattlesnake men¬
tioned east of this line was the massasauga. Approximately 275
references to rattlesnakes, mostly before 1880, have been accumu¬
lated by the author. To cite all the references to the several coun¬
ties would be superfluous. Only a few locations are spotted on the
map (Fig. 1), but every reference is included for the border of the
range. Maps showing the recent distribution occur in Knudsen
(1954.1) and Spaulding (1965).

Adams, — -A timber rattlesnake with eight rattles was killed on
the west side of Hixson Bluff (Friendship, 1869), now known as
Rattlesnake Mound, about five miles south of the village of Adams.

Buffalo.- — Ira Nelson came to the tovm of Nelson in 1855. Among
the first deaths was that of his daughter, who died from the bite of
a rattlesnake (Curtiss-Wedge, 1919 : 98) . Records of the timber rat¬
tlesnake exist for the towns of Alma, -Dover, Gilmanton, Glencoe,
and Mondovi. One killed in a field in Little Bear Creek Valley was
reported to be six feet in length and four inches in diameter. The
species was considered “quite scarce in this county” (Alma, 1874).
A rattlesnake five feet long was killed in a vacant lot in the village
of Alma (Alma, 1878,1), and one in a woodshed (Alma, 1878.4).

38 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

000«4lt-x0

Figure 1. Range of the Timber Rattlesnake and Massasauga.

Seventeen rattlesnakes were killed in an oatfield, in a space of 10
acres, near Mondovi (Mondovi, 1877.1).

Two specimens of the massasauga were taken at Nelson, town of
Nelson (Breckenridge, 1944:152).

Chippewa, — Records for the town of Eagle Point show several
persons to have been bitten, probably by massasaugas (Chippewa
Falls, 1872, 1876).

Clark. — Only one reference was found. On September 17, 1880,
an “enormous” rattlesnake, 44 inches in length, was killed in Neills-

1967-68] Schorger — Rattlesnakes in Early Wisconsin

39

ville (Neillsville, 1880), the only one ever seen in the vicinity. The
length shows that it was a timber rattlesnake.

A road-killed massasauga was found in the town of Dewhurst by
George Knudsen. It is quite common in the southwestern part of
the county.

Columbia. — Massasaugas were numerous, with many accounts of
them by travelers who crossed at Portage. In 1926, Pope (1926)
obtained two specimens which had been captured near by. One,
killed on October 9 along the canal in Portage (Portage, 1869),
gives some indication of the lateness of hibernation. Another was
killed in a barn in Portage (Portage, 1870).

The timber rattlesnake occurred along the Wisconsin River. On
September 26, 1886, a woman thrust her hand into a rock cavity in
the town of Westpoint, expecting to find nuts stored by squirrels,
and was bitten on a finger by a rattlesnake four feet long but with
only one rattle (Prairie du Sac, 1886). Another killed in the same
town had ten rattles (Prairie du Sac, 1877). A man was bitten in
the Baraboo Bluffs in the town of Caledonia (Portage, 1878).

During a period of high water, while men were working on an
improvement of the Wisconsin River at the mouth of the Baraboo
River, town of Caledonia, they killed 14 rattlesnakes. Other people
killed 12 in the same locality (Prairie du Sac, 1880).

Crawford. — Timber rattlesnakes have been found near Steuben
(Pope and Dickinson, 1928:71), and in the towns of Utica and
Wauzeka (Messeling, 1953).

Dane. — The timber rattlesnake occurred from Madison west¬
ward. James A. Jackson (1944:27). who came to Madison in 1853,
encountered while walking in the woods, locality not stated, a coiled
rattlesnake, sounding its rattle. Alvin R. Cahn, a student in zoology
in the University in 1914-1917, told me that while canoeing along
Maple Bluff, he found about a ‘'peck’' of rattlesnake bones in a
cavity exposed by a fall of rock. Apparently a slippage of rock at
some time had closed the cavity in which the snakes were hibernat¬
ing. In the western part of Section 3 town of Dane, is Rattlesnake
Bluff, so called from the former abundance of rattlesnakes (Cassidy,
1947 :200) . Following the battle of Wisconsin Heights in 1832, a
wounded soldier was laid on the ground at night at East Blue
Mounds, where the rattlesnakes gave warning (Parkinson, 1856:
361). In 1879, six large “yellow” rattlesnakes were killed at Black
Hawk Bluff (Lookout), town of Roxbury. There are other records
for the towns of Black Earth and Vermont.

The marsh which formerly covered most of the area between the
Yahara River and the capitol at Madison, contained massasaugas.

40 Wisconsin Academy of Sciences, Arts and Letters [VoL 56

Several adults and children were bitten, but none died. On May 24,
1881, a large massasauga was killed in front of the post office in
the village of Black Earth (Black Earth, 1881), This species
occurred also in the towns of Burke, Cottage Grove, Dunkirk,
Mazomanie, Oregon, Rutland, Springfield, Sun Prairie, and West-
port. They were killed in the county at least as late as 1892 (Madi¬
son, 1892, 1892,1).

Dunn. — With one exception, the records are for the southeastern
part of the county, and must pertain to the massasauga, Davis
(1911:170) making a preliminary railroad survey in 1857, at Elk
Creek found abundant a “variety of prairie rattlers.” Near Falls
City, town of Spring Brook, 35 rattlesnakes were once killed, the
heavy rains having driven them from the swamps (Menomonie,
1879). When Eugene Wiggins arrived at Falls City in 1855, these
snakes abounded (Curtiss^-Wedge, 1925:238). A man from
Menomonie, hunting prairie chickens, shot a rattlesnake which was
pointed by his dog (Menomonie, 1877).

Eau Claire. — A rattlesnake was killed in Augusta in 1870
(Augusta, 1870) and later two people were bitten near this village
(Augusta, 1878, 1880). A child and a woman were bitten at Eau
Claire (Eau Claire, 1859, 1872). There were no fatalities.

Fond du Lac. — Haas (1943:38), after he purchased a farm in
the town of Marshland in 1847, wrote that he had not met anyone
who had seen a rattlesnake. A large one, however, was killed a mile
east of Fond du Lac in June, 1875, undoubtedly a massasauga.
There was the comment: “This is a rare occurrence, as a rattle¬
snake is seldom found in this section of the state” (Fond du Lac,
1875),

Grant.— LAmher rattlesnakes, especially, were abundant. On
August 24, 1845, on an island at Potosi, when a member of Moore^s
(1946:39) party killed a rattlesnake he was informed that the
woods were full of them. There are several place names. Snake
Diggings took its name from a cave at Potosi which contained
rattlesnakes. A creek and a mound in the town of Hazel Green
bear the name Sinsinawa,* meaning rattlesnake. Rattlesnake Creek
rises in the northern part of the town of Bloomington and enters
Grant River 2.5 miles south of Beetown. An early account reports

* The origin of the name is uncertain. The word does not occur in the languages
of the Sioux, Chippewa, Winnebago, Fox, Sauk, or related Kickapoo. The Fox occu¬
pied the area prior to commercial lead mining. Very probably it is a corruption of
the Menomini name for the rattlesnake, sinawata. Schoolcraft (iL c. p. 346) used the
spelling Sissinaway for the mound. Mr. Buford Morrison of the Horton Agency, Hor¬
ton, Kansas, obtained the name Shen-weh-ah-gat from the resident Kickapoo. Mr.
Bernhard Richert of the Shawnee Agency, Shawnee, Oklahoma, has informed me that
the Sauk and Fox word for rattlesnake is Na’-to-we’-wuh, and Kickapoo, Na-to’— we’-a.

1967“68] Schorger — Rattlesnakes in Early Wisconsin

41

going from Beetown to Cassville, down Rattlesnake Valley and
across the Massasauga (Lancaster, 1844).

The timber rattlesnake has been reported from the towns of
Cassville, Hazel Green, Potosi, Waterloo, and Wyalusing.

The massasauga was found in the towns of Cassville, and Fenni-
more, and must have occurred in others. Doubtless it was a snake
of this species which bit a farmer on Balke’s Prairie, town of
Bloomington (Lancaster, 1848). Undetermined species are men¬
tioned for the towns of Marion, South Lancaster, Wingville, and
Woodman.

Grce?i.— Massasaugas, in 1836, raised their heads through the
puncheon floor of the cabin of David Bridge, town of Jefferson. A
Mr. Chadwick plowed a furrow 20 inches wide from his cabin to
the schoolhouse so that his children would not become lost in the
prairie, and : “On this furrow the children walked until the snakes,
pleased with the soft ground, took up their abode there, and then
they walked in the high grass by its side” (Bingham, 1877:167,
171). In 1875 A. W. Goddard, in Monroe, advertised for sale mens’
heavy brogans which were proof against rattlesnakes (Monroe,
1875).

Green Lake. — ^When Richard Dart came to Green Lake in 1840,
rattlesnakes were plentiful (Dart, 1910:255).

Iowa. — Timber rattlesnakes have been found in the towns of
Arena, Dodgeville, Highland, and Wyoming, where they are still
common locally. Specimens of the massasauga have been collected
at Mineral Point (Pope and Dickinson, 1928:70),

Jackson. — Three people, two of them children, were bitten near
Black River Falls (Black River Falls, 1867, 1871). Robert Ellarson
has informed me that the massasauga is still common along Hall
Creek, northwest of Merrillan.

Jefferson. — The massasauga has been recorded for the towns of
Lake Mills, Milford, Sumner, and Watertown. One was found under
a bed in Thure Kumlien’s cabin near Bussyville (Main, 1943:38).
S. W. Faville informed Hawkins (1940) that about 70 massasaugas
were killed about 1850 at a rocky den within a mile or two of
Faville Grove as they were coming out of hibernation.

(Except for the southwestern corner, the remainder of
the county was distinctly habitat of the massasauga. It occurred in
the towns of Lemon wier, Lisbon, Orange, and Necedah. Bertha
Thomson (1933:418) wrote of the vicinity of Necedah when a
child: “The rattlers were usually in the leaves, or old stumps and
logs, where the blueberries grew,” Robert Ellarson found a dead

42 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

massasauga in the road in the town of Finley, near the county
line. It is common along the Yellow River.

Rattlesnakes, species undetermined, occurred in the towns of
Lindina and Plymouth.

Kenosha. — A boy, about 20 months of age, living south of
Kenosha, was bitten on the foot by a “prairie” rattlesnake and re¬
covered (Southport, 1842). A. M. Jonsson wrote on December 9,
1843, from the town of Wheatland that the rattlesnakes were by
no means as large and venomous as they were thought to be in
Sweden (Stephenson, 1937:119).

La Crosse. — Both species occurred, but little is recorded of their
distribution. Haines (1848) in September, 1848, killed an “enor¬
mous” rattlesnake on a bluff of the Wisconsin shore opposite the
mouth of Root River, Iowa. In 1852, Ethan Roberts was told of
the attractiveness of the county, including “the large yellow rattle¬
snakes in the rocks and of massasaugas on the marshes” (Western
Hist. Co., 1881.1:465). Although the timber rattlesnake was com¬
mon, the only localities mentioned are La Crosse and Green’s Coulee
near Onalaska (Garland, 1917:32, 33, 49). Larson (1942:25), liv¬
ing on a farm in Jostad Coulee in the northern part of the town of
Hamilton, never saw more than three rattlesnakes.

Lafayette. — Rodolph (1900:354) settled in the town of Gratiot
in 1834. He wrote: “Another annoyance was the great abundance
of snakes, particularly rattlesnakes. I have never before or since
even in Florida or Louisiana seen anything like it.” Brunson (1900 :
290) mentions that in winter a rattlesnake in a cave in West Platte
Mound, near the county line in the town of Belmont, was crawling
about as in summer. A rattlesnake three feet long with six rattles
was killed near Darlington (Darlington, 1873). Smith (1838:25)
traveled south from Mineral Point to the Pecatonica, where, prob¬
ably in the town of Willow Springs, he found on the banks of the
river a “brown and yellow” rattlesnake “(Crotalus horridus)” be¬
tween four and five feet long, killed an hour or so previously.

Marquette. — On August 14, 1817, on ascending the Fox River
and arriving at Buffalo Lake, Keyes (1920:351) was informed that
rattlesnakes abounded in the country. Muir (1913 : 110) came to the
county in 1849, and while living on the farm at Fountain Lake (now
Ennis) in the town of Moundville, saw only one rattlesnake. He
mentions seeing a copperhead, a species never known to occur in
Wisconsin.

Milwaukee. — In the early days hundreds of massasaugas were
killed on what was then a marsh at the foot of Mason Street in

1967-68] Schorger — Rattlesnakes in Early Wisconsin

43

Milwaukee (Olin, 1930:214). According to Haas (1943:38), they
were common in the Milwaukee region. Mrs. Carpenter (n.d.)
arrived in 1845. In going to school at Brookfield in the warm days
in spring it was common to see massasaugas on the ends of the
logs forming the corduroy road across a long swamp.

Monroe. — Cases of snakebite were reported from the towns of
Glendale, Lafayette, Le Grange, and Oakdale. A woman in the town
of Leon killed at her doorstep a rattlesnake with nine rattles
(Sparta, 1881).

Pepin, — In the town of Frankfort, timber rattlesnakes occurred
in the bluffs, while massasaugas abounded in the bottoms between
Dead Lake, at the northeastern corner of the town, and the Chip¬
pewa River (Curtiss-Wedge, 1919:1031).

Pierce. — There are sixteen references to rattlesnakes in the
county. The timber rattlesnake occurred in the towns of Clifton,
Hartland, Isabelle, Oak Grove, Trenton, and Union. The locality
and species of rattlesnake found along Rush River in a cabin be¬
longing to Harvey Seely are uncertain (River Falls, 1859.1). At
that time, a Harvey G. Seeley lived in the town of Salem, the only
clue to the locality.

Racine. — The massasauga was formerly numerous. Two speci¬
mens collected by Dr. Hoy at Racine, about 1858, are in the U.S.
National Museum (Pope and Dickinson, 1928:70). There are rec¬
ords for the towns of Burlington, Dover, and Mount Pleasant.

Richland. — Timber rattlesnakes were numerous in the northern
part of the town of Orion and in the town of Buena Vista. In 1889
they were plentiful in the Pine River Valley (Dodgeville, 1889).
Jackson (1961:117) killed one near Gotham, where it still occurs.
One said to have been five feet in length was killed in the town of
Westford (Reedsburg, 1874).

Rock. — The massasauga must have been more numerous than
the single record indicates. When Sayre (1920:424) came to Ful¬
ton in 1849, his fear of rattlesnakes vanished after killing one at
the bridge at Stebbinsville, a discontinued post office in the north¬
ern part of the town of Porter.

St. Croix. — The northern limit of rattlesnakes was in this county.
Breckenridge (1944:154) in 1939 examined two sets of rattles of
the timber rattlesnake in the possession of a farmer in the town of
Troy, and taken years before. A man in Emerald captured a rattle¬
snake which refused food of any kind during its captivity of eleven
weeks (Hudson, 1880).

44 Wisconsin Academy of Sciences, Arts and Letters [VoL 56

Sauk. — -Both the timber rattlesnake and massasauga were com¬
mon at the time of settlement (Biihler, 1923:326; Canfield, 1870:
40). The timber rattlesnake was especially numerous at Devil's
Lake and along Honey Creek, town of Honey Creek. The first year
that the Philip P. Grubb family lived in the town of Freedom, they
killed over 60 rattlesnakes (Cole, 1918:583). J. B. Fowler, on
August 3, 1877, shot a rattlesnake five feet three inches long. His
attention had been called to the snake by his cattle circling the
place where the snake was coiled (Baraboo, 1877.1).

The massasauga occurred on the prairies, and especially along
Otter Creek. In the town of Sumpter Knapp (1947 :14) was taught
how to tear down an old rail fence and kill rattlesnakes.

Trempealeau. — There are 18 early references to rattlesnakes in
the county. The timber rattlesnake was particularly abundant at
Mount Trempealeau. It is recorded for the towns of Caledonia,
Gale, Pigeon, and Preston. The snake mentioned for Tamarac
(Trempealeau, 1873.3) may have been the massasauga. The latter
occurred in the Trempealeau Valley, but there were no timber
rattlesnakes (Heuston, 1890:52-54),

Vernon. — In 1859, both species occurred in the town of Harmony
(Button, 1955:112). The timber rattlesnake was recorded for the
towns of Forest, Liberty, and Sterling, but most frequently from
the town of Kickapoo.

Wahvorth. — -The massasauga was abundant in the town of East
Troy (Burlington, 1882). Dwinell (1874), who settled on Spring
Prairie, town of Spring Prairie, in 1836 killed seven rattlesnakes
the first summer. They disappeared about 1850. During the harvest
season, 18 were killed on a farm in the town of Bloomfield (Lake
Geneva, 1876) . It is mentioned also for the towns of Delavan and
Lafayette. Specimens have been taken in the town of Richmond
(Pope, 1930:277).

Waukesha. — Unonius (1950:297) killed two rattlesnakes while
cutting wild hay at Pine Lake, town of Merton, He remarked that
the warning was feeble; people and stock, however, were seldom

bitten.

Wood. — On July 30, 1874, six rattlesnakes of the “black species,"
with four to seven rattles, were killed in the large marsh west of
Wisconsin Rapids (Grand Rapids, 1874). A few days afterwards
one entered the house of Silas Paine, although previously they
were unknown except along the Yellow River,

1967-68] Schorger — Rattlesnakes in Early Wisconsin

45

References

Aldous, C, M. 1938. Deer kills rattlesnake. Jour. Mam. 19:111.

Alma Buffalo Co. Jour. 1862. June 5.

Alma Express. 1864. Aug. 6.

- . 1877. 2. Aug. 23.

- . 1878.1. May 30,

- . 1878.2. June 6.

- . 1878.4. Aug. 8.

Anderson, T. G. 1882. Narrative. Wis. Hist. Colls. 9:147, 192.

Augusta Herald and Madison State Jour. 1870. Aug. 10.

Augusta Eagle. 1878. June 1.

- . 1878.1. July 18.

- . 1880. July 10.

Baraboo Republic. 1871. July 19.

- . 1877.1. Aug. 8.

Bingham, Helen M. 1877. History of Green County. Milwaukee. 310 p.

Black Earth Advertiser. 1881. May 26.

Black River Falls Jackson Co. Banner and Madison State Jour. 1867. Aug. 6,
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Breckenridge, W. J. 1944. Reptiles and amphibians of Minnesota. Minneapolis.

202 p.

Brown, C. E. 1934. Prairie stories. Madison. 12 p.

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to Wisconsin. Wis. Hist. Colls. 15:264-291.

Buhler, Jacob. 1923. A Swiss family in the new world. Wis. Mag. Hist. 6:
317-333.

Bunnell, L. H. 1897. Winona and its environs. Winona, 694 p.

Burlington Free Press. 1882. Sept. 26.

Burnett, Dr. W. I. 1854. Notes on the rattle snake. Proc. Boston Soc. Nat.
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Button, G, W, 1955. A Vermonter in the west, 1859. La Crosse Co. Hist. Soc.
Ser. 8:104-112,

Canfield, W. H. 1870. Outline sketches of Sauk County. Third sketch. Baraboo.
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Carver, J. 1838. Carver’s travels in Wisconsin. N. Y. 376 p.

Cassidy, F. G. 1947. The place-names of Dane County. Greensboro. 255 p.
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Chippewa Falls Herald. 1876. Aug. 25.

Cole, H, E. 1918. A standard history of Sauk County. Wisconsin. Chicago.

1128 p.

Cooke, W. W. 1940. A frontiersman in northwestern Wisconsin. Wis. Mag.
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Curtiss-Wedge, F. 1919. History of Buffalo and Pepin counties, Wisconsin.
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Darlington Democrat. 1873. Aug. 29,

Dart, R, 1910. Settlement of Green Lake County, Proc. Wis. Hist. Soc. for
1909. p. 252-272.

Davis, A. M. 1911. A preliminary railroad survey in Wisconsin, 1857. Proc.
Wis. Hist, Soc. for 1910. p. 165-170.

46 Wisconsin Academy of Sciences, Arts and, Letters [Vol. 56

Dodgeville Chronicle. 1878. Aug. 16.

- . 1889. Aug. 30.

Dwinnell, S. a. 1874. Wisconsin as it was. Reedsburg Free Press Sept. 17.
Eau Claire Free Press and Fond du Lac Commonwealth. 1859. Aug. 24.

- . 1872. Aug. 1.

Fonda, J. H. 1868. Early Wisconsin. Wis. Hist. Colls. 5:205-284.

Fond du Lac Journal. 1875. July 1.

Friendship Press. 1869. May 26.

Garland, Hamlin. 1917. A son of the middle border. N. Y. 467 p.

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Haas, Carl de. 1943. North America: Wisconsin hints for emigrants, [Fond
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Haines, R. B. 1848. Letter September 13. Copy Wis. Hist. Soc.

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Hennepin, L. 1903. A new discovery of a vast country in America. Chicago.
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Heuston, B. F. 1890. Original conditions and early history of Trempealeau
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Higley, W. K. 1889. Reptilia and batrachia of Wisconsin. Trans. Wis. Acad.
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Holford, C. N. 1900. History of Grant County, 'Wisconsin. Lancaster. 782 p.
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Hudson Republican. 1880. July 21.

Hyde, H. 1964. Treatment of poisonous snakebites. Wis. Med. Jour. 63: 340-
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Jackley, a. M. 1938. Badgers feed on rattlesnakes. Jour. Mam. 19:374-375.
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Jackson, J. A. 1944. Autobiography of James Albert Jackson, Sr., M.D. Wis.
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Keyes, W. 1920. A journal of life in Wisconsin one hundred years ago. Wis.
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Klauber, L. M. 1956. Rattlesnakes. Berkeley. 1476 p,

Knapp, G. N. 1947. Autobiop-aphy. Wis. Hist. Soc. MS. 220 p.

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- . 1954.1. Massasauga (Sistrurus catenatus catenatus). Wis. Consv. Bull.

19(10) : 35-36.

Lake City Publ. Co. 1892. Portrait and biographical album of Racine and
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- . 1848. July 29.

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Le Sueur, J. J. 1902. "Yoyage up the Mississippi. Wis. Hist. Colls. 16:177-200.
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Wis. Mag. Hist. 8:77-78.

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- . 1892. Sept. 3, Oct. 1

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47

Main, Angie K. 1943. Thure Kumlien, Koshkonong naturalist, Wis. Mag, Hist.
27:17-39.

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p. 141.

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119(5) :5, 17-18.

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Muir, John. 1913, The story of my boyhood and youth. Boston. 294 p,
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Olin, N. 1930. Reminiscences of Milwaukee in 1835-36. Wis, Mag. Hist. 13:
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Bull. Pub. Mus. Mil. 8(1):138 p.

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Seton, E. T, 1929. Lives of game animals. N. Y. 3(1) :412 p.

48 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

Smith, W. R. 1838. Observations on Wisconsin territory. Philadelphia. 134 p.
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Sparta Herald. 1881. Aug. 16.

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- . 1873.3. Sept. 12.

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1:419 p.

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1095 p.

- . 1881. History of Grant County, Wisconsin. Chicago. 1046 p,

- . 1881.1. History of La Crosse County, Wisconsin. Chicago. 862 p.

THE WILD HONEYBEE IN EARLY WISCONSIN

A. W. Schorger

It is not known exactly when the honeybee (Apis mellifica) was
brought to North America. The literature indicates that it was in¬
troduced first into Massachusetts, but the earliest records are for
Virginia. Williams (1844) listed honey and beeswax among the
commodities produced and available in Virginia, and gave their
prices as of 1621. Evidently bees had been brought in some time
previously. A letter of December 5, 1621, from the Virginia Com¬
pany of London reported that beehives, peacocks and pigeons were
being sent to the colony for preservation and increase (Kingsbury,
1933) . Swarms escaped to use hollow trees as hives, and by the end
of the 18th century honeybees were well established beyond the
Mississippi.

Unfortunately we do not know the rate at which bees spread
westward. Bradbury (1817) wrote that in 1810 they were found
in eastern Nebraska, and that they had moved 600 miles westward
in 14 years, approximately 40 miles a year. At this rate of progress
bees would have advanced from the coast of Virginia to the Missis¬
sippi in 20 years, which is improbable. In 1754 there were swarms
of bees at the forks of the Ohio (Pittsburg) (J.C.B., 1941), and in
1782 bees were kept by the Moravian Indians at Gnadenhutten on
the Muskingum (Zeisberger, 1885, 1:80). Although in 1776 wild
bees were reported to be abundant at Detroit (Hamilton, 1908),
Zeisberger (1885, 11:316) wrote in 1793 that no bees were found
in the woods at Fairfield on the Thames (near Detroit) and bees
brought there by an Indian from the Huron River, Ohio, swarmed
twice. The dates available show poor coincidence with longitude.
In 1804, two men from the Moravian Mission near Anderson,
Indiana, went with a Cherokee squaw to fell some bee trees which
she had found (Gipson, 1938). The U.S. Factory at Chicago paid
to the Indians thirty nine cents per pound for beeswax in 1805
(Peake, 1954). Flagg (1912) wrote from Edwardsville, Madison
County, Illinois (a prairie state), September 12, 1818, that more
wild honey was available in the territory than elsewhere in the
world. Bees progressed slowly in the virgin forest, but rapidly at
the margins of grasslands.

The date of the appearance of the honeybee in Wisconsin is un¬
certain. The U.S. Factory at Prairie du Chien purchased ''wax.

49

50 Wisconsin Academy of Sciences, Arts and Letters [Vol. 56

tallow, etc/’ to the amount of $70.88 during the first half of 1818
(J. W. Johnson, 1911). An 1825 inventory of the trading post at
Fond du Lac (Superior) appraised 10 pounds of wax at $2.00
(Anon., 1911). Although neither record indicated the source of the
beeswax, it probably came from near the Mississippi. The first
mention of wild bees in Wisconsin was in 1828. In January of this
year Fonda (1859) and a Frenchman, carrying mail from Chicago
to Green Bay, discovered in southeastern Wisconsin a bee tree, re¬
vealed by the claw marks of a bear and cut it down. Fonda ate so
much of the honey that he became ill. Subsequently he could not
eat honey without a feeling of nausea. In the same year honey was
so abundant in Grant County (Hollman, 1922) that bees must have
colonized the region before 1800.

V Collection of Honey by Indians

The Indians had collected honey long before the first white
settlers came to southern Wisconsin as shown by their ladders and
bee trees which had been cut. Except in the Lead Region, the tide
of immigration was unimportant until about 1840. The Indians
were on hand to exchange honey for pork and flour.

The Potawatomies in Walworth County used crude ladders to
reach the cavity containing the honey and opened it with hatchets
('Dwinnell, 1874). The earliest settlers in Waukesha County found
a great number of Indian ladders made from tall young trees,
their branches cut off to leave prongs eight to ten inches long
which served as rungs (Western Hist. Co. 1880.3:626). At times
the Indians sought assistance from the whites. Joseph H. Stickney
came to Waukesha County in 1839. His daughter described the
procedure (Martin, 1925) : “My father said when an Indian came
of an errand, he never failed to make his want known; he would
continue to act it out in pantomime until you caught his meaning.
Sometimes it was a bee-tree he had found, and he wanted the white
man to go with him with an axe and chop the tree down. First he
made the white man understand what he had found; he attracts
his attention, then bends over and imitates the bee as he flies from
flower to flower, buzz, buzz, buzz ; then he points, as away the bee
flies with his load to his home in the distant tree, then he says,
‘armo sispoquet’* ; ‘sispoquet’ meant bee sugar or honey. Then fa¬
ther gets his axe, the Indian shows him the way to the honey the
Indian divides with him ; then taking his half, vanishes among the
trees of the forest.”

* The Potawatomi were closely related to the Chippewa, in whose language honey
was amo sisihakwat.

1967-68] Schorger- — Wild Honeybee in Early Wisconsin

51

Collection of Honey by Whites

Cane sugar was an expensive item for the early settlers, and
maple sugar could be made only in particular areas. The cheapest
and most available sweetening was honey. In fact the only readily
marketable products were deer skins, furs, ginseng, honey, and
beeswax. Many of the settlers from the east were experienced bee
hunters and some became professional collectors of honey. Greening
(1942:213) wrote at Mazomanie in 1847: '‘Parties go bee hunting
for months together in Summer, they take wagons and a pair of
oxen, an ax and coffeepot, and that’s all except barrels for the
honey. When they come to a prairie they turn out the cattle, and
watch the flowers till they see a honey bee, catch it, put it into a
box, for its humming to attract other bees, then they let it go and
watch in what direction they fly, and then search all the hollow
trees on that side. And the tree, chop it down, smother the bees
and take the honey, barrel it up, then ditto, several times a day
perhaps. They shoot for meat, roast corn in a frying pan for coffee,
barter honey for flour from settlers, bake it in a pan, and sleep in
their wagons at night.”

The use of the box as described above is incorrect. The box con¬
tained honey which the bee consumed to the limit of its capacity.
When released, it flew directly to the bee tree. The standard pro¬
cedure in Sauk County for locating a bee tree is given by Brown
(1946) : “In the spring when the plum and apple trees were in
bloom he [Uncle Isaih] took a small box, put some honey in it and
caught a dozen bees or so and put them in the box, leaving a small
crack that would allow one bee to escape at a time. When ready to
‘hunt,’ he would open a small slide and let out one bee. It came out
laden with honey to be carried to the tree. When it first escaped,
it flew around in spirals until it reached a height of twenty or
thirty feet. Then it darted away in a straight line for the bee
tree . . . After Uncle Isaih had followed the direction taken by
the bee, until he was no longer sure of the direction, he opened the
slot and let out another bee which repeated the operation.” Bees
were released until the tree was found. Occasionally an entire day
was consumed in locating the tree, but the reward might be as
much as 100 pounds of honey.

The finder of a bee tree carved his initials on the tree. Under
unwritten pioneer law, this was a claim to ownership usually re¬
spected. Unonius (1950), writing of Waukesha County where he
arrived in 1841, said that the finder could not cut the tree without
the consent of the owner; but if the owner cut the tree, he had no
right to the honey. With the influx of Europeans, the traditional

52 Wisconsin Academy of Science, Arts and Letters [VoL 56

custom broke down and honey was taken without regard to
ownership.

Abundance

It has been said that “Wisconsin was one extensive apiary’’
(Cole, 1930). This was true only of the southern two-thirds of the
state. An early observation in the middle west was that bee trees
were most numerous in the woods bordering the prairies. The
reason for this lay in the profusion of flowers which existed on the
prairies from early spring until autumn. Honey could be obtained
from forest trees such as basswood and maples only during spring.
Sufficient honey usually could not be collected to more than last
the bees until the next flowering season. When clearings were
made in the woods and crops such as buckwheat and white clover
were raised, bees appeared. About 70 percent of the bloom in the
forests occurs before June 15, while on the prairie at least 25
percent of the bloom occurs after August 15 (Curtis, 1959). The
finding of bee trees by the early settlers is accordingly of ecological
significance since it shows the presence of prairie or oak opening.
Evidence for this is found in the title of the book by James Fenni-
more Cooper, The oak openings; or, the bee-hunter (1848).

In October 1834, E. Johnson (n.d.) and companions cut 31 bee
trees in four days near the “Big Spring” between Dodgeville and
Helena. After the honey was divided among the participants, he
kept of his share a sufficient amount to supply his family for a
year, and sold the remainder in Dodgeville for $75.00. A man in
Grant County found 75 bee trees between Lancaster and Beetown
(Western Hist. Co. 1881). Perkins (1842), living at Burlington,
stated that thousands of swarms were destroyed annually by the
Indians and whites and advised how the bees could be housed and
saved. In 1841 the inhabitants of Milwaukee County petitioned
the legislature to pass a law relating to wild bees. This petition
could not be found. It evidently sought to protect the bees from
destruction when a tree was cut ; however, “the committee had not
deemed it necessary to take any action upon the subject, and asked
to be discharged from its further consideration” (House Journ.
1841).

The census of 1840 recorded 1,474 pounds of beeswax produced
in the state. Grant County led with 399 pounds. Probably nearly
all of this wax was obtained from wild bees. The census of 1850
gave a combined production of 131,005 pounds of honey and bees¬
wax, indicating that bee culture was then well under way. (The
data are for the year prior to that in which the census was taken).

1967-68] Schorger — Wild Honeybee in Early Wisconsin

53

Bee Culture

An apiary in pioneer times usually began by the capture of a
swarm of wild bees. The simplest hive was a section of hollow
tree boarded at the top and bottom. As late as 1863, mention is
made of the transfer of a colony of bees from a hive of this kind
to a “patent’' one (Madison, 1863). The wild bee was the so-called
German, or black bee. Perkins (1842) wrote: “I wished to pur¬
chase some swarms and made considerable inquiry but notwith¬
standing the vast number of swarms which have been taken, yet
from the reckless manner [in which] they have been destroyed,
and the bad management of those kept, there is scarcely a swarm
to be bought in the country.” Adam Grimm (1927), settling near
Jefferson in the spring of 1849, found the country full of wild bees
and soon formed an apiary. These bees were black and vicious.
L. Teetshorn (Watertown, 1875) was convinced that the “native
or black bees” were superior to the Italian and was limiting his
apiary to them.

In 1847, Raeder (1929) found that bee keeping was thriving in
southeastern Wisconsin. A year later Ficker (1942) was in
Mequon, Ozaukee County, where bees were kept. They were con¬
siderably more productive than in Germany. Many kinds of
patented beehives were offered at Watertown in 1849 (Watertown,
1849) . A year earlier Mellberg recorded in his diary at Lake Kosh-
konong, “Hived a swarm of bees for Mrs. Devoe” (Barton, 1946).
A beehive was robbed at Kenosha in 1851 and thrown into the
river (Kenosha, 1851).

There was considerable early discussion of the relative values of
the German and Italian bees. The opinion prevailed that the latter
were the more docile and superior in the production of honey. I. S.
Crowfoot began an apiary in the town of Hartford, Washington
County, in 1856, and is said to have been the first to introduce the
Italian bee. He had as many as 900 hives at one time (Western
Hist. Co., 1881.1). The earliest specific date that has been found
for the Italian bee is 1864, when J. W. Sharp, Door Creek, Dane
County, offered Italian queens at $5.00 each (Madison, 1864).

The leading bee keeper was Adam Grimm (1927) of Jefferson.
He died in 1876, and on his tombstone is carved a straw beehive.
He had gone to Italy in the fall of 1867, returning in the spring of
1868 with hundreds of Italian queens. Some were sold subsequently
at $20.00 each. In January, 1871, he shipped 365 swarms to Utah
(Grimm, 1871) . Only a few people were keeping Italian bees at the
time. Grimm began the season of 1870 with 600 swarms which in¬
creased to 903 during the summer. His production of honey during
the year was 22,725 pounds, which was about one-tenth of the total
production in the state (Anon., 1871). Dr. Maxson of Whitewater

54 Wisconsin Academy of Science, Arts and Letters [Vol. 56

had 100 hives of imported Italian bees in 1874. Thirty hives were
taken to the Bark River woods, where, in three days, they produced
700 pounds of honey (Whitewater, 1874) . This would be at the rate
of 7.8 pounds of honey per hive per day.

Distribution

The places where bee trees were found are shown on the map
(Fig. 1). Below, by counties, is the information that has been
found.

1967-68] Schorger — Wild Honeybee in Early Wisconsin

55

Adams. — Two men cut down a tree in the town of Springville
from which 250 pounds of honey were obtained (Friendship, 1870).
Bee trees must have been found previously for the above amount
of honey was viewed as a record. Another tree, found by James
Needham, yielded 125 pounds of honey (Friendship, 1876).

Barron. — Apiaries were started in the towns of Vance Creek and
Arland, at unrecorded dates, by the capture of swarms of wild
bees. J. P. Carlson began raising bees in the town of Prairie Farm
about 1884 (Gordon, 1922).

Clark. — Although wild bees were undoubtedly present, no record
has been found. John R. Sturdevant, Neillsville, is credited with
having introduced the first swarm of bees into the county (Lewis
Publ. Co., 1891).

Columbia. — An early settler, staying at the cabin of William
Rowan at Poynette in 1837, reported “We had good coffee and
plenty of honey’’ (Butterfield, 1880). Beyond a doubt, only wild
honey was available at that time and place. A tree found in the
town of Fountain Prairie contained 65 pounds of honey (Portage,
1878).

Craivford. — In November 1830, Johnson (n.d.) found a colony
of bees in the root of a tree on the west side of the Kickapoo
River, town of Wauzeka.

Dane. — The fall of 1829, Johnson (n.d.) hunted for bee trees at
Blue Mounds. He took the honey, along with onions and potatoes
which he had raised, by ox team to Fort Winnebago for sale. Rose
Schuster Taylor (1945), born in 1863, daughter of Peter Schuster
who settled near Middleton in 1855, wrote: “Wild bees deposited
their delicious honey in hollow trees. We gathered it on cold days
when the bees could not fly and could not sting us since such bees
were truly wild. Many pounds of wild honey were added to our
supply which was used as a sugar substitute in sweetening as well
as for corn bread and griddle cakes. White sugar cost 15 cents a
pound, and brown sugar was only a little less.” The early hunting
for bee trees at Mazomanie has been mentioned.

Dodge. — In the town of Herman, in the fall of 1848, Reuben Judd

“took over thirty swarms of wild bees” (Western Hist. Co., 1880).

Dunn. — Two men, after an absence of eight days, returned to
Durand with over 500 pounds of strained honey obtained along
Wilson Creek in the center of the county (Durand, 1863). In 1864
Mrs. Thomas Huey came to the home of 0. Cockeram, town of
Lucas. Mrs. Cockeram “had some honey for supper which they told

56 Wisconsin Academy of Science, Arts and Letters [Vol. 56

us had been gotten out of a tree in the woods ^ which we thought
very wonderful then'’ (Curtiss-Wedge, 1925). That year wild
honey was reported to be very abundant and bee hunters were
prospering. Honey cost 30 cents a pound (Menomonie, 1864). In
1879, in the town of Dunn, many swarms of wild bees were found
in the woods (Menomonie, 1879) . A year later bee trees were found
in the town of Weston, the woods along Knights Creek being men¬
tioned (Menomonie, 1880).

Fond du Lac. — Government surveyors in the town of Calumet in
1834 noted that numerous trees had been cut by the Indians to ob¬
tain honey. Reuben Simmons, who settled in the town of Empire
in 1840, took butter, eggs, and honey, presumably wild, to Green
Bay (McKenna, 1912). At this time the Indians brought honey for
sale or exchange (Western Hist. Co., 1880.1). Titus (1936) adds
that the settlers obtained maple sugar and honey from the woods.

Grant. — Beetown, nine miles southwest of Lancaster, is said to
have obtained its name in 1827 when a large bee tree blew down,
exposing lead ore, one piece of which weighed 425 pounds (West¬
ern Hist, Co., 1881). Another version derives the name from local
mining activity (Lancaster, 1845). Hollman (1922) brought his
family to his cabin near Platteville April 9, 1828). Some men sud¬
denly left the cabin which was in a filthy condition : “in the other
corners were troughs full of honey in the comb, and kettles and
pans full of strained honey, which had been procured by the miners
from Tee trees’ found in the vicinity.”

James Grushong came to the Hurricane district, town of South
Lancaster, in 1836 when bees were so numerous that a bee tree
could be found almost anywhere (Western Hist. Co., 1881). About
two gallons of honey were obtained from a cave in the bluffs bor¬
dering the Mississippi, just below the entrance of the Wisconsin
(Platteville, 1841). Holford (1900) wrote: “Little sugar did they
have to buy; the wild bees of the woods had laid up in many a
hollow oak an abundant store of sweets gathered from the in¬
credible profusion of prairie flowers,”

Green. — The county seems to have been well supplied with wild
honey, John Dougherty established a trading post at the “diggings”
near Exeter in 1831, After the Black Hawk War was over he re¬
turned to the mines and “found his merchandise, which had been
left buried in the ground much injured by moisture; but a barrel
of metheglin which had been made early in the spring To keep’
was found so much improved that all present drank immoderately,
forgetting, until intoxication came, the unusual strength of its in¬
gredients.” There were enough bee trees around Monroe to furnish

1967-68] Schorger — Wild Honeybee in Early Wisconsin

57

sufficient honey for the inhabitants. In 1843 John Adams, while
looking for a bee tree in the town of Adams, discovered the Badger
Diggings. Honey Creek, which rises near Monroe and flows into the
Pecatonica, got its name from the felling of a bee tree to form a
bridge (Bingham, 1877).

Sylvester Hills came to the town of Albany in 1838. The sweets
required for the family were provided by maple sugar and wild
honey. T. B. Sutherland, who came with his family to the town
of Sylvester in 1843, mentioned the cutting of an oak to get the
honey in it (Union Publ. Co., 1884). According to Hiram Brown,
town of Albany, wild honey bees were quite plentiful between 1842
and 1850. He wrote that in 1838, a “swarm of my bees’’ settled in
the hollow limb of an oak which was later cut to obtain both bees
and honey (Butterfield, 1884).

Green Lake. — In 1840 the family of Richard Dart (1910) settled
near the Twin Lakes, town of Green Lake. He wrote: “We also
had splendid wild honey from the bee-trees.”

Iowa. — The large number of bee trees found in 1834 has been
mentioned (Johnson, n.d.). Foster (c. 1840) wrote from Helena:
“Some make a business of hunting for honey, furs and deer.” The
Jones family came in 1857 to the town of Arena, where “bee trees
were eagerly sought by the younger generation and bee keeping
was carried on as a side line by some of the more enterprising
farmers” (Jones, 1938).

Jackson. — Robert Douglas settled near Melrose in 1839. An In¬
dian brought him honey in the comb, obtained from a bee tree
(Polleys, 1948).

Jefferson. — Much attention was given in this county to hunting
for wild honey and bee keeping. William Ball was a noted bee
hunter at Jefferson in pioneer days. Buck (1876) reported that he
would find from one to three swarms a day, and that “fifty-two
swarms were taken up by us, upon the town site alone.” Cartwright
(1875), in the early I850’s, lived in the town of Sullivan in the
Bark River woods. Here, “Bees thronged in multitudes of swarms,
and their honey was very abundant. I commenced with my neigh¬
bor, Mr. Thomas, to hunt bees and we were very successful.’' A bee
tree was found in which a bear had made an unsuccessful attempt
at gnawing an opening. The tree when cut yielded over 160 pounds
of excellent honey.

The Coes settled in 1839 in the town of Ixonia where a man
named Smith was a very successful bee hunter (Coe, 1908). Hart
(1925-26) was born at Ft. Atkinson in 1840. Expert bee hunters
could find honey in the Bark River woods. The wife of Charles

58 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Rockwell, one of the pioneers of Ft. Atkinson, in the spring of
1838 traded pork and flour for the honey brought by two Pota-
watomies (Western Hist. Co., 1879).

Juneau. — In the early days, according to Kingston (1879), wild
honey could be obtained in any desired quantity. He wrote: “As
an instance of the abundance ... it may not be out of place to
state that Zach. Sheldon came up from Portage City in the fall of
1851, and at the end of a four weeks’ bee hunt, took home eight
barrels of strained honey.”

Kenosha. — In the fall of 1836, Kellogg (1924) came to the farm
of relatives near Kenosha. He was served biscuits and honey as his
uncle had found a bee tree. Quarles (1932) wrote from South Port
(Kenosha) on February 14, 1839, that 60 to 70 pounds of strained
honey were obtained from a tree.

La Crosse. — Manly (1927) and a trapping companion went down
the Black River into the Mississippi, then down to Prairie du Chien
(his chronology is awry and instead of May, 1847, it must have
been 1844). On the way they found two bee trees. About 1865,
when Hamlin Garland (1917) was a small boy, he was taken on a
visit to his grandparents in West Salem. Hot biscuits and honey
were served. “I am quite certain about the honey,” he wrote, “for
I found a bee in one of the cells of my piece of comb and when I
pushed my plate away in dismay grandmother laughed and said,
‘That is only a little baby bee. You see this is wild honey. William
got it out of a tree and didn’t have time to pick all the bees out of
it’.” In 1889 he visited the farm, at adjacent Neshonoc, of his
uncle, William McClintock, who was an expert in tracking wild
bees.

Marquette. — In the fall of 1865, James L. Jones of Packwaukee,
took 103 pounds of honey from a tree (Montello, 1865). John Muir
(1913), who came to Marquette County in 1849, wrote that honey¬
bees were not seen until several years later. They were probably
overlooked. In 1860 his parents moved to the Hickory Hill farm,
town of Buffalo. After hearing men on the farm talk of “lining”
bees with a box containing honey, he tried it, and traced bees to a
hollow, bottom log in a fence. Someone had chopped a hole in the
log and removed the honey. In May, 1879, Christopher Kellogg of
Buckhorn found a bee tree, hived the bees, and took 25 pounds of
honey (Westfield, 1879).

Milwaukee. — Honey Creek rises in Sec. 26, town of Greenfield,
and flows north into the Menomonee. As early as 1841 the legisla¬
ture was petitioned to protect the wild bees in the county.

1967-68] Schorger — Wild Honeybee in Early Wisconsin 59

Outagamie. — Mrs. Ellen Van Tassel came to Hortonville with
her parents in 1852. They easily found bee trees, so that honey was
available in quantity (Ware, 1917).

Ozaukee. — Cigrand (1916) wrote: “The Indians gathered honey
which was plentiful in the hollow trees of this part [Sauk Creek]
of Ozaukee County. They strained the honey and then poured it
into large hollow gourds, corked it and then in canoes paddled into
Lake Michigan.” The honey was taken to Milwaukee and sold.

Pierce. — The dates found for bee trees are so late that the
swarms could have been escapes from apiaries as well as original
wild bees. A bee tree found September 4, 1877, at Lost Creek, town
of El Paso, yielded about 60 pounds of honey. At the same time a
man was reported hunting bees (Ellsworth, 1877). A number of
bee trees were found in August 1880, in the town of Maiden Rock
(Ellsworth, 1880).

Racine. — Henry Trowbridge came to Racine in 1836. Wild honey
was obtainable in the woods (Lake City Publ. Co., 1892). The
winter of 1837-38, a man living in the town of Caledonia traded
an ox for a barrel of flour. Having found a bee tree he invited his
neighbors to partake of biscuits and honey (Kellogg, 1924). The
abundance of bee trees at Burlington has been mentioned (Perkins,
1842).

Richland. — Johnson (n.d.) in 1840 was living in the town of
Richwood, nine miles below Muscoda. At Christmas he cut down
what he thought was a coon tree but it contained a swarm of bees.
He sawed off a section containing the bees and placed it in the root-
house where he had his bees. It seems probable that many people
had swarms of wild bees at an early date. In November 1843,
Samuel Swinehart and Thomas Parrish explored Pine River and
feasted on the honey found in a tree. According to Israel Janney,
wild bees ,were plentiful in 1846, and hunting them for the honey
was profitable. James M. Cass came to the town of Richland in
1851. Some honey spilled in one of the wagons attracted the wild
bees. The bees were followed and two swarms which were found
yielded 150 pounds of honey. Honey was also plentiful in 1845 in
the town of Rockbridge (Union Publ. Co., 1884.1).

Rock.— IjQvi St. John, who came to Janesville in 1836, wrote:
“I have frequently visited their [Indian] camps, gone into their
wigwams and bought honey and maple sugar of them” (Guernsey,
1856). Ogden (1838) recorded in his diary finding several bee
trees, at Milton in the fall of 1838 and in the spring of 1839.

60 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Sank, — Honey Creek rises in the northwest corner of the town
of Honey Creek and flows southeast into the Wisconsin River. It
is conjectured that the name was derived from the abundant
amount of honey collected by professional bee hunters (Western
Hist. Co., 1880.2). Opinion differs as to the amount of honey to be
found along the creek. F. J. Finn thought the supply was limitless.
An early settler, under urgent pressure to pay for his land, col¬
lected, with the aid of his wife, so much honey that it brought him
over $100 in sales to neighboring settlements. Mrs. Henry Keifer,
who arrived in 1846 after Honey Creek had already been named,
reported bee trees here and there, but not in profusion. A Mr.
Jassop of Ironton is also credited with payment for 40 acres of
government land with the proceeds of the sales of wild honey ( Cole,
1918).

Bee trees were so common in the town of Lavalle that honey could
be obtained with little difliculty. A bee hunter in the town of Iron-
ton is reputed to have taken to market 1500 pounds of honey in a
single load (Western Hist. Co., 1880.2). Edmond Rendtorff (1861)
came to Sauk City in 1840. Although he found wild honey he did
not know the procedure for securing it. A bee tree found on Web¬
ster’s Prairie, town of Delton, contained 135 pounds of honey in
the comb (Baraboo, 1869). In the fall of 1886, four bee trees were
found near Cassel Prairie, town of Troy (Prairie du Sac, 1886).

Sheboygan. — In the town of Lima, in 1839, A. G. Dye frequently
accompanied Indians to fell bee trees which they had found. Honey
was also obtained in quantity in the towns of Russell and Lyndon
(Zillier, 1912). Joseph Benedict wrote on November 25, 1845, that
there was plenty of wild honey (Buchen, 1944).

Trempealeau. — A farmer near Trempealeau reported honey
stolen from a tree near his home (Arcadia, 1878) . Bee keeping must
have been established at this time because F. A. Goodhue of Arcadia
had 25 swarms for sale at $5.00 each (Arcadia, 1879). At Inde¬
pendence two young men found a bee tree after a search of several
days (Independence, 1878).

Vernon. — In the early days at Kickapoo Center, according to
Mrs. Cyrus D. Turner, the best fare was “pancakes with pumpkin
butter or wild honey” (Union Publ. Co., 1884.2).

Walworth. — Honey Creek rises in the town of Troy and flows
east-southeast into the Fox River. Its name was bestowed in the
fall of 1835 when Jessie Weacham and Adolphus Spoor found honey
which the bees had collected from the prairie flowers (Western
Hist. Co., 1882). Dwinell (1874), settling in the town of Spring
Prairie in 1836, found that the Indians were accustomed to collect-

1967-68] Schorger — Wild Honeybee in Early Wisconsin

61

ing wild honey. In 1845, the hollow oaks in the town of East Troy
contained swarms of bees which collected honey from the woods
and prairies (W.H.M., 1882). Joseph Nichols was a celebrated bee
hunter at Whitewater in 1837. Having accumulated about 200
pounds of honey, he drew it to Milwaukee on a hand sled and ex¬
changed it for provisions. That year an Indian, as a reward for
being fed, brought Mrs. Norman Pratt a pail of honey (Cravath,
1906),

Waukesha, — Almon Welch settled in the town of Vernon in 1837.
In the fall of 1839, he and N. K. Smith found 40 swarms of bees.
The honey was sold in Milwaukee for $60.00. His share went far
towards paying for his claim (Western Hist. Co., 1880.3). In 1840
Charles D. Parker attended a school in the town of Muskego. As
usual the teacher boarded around he reported, ‘‘but there was no
butter or milk in most places. Honey was substituted for both’’
(Showerman, 1926). Unonius (1950) has described the method of
locating bee trees by the settlers near Pine Lake. When a proper
tree was found, it was left until winter. A section containing the
swarm was cut off and taken home. Barker (1913) arrived in
Milwaukee June 14, 1845, and the family settled in the town of
Brookfield. His brother was a good hunter and supplied venison
and wild honey.

The first swarm of bees owned by William Addenbrooke, town
of Mukwonago, was captured in the woods about 1860. For a time
he was in partnership with George Grimm, son of Adam Grimm
of Jefferson. In 1879 Addenbrooke had 150 swarms of pure and
hybrid Italian bees (Western Hist. Co. 1880.3).

Waupaca, — Honey Creek is a small stream emptying into the
Pigeon River at Clintonville. The creek was named by N. C. Clin¬
ton, who came to the site of the town in 1855. An enthusiastic bee
hunter, he found many bee trees on the banks of this stream (Wake¬
field, 1890). In the fall of 1883, Jim Turney found three bee trees
near New London (New London, 1883).

Waushara, — The first land claim in the town of Leon was made
in 1849 by a bee hunter named Worden. Evidently he was a member
of the exploring party which in the fall of that year hunted game
and bees near a lake which they called Lone Pine, possibly Pearl
Lake. Isaac and William Warwick settled in the town of Marion in
September 1848, and in the following spring they obtained a large
amount of honey from a bee tree (Acme Publ. Co., 1890) .

Winnebago, — Lockwood (1847) recorded in his diary on October
1, 1847, that he went with a local resident into the oak timber south
of Oshkosh in search of bee trees.

62 Wisconsin Academy of Science, Arts and Letters [Vol. 56

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241.

WISCONSIN PINELAND AND LOGGING MANAGEMENT

George W. Sieber
Department of History
Wisconsin State University — Oshkosh

In the second half of the nineteenth century, sawmills located in
the middle and lower districts of the Mississippi Valley did not
have adequate forest resources near them. They were dependent
upon the northern pineries located on the rivers St. Croix, Chip¬
pewa, Black, Wisconsin, and their tributaries. The downriver firms
could buy logs from independent logging contractors, but in order
to make certain of their supply, some of the larger companies in¬
vested in pinelands and stumpage themselves. The purchaser of
stumpage had the right to cut timber, but did not retain the land.
Companies with timberlands at their disposal either expanded op¬
erations and sent men into the woods, or contracted to have their
timber cut by independent loggers.

This article describes problems of absentee land ownership and
the business techniques of pineland management and contract log¬
ging experienced by one of the largest sawmill firms on the Missis¬
sippi, W. J. Young & Company of Clinton, Iowa. Established in
1858, the capital of the firm totaled a little over $1,000,000 in
1882,^ including the value of over 60,000- acres of pinelands lo¬
cated mostly in the vicinity of the Chippewa and Flambeau rivers.^

W. J. Young obtained the pinelands through his association with
John McGraw of Ithaca, New York. Already a wealthy lumber¬
man and timber owner, McGraw bought out Young’s earlier part¬
ners, but left the management of the firm in the Clinton lumber¬
man’s hands. On becoming Young’s partner, McGraw agreed to sell
him approximately 31,000 acres of pinelands at $9 an acre, and
to contribute a like amount of forested area to the firm as an equal
partner. After McGraw’s death. Young purchased his partner’s

1 N. Y. Court of Appeals. In the Matter of the Estate of John McGraw, Deceased,
and also in the Matter of the Estate of Jennie McGraw-Fiske, Deceased. Return to
the Court of Appeals. (5 vols. ), v. 3, Testimony, pp. 1487-88, cited hereafter as
McGraw— Fiske Testimony.

^Ihid., p. 1080. W. J. Young' & Co. Pineland Reg-ister, not pag-inated.

^W. J. Young- to John Dean, Minneapolis, Minn., Mar. 24, 1880, LPB 57, p. 104. To
D. A. Park, Minneapolis, Minn., Mar. 15, 1880, LPB 57, p. 101. Note: Unless otherwise
stated, manuscript sources are from the W. J, Young & Co. special collection at the
University of Iowa, and Young- wrote his correspondence at Clinton. The notes desig--
nate the particular record, letter-press-book (LPB), or box where information is found.

65

66 Wisconsin Academy of Science, Arts and Letters [Vol. 56

interest for $7,000,000 and the pinelands were among the assets
which continued under his direction.^

The task of looking after the pinelands was so extensive that
Young had to hire an agent to manage that end of the business.
Daniel Page Simons was originally from Dryden, New York, had
served in the Union army over three years, and thereafter had
become a timber cruiser (a person who located lands and estimated
their value) and agent for large land holders.^ Simons had charge
of the W. J. Young & Company pinelands beginning November 1,
1876.® He arranged for the payment of taxes, guarded against tres¬
pass, and handled financial affairs with logging contractors for a
salary of $100 per month.

The Clinton firm did not engage in logging operations itself, but
Simons made contracts with independent loggers to cut the com¬
pany’s timber. Simons’ contracts usually stated that the logger
agreed to cut, haul, bank, and prepare rollways or landings by the
stream for all marketable pine timber on lands which were accu¬
rately described according to township, range, and section. All of a
season’s operations should take place prior to the first day of May.
Logs were to be cut into lengths as directed by Young from time
to time, and if he so required, the logger had to cut one fourth of
the whole amount into logs 26 feet, 6 inches to 40 feet, 6 inches.
Contracts specified a minimum diameter for the small end of logs,
usually 12 inches in the 1870’s. Logs were to be ready for driving
in the spring, plainly marked, and all of the operations done in a
'‘good and workmanlike manner.”^

Young agreed to pay a higher price per thousand (M) feet for
long logs than for shorter ones. Long logs were 26 feet or more in
length. Payments were made only upon the certificate of a scaler,
or person who estimated the number of board feet in logs by
measuring them with a scale rule calibrated to allow for the taper¬
ing of the trunks. The logs, straight and sound, were to be scaled

* McGraw-Fiske Testimony, v. 3, pp. 1073, 1396.

^ Ihid., p. 1505.

" W. J. Young- to D. P. Simons, Eau Claire, Wis., June 5, 1878, W. J. Young & Co.
papers, microfilm collection. Main Library, University of Iowa.

LSee Simons’ financial statements. Box 162.

** Contract between W. J. Young & Co., and Elias Moses, Minneapolis, Minn., Nov. 9,
1876, Box 91-A. Contact with W. F. Price, Black River, Wis., Nov. 20, 1877, Box 162;
and with Philander A. Viles, Eau Claire, Wis., Nov, 13, 1877, Box 162.

1967-68] Sieber — Wisconsin Pineland and Logging

67

by Scribner's rule, which tended to read a smaller number of board
feet in logs of large diameters than did the Doyle scale.^

Young paid the wages of the scaler and boarded him without
charge. Contractors received $1 per thousand feet on the tenth day
of each month for logs banked the previous month; followed by
$.50 per thousand feet on a designated day in April, and another
$.50 per thousand feet when the landings were broken and the logs
fully ready for the spring drive. The balance was due in two equal
payments, on the tenth day of the following September and
October.!®

A logger contracted to carry out his operations for a stipulated
sum per thousand feet. One of the most important variables in
determining logging prices was the location of the timber in rela-
tion to the stream — the closer the distance, the easier the job, and
the lower the price. If a logger underestimated his expenses he
would sustain a loss. Young was not disposed to adjust his pay¬
ments to cover such a deficit. In 1880 he refused to do so, saying
that it would encourage others to make similar claims, and that
the practice might tend to destroy the force of contracts.!!

Of the clauses in the contracts. Young considered the most im¬
portant to be the ones stating that logs should not be shorter than
12 feet, 4 inches; that as many long logs be cut as practical; and
that the ends of the logs be ‘Tutted" or crosscut square. He wanted
the logs trimmed smooth in the woods where it could be done
cheaper than in the mill, and he would not have to run the risk of
breaking machinery on immense limbs. Having said this. Young
allowed Simons to add to the contracts whatever provisions he
thought desirable.!^

Young instructed Simons to make contracts with the “best" and
“safest" loggers, because “good men are worth a premium." By
safe loggers. Young meant responsible persons who did good
work.!® Tpg Clinton lumberman did his part by arranging for each
logging camp to receive copies of the Northwestern Netvs, sl tem¬
perance publication.!^

® Both rules were orig-inally used by log-gers in the northeastern states. The State
of Minnesota made the Scribner rule the standard in 1854, and Wisconsin adopted it
in 1871. According to the Northwestern Lumberman trade journal (Aug. 21, 188 6,
p. 3), the Doyle scale eventually replaced the Scribner rule, and beginning in 1872,
Scribner’s Lumber and Log Booh actually contained the Doyle scale. See also : Robert
P. Pries, Empire in Pine, the Story of Lumbering in Wisconsin 1830—1900 (Madison:
State Historical Society of Wisconsin, 1951), p. 38 ; and William G. Rector, Log Trans¬
portation in the Lake States Lumber Industry 1840-1918 (Glendale: Arthur H. Clark
Co., 1953), pp. 81-82.

Contracts in Box 162 as cited in note 8.

^W. J. Young to D. P. Simons, Eau Claire, Wis., July 24, 1880, LPB 57, p. 195.

^W. J. Young to D. P. Simons, Eau Claire, Wis., Sept. 29, 1880, LPB 57, p. 246.
To C. G. Bradley, Osceola, Wis., Jan. 8, 1864, LPB 5, p. 150.

^W. J. Young to D. P, Simons, Eau Claire, Wis., Mar. 1, 1879, LPH 57, p. 16.

W. J. Young to E. W. Brady, Davenport, la., June 22, 1883, LPB 57, p. 283.

68 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Young insisted that Simons receive a monthly report from the
scalers of each of the several logging camps which might be oper¬
ating simultaneously in different areasd^ The contractors and
scalers were not supposed to receive their pay until the reports
were complete/^ identifying the location of the camps by township,
range, and section ; and containing a record of the number of logs
cut in various lengths ; and the amount of board feetd"^

From the reports, the company compiled surprisingly intricate
statistics that represented the prospective stock down to the last
log and its length. This information was useful because the com¬
pany took orders for future delivery based on the prospective sup¬
ply of logs. In the case of long timbers, moreover, the firm took
orders based on the prospective supply of logs of each specific
length. Before accepting an order calling for a large number of
timbers for a railway bridge, for example, the company wanted to
be sure of its supply of long logs.^®

The cutting of logs into various lengths was not a haphazard
undertaking, but was carefully planned. After trees were felled in
the woods, the ‘‘buckers” sawed the logs into the proper lengths,
which could be a few inches longer than the lumber to be sawed at
the mill. It was not however, permissible to saw logs a few inches
too short. Logs were always cut into sizes from which lumber
could be sawed into even lengths of 12, 14, 16, 18, 20, 22, or more
feet. Odd lengths of lumber were seldom sawed, and only by spe¬
cial order. In order to saw lumber 19 feet in length, for example,
it was necessary to use a log meant for boards 20 feet long.

Young gave considerable attention to figuring out the amount of
the various lengths of logs needed to match the demand of the
market. He frequently suggested to the independent dealers and
loggers the lengths that they should provide for the Clinton trade.
He once stated that 16 feet was the best average length, but the
situation was changeable each year depending upon supplies on
hand. In 1863, for example, he called for more 12 and 14 foot
lengths, and suggested that out of every million feet, a good pro¬
portion would be one-sixth in 12 foot lengths, one-sixth in 14 foot
lengths ; and that 32 foot lengths would be very convenient for long
timber.^®

J. Toung- to D. P. Simons, Kau Claire, Wis., Mar. 1, 1879, LPB 57, p. 16.
Georg-e W. Forrest, Clinton, la., to D. P. Simons, Eau Claire, Wis., Feb. 6, 1882,
LPB 57, p. 574.

W. J. Young to D. P. Simons, Phillips, Wis., Jan. 12, 1880, LPB 57, p. 78. George
W. Forrest to D. P. Simons, Eau Claire, Wis., Jan. 23, 1883, LPB 57, p. 827,

George W. Forrest, Clinton, la., to D. P. Simons, Eau Claire, Wis., Feb. 8, 1887,
LPB 57, p. 324.

’^W. J. Young to B. H. Hollway, Onalaska, Wis., Jan. 1, 1863, LPB 3, p. 514.

1967-68] Sieber — Wisconsin Pinelctnd and Logging

69

With estimates of the lengths of logs needed for the coming
season, Young could apportion work to loggers according to a sys¬
tematic plan. Young estimated the lengths of logs he needed for
1877 by averaging the amount of each length sawed at his mills
during April and October, 1876. April was an early sawing month
of low productivity; October was a late season time of large pro¬
duction. To obtain an average monthly figure for each length,
Young added the amount sawed in April and October and divided
by two. Then he multiplied by 12 to record the amount for a year.
The total amount of the cut would vary, however, from year to
year, and the important thing was the proportion of each length to
be sawed. Therefore, Young figured the percentage of each length,
and the proportion of each he would need, based on his estimate
of the next year's total cut. The total cut for the year varied with
Young's view of the lumber market, and his estimate of the num¬
ber of feet of logs needed. Other factors, however, could enter into
the decision to cut timber on specific tracts of land. A contract
for stumpage could contain a time limit for the felling of trees,
and a provision that Young would in the meantime pay the taxes.
Moreover, logging operations were planned so that no small areas
were left uncut; it was too expensive to send a logging force into
an area a second time for an insignificant amount of timber.-®
During the season of 1876-1877, Young's contracts for opera¬
tions in the woods called for more than double the amount of board
feet cut any previous year. The Chippewa Herald reported that
Young's logging contract with Elias Moses of Minneapolis was
probably the largest of its kind ever made in the West. Out of a
tract estimated to contain 250,000,000 feet of timber, Mr. Moses
expected to cut 20,000,000 feet the first winter; and thereafter
20,000,000 to 50,000,000 feet per year. The contract covered only
the delivery of the logs to the banks, and not the driving.^^ Moses
fell short of his goal for the season of 1876-1877, but his loggers
did cut, haul, and bank 67,033 logs which totaled 15,079,060 feet.
Allowing $2.50 for logs under 26 feet in length, and $3 for longer
ones. Young paid Moses $39,099.63 in seven payments on and be¬
tween February 7, and October 10, 1877.^^

Taxes and assessments were persistent problems, particularly
for absentee land owners, because local assessors might try to dis¬
criminate against them. These matters were complicated, however,
and at times complaints by land owners lacked substance. Never¬
theless, as an agent for absentee land owners, D. P. Simons spent

20 W. J. Young- to D. P, Simons, Eau Claire, Wis., Nov. 11, 1881, LPB 57, p. 512.

21 The Ghippexoa Herald as quoted in the Glinton Age, Dec. 8, 1876.

22 W. J. Young- to J. M. Williams, Minneapolis, Minn., Sept. 8, 1877, LPB 52, p. 651;
and Oct. 11, 1877, LPB 53, p. 24.

70 Wisconsin Academy of Science, Arts and Letters [Vol. 56

much of his time traveling around Wisconsin townships, examin¬
ing local taxation procedures, and seeking refunds or adjustments
of taxes on lands that he held to be unfairly assessed. Simons
stressed that lands had to be properly assessed as the only way to
avoid something 'dike confiscation more than legal taxation.”
Simons explained in 1884 that town officers tended to levy a par¬
ticularly high tax on the lands of non-residents, but that he had
obtained a reduction of about $5,000 on lands already cut.^^

Taxes were higher than ever in 1884, Simons said, but not over
one and one-half per cent of the actual value of the land. The com¬
pany reduced the valuation of some lands by removing pine, but
Simons believed that tax officers tended to increase the rate on
what remained. The only solution was to keep a close watch on the
officials, who changed frequently, and necessitated “continued and
constant care to get them half right. Sometimes Simons claimed
that he encountered outright corruption and he cooperated with
other large interests in seeking tax reductions.^®

The management of pinelands involved numerous negotiations of
right-of-way. Young authorized Simons, for example, to allow other
parties to build a dam on his land providing they paid for all tim¬
ber used in construction and for any fiood damage to other trees.
Also, free of charge, W. J. Young & Company could use the dam
to store water and increase the probability of successful drives.^®

Simons guarded against trespass on the part of loggers who
might encroach on his employer’s lands, and he watched for steal¬
ing on a small scale. In 1884 Simons caught a man who had cut
several choice trees and hauled them to sell to a railroad for bridge
timber. The agent believed that this arrest would help discourage
such happenings.^’^

Once Simons was himself caught in the middle of a trespass dis¬
pute between two of his employers. Simons was negligent and
allowed Young’s loggers to cross over a boundary and cut over
2,000 acres that belonged to Henry W. Sage. Sage sent another
agent, Mr. Emery, to accompany Simons and survey the loss, and
the lands that Young offered in exchange.^® Simons accepted
Emery’s judgment of the lands, but Young became considerably

D. P. Simons, Eau Claire, Wis., to W. J. Young- & Co., July 19, 1884, Box 173.
2*D. P. Simons, Eau Claire, Wis., to W. J. Young- &. Co., Jan. 28, 1884, Box 162.

^ D. P, Simons, Merrill, Wis., to W. J. Young- & Co., Feb. 20, 1884, Box 162, See also:
Paul Wallace Gates, The Wisconsin Pine Lands of Cornell University : A Study in
Land Policy and Absentee Oivnership (Ithaca: Cornell University Press, 1943), pp.
163-164.

2«W. J. Young to D. P. Simons, Eau Claire, Wis., Jan. 27, 1883, LPB 57, p. 831; and
Mar. 3, 1883, LPB 57, 857.

27 D. P. Simons, Eau Claire, Wis., to W. J. Young, Sept. 1, 1884, Box 162.

28 D. P. Simons, Eau Claire, Wis., to W. J, Young, Apr. 24, 1878, W. J. Young & Co.
Papers, Microfilm Collection, University of Iowa.

1967-68] Sieber — Wisconsin Pineland and Logging

71

irked when the number of board feet that actually materialized in
logs and lumber from Sage’s land failed to approach the estimate.-^
The Clinton millman complained to Simons, . . you were in Mr.
Emerys [sic] hands, as clay in the hands of the potter, and to our
humiliation we find ourselves adrift without a pilot compass or
rudder.

Young suspected that Sage had written Simons about the matter,
and he requested that any such letters be sent to him to be laid
before Douglass Boardman, a mutual acquaintance of everyone in¬
volved, and a sort of adviser-arbiter in the dispute.^^ Young, who
suspected that Sage might exercise undue influence with Simons,
wrote to Boardman that the agent had showed '‘great weakness,
and I am afraid a sprinkling of negligence — an unpardonable ig¬
norance in treating this matter of land exchange. Emery has
clearly outgeneraled him, and H. W. Sage is a good prompter.

Without the backing of Simons, Young was humiliated. Since
many of the logs had turned out to be of poor quality, the Clinton
lumberman honestly felt that Sage asked too many acres in ex¬
change. Young decided to settle on Sage’s terms, but he wrote to
Boardman that he might have some sworn experts examine the
lands all over again and publish the results, along with all the cor¬
respondence in the matter, as a “precedent for lumbermen.” As
for Sage, Young commented, “He has tried to insult me in this
whole matter, but he may realize that there is a God in Israel yet.
I have no desire to retaliate on Sage but he must do right or I
will tell him he has done wrong.”^^ A month later the parties
settled the matter and Young wrote, “Mr. S. is a curious man, but
no doubt thinks he is all right, a good many kind traits but hard in
a trade.”^^

Owners of large tracts of pine who held their lands for a consid¬
erable length of time could make immense profits through the ap¬
preciation of stumpage values. On November 1, 1890, W. J. Young
<& Company sold 41,251.26 acres to the Mississippi River Logging
Company. The contract stipulated that the logging company should
not cut more than 30,000,000 board feet of timber in any one year

2^* W. J. Young- to Doug-lass Boardman, Ithaca, N. Y., Apr. 20, 1878, W. J. Young &
Co., Papers, Microfilm Collection, University of Iowa.

30 W. J. Young to D. P. Simons, Eau Claire, Wis., June 5, 1878, W. J. Young Papers,
Microfilm Collection, University of Iowa.

SI IMd.

s^W. J. Young to Douglass Boardman, Ithaca, N. Y., June 5, 1878, W. J. Young &
Co. Papers, Microfilm Collection, University of Iowa.

33 lUd.

3*W. J, Young to Douglass Boardman, Ithaca, N. Y., July 6, 1878, W. J. Young &
Co. Papers, Microfilm Collection, University of Iowa. Interestingly enough. Young there¬
after had an agreeable relationship with Sage, who loaned him large sums of money
personally, and arranged for him to borrow other funds from Cornell University.

72 Wisconsin Academy of Science, Arts and Letters [VoL 56

until it finished paying for the land, and meantime should pay the
taxes. W. J. Young & Company reserved the right to profit from
one-tenth of all minerals that might be discovered. The Clinton
firm received $694,887.50, or approximately $16 per acre for land
valued at approximately $9 an acre in 1875.

Profits awaited the owners of pinelands whether the timber be
cut or held for speculation. Surprisingly systematic procedures
could increase those profits, particularly if the absentee land owner
found a loyal, talented agent to stand guard and handle the admin¬
istrative details for a salary of $100 per month.

Ledg-er F, 1890-91, p. 53. See also: Mississippi River I.iOg-g-ing Co. papers, Box 1,
1871-99, in the Minnesota Historical Society, St. Paul.

MARY MORTIMER:

CONTINUITY AND CHANGE AT
MILWAUKEE FEAAALE COLLEGE

Walter F. Peterson
Lawrence University

During the early years of the nineteenth century it was hardly
an open question whether it was worthwhile to educate a girl or
whether her fragile mind could stand it. However, under the leader¬
ship of such reformers as Emma Willard, Catharine Beecher, and
Mary Lyon, ambitious women began to fight the traditional social
attitudes which insisted that woman's place was in the home and
that women were inherently inferior intellectually to men. Despite
these obstacles, schools for young women were established by
Emma Willard at Troy, New York, in 1821, by Catharine Beecher
at Hartford, Connecticut, in 1828, and by Mary Lyon at Mount
Holyoke, Massachusetts, in 1838.

Milwaukee Female College, during the period 1853-1873, stood
as an extension of the educational concepts of the second quarter
of the century, as they related to women, and as an institution
caught in the currents of change in the third quarter of the nine¬
teenth century. The key figure in this study of continuity and
change is Mary Mortimer.

Born in Trowbridge, Wiltshire, England, on December 2, 1816,
Mary emigrated with her family to New York five years later.
Until she became of age her educational ambitions were frustrated
by a tight-fisted guardian and her only formal education was a
brief time in the common schools and a short span at an academy
at Auburn, New York,^ In July of 1837, when she was legally able
to take possession of her share of the family estate, she entered
Madam Ricord’s Seminary at Geneva, New York. A friend later
wrote, “It was at Geneva that she first began really to live."^ Mary
threw all her energies into her studies. She tackled Chemistry,
Mental Science, Latin and Paley's Natural Theology. Then she
turned to Algebra, Evidences of Christianity, Ancient History and

1 William W. Wight, Annals of Milwaukee College, 18^8-1891 (Milwaukee, 1891),
p. 3, Also, Milwaukee Sentinel, July 23, 1877, p. 2.

^Minerva Brace Norton, Mary Mortimer: A True Teacher (New York, 1894), p. 9.
This volume is particularly valuable because it includes a great deal of Mary Morti¬
mer’s correspondence, all of which, unfortunately, has since been destroyed.

73

74 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Astronomy. Finally there was Rhetoric, Moral Philosophy, History
of Modern Europe, Geometry, French, and Butler’s Analogy.
Through concentrated effort she succeeded in completing the four
year course in two years.^

Her deep interest in the areas of Moral Philosophy and Evi¬
dences of Christianity and her intense discussions with her teach¬
ers drove her to think earnestly about the state of her own soul.
During her first year at Madam Ricord’s Seminary, Mary Mortimer
experienced a religious conversion and joined the Presbyterian con¬
gregation at Geneva.^ She was also inspired by her teachers to
become a teacher. Miss Thurston, a teacher at the seminary wrote,
“In accordance with my advice she decided to remain in school
another year and to adopt teaching as her life work. Her great
desire now was to dedicate her time and talents to that employ¬
ment in which she could be most useful,”^

For the next ten years Mary Mortimer gained experience for her
life’s work teaching in various places in western New York. During
this time she grew from a novice assistant at Geneva to a capable
administrator of Le Roy Seminary, during the time when the regu¬
lar principal traveled in Europe on a leave of absence. She regarded
teaching as her calling and her vineyard was the education of
women. In 1857 she wrote to a friend, ‘T shall never lose my inter¬
est in female education, to which I long ago desired to dedicate my
life. I have no wish or thought of changing that dedication.”® In
1848, while travelling through Michigan, she came to the conclu¬
sion that her calling, her vocation, should be exercised in the West.
“Indications have pretty nearly decided me to remain somewhere
in this western land. I trust I have been and still am watching the
indications of Providence and desiring to be led in the path of
duty.”^

It was, then, only natural that when an opportunity came to es¬
tablish a school in Milwaukee that she would take it. This oppor¬
tunity appeared in the person of Miss Catharine Beecher. For
nearly twenty years Miss Beecher had been developing a plan for
women’s education. In Mary Mortimer she and her sister, Harriet
Beecher Stowe, were happy to find one of the “original, planning
minds’’^ to help with this undertaking. Miss Beecher stated her

pp. 12-13.

* For a detailed account of the conversion of Mary Mortimer see Walter F. Peter¬
son, “Mary Mortimer: A Study in Nineteenth Century Conversion,” Journal of Presby¬
terian History, June, 1963, pp. 80-88.

5 Norton, pp. 23-4.

<^Ihid., pp. 176-177.

'ilhid., p. 77.

^Dictionary of American Biography, XIII, p. 252,

1967-68]

Peterson— Milwaukee Female College

75

philosophy in succinct fashion: '1 am one who believes that
‘woman’s wrongs’ are to be righted, not by putting her into the
profession of the other sex, but in fitting her for her profession,
and giving her employment in it.”^ This was to be accomplished
by establishing endowed non=sectarian colleges for women in the
Westd^ In a letter to Mary Mortimer, Catharine Beecher indicated
what the aims of such schools should be. “First, an effort to over¬
come sectarianism; second, opposition to large boarding houses;
third, more thorough and practical education for girls ; fourth, bet¬
ter positions for female teachers.

Mary Mortimer was selected to implement this plan and Milwau¬
kee was ripe for the experiment. The growing city had no college,
no high school and the public schools were poorly staffed, crowded
and generally inadequate.^^ In 1849 she arrived in Milwaukee. With
the cooperation of Mrs. Lucy A. Parsons, founder of a private
seminary, the plan was put into operation. By 1851 the Normal
Institute and High School of Milwaukee, a new institution with a
college charter had been established. Mary Mortimer was one of
four co-equal department heads and leading Milwaukee citizens
were members of its board of trustees. During the next two years
the school acquired a new building and a new name — Milwaukee
Female College.

By 1852 the “Collegiate School” had been^ divided into four
classes but the program allowed for a great deal of flexibility. The
Preparatory Class covered “an indefinite period of time, according
to the health, capacity and circumstances of the pupil.” The Junior,
Middle and Senior classes covered the period of one year each. “The
Course of Study has been arranged with a view to ordinary capaci¬
ties, Those who are below this standard must acquire a fuller
preparation before leaving the Preparatory Class; those who are
above, can carry on the study of one or more branches more ex¬
tensively than their class. During the academic year 1852-1853,

» Catharine Beecher, Suggestions in Regard to the M. F. College, undated MS, State
Historical Society of Wisconsin Manuscript Collections.

Catharine Beecher and Mary Mortimer spent the summer of 1852 at the home of
Harriet Beecher Stowe drafting plans for the American Woman’s Educational Asso¬
ciation. This association was to further “The Plan” by securing an endowment for
Milwaukee Female College and other similar schools to be established particularly in
the West. This grand scheme largely failed due to the inability of the association to
raise the necessary funds.

11 Wight, p. 3.

12 Bayrd Still, Milwaukee: The History of a City (Madison, 1948), pp. 216—218.

1® First Annual Catalogue of Officers and Pupils of the Milwaukee Normal Institute
and High School (Milwaukee, 1851).

14 First Annual Report of the Officers and Pupils of the Milwaukee Female College
(Milwaukee, 1853), p. 12.

76 Wisconsin Academy of Science, Arts and Letters [Vol. 56

2 students were enrolled in the Senior Class, 10 in the Middle Class,
32 in the Junior Class and 61 in the Preparatory ClassJ^

Milwaukee Female College, according to the Beecher plan, was
to be unique in the annals of higher education. Miss Beecher pro¬
posed that the faculty be composed of co-equal teachers, instead of
a principal and subordinate teachers. This was supposed to result
in '‘increased thoroughness in the course of instruction, and, by
the division of responsibility, greater security in the health of
teachers. In point of fact, this plan was attempted only very
briefly. Catharine Beecher as a theoretician was primarily inter¬
ested in the administrative organization and such peripheral as¬
pects as the architectural appearance of the college. Mary Mor¬
timer, however, was essentially a practical educator interested in
the curriculum and instruction. After a brief experimental period
Mary Mortimer came to be viewed by the faculty, the board of
trustees, and also the community as the head of the school and
she proceeded to direct the affairs of Milwaukee Female College.

Mary Mortimer succeeded in imparting to her students an en¬
thusiasm for study and a strong religious impulse. Her methods,
born of independence of thought and a spirit of skepticism, were
rather unique for that day. Because her religious conversion had
been through intellectual rather than emotional processes, Mary
Mortimer was convinced of the importance of developing a skepti¬
cal, searching mind in her students. A former student recalled that
this was particularly true in Bible and Mental Philosophy classes.
“How often would she suffer us to lose ourselves in the labyrinth
of metaphysics, and then carefully seek to guide our way into the
light of truth. It seemed sometimes as though she sought to make
us stronger doubters, that we might prove more sincere. This
teaching technique, used so effectively by Mary Mortimer, but such
a far cry from the Mount Holyoke of Mary Lyon,^® was carried
directly from her academic and religious experience in New York.

With the support of the trustees, Mary Mortimer in 1852 intro¬
duced a course of study which divided the curriculum into three
departments: The Department of Mathematics and Natural Sci¬
ences ; the Department of Geography, History and Mental Science ;
and the Department of Languages, Belles Lettres and Composi¬
tion.^^ While the curriculum was basically patterned after the of-

pp. 5-9.

iG First Annual Catalogue, p. 11. Also, Catharine Beecher, An Appeal to American
Women in Their Own Behalf (Milwaukee, 1851), p. 2.

1'^ Norton, p. 157.

In the Memorial: Ttventy -Fifth Anniversary of the Mt. Holyoke Female Seminary
(South Hadley, 1862), p. 51, it is interesting- to note that they had maintained an
exact count of the number of students who had been converted prior to g-raduation.
Nothing like this ever occurred at Milwaukee Female College.

19 First Annual Report, p. 13.

1967-68]

Peterson — Milwaukee Female College

77

ferings at Miss Ricord's Seminary in New York, it placed greater
emphasis on history and English literature and introduced French
into the offerings.

Milwaukee gave increasing support to Milwaukee Female Col¬
lege in terms of raw numbers. During the term beginning in 1852,
the total student enrollment had been 105, but by 1856 this number
had increased to 256. However, a breakdown of this total by classes
would indicate that the concept of a complete collegiate education
for young women received only token endorsement. Only 6 of the
total number were seniors, 11 were middlers and 49 were juniors.
The remaining 190 students were all in the Preparatory Class.^^^
But those few students who did complete the entire course re¬
ceived an excellent education for that day for Mary Mortimer
insisted on high standards. The senior year was brought to a close
by a public oral examination to which the local clergy, professional
men, and others were invited. The guests as well as the faculty
committee and trustees were free to ask questions of the candidates
for graduation and enter into the discussions. During one of the
oral examinations a theological discussion developed “on the ques¬
tion of our moral capacity for doing good when evil passions have,
as it were, overcome and consumed our good promptings. Here
there was a terrible battle in which the class, the divines of the
committee, and Miss Mortimer all seized weapons.

Catharine Beecher had provided the inspiration for the estab¬
lishment of Milwaukee Female College. She had also induced Mary
Mortimer to become a member of the faculty, to implement her
plan, and to represent her in relations with the trustees and the
community. The trustees were a most competent group. Men such
as Alexander Mitchell, Milwaukee’s leading banker, and Increase
Lapham, who had gained an international reputation as a scien¬
tist, provided stability and inspired confidence. However, the ab¬
sentee direction of the institution was most unsatisfactory for both
the trustees and for Mary Mortimer. Mary Mortimer was supposed
to direct the college at the same time that she was supposed to be
co-equal with the other members of the Board of Instruction. She
also found it difficult to interpret the long, demanding, querulous
letters of Catharine Beecher to the trustees.^^ Catharine Beecher
proved to be an incredibly difficult woman. By 1857 the role of
mediator became too much for Mary Mortimer and she took a posi¬
tion as principal of a female seminary at Baraboo, Wisconsin.

^ Fifth Annual Catalogue, p. 9.

Wight, p. 16.

^Catharine Beecher to Increase Lapham, Feb. 18, 1856. Writing from Columbus,
Ohio, Miss Beecher in this letter insisted on the erection of a building M^hich the trus¬
tees did not think necessary, the importation of her own carpenters and workmen
from the East and an immediate reply to a long list of questions. State Historical
Society of Wisconsin Manuscript Collections.

78 Wisconsin Academy of Science, Arts and Letters [Vol. 56

In 1866 the trustees invited Mary Mortimer to return to Milwau¬
kee Female College on her own terms. She now had an opportunity
to introduce further curricular changes. Through the division of
the Department of Geography, History and Mental Science into
the Department of Moral and Mental Science, in which she was the
sole teacher, and the Department of Geography and History,
greater emphasis was placed on history than was usual at that
time. Also, the amount of time devoted to Latin and classical litera¬
ture was reduced to provide more time for English literature. It
is also significant that German, as a companion to French, was
introduced as a modern language.^^

Although the curriculum of Milwaukee Female College, with its
emphasis on history, English literature, and modern languages,
was liberal for that day, Mary Mortimer held to the traditional,
highly structured pattern and failed to anticipate the introduction
of elective studies as was found at Smith College when it opened
its doors in 1875.^^ However, this is not to suggest that she did
not maintain widespread professional relationships. In fact, she
chose as her successor Professor Charles S. Farrar, an acquaint¬
ance of some years, who had been the first chairman of the Science
Department when Vassar opened in 1865.^^ In her own quiet way
Mary Mortimer worked to keep abreast of Eastern educational de¬
velopments and to open intellectual doors in the Milwaukee area.

Mary Mortimer was always conscious of the importance of a
strong board of trustees and the relation of the board to the com¬
munity. In 1872, Milo Parker Jewett, first president of Vassar
College, who had moved to Milwaukee after personal differences
with Matthew Vassar, was elected vice president of the board of
trustees. He was to provide enlightened educational leadership for
the trustees and the college until his death in 1882.^^ While Mary
Mortimer “had no sympathy with so-called ‘women’s rights,’
she did feel that leading women of Milwaukee should be on the
board of a woman’s college and would provide a valuable link with
the community. To that end, she had three Milwaukee women ap¬
pointed to the board in 1869 and this number was increased to 5
in 1872.-8

In the spring of 1874, Mary Mortimer retired from Milwaukee
Female College. During her years at the college she had contact

^Annual Catalogue, (1866-1867), p. 14.

Henry M. Tyler, “The Curriculum,” in L. Clark Seelye, The Early History of
Smith College, 1871-1910 (Boston, 1923), pp. 162-164.

^ Wight, p. 33. For Vassar curriculum in this period see Henry Noble MacChacken,
The Hickory Limh, (New York, 1950), pp. 55-56. 58-59.

^Dictionary of American Biography, vol. X, p. 70.

^Milwaukee Sentinel, July 23. 1877, p. 2.

^Annual Catalogue (1869-1870), p. 3, and Annual Catalogue (1872-1873), p. 3.

1967-68]

Peterson — Milwaukee Female College

79

with some 1,500 young women, about 200 graduating from the
colleger® At her death, in 1877, a former colleague paid a fitting
tribute to Mary Mortimer. “As a teacher of young ladies, Miss
Mortimer has stood in the front rank for more than thirty years
and especially in the West her influence has been widely extended
and deeply felt.'’®® Her successor as principal of Milwaukee Female
College, Professor Charles S. Farrar, may have thought it politic
but his words were nonetheless true when he said, “That life was
the most important in the founding and development of this insti¬
tution; and whatever in the future it may become, the chief per¬
sonal story will be that of Mary Mortimer."®^

29 Lilian Bacon Mallory, “Milwaukee College” in J. W. Stearns, ed., The Columbian
History of Education in Wisconsin (Milwaukee, 1893), p. 702
^Milwaukee Sentinel, November 14, 1877. p. 2.

^ Ibid., July 18, 1877, p. 8. The institution for which Mary Mortimer set a high stand¬
ard of academic leadership and excellence was merged in 1895 with Downer College
of Fox Lake, Wisconsin, to form Milwaukee-Downer College. On July 1, 1964, Mil¬
waukee— Downer College and Lawrence College, Appleton, Wisconsin, merged to become
Lawrence University.

A PROPOSITIONAL INVENTORY OF
EXECUTIVE-LEGISLATIVE CONFLICT*

A. Clarke Hagensick
Associate Director, Institute of Governmental
Affairs, University Extension

and

Associate Professor of Political Science
The University of Wisconsin-Milwaukee

The tradition of separation of powers in American government
focuses attention on executive-legislative relations and how they
are shaped by underlying doctrines. Such attention ranges from
The Federalist Papers through the standard textbooks on American
government. Yet nowhere is there set forth in a systematic way
the fundamental characteristics of executive-legislative relations.
This study draws upon the generally accepted wisdom to present
six tentative propositions on the nature and scope of executive-
legislative conflict. Milwaukee County is used to illustrate these
propositions, because its relatively recent creation of a chief execu¬
tive office dramatized over a short time the fundamental conflicts
between the executive and legislative establishments.'^

Since 1960 Milwaukee County has had an elective county execu¬
tive. He is chosen for a four-year term at the Spring nonpartisan
election. A board of supervisors wields county legislative powers.
Its 24 members are elected concurrently with the executive for
four-year terms. Prior to the creation of the chief executive office,
the board necessarily and partially performed some of the functions
granted to the new office. These were usually handled through
administrative oversight activities of board committees. Percep¬
tions of the inability of a legislative body to provide effective execu¬
tive leadership contributed to demands for the creation of the chief
executive office for the county.

I. Constituency differences produce executive-legislative conflicts.
The prevailing basis for the selection of chief executive officers is
through election within the political jurisdiction as a whole. Legis-

* The Research Committee of the Graduate School, University of Wisconsin, provided
summer salary support for this study. Michael C. Quinn gave invaluable assistance in
gathering and analyzing data used in the study.

1 The position of Milwaukee County Executive was created in 1959, and the first
incumbent was elected in 1960. In 1968 the Board was increased to 25 members.

81

82 Wisconsin Academy of Science, Arts and Letters [Vol. 56

lators are generally elected from subdivisions of political units,
usually single member districts in which only one legislator is
elected in each district. Milwaukee County follows the normal pat¬
tern. Its chief executive is elected by plurality vote of the entire
county. Twenty-four supervisors are selected from as many dis¬
tricts within the county.

Nelson Polsby cites constituency differences as an important in¬
gredient in Presidential-Congressional relations, and argues that
conflict is heightened by a tendency of each participant to over¬
represent his constituency.- Similarly, Wilfred Binkley comments
upon the typical presidential tactic of using messages to Congress
as a device to communicate with “the larger audience, the American
people.”^

The conflict between the executive and legislative establishments
is often translated into executive defense of “general” interests
and legislative protection of “local” interests. As representatives
of a specific area, legislators are expected to guard that area's in¬
terests. It is presumed that the executive rises above “selfish” in¬
terests and fights for programs of value to the entire jurisdiction.
This point is highlighted by Don K. Price when he states that “the
greater threat of political interference . . . had typically throughout
all our history come less often from the President . . . than from
local interest groups exerting pressure on members of the
Congress.”^

Two results are implied from the basic proposition, and these are
seen in the Milwaukee County situation. One is that a shrewd chief
executive can use the localized reward system in bargaining with
legislators for support of the executive's general program. For
example, in an effort to replace a Park Commissioner who opposed
the County Executive's program to reorganize the park system,
the executive nominated a south side suburbanite. This was inter¬
preted as an effort “to curry favor from supervisors who have been
lukewarm to his program.”®

A second result is that the legislator's constituency does not
tend to be particularly concerned about the issues that produce the
greatest stress between the executive and legislative branches.
Even if the executive's constituency supports him and his program,
that support may not be translated into effective pressures upon
legislative representatives. A study of Congressional representation
showing that on a host of important national issues congressmen

^Congress and the Presidency, Eng-lewood Cliffs, N. J. (Prentice-Hall), 1964, p. 102.
^President and €ongress, 3rd Ed., New York (Random House), 1962, p. 238.

* Government and Science: Their Dynamic Relation in American Democracy, New
York (Oxford U.P.), 1962, p. 63.

^ Milwankee Journal, January 13, 1965.

1967-68] Hagensick — Executive-Legislative Conflict

83

hear nothing from their constituents illustrates this point.® Milwau¬
kee County Board members have similar experiences. Several in¬
dicated that they hear very little from their constituents on the
issues pushed by the executive. They were much more likely to be
approached by constituents on matters of localized district concern.

This is also implied by a comparison of the votes by district on
two constitutional amendments pertaining to the office of county
executive and support for the executive by representatives of those
districts. One provided constitutional authorization for the county
executive office, instead of its statutory base theretofore, and the
other granted the executive veto powers over county board enact¬
ments. They were approved by overwhelming majorities in Novem¬
ber, 1962, and every district supported them. Moreover, there was
a significant positive correlation between the level of district sup¬
port for the referenda and the social rank of the districts."^ That is,
the districts ranked highest in terms of the educational achieve¬
ment and income of their residents gave greatest support to the
referenda. One might expect a similar correlation on roll call votes.
Then representatives of districts of highest social rank would give
the executive greatest support and vice versa. However, there was
no such correlation.

Similarly, the county executive had no opposition in running for
reelection in 1964. Yet the strong support of the executive’s con¬
stituency did not increase support for his program by county board
members. Analysis of roll call votes within the board after the elec¬
tion did not show significant differences in the pattern of support
for the executive’s program. Board members clearly did not receive
the same signals from their individual constituencies as the county
executive did from those constituencies as a whole.

In this connection the use of a nonpartisan ballot for the election
of the executive and the supervisors may heighten the impact of
constituency differences between the two branches. It is alleged that
on the national and state levels political parties modify or cloak the
separation of the executive and legislative branches. Lacking the
overt participation of parties, the county may illustrate more
sharply the basic executive-legislative relationship.

II. Legislators representing a relatively large and readily identi¬
fiable minority ivithin the political jurisdiction will tend to guard
legislative prerogatives against presumed executive encroachment.
The bloc of legislators must be large enough so that, by maintaining
a solid front against the often quiescent majority, it may safeguard

® Lewis Anthony Dexter, “The Representative and His District’’ in Nelson W. Polsby,
Robert A. Dentler and Paul A. Smith, Politics and Social Life, Boston (Houghton
Mifflin), p. 496.

Based on Shevky-Bell Social Rank Index.

84 Wisconsin Academy of Science, Arts and Letters [Vol. 56

the group’s interests. Developing expertise in using the legislative
process to that end is also typical of a strong minority bloc within
a legislature. This stake in the legislative process is likely to make
the group antagonistic toward the exercise of executive power.
Within Congress the South traditionally fit into this political role.
State politics provide similar examples.

The southern half of Milwaukee County has developed a political
self-consciousness consistent with its minority status within the
entire county. This area contains eight of the county’s 24 districts,
and its spokesmen frequently defend south side interests. From
one member of its county board delegation came the threat: ‘T
won’t go for one dime for swimming pools until the south side is
taken care of.”^ From another: ‘T feel that the county executive
has been neglecting the south side in making his appointments.”^
While other supervisors might complain about similar issues, it is
unlikely that their complaints would stress ‘‘north side” or “west
side” interests.

In 1960 the leader of the south side bloc became chairman of the
county board. It was possible for him to build a majority coalition
because of 1) his long tenure on the board, 2) his previous position
of vice chairman of the board, and 3) the failure of his opponent to
expend much energy in mobilizing support.’^ The coalition formed
to select the chairman did not remain intact. Frequently, the chair¬
man was defeated on issues he strongly supported. However, his
selection meant that the board was led by the leader of a faction
which had greatest fears of expanded executive authority. The ex¬
tent of this opposition was reflected in several ways. Table I illus¬
trates that on county board roll call votes on issues of concern to
the executive the south side bloc provided his greatest opposition.^^
This bloc does not exhibit the same solidarity in other important
issue-areas, such as welfare or appropriations. It appears that the
executive issues raised special threats to the sectional interests of
this minority group.

Another illustration of the bloc’s antagonism toward the county
executive came during the battle for ratification of the aforemen¬
tioned constitutional amendments. While the county executive cam¬
paigned for approval of the amendments, the board chairman
opposed them. As part of his efforts, the eight south side super¬
visors, joined by three other board members, issued a manifesto

^Milwaukee Journal, September 21, 1960.

^Milwaukee Journal, July 19, 1962.

Milwaukee Journial, April 27, 1960.

This was based on roll call votes in which more than one supervisor voted in the
minority. Only those votes in which the executive’s position was clearly known through
speeches, messages to the board and/or newspaper accounts, are tabulated.

1967-68] Hagensick — Executive-Legislative Conflict 85

Table I. Support of County Executive’s Position By County
Supervisors on Divisive Roll Call Votes, 1960-1963 (In

Percentages; Supervisors Identified By District
Number; South Side Districts in Italics)

N = 84

1st Quartile

2nd Quartile

3rd Quartile

4th Quartile

1 — 77.4

7—67.5

22 — 54.9

19 — 38.3

9—77.1

3—64.1

13 — 51.8

14 — 38. 1

4—76.2

15 — 63.8

6—51.2

24 — 37.8

5—71.4

18 — 61.4

12 — 43.9

23 — 37.7

20 — 70.7

16 — 58.2

10 — 42.8

11 — 33.7

21 — 70.2

2 — 56.2

17 — 40.2

8—26.5

denouncing the effort to grant veto powers to the executive. One
of the signers argued : “The veto is a tremendous weapon. It results
in one man rule and we just aren’t ready for it,”^^

Without question, executive veto powers posed a critical threat
to the chairman and his followers. Given the cohesiveness of that
group, it was often possible for it to pick up the necessary four or
five votes from other supervisors to pass measures of special in¬
terest to the bloc. To muster sixteen votes to override a veto seemed
to be an almost hopeless task.

Other issues viewed as threats to legislative prerogatives drew
heavy attack from the south side bloc. These included administra¬
tive reorganization proposals, suggestions by the executive for re¬
organization of the board’s committee structure, and appointments
to county boards and commissions. These were expected to make
the executive more powerful at the expense of the board. Therefore,
the predominant minority bloc on the board was quick to defend
its legislative interests.

III. Institutional role playing is an important source of executive-

legislative conflict. Because of institutional pride, traditions and
customary methods of operation, organizations develop sets of roles
for their members designed for organizational self-preservation
and the maintenance or increase of the organization’s importance.
Legislative bodies often are more susceptible to the manifestations
of institutional role playing than other organizations. Their multi-
membership with an absence of hierarchical authority contributes
to that result. So too does the complicated system of rules used by
legislatures to carry on their business.

As William S. White notes, the Senate’s censure of Joseph R.
McCarthy for conduct unbecoming a Senator illustrated institu-

Milwaukee Sentinel, September 15, 1962; Milwaukee Journal, October 30, 1962.

86 Wisconsin Academy of Science, Arts and Letters [Vol. 56

tional self-preservation.^^ Another context in which legislative role
playing typically develops is when one chamber of a bicameral
legislature takes action affecting the other without proper notice.
The recriminations that result are reminders that legislative bodies
are quick to feel alleged slights and defend against them. The an¬
tagonism is intensified if the legislature preceives an outside threat
to its status and prestige.

Institutional role playing within the Milwaukee County Board
was illustrated following the grant of executive veto powers. The
first use of this new power occurred with a veto of an ordinance
providing music in the Courthouse. The board fell one vote short
of the two-thirds majority necessary to override. Thereupon, one of
the board’s leaders switched his vote. The action was interpreted as
a victory for supervisors who “sought to establish a strong prece¬
dent” in their relationship with the executive. That two other
board members supported the measure only after it was vetoed
gives additional weight to the impact of institutional role playing.’^
The final act in the Courthouse musical came a year later when
funds were necessary to continue the program. A board committee
voted unanimously to kill the measure. The County Executive’s
entrance into the committee room prompted a supervisor to remark :
“He won’t be able to veto it this time.”^^

Institutional role playing is intensified when members of the
organization feel that their power is threatened. For example, the
county treasurer brought together municipal officials to discuss
property tax billing. The County Executive commended the treas¬
urer for his initiative. The board chairman was not enthusiastic
and said : “The only thing wrong with it was that the county board
was not notified.” The treasurer hastily apologized for his slight of
legislative prerogatives.^^ Similarly, a group of supervisors threat¬
ened to boycott a ground-breaking ceremony simply because the
executive’s office scheduled it. The suggestion of a boycott at a cere¬
monial rite illustrates how quickly legislators may feel that they
are being upstaged.

Institutional orientation is also exemplified by the executive. The
first county executive previously served as chairman of the county
board. In that capacity he occupied a middle-of-the-road position
in a consensus-oriented legislative body. He presented policy goals
to the board, but he tended to emphasize goals which did not sharply

^Citadel: The Story of the U. S. Senate, New York (Harper’s), 1956, pp. 126-33.
See also Donald R. Matthews, U. S. Senators and Their World, Chapel Hill (Univ, of
North Carolina Press), pp. 101-102, for a discussion of “institutional patriotism.”

Milwaukee Sentinel, May 1, 1963.

15 Committee meeting', July 7, 1964, Three of the four members of the committee who
were on the board the previous year had voted to override the veto.

Milwaukee Journal, July 22, 1962.

1967-68] Hagensick- — Executive-Legislative Conflict

87

divide it. He had the added advantage of speaking from within
the board. As county executive his role vis-d-vis the board became
markedly different. As policy initiator, he was no longer circum¬
scribed by his legislative leadership role. Moreover, a county-wide
constituency added impetus for the presentation of bold, far-
reaching changes in county policies. Consensus was clearly more
difficult to attain on these recommendations.

The change in roles is seen in a comparison of the support he
received from county supervisors first as board chairman and then
as executive. He was supported as board chairman in a range of
55 to 72 per cent. As executive, the range was 27 to 77 per cent.
Apart from some limitations in the data, the controversial issues
pushed by the county executive are an important cause of this
change in support.^^ Differences between the roles of a chief execu¬
tive and a legislative chairman contribute to the marked change.
The strong executive pushes controversial divisive issues, because
successes on those issues produce a noteworthy administration. This
is similar to Polsby's argument that different conceptions of time
underlie congressional-presidential relations. The President is in¬
terested in what he can accomplish in the limited time remaining
in his tenure. Congressmen, especially those in leadership positions,
usually expect to be around for many years. With that gauge the
need for haste cannot seem pressing^® The executive then must
force his issues against what appears to him to be a lethargic
legislature.

Another ingredient of institutional role playing in executive-
legislative relations is that awareness of the conflict tends to in¬
crease the conflict. Battle lines become hardened as anti¬
administration groups develop in the legislature. When the Mil¬
waukee County Executive attacked his opponents in a strongly
worded message to the board early in 1965, anti-executive legisla¬
tors lashed back with denials of the charges and accusations against
the executive. More significantly, legislators who normally sup¬
ported the executive also bristled against what they considered to
be unjustified attacks. Thus, in taking the battle to the legislature,
the executive discovered that the opposition stiffened and allies
took umbrage.

Finally, if an executive's program is enthusiastically received in
the legislature, it is never long before “rubber stamp" charges are

17 The findings are from an analysis of divisive roll call votes (84 on issues of
concern to the county executive. 1960-1964, and 327 taken at random when he served
as chairman, 1956-1960). Only the votes of the 15 supervisors who served in both
periods are tabulated. The data from the two periods are not completely comparable.
As part of the board, the chairman votes on many measures in which he may not have
strong feelings. As executive, only those issues in which he declared an interest could
be tabulated.

'Congress and the Presidency, op. cit., pp. 103-103.

88 Wisconsin Academy of Science, Arts and Letters [Vol. 56

heard. Typically, the “rubber stamp” charge is an appeal for legis¬
lative pride to assert itself against the challenge of executive
domination.

IV. The legislative committee structure tends to strengthen
specific interests of legislators at the expense of the executive's
general program. Legislatures establish committees to handle their
workload. Legislators generally attempt to serve on committees in
which they have strong personal or constituency interests. There
is a great degree of continuity within the committee system. After
obtaining a preferred committee assignment, a legislator is likely
to retain it throughout his legislative career. This occurs because
of 1) the legislator’s interest in the committee, 2) the investment
he makes acquiring expertise in the committee’s subject matter,
and 3) the importance of seniority within the committee.

Prior to the creation of the county executive office in Milwaukee
County, board committees dabbled in day-to-day administration in
varying degree depending on the composition of the committee,
the administrative departments under its surveillance and so on.
Moreover, it has been traditional to have county board represen¬
tation on some of the boards and commissions which administer
county activities. For example, a supervisor is appointed chairman
of the county welfare board. Supervisors also serve on a number
of other administrative boards. Participation on these administra¬
tive bodies intensifies the legislator’s involvement in his subject
matter interests.

The impact of the board’s committee structure on executive-
legislative relations in Milwaukee County is most clearly seen in
two committees chaired by the chairman of the board : the airport
committee and the civil defense committee. Through his position
on these committees, he made strong efforts to push policies an¬
tithetical to the executive’s general program. The airport committee
took the lead in making the airport division a separate department.
This was strongly opposed by the county executive, who argued
that it was inconsistent with his administrative reorganization
proposals.

Warfare over civil defense in Milwaukee County has raged con¬
tinuously during the past several years, and much of it was related
to executive-legislative conflict. Who should appoint the civil de¬
fense director? Who should be the head of government in the event
of a disaster? In both cases the alternatives presented were the
county executive or the county board chairman. The executive won
these battles, but only after spirited wrangling, made more compli¬
cated because of the board chairman’s control of the civil defense
committee.

1967-68] Hagensick — Executive-Legislative Conflict

89

V. A recurring element of executive-legislative conflict is com¬
petition between each sphere in controlling administration. This
proposition is closely related to the preceding one. All of the ex¬
amples presented there included efforts to control parts of the
county administrative structure through the operations of legisla¬
tive committees. Yet the battle to control the bureaucracy trans¬
cends the committee system. It is a central component in the rela¬
tionship between the legislative and executive establishments.

The issue of controlling administration has two corollaries. The
first is that legislators tend to be suspicious of administrative ex¬
perts, while chief executives are more likely to respect and utilize
them.^^ The legislator therefore may view fragmentation of admin¬
istrative authority as a defense against the exercise of power by
the bureaucratic expert. Chief executives have been the principal
forces behind administrative reorganization efforts. These include
the centralization of agencies, elimination of boards and commis¬
sions and increases in the executive’s own staff.

A second corollary is that fragmentation of administration en¬
hances the “errand boy” role of legislators. Legislators spend a
great deal of time performing services in which they act as inter¬
mediaries between constituents and administrative agencies. The
legislator may feel that he can perform the service function more
effectively within a fragmented bureaucracy, where there will be
more points of access. In a centralized administration he may have
to negotiate for access with a chief executive who is unsympathetic
to the individual demands of the legislator’s constituent. Moreover,
a chief executive may feel that intercession by legislators may be
disruptive intrusions upon efficient administrative procedures.

Competition between the executive and legislature for control
of the Milwaukee County bureaucracy has frequently occurred. On
several occasions legislators have suggested that the county board
or its chairman should make appointments to various county offices
now made by the executive. The board was especially irked by the
executive’s power to choose the supervisors who represent the
board on various county commissions.

Attempts by legislators to make certain county administrative
functions directly accountable to the board have been numerous.
The use of the board’s committee structure to this end has been
cited. The chairman of the airport committee (the board chairman)
has participated extensively in matters relating to airports which
normally are executive responsibilities. For example, a threatened
cutoff in federal airport funds for the county prompted him to write

See David Booth, A Guide to Local Politics, East Lansing (Michigan State Univ. ),
1961, p. 16.

Milwaukee Journal, April 15, 1964.

90 Wisconsin Academy of Science, Arts and Letters [VoL 56

to the Federal Aviation Administration explaining why funds
should not be withheld. Later he and the airport director met with
FA A officials to forestall the cut.'^^ At no point was the county
executive involved.

In another area of administration, the county board passed an
ordinance requesting state legislative authorization of a county de¬
partment of children’s services. Included was the provision that the
department’s director be solely responsible to the board. As inter¬
preted by one commentator, the supervisors would be '‘in the bus¬
iness of administration.”^^

Along with efforts to exert control over county agencies, the
board has refused to enact any of the executive’s administrative
reorganization proposals. In brief, these proposals recommend a
reduction in county agencies from more than 40 to 11. The heads
of the revamped departments would be appointed by the executive
with confirmation by the board. At present, many agency directors
are selected by a board or commission or are covered by merit civil
service provisions rather than subject to appointment by the
executive.

The board has responded to the executive’s requests for profes¬
sional staff assistance grudgingly. For the first four years of the
executive’s tenure, he had just one staff aide in addition to clerical
assistance. A request for a planning analyst in his office was re¬
jected in 1961.^^ Claims that the county executive was “empire
building” marked board response to this request.

These instances are actions of the legislative body to maintain
or increase its control over administration. Most moves by the
executive to strengthen his position in this area have been chal¬
lenged by the board, often because they were interpreted as threats
to its power and influence.

This is not to suggest that the executive has been completely
thwarted in his effort to control the county bureaucracy. Through
his budget-making authority he has had considerable leverage
against administrative agencies. He has used this power effectively
both at budget-review sessions and also through an executive order
directing all county agencies to notify him of new programs that
would affect their budgets.

VI, The presence of a threat to the political jurisdiction from an
outside source tends to reduce executive-legislative conflict. The
notion in American foreign policy that “politics ends at water’s
edge” is applicable also in the realm of executive-legislative rela-

^ Milwaukee Journal, December 16, 1964.

^WTMJ-TV Editorial, March 23, 1965.

^Milwaukee Journal, Feb. 8, 1961. Not until 1964 did the board authorize a second
staff aide.

1967-68] Hagensick— Executive-Legislative Conflict

91

tions. On local levels, of course, the ''outside threat” may be an
adjoining municipality or county, a state or federal agency or even
private economic interests. Any of these may pursue policies at
variance with the self-interest of the governmental unit involved.

In Milwaukee County a number of issues arising from diverse
sources have demonstrated that the legislature and executive are
able to put aside institutional differences at least temporarily to
meet an outside threat. Executive-legislative relations were at low
ebb before the shift of the Milwaukee Braves baseball franchise to
Atlanta was announced in October, 1964. As the county appealed
to baseball leadership for a veto of the shift, the "feud” between
the executive and the board chairman was set aside. They led a
delegation to a hearing before the Baseball Commissioner to plead
Milwaukee's case for retention of its franchise. Subsequently, the
major branches of county government maintained their newly-
formed alliance to investigate what legal remedies could be pursued
to retain or regain major league baseball in Milwaukee.

Conclusion

The creation of a new chief executive office in a political juris¬
diction telescopes into very short time many of the fundamental
issues involved in executive-legislative relations. As such, it ex¬
emplifies the recurring conflicts and problems involved in the
separation of powers. Differing constituencies represented by an
executive elected at-large as against legislators elected on a district
basis produce a contrast between a generalized public interest and
local public interest. When the legislature contains an influential
minority bloc, it will perceive the chief executive as an especial
threat to the minority interest represented.

In addition to differences in conflicting perceptions of individual
or group interests, institutional role playing frequently affects the
relationship between the establishments. This is seen most clearly
as legislative or executive jealousies are aroused or when customary
standards of behavior are ignored. The legislative committee struc¬
ture produces an additional source of conflict with the executive.
Insofar as legislators attain committee assignments because of their
personal or constituency interest in the committee's subject-matter,
it further intensifies executive-legislative conflict.

Competition for control over the jurisdiction's bureaucracy is
another source of potential conflict between a chief executive and a
legislature. The clash between an executive's predilection for cen¬
tralization as against the legislature's acceptance of fragmentation
is seen with distinct clarity in a political unit in which the legisla¬
ture has, in the absence of a chief executive office, become ac-

92 Wisconsin Academy of Science, Arts and Letters [Vol. 56

customed to direct and continuing participation in the administra¬
tive process.

Finally, outside sources may mollify executive-legislative con¬
flict. A clearly perceived threat to the political unit from an outside
source may produce an armistice in the executive-legislative strug¬
gle — at least until the outside threat is removed.

Given the separation of powers between governmental establish¬
ments, friction is perhaps inevitable. This analysis is an effort to
indicate the form it will take and the reasons for its occurrence.
Through additional empirical examination of executive-legislative
relations the propositions may be further tested and assessed.

THE HANDWRITING ON THE LAND

Robert A. McCabe

Wildlife ecology, like any other science, attempts with its tools
to categorize and to predict ; and like other sciences, it is only par¬
tially successful. Predicting behavior of wildlife populations and
their interaction with manufacturing and industrial growth strains
both the limited source material and the limited skill of one fool¬
hardy enough to attempt such an appraisal.

Economists have numerical data with which to extrapolate
growth curves for various aspects of manufacturing and industry.^
To much lesser extent we have crude indices of wildlife population
trends which can be used for prediction — at least short-term. A
study of the interaction between wildlife on one hand and manu¬
facturing on the other enjoys no basic data to draw on. The
relationships are a matter of conjecture but by no means pure
guesswork.

No wildlife population can materially affect a major industry,
but there is a likelihood that major industries do affect wildlife.
This one-way cause-and-effect relationship we will explore. This
is not to imply that the effect is always negative, but it is likely to
be the more common.

In a recent international symposium on Man's Role in Changing
the Face of the Earth, ^ Dr. Paul B, Sears, as chairman of one of

R

the sessions, began by putting on the blackboard the notation p- =

f (C), where R represented resources, environment, or land; P,
human population; and C, culture. This social equation said (p.
423), “The sum total of resources and the population among which
the resources have to be divided are a function of the pattern of
culture.” Unfortunately, the impact of this formula was not pur¬
sued in the reported discussion, and only one other participant in
the symposium questioned it by asking somewhat rhetorically,
“What is culture?”* Apparently assuming the answer had no rele¬
vancy to the basic tenets, the questioner rushed on to even more
abstract involvement.

* From Webster’s unabridged 3rd new international dictionary : The total pattern
of human behavior and its products embodied in thought, speech, action and artifacts
and dependent on man’s capacity for learning and transmitting knowledge to succeed¬
ing generations through the use of tools, language and sytems of abstract thought.

93

94 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1 suggest a closer look at this generalized man-resource formula,
for it appears to be adequate as far as it goes, but needs to go
further. The relationship of available resources and numbers of
of people interacting with them is more than a function of culture.
These interactions have a feedback influence on culture. Indeed
numbers of people and quantity of resources are influenced by
still another major factor, technological advance. This substitu¬

tion

y = C is offered as a better expression of culture and

resource ecology. Where R represents resources; P, human popula¬
tion; T, rate of technological advance; t, time in history; and C,
culture. What this social formula says is that culture is a function
of the rate at which technology allows a given population in time
to exploit its resources.

Thus these resources influence culture when technology makes
them available to the population. An increase in the population
increases the chance for more minds to contribute to technological
breakthrough. Technology (which includes knowledge and learning)
is not and was not acquired evenly in historical time but increased
by fits and starts. This accounts for our use of the exponent t in the
formula and allows us to take stock at various times in history.
An increase in the number of people alone, as in the denominator
of the fraction, does not necessarily mean a greater impact on R,
the resources. In the periods prior to the Civil War, and even World
War I, our expanding population, plus relatively limited technol¬
ogy, did not appear to be making serious inroads on the resources,
but the population in the last two decades with concomitant techno¬
logical advances gives us cause to stop and reflect on whether the
interaction of R and rapidly-growing T fostered by an expanded
P is not creating a negative effect on C.

One needs only to be aware of the comparatively advanced cul¬
tures of the Mayans, Greeks, Romans, Egyptians and Babylonians
to realize that somewhere an imbalance occurred in the resources-
people-technology side of the equation to cause these cultures to
take on negative aspects until C was reduced to a low status or to
zero. Admittedly, it is difficult to recognize when and to what degree
negative values are introduced by man to debase our culture. In
part, the difficulty occurs because we tend to lower standards
instead of confronting the more arduous mastering of deterioration.

Up to a point the greater our technology, the better we are able
to exploit and to increase our resource base, and the better or
greater our culture becomes. Our resources, both available and as
yet unknown, are in a sense finite, and when the values for P and
T increase rapidly, the drain on R reduces its ability to sustain
itself, and the result can only be a lessening of the magnitude of C.

1967“-68] McCabe — The Handwriting on the Land

95

To illustrate: When the number of people demanding a given
manufactured product is low, and the technical know-how in gather¬
ing raw materials or disposing of associated wastes is also low, the
resulting degradation of the resource used, or the resource han¬
dling the waste burden is not seriously impaired. When the demand
increases, technology is stepped up to meet the demands of procure¬
ment and production, but frequently the companion technology
necessary to relieve the environment of the resulting waste burden
is overlooked. Industry's basic charge is to stockholders, not to the
environment, I asked a very good friend of mine who sits on the
board of directors for two manufacturing companies if he knew
of any company that by its own free will, without public pressure,
substantially reduced or eliminated stock dividends to cover costs
which would free an environment of the company's own metabolic
wastes. His answer : 'Tt would not be in the interest of good busi¬
ness. In spite of the fact that each waste-producing industry would
like to do the necessary clean-up, it cannot afford to do so." The
eighteenth century Scottish philosopher David Hume^ considered
avarice the spur of industry. I do not agree fully with this appraisal,
but if avarice be the spur of industry, then let wisdom and con¬
science be its bit and bridle.

In some cases pollution is so acute or extensive that not only is
recreation impaired, but public health and safety are in jeopardy.
The informed citizen must assume the attitude that it is in the
interest of national health, that all polluters, public and private, be
forced into clean-up practices. As stockholders in our future, we
cannot afford to accept less.

I firmly believe that our social equation as of this time indicates
an imbalance in values that is causing C to take on some negative
characteristics. When a resource is so badly abused that the public
ernment steps in (often with subsidy) to remedy and repair. The
mutterings from some onlookers are concerned with ''creeping so¬
cialism." If this is bad, blame “resource disregardism." And, if
there is any doubt as to whether the public is aware of the serious¬
ness of pollution, one needs only to be cognizant of the pending
and recently enacted remedial legislation at both the state and
federal levels.

Before we leave our equation, mention should be made of the
component space. It is the one aspect of our resources that tech¬
nology cannot physically increase. Space use will increase with
population, and whatever is available will likely suffer quality
deterioration.

Moving a polluting industry from one part of the country (space)
to another does not, from a political or economic point of view,
alter the inevitability of degradation in the U. S. A. Temporary eco-

96 Wisconsin Academy of Science, Arts and Letters [Vol. 56

nomic gains mask but do not stay the ultimate loss to an environ¬
ment which will be called upon to support more and more people.

The control of our human population growth (P) could prolong
and perhaps insure the positive values of our culture (C). To exer¬
cise this control through a limited birth rate, a pill to adjust intel¬
lectual attitudes as well as reproductive physiology will be needed.
Birth control implications, however pertinent here, are beyond the
purview of this symposium.

I have said before,^ and it is germane here, that society tends
to default on resource jeopardy when the debasing activity is dis¬
tant in either time or space. We are jarred from a complacent atti¬
tude only when our sight, hearing, sense of smell, and intelligence
are offended by resource misuse or overuse, and are goaded into
action by infringement on our health, economy or recreation. To
act for what is right when it affects our neighbor and not ourselves
is not only a biblical dictate, but evidence of conservation aware¬
ness.

The foregoing exercise in letter symbolism allows us to think
about the people-resource interaction when precise, statistically sig¬
nificant data are lacking, and we are left with the unmeasured
obvious.

Fish and other wildlife and the habitats they occupy are an
important part of our state and national resource base. They are
in an economic sense essential to a rapidly-growing enterprise called
tourism, or outdoor recreation. No major wildlife group is com¬
pletely free from infringement, but not all are affected uniformly.
Without magnifying problems of particular species, the following
groups are involved:

Waterfowl and water birds
Upland game birds
Songbirds
Small mammals
Large mammals

Furbearers

Fish

Amphibians

Reptiles

Invertebrates

These categories of wildlife are affected in four major ways:
loss of cover, loss of food, direct killing, and impairment of pro¬
ductivity. The following are examples of this industry-wildlife
interaction.

Fish. — In the inland aquatic environment which is limited and
to a considerable extent a functional part of our industrial sewer¬
age system, fish suffer loss of protective plant life in waterways,
the death of small animal life on which they feed, are killed directly
by poisons or by lack of oxygen used by oxygen-demanding pollut¬
ants, or are limited in reproducing when spawning habitat is denied
them or is destroyed by siltation. Loss of biological and physical

1967-68] McCabe — The HandwrifAng on the Land

97

aspects of fish environments to industrial pollutants are widespread
and readily documented. They remain to be corrected.

Birds. — In a zealous effort to recoup losses of cellulose fiber ini¬
tiated by the period of ‘'cut out and get out” harvesting of our vir¬
gin forests and completed by the holocausts that followed, we were
oversold on reforestation. Most reforestation of softwoods is nec¬
essary and beneficial. Some of the land that could not be planted
has now grown to other forest trees, making reforestation today
difficult if not impossible on these lands. Forest fire control is virtu¬
ally complete. The bird affected by these efforts is the sharp-tailed
grouse, a bird adapted to forest openings and young forests. In
pristine times fire created the openings and natural reseeding pro¬
duced the young forests. Today, without fire, frost pockets, high
water tables, and plant competition have maintained scattered open¬
ings for sharp-tails. Because these areas are relatively accessible,
they are vulnerable to the mechanical tree planter. To stop the pon¬
derous social, governmental, and industrial complex dedicated to
covering our northland with trees is difficult but not impossible.
There are encouraging signs that industry and some governmental
foresters are recognizing varied uses of forests, particularly the
recreational values of forests, and the sharp-tailed grouse is impor¬
tant to part of such activity. While the foot is on the brake, we can
only hope that the vehicle will stop before it causes the demise of
one of Wisconsin's handsomest native grouse.

Mammals. — The common muskrat is one of the most important
furbearers in the state. Destruction of food and cover causes con¬
tinual shrinkage of habitat and numbers of this animal. Aquatic
habitats, particularly wetlands and marshes, are subject to sani¬
tary land fill (dumps) ; industrial fill for building sites; drainage
for agriculture; cesspools for villages and cities; and graveyards
for automobiles, machinery, and construction debris. One by one
these lowland communities, because they are not understood or
appreciated, are destroyed and with them the muskrat. In some
cases when the desecration is from industrial pollutants, the musk¬
rat is poisoned directly.

What kind of yardstick shall we use to appraise natural losses
against commercial gains? Whatever it is, the results may be the
same, but the sad fact is that at present we are using no yardstick
at all.

A University of Chicago scientist grew flour beetles in a jar
under controlled conditions and found that up to a point (55/per
gm flour) further increase fouled their environment and caused
cannibalization of the very young. Flour beetles in a jar appear to
be a far cry from people and our land, but the ecology is not, John

98 Wisconsin Academy of Science, Arts and Letters [Vol. 56

W. Gardner, the ex-Secretary of Health, Education, and Welfare,
said {Time Magazine, October 1, 1965) :

“We are living in our own filth/’ The article continues: “U. S.
rivers and streams, like the muddy Missouri, used to be contami¬
nated with nothing worse than silt, some salt, and the acids from
mines. Now they are garbage dumps. Raw sewage, scrap paper,
ammonia compounds, toxic chemicals, pesticides, oil and grease
balls as big as a human fist — these are the unsavory contents of
thousands of miles of U. S. waterways.

“Industry now pours at least twice as much organic material
into U. S. streams as the sewage of all the municipalities combined.
Americans who once could be excused a superior attitude about san¬
itation after traveling abroad, now come home to find that their
own drinking water may come from rivers into which steel mills
pour pickling liquors, paper mills disgorge wood fibers that decay
and use up oxygen, and slaughter-houses dump the blood, fat, and
stomach contents of animals. Pollution has become such a problem
that it is all but impossible to calculate the probable cost of clean¬
ing up the streams. A conservative estimate: at least $40 billion
over the next decade.”

Although certain industries are more directly involved in ad¬
versely affecting natural resources, particularly wildlife, my mis¬
sion is not to point an accusing finger. Industry, in defending itself,
calls attention to municipal infractions; pesticide interests chal¬
lenge the adverse effects of agricultural fertilizers ; water polluters
focus attention on air polluters. Each offender advertises recent
improvements in its own back yard. May these improvements catch
up with, neutralize, and then correct the deleterious effects on
human and wild environments.

In our zeal to make life better, easier, more pleasant and more
profitable we are, like flour beetles, destroying these environments
while the gears of technology having writ move on.

Literature Cited

1. Landsberg, Hans H. 1964. Natural resources for U. S. growth. The Johns

Hopkins Press, Baltimore (paperback), 257 pp.

2. Thomas, William L., Jr. (Editor). 1956. Man’s role in changing the face

of the earth. University of Chicago Press, Chicago. 1193 pp.

3. Hume, David. 1898. Essays moral political and literary. Longmans, Green

and Co., London, New York and Bombay. Vol. 3 503 pp.

4. McCabe, Robert A, 1965. A place for battle: the St, Croix. Wisconsin

Academy Review, 12(4):66-68.

5. Park, T. 1948. Experimental studies of interspecies competition. I. Com¬

petition between populations of flour beetles TriboUum confusum Duval
and TriboUum castaneum Herbst. Ecological Monographs, 18: 265-308.
This paper was originally read at the Ninety-Eighth Annual Meeting of
the Academy held at Lawrence University as part of a panel entitled “Wis¬
consin’s Manufacturing and Potential for Industrial Growth and Its Probable
Effects.

RADIOCARBON DATES OF WISCONSIN

Robert F. Black^ and Meyer Rubin^

Introduction

During the last 15 years, radiocarbon dating (Libby, 1961) has
done more to change the correlation and reconstructed chronology
of the Wisconsinan Stage of glaciation in the upper Mississippi
Valley than any other method. The classical chronology, which was
dependent on correlation of events between the Lake Michigan and
Des Moines Lobes, was shown to be inadequate and inaccurate and
was discarded in Illinois in favor of a new radiocarbon-supported
chronology for the Lake Michigan Lobe alone (Frye and Willman,
1960). Discrepancies between the new and classical chronologies
are readily apparent (table 1). The new chronology has been ap¬
plied to Wisconsin (Frye, Willman, and Black, 1965) and to other
places. However, not all events recorded from the Wisconsinan
Stage in states adjoining Wisconsin have been recognized in Wis¬
consin, and vice versa. Furthermore, some of the major events
affecting Wisconsin, according to the variety and distribution of
land forms and deposits left behind, have not yet yielded organic
matter for radiocarbon dating and can only be dated relative to
other events.

This paper lists (table 2) the radiocarbon dates older than 5,000
years from Wisconsin and discusses the significance of some in our
interpretation of the glacial history of the state. Where samples
have been re-dated by better methods, only the latter are given.
Representative dates are shown on a map of Wisconsin (figure 1).
The dates fall into natural groups (table 2) that are correlated
with the new chronology even though some dates may be inter¬
preted in different ways.

It is readily apparent that final answers on all events of the
Wisconsinan Stage in Wisconsin are not yet in hand. However, no

1 Professor of Geology, University of Wisconsin-Madison. Field work leading to this
paper was supported in part by National Science Foundation Grant GP-2820, in part
by the Research Committee of the Graduate School from funds supplied by the Wis¬
consin Alumni Research Foundation, in part by the Wisconsin State Highway Com¬
mission, in part by the National Park Service, Department of the Interior, and in part
by the Wisconsin State Geological and Natural History Survey.

2 Geologist, U. S. Geological Survey, Washington, D. C.

99

100 Wisconsin Academy of Science, Arts and Letters [Vol. 56

(years)

0

5,000

10,000

15,000

20,000

25,000

30,000

65.000 to
70,000

Table 1

NEW CLASSICAL

(Glacial) (Non-Klaclal) (Glacial) (Non-glaclal)

(RECENT)

(RECENT)

Valderan

Woodfordlan

Twocreekan

Valders

Mankato

Cary

Tazewell

Farmdallan

Two Creeks

Rowmanvllle

St. Charles

Gardena

Farm Creek

Altonlan

(Frye and Wlllman, 1960)

Farmdale

///

(Leighton, 1965)

Two contrasting- classifications of the Wisconsinan Stag-e.

Table 2. Some Radiocarbon Dates From Wisconsin=^'

1967-68]

Black and Rubin — Radiocarbon Dates

101

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Table 2. Some Radiocarbon Dates From Wisconsin* — Continued

102 Wisconsin Academy of Science, Arts and Letters [Vol. 56

CO

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Table 2. Some Kadiocarbon Dates From Wisconsin* — Continued

1967-68] Black and Rubin — Radiocarbon Dates

103

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I. Some Radiocarbon Dates From Wisconsin* — Continued

104 Wisconsin Academy of Science, Arts and Letters [Vol. 56

c iS

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L Some Radiocarbon Dates From Wisconsin* — Continued

1967-68]

Black and Ruhin— Radiocarbon Dates

CO
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On a

105

*Excludes younger archeologic dates, those samples well above the bottom of lake deposits, and some solid carbon dates of doubtful
validity Letter prefix denotes laboratory where sample was run: W = U. S. Geological Survey; Y = Yale U. ; M = U. of Michigan; C = U.
of Chicago; L = Lamont; WIS = U of Wisconsin; UCLA = U. of Cal. at L. A.; Gro and GrN = Groningen ; A = Arizona; Tx = U. ot lexas;
and SM = Socony Mobil. See various issues of “Radiocarbon” for futher details on individual samples.

106 Wisconsin Academy of Science, Arts and Letters [Vol. 56

consin and the approximate boundaries between drifts of Valderan, Wood-
fordian and Rockian age.

attempt is made here to review in detail the entire Wisconsinan
chronology for the state nor the events that are not recorded by
radiocarbon dates.

Discussion

Pre-Rockian

Four dates are indefinite but are more than 33,000 radiocarbon
years old (table 2). Two samples (W-1370 and Nuclear Science
and Engineering Laboratory) from the vicinity of Marshfield,
Wood County, are of finely disseminated organic matter in silty-

1967-68]

Black and Rubin — Radiocarbon Dates

107

clay pond deposits (interpretation by Black of samples from power
aug:ering by T. E. Berg) on bedrock and beneath a single drift
sheet that surely is Wisconsinan in age (Hole, 1948), The date of
more than 45,000 radiocarbon years may be interpreted to mean
that the fluctuations of the Altonian ice in Illinois (table 1) (Frye,
Willman, and Black, 1965) were not represented in central Wis¬
consin from the time of existence of the pond to the advance of the
ice that left the overlying till The same interpretation is possible
for the situation in St, Croix County. There basal till with erratic
spruce dated at 29,000 (W-747) and 30,650 (Y-572) radiocarbon
years also seems to have incorporated peat (W-1758) from former
ponds that is dated at greater than 45,000 radiocarbon years. The
wood (W-747 and Y-572) is thought to date the time the ice ad¬
vance destroyed the spruce forest on a residual soil rich in chert;
the older peat (W-1758) now overlies the younger wood (Y-572)
in the till and is thought to represent a pond deposit overrun and
picked up by the ice.

If the different kinds of organic matter were transported by the
ice only once, they would imply that central and west-central Wis¬
consin were ice-free from more than 45,000 radiocarbon years ago
until about 31,000 radiocarbon years ago. Such a situation existed
in Ontario (Dreimanis, Terasmae, and McKenzie, 1966) , but appar¬
ently not in northern Illinois (Kempton, 1963 and 1966). Obviously
other interpretations are possible, and data are not now sufficient
to reconcile them.

Similarly, the spruce and willow fragments (W-1598) from Polk
County are more than 38,000 radiocarbon years old, but they tell us
little about the chronology of the area. The 175-180 feet of drift
overlying the sample is so poorly recorded in the well records that
almost any interpretation is possible.

Rockian

The three dates of 29,000 to 31,800 radiocarbon years from
spruce (W-903, W-901, and W-638) in drift of Walworth and Wau¬
kesha Counties, the two comparable dates of spruce (W-747 and
Y-572) from St. Croix County, and the comparable date of spruce
(GrN-2907) of the paleosol from the base of the loess in Grant
County are believed to represent the time of a brief ice advance,
called Rockian by Black (1960b and 1962), that occurred simul¬
taneously from the Des Moines Lobe on the west and from the
Lake Michigan Lobe on the east (Black, 1964b), This time is
latest Altonian (Frye, Willman, and Black, 1965) (table 1) and
is recorded also outside Wisconsin (White and Totten, 1965). The
wood in St. Croix County is in the basal till which is rich in
disseminated organic matter, clay, and residual chert (Black,

108 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1959a; Black, Hole, Maher, and Freeman, 1965) ; the wood in Wal¬
worth and Waukesha Counties is in oxidized sandy till and in
overridden gravelly outwash. All the wood is erratic and conceiv¬
ably could have been picked up and transported more than once
by ice or water. Hence, other interpretations are possible, but the
deposits can only be younger than the included wood — not older
as had been proposed decades ago (Alden, 1918) .

The dated paleosol (GrN-2907 and Gro-2114) from Grant County
occurs only in a few isolated thin patches which are disrupted and
moved. Spruce (GrN-2907) occurs as small angular fragments of
charcoal (Hogan and Beatty, 1963) and provides a more realistic
date than does the bulk sample (Gro-2114). Clays in the paleosol
are similar mineralogically to those in the residuum from the dolo¬
mite below (Akers, 1961), and general alteration of the mineral
fragments (Hogan and Beatty, 1963) is not severe. The paucity
of paleosol below the loess in southwest Wisconsin, if the area had
never been glaciated, is difficult to explain. That it was glaciated
now seems accepted (Black, 1960a; Frye, Willman, and Black,
1965; Trowbridge, 1966) and disagreement now is concerned more
with timing — Frye and Willman and Trowbridge suggest a Ne¬
braskan time ; Black concludes that Rockian ice from east and west
joined in the center of the state, with relatively thin inactive ice
formed in large part by local accumulation covering the Driftless
Area. Earlier glaciation is also recognized as a strong likelihood.

Positive evidence of glaciation of the Wisconsin Driftless Area
(Frye, Willman, and Black, 1965) comes from some fragments of
Precambrian igneous and metamorphic rocks and particularly
Paleozoic chert and sandstone that rest on younger formations.
Erratics of sedimentary rocks are especially abundant in the cen¬
tral and northern parts of the area (Akers, 1964). Sparse igneous
erratics occur in isolated kame-like deposits south of Taylor in
the northern part of the area and in fresh gravel on the upland
beneath thick loess at Hazel Green, Richland County. Igneous
erratics are also found in terraces tens of feet above the Wiscon¬
sin River. One deposit north of Muscoda contains 10-foot angular
clasts in foreset beds that demonstrate a northeastward flow of
water and a probable ice front. Large sand bodies in the Kickapoo
River valley have come off dolomite uplands and contain glauconite
above any known source. Anomalous rubbles on the upland (Akers,
1964) also have anomalous clay minerals (Akers, 1961). Thus, in
an area of 10,000 square miles in southwest Wisconsin we see an
absence or paucity of chert and clay residuum on bedrock, an ab¬
sence of loess older than 30,000 years, and an almost complete
absence of older paleosols. Moreover, shale with thin seams of un¬
weathered dolomite (Maquoketa Formation) caps East Blue Mound,

1967-68] Black and Rubin — Radiocarbon Dates

109

with only small fragments of the silicified Niagaran dolomite scat¬
tered on the broad flat upland (Black, Hole, Maher, and Freeman,
1965, p. 75-76). This is also an incongruous situation. No grada¬
tional processes other than glaciation seem competent to strip the
Niagaran from so broad an area of shale and remove it and the
chert residuum of the Paleozoic formations from all surrounding
stream valleys.

A pre-Wisconsinan age for some of the peculiar features or
deposits in the area can neither be confirmed nor denied (Akers,
1964; Black, Hole, Maher, and Freeman, 1965; Frye, Willman,
and Black, 1965) but it is suggested (Frye, Willman, and Black,
1965; Palmquist, 1965; Trowbridge, 1966). The supposed front at
Muscoda, relatively thick cherty residuum on the dolomite uplands
near LaValle, Sauk County, red-brown stony drift mostly in Green
and western Rock Counties, and some Windrow deposits (Black,
1964c; Andrews, 1958) still offer the most promise of being early
Altonian or pre-Wisconsinan. No way yet has been found to date
these isolated deposits adequately.

Farmdalian

The time of the Farmdalian deglaciation, which is recorded so
well in Illinois (Frye, Willman, and Black, 1965), is represented
in Wisconsin by one date only (24,800 ± 1,100 years B.P.) at the
base of loess in the Driftless Area. This date is from a bulk or¬
ganic soil sample which differs significantly from the date (29,300
it 700 years B.P.) of fragmented spruce contained in it. No sig¬
nificant breaks in loess deposition from the dated paleosol at the
base to the present surface have been found (Glenn, Jackson, Hole,
and Lee, 1960; Hogan and Beatty, 1963).

Farmdalian time in Wisconsin was at least partly a time of very
cold climates and accompanying permafrost and periglacial phe¬
nomena (Black, 1964a and 1965). However, dating of events is
difficult as no trace of woody material has been found. Presumably,
the thick outwash gravel in southeastern and southern Wisconsin
was formed at this time while ice remained in the northern part of
the state (Black, 1960b). The Farmdalian was a time of ice ad¬
vance in Ontario (Dreimanis, Terasmae, and McKenzie, 1966).

Woodfordian

Woodfordian time is represented in Wisconsin by two dates in
the Driftless Area. One, of caribou bone, is 17,250 radiocarbon
years; the other, a bulk sample of loess (GrN-3624), is 19,250
radiocarbon years. Their significance and relationship to the promi¬
nent Cary (late-Woodfordian) front or the chronology of glacial
events are not known.

110 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Drift of middle and later Woodfordian age makes up the surface
of more of the state than any other, yet it has no known organic
remains. Early Woodfordian deposits are thought to be present
(Black, 1959a) but have not been dated for lack of organic matter.
Isochronous boundaries (Alden, 1918) at the front or within the
Woodfordian drift sheet are exceedingly tenuous. Woodfordian
time in Illinois is represented by tens of moraines and numerous
radiocarbon dates (Frye, Willman, and Black, 1965). Clearly the
Woodfordian in Illinois and Wisconsin is multiple and is composed
of many pulsations of the ice front, some having only limited move¬
ment, but others consisting of retreats or advances up to 100 miles.
The outermost Cary of presumed late-Woodfordian age is not rep¬
resented everywhere in either Wisconsin or Illinois by the same
pulse. Although its border from the Plains to the Atlantic Ocean
has been described and mapped for decades as the break between
deposits of the First and Second Glacial Epochs (Chamberlin,
1878 and 1883), we still have much to learn about it. Without a
single radiocarbon date related to the advances of the Woodfordian
ice in Wisconsin, and few to record its destruction, we have been
dependent on morphology of forms and direction indicators to sep¬
arate pulsations. These are applied with difficulty in many places
but generally seem better than lithology or texture of the material
involved in any one sublobe (Oakes, 1960). Lithology helps to dis¬
tinguish major lobes (Anderson, 1957).

Post-Cary or late Woodfordian events which are pre-Twocreekan
are much less well known in Wisconsin than elsewhere. Moraines
assigned to Mankato and Port Huron in Minnesota and Michigan,
for example, are presumed to be present in Wisconsin, behind the
Cary front. However, the correlation of moraines in Wisconsin with
type localities has not been done, and deployment of such ice in
the state is conjectural.

Deglaciation of the Woodfordian ice in Wisconsin may be time
transgressive, being earlier in the south than in the north. Re¬
vegetation presumably took little time after deglaciation, forest
trees coming in last but perhaps even growing on stagnant buried
glacial ice. A peat mound on Cary drift in Jefferson County has
spruce (WIS-48) at the base dated at 12,800 radiocarbon years
(Ciolkosz, 1965). In Waushara County, a date of 12,800 radio¬
carbon years was obtained on organic matter (UCLA-632) in
marly gyttja four feet above the base of undisturbed marsh de¬
posits (Park, 1964, p. 8), but spruce (UCLA-631) at the base of
the same deposit and higher on the flank of the kettle was dated at
11,600 radiocarbon years. Three other dates on peat (W-820,
W-641, and W-762) in basal pond deposits in Waushara County
are 10,420, 12,000 and 12,220 radiocarbon years respectively. One

1967-68]

Black and Rubin — Radiocarbon Dates

111

(W-1183) in Winnebago County is 12,060 years. The dates of or¬
ganic matter are suggestive of transgression, but the main evidence
for the time transgressive deglaciation is morphologic— that is, the
widespread evidence of ice stagnation and the youthful lakes and
other features in the north. The time difference may be several
thousand years for all buried ice to melt out.

Twocreekan

The Twocreekan interval is named from Two Creeks, Wiscon¬
sin, where a buried soil and organic remains were recognized
in lacustrine deposits along the exposed bluff of Lake Michigan
(Goldthwait, 1907 ; Black, Hole, Maher, and Freeman, 1965) . This
is the most dated interval in Wisconsin, the latest dates yielding
an average of 11,850 radiocarbon years (Broecker and Farrand,
1963). A number of dates (Thwaites and Bertrand, 1957) de¬
rived by the original solid-carbon method were as much as several
thousands of years in error, according to re-runs by better methods.
Many samples (e.g., C-308, C-365, and C-366) dated years ago have
not been re-run.

The general range of Twocreekan time from 11,000 to 12,500
years proposed by Frye and Willman (1960) is distinctly longer
than the interval represented at Two Creeks. There, only an in¬
cipient soil profile was formed under trees of which the oldest by
tree-ring count was only 142 years (Wilson, 1932 and 1936). Sev¬
eral other localities in east-central Wisconsin contain the Two
Creeks horizon in situ, and logs from it are incorporated in the
overlying Valderan till. These also tend to cluster close to 11,850
years ago so the span of Twocreekan time in central and northern
Wisconsin likely is less than in southern Wisconsin. This is to be
expected, because deglaciation through several hundred miles of
latitude of an ice lobe the size of that which occupied the Lake
Michigan area during late Woodfordian time cannot be accom¬
plished overnight.

The sample (WIS-48) dated 12,800 radiocarbon years from Jef¬
ferson County attests to the early development of the spruce forest
in the southern part of the state. Similar dates (samples W-641,
W-762, and UCLA-632) from Waushara County confirm that de¬
glaciation of the Woodfordian ice from those areas, and, hence, the
beginning of Twocreekan time, must have taken place about 12,000
to 13,000 radiocarbon years ago. The carbonate date (UCLA-632)
of 13,700 radiocarbon years is likely too old, according to associated
organic matter that has an age of 12,800 radiocarbon years.

Destruction of the Twocreekan forests by rising lake waters and
by Valderan ice at about 11,850 radiocarbon years ago should
mark the close of Twocreekan time rather than the 11,000 years

112 Wisconsin Academy of Science, Arts and. Letters [Vol. 56

proposed. Probably the entire area of Wisconsin was free of sur¬
face ice during Twocreekan time, and only the northeastern part
was again covered by glaciers. Consequently over most of the state,
the effects of Twocreekan soil formation and geomorphic processes
were merged and obliterated by the same processes that continued
down to the present day in all but rare situations where quick
burial took place. Aggrading stream valleys retain Twocreekan
material (W-1391) (Andrews, 1966), as does the rock shelter un¬
der the Natural Bridge in Sauk County (M-812) (Black, 1959b;
Wittry, 1959). Man was associated with the shelter, leaving re¬
mains of his wood fires (Black and Wittry, 1959; Wittry, 1964).
The climate in northeastern Wisconsin at the time was perhaps
similar to that of today in northern Minnesota (Roy, 1964). Pollen
analysis of Twocreekan material shows spruce forests dominated
(Black, Hole, Maher, and Freeman, 1965; West, 1961).

Valderan

Distribution of the Valderan ice in Wisconsin seems limited to
the northeastern part of the state bordering Lake Michigan (Black,
1966). Whereas the ice was formerly thought to extend across
northern Wisconsin (Leverett, 1929) and to correlate with red
clayey till in eastern Minnesota, this is clearly incorrect (Wright
and Ruhe, 1965). Unfortunately we have no radiocarbon dates in
Wisconsin directly reflecting either the rate of advance or retreat
of the ice. Although the trees at the dated Twocreekan localities
apparently were living when drowned by rising lake waters or
were knocked over by the advancing glacier, the sites are too close
together and the dates are too imprecise to record the date of
advance; we do not yet have any dates in Wisconsin that record
its retreat.

Valderan ice at one time occupied the eastern part of Lake Supe¬
rior and the northern part of Lake Michigan, radiating from a cap
on the peninsula between them (Black, 1966) . Parts of both those
lakes must have had only seasonal ice and open water from the
latter part of Woodfordian time to the present. Other very local
caps on Michigan's Upper Peninsula, as in the Huron and Porcu¬
pine Mountains, may have formed at the same time and survived
after Lake Michigan was entirely freed of ice. That ice seems to
have stopped short of Wisconsin. Buried ice from earlier glacial
advances into northern Wisconsin survived through the Valderan.

Post-Valderan

No radiocarbon dates from Wisconsin record the withdrawal of
the Valderan ice, and we can only infer from evidence elsewhere
that it likely left the state about 10,000 radiocarbon years ago. Only

1967-68] Black and Rubin— Radiocarbon Dates

113

one date (SM“16) (7,650 radiocarbon years B,P.) from the bottom
of a kettle lake in northern Wisconsin is older than 5,000 radio¬
carbon years. It is a minimal date for organic accumulation, but
time must also be allowed for the thaw of the buried ice to produce
the kettle. Drumlins made by Valderan ice have been dropped into
some lakes by post- Valderan thaw of buried ice of Woodfordian
and Rockian age. The ice blocks of different sizes and depth of
burial presumably melted out during a relatively long period of
time-many hundreds to several thousand years. Hence, radio¬
carbon dating of pond sediments can provide minimal dates only
for withdrawal of the Valderan ice.

The Columbia County dates (W-1138 and W-1139) of about 6,000
radiocarbon years record the rapid alluviation of the Wisconsin
River valley south of Portage and are of comparable age to a
paleosol (W-1017) exposed beneath dunes along the shore of Lake
Michigan in Kenosha County, Although that was a time of increas¬
ing temperature and dryness, the altithermal, actual significance
of the dates is not yet known.

References Cited

Akers, Ronald H., 1961, Clay minerals of glacial deposits of west-central
Wisconsin: M.S. thesis, Univ. Wis., 82 p,

Akers, Ronald H., 1964, Unusual surficial deposits in the Driftless Area of
Wisconsin: Ph.D. thesis, Univ. Wis., 169 p,

Alden, W, C., 1918, The Quaternary geology of southeastern Wisconsin: U.S.
Geol, Survey Prof. Paper 106, 356 p,

Anderson, R. C., 1957, Pebble and sand lithology of the major Wisconsin
glacial lobes of the central lowland: Geol. Soc. Amer. Bull. v. 68, p, 1415-
1450.

Andrews, George W., 1958, Windrow formation of upper Mississippi Valley
region — a sedimentary and stratigraphic study: Jour. Geol. v. 66, p. 597-
624.

Andrews, George W., 1966, Late Pleistocene diatoms from the Trempealeau
Valley, Wisconsin: U. S. Geol. Survey Prof. Paper 523-A, 27 p.

Black, Robert F., 1958, Glacial geology Lake Geneva area, southeast Wis¬
consin: Geol, Soc. Amer. Bull. v. 69, p. 1536.

Black, Robert F., 1959a, Friends of the Pleistocene: Sci., v. 130, p. 172-173.
Black, Robert F., 1959b, Geology of Raddatz rockshelter, Sk5, Wisconsin:
Wis. Archeologist, v. 40, p, 69-82.

Black, Robert F., 1960a, ^‘Driftless Area’^ of Wisconsin was glaciated: Geol.
Soc. Amer, Bulk, v. 71, pt, 2, p. 1827.

Black, Robert F., 1960b, Pleistocene history of Wisconsin: Lake Superior
Inst, of Geology, Abstracts, p. 13.

Black, Robert F., 1962, Pleistocene chronology of Wisconsin: Geol. Soc. Amer.
Spec. Paper 68, p. 137.

Black, Robert F., 1964a, Periglacial phenomena of Wisconsin, northcentral
United States: Vlth Intern. Congress on Quaternary, Report v. 4, Peri¬
glacial sect., p. 21-28.

Black, Robert F., 1964b, The physical geography of Wisconsin: Wis. Blue
Book, p. 171-177.

114 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Black, Robert F., 1964c, Potholes and associated gravel of Devils Lake State
Park; Wis. Acad. Sci. Arts and Letters, Trans., v. 53, p, 165-175.

Black, Robert F., 1965, Ice-wedge casts of Wisconsin: Wis. Acad. Sci., Arts
and Letters, Trans., v. 54, p. 187-222.

Black, Robert F., 1966, Valders glaciation in Wisconsin and upper Michigan
A progress report: Mich. Univ., Great Lakes Research Div., Pub. No. 15,
p. 169-175.

Black, Robert F., Francis D. Hole, Louis J. Maher and Joan E. Freeman,
1965, Guidebook for Field Conf. C., Wisconsin: Internal. Assoc. Quater¬
nary Res., 7th Congress, p. 56-81.

Black, Robert F., and Warren L. Wittry, 1959, Pleistocene man in south-
central Wisconsin: Geol. Soc. Amer. Bull., v. 70, p. 1570-1571.

Broecker, Wallace S., and William R. Farrand, 1963, Radiocarbon age of
the Two Creeks Forest Bed, Wisconsin: Geol. Soc. Amer. Bull., v. 74,
p. 795-802.

Chamberlain, T. C., 1878, On the extent and significance of the Wisconsin
kettle moraine: Wis. Acad. Sci., Arts and Letters, Trans., v. 4, p. 210-234.

Chamberlin, T. C., 1883, Terminal moraine of the second glacial epoch: U. S.
Geol. Survey Third Ann. Rep., p. 291-402.

CiOLKOSZ, Edward J., 1965, Peat mounds of southeastern Wisconsin: Soil
Survey Horizons, v. 6, no. 2, p. 15-17.

Dreimanis, a., j. Terasmae, and G. D. McKenzie, 1966, The Port Talbot
Interstade of the Wisconsin glaciation: Canadian Jour. Earth Sciences,
V. 3, p. 305-325.

Frye, John C., and H. B. Willman, 1960, Classification of the Wisconsinan
Stage in the Lake Michigan glacial lobe: Ill. Geol. Survey Circ. 285, 16 p.

Frye, John C., H. B. Willman, and Robert F. Black, 1965, Glacial geology
of Illinois and Wisconsin: in Quaternary of the United States, Princeton
Univ. Press, p. 43-61.

Glenn, R. C., M. L. Jackson, F. D. Hole, and G. B. Lee, 1960, Chemical
weathering of layer silicate clays in loess-derived Tama silt loam of
southwestern Wisconsin: Eighth Nat. Conf. Clays and Clay Minerals,
Pergamon Press, p. 63-83.

Goldthwait, James W., 1907, The abandoned shore lines of eastern Wiscon¬
sin: Wis. Geol. Survey Bull. 17, 134 p.

Hogan, J. D., and M. T. Beatty, 1963, Age and properties of Peorian loes.3
and buried paleosols in southwestern Wisconsin: Soil Sci. Soc. Amer.,
Proc., V. 27, p. 345-350.

Hole, Francis D., 1943, Correlation of the glacial border drift of north-
central Wisconsin: Amer. Jour. Sci., v. 241, p. 498-516.

Kempton, John P., 1963, Subsurface stratigraphy of the Pleistocene deposits
of central northern Illinois: Ill. Geol. Survey Circ. 356, 43 p.

Kempton, John P., 1966, Radiocarbon dates from Altonian and Twocreekan
deposits at Sycamore, Illinois: Ill. Acad. Sci. Trans., v. 59, no. 1, p. 39-42.

Leighton, Morris M., 1965, The stratigraphic succession of Wisconsin loesses
in the upper Mississippi River valley: Jour. Geol. v. 73, p. 323-345.

Leverett, Frank, 1929, Moraines and shore lines of the Lake Superior region:
U. S. Geol. Survey Prof. Paper 154-A, 72 p.

Libby, W. F., 1961, Radiocarbon dating: Sci., v. 133, p. 621-629.

Oakes, Edward L., 1960, The Woodfordian moraines of Rock County, Wis¬
consin: M.S, thesis, Univ. Wis., 61 p.

Palmquist, Robert C., 1965, Geomorphic development of part of the Driftless
Area, southwest Wisconsin: Ph.D. thesis, Univ. Wis., 182 p.

1967-68] Black and Rubin — Radiocarbon Dates

115

Park, Richard A., 1964, Paleoecology of the late-glacial and post-glacial
Luedtke marsh deposit, Waushara County, Wisconsin: M.S. thesis, Univ.
Wis., 46 p.

Roy, Edward C., Jr., 1964, Pleistocene non-marine Mollusca of northeastern
Wisconsin: Sterkiana, no. 15, p. 5-75.

Thwaites, F. T., and Kenneth Bertrand, 1957, Pleistocene geology of the
Door Peninsula, Wisconsin: Geol. Soc. Amer. Bull., v. 68, p. 831-880.
Trowbridge, A. C., 1966, Glacial drift in the “Driftless Area” of northeast
Iowa: Iowa Geol. Survey Rep. Investigations 2, 28 p.

West, R, G., 1961, Late- and postglacial vegetational history in Wisconsin,
particularly changes associated with the Valders readvance: Amer. Jour.
Sci., V. 259, p. 766-783.

White, George W., and Stanley M. Totten, 1965, Wisconsin age of the Titus¬
ville till (formerly called “Inner Illinoian”), northwestern Pennsylvania;
Sci. V. 148, p. 234-235.

Wilson, L. R., 1932, The Two Creeks Forest Bed. Manitowoc County, Wis¬
consin: Wis. Acad. Sci., Arts and Letters, Trans., v. 27, p. 31-46.
Wilson, L. R., 1936, Further fossil studies of the Two Creeks Forest Bed,
Manitowoc County, Wisconsin: Torrey Bot. Club Bulk, v. 66, p. 317-325.
WiTTRY, Warren L., 1959, The Raddatz rockshelter, Sk5, Wisconsin: Wis.
Archeologist, v. 40, p. 33-69.

WiTTRY, Warren L., 1964, Earliest man in the upper Great Lakes area:

Cranbrook Inst. Sci. News Letter, v. 34, no. 3, p. 34-40.

Wright, H. E., Jr., and R. V. Ruhe, 1965, Glaciation of Minnesota and Iowa:
in Quaternary of the United States, Princeton Univ. Press, p. 29-41.

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lit;:

GEOMORPHOLOGY OF DEVILS LAKE AREA, WISCONSIN

Robert F. Black^
Professor of Geology
University of Wisconsin — Madison

Introduction

Devils Lake Park in the Baraboo Range, Sauk County, presently
contains about five square miles of scenic cliffs, wooded hills and
Devils Lake itself (Fig. 1). Topographically the Devils Lake area
is mostly a rolling upland near 1400 feet above sea level, cut by a
steep walled L-shaped gorge whose floor is generally 500 feet be¬
low the summit. The north-trending part of the gorge is occupied
by Devils Lake, held in on the north and on the southeast by end
moraines. Within, but especially adjacent to, the Park many glacial
phenomena are beautifully preserved. Ancient rocks of Cambrian
and Precambrian age crop out locally in bizarre forms. The record
in the rocks and in glacial and periglacial features of the Devils
Lake area is especially rich. The geomorphic development of the
area, resulting in the present landscape, covers many hundreds
of millions of years of geologic time and is truly an intriguing
story.

As a field laboratory in earth history, this area has been one of
the most valuable and fascinating in the upper Mississippi Valley
region. Besides being one of the most popular parks in Wisconsin,
the Devils Lake area has been the locus of field trips for many
hundreds of geology students each year. In spite of the great
amount of study given the area by scientists over the decades, new
information continues to appear.

The last major summaries of the surhcial geology of the Devils
Lake area are those of Salisbury and Atwood (1900), Weidman
(1904), Trowbridge (1917), and Alden (1918). They are now out
of print and found only in the larger libraries. However, Smith
(1931), Martin (1932), Thwaites (1935; 1958), and Powers
(I960), have discussed some of the prominent land forms in the

’ The field work leading- to this paper was supported in part by National Science
Foundation Grant GP-2820, in part by the Research Committee of the Graduate School
of the University of Wisconsin from funds supplied by the Wisconsin Alumni Research
Foundation, in part by the Wisconsin State Hig-hway Commission, in part by the
National Park Service, Department of Interior, and in part by the Wisconsin State
Geological and Natural History Survey.

117

118 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 1. Topographic map of the Devils Lake area, showing border of the
late-Woodfordian (Cary) drift and some proglacial drained lakes. Portions
of the U. S. Geological Survey quadrangle maps Baraboo and North Freedom.
Scale one mile to the inch.

geomorphic evolution of the Baraboo Range. For the benefit of
the many Park visitors this paper outlines the geology of the area,
and then describes in more detail some of the specific features to be i
seen. Devils Lake, the moraines, periglacial features, drained lakes,
stagnant ice features, pot holes, and erratics are singled out. Many
facets of the geologic history are still missing, thus making differ- '
ent interpretations equally viable. Hopefully this summary will
accelerate the search for additional clues.

It is urged that the many striking features illustrated in the
Devils Lake area be seen and appreciated, but not destroyed, by i
the thousands of visitors each year who come to Devils Lake Park.
Pressure of man’s use continues to increase each year, now to the i
point where even the durable rocks need protection. In their zeal,
geology students particularly have contributed disastrously to the i
natural attrition of certain exposures of the bedrock. Every geolo¬
gist who has written extensively on the Devils Lake area has em- i

1967-68] Black — Geomorphology of Devils Lake Area 119

phasized the uniqueness of the glacial, periglacial and bedrock
phenomena present. No other location in the midwest has such a
rich variety of unique features in so small an area near major
centers of population. As a tourist area and as the scientist’s field
laboratory, it is certainly unrivaled for hundreds of miles around.
Hence, every effort must be made to preserve, not just the features
in the Park, but the many glacial and bedrock features adjoining
it as well for future use of all mankind. Once destroyed, they can
not be replaced.

Outline of Geology

The story of Devils Lake Park must begin about a billion years
ago, in middle Precambrian time, with the deposition in shallow
seas of many hundreds of feet of very clean, welLwinnowed quartz
sand of medium-grain size. Subsequent burial in the earth’s outer¬
most crust and accompanying alteration during late Precambrian
time lithified the rounded to subangular sand grains into the brittle
Baraboo Quartzite in which the gap containing Devils Lake has
been cut. The lithification involved little or no crushing of the sand
grains — only deposition of secondary silica cement in interstices.
This makes the total rock very hard yet brittle so it breaks across
grains. The individual grains of sand and some pebbly and bouldery
zones are still easily distinguished today.

Large joint blocks are commonplace and lead to the formation
of extensive talus and jagged cliffs (Fig. 2). The characteristic
pink, red, and lavender hues are attributed to finely disseminated
iron oxides in very small amounts. Oscillation and current ripple
marks, mud cracks, and cross-cutting stratification typical of the
former marine environment are widespread in the Park.

Perhaps in part during metamorphism of the sand to quartzite
and certainly afterwards the area was folded into a large basin or
syncline by a mechanism not fully understood. Perhaps more than
one episode of regional stress produced the minor structures now
visible in the Baraboo Range. The basin is 25 miles long, 10 miles
wide, and trends east-northeast. Devils Lake lies in the center of
the south limb of the basin where the gentle north dips and local
gentle undulations of the quartzite are readily discerned in the
cliffs of the West and East Bluffs overlooking the Lake (Fig. 3).
Local very gentle south dips of the quartzite are found in the cliffs
two to three miles east of Devils Lake.

Fracture cleavage — a parallel splitting of the quartzite easily
confused with bedding — dips northward at Devils Lake at angles
greater than the bedding planes of the quartzite (Weidman, 1904;
Hendrix and Schaiowitz, 1964). It too aids in the formation of
joint blocks, talus, and jagged cliffs. Such fractures are considered

120 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 2. Talus surmounted by castellated cliffs of Baraboo Quartzite on the
south face of East Bluff of Devils Lake.

‘‘normar’ in their orientation with respect to the stresses that are
inferred to have produced the syncline (Hendrix and Schaiowitz,
1964). So are minor drag folds in thin argillites (clayey zones) at
Devils Lake, but other minor structures including small folds,
slip cleavage, and shears are considered ‘"reverse” by Hendrix and
Schaiowitz (1964). The normal minor features are confined to
thin silty argillite layers interbedded with quartzite, whereas the
reverse minor features are in thick argillite beds. Extensive ex¬
posures of the reverse structures are in Skillet Creek, about one
mile northwest of Devils Lake; a small outcrop is just inside the
present northwest entrance to the Park (Hendrix and Schaiowitz,
1964). (Both exposures are rapidly being destroyed by the promis¬
cuous hammering of geology students who do not realize that more

1967-68] Black — Geomorphology of Devils Lake Area 121

Figure 3. View westward of West Bluff of Devils Lake, showing talus and
jointed beds of the Baraboo Quartzite dipping 10° northward.

can be seen on the weathered surface than on a fresh one. This
must cease if we are to preserve these structures. )

The details of the folding mechanism of the quartzite are inter¬
esting but not especially germaine to the problem of the present
day surface features in the Devils Lake area even though the re¬
sulting structures are. As pointed out, the fracturing of the rock
has made frost work especially effective in the formation of pin¬
nacles, talus slopes, and bizarre forms (Figs. 2 to 5).

The younger Seeley Slate Formation and the overlying Freedom
Formation of iron-bearing slate, chert and dolomite of Precambrian
age are on the Baraboo Quartzite southwest of North Freedom
but have not been recognized in the Park. They were the object of
a flourishing subsurface iron ore exploration program in the late

122 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1800’s (Weidman, 1904), but no mines are operating today. They
do not contribute to any notable or striking surface features.

Some time after the folding and uplift of the Baraboo Range,
sub-aerial erosion (Trowbridge, 1917) and probably marine shore
erosion also (Thwaites, 1958) developed relief of a thousand feet
between the top of the Range and the surrounding beveled Pre-
cambrian igneous and metamorphic rocks. Such relief was due
almost entirely to the great resistance to weathering and erosion
of the quartzite.

Igneous rocks crop out in Baxter Hollow (quartz diorite) ; in
three isolated bodies northeast, north and northwest of Denzer
(rhyolite and diorite) ; at the Caledonia Church on Highway 78,
southwest of the east nose (rhyolite) ; and in a larger body at the

Figure 4. Devils Doorway on the south-facing slope of East Blulf of Devils
Lake, formed by periglacial frost action from well- jointed Baraboo Quartzite.

1967-68] Black — Geomorphology of Devils Lake Area

123

Figure 5. Balanced Rock, a joint block of Baraboo Quartzite, isolated by peri-
glacial frost action and rock falls of adjacent material. On the west-facing
slope of East Bluff of Devils Lake.

Lower Narrows of the Baraboo River, on the north side of the
Range (rhyolite) (Weidman, 1904). However, in most of the area
surrounding the Range, the igneous rocks are buried beneath thick
accumulations of sand of Upper Cambrian age (500 to 550 million
years old).

During the development of the relief, beveling of the upland
quartzite obliquely across the bedding produced surfaces which
look smooth to the eye and have long been called peneplains (Trow¬
bridge, 1917). The interpretation that the region was in the end-
stage of one or more cycles of erosion is now discredited (Thwaites,
1958 and 1960). Nonetheless the mode of beveling of the resistant
quartzite at such marked elevations above the surrounding plains
is not truly understood. Certainly toward the close of the erosion
cycle marine waters again inundated the quartzite.

Thick accumulations of sand were piled around the Range which
for a time stood as islands in the shallow seas, shedding their char¬
acteristic pinkish rocks into the surf zone to be transported down-

124 Wisconsin Academy of Science, Arts and Letters [Vol. 56

wind and along shore to inevitable burial (Raasch, 1958). Thus we
find pebbles and boulders of rounded quartzite scattered thickly
and widely through the sands lapping onto the quartzite to the
south but only a short distance to the north. Quartzite pebbles are
found locally from the Cambrian basal conglomerate up to the
Platteville Formation of Ordovician age. The Cambrian sands not
only lapped onto the flanks and filled the center of the syncline, but
they also filled channels cut into the quartzite by ancient streams.
The angular unconformity of the sands with respect to the beveled
quartzite is striking in many places as is also the abrupt textural
and compositional change in the basal conglomerate. A possible
wave cut terrace lies on the northeast part of Happy Hill, six miles
west of Devils Lake (Thwaites, 1958).

Gaps cut entirely through the Range are common in the narrow
steeply dipping north flank. Only one is known, that with Devils
Lake, in the broad south flank (Trowbridge, 1917 and Alden, 1918,
p. 105-107). Some, such as at least part of Devils Lake Gap, are
definitely pre-late Cambrian in age for they contain Cambrian sand¬
stone; others likely are post-Paleozoic and still others were modi¬
fied by streams as young as Pleistocene to the Recent age.

Hanging valleys in the quartzite of the south flank are anomalous
also. They are broad and gently dipping in their upper reaches and
plunge precipitously to the buried Precambrian surface hundreds
of feet below. Some are filled partly with Cambrian sandstone so
date from the Precambrian erosion cycle; some also have narrow
notches cut into them that may postdate the Paleozoic. The dis¬
tribution of the hanging valleys in the Baraboo Range is not known
nor is their origin. Pine Hollow in the southwest corner of the
Park, southwest of Devils Lake, is typical (Thwaites, 1958).

Cambrian sandstone crops out near the northeast and northwest
corners of Devils Lake, in the gorge east of Devils Lake and con¬
tinuing eastward to Parfreys Glen, near Koshawago Springs and
along Messenger Creek southwest of the Lake to the headwaters
of Pine Hollow, and in a considerable area in Skillet Creek. It has
not been found in the deep valley under Devils Lake itself which is
filled with glacial sediments. Cambrian sandstone also is common
along Highway 12 where it crosses the south limb, and continues
westward to Baxter Hollow where it produces striking cliffs.

Paleozoic sediments continued to be deposited around and over
the Baraboo Range probably until Silurian or possibly Devonian
time (Wanenmacher, Twenhofel, and Raasch, 1934) with erosional
intervals such as that below the St. Peter Formation (Thwaites,
1961). However, in the Park only the Upper Cambrian sandstone
lies on the quartzite. The oldest units exposed is the Galesville For
mation of the Dresbach Group (Ostrem, 1967). It is thickbedded

1967-68] Black — Geomorphology of Devils Lake Area 125

and mostly white or very pale yellow. It is the unit that develops
striking cliffs and steep slopes in Baxter Hollow west of the Park.
The next younger formation is the dolomitic, fine-grained Franconia
Sandstone that forms local benches, cliffs, and crags that are green¬
ish grey in contrast to the Dresbach. Still younger rocks are more
distant from the Park today although they may have been present
in the geologic past (Ostrem, 1966). Chert nodules and clay on top
of the quartzite west of Devils Lake are thought to have been “let
down’' during weathering of the dolomitic formations of Ordovician
age (Thwaites, 1958).

Peneplaination of the upper quartzite surface also has been
attributed to the erosion cycles that removed the Paleozoic strata
from the top of the Range. Thwaites (1958 and 1960) discards
those hypotheses in the same way as he discards that for the
Precambrian.

Between the time of deposition of the post-Cambrian strata and
the Pleistocene, or Great Ice Age, geologic events in Devils Lake
Park are obscured. The latter part of that interval, encompassing
at least 300 million years, must have been largely a time for erosion
as no rocks are left behind. If the interpretation is correct that the
upland surface of quartzite is only the recently exhumed Precam¬
brian surface protected during much of the time by a cover of
Paleozoic sediments, then the topography of Devils Lake Park has
changed considerably during the last 550 million years even though
present day topography in the Park may be essentially the same
as it was 550 million years ago. The small amount of Cambrian
sandstone in the present Park does not make striking erosional
features as it does farther west, especially in Baxter Hollow, or
eastward in Parfreys Glen.

It seems clear that at least part of the Devils Lake gorge was cut
by an ancient stream in Precambrian times, otherwise Cambrian
sandstone would not infill it, but perhaps not all was cut then. Some
writers attribute the north part of the gorge to the Paleozoic cycles
of erosion (Thwaites, 1958), and the writer does not believe that
an early Pleistocene time for cutting part of the gorge can yet be
ruled out.

Potholes on the East Bluff are attributed by different people to
the stream work associated with the cutting of the gorge during
the Precambrian, the Paleozoic, the Cretaceous, or the Tertiary
yet they too may only be Pleistocene (Black, 1964), However, at
the east end of the Baraboo Range one pothole in a group of about
25 in the quartzite has Cambrian sandstone firmly adhering to the
inside walls so it was cut indisputedly in late Precambrian or early
Cambrian time. (These were called to my attention by A. C. Trow¬
bridge.) Different kinds of potholes are present at that site, and

126 Wisconsin Academy of Science, Arts and Letters [Vol. 56

all may not be of the same age nor are they necessarily the same
age as those at Devils Lake. Several are altered by glacial ice
of late-Woodfordian (Cary) age, but if all were, the evidence is
obscured by post-glacial weathering.

The pebbly loam with much expandable clay on top of East Bluff
must be the source for the Windrow gravel which Andrews (1958)
considers Cretaceous, but again a Pleistocene age for the deposit
cannot yet be ruled out (Black, 1964). The gravel has been found
in and around the potholes. No way has yet been found to date the
deposits or cutting of potholes satisfactorily. Their place in the
history of events must await new evidence. Regardless of their age,
however, loose blocks with potholes have been moved about on the
upland, and angular quartzite blocks lie on top of the pebbly clay.
Glacial ice must have accomplished this for blocks up to 85 tons
seem to have moved upslope (Black, 1964). The area lies imme¬
diately west of the prominent Cary end moraine on the upland. This
is correlative with the moraines that plug the southeast and north
parts of the Devils Lake gorge. These features are perhaps only
13,000-16,000 years old (Black, Hole, Maher, Freeman, 1965). They
themselves do not prove that earlier ice went no farther into the
Driftless area, and much evidence has now been amassed to indi¬
cate ice did go further west (Black, 1960; Black, Hole, Maher, and
Freeman, 1965; Frye, Willman and Black, 1965).

Much of the talus and the pinnacled cliffs around Devils Lake
(Figs 2 to 5) are associated with the late Wisconsinan Stage of
glaciation (Smith, 1949; Black, 1964; and Black, Hole, Maher, and ,
Freeman, 1965). Whether the area was glaciated more than once
is not proved but is suggested by distribution of erratics and bur- ■
ied organic matter (Weidman, 1904; Alden, 1918, p. 177-178; I
Thwaites, 1958; Black, 1964; and Black, Hole, Maher and Free- I
man, 1965). For example, organic matter from a depth of 130 feet |
in glacial deposits at Baraboo was submitted by F. T. Thwaites to '
Wilson (1936, p. 43) for identification; he found leaves of several ]
dicotyledonous plants, some probably Vaccinium, and one species |
of moss, identified as Campylium stellatum. Thus the story of the |
geomorphic development of Devils Lake Park jumps quickly from
the Paleozoic to the Pleistocene or even late Pleistocene. |

Since glaciation, gravity and frost have moved many large blocks i
of quartzite down slope although the present rate is very slow, i
Man's unsightly activities are now most important. Railroad and j
other construction, and abortive attempts at farming in the last ,,
century have left their mark. Large pits for aggregates have been
opened in glacial materials and in bedrock, and increasing pressure i
from tourists and students is showing. The need for judicious con-

1967-68] Black — Geomorphology of Devils Lake Area

127

trols is painfully obvious and cannot long be withheld if we are to
preserve many of the striking features.

Description of Specific Features
Devils Lake

Probably most tourists are interested in Devils Lake itself (Fig.
6) and spend most of their time in and around it. It is well known
for its trout fishing. The lake is about 1.3 miles long, 0.4 to 0.6
miles wide, and generally 30 to 40 feet deep. A shallow shelf extends
southward from the north shore a distance of about 500 feet; a
narrower shelf surrounds the south end. The east and west sides
drop abruptly into deep water. The water is soft and clear — on
the border between eu trophic and oligo trophic (Twenhofel and
McKelvey, 1939).

The lake has only two small streams entering it — Messenger
Creek on the southwest and the smaller creek from Hells Canyon
on the northeast. The total drainage basin is only about 5.5 square
miles. No streams flow out of Devils Lake. Evaporation and seep¬
age control the losses. The water table is perched at the general

Figure 6. Air view looking southward of Devils Lake, its morainal plug in the
foreground, the quartzite bluffs, and the distant broad flat of the Wisconsin
River at the Sumpter Badger Ordnance Works.

128 Wisconsin Academy of Science, Arts and Letters [Vol. 56

lake level presumably by the fine sediments and organic matter in
the lake basin.

The sediments around the north and south shores are mostly
clean, light-yellow, medium-grain sand with some pebbles of gla¬
cial origin. These become finer and darker as water depth increases.
The bottom of the lake, below about 25 feet of water, is covered
with fine black muds (Twenhofel and McKelvey, 1939) . Near shore
a black soupy liquid or sludge is present up to about three feet
in thickness. It is very rich in aerobic and facultative bacteria.
The sediments below the sludge are black, porous silts and clays
with 15 to 20 percent organic matter. Bacteria are not abundant
below the sludge. Little or no carbonate is present. Most inorganic
matter is silica. Few macroscopic animal remains are present;
microscopic tests and skeletal materials are diatoms and sponge
spicules, and these are not abundant.

The thickness of organic-rich sediment is not known, but is more
than 10 feet. Glacial out wash sand and gravel near the south end
of the lake was penetrated in a well to a depth of 383 feet without
reaching bedrock (Thwaites, 1958).

Moraines

The most important glacial feature in the Devils Lake area is
the end moraine of Cary age (late Woodfordian) depicted on Fig¬
ure 1. This end moraine can be traced with only minor breaks
through the area, in an irregular looping course. It is an extension
of the Johnstown moraine to the south and others traceable along
the entire front of the Green Bay lobe and farther (Alden, 1918).
The description of the moraine in the Park, initially given in detail
by Salisbury and Atwood (1900, p. 93, 94, and 105-111), has stood
the test of time. Because of its length, it will not be quoted here.
The moraine marks the still stand of the outer edge of the ice sheet.

It is only in part synchronous with the more massive till-covered |
outwash and deltaic deposits plugging the valley and enclosing ;
Devils Lake on the north and on the southeast (Thwaites, 1958). j

Terminal moraine plugs, such as occupy the gorge north and j
southeast of Devils Lake, are unusual individually but together com- j
prise a unique situation. Having such a prominent well-defined end
moraine extending for so many miles from those plugs makes the ;
situation even more astounding. The moraine outlined in Figure 1 j
is certainly one of the best to be found anywhere in the world.
Having it so readily accessible to centers of population with so j
many other features nearby makes it especially attractive. As rec- '
ognized by Salisbury and Atwood (1900), the striking loops show ^
clearly the inability of the ice to surmount topographic obstacles
of negligible relief because of restricted flow over and around but- j

1967-68] Black — Geomorphology of Devils Lake Area 129

tresses up ice. Nowhere are similar features so well displayed
amongst so many other phenomena of intriguing historical conno¬
tation.

The uniformity of height (15 to 50 feet) and width (100 to 200
feet) of the moraine on flat surfaces and the asymmetry of the
moraine on hill sides (only 10 to 15 foot abrupt faces on the uphill
side and 50 to 100 foot faces on the downslope side) are in them¬
selves very unusual over such broad distances. Furthermore the
position of the moraine from its high point on Devils Nose south-
westward to the level of the plain records precisely the distal slope
of the ice front during at least the latter part of the deposition
of the moraine. Recording of such gradients is a rare occurrence
almost anywhere because of concurrent or post-glacial destruction
by flowing water and mass movements. Thus, in the area of Devils
Lake are numerous textbook examples of glacial features.

The moraine is but a small part of the end moraine of essentially
similar age that has been traced throughout Wisconsin, from
the Minnesota border near Hudson to the Illinois border south of
Lake Geneva, and also from the Great Plains to the Atlantic Ocean
(Chamberlin, 1883). This moraine was designated the terminal
moraine of the Second Glacial Epoch (Chamberlin, 1878; 1883).
He considered it to be the boundary between older and younger
drift and, as such, to be the most important time break in the
Pleistocene in the state. Field work in Illinois has not supported
this viewpoint (Frye, Willman, and Black, 1965). Unfortunately
we do not have a single radiocarbon date recording the advance of
ice to this end moraine in Wisconsin. From evidence in Minnesota
and Illinois, it likely was formed 13,000 to 16,000 years ago. In
many places outside the Park, the moraine appears more massive
than it is within the Park. Yet its massiveness commonly may be
attributed to bedrock elevations on which it is found or to the over¬
riding and pushing up of material from below (Alden, 1918).

Outside of the two plugs containing Devils Lake, the end moraine
in the Park is generally only 15 to 50 feet high. Locally the front
is fully 80 feet high as at the easternmost loop at Sauk Point (Fig.
1 ) , It is accentuated there because of the high level of the Baraboo
Quartzite on which it is built and the low plain stretching to the
west which was occupied by outwash and a former glacial lake
(Ott Lake, Fig. 1). The more massive moraines containing Devils
Lake rise 90-130 feet above Devils Lake and even higher above the
valley floors north and east. Their massiveness is due mostly to out¬
wash and deltaic deposits (Thwaites, 1958, p. 150) deposited in
front of the advancing ice. Only a thin local veneer of till was de¬
posited directly on these deposits by the ice. The deepest well in
the gorge, 383 feet, did not reach bedrock.

130 Wisconsin Academy of Science, Arts and Letters [Vol. 56

From a car, views of the end moraine are particularly good along
County Highway DL northeast of Devils Lake and at the extreme
northwest corner of the park where Highway 159 crosses the
moraine (Fig. 7). There the abrupt steep slope to the northeast
was formerly occupied by ice which built the small moraine ridge
with its smooth outwash plain to the west or front of the moraine.
These provide a classic example of the relationship of the ice sheet
to its proglacial fluvial and lacustrine deposits. At the easternmost
loop, by Sauk Point, the moraine and its relationship to the quartz¬
ite and proglacial lakes are accessible and readily discernible.
Stratiflcation and texture of outwash dipping westward, unassorted
sandy till, and shear planes inclined steeply up ice to the east are
especially well displayed in the gravel pit shown in Figure 8. Imme¬
diately below the pit is Ott Lake Basin, a former proglacial lake,
and weathered outcrops of the Baraboo Quartzite. Retreatal mo¬
raines are also common behind the outer moraine south of the
gravel pit. The abrupt interlobate junction of ice from the north

Figure 7, Outwash plain in front of the late-Woodfordian (Cary) terminal
moraine as viewed northeastward from Highway 159, about one-half mile east
of Highway 12.

1967-68] Black — Geomorphology of Devils Lake Area

131

and south sides of the Range is clearly portrayed in the moraines
northeast of the pit.

The gravel pit (Fig. 8) at the easternmost loop of the moraine
contains a wide variety of material typical of much of the moraine
and associated outwash. A count of the 6- to 18-inch boulders
shows :

Percent

dolomite 30

gabbro 26

Baraboo Quartzite 16

Cambrian sandstone 12

granite 6

diabase 5

dense intermediate mafic
rock 4

rhyolite porphyry 1

100

The till zones are sandy, brown, yellow-brown, to dark red-brown.
Sandy, bouldery outwash has been overridden locally. Native cop¬
per has been found in the pit and presumably had its source from
Keweenaw Peninsula in Upper Michigan. Ordovician and Silurian
dolomite and oolitic chert of the Ordovician Prairie du Chien Group
are readily identifiable.

For hikers the views of the moraine are particularly good near
the Devils Nose on the South Bluff southeast of Devils Lake, on the
southern part of East Bluff (Fig. 9) extending eastward to the
extreme eastern loop of the moraine at Sauk Point, and also at the
north tip of the north loop. From the north loop one has a striking
view of the Baraboo Valley, the city of Baraboo, and the Lower
Narrows gap of the Baraboo River through the North Range. Views
to the Wisconsin River Valley are superb from the south rim of
the East Bluff at Devils Lake to the vicinity of Parfreys Glen.
Excellent views of the plugs containing Devils Lake may be had
from all of the bluffs rising above them.

Concentric moraines arc around the extreme north end of the
north loop of the Cary end moraine, in section 9, north of Hanson
Marsh (Fig. 1). These show beautifully the building of ridges at
the edge of the ice as it struggled to maintain its position around
that high point. Probably during the initial advance the ice went
over the inside of the loop for erratics are to be found in it. How¬
ever, their presence can also be attributed to water transportation
and even gravity movement from the steep face of the ice that must
have developed there. As the terrain inside the loop is precipitous,

132 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 8. Gravel pit at the east loop of the late-Woodfordian (Cary) end
moraine at Sauk Point, looking northeastward. Stratified outwash dips gently i
to the left; till and drift partly bedded is inclined steeply to the right, reflecting I
ice push and possible shear and flowage as the ice attempted to override its
moraine. i

I

boulders could have bounced and rolled practically across the loop i
on a vegetation-free surface or on an ice-covered surface. At any |
rate the successive arcs are each slightly lower than their prede- !
cessor. The first two are separated by a gap only 60 to 100 feet i
across and 10 to 20 feet deep. The later ones are lower and less ;
regular. The features at the nose of the arc are among the best !
developed anywhere. When coupled with the beautiful views of the =
Baraboo Valley to north and west and of the drained lakes and
other features to the south, this can be considered truly one of the
grand overlooks of the Devils Lake area. j

1967-68] Black — Geomorphology of Devils Lake Area

133

Figure 9. Top of the late-Woodfordian (Cary) end moraine on the East Bluff
of Devils Lake, about one-quarter mile southwest of Hig-hway 113, looking-
northeastward.

For additional details on the moraine, the reader is referred to
the original works of Salisbury and Atwood (1900), Trowbridge
(1917), and of Alden (1918). All emphasize its uniqueness.

Periglacial Features

Periglacial processes are those particularly involving frost action
(especially frost riving in the Baraboo area) and gravity move¬
ments. Within the Baraboo area Smith (1949) lists three groups
of features attributable to periglacial processes: 1) stabilized talus,
2) block concentrations and block-strewn slopes, and 3) choked
valleys and block cascades. Talus occurs in the vicinity of Devils
Lake, and block concentrations, choked valleys, and talus slopes
are west and northwest of Devils Lake and also on the south flank
of the Baraboo Range south of the lake. Pinnacles and monuments
on the cliffs of Devils Lake and wind-polished surfaces north of
the lake are also considered periglacial in origin.

134 Wisconsin Academy of Science, Arts and Letters [Vol. 56

The talus accumulations around Devils Lake are among the most
striking features of the Park (Figs. 2 and 3). They are better dis¬
played there than anywhere else in the Baraboo Range. Other good
locations are in the gorge north of North Freedom (Salisbury and
Atwood, 1900, p. 67), and also along the bluffs of the Lower Nar¬
rows northeast of Baraboo. Talus is best developed on the East,
West and South Bluffs of the lake. Where the Cary ice stood in the
southeast gap it presumably removed much of the talus that appar¬
ently was there before. On the bluffs above the lake the talus is
almost continuous laterally, being interrupted locally by dipping
ledges of the quartzite. It is partly covered by irregular patches of
forest. The talus on the south-facing slope of East Bluff attains
maximum height and continuity of exposure. On the north-facing
slope of the South Bluff the talus is covered largely by trees, and
the slope is slightly less steep.

The talus is composed of heterogenous angular irregular blocks
of quartzite more or less firmly wedged together. The blocks com¬
monly are more than six feet on a side. No marked vertical zoning
of large blocks is apparent. Occasional erratic boulders up to 90
feet above the lake level (Salisbury and Atwood, 1900, p. 133) may
be found in the talus. During the 1930’s the Civilian Conservation
Corps brought in foreign material to surface trails, and the prac¬
tice continues. Erratics of foreign debris should be found in the
talus up to the level of the divide between Messenger and Skillet
Creeks, if the interpretation of that divide as the outlet of glacial
Devils Lake is correct (Trowbridge, 1917).

The maximum height of talus is about 300 feet; the maximum
inclination of the slope is about 36°. The hydrographic map of
Devils Lake (Juday, 1914, map 8) suggests that the talus extends
30 feet out from shore below water level. According to unpublished
data of F. T. Thwaites, the talus may extend to depths of as much
as 285 feet below lake level. However, its subsurface distribution is
not known. Surmounting the talus at many places around the lake
are nearly vertical rock bluffs, some tens of feet high and present¬
ing a jagged appearance.

Many of the talus blocks as well as the rock surfaces and ledges
above them are partly covered with lichens and show some weather¬
ing stains. No clear indications of movement are available. The
vegetation seems stabilized on the slopes. Few blocks are seen on
snow surfaces in winter, and isolated loose blocks in the forest at
the foot of the bluffs are also relatively uncommon. The frost-rived
bluffs and ledges above the talus show many loose blocks and pin¬
nacles (Figs. 3 and 4) apparently in unstable situations, yet few
seem to have collapsed in historical time. The angularity and weight
of the blocks permit them to stand as relatively permanent features.

1967-68] Black — Geomorphology of Devils Lake Area 135

Other signs of inactivity recorded by Smith (1949) indicate that
the formation of talus blocks now is an exceedingly slow process.
How much of the talus originated prior to the advance of the Cary
ice into the north and southeast gorges is not known. If the talus
does extend many tens of feet below the surface of the lake, it
seems likely it has been covered by outwash from the Cary front.
In an abandoned quarry on the northeast side of Devils Lake a
thin veneer of talus is separated from the bedrock by about five
feet of stony soil containing small blacks and rock fragments scat-
tered through an earthy matrix (Smith 1949, p, 202), The contrast
between the talus and underlying material is striking and points,
according to Smith, to a marked change in conditions of weather¬
ing when talus accumulation began.

Smith (1949) did not discuss the effects that high glacial lake
levels might have had on the formation of talus in the gorge. If
Trowbridge (1917) is correct that Devils Lake was up to the level
of the divide between Messenger and Skillet Creeks, then the bulk
of the talus in the area would have been covered by the glacial
lake waters, and the lake level would have been near the base of
the present cliff in many places. Would frost action which is com¬
monly more severe at the water level of a lake have been instru¬
mental in the formation of some of the talus? This is a question
that we cannot yet answer. The lack of erratics of obvious foreign
sources among the talus blocks where they surely were covered by
lake waters is difficult to explain unless the talus has come down
on top of such material to hide it. However, small particles could
have been flushed through the coarse openings of the large talus
blocks.

Around Devils Lake talus is more abundant outside the area
covered by the Cary ice than inside. However, pinnacled slopes and
jagged angular cliffs are just as common along the Baraboo bluffs
to the east of Devils Lake and also in the Lower Narrows of the
Baraboo River to the northeast of Baraboo where the Cary ice
definitely overrode the surface as they are in the cliffs around
Devils Lake. Talus also is abundant in the Lower Narrows. In the
St. Croix Dalles area of western Wisconsin Cary ice clearly went
through the gorge and the pinnacles in cliffs of basalt have de¬
veloped subsequently. Pinnacles may form rapidly. As a conse¬
quence, we can not say specifically when some of the talus or the
pinnacled cliffs of the Devils Lake area were produced. Some of
the material must have been derived in pre-Cary times; some of
the monuments such as Devils Doorway (Fig. 4), Elephant Rock
and Balanced Rock (Fig, 5) possibly were produced during or im¬
mediately after Cary glaciation.

The narrow depressions (Fig. 10) along the bases of many talus

136 Wisconsin Academy of Science, Arts and Letters [Vol. 56

slopes are peculiar. They are elongate, discontinuous channels 15
to 20 feet wide and 5 to 15 feet deep. Thwaites (1935, p. 404; 1958, j
p. 153) attributed them to settling of the finer sediment into inter¬
stices of the talus, but their origin is conjectural. On the hottest
days cold air drains down through the talus into some depressions,
and at times running water may be heard in the talus above them.

The smaller block concentrations and block strewn slopes on the j
south-facing fiank of the Baraboo Range south of Devils Lake and j
the choked valley in the depression southwest of the Lake contain
angular blocks similar to those in the talus slopes of Devils Lake.

The blocks are distributed in elongate bodies. Locally many are
covered with forests, and interstices of the large blocks are filled
with soil.

The locality less than a mile northwest of Devils Lake presents
a problem (NE l^, sec. 14) (Smith, 1949, p. 204). Smith records
shattered blocks and boulders of quartzite, sandstone and conglom¬
erates occurring along a shallow valley and adjoining gentle slopes.
Some of the blocks are almost buried in the soil, but others appear

Figure 10. Elongate depression in drift at the base of talus along the south¬
facing slope of East Bluff, Devils Lake.

1967-68] Black — Geomorphology of Devils Lake Area 137

to be largely above the ground surface. Locally the blocks are
jumbled together. This area is very close to the Cary ice front
where it butted against the northeast corner of West Bluff. Some
glacial drainage went around the end of the bluff and may have
affected this particular area. Smith concluded that this material
was produced in the same way as that of the talus on the south
flank of the Baraboo Range south of Devils Lake. However, the
materials and history of these localities are different. Concentra¬
tion of blocks by running water could not have been important or
the angular blocks would have been more rounded.

A small ridge of Baraboo Quartzite juts above the level of the
south fork of Messenger Creek in the extreme southeast corner of
section 23, southwest of Devils Lake. The relatively flat top of the
ridge reaches an elevation of about 1100 feet, but a large isolated
pinnacle rises fully 20 to 30 feet above the level of the ridge, and
isolated rocks and smaller pinnacles are also present to the north.
The origin of these features is conjectural. They may have been
formed in glacial Devils Lake, assuming it had reached this gen¬
eral level.

Wind work is not common in the Park. A thin accumulation of
loess has been brought in by wind and deposited over the upland
surface. This loess probably is latest Wisconsinan to Recent in age
according to the immaturity of weathering. Such deposition is
common on uplands adjacent to abandoned lake areas or glacial
outwash valleys like those around the Baraboo area. The sources
of the loess could well have been Glacial Lake Baraboo to the north¬
west and the outwash apron in the Wisconsin River Valley to the
south.

Wind-polished and fluted surfaces may be seen outside the Park
on quartzite knobs that rise above the early Paleozoic formations
south of Baraboo. There ventifacted, furrowed surfaces suggest
strong winds blew from the west-northwest. Polishing of the cor¬
ners and faces of some of the upland cliffs of the Baraboo area have
been attributed to wind work, but we cannot exclude water work
and chemical action from such alteration.

Drained Lakes

Proglacial lakes were formed immediately in front of the
moraine in several places in the Park area (Fig, 1). All of these
former lakes (except Devils Lake discussed earlier) have been
drained, but the sediments remain behind as mute testimony of
their former existence. One unnamed lake formerly existed 1,3
miles southeast of Devils Lake on the north side of Devils Nose.
Cary ice butted against the ridge leaving its end moraine which

138 Wisconsin Academy of Science, Arts and Letters [Vol. 56

may be traced around and across the nose (Fig. 1). Where the
moraine crosses the gully in the east half of section 30, it is a sym¬
metrical ridge about 45 feet high and 100 feet wide. It is breached
at the gully, and a smooth plain extends to the southwest. That
plain is underlain with 10 to 30 feet of silty sands and some clay
and gravel. From the moraine down the gully to the north one
sees numerous very large foreign erratics, but from the moraine
up the gully to the west and southwest only the Baraboo quartzite
blocks and small amounts of fine pea-sized foreign gravel are seen.

Similar but larger lakes were present in sections 16, 17 and 18
northeast of Devils Lake (Fig. 1). Peck and Steinke glacial lakes
were named early (Salisbury and Atwood, 1900; Trowbridge,
1917). Glacial Ott Lake is the name given here for convenience to
the easternmost and smallest basin in the Sauk Point Loop. At one
time those basins probably were merged into one lake which would
have drained into glacial Devils Lake via Hells Canyon. As the ice
border retreated somewhat from the end moraine shown on Figure
1, the water level would have dropped and the lakes would have
become separated from each other. Ott Lake in the southeastern
part of section 16 would have been the first to be drained or filled
with outwash. Peck and Steinke Lakes, farther west at lower levels,
remained longer.

Just how long the lakes were able to survive is not known. How¬
ever, in sections 9 and 16, northeast of Devils Lake is the low
swampy area known today as Hanson Marsh. It was a lake that
survived for many centuries (Bachhuber, 1966). Ice at its furthest
extent, at the position shown in Figure 1, covered the area of the
marsh but withdrew almost immediately thereafter to build an end
moraine on the ridge to the west and another to the north, sur¬
rounding Hanson Marsh and forming a lake. Rhythmically-banded
lacustrine sediments at least 25 feet thick were laid down in the
lake along the ice margin. Bachhuber (1966) has counted repre¬
sentative samples of the supposed varves which represent at least
600 years of time. These are only part of that lacustrine sequence.
Similar studies have not been attempted for Steinke, Peck, or Ott
Lakes.

While the ice stood around the old lake at Hanson Marsh, spruce
forests to the west of the area were contributing pollen to the lake
sediments. The pollen sequence throughout the deposits shows
clearly the post-glacial climatic changes as reflected in the local
vegetation. In brief they record a transition from spruce to pine
to mixed hardwoods and other deciduous trees (Bachhuber, 1966).

At its maximum, expanded Devils Lake may have reached an
elevation of 1155 feet, enough to drain the lake to the northwest
down Skillet Creek (Trowbridge, 1917), (Elevations cited by

1967-68] Black — Geomorphology of Devils Lake Area 139

Trowbridge differ from those cited here because of availability of
more accurate maps today.) Thwaites (1960) does not accept any
available evidence that the lake actually overflowed through Skillet
Creek even though Trowbridge (1917) found erratics in Messenger
Creek on the lake side within 16 feet of the Skillet Creek divide.

The Cary moraine at the north end of West Bluff has left its
mark up to 1050 feet in elevation only. How much higher the Cary
ice went is not known, although the writer has found large igneous
and dolomite boulders up to 1160 feet on the northwest side of
that nose (SW 1/4, SE l^, NE l^, sec. 14, T 11 N, R 6 E). Farther
south along Highway 123, cobbles of igneous erratics are found to
elevations of 1210 feet. Thus, this writer would agree with Trow¬
bridge (1917) that Devils Lake overflowed into Skillet Creek.
Furthermore, the cutting of lower Skillet Creek valley in quartzite
must have been accomplished by far more discharge than is avail¬
able from the present drainage area. The monuments and jagged
spurs now present in the gorge are considered to reflect frost
processes like those in the bluffs around Devils Lake itself.

At its maximum advance, over 11 miles of the Cary ice front was
contributing water to the lakes in the Devils Lake area (Trow¬
bridge, 1917, p. 364). His argument is that ice was brought to the
terminous at a rate of 6 inches a week or 26 feet a year. Assuming
ice was melted along the entire front in a zone 100 feet wide by
26 feet deep, we get a measure of the minimum amount of water
that could reach the lakes. Surely some melt water farther back
from the front would also reach the area. Trowbridge concludes
that the annual discharge to the lakes would be at least on the
order of 1.5 billion cubic feet.

The Devils Lake basin itself has a capacity to the discharge level
at Skillet Creek of about 7.5 billion cubic feet. Thus the main lake
basin should have been filled to overflowing in only five years. The
upper lakes would have held a relatively small amount before
draining into Devils Lake. With the ice standing in the area more
than 600 years surely Devils Lake overflowed for considerable time
through Skillet Creek and later possibly past the north end of West
Bluff, at the margin of the ice, even though no features or deposits
there prove this unequivocably.

Trowbridge has supported these rough estimates of water volume
by a check on the amount of material deposited from the glacial
waters. Trowbridge calculated that six miles of the ice front
drained into Steinke Lake, depositing over 2.5 billion cubic feet of
debris. In Peck Basin its 0.5 miles of ice front contributed at least
142 million cubic feet of debris. The Devils Lake gap between the
two morainal dams contains over 2 billion cubic feet of debris.
Thus, it would seem clear that these lakes must have had more than

140 Wisconsin Academy of Science, Arts and Letters [Vol. 56

enough water to drain through the headwaters of Skillet Creek,
the lowest divide available if the Cary ice stood higher than 1155
feet against the north end of West Bluff.

Unfortunately it is not yet possible to date deposition of the
large foreign boulders deep in the soil up to at least 1160 feet ele¬
vation on the northwest side of West Bluff. The dolomite erratics
are very little etched; gabbro and other coarse textured boulders
are not disaggregated so it is assumed they were left by ice im¬
mediately preceding the building of the prominent end moraine.
A large fresh gravel kame (SE SEi/4, sec. 25, T 12 N, R 5 E)
is 3.5 miles west of the same front at Baraboo, and a deep kettle
hole (SW %, NE sec. 15, T 10 N, R 6 E) is one mile beyond
the front. They attest to an advance of the ice beyond the promi¬
nent end moraine. Thwaites (1958) and earlier writers, except
Weidman (1904), do not accept these features as resulting from
direct glacial action. However, about 40,000 cubic yards of foreign
erratics from gravel to boulders have been removed from the kame
which shows typical cross stratification and irregular sorting — ^far
too much material to be ice rafted and deposited with such internal
fabric. Till is present below the kettle which is as perfectly de¬
veloped as any.

In the Steinke Lake sediments Salisbury and Atwood (1900,
p. 120 and plate 28, p. 108) noticed laminated silts and clays in
which marked deformation of certain horizons were present. Lo¬
cally more than 60 feet of these deposits were excavated, but the
exposures are now covered. Salisbury and Atwood (1900, p. 134)
outline the history of that lake briefly. Because the basin is en¬
closed to the south, east, and west by quartzite, it was in a logical
position to receive and hold water. The first lake formed against
the ice to the north had no outlet until the water rose to the level
of the lowest divide on the southwest side where the water over¬
flowed to the west and northwest into Devils Lake via Hells Can¬
yon. Sediments borne into the basin by the glacial drainage were
deposited as deltas and outwash in the lake. The coarser particles
were left near the ice; the finer ones were carried farther away.
Continued melt water from the ice front brought more and more
sediment into the lake until its delta front extended completely
across the lake, filling it to the level of the outlet. Later drainage
followed the retreating edge of the ice directly westward into
Devils Lake.

Other drainage modifications in the area accompanied and fol¬
lowed the lakes. An example is that of Skillet Creek, the small
tributary to the Baraboo River, which flows northwesterly from
the southwest divide of Devils Lake, Before glaciation of the dis¬
trict, Skillet Creek probably flowed in a general northeasterly di-

1967-68] Black — Geomorphology of Devils Lake Area 141

rection to the Baraboo River (Salisbury and Atwood, 1900, p. 138).
The Cary ice blocked the stream forcing it to seek a new course.
The only course open was to the north and west in front of the
advancing ice. Drainage from the ice, depositing glacial-fluvial
and then glacial-lacustrine materials, forced the stream farther
westward until finally it reached its position across the Cambrian
sandstone plain well to the north and west of its former route.
In that position ancestral Skillet Creek began to downcut after
deglaciation and drainage of glacial lake Baraboo that inundated
up to 980 feet elevation the lowland where Baraboo is now located.
It superimposed itself on the bedrock and cut a new gorge. Such
superposition could have occurred only after the cessation of over¬
flow of water into Skillet Creek from the glacial lake occupying
the gorge of Devils Lake. To drain Lake Baraboo it was necessary
to clear the ice from the east nose of the Baraboo Range proper,
near Portage (Bretz, 1950). The position of the lower part of Skil¬
let Creek well on the westward flank of the outwash apron of Cary
age can be attributed to the initial topography left during the
draining of Lake Baraboo.

Stagnant Ice Features

The retreat of the ice from Sauk Point at the crest of the Bara¬
boo Range was by melting in situ for it left behind typical ice
stagnation features with knob and swale topography. Many knobs
are small kames of poorly sorted but water worked sand and
gravel; the depressions are almost invariably kettles produced by
the melting out of buried ice blocks in the debris. The stagnant
buried ice area formed at the junction of an advancing lobe from
the north and another from the south — a kettle interlobate moraine
of very small size when compared to the Kettle Interlobate Moraine
of eastern Wisconsin. Yet, its origin would have been similar. Sec¬
tion 15 at Sauk Point (Fig, 1) contains the better features of this
ice stagnation interlobate area. Relief is generally only 10 to 30
feet between the knobs and adjacent kettles. It is readily viewed
from the east-west highway extension of County Highway DL.

Behind the end moraine as mapped through the area of Figure
1, numerous ice stagnation features may be seen. These are par¬
ticularly well-developed on the flanks of the Baraboo Range to the
north toward the city of Baraboo and also to the south and east
toward the Wisconsin River. Many knobs are kames ; many swales
are kettles. Such ice stagnation features on the steeper slopes of
the Baraboo Range are generally nowhere as well developed or as
large as those of the lowlands. It is in the lowlands that the larger
ice blocks were buried more readily.

142 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Potholes

Black (1964) has described potholes on the East Bluff of Devils
Lake overlooking- the late-Woodfordian (Cary) moraine which
plugs the southeast gorge. The potholes are carved in bedding
plane surfaces of the Baraboo Quartzite in situ and also in loose
blocks of the quartzite that are scattered irregularly on the beveled
upland surface. Polished chert-rich gravel of the Windrow Forma¬
tion is associated with some potholes and has been found in them
(Salisbury, 1895, p. 657).

On July 29, 1964, after the pothole paper was submitted for
publication. Black used a power auger to drill 12 holes through
the quartzite rubble scattered over the higher part of East Bluff.
Most holes were less than five feet deep; the deeper holes pene¬
trated only six to eight feet. All encountered a silty clay zone with
5 to 10 percent well-rounded and polished pebbles of the Windrow
type scattered uniformly throughout. The zone is reported to be
16 feet thick in a dug well (Salisbury, 1895) near the junction of
the trails at the apex of the bluff. The clay is mostly of the ex¬
pandable type — swelling greatly with a sodium-rich water soften¬
ing agent. Such clay is not common in Tertiary or Mesozoic de¬
posits in the upper Mississippi River area which have kaolin — a
non expanding clay.

Through the years most writers have attributed the potholes and
associated gravels to streams of Paleozoic, Cretaceous, or Tertiary
age that flowed across a continuous upland surface at and above
the level of the rim. The writer thought no one seriously had con¬
sidered them to be glacial prior to publication of his paper (Black,
1964), but subsequently he found that Powers (1960) raised the
question without attempting to answer it. We know now that at
least one of the potholes at the extreme east nose of the Baraboo
Range, which contains Cambrian sandstone firmly adhering to its
walls, must have been produced in late Precambrian or earliest
Cambrian times. Not all of the other potholes of that locality can
be ascribed necessarily to the same time of formation even though
it would be most logical. By analogy it would be logical also to
suspect that the potholes on the East Bluff of Devils Lake were
produced at the same time, but this does not prove it.

Regardless of when the potholes were formed, it is clear that
the loose blocks in which we find potholes have been moved subse¬
quent to grinding of the potholes. Some blocks have been split, and
one side or bottom of its pothole is now gone. Others are turned
on their sides or are upside down. These are scattered along with
other loose blocks of the Baraboo Quartzite over the chert-rich,
gravelly clay (Windrow Formation) on the upland. The splitting
off of the blocks from bedrock and movement of the loose blocks

1967-68] Black — Geomorphology of Devils Lake Area 143

to their present location can most easily be explained on the basis
of movement by glacial ice or possibly by melt waters associated
with ice. The hundreds of blocks of Baraboo Quartzite on top of
the Windrow Formation cannot be explained by simple down
weathering in place, and no quartzite nearby is higher. Such blocks
of the quartzite on top of the Windrow Formation must be consid¬
ered true glacial erratics.

The large erratic to the north of the pothole area is so described
by Black (1964). It weighs 85 tons and must have been moved
upslope to its present resting place. This surely could only be ac¬
complished by ice. Other smaller but impressive quartzite erratics
may be seen on the South and West Bluffs of Devils Lake as well
(Fig. 11). No mechanism of erosion of the smoothly beveled up-

Figure 11, “Erratic” of Baraboo Quartzite on the highest part of the South
Bluff of Devils Lake.

144 Wisconsin Academy of Science^ Arts and Letters [Vol. 56

land surface, known to the writer, can leave behind such large
loose blocks to rise above the general level.

These various phenomena would imply that glaciation of the
Devils Lake area had occurred some time prior to the late-
Woodfordian (Cary) advance. This seems certain from a variety of
evidence that cannot be detailed here. However, for example, dolo¬
mite and igneous rock erratics are found 100 feet above the Cary
moraine at the north end of West Bluff. Moreover, a kame deposit
3.5 miles west of the front near Baraboo has more than 40,000
cubic yards of gravel, and a deep kettle with till lies one mile west
of the front at the Badger Ordinance Works south of the Park.
They attest to nearby extensions of glacier ice beyond the end
moraine of the Cary as recorded on Figure 1. The freshness of
dolomite and igneous erratics, the lack of erosion and filling of the
kettle, the amazing freshness of the igneous outcrops near Denzer,
the youthfulness of the loess on the upland, and other criteria
would suggest that the time of such glaciation did not long pre¬
cede that of the Cary. Unfortunately, this is a very perplexing
problem for which we have relatively little information to go on,
and it cannot be discussed further here.

Erratics

For convenience, erratics within the area of Devils Lake Park
may be classified into two groups. One contains those rocks, such
as igneous and highly metamorphosed materials, that could have
originated only from a point far to the north. The other contains
those rocks of local derivation which are in anomalous situations.
This section is concerned largely with the second group — the large
mass of debris brought in by the Cary ice and dumped inside the
end moraine is clearly of glacial origin. At Devils Lake erratics
have been washed out from the terminal area of the ice that
blocked the north and southeast gaps. Erratics have been carried
by drifting ice at least 90 feet above the present lake level (Salis¬
bury and Atwood, 1900, pp. 133). Trowbridge (1917, p. 366) in
one hour found 103 erratic boulders in the valley of the north fork
of Messenger Creek and one diabase cobble on the west slope of
the divide in the drainage of Skillet Creek. He found igneous rock
erratics 164 feet (202 feet in his paper reflects use of now out¬
dated topographic maps) above the present level of Devils Lake
and only 28 feet below the divide. Other glacial cobbles occurred
within 16 vertical feet of the divide. Thus the origin of erratics
behind the end moraine and those carried out from the terminous
by outwash waters and floating icebergs in the proglacial lakes
are readily explained. These are recognized easily because of their
obvious foreign source.

1967-68] Black — Geomorphology of Devils Lake Area 145

In the second group of rocks, however, we find various local
materials which are distributed in the area in such a way that it
is far more difficult to prove that they obtained their present loca¬
tions on the basis of glacial ice directly. In this group are placed
the large Baraboo Quartzite erratic blocks and fragments which
occur on East Bluff on top of the Windrow Formation and also
those which occur on the South and West Bluffs and on Sauk Hill
on the Baraboo Quartzite itself. To this group is added also the
Paleozoic cherts which lie outside the end moraine. These categories
require additional comment.

It is difficult not to accept as glacial erratics the angular quart¬
zite rubble on top of the Windrow Formation on East Bluff. If one
accepts the 85-ton Baraboo Quartzite block near the block fields
north of the pothole area on the East Bluff as a glacial erratic, then
it would seem to the writer that we must also accept similar large
angular blocks of the Baraboo Quartzite on the South (Figure 11)
and West Bluffs and on Sauk Hill as well. They lie on rounded rela¬
tively smooth upland surfaces, protruding through the loess cap
which is a few inches to two or three feet thick. These blocks are
loose and rest directly on the quartzite. Hence, they have not
attracted attention by previous workers in the area. However, no
process of planation by sea or streams could leave such large
angular fresh blocks behind to rise above the smoothly planed
surfaces. At least it seems unusual to this writer to see such large
angular blocks rising above the general level of a truncated sur¬
face that is supposed to be exhumed from beneath hundreds of
feet of Paleozoic sandstones and dolomites. These are the highest
surfaces in the area. The blocks can not have been let down from
a higher cover or plucked out of the upland by any means other
than glacial ice. To the writer it is far easier to explain such loose
blocks as having been brought in some time after the exhumation
of the upper surfaces. The logical time to do this is during the
Pleistocene, by glacial action. Many blocks are angular with very
sharp corners; relatively little pitting has taken place, and frost
riving is minimal. A Wisconsinan age for them would seem most
logical, yet an earlier Pleistocene age is possible.

Associated with the erratic blocks of Baraboo Quartzite on the
South Bluff are distinct channels in the upland surface which are
also peculiar. One due south of the lake crosses through the crest
of the range and has steep overhanging banks 10 to 15 feet high
(Fig. 12). Corners of the blocks are very sharp, A few blocks pre¬
sumably derived by frost action lie at the foot of the bank, but
hundreds of cubic yards of material have been removed from the
largest channel. No accumulation of such debris is seen either to
the north or to the south. Where has it gone? Are such features

146 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 12. Overhanging wall of Baraboo Quartzite on top of the South Bluff
of Devils Lake, looking northward. Part of a channel possibly cut during the
Pleistocene.

related to the Paleozoic or Mesozoic erosion cycles that have affected
the area, or is this again something that may be attributable to a
pre-Cary glacial event? Did glacial water flow across the upland
which is higher than the divide at the head of Pine Hollow? Pine
Hollow has some rare foreign rocks such as schist and rounded
Windrow-type pebbles among the angular quartzite, sandstone,
sandy dolomite, and chert. Glacial water may have aided in cutting
it. However, we have really no basis for saying one way or the
other except for the relative freshness of the edges and faces of
the Baraboo Quartzite exposed in these peculiar features. We have
inserited at least one late-preCambrian or early Cambrian pothole,
but it is a very small feature obviously protected by the Cambrian

1967-68] Black — Geomorphology of Devils Lake Area

147

sandstone» No sandstone was seen anywhere in association with
the loose angular blocks of the Baraboo quartzite on the upland
or with the sharp channels. Hopefully, more detailed subsurface
study will provide additional clues to the perplexing origin of these
features.

The chert erratics present another puzzling situation. Chert be¬
hind the end moraine of Cary age clearly can be explained as having
been brought in by ice. It has been customary to explain chert,
locally identifiable as Ordovician— Silurian in age to the west of
the Cary terminous, as having been “let down’^ during the weather¬
ing and removal of the Paleozoic formations that once overlay the
Baraboo Quartzite (Thwaites, 1958 and 1960). However, the abun¬
dance of chert of Silurian age is puzzling. One would expect that
the younger formations which would be removed first in the
Paleozoic-Mesozoic-Tertiary weathering cycles would be essentially
absent from the upland in contrast with chert of the underlying
older formations. Detailed statistical sampling has not been done,
but yet we find considerable Silurian chert. This seems incongruous
because there is no difference in size nor in weatherability of the
Ordovician-Silurian chert. Is it possible that the chert has not been
simply let down but has actually been brought in by ice of an earlier
glaciation that did not have abundant igneous materials in the ice?
Again we have no basis for discussion, of such a problem, because
the evidence is still too meager to constrain our thinking,

Eeferences Cited

Alden, William C., 1918, The Quaternary geology of southeastern Wisconsin:

U. S. GeoL Survey Prof, Paper 106, 356 p.

Andrews, George W., 1958, Windrow formation of upper Mississippi Valley
region — a sedimentary and stratigraphic study: Jour. GeoL, v, 66, p, 597-
624.

Bachhuber, Frederick W., 1966, Pollen analysis from Hansen Marsh — An
upland 'site, southcentral Wisconsin: M. S. thesis, Univ. Wis., 30 p.
Black, Robert F., 1960, ‘‘Driftless area^’ of Wisconsin was glaciated: GeoL
Soc. Amer. Bull. v. 71, pt. 2, p, 1827.

Black, Robert F., 1964, Potholes and associated gravel of Devils Lake State
Park: Wis, Acad. Sci., Arts and Letters Trans., v, 53, p. 165-175.

Black, Robert F., Francis D, Hole, Louis J. Maher, and Joan E. Freeman,
1965, Guidebook for Field Conference C, Wisconsin: Intern. Assoc. Qua¬
ternary Res., 7th Congress, p. 56-81.

Bretz, J. Harlen, 1950, Glacial Lake Merrimac: Ill. Acad. Sci. Trans, v. 43,

p. 132-136.

Chamberlin, T. C., 1878, On the extent and significance of the Wisconsin
kettle moraine: Wis, Acad. Sci., Arts and Letters Trans., v. 4, p. 201-234.
Chamberlin, T. C., 1883, Terminal moraine of the second glacial epoch: U. S.

GeoL Survey Third Ann. Rept., p, 291-402.

Frye, John C., H. B, Willman, and Robert F. Black, 1965, Glacial geology
of Illinois and Wisconsin: in Quaternary of the United States, Princeton
Univ. Press, p. 43-61,

148 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Hendrix, T. E., and M. Schaiowitz, 1964, Gravitational structures in the
Baraboo Quartzite near Baraboo, Wisconsin: Geol. Soc. Amer. Bull.,
V. 76, p. 1045-1050.

JUDAY, Chancey, 1914. The inland lakes of Wisconsin: Wis. Geol. and Nat.
Hist. Survey Bull. 27, 137 p.

Martin, Lawrence, 1932, The physical geography of Wisconsin: Wis. Geol.

and Nat. Hist. Survey Bull. 36, 608 p. Reprinted, 1965, Univ. of Wis. Press.
Ostrem, M. E., 1966, Cambrian stratigraphy in western Wisconsin: Wis. Geol.
and Nat. Hist. Survey Information Circ., No. 7, Guidebook for Ann. Field
Conf. of Mich. Basin Geol. Soc., 79 p.

Ostrem, M. E., 1967, Paleozoic stratigraphic nomenclature for Wisconsin: Wis.

Geol. and Nat. Hist. Survey Information Circ., No. 8.

Powers, William E., 1960, Physiography of northern Illinois and southern
Wisconsin: Ill. Geog. Soc., Bull. (Dec.), p. 8-32.

Raasch, G. 0., 1958, Baraboo monadnock and palaeo-wind direction: Jour.

Alberta Soc. Petroleum Geologists, v. 6, p. 183-187.

Salisbury, R. D., 1895, Pre-glacial gravels on the quartzite range near Bara¬
boo, Wisconsin: Jour. Geol., v. 3, p. 655-667.

Salisbury, R. D., and W. W. Atwood, 1900, The geography of the region about
Devil’s Lake and Dalles of the Wisconsin: Wis. Geol. and Nat. Hist. Sur¬
vey Bull. 5, 151. p.

Smith, Guy-Harold, 1931, Physiography of Baraboo Range of Wisconsin:

Pan-Amer. Geologist, v. 56, p. 123-140.

Smith, H. T. U., 1949, Periglacial features in the Driftless Area of sourthern
Wisconsin: Jour. Geol., v. 57, p. 196-215.

Thwaites, F. T., 1935, Physiography of the Baraboo district, Wisconsin:

Kansas Geol. Soc. Guidebook, 9th Ann. Field Conf., p. 395-404.
Thwaites, F. T., 1958, Land forms of the Baraboo district, Wisconsin: Wis.

Acad. Sci., Arts and Letters Trans., v. 47, p. 137-159.

Thwaites, F. T., 1960, Evidences of dissected erosion surfaces in the Driftless
Area: Wis. Acad. Sci., Arts and Letters Trans., v. 49, p. 17-49.
Thwaites, F. T., 1961, the base of the St. Peter sandstone in southwestern
Wisonsin: Wis. Acad. Sci. Arts and Letters Trans., v. 50, p. 203-219.
Trowbridge, Arthur C., 1917, The history of Devils Lake, Wisconsin: Jour.
Geol., V. 25, p. 344-372.

Twenhofel, William H., and Vincent E. McKelvey, 1939, The sediments of
Devils Lake, a eutrophic-oligo trophic lake of southern Wisconsin: Jour.
Sed. Petrology, v. 9, p. 105-121.

Wanenmacher, J. M., W. H. Twenhofel and G. 0. Raasch, 1934, The
Paleozoic strata of the Baraboo area, Wisconsin: Am. Jour. Sci., v. 228,
p. 1-30.

Weidman, Samuel, 1904, The Baraboo iron-bearing district of Wisconsin:

Wis. Geol. and Nat. Hist. Survey Bull. 13, 190 p.

Wilson, L. R., 1936, Further Fossil Studies of the Two Creeks Forest Bed,
Manitowoc County, Wisconsin: Torrey Bot. Club Bull., v. 66, p. 317-325.

EVIDENCE FOR FAULT ZONES IN THE
BEDROCK OF MILWAUKEE COUNTY

Carl A. R. Distelhorst and A. G. Milnes

Because of the irregular cover of Pleistocene deposits, little is
known about the overall geologic structure of the bedrock in Mil¬
waukee County, Recently, however, a compilation has been made
of the geologic information obtained from the innumerable bore¬
holes located in the county (Distelhorst 1967) and on this basis a
more detailed picture of the bedrock structure can be constructed.

Those boreholes which penetrated sufficiently deeply intersected
a prominent and unmistakable contact, that between the Niagran
Dolomite (Silurian) and the underlying Maquoketa Shale (Ordo¬
vician) . This contact has been logged and recorded in many holes
throughout the county and thus provides a convenient horizon for
determining the bedrock structure. At each borehole, the height
above sea level of this contact was calculated from the information
compiled from the well log records of the Wisconsin Geological and
Natural History Survey. These heights were placed in height
groups each spanning 100 feet (see Fig. 1) . The symbols represent¬
ing the various height groups form definite broad bands across the
county map, and enable rough stratum contours at 100 foot inter¬
vals to be constructed.

The most striking feature of the distribution of height group
symbols on Fig. 1 is a line of discontinuity which runs southwest-
wards from the mouth of Milwaukee River. South of this line the
stratum contours seem to have been displaced towards the east.
Closer inspection reveals that in a number of the boreholes situated
close to this line, the Silurian-Ordovician boundary shows an
anomalous position, either much higher or much lower than that
to be expected from neighboring wells (Fig. 1). This line probably
represents a fault zone in which the strata have become much dis¬
turbed. The downthrown block is to the northeast, but on the
present data the true direction of movement and hence the amount
of movement this zone represents cannot be determined.

The same procedure has been carried out for the Devonian-
Silurian boundary, another easily logged contact but one only found
in the northeast corner of the county (Fig. 2). Again there is sug¬
gestion of displacement of stratum contours (though the evidence

149

150 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1967-68] Distilhorst and Milnes — Fault Zones in Bedrock

151

Figure 2. Structure contour map of the Devonian-Silurian boundary in north¬
ern Milwaukee County, Wisconsin. Ornament as in Fig, 1. Height group sym¬
bols: the Devonian-Silurian boundary lies between 300 and 400 feet (symbol
A), 400 and 500 feet (symbol B), 500 and 600 feet (symbol C), or above 600
feet (symbol D), above mean sea level.

is much more tenuous) and a succession of anomalous readings,
along a narrow zone running eastnortheast-westsouthwest. Anom¬
alous heights are also obtained for the Silurian-Ordovician bound¬
aries in other wells along the same line (two of the three most
northerly wells shown on Fig. 1 are lower than the stratum contours
to the south would indicate). Another fault zone is thus postulated
in the extreme north of the county, in this case with the down-
thrown block to the south.

The northernmost of these two fault zones is probably a con¬
tinuation of the north-east striking fault known to exist under
the town of Waukesha to the west (Foley et al., 1953). The south¬
ern fault zone was previously thought of as a fold structure (Foley,
op. cit.) but this was on the basis of structure contours drawn on
the top of the St. Peter Sandstone, a much less well defined horizon

152 Wisconsin Academy of Science, Arts and Letters [Vol. 56

than the one used here. There is, however, good indication of a
slight monoclinal warping in both the Devonian-Silurian and
Silurian-Ordovician contacts. In both Fig. 1 and Fig. 2, two of the
stratum contours lie closer together than the others.

References

Distelhorst, C. a. R., (1967) : Bedrock Formations of Milwaukee County.

M.S. Thesis, University of Wisconsin-Milwaukee.

Foley, F. C., W. C. Walton and W. J. Drescher (1953) : Ground-Water
conditions in the Milwaukee-Waukesha Area, Wisconsin. U.S. GeoL Surv.
Water Supply Paper 1229, 96 pp.

THE DISTRIBUTION OF IRON IN LAKE SEDIMENTS

Jerome O, Nriagu*
Dept of Geology,
University of Wisconsin, Madison

Abstract

In Lake Mendota sediments, iron is present as the sulfides
(hydrotroilite and/or greigite), in organic material^ detrital ma¬
terial, magnetic spherules and as 'acid-soluble’ iron which has been
shown to be coprecipitated with calcite (the predominant mineral
phase in the lake muds) by adsorption. A small fraction of the
acid-soluble iron may also be tied up as the polyphosphates.

Introduction

The cycling of iron within and through the aquatic ecosystems
has been extensively investigated because the amounts and kinds
of ions or molecules containing iron in the ferrous or ferric states
are directly relatable to the pH and Eh of the water in which they
occur (Mortimer, 1941-42; Hutchinson, 1957; Gorham, 1958; Hem,
1959). None of these investigations however has reported on the
influence of mineralogy on the partitioning of iron between the
lake water and the solid phases in the bottom sediments with which
the water is in contact. This important variable which would
greatly influence the amount and rate of iron leaching from the
lake muds has been neglected because of the complete lack of infor¬
mation on the nature of iron in lake sediments (Hutchinson, 1957) .
The present investigation is aimed at providing some information
on the various forms of iron in the bottom muds of Lake Mendota,

Sampling and Field Observations

- The grab samples used in the analyses were obtained with an
Eckman dredge along a traverse from Picnic Point to Maple Bluff
(Fig. 1). The core samples were obtained near University Bay
(Fig, 1) with a three-inch diameter piston corer mounted on the
Water Chemistry Research Vessel, Kekule. Water depths were de¬
termined with a fathometer. Detailed information is available on

Present address : Dept, of Geoiog-y, University of Toronto, Toronto 5, Canada.

153

154 Wisconsin Academy of Science, Arts and Letters [Vol. 56

the physical properties of Lake Mendota sediments (see for exam¬
ple, Twenhofel, 1933 ; Hanson, 1952 ; Murray, 1956) ; no attempt
will be made to describe these properties in this report.

The often reported knife-sharp nature of the contact between the
sludge and the marl was not observed in any of the cores used in
this study. In all the core sections examined, the sludge passed
gradationally into the lake waters at the top and at the bottom
graded into marl over a zone ranging from five to ten centimeters
marked by a gradual lightening of color. Apparently the false im¬
pression of a knife-sharp contact (as reported by Hanson, 1952,
and Murray, 1956) was created by compression of the core section
during sampling. Emery and Dietz (1941) reported that gravity
corers gave shortenings of up to 60 percent in some marine sedi¬
ments off the coast of California. Core shortening of comparable
magnitude is quite possible in these sediments considering the high
fluidity of the sludge (water content up to 85 percent). Murray
(1956) describes the core : — “when the core and liner were removed
from the steel tube, the water above the sediments in the liner

1967-68] Nriagu — Distribution of Iron in Lake Sediments 155

remained clear despite the agitation of the water and sediments in
the sampling process”. This may be considered a measure of the
degree of compression of his core samples. Furthermore, the thick¬
ness of the sludge measured directly along the core column or de¬
termined on the basis of sulfur content of the sediments, is gen¬
erally much greater than that reported by Murray and may be
regarded as another evidence that the cores studied by him were
compressed. The core section obtained for this study almost com¬
pletely filled the core barrel showing that the shortening of the
column was very small.

Laboratory Investigations

All the samples were stored frozen until required for analysis.
To minimize the air oxidation of the ferrous iron, the interval
between the collection of the samples and the determination of the
ferrous iron was kept at less than a week. The analysis for ferrous
iron was made on wet samples; the total iron content was deter¬
mined on oven-dried samples.

Ferrous Ironb The analysis for acid-soluble (ferrous) iron con¬
sists of boiling the sediment sample with IN. HCl, filtering off the
iron in the solution and reducing the filtrate with a 10 percent solu¬
tion of hydroxylammonium hydrochloride. The iron in the extract
is then determined by the o-phenanthroline method using a Beck¬
man Model B spectrophotometer with a wavelength setting of 519
m/x (a make up from a filtrate was used as the standard in the
reference cell) . The details of the procedure used in the analyses
are given in Standard Methods (Am. Publ. Health Assoc. 1960).

Total Iron, To obtain the total iron, a weighed portion of the
oven-dried sediment is digested with a mixture of HNO3, HCIO4
and HF and the iron in the acid extract determined by the 0-
phenanthroline method. The experimental details involved in the
analysis are given in Black (1965).

Sulfide Sulfur, Sulfide sulfur was analysed by the evolution
method of Kolthoff and Sandell (1952). An excess of dilute HCl is
added to a weighed amount of the sediment sample in a distillation

1 Bailing in dilute acid will also dissolve a considerable quantity of ferric iron, if
any happens to be present. It is how'ever unlikely that ferric iron is present consider¬
ing the fact that these samples were drawn in the late stages of stratification when
the bottom sediments are known to be highly reducing. Ferric iron precipitated in
bottom muds during the seasonal overturns has been shown (see Hutchinson, 1957 ;
Mortimer, 1942/43) to be almost completely reduced to the ferrous state early, fol¬
lowing the development of anoxic conditions in the hypolimnion. In addition, any
ferric iron settling through the oxygenated epilimnion should in fact be reduced in
the anaerobic hypolimnion before it can get to the bottom muds — ferrous and ferric
ions in aquatic ecosystems are very sensitive to Eh-pH changes (J. D. Hem, 1959).
It is simply unlikely that ferric iron should exist as a stable phase (except as pyrite,
which however was not isolated in any of the samples) in the presence of relatively
large concentrations of dissolved sulfide associated with these bottom muds.

156 Wisconsin Academy of Science, Arts and Letters [Vol. 56

flask. The acidified suspension is boiled gently for one hour on a
hot plate and the liberated H2S absorbed in a zinc acetate-sodium
acetate mixture and subsequently determined volumetrically using
a standard iodine solution as the titrant and starch solution as the
end point indicator.

Total Sulfur, Analysis for total sulfur was by dry combustion to
the sulfate followed by reduction and subsequent conversion of the
sulfuric acid to H2S. The precautions and experimental details for
the determination of SO4 as hydrogen sulfide (using a reducing
mixture of hydriodic acid, hypophosphorus acid and formic acid)
are given in Black (1965).

Results and Discussion

The analytic data for the dredge samples are given in Table 1 ;
the data for the core samples are presented in Table 2. All depths
are apparent depths, no corrections for core shortening have been
made.

In all the samples examined, (see Tables 1 and 2) , the iron occurs
predominantly in the acid-soluble (ferrous) state. The difference
between the total and acid-soluble iron at any given location can
be accounted for as iron in organic matter, in detrital sediments
and in the magnetic spherules (for a discussion of these spherules,
see Nriagu, 1967). It was not possible to differentiate between
these forms of iron owing to the diflSculties involved in removing
organic material from these sediments. Since these forms may be
regarded as the “inactive” iron in the sediments (and because not
much is known about their form anyway), no further discussion
will be made about them.

Table 1. Sulfide-S, Total-S, Acid-soluble Fe and Total Fe Content
OF Dredge Samples of Lake Mendota Sludge*

Sample No.

Depth of
Water
(Ft.)

Sulfide

SuLFURf

Total

SuLFURf

Acid-

soluble

iRONf

Total

iRONf

901/7 .

35

0.3

0.5

5.5

901/6 .

50

1.0

1.9

13.7

17.0

901/1 .

50

1.3

1 .9

16.5

22.5

901/5 .

60

1.6

2.2

16.3

21.0

901/2 . . . . .

75

3.5

4.2

21.6

24.0

901/4 .

77

3.3

4.0

19.4

23.6

901/3 .

83

3.6

4.3

22. 1

25.0

tExpressed as mg/gm dry weight.
*Samples were drawn on 9/1 /66.

1967-68] Nriagu — Distribution of Iron in Lake Sediments

157

Table 2. Sulfur and Iron Contents of Core Samples.
Depths of Water = 33 ft.

Sample Number

Depth

Beneath

Mud

Surface

‘“Acid-

Volatile

Sulfur

*Total

Sulfur

“Acid-

Soluble

Iron

“Total

Iron

1 .

5-5cm

0.6

1.3

15.6

20.4

2 .

5-10

1.4

1.5

15.1

20.0

3 .

10-15

1.7

1.8

16.8

20.6

4 .

15-20

1.9

2.0

15.9

—

5 .

20-25

1.4

1 .9

15.7

22. 1

6 .

25-30

1.3

1.8

16.4

22.2

7 .

30-35

0.9

1 . 1

15.2

21.9

8 .

35-40

0.5

0.9

14.6

20.4

9 .

40-45

0.5

0.7

14. 1

19.4

10 .

45-50

0.4

0.6

11.7

16.0

12 .

55-60

0.2

0.5

9. 1

12.5

14 .

65-70

0. 1

0.4

4.3

7.9

16 .

75-80

0. 1

0.3

3.2

7.0

18 .

85-90

0. 1

0.2

3.1

7.2

*Concentration expressed as mg/gm of the dry sediment.

The iron sulfide in these sediments is a black amorphous, acid-
soluble substance believed to be hydrotroilte, FeS.nHoO (and/or
greigite, Fe3S4) and is responsible for the color of the sludge. No
pyrite or marcasite was isolated from these lake muds.

A feature shared by the sludge and marl is that the iron content
is very much greater than should be required to hold all the sulfur
present as FeS (for instance, the molar ratio of sulfide sulfur to
acid soluble iron varies from 1:8 to 1 :4) ; only a small part of the
total iron in the sediments can be present as FeS even though a
significant proportion of the sulfur in the sediment may be so
contained. It is not clear however in what form this excess iron^
exists in the lake sediments. The question of mineralogy is im¬
portant because it affects the aqueous chemistry of the solid (min¬
eral) phase and would influence to a great extent the amount and
rate of iron leaching from the sediments. The problem in dealing
with the excess iron in the sediments is that the iron mineral
phase (s) present cannot be detected by X-ray diffractometry or
microscopic techniques.

Ferrous carbonate has been suggested as the solid form in which
the excess iron exists in Lake Mendota sediments (Murray, 1956;
Lee, 1962; Gardner and Lee, 1965). With the available chemical

* Excess iron is used here as an operational term referring- to the acid soluble iron
in these sediments not accounted for as the sulfides. It g-ives an indication of the
‘reactive’ iron in the sediments other than the sulfides.

158 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Table 3. Average Analysis of Water from Lake Mendota

(ppm)

Molarity

Alkalinity .

142

0.0014*

Ca .

30

0.00075

Mg .

24

0.0001

Fe .

0.1

0.000002

Mn .

0.05

—

Chlorides .

5.0

0.00014

Sulfates . . .

17.0

0.00018

Ammonia Nitrogen .

0.08

—

Organic Nitrogen .

0.6

—

Silica (Si0 2) .

1.0

—

pH .

8.0

*RecalcuIated to HCO 3“.

analyses of the lake water it is possible to calculate whether or not
the waters of the lake are in equilibrium with respect to siderite.
An average analysis of Lake Mendota water (courtesy of the Water
Chemistry Laboratory, Univ. of Wise.) is shown in Table 3.

Recalculation of these, assuming that at pH 8, the titration alka¬
linity equals the bicarbonate alkalinity and ignoring ammonia, or¬
ganic nitrogen, Mn, and Si02, shows that the ionic strength is about
0.005 ; mpe+ + = 10-^-^ ; mHco“ = lO-^-^L

The activity coefficient 7, for HCO it and Fe^+ can be computed
from the Debye-Huckel equation:

— log yi = A • V I ^

1 + ai • B • V I

where A and B are constants relating to the solvent (in this case
water) ; z is the ionic charge; I is the ionic strength of the solution;
and ai represents the effective diameter of the Jon in solution. Sub¬
stituting I = 0.005 and the empirical values for the constants at
25 °C (Carrels and Christ, 1965, p. 61-62) into the above equation,
we obtain:

yFe2+ = 0.72 = 10-0-12
yHCOg- = 0.93 = 10-0-03

Precipitation of siderite is generally believed to be controlled by
the reaction :

FeC03(s) + H+ = Fe2+ -f HCO3- (2)

.‘. K = aFe2+ * anco «

- p — (^7

^FeCO g ■ -F

where a represents the activity of the ion.

1967-68] Nriagu — Distribution of Iron in Lake Sediments

159

Assuming apecog = 1, the equilibrium may be stated in terms of
molarities m, and the activity coefficient y :

K

mFe2+ • mnco “ = —

^ yre"'^ • yHOOs

The equlibrium constant, K for this reaction calculated from free-
energy data is 0.46 (Latimer, 1952).

ForyFe2+ = 10-012 ;yHC03- = 10-0 03
aH+ = 8.0 ; mnco 3 “ = 10-®-^ ; and K = lO-o-^^^ then
mpe2+ = 10-5-4

The iron content of the lake (Table 3) is 10“ ^ therefore accord¬
ing to these calculations, the lake appears to be in equilibrium (just
saturated with respect to FeCOs. Thus within the limits of the
assumptions used in these calculations, there is a likelihood of sider-
ite being precipitated in the lake.

The preceding calculations represent the conditions during the
overturn when a homogeneous chemical system is established in
the lake and the pH is 8.0. During the periods of thermal strati¬
fication however, the pH of the hypolimnion near the mud-water
interface commonly falls to 6. 5-7.0 (Open File Report, Water
Chemistry Dept). Substituting this pH range in equation 4, ,we
find :

mFe2+ = 10-^-no 10-4-4

But the iron content of the lake water (Table 4) is lO-^-'^ indicat¬
ing that during the periods of bottom anoxia the interstitial water
in contact with the sludge is undersaturated with respect to fer¬
rous carbonate. The large difference in value between the observed
and the calculated mFe2+ suggests a high degree of instability and
a strong possibility of solution of the ferrous carbonate.

If the excess iron exists as the ferrous carbonate, its stability
must therefore be due to its inability to equilibrate with the dy¬
namic variables of the interstitial water. Experiments on the solu¬
tion kinetics of carbonates in dilute carbonic acid solutions indi¬
cated that the rates of the solution were dependent only on the
rate of diffusion of the ions in the aqueous phase (Weyl, 1958).
Possible factors (all of which are manifestations of surface phe¬
nomena) in the lake that may inhibit simultaneous solution of
FeCOs, thereby engendering non-equilibrium behavior, include ad¬
sorbed protective coating on the grains, isolation of the siderite
grains as mechanical inclusions within the flocculent particles, and
low temperature (which retards the rate of solid diffusion). There
is however no evidence from these sediments to suggest that any

160 Wisconsin Academy of Science, Arts and Letters [Vol. 56

of the processes is keeping either the calcite grains or the other
chemical precipitates (e.g. the sulfides) out of reaction.

It is reasonable to suppose that if the excess iron is precipitated
as the carbonate, the grains have the same size range as the calcite
crystals. The grain size of the calcite crystals was determined by
the method discussed in Henry, Lipson & Wooster (1951, p. 212-
213). Diffraction photographs were taken of marl samples which
were mounted on glass fiber with the maximum care to avoid crush¬
ing the calcite grains. Each sample gave well defined powder lines
without spots showing that the crystals have a size range of 0.001
tot 0.0001 mm diameter. If therefore ferrous carbonate is present
in these sediments, it would be as very fine particles with high spe¬
cific surface area, a condition that would favor rapid decomposi¬
tion in air. The decomposition of FeCO.s in air is controlled by grain
size and temperature (Seguin, 1966). Aeration of Lake Mendota
sediments would thus be expected to cause a notable reduction in
the excess iron, if present as the ferrous carbonate. Table 4 how¬
ever shows that this is not the case. Furthermore, it has been shown
by Gardner (1964) that after the initial rapid uptake of oxygen
(probably due to the reaction of oxygen with the sulfides), the oxy¬
genation of the iron compounds in the lake sediments becomes a
linear function of time. This clearly should not be expected if the
iron were precipitated as the easily oxidizable ferrous carbonate.
These two observations thus cast considerable doubt on the sugges¬
tion that the excess iron has been precipitated as FeCOs (siderite).

Table 4. Comparison of Rates of Oxidation of Iron Sulfides
AND Excess Iron of the Sludge in Air*

Sample No.

SULFIDE-S

Acid-Soluble Iron

Initial

(ppm)

After

Aeration

Initial

(ppm)

After

Aeration

(ppm)

901/1 .

1,300

70

16,500

11,200

901/3 .

3,600

204

22,100

14,700

901/4 .

3,300

150

19,400

12,300

901/5 .

1 ,600

190

16,300

12,900

*To obtain the data, the sludge samples were aerated until the dark color had changed
to grey-brown. The sulfide sulfur and acid-soluble iron were determined again. The
aeration procedure consisted of exposing the periodically stirred, wet sludge to the
laboratory air. Distilled^water was added to keep the samples continuously moist.

1967-68] Nriagu — Distribution of Iron in Lake Sediments

161

If therefore the excess iron is not precipitating as FeCOa, the
alternative phenomena which may account for it include:

1. Formation of organic complexes and chelates

2. Formation of solid phases other than siderite

3. Coprecipitation of the iron with the calcite.

To estimate the amount of iron in the sludge associated with
organic compounds and chelates, the sludge samples were leached
with acetone, the extract carefully evaporated to dryness and the
residue analysed for iron by the o-phenanthroline method. The
results are given in Table 5 below.

Table 5. Iron Content of Acetone Extracts from Sludge Samples

Sample No.

Depth of
Water
(Ft.)

Fe in
Acetone
Extract
(ppm)

Initial

Acid-

Soluble

Fe (ppm)

901/5 .

60

13

16,300

901/2 .

75

26

21,600

901/4 .

77

21

19,400

901/3 .

83

19

22,100

The data above show that acetone-soluble organic compounds and
chelates not bound up as solid and particulate organic matter ac¬
count for only a very small fraction of the iron in the sludge. Ap¬
parently, not much of the excess iron is associated with the yellow
coloring matter observed when organic muds are treated with ace¬
tone as has been suggested by J. D. Hem (1959).

In addition to the carbonates, other iron minerals (possible in
these sediments) that may exist metastably in aqueous environ¬
ments are silicates and phosphates (see Weyl, 1966). Iron silicates
may be ruled out because most of them are insoluble in dilute acids.
The occurrence of phosphorus not associated with organic matter
or clay minerals has been reported in Lake Mendota sediments
(Wentz, 1967). It seems reasonable to suggest that a part of this
phosphorus is combined as acid-soluble polyphosphates, the meta¬
stability of which may be due to biologic and/or chemical factors.
The presence of ferric phosphate should not be affected by aeration
which is in accordance with the data presented in Table 5. The
mean phosphate content of the lake sediments is in the range 1 to
2 mg/gm on a dry weight basis (Sawyer et ah, 1944; Kaneshige,
1952; DeMno, 1967, Pers. Comm,). Clearly this quantity is insuffi¬
cient to account for all the excess iron in the samples examined.

162 Wisconsin Academy of Science, Arts and Letters [Vol. 56

The final possibility is that the greater part of the excess iron
is coprecipitated with calcite by adsorption. Coprecipitation of iron
with calcite would mean that the activity of the solid carbonate
phase is not one as has been assumed in the calculation but less than
one. From Equation 3 one finds that the effect of lowering aFeco ^ is

to increase the value of the calculated mFe2 + . This effect would of
course decrease the difference between the calculated and observed
mFe2+ and hence should be particularly significant during periods
of stratification when the pH falls to 6. 5-7.0. The smaller this dif¬
ference is, the less undersaturated the water is relative to FeCOs
and the less likelihood of the solid FeCOs going into solution. It is
thus possible to account, at least partially, for the stability of the
excess iron during periods of bottom anoxia by coprecipitation
which has decreased the activity of the solid carbonate phases in
the aqueous system.

Such a coprecipitation would also reasonably explain the data
of Table 6 ; the apparent stability being due to the very slow diffu¬
sion of iron through the calcite grains which thereby controls its
rate of oxidation. Furthermore, the chemical reaction rates of a
constituent in solid solution has been shown to approximate a
zero’ th order process (Crocket et ah, 1966) . This may be the expla¬
nation for the linear rate of oxygen uptake observed during the
manometric oxygenation of these lake sediments (Gardner & Lee,
1965).

In conclusion, the available evidence suggests that the greater
fraction of the excess iron in Lake Mendota sediments has been
coprecipitated with calcite (the predominant single mineral in the
sediments) rather than precipitated as the pure compound, FeCOs,
as has been suggested by Murray (1956), and Lee (1962). A small
part of the excess iron may also be tied up as polyphosphates. Fur¬
ther experiments are necessary to evaluate the exact mechanism
of coprecipitation, particularly the influence of organic complexing
and biochemical processes.

The influence of the nature of solid phase (with which the lake
water is in contact) on the cycling of iron in the lake is obvious.
Iron structurally incorporated into calcite is of course not a read¬
ily exchangeable iron. Most of the iron released from the sediments
during thermal stratification must therefore come mainly from
sources of 'available’ iron other than the excess iron, notably the
sulfides, (the phosphates) , and perhaps, in the early stages of strat¬
ification, the oxides also. Evidence that part of the iron leached
from the sediments has resulted from the dissolution of iron sul¬
fides comes from the observed coexistence of free H2S and Fe^+ in
the hypolimnion during the latter stages of bottom anoxia. Since
iron sulfides can only dissociate under certain restricted conditions

1967-68] Nriagu — Distribution of Iron in Lake Sediments 163

(low Eh, and pH less than 7.0; see Garrels and Christ, 1965) it
follows that dislocation of iron in the lake 'by dismutation must be
very small. Consequently, the lake sediments must be acting as a
large sump for iron, a suggestion which has already been confirmed
by Rohlich (1963) who reported an iron retention in the lake of
over 80 percent.

Acknowledgements

I would like to express my gratitude and appreciation to Dr. C. J.
Bowser of the Geology Department for his help and guidance
throughout this study and the preparation of the manuscript. Ap¬
preciation is also expressed to Professor G. F. Lee of the Water
Chemistry Laboratory for his help in obtaining the samples used
in the analysis. The work was done with the financial assistance
from African-American Institute (New York) and the Nigerian
Government.

Bibliography

Black, C. S., (1965) (Editor), Methods of Soil Analysis: Univ. of Wise.
Press; Amer. Soc. of Agronomy Inc.

Emery, K. 0. and R. S. Dietz, (1941) Gravity coring instrument and mechan¬
ics of sediment coring: Bull. Geol. Soc. Amer. 52, p. 1685-1714.

Gardner, W. S., (1964) The oxygenation of lake sediments: M.S. thesis. Wa¬
ter Chem. Dept., Univ. of Wise.

- and G. F. Lee, (1965) Oxygenation of lake sediments: Int. Jour. Air

and Water Poll, 9, p. 533-564.

Garrels, R. M. and C. L. Christ, (1965) Solutions, Minerals and equilibria:
New York, Harper and Row, 450 pp.

Hanson, G. F., (1952) Some observations on the sediments of University
Bay, Lake Mendota, Wisconsin: Univ, Wise. Comm, on Lake and Stream
Investig. Rept. (mimeo), 22 pp.

Hem, J. D., (1959) Chemistry of iron in natural waters: U. S. Geol. Survey,
Water Supply Papers, 1459A-1459I.

Henry, N. F., H. Lipson and W. A, Wooster, (1961) An interpretation of
X-ray diffraction photographs: London, McMillan & Co. Ltd. 282 pp.

Hutchinson, G. E., (1957) A treatise on Limnology: New York, John Wiley
& Sons, Vols. 1 and 2.

Kaneshige, H. M., (1952) Chemical analysis of bottom muds of Lake Men¬
dota: (Unpubl.) M.S. Thesis, Civil Engr. Dept. Univ. of Wise.

Latimer, W. M., (1952) Oxidation Potentials: 2nd Edition, New York, Pren¬
tice-Hill, p. 234-241.

Lee, G. F., (1962) Studies on iron, manganese, sulfate and silica balances and
distribution for Lake Mendota, Wisconsin: Trans. Wise. Acad. Sci. Arts
and Letts. 51, p. 141-155.

Mortimer, C. H., (1941-42) The exchange of dissolved substances between
mud and water in lakes: Journ. Ecology, 29, p. 280-329.

Murray, R. C., (1956) Recent sediments of three Wisconsin lakes: Bull. Geol.
Soc. Amer. 67, p. 884-911.

Nriagu, J. 0., (1967) The distribution of sulfur and iron in Lake Mendota
sediments: M.S. Thesis, Dept, of Geology, Univ. of Wise,

164 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Rohlich, G. a., (1963) Origin and quantities of plant nutrients in Lake Men-
dota; in D. G. Frey (Ed.), Limnology in North America, Madison, Univ.
of Wise. Pi’ess, p. 75-88.

Sawyer, C. N., J. B, Lackey and T. A. Lena, (1944) Investigations of odor
nuisance occurring in Madison lakes: (mimeo) Kept, of Governors Com¬
mittee on Lakes, Madison, Wise.

Seguin, M., (1966) Instability of ferrous carbonate in air: Amer. Journ. Sci.
26J,, p. 562-568.

Vogel, I. I., (1961) Quantitative Inorganic Analysis: London, Longman Green
& Co., 1216 pp.

Wentz, D. A., (1967) Available phosphorus in lake sediments: (unpubl.)
M.S. Thesis, Water Chem. Dept., Univ. of Wise.

Weyl, P. K., (1958) The solution kinetics of calcite: Journ. Geol. 66, p. 163-
176.

- (1966) Environmental stability of the earth’s surface. Chemical con¬
siderations: Geochim. et Cosmochim. Acta, 30, p. 663-679.

EFFECT OF FLOODING, DRAINAGE AND pH
ON TRANSFORMATIONS OF Mn AND
Fe IN 19 WISCONSIN SOILS'

H. Graven and O. J. At toe'

The results of a previous study (2) indicated that limited pe¬
riods of flooding or waterlogging can greatly increase the amount
of exchangeable Mn in soils and cause acute Mn toxicity in alfalfa.
Owing to its dependence on the redox potential of the soil, Mn
availability is conditioned by the amount of easily decomposable
organic matter present and by seasonal variations in temperature,
moisture and microbial activity. Flooding of soils is generally fol¬
lowed by a rapid rise in water soluble and exchangeable Mn and a
somewhat slower rise in these forms of Fe (1, 7, 10). Harter and
McLean (3) found that a moisture content intermediate between
field moisture capacity and complete saturation was sufficient to
fill all except the largest pores of a clay loam soil and cause nearly
as large an increase in exchangeable Mn as complete flooding.
Although some disagreement exists concerning the relation between
soil pH and the content of exchangeable Mn (5, 12, 14), liming
acid soils is generally known to lower the amount of this constitu¬
ent. Oxidation of Mn in soils may be due largely to microbial activ¬
ity (9, 11, 13), but greater difficulty has been encountered in dem¬
onstrating microbial oxidation of Fe (4) because this process also
proceeds in sterilized soils.

The present study was initiated to obtain information on the
effects of flooding and drainage on the transformations of Mn and
Fe in a number of important Wisconsin soil types representing a
wide range in pH, texture and other properties.

Methods and Materials

Twenty-gram samples of the air-dry soils were flooded in
weighed 100-ml glass bottles by the addition of 40 mis of distilled

1 Contribution from the Department of Soils, University of Wisconsin, Madison. Pub¬
lished with the approval of the Director, Wisconsin Agr. Exp. Sta. Part of a disserta¬
tion submitted by the senior author in partial fulfillment of the requirements for the
Ph.D. degree. University of Wisconsin.

2 Research Assistant and Professor of Soils, respectively.

165

166 Wisconsin Academy of Science, Arts and Letters [Vol. 56

water. Exchangeable Mn and Fe were determined after 0, 5, 15,
35 and 75 days of flooding. For reference purposes, similarly
treated samples were flooded in 800-ml beakers at the same time.
After 75 days of flooding, the latter samples were thoroughly
stirred, transferred to Buchner funnels and the excess water re¬
moved by suction. Suction was maintained for 45 minutes after
the surface water had disappeared. Twenty-five grams of this moist
soil was subsampled into weighed glass bottles. The moisture con¬
tent was determined from separate samples. The bottles were
stored open to the atmosphere at room temperature and the soils
were maintained near field moisture capacity (FMC) by watering
to the initial weight every third day. Exchangeable Mn and Fe
were determined after 0, 5, 15, 35 and 75 days at FMC.

Exchangeable Mn and Fe were extracted with N Mg(N03)2;
easily reducible Mn was extracted with N NH4OAC (pH 7.0) con¬
taining 0.2% hydroquinone ; and total Mn was brought into solu¬
tion by use of HF-HCIO4. Mn and Fe were determined colori-
metrically (6) using NasH^IOe oxidation and orthophenanthroline
methods, respectively. Organic matter was determined by the
Walkley-Black method (15). Cation exchange capacity was
determined flame-photometrically following saturation with
N Ca(0Ac)2 at pH 7.0 and displacement with N Mg(OAc)2. Ex¬
changeable cations were displaced with N NH4OAC adjusted to pH
7.0 and determined flame-photometrically. The clay content was de¬
termined by the pipette method (8). Soil pH was determined on a
thin soil paste.

Results and Discussion |

The values for the chemical properties of the individual soils are |
given in Table 1, and the range and average values for these j
properties by pH groups are given in Table 2. The relatively high |
values for exchangeable Mn in many of the soils prior to flooding j
were no doubt due to the well-known effect of drying on increasing j
the content of this constituent. The linear correlation coefficients I
obtained for pairs of these properties, as indicated in Table 3, show
a close relation among the three forms of Mn. A close relation was
also found between the clay content and the properties of exchange¬
able Mn, cation exchange capacity and total cations. Similarly,
there was a close relation between cation exchange capacity and
the contents of organic matter and total cations.

The data presented in Fig. 1 shows that flooding for as little as 5 ;
days caused a marked increase in the values for exchangeable Mn
in all three soil pH-groups and in exchangeable Fe in the pH ;
5.3-group. The Mn values reached a maximum after 15 to 35 days i
and declined somewhat with further flooding. The reason for the i

Table 1. Certain Chemical Properties of the 19 Wisconsin Soils

1967-68] Graven and Attoe — Transformations of Mn and Fe

167

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168 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Table 2. Range and Average Values for Certain Chemical Properties of
Three pH Groups of 19 Wisconsin Soils

Soil Property

pH 4. 8-5. 5
(6 Soils)

pH 5 . 6-6 . 0
(8 Soils)

pH 6. 3-7.4
(5 Soils)

Range

Ave.

Value

Range

Ave.

Value

Range

Ave.

Value

pH .

4.8- 5.5

5.3

5.6- 6.0

5.8

6.3- 7.4

6.8

Exch, Mn, ppm .

Easily reducible Mn,

5-49

30

11-82

43

1-67

35

ppm .

17-481

211

112-653

359

2-722

422

Total Mn, ppm .

Cation exch. cap.,

236-981

595

507-1641

836

120-1587

911

meq/100 g .

1 .9-24.3

10.8

3.1-18.4

10.8

6.4-21.5

15.0

Organic matter, % .

0.4-3. 7

1 .9

1 . 1-3.2

1.9

1.6-2. 7

2.2

Clay, % .

3.5-27.7

14.3

3.4-28. 1

14.5

9.2-23.9

18.3

Total cations, meq/100 g

0.9-14.2

6.4

1.9-14.3

8.0

6.3-17.4

12.9

Table 3. Linear Correlation Coefficients* for the Relation Between
Various Chemical Properties of the 19 Soils

Soil

pH

Cation

Exch.

Cap.

Or¬

ganic

Matter

Total

Ca¬

tions

Total

Mn

Easily

Reduc.

Mn

Exch.

Mn

Clay .

.06

.88

.49

.91

.45

.48

.66

Soil pH .

.03

. 14

.33

.07

. 16

.18

Cation exch. cap. . .

.68

.84

.44

.41

.55

Organic matter. . . .

.55

.35

.29

.24

Total cations .

.52

.54

.53

Total Mn .

.93

.84

Easily reduc. Mn. .

.84

Significant at respective levels if value is equal to or greater than following:
5%==*=. 46 and 10% = . 39. All values shown are positive.

decline is not clear but it may be due to absorption of Mn by a
larger population of anaerobic microorganisms present during the
latter part of the flooding period. The highest values for exchange¬
able Mn were attained in the 5.8 and 6.8 pH-groups. In contrast,
the values for exchangeable Fe continued to rise all through the
75 days of flooding, and the highest values were attained for soils
in the pH 5.3-group and the lowest in the pH 6.8-group. Drainage
of the pots and a return to field moisture capacity resulted in an
abrupt drop in exchangeable Fe in all groups and in exchangeable
Mn in the pH 6.8-group. The Mn values for the pH 5.3-group
declined very slowly and those for the pH 5.8-group were inter-

1967-68] Graven and Attoe — Transformations of Mn and Fe 169

Figure 1. Effect of flooding, drainage and pH on the contents of exchangeable
Mn and Fe for the three pH-groups of soils.

mediate. These results suggest that the most harmful effects on
plant growth of relatively long periods of flooding or waterlogging
could be for soils near the neutral point because of the very wide
ratio of exchangeable Mn to Fe under these conditions. However,
for rather short periods of saturation the most harmful effect
could be for the more acid soils in which the exchangeable Mn re-
mained at a very high level for extended periods following drain¬
age. The results also suggest the need for caution in the amounts
of water added to potted plants.

The data presented in Table 4 show that the exchangeable Mn
values after 15 days of flooding were highly correlated with both
total and easily reducible Mn. The decrease in this constituent dur¬
ing 35 days at FMC after 75 days of flooding was closely correlated
with soil pH and % clay. The importance of the clay content in
this relation appears to be due in part to the fairly close association
between it and the forms of Mn present prior to flooding. The
exchangeable Fe values after 35 days of flooding were highly cor¬
related with soil pH (negatively) and % organic matter. Appar¬
ently, the presence of organic matter favored the reduction and

170 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Table 4. Coefficients of Multiple Correlation and Standard Partial
Regression Coefficients for the Relation Between Selected
Soil Properties of the 19 Soils in Relation to
Flooding and Drainage

Standard Partial

Coefficients

Independent Variables

Regression Coefficients
FOR Respective Variables

OF Multiple
Correlation

Exchangeable Mn after 15 days of flooding

Easily reducible Mn; total Mn .

Easily reducible Mn; soil pH .

Soil pH; % clay; % organic matter.

.44

.54

—

.96**

.97

— . 14

—

.96**

— .04

.37

.24

.52*

Decrease in exchangeable Mn during 35 days at EMC after 75 days of flooding

Soil pH; % clay . I .38 .53 - 1 .68**

Soil pH; % clay; % organic matter. . j .40 .63 — 18 | .69**

Exchangeable Fe after 35 days of flooding

Soil pH; % organic matter . j — .72 .57 - I .86**

Soil pH; % clay ;% organic matter. . 1 — .73 .13 .50 | .87**

Exchangeable Fe after 5 days at EMC following 75 days of flooding

Soil pH; % clay .

— .55

.41

S —

.67**

Soil pH; % clay; % organic matter. .

— .54

.44

— .07

.67**

*Significant at 5% level.
**Significant at 1% level.

transformation of the Fe in the ferric oxides to the more soluble
and exchangeable ferrous forms. The Fe values after 5 days of
drainage were closely correlated with soil pH (negatively) and
% clay.

The increases in water soluble and exchangeable Mn caused by
flooding and decreases after drainage may be represented by the
following equation:

(Reduction)

Mn02 + 4H+ + 2e- Mn++ 4- 2H2O

(Oxidation)

Flooding and low soil pH values tend to shift the reaction to the
right and drainage and relatively high pH values tend to shift it
to the left. A similar equation may be used to represent the trans¬
formations of Fe under conditions of flooding and drainage. In this
case FeaOs would be reduced to Fe+ + . For soils near the neutral
point, this reaction appears to proceed less readily than for MnOo.

Summary

Flooding gave large increases in exchangeable Mn in nearly all
of the 19 soils studied. After drainage and a return to field mois¬
ture capacity, an abrupt decrease occurred in this constituent for

1967-68] Graven and Attoe — T-^ans formations of Mn and Fe 171

soils in the range of pH 6.0 to 7.4 and a much slower decrease in
the more acid soils. In contrast, flooding resulted in a much slower
increase in exchangeable Fe for the soils in this pH range and a
more abrupt decrease in most of the soils after drainage. An im¬
portant benefit derived from liming acid soils appears to be in
causing a more rapid decline in excessive amounts of exchangeable
Mn following drainage of saturated soils. The data reported help
understand the response of plants to waterlogging and drainage of
soils and emphasize the harmful effects that may result from over¬
watering of potted plants.

References Cited

1. Clarke, F. E., and J. W. Resnicky. 1956. Some element levels in the soil

solution of a submerged soil in relation to rate of organic matter addition
and length of flooding. Proc. 6th Int. Cong. Soil Sci. C: 545-548.

2. Graven, E. H., 0, J. Attoe, and Dale Smith. 1965. Effect of liming and

flooding on manganese toxicity in alfalfa. Soil Sci. Soc. Amer. Proc.
29:702-706.

3. Harter, R. D, and E. O. McLean. 1965. The effect of moisture level and

incubation time on the chemical equilibria of Toledo clay loam soil.
Agron. J. 57:583-588.

4. Hodgson, J. F. 1963. Chemistry of the micronutrient elements in soils.

Advances in Agronomy, Academic Press Inc., N. Y. 15:119-159.

5. Hoff, D. J. and R. J. Mederski. 1960. The chemical estimation of soil

manganese. Trace Elements, Academic Press Inc., N. Y. pp. 109-116.

6. Jackson, M. L. 1958. Soil Chemical Analysis, Prentice-Hall, Inc.

7. Jeffrey, J. W. 0. 1960. Iron and the Eh of waterlogged soils with particu¬

lar reference to Paddy. J, Soil Sci. 11:140-148.

8. Kilmer, V. J. and L. T. Alexander. 1949. Methods of making mechanical

analysis of soils. Soil Sci. 68:15-24.

9. Keeper, G. W. and R. J. Swaby. 1940. The oxidation of manganous com¬

pounds by microorganisms in the soil. Soil Sci. 49:163-169.

10. Mandal, L. N. 1961. Transformation of iron and manganese in waterlogged

rice soils. Soil Sci, 91:121-126.

11. Mann, P. J. G, and J. H. Quastel, 1946. Manganese metabolism in soils.

Nature 158:154-156.

12. Morris, H. D. 1948. The soluble Mn content of acid soils and its relation

to the growth and Mn content of sweet clover and Lespedeza. Soil Sci.
Soc. Amer. Proc. 13:362-371.

13. Mulder, E. G. and F. C. Gerretsen. 1952, Soil manganese in relation to

plant growth. Advances in Agronomy, Academic Press, Inc., N. Y, 4:
221-277.

14. Tisdale, S. L. and B. R. Bertramson. 1949. Elemental sulphur and its

relationship to manganese. Soil Sci. Soc. Amer, Froc. 14:131-137.

15. Walkley, a. and I. A. Black. 1934. An examination of the Degtjareff

method for determining soil organic matter and a proposed modification
of the chromic acid titration method. Soil Sci. 37:29-38.

J

THE FRAGIPAN IN SOILS OF NORTHEASTERN WISCONSIN'

Gerald W. Olsorr and Francis D. Hole^
Wisconsin Geological and Natural History Survey
The University of Wisconsin, Madison

A brittle subsoil horizon called the fragipan occupies large areas
in upland soils of northern Wisconsin, particularly in the north¬
east. This horizon restricts root growth (Figure 1) and movement
of water, and is therefore of importance to an understanding of
the ecology of forests and hydrology of drainage basins of northern
counties. This is a report on recent investigations of the fragipan
in Florence, Menominee, Oneida and Bayfield Counties, Wisconsin
(Milfred, Olson and Hole, 1967; Olson, 1962; Hole, Olson et ah,
1962; Ableiter and Hole, 1961; Hole and Schmude, 1959). This
work was supported by the Wisconsin Geological and Natural His¬
tory Survey.

Definition

Among existing definitions of the fragipan (Soil Survey Staff,
1951 ; Carlisle, et ah, 1957 ; Committee on Terminology, 1960 ; Soil
Survey Staff, 1960), the following is one of the most complete. ‘‘A
fragipan (modified from L. fragilis, brittle; and pan: meaning
brittle pan) is a loamy subsurface horizon, often underlying a B
horizon. It is very low in organic matter, has high bulk density
relative to the overlying horizons, is seemingly cemented when dry,
having hard or very hard consistence. When moist, a fragipan has
moderate or weak brittleness (tendency for a ped or clod to rup¬
ture suddenly when pressure is applied rather than to undergo
slow deformation). It is usually mottled, is slowly or very slowly
permeable to water, and has few or many bleached fracture planes
that form polygons” (Soil Survey Staff, 1960). Platy structure and
vesicular porosity are also characteristic of many fragipan hori-

1 Published with the pei'mission of the Director of the Wisconsin Geological and
Natural History Survey and the Director of the Wisconsin Agricultural Experiment

Station, Madison.

2 Formerly Research Assistant, Wisconsin Geological and Natural History Survey,
now Assistant Professor of Soil Science, in Resource Development, Cornell University,
Ithaca, N. Y.

2 Professor of Soil Science, in charge, Soil Survey Division, Wisconsin Geological and
Natural History Survey.

173

174 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 1. Photograph of the typical shallow rooting system of a tree growing
in a soil with a strongly developed fragipan.

zons. A slab of fragipan will slake when immersed in water. The
horizon designation is the letter x, added to the main horizon desig¬
nation, as in A'x and B'x of the lower solum of a double (bisequal)
soil profile (Figure 2).

Previous Work

Whitson (1927) referred to a ‘^hardpan” in the subsoil of Kennan
soils of northern Wisconsin, but did not describe it. Nygard, et ah,
(1952) referred to “a gray partially gleyed, vesicular pan’' lying
below the Podzol solum in soils of the Iron River and Munising
soil series in northern Wisconsin, Michigan and Minnesota. The
Soil Survey Staff of the U. S. Department of Agriculture (1951)

1967-68]

Olson and Hole — Fragipan in Soils

17e5

Figure 2. Cut-away block diagram of a bisequal soil, Goodman silt loam (Alfic
Haplorthod) , showing two fragipan horizons with extensions of the upper one
down a polygonal system of cracks in the lower, somewhat finer textured one.

assigned the term fragipan to such nonindurated pans, as distinct
from indurated pans (hardpans) and clay pans. During the past
decade a number of workers (Carlisle, et al., 1957; Grossman,
et ah, 1959; Yassoglou and Whiteside, 1960; Jha and Cline, 1963;
McCracken and Weed, 1963; Daniels, et aL, 1966) have investi¬
gated the characteristics and genesis of the fragipan in detail. This
horizon has developed under a variety of climates in a wide range
of materials, including Wisconsin loess, glacial till and lacustrine
deposits in the Lake States, residuum from sandstone, shale and
limestone farther south, and unconsolidated coastal plain sediments
in southeastern states. The fragipan does not occur in desert soils
(Aridisols) and other soils in which calcium carbonate and other
carbonates and salts either create a friable soil condition or cement
the soil into a hardpan (duripan) ; or in which claypans have
formed under the influence of the sodium ion. The fragipan does
not occur in prairie soils (Mollisols) in which root growth and
accumulation of organic matter have favored a more porous and
friable soil condition than is required for the formation of the pan.

176 Wisconsin Academy of Science, Arts and Letters [Vol. 56

The fragipan occurs in forest soils grouped in the soil orders, Incep-
tisols, Spodosols, Alfisols and Ultisols (Soil Survey Staff, 1960).
The investigations indicate that fragipan horizons have developed
by compaction of a mixture of noncalcareous silts and sands with
relatively little clay (Figure 3) and low content of organic matter.
Weak cementation is thought to be by films of clay, silica, alumina,
or iron oxide but this has not been definitely proved. The fragipan
has formed at sites where biologic activity in the subsoil has been
so slight that the fragipan has not been disturbed appreciably since
its formation. The fragipan has developed in the lower part of
many bisequal soils (Gardner and Whiteside, 1952), that is to say,
below a Podzol (Spodosol, Soil Survey Staff, 1960) sequum of hori-

Figure 3. Textures of fragipan horizons plotted on a soil textural triangle.
Solid lines represent data for soil samples from northeastern Wisconsin. Dotted
lines represent data for fragipans from other parts of the country (Olson,
1962). Arrowheads indicate the fragipan sample taken at the greatest depth
in each soil from which two or more subhorizons were sampled.

1967-68]

Olson and Hole — Fragipan in Soils

177

zons (Figure 2) and in a sequum consisting of an eluvial (A'2x)
horizon and an illuvial (B'2x) horizon. Bisequal soils of Wisconsin,
including Alfic Haplorthods like the Goodman silt loam (Figure 2),
have been discussed elsewhere (Carroll, 1959; Ableiter and Hole,
1961; Hole, et al., 1962; Beaver, 1963, 1966; Ranney, 1966). The
fragipan has been recognized in some soils on all continents except
Antarctica.

Geographic Distribution in Wisconsin

The fragipan occurs extensively in Wisconsin north of a bound¬
ary (A-A', Figure 4) which approximates the isotherm for the
average annual air temperature of 41° F (5° C) or about 43° F,
(6° C) in the soil at a depth of 50 cm. For the most part the bound¬
ary lies west of the carbonate-rich glacial drift of northeastern
Wisconsin. The postulated A-A' boundary of Figure 4 first ap¬
peared on a map published by Nygard, et ah, (1952) to indicate
the souhern limit of a zone of Podzol, Brown Forest and Brown
Podzolic soils, now classified as Spodosols and Inceptisols (Soil Sur¬
vey Staff, 1960). The boundary closely parallels and lies somewhat
north of range limits for several species of plants, such as the or¬
chid Habenaria obtusata (Curtis, 1959), within the northern mesic
conifer-hardwood forest. Weaker fragipans occupy large areas in
a 40-mile-wide zone to the south (A-A' to D-D', Figure 4; Carroll,
1959; Beaver, 1963), In both zones the fragipan commonly occurs
in acid bisequal soils.

Procedures

In the course of soil survey operations in the four Wisconsin
counties indicated in Figure 4, soil scientists working in the cooper¬
ative soil survey have described and analyzed soil profiles that in¬
cluded fragipan horizons. Observations were made of limitation of
root growth by the fragipan (Figures 1 and 2) and, in early spring,
of movement of water. Portions of this horizon that were experi¬
mentally exposed to weather for one to two years slaked differen¬
tially. Resistant gravel-size fragments were collected for analysis
from the slaked mass. Bulk density of representative soil peds was
determined by the method of coating dry peds with paraffin. Poros¬
ity was calculated from bulk density and an assumed specific grav¬
ity of mineral matter of 2.65. Estimates of root distribution were
made by weighing oven-dried roots by horizons from columns of
soil 6 inches (15 cm) in diameter. Particle size distribution analy¬
ses were made by the hydrometer method (Day, 1956). Meaning¬
ful calculations of pedogenetic gains and losses of soil plasma by
the index mineral method were found to be impossible, because of

178 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 4. Index map of Wisconsin showing* four generalized boundaries:
(A-A'), the southern limit of most strongly developed fragipans in Wisconsin;
(B-B'), the eastern limit of carbonate-rich glacial drift of the Grantsburg
glacial lobe; (C-C'), the western limit of carbonate-rich glacial drift of the
Green Bay glacial lobe (after Thwaites, 1943) ; (D-D'), the southern limit of
the zone of bisequal soils (in part after Carroll, 1959 and Beaver, 1963). The
four counties in which fragipans have been studied by one or both of the
authors are labeled: F, Florence; M, Menominee; 0, Oneida; Y, Bayfield.

the heterogeneity of the Pleistocene deposits from which the soils
formed. Contents of reductant-soluble iron oxide and Na2C03-
extractable silica and alumina, cation exchange capacity, exchange¬
able cations, contents of total carbon and nitrogen, and X-ray dif¬
fraction studies of clay minerals were made by methods outlined
by Jackson (1956, 1958). Soil reaction, and contents of available

1967-68]

Olson and Hole — Fragipan in Soils

179

nitrogen, available phosphorus and available potassium were deter¬
mined by the State Soil Testing Laboratory, and in the Cooperative
Subsoil Fertility Project at the University of Wisconsin, Madison,
Thin sections were prepared by the method of Buol and Fadness
(1961) and studied with the petrographic microscope.

Results and Discussion

The fragipan is recognized in the field by its compactness in
place and brittleness in hand specimen. The search in the labora¬
tory for other properties that definitely differentiate this pan from
other soil horizons in the same soil profiles has been rather unsuc¬
cessful, here (Table 1) as in other laboratories. The fragipan in
northeastern Wisconsin is a nonuniformly compacted, texturally
heterogeneous, slightly mottled, platy and vesicular horizon that
formed in a cold acid subsoil layer with sparse root growth. Since
its formation, the pan has been little disturbed by plant roots and
animals. Mottles are few, faint to distinct, and medium, reddish
brown to yellowish red in moist color (SYR 4/4-5/8). The tex¬
tural range of the fragipan (Figure 3 and Table 1) is from sandy
loam to silt loam, which is in the middle of the wider range from
loamy sand to silty clay loam reported from other parts of the
United States (Olson, 1962), Heterogeneity of particle size and a
low content of clay apparently favor the compaction necessary to
form a fragipan of bulk density between 1.42 and 1.97 (Table 1).
Weather-resistant gravel-size (2 to 3 cm in diameter) fragments
from the IIB'2x horizon of a Superior sandy loam contained 10%
clay, 39,% silt and 51% sand as compared with 24% clay, 45% silt
and 31% sand in the bulk sample from the same horizon. The rela¬
tively low content of clay and its peculiar distribution in the fragi¬
pan allow for the essential bridges and menisci of clay at many
points of contact between silt and sand grains. The amount of clay
present is not sufficient to cause noticeable disruption of the pan
by expansion and contraction during wet-dry cycles. Platy struc¬
ture and vesicularity may be inherited from the condition of the
horizon before it consolidated to the point that small plant roots
could no longer invade the prisms, blocks and coarse plates. Plati-
ness and vesicularity have been observed in the field and laboratory
to be formed by freeze-thaw cycles (Olson, 1962). Some voids in
the A'2x horizon may have resulted also from eluviation of clay
from it during a pre-fragipan stage of development by percolating
water and possibly also by advancing freezing fronts. Fragments
of clay films deposited by running water on ped surfaces were
observed in thin sections of fragipan horizons. These are presum¬
ably remnants, particularly in the upper fragipan, of formerly more

Table 1. Ranges of Some Properties of Soil Horizons from Several Northeastern Wisconsin

Silty Soil Profiles^ with Fragipans.

180

Wisconsin Academy of Science, Arts and. Letters [VoL 56

<0-

<

8^2

D

CD

O O O O
00 00 — u-'

^ — — —
I I I I I

Vn V/^ Its

sD ^ t\ csi

O 00 00 O O
O nO sD

Coarse
and Very

Coarse

0-9

1-9

1-16

1-23

0-48

-Q

C

cd

Me¬

dium

rq oi rq rq O

1 7777-?

rq rq — ' — ' — 1

z

o

H

D

£

CO

Fine

, — ^ sO VO Its

1 ro 03 rq Vo Vo

1

5

H

c/3

Q

Very

Fine

— ' OC — o

1 ol ol 0,1 ro ro

1 sO ro 03 t\ ol

LD

N

c5

ID

U

Coarse

sD Vo O' O' ro

1 ro ro ro ro Tf

1 1 1 1 1 1

1 VO — 00 00 O

k

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cx

Silt

Medi¬

um

11-45

9-33

6-33

5-31

0-26

Fine

2-12
4-9
2-1 1
2-21
0-14

O 00 O

^ r-)

I I I I I
IS,. >j-N

O'

—

—

I I I I
rs •—

O' ....

o o o o

I I

O' O

i/^ Wn ^ ^

I I I I I
t\ sO O 00

D Z
CO.

O r-i c\ sD r^)
^ O' 00 O'

I I I I I
0,1

O O ^ lo VO

Z .4^ I
o V ^
N z c.

< U1

oo^Q

I

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T5
C

CO

'I' vr^ O +

0<CQ<CQU

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i S

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o o

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T5 ^

iz

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(Z) cQ
c"?

8 i

Of;:

o —
X '

n2

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■55 g

o w

u!t3

x|

bC^
O C
03 2

I g

cd 4-1
DO '55

c-n
js t
£ o
XZ

O

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X D)
2 <D)

W X
c/) 2
^ a
qn

O

o
C

a o

i'c

h”

S'

Vo

O'

sandy loam (Olson, 1962).

^Bhir is the Podzol B that is dark with humus (h) and iron oxide (ir).
3gm of oven-dried roots per ft. ^ X 1 in. thick, average per horizons.

1967-68]

Olson and Hole — Fragipan in Soils

181

0)

S

pH

4.4- 6. 5

4. 3-6. 2

4.9-5. 7

4. 5- 6. 5

4.8-6. 1

1 w

D H

Q Z D

UJ < J UJ

%

0.2-1 .6

1. 0-2.0

0.9-1. 7

1.3-2. 7

0. 3-2.0

8l“q

” H 5

%

tr-0 . 03

tr-0.04

0.02-0.04

0.01-0.03

tr-0.06

Na 2CO 3
Extract-
able
S1O.2

%

0.03-0.38
0.1 -0.05
0.02-0.06
0.02-0.07

0 . 04-0 . 08

Base

Satura¬

tion

%

23-67

8-50

19-49

35-67

30-67

(/} -

Na

ro ro ro

00000

1 1 1 1 1

00000

z

0

H

<

U

UJ

me/lOOg

0. 1-8.0
0. 1-0.2
0. 1-0.3
0. 1-0.3
0. 1-0.2

<

U-'

0

Z

<

z

u

X -

M

2

0.2-13.5

Tr-0.4

0. 1-1.3
0.4-3. 2

0. 1-T.6

UJ

Ca

1.0-29.5
0. 8-2.6
0.8-3. 4
0.7-4. 3

0. 7-6.0

C.E.C.

me/lOOg

6- 75

7- 18
4-10
4-12
2-15

D

%

1.2-15.6

1 . 0-5 . 3
0.3-1 .5

0. 1-1 . 1
Tr-0.3

Hori¬

zons

A .

Bhir. . . .

A’x .

B’x .

C .

182 Wisconsin Academy of Science, Arts and Letters [Vol. 56

extensive clay films. Eluviation of some clay from the lower sequum,
particularly in the A'2 horizon, and distribution of remaining clay
as “bridges” between skeletal grains improved the texture and fab¬
ric with respect to pan formation, in many of the soils studied.
No definite evidence was found of chemical cementation of the frag-
ipan by silica, alumina or iron oxide. Laboratory experiments sug¬
gested that clay and possibly iron oxide are cementing materials.
The wide dissemination of a cementing material through the hori¬
zon would seem to be a prerequisite to brittleness.

Proposed Genetic Sequence

It is possible that the first major horizonation to develop in these
soils was the slight textural differentiation of the lower sequum
into A'2 and B'2 horizons, the first of which included the zone now
occupied by the upper sequum. This differentiation took place dur¬
ing a moist period and may have reached a climax when the lower
sequum had settled and become compact enough to be somewhat
impervious to eluviation and root penetration, except along polyg¬
onal shrinkage cracks. These may have formed by desiccation dur¬
ing the relatively warm-dry period of 6,000 to 4,000 years ago
(Smith, 1965), and may have been enlarged by eluviation since
then. The upper Podzol (Spodosol) sequum may represent a state
of equilibrium with the northern hardwood-hemlock forest during
the last two thousand years. Disturbance of the forest by lumber¬
ing during the past century has apparently weakened the Podzol
sequum at many sites (Milfred et ah, 1967).

It is proposed that the fragipan is most extensive in northeast¬
ern Wisconsin because (1) textural heterogeneity of initial drift
materials was commonly favorable for development of compact
horizons, (2) the maximum effective precipitation has been in the
northeast, where the resulting notable percolation of water through
the soil in the early stages of pedogenesis and particularly after
periods of drouth hastened eluviation of fines and settling of the
subsoil into a compact state, and (3) the subsoil was colder than
in more southern landscapes and therefore had unfavorable tem¬
peratures for root growth and related faunal activity both before
and after compaction of the fragipan.

Tree-Throw and Perching of Water
Related to the Fragipan

The mottling in the fragipan suggests that an early stage of
development of perched gley, known in Germany as pseudogley
(Miickenhausen, 1956) has been reached. While digging trenches
in these soils in early spring, the writers have observed seepage of

1967-68]

Olson and Hole — Fragipan in Soils

183

water over the surface of the fragipan into the trenches. In studies
of the under surfaces of root masses of wind-throvv^n trees, par¬
ticularly of large maple trees (Acer saccharum), we have noted
extensions below the horizontal roots (Figures 1 and 2) of a polyg¬
onal arrangement of vertical sheets of small roots delineating cells
about 10 cm deep and 10 cm across. This diameter matches the aver¬
age diameter of the polygons observed in the upper part of the
B'2x horizon (Figure 2).

The frequency of tree-throw on forest soils of northern Wis¬
consin, as evidenced by the presence of about 140 cradle-knolls per
acre in some areas (Milfred, et ah, 1967), is attributable to the
restriction of root growth caused in part by the fragipan, and in
considerable part by the cool temperatures prevailing in the subsoil.

The perching of water over the fragipan creates reducing con¬
ditions in saturated horizons, and can be expected to increase sur¬
face runoff in wet seasons, particularly in early spring. Seepage
water moves laterally downslope over the pan, and has been ob¬
served to initiate slumping of subsoils in highway cuts. Some seep¬
age water also has moved vertically down through prism joints
and a few porous inclusions in the fragipan.

Literature Cited

Ableiter, J. K., and F. D. Hole, 1961. Soil Survey of Bayfield County, Wis¬
consin. Series 1939. No. 39. U.S.D.A.

Beaver, Albert J. 1963. Characteristics of some Bisequal Soils in the Valderan
Drift Region of Eastern Wisconsin. M.S. Thesis, University of Wisconsin,
Madison.

- . 1966. Characteristics and Genesis of some Bisequal Soils in Eastern

Wisconsin. Ph.D. Thesis, Universty of Wisconsin, Madison.

Buol, S. W., and D. M, Fadness. 1961. New Method of Impregnating Fragile
Material and Thin Sections. Soil Sci. Soc. Am. Proc. 25:253.

Carlisle, F. J., E, G. Knox, and R. B. Grossman. 1957. Fragipan Horizons
in New York Soils: I. General Characteristics and Distribution. Soil Sci.
Soc. Am. Proc. 21:320-321.

Carroll, Paul H. 1959. Genetic and Mophological Characteristics of Gray
Wooded Soils in Taylor County, Wisconsin. Unpublished Report, U.S.D.A.,
S.C.S., Madison, Wisconsin.

Committee on Terminology. 1960. Supplementary Report of Definitions ap¬
proved by the Committee on Terminology. Soil Sci. Soc. Am. Proc. 24:
235-236.

Curtis, J. T. 1959. The Vegetation of Wisconsin. The University of Wiscon¬
sin Press, Madison 53706.

Daniels, R. B., W. D. Nettleton, R. J. McCracken, and E. E. Gamble. 1966.
Morphology of Soils with Fragipans in Parts of Wilson County, North
Carolina. Soil Sci. Soc. Am. Proc. 30:376-380.

Day, P. R. 1956. Report of the committee on physical analysis, 1954-55. (Pro¬
cedure for particle size analysis by the hydrometer method.) Soil Sci. Soc.
Am. Proc. 20:167-169.

184 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Gardner, D. R., and E. P. Whiteside. 1952. Zonal Soils in the Transition
Region between the Podzol and Gray-Brown Podzolic Regions in Michi¬
gan. Soil Sci. Soc. Am. Proc. 16:137-141.

Grossman, R. B., F. B. Fehrenbacher, and A. H. Beavers. 1959. Fragipan
Soils of Illinois: 1. General Characterisctics and Field Relationships of
Hosmer Silt Loam. Soil Sci. Soc. Am. Proc. 23:65-70.

Hole, F. D., and K. 0. Schmude. 1959. Soil Suiwey of Oneida County, Wis¬
consin. Bui. 82. Wis. Geol. and Nat. Hist. Survey, University of Wiscon¬
sin, Madison.

- , G. W. Olson, K. 0. Schmude, and C. J. Milfred. 1962. Soil Survey

of Florence County, Wisconsin. Bui. 84. Wis. Geol, and Nat. Hist. Survey,
University of Wisconsin, Madison.

Jackson, M. L. 1956. Soil Chemical Analysis — Advanced Course. Pub. by the
author. Dept, of Soils, University of Wisconsin, Madison.

- . 1958. Soil Chemical Analysis. Prentice-Hall, Inc. Englewood Clilfs,

N. J.

Jha, P. P., and M. G. Cline, 1963. Morphology and Genesis of a Soil Brun
Acid with Fragipan in Uniform Silty Material. Soil Sci. Soc. Amer. Proc.
27:339-344.

McCracken, R. J., and S. B. Weed. 1963. Pan Horizons in Southeastern Soils:
Micromoi-phology and associated chemical, mineralogical and physical
properties. Soil Sci. Soc. Am. Proc. 27 : 330-334.

Milfred, C. J., G. W. Olson, and F. D. Hole. 1967. Soil Resources and Forest
Ecology of Menominee County, Wisconsin. Bui. 85. Wis. Geol. and Nat.
Hist. Survey, University of Wisconsin, Madison.

Muckenhausen, E. 1956. Die Einteilung der Wasserbeeinflussten (Hydromo-
phen) Boden Deutschlands. 6th Congr. InPl. Soil Sci. Soc. E: 111-114.

Nygard, I. J., P. R. McMiller, and F. D. Hole. 1952. Characteristics of some
Podzolic, Brown Forest and Chernozem Soils of the Northern Portion of
the Lakes States. Soil Sci. Soc. Am. Proc. 16:123-129.

Olson, G. W. 1962. Morphology and Genesis of Fragipans of some Soils of
Northeastern Wisconsin. Ph.D. Thesis, University of Wisconsin, Madison.

Ranney, Richard W. 1966. Soil Forming Processes in Glossoboralfs of West-
Central Wisconsin. Ph.D. Thesis, University of Wisconsin, Madison.

Smith, P. W. 1965. Recent Adjustments in Animal Ranges. In H. E. Wright,
Jr., and D. G. Frey. The Quaternary of the United States. 633-642. Prince¬
ton University Press, N. J.

Soil Survey Staff. 1951. Soil Survey Manual. U.S.D.A. Handbook. No. 18.

- . 1960. Soil Classification: A Comprehensive System; The Seventh Ap¬
proximation. U.S.D.A.

Thwaites, F. T. 1943. Pleistocene of part of Northeastern Wisconsin. Bui.
Geol. Soc. Amer. 54:87-144.

Whitson, A. R. 1927. The Soils of Wisconsin. Wisconsin Geological and Natu¬
ral History Survey Bui. 68, University of Wisconsin, Madison.

Yassoglou, N. j., and E. P. Whiteside. 1960. Morphology and genesis of some
Soils containing Fragipans in northern Michigan. Soil Sci. Soc. Am. Proc.
24:396-407.

A COMPARISON OF RED CLAY GLACIO-LACUSTRINE
SEDIMENTS IN NORTHERN AND EASTERN WISCONSIN’

G. W. Petersen, G. B. Lee and G. Chesters

Abstract

Selected red, clayey, glacio-lacustrine sediments from the south¬
ern shores of Lake Superior in northern Wisconsin and from the
Fox and Wolf river drainage basins in eastern Wisconsin were
compared because field observations suggested that a similarity
exists between these sediments, and also between the soils formed
from them.

Close similarities were found in the particle size distribution of
the carbonate-free clay fractions, and in the types and amounts of
mineral species in each clay fraction, supporting the theory that the
finer fractions of the sediments in both areas were derived from
a common source and were deposited in a similar manner. The
source of the noncarbonate clay fraction is most likely the Lake
Superior basin which is believed to have discharged fine-grained
sediments into the Lake Michigan basin during the retreat of Cary
ice. Differences between the sand, silt, carbonate, iron oxide and
feldspar contents of the sediments from the two areas are likely
due to the influence of local bedrock and earlier drift. The lower
carbonate content of deposits in northern Wisconsin may account
for the greater leaching depth and degree of podzolization observed
in these soils.

Introduction
Present Investigation

Deposits of red, clayey, glacio-lacustrine sediments, interjacent
with Valders till and other drift deposits, are found along the Fox
and Wolf River Valleys of eastern Wisconsin and to a lesser extent

Contribution from the Department of Soils, University of Wisconsin, Madison.
Published with the approval of the Director, Wisconsin Ag-ricultural Experiment Sta¬
tion. Supported in part by a grant under Title IV of the National Defense Education
Act of 1958.

* Mr. Petersen is a former Research Assistant ( now Assistant Professor, Department
of Agronomy, Pennsylvania State University, University Park, Pennsylvania) and
MESSRS. Lee and Chesters are Associate Professors of Soils, University of Wisconsin,
Madison.

185

186 Wisconsin Academy of Science, Arts and Letters [Vol. 56

along the western shores of Lake Michigan. Similar deposits are
located along the shores of Lake Superior in Minnesota, Wisconsin
and Michigan (Fig. 1).

The calcareous nature of these deposits and their similarity in
color and texture suggests that their fine fractions were derived
from a common source, namely, the Lake Superior Basin (Murray,

100

H=

200

Northern red clays
Eastern red clays

SCALE IN MILES |

Scale 1:2,500,000

Figure 1. Sample sites and clayey drift areas. Glacial till and glacio-lacustrine
deposits are not differentiated.

1967-68] Petersen — Glacio-Lacustrine Sediments

187

1953). However, detailed analyses of the mineralogy and the types
and amounts of carbonates and free iron oxides in the silt and clay
fractions have not been reported. Because of the pedological impli¬
cations of these characteristics, and their influence on the genesis
and classification of soils formed from these deposits, an investiga¬
tion of fine-textured, glacio-lacustrine sediments from eastern and
northern Wisconsin was undertaken.

Previous Investigation

The most recent glacial till deposits in eastern Wisconsin (desig¬
nated “Valders’' by Thwaites, 1943 and more recently “Valderan”
by Frye and Willman, 1960) consist mainly of calcareous, reddish-
brown clay to clay loam or loam till (Lee, et al, 1962). Valders till
differs from the Cary till in this area in having a reddish-brown
rather than a yellowish-brown color; in part of the region the
Valders till has a much higher clay content. The interval of deglaci¬
ation between Cary and Valders advances is marked by the Two
Creeks Forest Bed, whose radiocarbon age is 11,850 db 100 years
before present (Broecker and Ferrand, 1963).

Alden (1918) described the red drift of eastern Wisconsin as a
dense calcareous clay, the bulk of which was silt and clay of such
fineness that it could have been transported long distances in sus¬
pension. The high concentration of silt and clay has led several
investigators to suggest that the red sediments are lacustrine de¬
posits (Chamberlin, 1873-77, p. 214-228), or glacial till deposits
that were derived from lake clays (Alden, 1918; Thwaites, 1943;
Murray, 1953). These red sediments may also have been partially
derived from the earlier Cary deposits that were eroded and re¬
deposited by the Valders ice sheet (Murray, 1953). However, the
high silt and clay content and red coloration, attributed to the pres¬
ence of hematite both as discrete particles and particle coatings
(Murray, 1953), suggests that these red sediments were derived
from the red sandstones and shales of the Lake Superior region
(Alden, 1918). Murray (1953) suggested that as Cary ice retreated
silts and clays from the Superior basin (Glacial Lake Keweenaw)
were siphoned into the Lake Michigan basin (Early Lake Chicago)
and the Green Bay lowlands. With the advance of Valders ice the
silts and clays were eroded, mixed with other materials, trans¬
ported to surrounding uplands and deposited as glacial till. As
Valders ice retreated, melt waters in front of the ice formed large
glacial lakes into which silts and clays were deposited (Thwaites,
1943; Murray, 1953). Thus the drift in eastern Wisconsin includes
glacial till and glacio-lacustrine deposits, as well as glacio-fluvial
and eolian sediments. Many of the red clay deposits in northern
Wisconsin are also of glacio-lacustrine origin (Ableiter and Hole,

188 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1961) and probably were deposited in glacial Lake Duluth follow¬
ing the ice retreat in that region (Leverett, 1929; Murray, 1953).

Experimental Procedures
Source of Samples

Reddish-brown, calcareous, glacio-lacustrine clays (Fig. 1) are
the parent materials of certain ‘‘red clay’’ soils in northern and
eastern Wisconsin and adjacent states. The topography of these
soils is level or slightly undulating. In contrast to the red clay soils
of the eastern part of the state, those of the north are leached to a
greater depth and are more strongly podzolized. In both areas red
clay lacustrine soils may be interjacent with soils formed in clayey
glacial till, glacio-fluvial sediments, beach deposits or dunes.

The two pairs of sediment samples chosen for detailed investiga¬
tions are described in Table 1. One pair (designated Oshkosh I and
II) was selected to represent the glacio-lacustrine parent materials
of the Oshkosh soils of eastern Wisconsin (Lee et al., 1962). The
other pair (designated Hibbing I and II) was selected on the basis
of field observations and other preliminary investigations to repre¬
sent parent materials of Hibbing, Ontonagon and related soils in
northern Wisconsin. All samples were obtained below soil sola and
appeared to be unaffected by pedalogic weathering.

Analytical Techniques

The samples were dispersed and their sand (2,000 to 50 /x), silt
(50 to 2 fi) and clay (< 2/x) contents determined as outlined by
Day (1956).

table 1. Location and Description of Sample Sites

Sample

Location

Depth,

In.

Color*

Texture

Eleva¬

tion,

Ft.

Hibbing I

SEX SWX, sec. 31, T. 47
N., R. 5 W., Bayfield
County .

32 to 44

2.5YR 4/4

silty clay

675

Hibbing 1 1

NWXNWX. sec. H.T. 47
N., R. 5 W., Bayfield
County .

30 to 48

2.5YR 4/4

clay

900

Oshkosh 1

NWXSWX, sec. 33, T. 20
N., R. 17 E., Winne¬
bago County .

35 to 51

5YR 5/3

clay

750

Oshkosh 1 1

SWX SWX, sec. 18, T. 22
N., R. 17 E., Outagamie
County .

19 to 45

5YR 5/3

clay

790

Munsell color system.

1967-68] Petersen — Glacio-Lacustrine Sediments

189

Following sample dispersion, the sand fraction was separated
by filtration through a U. S. Standard 300 mesh sieve. Separation
of the silts and clays was achieved by sedimentation (Janke, Ph.D.
Thesis, Univ. of Wisconsin, 1962) ; in this method carbonates and
iron oxides were not removed prior to fractionation. The clay frac¬
tions were further separated into coarse (2.0 to 0.2/x), medium (0.2
to 0.08/x) and fine (<0.08/x) particle size ranges (Kittrick and
Hope, 1963), and leached of salts with water and acetone.

The carbonate percentage of each of the sand, silt and clay frac¬
tions was determined by treatment with excess acid and the quan¬
tity of acid required to neutralize the carbonates was determined
by titration (USDA Handbook 60, 1954). Because this procedure
does not differentiate between carbonates, carbonate contents were
expressed as CaCOs.

Iron oxides were extracted from the separated fractions by a
citrate-dithionite technique (Aguilera and Jackson, 1953) and de¬
termined by the colorimetric orthophenanthroline procedure (Jack-
son, 1958, p. 389-390).

Clay suspensions, dried on glass slides to obtain parallel-
orientation (Jackson, 1956, p. 183-188), were analyzed on a North
American Philips X-ray diffractometer- equipped with copper tar¬
get and nickel filter. After removal of iron oxides, clay fractions
were magnesium saturated and solvated with 10 percent aqueous
glycerol, to expand the d-spacing of montmorillonite to 18 A. This
was followed by potassium saturation to collapse montmorillonite
to 14.2 A and some of the vermiculite from 14.2 to 10 A. The terms
montmorillonite and vermiculite, as used here, refer to specific
mineral species as differentiated by their lattice swelling character¬
istics. Following potassium saturation, the samples received suc¬
cessive heat treatments at 350 to 540 °C. Heating the samples to
350 °C collapsed any remaining vermiculite to 10 A. Chlorite was
identified by a 14.2 A d-spacing that persisted through each of the
heat treatments and was enhanced by the 540 °C treatment. Mica
was identified by a 10 and 5 A d-spacing that persisted throughout
the heat treatments. Interstratihed clay minerals, which exhibit
average d-spacings of 24 to 32 A rather than a spectrum of sharply-
defined peaks diagnostic of a mixture of discrete mineral species,
collapsed to 10 to 14 A on heating. Quartz was indicated by 4.1 and
3.35 A d-spacings that persisted throughout the heat treatments.

Because silt fractions contain only small amounts of expansible
layer silicates, no attempt was made to expand them prior to X-ray
diffraction analysis, nor were heat treatments deemed necessary.
The silt fractions (50 to 2/x) were ground to <5/x and slides were

2 The authors express their g-ra^titude to Dr. S. W. Bailey of the University of Wis¬
consin Geology Department for the use of X-ray diffraction facilities.

190 Wisconsin Academy of Science, Arts and Letters [Vol. 56

prepared by aqueous glycerol solvation. Feldspars were identified
by their 3.24 and 3.18 A diffraction peaks; carbonates were distin¬
guished by the 3.03 A d-spacing for calcite and 2.88 A for dolomite.

Interpretation

Investigations were initiated to test the hypothesis that the red,
clayey sediments of northern and eastern Wisconsin are derived
from a common source (Murray, 1953). These red sediments are
probably transported silt and clay-sized fragments from red sand¬
stone and shales of the Lake Superior region (Alden, 1918). The
sediments should not be identical, however, because of the influ¬
ences of local bedrock and older drift in supplying materials like
carbonates and feldspars to the sediments.

Evidence that the samples collected at the Hibbing and Oshkosh
sites (Fig. 1) are typical of glacio-lacustrine deposits follows:
Field estimates of texture indicate that the parent materials of
Hibbing and related soils such as Ontonagon and Pickford have
a high content of clay and a relatively low sand content. Analysis
of a C2 horizon of an Ontonagon soil from Ontonagon County,
Michigan, (Soil Survey Laboratory Staff, 1952) showed this ma¬
terial to contain 66.5 percent clay, 21.3 percent silt and 12.2 percent
sand. Analysis of the C2 horizon of a less well drained soil formed
from similar deposits (Pickford), also in Ontonagon County, Mich¬
igan (Soil Survey Laboratory Staff, 1952), showed it to contain
66.8 percent clay, 28.8 percent silt, and 4.4 percent sand. Carbon¬
ate content of this soil was 13 percent. Field observations made
in the Valders drift region of eastern Wisconsin indicate that
the parent materials of Oshkosh soils typically consist of reddish-
colored calcareous, clays or silty clays. Laboratory analysis of 28
samples (Lee, et al., 1962) showed that 26 of them ranged in clay
content from 40 to 82 percent and that half of them contained in
excess of 60 percent clay. All samples contained less than 5 percent
sand. Average carbonate content for 26 of the samples (Janke,
Ph.D. Thesis, University of Wisconsin, 1962) was 27.5 percent;
most samples ranged from 20 to 30 percent.

The particle size distribution of samples selected for detailed
study are shown in Tables 2 and 3. When these samples are placed
in the USD A Textural Classification (Soil Survey Staff p. 205-
213), the Hibbing I sample is placed in the silty clay textural class
because of its relatively high silt content ; the other three samples
are classified as clays. Each of the samples is characterized by a
high clay and low sand content, although the Hibbing II sediment
contains appreciably more sand than the other three sediments.
The Oshkosh samples contain about one and a half times as much
coarse clay (2 to 0.2/x) as the Hibbing, while the contents of medium

1967-68] Petersen — Glacio-Lacustrine Sediments

191

(0.2 to 0.08/x) and fine clay (<0.08/x) were very similar in all
samples when determined as a percentage of the intact sediment
(Table 3). The similarities in particle size distribution are even
more striking when expressed as percentages of the total clay
fraction ; the Hibbing samples contain slightly less coarse clay, and
slightly more fine clay than the Oshkosh samples. Thus the hypo¬
thesis that the northern and eastern red clay sediments of Wiscon¬
sin are derived from a common source, but are not identical be¬
cause of the influence of local source materials is feasible and
will be tested more rigorously in this discussion.

The carbonate content of all fractions except the fine clay (Table
4) show that the Oshkosh samples are generally more calcareous
than the Hibbing. Expressed as a percentage of the particular frac¬
tion, the coarse clays and silts of the Oshkosh sediments contain
in excess of 30 percent carbonate. The carbonate contents of the
medium and fine clay fractions are much lower than those of the
silts and coarse clays. A possible explanation of this phenomenon
is the increased solubility of the finer-sized carbonates resulting
from their greater surface area. Increased solubility of carbonates

Table 2. Particle Size Distribution of Sediment Samples

Sample

Sand

(2000 TO 50 fx)

Silt

(50 TO 2 fx)

Clay

(<2m)

%

%

%

Hibbing I .

3.2

41.8

55.0

Hibbing II .

12.4

32.6

55.0

Oshkosh I .

0.2

28.8

70.9

Oshkosh II .

3.0

22.0

74.9

Table 3. Particle Size Distribution of Clay Fractions
OF Sediment Samples

Coarse Clay

Medium Clay

Fine

Clay

Sample

2.0 TO

0.2 m)

(0.2 TO

0.08 m)

(<o.

■ o

00

1=

A

B

A

B

A

B

or

/o

%

%

%

%

%

Hibbing I .

32.4

58.9

14.0

25.5

8.6

15.6

Hibbing 11 .

32.0

58.2

14.3

26.0

8.7

15.8

Oshkosh I .

45.3

63.8

17.9

25.2

7.7

10.9

Oshkosh II .

48.3

64.5

16.8

22.4

9.8

13.1

A Expressed as a percentage of the total sediment sample.
B Expressed as a percentage of the total clay fraction.

192 Wisconsin Academy of Science, Arts and Letters [Vol. 56

in cold glacial lake waters also causes a more rapid dissolution of
the finer-sized carbonate particles.

The fact that the parent sediments of Hibbing soils contain less
than half as much carbonate as the sediments from which Oshkosh
soils are formed (Table 5) may have pedological significance. Field
observations of these soils indicate that Hibbing soils are leached
(of carbonates) to greater depths and exhibit evidences of podzoli-
zation, such as albic tonguing (stripping of oxide coatings from
soil particles) to a greater degree than do Oshkosh soils.

The carbonate data were used to determine if the particle-size
distribution of the sediments from the two areas are more nearly
alike when determined on a carbonate-free basis. The data in Table
5 shows that the noncalcareous clay (<2/x) contents of the sedi¬
ments from the two areas are almost identical. Also the distribu¬
tion of clays (Table 6) into the three particle-size ranges are very
similar: 56 it 1 percent coarse clay, 28 zh 2 percent medium clay
and 15.5 zt 1.5 percent fine clay expressed on the basis of the total
clay fraction. The elimination of carbonate from the particle-size
distribution improves greatly the similarity in distribution of clay

Table 4. Content of Carbonates (as CaCOs) in Sediment Samples

Sample

Carbc

)NATE (as CaC

30 3) IN

Sand

Silt

Coarse

Clay

Medium

Clay

Fine Clay

A

B

A

B

A

B

A

B

A

B

%

%

%

%

%

%

%

%

%

%

Hibbing I .

0.5

15.4

8.0

19.2

4.3

13.3

0.5

3.5

0. 1

1.7

Hibbing 11....

2.4

19. 1

6.2

18.9

3.2

10. 1

0.6

4.0

<0.1

0.2

Oshkosh I .

0. 1

52.5

9.8

33.9

16.2

35.8

2.2

12.6

<0. 1

0.8

Oshkosh 11....

2.4

78.5

8.2

37.3

17.5

36.2

1.6

9.3

0.5

4.8

A CaCO 3 expressed as a percentage of the total sediment sample.
B CaCOs expressed as a percentage of the particular size fraction.

Table 5. Contents of Noncalcareous Sand, Silt and Clay and Carbonate
(as CaCOs) IN Sediment Samples

Sample

Sand

Silt

Clay

CaCo3

%

/o

%

%

Hibbing I .

2.7

33.8

50.1

13.4

Hibbing II .

10.0

26.4

51.1

12.5

Oshkosh I .

0. 1

19.0

52.5

28.4

Oshkosh II .

0.6

13.8

55.4

30.2

1967-68] Petersen — Glacio-Lacustrine Sediments

193

size particles for the two areas (compare Table 2 with 5 and 3
with 6).

The iron oxide percentages of the sediment separates are shown
in Table 7. When iron oxides within individual size fractions are
expressed as percentages of the total sediment sample, these per¬
centages are partially a reflection of the relative sample weights
of each fraction within that sample. Therefore, it is also desirable
to examine the iron oxide percentages within a particular size frac¬
tion when making comparisons between samples. Expressed as a
percentage of the total sediment, the iron oxide percentages found
in the sand fraction are extremely low; however, the Oshkosh I
sample is 9% iron oxide probably in the form of concretions. The
iron oxide percentages of the silt, coarse clay and medium clay
fractions of the Hibbing sediments are somewhat higher than those

Table 6. Particle Size Distribution of Clay Fractions of Sediment
Samples Expressed on a Carbonate*-Free Basis

Sample

Coarse

i Clay

Medium Clay

Fine Clay

A

B

A

B

A

B

%

%

%

%

%

%

Hibbing I .

28. 1

56.1

13.5

26.9

8.5

16.9

Hibbing II .

28.8

56.4

13.7

26.8

8.6

16.8

Oshkosh I .

29. 1

55.5

15.7

29.9

7.6

14.5

Oshkosh II .

30.8

55.6

15.2

27.5

9.3

16.8

*A11 carbonate expressed as CaCO 3.

A Expressed as a percentage of the total sediment sample.
B Expressed as a percentage of the total clay fraction.

Table 7. Content of Iron (as Fe^Os) in Sediment Samples

Iron (as Fe20 3) in

Sample

Sand

Silt

Coarse

Clay

Medium

Clay

Fine Clay

A

B

A

B

A

B

A

B

A

B

%

%

%

%

%

%

%

%

%

%

Hibbing I . .

0.02

0.78

0.63

1.51

0.60

2.14

0.40

2.96

0.03

0.39

Hibbing II .

0.07

0.53

0.38

1.16

0.92

3.20

0.44

3.24

0.03

0.39

Oshkosh I . .

0.02

8.87

0.24

0.84

0.38

1.31

0.34

2.17

0.04

0.46

Oshkosh 1 1 .

0.05

1.63

0.23

1.06

0.39

1.26

0.33

2. 18

0.03

0.37

A Ee 2O 3 expressed as a percentage of the total sediment sample.
B Fe20 3 expressed as a percentage of the particular size fraction.

194 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Table 8. Content of Iron (as Fe203) in Carbonate* -Free
Sediment Samples

Iron (as Fe20 3)

IN

Sample

Sand

Silt

Coarse

Clay

Medium

Clay

Fine

Clay

A

B

A

B

A

B

A

B

A

B

%

%

%

%

%

%

%

%

%

%

Hibbing I . .

0.02

0.92

0.68

1.87

0.63

2.47

0.40

3.06

0.03

0.40

Hibbing 1 1 .

0.07

0.66

0.41

1.43

0.95

3.55

0.44

3.37

0.03

0.39

Oshkosh I . .

0.02

18.67

0.27

1.27

0.45

2.04

0.35

2.48

0.04

0.46

Oshkosh 1 1 .

0.05

7.58

0.25

1 .69

0.47

1.97

0.34

2.40

0.03

0.39

*All carbonates expressed as CaCOa.

A Fe 2O 3 expressed as a percentage of the total sediment sample.
B Fe 2O 3 expressed as a pereentage of the particular size fraction.

of the Oshkosh sediments ; the fine clay fractions are very similar.
When the iron oxide percentages are calculated on a carbonate
free basis (Table 8), the iron concretions in the sand fractions of
the Oshkosh sediments become more apparent and slightly more
iron oxides are present in the coarse and medium clay fractions of
the Hibbing samples, which may be attributed to their closer prox¬
imity to the iron source of these red sediments.

X-ray diffractograms (summarized in Table 9) indicated similar
clay minerals in the Oshkosh and Hibbing sediment samples in each
of the clay fractions. These similarities are particularly evident in
the fine and medium clay fractions. The fine clay fractions of all
the sediment samples consist predominately of montmorillonite
with some interstratified material and small amounts of mica,
whereas the medium clays consist of montmorillonite, with meas¬
urable amounts of mica, vermiculite, chlorite and some interstrati¬
fied materials. Both the Hibbing and Oshkosh coarse clay fractions
are comprised of mica, vermiculite, chlorite, montmorillonite, in¬
terstratified materials, quartz, feldspars, calcite and dolomite. Al¬
though some differences exist in diffraction peak intensities between
the same size fractions of different samples, this is not necessarily
indicative of differences in quantitative mineralogy since peak in¬
tensities are controlled by crystal orientation and degree of
crystallinity as well as concentration.

Diffractograms of silt fractions showed them to consist largely
of quartz, feldspars, calcite and dolomite, with some chlorite and
mica. Hibbing silt fractions showed slightly more intense feldspar

Table 9. Semi-Quantitative Mineral Compositions of Clay and Silt Fractions of Sediments

1967-68]

Petersen — Glacio-Lacustrine Sediments

>

I I I I III

+ + + + + + + 4-

+ + + +

I i I

I 1 I

I I I I I I I

++++ ++++ INI
++++ +

I I 1 ++++ +S++ i:+ +

+ +++

+ + +

I 1 I

+

+ +

I 1 I I

++-+ 1 I

++++ +++

(- (-■ CO </)

.5 .5 o o

— — CO CO

HOO

bJD bC-C _C

.5 .S o o

X!

X X X X

— CO CO

:n:roo

bX) bCX X

C r CO

h h o o
X X X X
XXXX

— — CO CO

xxoo

bc bcx X

r- CO CO

X .5 O O

XXXX

XXXX

— — CO CO

i:xoo

>-

_co

U

(U

C

£

u

iS

CJ

u

195

*Mt = ]Vlontmorillonite; Vt = Vermiculite; Chi = Chlorite; Int = Interstratified; Fids = Feldspars; Carb = Carbonates.
- = Undetected ; tr= Trace; +=Low; + + =Moderate; + + -b = Abundant ; + -h + += Dominant.

196 Wisconsin Academy of Science, Arts and Letters [Vol. 56

peaks and the Oshkosh samples more intense carbonate peaks,
which is likely indicative of the effect of local drift and bedrock.

Because calcite is dissolved during- the citrate-dithionite pre¬
treatment of clay samples to remove free iron oxides (Petersen
et al., 1966), water smears of untreated clay fractions were ana¬
lyzed by X-ray diffraction. Insufficient carbonates were present in
the fine and medium clays to show characteristic diffraction peaks
of either dolomite or calcite. However, water smears of the coarse
clay fractions showed characteristic peaks of both minerals, being
more intense in the Oshkosh samples. This corresponds to the in¬
tensities of carbonate peaks in the various silt fractions and sup¬
plies further evidence for the local addition of carbonates.

References Cited

Ableiter, J. E. and F. D. Hole. 1961. Soil survey of Bayfield County,
Wisconsin. 77 p.

Aguilera, N. A. and M. L. Jackson. 1953. Iron oxide removal from soils and
clays. Soil Sci. Soc. America Proc., v. 17, p. 359-364.

Alden, W. C. 1918. Quaternary geology of southeastern Wisconsin. U.S. Geol.
Survey Prof. Paper 106, 356 p.

Broecker, W. S. and W. R. Farrand. 1963. Radiocarbon age of the Two
Creeks Forest Bed, Wisconsin. Geol. Soc. America Bull., v. 74, p. 795-802.
Chamberlin, T. C. 1873-77. Geology of eastern Wisconsin: Geology of Wis¬
consin. v. 2, p. 219-228.

Day, P. R. 1956. Report of the Committee on Physical Analyses, 1954-55. Soil
Sci. Soc. America Proc., v. 20, p. 167-169.

Frye, J. C. and H. B. Willman. 1960. Classification of the Wisconsin Stage in
the Lake Michigan glacial lobe. Illinois Geol. Survey Circ. 285, p. 1-16.
Jackson, M. L. 1956. Soil chemical analysis — advanced course. Published by
the author. Dept, of Soils, Univ. of Wis., Madison, Wisconsin, 991 p.

- . 1958. Soil chemical analysis. Prentice-Hall, Englewood Cliffs, N. J.,

498 p.

Kittrick, j. a. and E. W. Hope, 1963. A procedure for the particle-size
separation of soils for x-ray diffraction analysis. Soil Sci., v. 96, no. 5,
p. 319-325.

Lee, G. B., W. E. Janke and A. J. Beaver. 1962. Particle-size analysis of
Valders drift in eastern Wisconsin. Science, v. 138, no. 3537, p. 154-155.
Leverett, E. 1929. Moraines and shore lines of the Lake Superior basin. U.S.

Geol. Survey Prof. Paper 154, p. 1-72.

Murray, R, C. 1953. The petrology of the Cary and Valders tills of north¬
eastern Wisconsin. Am. Jour. Sci., v. 251, p. 140-155.

Petersen, G. W., G. Chesters and G. B. Lee. 1966. Quantitative determina¬
tion of calcite and dolomite in soils. Jour. Soil Sci., v. 17, no. 2, p. 328-338.
Soil Survey Staff, USDA. 1951. Soil Survey Manual. USDA Handbook 18.

U.S. Govt. Printing Office, Washington, D. C. 503 p.

Thwaites, F. T. 1943. Pleistocene of part of northeastern Wisconsin. Geol.
Soc. America Bull., v. 54, p. 87-144.

U.S.D.A. 1952. Field and laboratory data on some Podzol, Brown Podzolic,
Brown Forest, and Gray-Wooded Soils in northern U.S. and southern
Canada. Soil Survey Laboratory Memo. No. 1, Beltsville, Md.

LIGHT PENETRATION STUDIES IN THE MILWAUKEE
HARBOR AREA OF LAKE MICHIGAN'

Carroll R. Norden
Department of Zoology
University of Wisconsin — Milwaukee

Introduction

Biologists have long recognized the importance of light in natural
waters and its relationship to biological productivity. A number of
papers have been published bearing on light penetration, such as
that of Birge and Juday (1929) which concerned submarine illu¬
mination in a number of Wisconsin lakes, the work of Sauberer
(1939) on several Alpine lakes, and Strickland’s (1958) review of
solar radiation in the oceans. Of particular interest in this study
was the work of Chandler (1942) on light penetration and its rela¬
tion to turbidity in Lake Erie, as well as the studies by Beeton
(1958, 1962) on light transmission in the Great Lakes,

Five stations were visited in this investigation which were
located in Lake Michigan, near the Milwaukee harbor (Fig. 1). It
is at this point that the Milwaukee River empties into Lake Michi¬
gan. Just before the river enters the lake, its waters are joined by
the Kinnickinnic and Menomonee Rivers. All three flow through
the highly industrialized section of southeastern Wisconsin. By the
time their waters reach Lake Michigan, they have been subjected
to a wide variety of influences from farms, cities and industries.

The purpose of this study was to discover the extent to which
the highly turbid waters of the Milwaukee River influences light
pentration in nearby Lake Michigan and to interpret the relation¬
ship of the Secchi disc determinations to photometer measurements.

Equipment and Methods

Light penetration measurements were made with a submarine
photometer, number 268 WA, from the G. M. Manufacturing and
Instrument Corporation. Light intensity was recorded in micro¬
amperes as registered on the microameter in the boat and the micro-
ameter readings were converted to footcandles. The photometer was
calibrated by the Electrical Engineering Department of the Univ-

1 Contribution No. 2, Center for Great Lakes Studies, University of Wisconsin —
Milwaukee.

197

198 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 1. Lake Michigan at Milwaukee, Wisconsin showing the location of the
principal stations.

ersity of Wisconsin using an incandescent tungsten filament lamp.
Paired surface and subsurface light intensity measurements were
made at one meter depth increments at each station and the data
presented are of total visible light.

A standard Secchi disc, with a diameter of 20 centimeters divided

1967-68]

Nor den — Light Penetration Studies

199

into black and white quadrants was used to measure transparency.
The greatest depth in feet at which the Secchi disc was visible was
determined and then the photometer was lowered to that depth and
light intensity measured.

Five stations were established near the mouth of the Milwaukee
River. An attempt was made to visit them at weekly intervals be¬
tween June and November, 1965.

Station 1 was located inside the seawall at the mouth of the
Milwaukee River (Fig, 1), where the depth of the water was 6
meters.

Stations 2 and 3 were located one and one-half and three miles
from shore, east of the mouth of the Milwaukee River. The maxi¬
mum depth at station 2 was 15 meters; at station 3, 24 meters.
Most of the comparative data was obtained at these three stations.

Station 4 was located north of the harbor entrance, about two
miles from shore and in water 17 meters deep.

Station 5 was located south of the harbor entrance, two miles
from shore and in water 11 meters deep. Light penetration meas¬
urements made at stations 4 and 5 showed no significant differences
from those taken at stations 2 and 3.

Except for some special studies on diurnal changes in light in¬
tensity, the data presented in this study were taken on fairly clear,
calm days between the hours of 0800 and 1600.

Weather conditions made it impossible to obtain measurements
as often as desired and prevented all stations from being visited at
weekly intervals. Certain segments of the data were omitted which
were incomplete or of questionable value.

Effect of River Water on Light Penetration

The amount of incident light penetrating to various depths is
greatly reduced in the harbor as compared to that in Lake Michi¬
gan, one and one-half miles from shore (Fig. 2). In the harbor,
penetration of one percent of the incident light to a depth greater
than one meter occurred only in summer. Light penetration in the
fall was drastically reduced. This is believed to be due in part to
the angle of incident light falling on the surface of the water and
in part to the greater turbidity of the water caused by wave action
and heavy rainfall.

Light penetration to the 5 meter depth in the harbor is negligible
(Table 1), whereas about 10 percent of the total visible light pene¬
trates to that depth at station 2 (one and one-half miles from
shore) and nearly 15 percent at station 3 (three miles from shore) .

The transmission of incident light to a depth of 5 meters com¬
pares favorably with measurements reported by Beeton (1962)

200 Wisconsin Academy of Science, Arts and Letters [Vol. 56

for another area of open Lake Michigan. He wrote that the percent
transmission of various wave-lengths of light to the five meter
depth fell approximately between 3.5 percent for the red and 25.
percent for the green. In the present study, the penetration of
total surface illumination ranged from about 7 to 19 percent (Table
1). Light transmission was 10 to 14 percent greater at station 2 in
Lake Michigan during July and August than in western Lake Erie
at the five meter depth during a comparable period (Chandler,
1942).

The curves at station 2 (Fig. 2) tend to be somewhat irregular,
particularly in the fall of the year. This indicates that the column
of water is not optically homogeneous throughtout and may be
caused by concentrations of plankton at certain depths or to tur¬
bidity differences resulting from water movements.

The optical properties of waters can be described by vertical
extinction coefficients (K),

K = 2.30 (log /, h — log / (h + 1) )

I, h and 7 (h + 1) = light intensity at depths h meters and (h
+ 1) meters. The 2.30 compensates for the use of base — 10
logarithms.

% of Incident Light

.01 .1 1 10 100

Figure 2. Relation between depth and total visible light expressed as a per¬
centage of the light falling upon the surface of the water.

1967-68]

Norden — Light Penetration Studies

201

The coefficient indicates the rate of decrease of light as the depth
increases. It is based on Lambert’s Law and represents the percent¬
age of original light held back at each depth.

The average vertical extinction coefficients for the curves in
Figure 2 are given in Table 1. The extinction coefficients for the
turbid waters of the Milwaukee harbor are particularly high, most
of them surpassing those of western Lake Erie (Beeton, 1962),
although not as high as Little Star Lake, Wisconsin (Whitney,
1938).

Light penetration in the waters of Lake Michigan at Milwaukee,
Wisconsin (stations 2 and 3), as indicated from the vertical ex¬
tinction coefficients, was less than those reported by Beeton (1962)
for another area of Lake Michigan. These data (Table 1) suggest
that the turbid waters from the Milwaukee River are influencing
light transmission at one and one-half, and as far as three miles
from shore.

Transparency

Forty-nine Secchi disc measurements were compared with the
photometer readings made at the same depths and are expressed as
the percentage of surface light present at the depth of Secchi disc
extinction. The 22 Secchi disc readings made in the harbor were
consistently shallower than the 27 readings obtained from Lake
Michigan proper (Fig. 3). However, the average percentage of
surface light intensity at Secchi disc depth was 28.1 percent, in
the harbor and 16.5 percent in Lake Michigan. Beeton (1958) re¬
ported 14.7 percent from Lake Huron, whereas Poole and Atkins
(1929) reported 15.8 percent from the English Channel, and Clarke
(1941) gave a value of 15.2 percent for the Atlantic Ocean.

This disparity between percentage of surface light intensity in
the harbor and in Lake Michigan at Secchi disc extinction is un-
doubtably due to the highly turbid waters which are concentrated
at the mouth of the Milwaukee River. Sauberer (1939) reported
similar results from turbid waters. He obtained a greater percent¬
age of surface light at the Secchi disc depth which he attributed to
the suspended materials which caused diffusion and scattering of
light. Chandler (1942) showed that in Lake Erie, Secchi disc meas¬
urements were inversely related to turbidity.

Several investigators (Riley, 1941; Halicki, 1958) have at¬
tempted to derive a value at which Secchi disc reading could be
converted into the depth at which one percent of the surface light,
as determined by photometer measurements, was present. This is
termed the euphotic depth and Strickland (1958) reported that this
should be about 2.5 times the Secchi disc depth. Riley (1941) used

202 Wisconsin Academy of Science, Arts and Letters [Vol. 56

% Surface Light

Figure 3, Relationships of Secchi disc readings to percentage transmission of
surface light at that depth.

a conversion factor of 3 for the Atlantic Ocean and Halicki (1958)
obtained 4.3 for western Lake Erie.

In this study, factors of 3.1 were obtained for Lake Michigan
at stations 2 and 3 (Table 1) . However, a value of 2.1 was obtained
at station 1, in the harbor.

Several authors (Jones and Wills, 1956; Halicki, 1958; Vollen-
weider, 1960; Graham, 1966) have indicated that conversion factors
and values derived from Secchi disc readings are applicable only
within the specific body of water. These data suggest that the

Table 1. Vertical Extinction Coefficients, Percentage of Incident Light at the 5 Meter Depth, Secchi Disc
Readings, and the Euphotic Depth for Stations 1, 2 and 3 in Lake Michigan at Milwaukee, Wisconsin

1967-68]

Norden — Light Penetration Studies

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Time

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0815-1040

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203

204 Wisconsin Academy of Science, Arts and Letters [Vol. 56

harbor water and Lake Michigan water should be treated as distinct
entities in so far as Secchi disc measurements are concerned.

Summary

The higher percentage transmission of surface light intensity at
Secchi disc depth (16.5 percent) as compared with previous reports
for other bodies of water (Poole and Atkins, 1929; Clarke, 1941)
and particularly with Lake Huron (Beeton, 1958) indicates that
Milwaukee River water had some effect on light penetration in
Lake Michigan, at least to a distance of one and one-half miles from
shore.

Vertical extinction coefficients substantiate this conclusion. The
percentage of surface light held back at each depth gradually in¬
creased, from .37, three miles from shore to .44, one and one-half
miles from shore at Milwaukee, Wisconsin. The increase is even
greater in the Milwaukee harbor where the average vertical ex¬
tinction coefficient was 2.29. All three values are higher than that
reported from another area of Lake Michigan (Beeton, 1962).

The percentage transmission of surface light intensity at Secchi
disc depth is greater in turbid water.

Further studies are necessary in order to determine more pre¬
cisely the integration of Milwaukee River water into the waters of
Lake Michigan.

Acknowledgements

The author is indebted to Dr. Arthur T. Tiedemann of the Uni¬
versity of Wisconsin for calibrating the photometer and to Mr.
Donald Martinson and Mr. Gerald Ludwig for assistance in gather¬
ing the data. I should also like to thank Drs. Alfred Beeton and
John Blum of the University of Wisconsin-Milwaukee for their |
comments on the manuscript.

The work was partially supported by the Wisconsin Alumni Re¬
search Foundation.

Literature Cited

Beeton, A. M. 1958. Relationship between Secchi disc readings and light
penetration in Lake Huron. Trans. Am, Fish. Soc., 87 (1957) : 73-79. ;

- . 1962. Light penetration in the Great Lakes, Great Lakes Res. Div.,

Inst. Sci. and Tech., Univ. Mich., Pub. No. 9, pp. 68-76.

Birge, E. A. and C. Juday. 1929. Transmission of solar radiation by the wa¬
ters of inland lakes. Trans. Wis. Acad. Sci,, Arts and Lett., 24: 509-580. If
Chandler, D. C. 1942. Limnological studies of western Lake Erie, IL Light j
penetration and its relation to turbidity. EcoL, 23 (1) : 41-52. |;

Clarke, G. L. 1941. Observations on transparency in the southwestern section
of the North Atlantic Ocean. J. Mar. Res., 4: 221-230.

1967-68]

Norden — Light Penetration Studies

205

Graham, J. J. 1966. Secchi disc observations and extinction coefficients in the
central and eastern North Pacific Ocean. Limnol. and Oceanogr., 11(2):
184-190.

Halicki, P. j. 1958. A comparison of two methods of determining transpar¬
ency in natural waters. Trans. Am. Microscop. Soc., 77 (4) : 428-434.

Jones, D. and M. S. Wills. 1956. The attenuation of light in sea and estuarine
waters in relation to the concentration of suspended solid matter. J. Mar.
Biol. Assoc. U. K., 35: 431-444.

Poole, H. H. and W. R. G. Atkins. 1929-30. Photo-electric measurements of
submarine illumination throughout the year. J. Mar. Biol. Assoc., U. K.,
16: 297-324.

Riley, G. A. 1941. Plankton studies. IV. Georges Bank. Bull. Bingham
Oceanogr. Coll., 7 : 1-73.

Sauberer, F. 1939. Beitrage zur Kenntnis des Lichtklimas einiger Alpenseen.
Int. Rev. Hydrobiol. und Hydrographie., 39: 20-55.

Strickland, J. D. H. 1958. Solar radiation penetrating the ocean. A review
of requirements, data and methods of measurement, with particular refer¬
ence to photosynthetic productivity. J. Fish. Res. Bd. Canada, 15(3):
453-493.

Vollenweider, R. a. 1960. Beitrage zur Kenntnis optischer Eigenschaften
der Gewasser und Primarproducktion. Mem. Int. Ital. Idrobiol. Pallanza.,
12: 201-244.

Whitney, L. V. 1938. Transmission of solar energy and the scattering pro¬
duced by suspensoids in lake waters. Trans. Wis. Acad. Sci. Arts, and
Lett., 31: 201-221.

THE MOVEMENT, RATE OF EXPLOITATION AND
HOMING BEHAVIOR OF WALLEYES IN LAKE
WINNEBAGO AND CONNECTING WATERS,
WISCONSIN, AS DETERMINED BY TAGGING

Gordon R, Priegel
Fishery Biologist
Wisconsin Conservation Division
Oshkosh. Wisconsin

Introduction

The walleye, Stizostedion vitreum vitreum (Mitchill) in Lake
Winnebago and connecting waters is the most sought-for sport fish
especially during the spawning run in the rivers and during the
ice fishing season on Lake Winnebago. Various studies concerning
the walleye in these waters have been initiated to further contribute
knowledge that will lead to improved management practices and
provide for a sustained annual yield in the future. The tagging
study is one phase of this comprehensive program.

The water areas involved in the study include Lake Winnebago
and Big Lake Butte des Morts on the 107-mile-long Fox River and
lakes Poygan and Winneconne on the 216-mile-long Wolf River.
The Wolf River joins the Fox River in Big Lake Butte des Morts,
10 river miles above Lake Winnebago and then enters the lake as
the Fox River at Oshkosh (Figure 1). The Fox River also flows
out of Lake Winnebago at Neenah and Menasha and flows 39
river miles north to Green Bay, Lake Michigan. The runoff water
from 6,000 square miles enters Lake Winnebago.

Lake Winnebago has an area of 137,708 acres with a maximum
depth of 21 feet and average depth of 15.5 feet. The lake is roughly
rectangular in shape : 28 miles long and 10.5 miles wide at its wid¬
est point. The smaller upriver lakes (Poygan, Winneconne and Big
Lake Butte des Morts) have areas of 14,102; 4,507 and 8,857 acres,
respectively. The depths of these smaller lakes are similar with
maximum depths not exceeding 11 feet which are located in the
river channels. All four lakes have many characteristics common
to shallow eutrophic lakes.

Spawning walleyes from Lake Winnebago must migrate through
one or more of the smaller upriver lakes to enter either the Wolf
or Fox rivers to spawn. Walleyes from Lake Winnebago travel as
far as 90 miles up the Wolf River and when water levels permit

207

208 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 1. Lake Winnebago and connecting water areas involved in the tagging
study.

passage over the Eureka dam some 40 miles up the Fox River to
spawn in adjacent grass and sedge marshes.

Objectives !

Objectives of the tagging program which was initiated in Sep- :
tember, 1960, were to obtain information on angler exploitation,
angler exploitation in relation to migration, extent of migration,
times of migration, homing tendencies of spawning fish, effects
of dams, effect of tagging on growth, suitability of various type
tags and effect of entrapment gear on tag recoveries.

1967-68] Priegel — Walleyes in Lake Winnebago

209

The original plan was to tag as many walleyes as possible during
the fall, 1960, in Lake Winnebago and to tag 1,000 walleyes in
Lake Poygan during the winter, 1960-61. The tagging program,
however, was continued through the spring of 1964 and includes
additional tagging areas.

Methods

Capture Methods

On Lake Winnebago during the fall, 1960-62 walleyes were ob¬
tained from commercially fished Lake Erie type trap nets and 45-
foot trawls. During this period, an A.C. shocker unit was also used
during daylight hours to obtain walleyes in areas inaccessible to
commercial fishermen. Trap nets were also set at the mouth of the
Fox River off Oshkosh from January through April, 1964.

All walleyes tagged on marshes adjacent to the Wolf and Fox
rivers were captured with an A.C. shocker unit during the spawn¬
ing period, 1962-63.

On lakes Poygan and Winneconne, walleyes were captured in
commercially fished hoop and trap nets set under the ice from
January-March, 1961 and 1963.

Tagging Methods

The normal procedure for tagging fish was to remove them from
source of entrapment, place in a holding tank, measure them (total
length in inches to the nearest tenth), tag them and release them
in the same approximate area of entrapment.

All of the fish except 994 fish tagged with plastic dart tags
(Yamashita and Waldron, 1958) were marked on the upper jaw
with either monel-metal or aluminum strap tags passing around
the maxillary and premaxillary (Shetter, 1936). Eschmeyer and
Crowe (1955) demonstrated no statistically significant difference
in the rate of recovery among walleyes tagged in the upper and
lower jaws with No. 3 strap tags.

The plastic dart tags were 0.0625 inch in diameter and 2.5 inches
long. Five colors were used — orange, white, red, green and yellow.
The tags were stamped with a serial number near the distal end of
the shaft. The dart tags were inserted into the epaxial musculature
immediately below the spiney dorsal fin where the %-inch barb
pierced through the interspinous bones so when the tag was tugged
the barb contacted the interspinous bones.

Recapture Methods

Recaptures of tagged walleyes were reported voluntarily by
anglers and commercial fishermen ; no rewards were offered. Fish¬
ermen were alerted to the presence of tagged walleyes by the local

210 Wisconsin Academy of Science, Arts and Letters [Vol. 56

press, radio, television and posters at boat liveries, resorts and
public access points. To stimulate combined cooperation, all reports
of recapture were acknowledged with a form letter giving locality,
date and length of fish at tagging. In addition to the recaptures
reported by fishermen, tagged walleyes were recaptured by project
personnel during field operations which included electrofishing on
the spawning marshes.

Results

The number of walleyes tagged and the number recovered each
succeeding year after tagging are shown in Table 1. Of 14,885
tagged in the five years, 3,237 or 21.8% have been reported caught
by anglers during a six-year span, 1961-66. Recoveries the first year
after each tagging period were consistently the highest.

April was the peak month for tag returns from anglers every
year, 1961 through 1966, except in 1962, when May was the peak
month. January and February were also high tag return months
in 1961 and 1964 (Figure 2).

In addition to angler returns 372 tagged fish were recaptured
by private and state commercial fishing crews in nets and trawls
and by project personnel with electro gear. All of these fish were
returned to the water after the length, tag number, date and local¬
ity of capture were recorded. Anglers eventually reported catching
90 (24.2%) of these 372 fish.

The size of the tagged walleyes ranged from 10.2 to 28.6 inches
in total length with 49.2% falling in the 15- to 19-inch groups
(Table 2). Smaller-sized walleyes were available, but the intent
was to tag only walleyes over 10 inches which were assumed more
vulnerable to the angler. For example, only 2.5% of the 10-inch
group and 6.5% of the 11-inch group were recaptured while for
lengths 12 through 26 inches the return ranged from 17.3 to 30.0%
for each one-inch group.

In Lake Winnebago, 102 recaptured walleyes were measured by
project personnel after they had carried monel jaw tags over one
growing season. Their growth during this period is compared with
the average annual increment of untagged fish in the population
(Priegel, in press) in Table 3. Although the percentage of the
normal increment attained by the tagged fish varied vddely, the
average of 53.7 for the group suggests a marked retardation of
growth as a result of the presence of monel jaw tags. Several in¬
vestigators (Rose, 1949; Smith, Krefting and Butler, 1962; Patter¬
son, 1953; and Eschmeyer and Crowe, 1955) have shown that the
presence of jaw tags tends to retard the growth rate of walleyes.
Retardation of growth was also noted in the Lake Winnebago
walleye,

Table 1. Number and Percentage (in parentheses) of Tagged Walleyes Reported by Anglers in Lake Winnebago

AND Connecting Waters, 1960-66

1967-68] Priegel — Walleyes in Lake Winnebago

. O

sD

— — ro

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211

*Months are numbered consecutively from January (1) to November (11).

'’*Less than 0.05 percent.

fFall tagging (9, 10, II): First year extends from date of tagging through December 31 of the following year.

FREQUENCY OF CAPTURES (PERCENT)

212 Wisconsin Academy of Science, Arts and Letters [Vol. 56

TIME (MONTHS)

Figure 2. Walleye returns by months for the years 1961-66.

Anglers returned 418 (35.3%) of the 1,183 No. 4 Monel tags used
which was superior to the return of No. 3 monel tags (19.4%).,
aluminum strap tags (21.5%) and plastic dart tags (23.0,%). The
serial numbers on the monel tags were easily distinguished while
on the aluminum strap tags serial numbers were difficult to distin-

1967-68] Priegel — Walleyes in Lake Winnebago

213

Table 2. Length Frequency of Tagged and Angler Recaptured Walleyes
IN Lake Winnebago and Connecting Waters

Length Groups in
Inches (T.L.)

Tagged Fish

Recaptured Fish

Percentage
Captured
FOR Length
Groups

Number

Percent

Number

Percent

10.0-10.9 .

239

1.6

6

0.2

2.5

11.0-11.9 .

1,117

7.5

73

2.3

6.5

12.0-12.9 .

1,180

7.9

204

6.3

17.3

13.0-13.9 .

1,200

8.1

265

8.2

22.1

14.0-14.9 .

1 ,222

8.2

279

8.6

22.9

15.0-15.9 .

1,469

9.9

357

11.0

24.3

16.0-16.9 .

1,663

11.2

390

12.0

23.4

17.0-17.9 .

1,378

9.3

266

8.2

19.3

18.0-18.9 .

1,406

9.4

345

10.7

24.5

19.0-19.9 .

1,395

9.4

372

11.5

26.2

20.0-20.9 .

1,154

7.7

325

10.0

28. 1

21.0-21.9 .

705

4.7

184

5.7

26. 1

22.0-22.9 .

421

8.9

105

3.2

24.9

23.0-23.9 .

204

1.4

42

1.3

20.6

24.0-24.9. . .

84

0.6

15

0.5

17.9

25.0-25.9 .

33

0.2

6

0.2

18.2

26.0-26.9 .

10

0. 1

3

0.1

30.0

27.0-27.9 .

4

*

28.0-28.9 .

1

*

Total .

14,885

3,237

21.8

*Less than 0.05 percent.

guish because the digits were embossed and wore down within one
year. Many of the aluminum strap tags were paper-thin when re¬
turned to us so it was conceivable that after one year considerable
tag loss could occur.

There was no evidence of tag loss for plastic dart tags and all
fish examined showed that the plastic dart tags were solidly em¬
bedded. Some infection around the tag was reported by the anglers
but never observed by project personnel. Plastic dart tags are easy
to apply, but the legend became difficult to distinguish after one
summer season. The tags used had a serial number near the end of
the shaft; however, it would have been more beneficial to have a
serial number at the end of the shaft near the barb to enhance
distinguishing the legend. The lighter colors (orange, white and
yellow) are preferred (as against green and red) as it is easier to
distinguish the legend.

Tagging during the fall of 1960 and 1962 on Lake Winnebago
provided sufficient data to evaluate the use of trap nets, trawls and
an A.C. shocker unit as means of capturing walleyes for tagging
studies. Of the 10,691 walleyes tagged during this period, trawling

214 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Table 3. Growth of Tagged Walleyes Recovered
After One Growing Season

LeNGI'H
Groups in
Inches

Number

OF

Fish

Average Total
Length (Inches)

AT Time of

Averagi

MENTS (

i Incre-
Inches)

Increment of
Tagged Fish
(Percentage
OF Increment
OF Untagged
Fish)

Tagged

Fish

Un¬

tagged

Fish

Release

Recovery

12.0-12.9. . . .

5

12.6

13.4

0.8

1.8

44.4

13.0-13.9. . . .

1

13.3

14.4

1 . 1

1.8 -

61.1

14.0-14.9. . . .

4

14.3

15.4

1.1

1.3

84.6

15.0-15.9. . . .

6

15.6

16. 1

0.5

1.2

41.6

16.0-16.9. . . .

6

16.7

16.9

0.2

1.2

16.6

17.0-17.9. . . .

14

17.6

17.9

0.3

1.0

30.0

18.0-18.9. . . .

14

18.6

19. 1

0.5

1.0

50.0

19.0-19.9. . . .

20

19.5

20.2

0.7

0.8

87.5

20.0-20.9. . . .

11

20.4

20.8

0.4

0.8

50.0

21.0-21.9....

9

21.5

21.7

0.2

0.6

33.3

22.0-22.9. . . .

6

22.5

23.0

0.5

0.4

125.0

23.0-23.9. . . .

6

23.5

23.8

0.3

0.4

75.0

Total or
weighed av¬
erage .

102

53.7

accounted for 1,555 (14.6%), trap netting for 4,087 (38.2%) and
electrofishing for 5,049 (47.2%).

The angler return of walleyes tagged while electrofishing was
25.5% and it was 23.5% for walleyes captured in trap nets. Only
6.4% of the walleyes taken by trawling gear were returned by
anglers. The pressure on a few walleyes exerted by 200-800 pounds
of commercial species when the trawl is lifted, the expansion of the
swim bladder and the handling of the fish during tagging opera¬
tions most likely resulted in a substantial mortality of trawl-caught
walleyes used for tagging. 1

Hopp’s Marsh and marshes near the city of Berlin on the Fox '
River and Spoehr’s Marsh, Hortonville Marsh and Colic Slough on
the Wolf River were electrofished to recover tagged walleyes during
the spring, 1961-66. Fifty-six tagged walleyes were recovered on ,
these spawning marshes, 16 on Fox River marshes, and 40 on Wolf '
River marshes. Of the 16 recovered on Fox River marshes, nine
had originally been tagged on Hopp’s Marsh while the other seven ,
were tagged on Lake Winnebago, Four of the 40 recaptured on
Wolf River marshes were originally tagged on Spoehr’s Marsh, i
while 8, 5 and 23 were tagged on lakes Poygan, Winneconne and
Winnebago, respectively. Angler returns indicate that same pat¬
terns with fish originally tagged in Lake Winnebago being recap¬
tured in both the Wolf and Fox rivers (Table 4) ; however, angler i

Table 4. The Number and Percent (in Parentheses) of Angler Returns By Recapture Location

1967-68] Priegel — Walleyes in Lake Winnebago

215

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216 Wisconsin Academy of Science, Arts and Letters [Vol. 56

returns from the Wolf River were 13.1% as compared to 5.6% from
the Fox River. Only three (0.9%) of the 315 angler returns from
fish originally tagged in lakes Poygan and Winneconne, were re¬
captured in the Fox River, as compared to 182 (59.1%) from the
Wolf River.

Of the 12,822 walleyes tagged and released in Lake Winnebago,
anglers reported capturing 2,828 of which 2,061 (73.9%) were
reported taken in Lake Winnebago (Table 4.) The remaining 767
recaptured walleyes were taken in Lake Poygan (1.1%), Lake
Winneconne (4.2%), Big Lake Butte des Morts (1.2%), the Wolf
River (13.1%) and Fox River (5.6%) and below the outlet dams
at Neenah and Menasha (1.9%). Walleyes were recaptured
throughout the year in these water areas connecting into Lake
Winnebago.

Anglers reported capturing 141 walleyes of 725 originally tagged
and released in Lake Poygan, of which 89 (62.4%) were taken in
the Wolf River during the spawning migration. Only 20 (14.9%)
were recaptured in Lake Poygan.

Of the 781 walleyes tagged and released in Lake Winneconne,
anglers reported capturing 174 of which 94 (54.0%) were reported
taken in the Wolf River during the spawning migration. The num¬
ber recaptured in Lake Winneconne was 24 (13.8%) with 36
(20.7%) being recaptured in Lake Winnebago.

On the Fox River marshes, 322 walleyes were tagged. Anglers
reported recapture of 76, with 44 (58.0%) being caught in the
Fox River and 21 (27.6%) being caught in Lake Winnebago. Nine
(11.8%) were recaptured in Big Lake Butte des Morts and one
each in Lake Poygan and below the Neenah-Menasha dams.

On the Wolf River marshes, 235 walleyes were tagged, with
anglers reporting recapture of 18 of which 13 (70.5%) were taken
on the Wolf River. Two were recaptured in Lake Winneconne and
one each in lakes Winnebago, Poygan and Big Lake Butte des
Morts^

Migration of walleyes out of Lake Winnebago into the upriver
lakes and rivers during the late fall and winter was expected but
the extent was unknown. During tagging operations in January
and February, 1961 on lakes Poygan and Winneconne, 12 walleyes,
previously tagged in Lake Winnebago during the fall, 1960, were
taken while in January and February, 1963, nine walleyes previ¬
ously tagged in Lake Winnebago during the fall, 1962, were taken
in commercially fished nets. Angler returns of walleyes tagged in '
Lake Winnebago during the fall of 1960, 1961 and 1962 and taken
through the ice in the upriver lakes during the following winter
were 19.9, 3.3 and 4.7% respectively of the total annual returns
from the upriver lakes. Angler returns also indicate that of the |

1967-68] Priegel — Walleyes in Lake Winnebago

217

walleyes tagged in lakes Poygan and Winneconne, only 14.9% and
13.8%, respectively, are caught in these lakes. Angler returns of
Lake Poygan tagged walleyes were from the Wolf River (62.5%)
and Lake Winnebago (9.2%) and for Lake Winneconne 54.0%
were from the Wolf River and 20.7% from Lake Winnebago. Net
and angler returns would indicate a sufficient migration of wall¬
eyes out of Lake Winnebago during the late fall and winter into
the upriver lakes.

Following the first year after tagging, 28 walleyes that were
tagged and released in Lake Winnebago were caught by anglers
below the outlet dams of Neenah and Menasha. Water levels were
unusually high during the spring of 1961, and may account for
this migration over the dams the first year after tagging. For the
entire six-year period 57 tagged ymlleyes were reported taken by
anglers below the outlet dams.

The average distance traveled for 2,559 walleyes that were
originally tagged in Lake Winnebago and for which exact locations
of recaptures were known was 18.8 miles. The maximum distance
traveled was 97 miles from Oshkosh, Lake Winnebago to Leeman,
Wolf River. Of 2,559 recaptures, 340 (13.3%) were taken within
the same area as tagged, 226 (8.8%) were within two miles of the
tagging site, 298 (11.6%) were within 2 to 5 miles, 607 (23.7%)
5 to 10 miles, 550 (21.5%) from 10 to 25 miles and 538 (21.0,%)
from 25 to 97 miles. The average distance traveled for 115 wall¬
eyes from Lake Poygan, 143 from Lake Winneconne, 70 from Fox
River marshes and 17 from Wolf River marshes, was 28.9, 28.2,
22.1 and 33.2 miles, respectively.

Eschmeyer (1942) recovered four walleyes tagged in the Norris
Reservoir at an average distance of 4.8 miles. Most of these tagged
in Houghton Lake, Michigan, by Carbine and Applegate (1946)
were recovered at an average distance of 3 miles, but three had
gone 130 miles downstream. Doan (1942) recovered 22 specimens
in western Lake Erie, most of them about 20 miles away but one
had gone 200 miles to the east end of the lake. The average distance
of travel at Lac la Rouge, Saskatchewan (Rawson, 1957) was 3.5
miles for 281 recaptures with one specimen going upstream 65
miles. The general pattern of rather limited movement in walleye
populations with a few long-distance wanderers does not apply
to the Lake Winnebago walleye population because of the distance
traveled during spawning migrations.

During the course of the study, nine fish originally tagged in
Lake Winnebago were recaptured in Lake Puckaway, a distance of
68 river miles from Lake Winnebago. These fish had to pass over
four low-head dams in the Fox River : Eureka, Berlin, White River
and Princeton. One walleye tagged during April, 1963, in Hopp's

218 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Marsh, Fox River was also recaptured in Lake Puckaway during
June, 1963.

Discussion and Significance

The return of 3,237 or 21.8% of the 14,885 tagged walleyes by
anglers over a six-year period definitely demonstrates the effective¬
ness of angler exploitation in this large and extensive water area
especially when one considers the voluntary return. There were no
closed seasons nor minimum size limits in effect for walleyes during
the study period. Herman (1947) reported a recovery of 9.3% of
3,694 walleyes tagged from 1944-46 in the Wolf River, Wisconsin,
after three years ; however, at that time, the season was closed from
February 1 until after the peak of spawning in April and a 13-inch
minimum size limit was in effect during this period. Patterson
(1953) after one year reported a recovery of 20.5% for 984 wall¬
eyes tagged in Escanaba Lake, Wisconsin, where there was a 100%
creel census, no closed seasons, no minimum size limit and no bag
limits during this period. Hubley and Jergens (1959) recovered
5.7% of the 1,784 walleyes tagged in the spring of 1958 within
seven months after tagging in a 40-mile stretch of the Upper
Mississippi River. Eschmeyer and Crowe (1955) reported that
from the grand total of 11,354 walleyes that had been jaw-tagged in
Michigan during the period 1939-52, 12.2% were recovered. In
Blackduck River, Minnesota, sport and commercial fishermen re¬
turned 25.1% of 4,697 walleyes tagged in 1949 after three years
(Smith, Krefting, and Butler, 1952).

Angler returns were consistently higher the first year after
tagging for each tagging period and location (Table 1). Lack of
returns two or more years later is probably due to tag loss and fish
mortality because of high first-year returns in different years.

The length frequency of walleyes when tagged and at the time
of angler recapture is based on the size at tagging, as the error in
using lengths provided by the anglers when the fish was captured
is too great or in many cases the length was not provided by the
angler. Walleyes over 12 inches were more vulnerable to the anglers
(Figure 3). The fact that 44% of the angler returns occurred in
April and May during the spawning migration accounts in part for
the greater vulnerability of larger size walleyes. .The average size
of male walleyes at maturity is 12.7 inches for females it is 17.3
inches (Priegel, in press).

Frequently the question regarding the taking of female walleyes
during the spawning migration before they had a chance to spawn
comes up for discussion. Tag returns from anglers which provided
date of capture during the spawning period on the Wolf River in
1961, 1962 and 1963, were tabulated from ice-out to May 1 to de-

1967-68] Priegel — Walleyes in Lake Winnebago

219

TOTAL LENGTH CINCHES)

Figure 3, Length frequency of tagged (solid line) and angler recapture
(broken line) walleyes in Lake Winnebago and connecting waters.

termine when the majority of female walleyes were caught— -before
or after spawning. All fish over 19 inches are considered females as
determined from age and growth studies (Priegel, in press). The
percent of tagged female walleyes taken after the peak spawning
period was 68.5, 84.2 and 80.7 for the years 1961, 1962 and 1963,

220 Wisconsin Academy of Science, Arts and Letters [Vol. 56

respectively (Table 5). Based on these tag returns it is reasonable
to conclude that proportionately more untagged females are taken
also after the peak spawning period. The same situation was noted
for male walleyes as the percent of tagged male walleyes taken
after the peak spawning period was 62.6, 89.2 and 87.1 for the
years 1961, 1962 and 1963 respectively.

Angler exploitation of tagged walleyes was consistently higher
during the spawning migration period than during the non-
migratory season. April and May in 1961 through 1965 were the
months during the spawning migration while in 1966, March and
April were used because of the early spring breakup (Table 6).
Angler returns of tagged walleyes during the spawning migration
ranged from 33.3% of the total in 1961, to 63.9% of the total in
1966.

January and February were also high tag return months in 1961
and 1964, due to intensive winter angling pressure on Lake Winne¬
bago, and the availability of walleyes tagged during the fall. The
periods of highest tag returns coincide with the best fishing months
and periods of heaviest fishing pressure.

There is currently no closed season on walleyes in Lake Winne¬
bago and connecting waters; however, in the future, if a closed
season would be essential to preserve the walleye fishery, a closed
season during April and May would be most beneficial. April and
May were consistently the high tag return months during the
study period, 1961-66.

The tendency of the walleye to return to specific spawning areas
in lakes and streams has been noted by several investigators:
Stoudt, 1939; Stoudt and Eddy, 1939; Eschmeyer, 1950; Smith,
Krefting and Butler, 1952; Eschmeyer and Crowe, 1955; Rawson,
1957 ; Olson and Scidmore, 1962 ; and Crowe, Karvelis and Joeris,
1963. All observed that stream-spawning walleyes tagged on specific
spawning grounds tended to return to them. The tendency for
spawning walleyes to return to the spawning area where they had
been marked in previous years, or at least utilize the same major
river was also noted in the Lake Winnebago area. On Hopp’s Marsh,
Fox River, 9 of 13 recaptures taken while electro-fishing during the
spawning period were originally tagged and released on Hopp’s
Marsh. On Spoehr’s Marsh, Wolf River, 4 of 27 recaptures taken
while electrofishing during the spawning period were originally
tagged and released on Spoehr’s Marsh. None of the 322 walleyes
tagged during the spawning period in 1962 and 1963 on Fox River
marshes were ever recaptured by anglers or project personnel in
the Wolf River or adjacent marshes although 24% were returned
by anglers from Lake Winnebago and the Fox River. A single fish
was returned from Lake Poygan. None of the 235 walleyes tagged

Table 5. Tagged Walleyes Taken During the Spawning Season in the Wolf River 1961-63.
Fish Over 19 Inches are Considered Females. Peak Spawning is in Parenthesis

1967-68] Priegel — Walleyes in Lake Winnebago

221

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222 Wisconsin Academy of Science, Arts and Letters [Vol. 56

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1967-68] Priegel — Walleyes in Lake Winnebago

223

on Wolf River marshes .were ever recaptured in the Fox River or
adjacent marshes although 8% were returned from the Wolf River
and downstream lakes.

The loss of 57 walleyes (1.8% of all tag returns) for the entire
six-yeS'i* period over the outlet dams at Neenah and Menasha must
be considered negligible when considering the large, extensive water
area involved in this study.

Literature Cited

Carbine, W. F,, and Vernon C. Applegate, 1946. Recaptures of tagged wall¬
eyes, Stizostedion v. vitreum (Mitchill), in Houghton Lake and the Mus¬
kegon River, Roscommon County, Michigan. Copeia., 2:97-100.

Crowe, W. R., Ernest Karvelis, and Leonard S, Joeris. 1963. The movement,
heterogeneity and rate of exploitation of walleyes in Northern Green Bay,
Lake Michigan, as determined by tagging, Internat. Comm. N, W. Atl,
Fish., Spec, Publ. No, 4:38-41.

Doan, Kenneth H. 1942. Some meteorological and limnological factors in the
abundance of certain fishes in Lake Erie. Ecol. Monog., 12:293-314.
Eschmeyer, Paul H, 1950. The life history of the walleye Stizostedion vit¬
reum vitreum (Mitchill) in Michigan. Mich. Dept. Cons., Inst. Fish. Res.,
Bull. No. 3, 99 pp.

- , and Walter R. Crowe. 1955. The movement and recovery of tagged

walleyes in Michigan, 1929-1953. Mich. Dept. Cons., Inst. Fish, Res., Misc.
Publ. No. 8, 32 pp.

Eschmeyer, R. W. 1942. The catch, abundance and migration of fishes in Nor¬
ris Reservoir, Tennessee, 1940. J. Tenn. Acad, Sci., 17:90-115.

Herman, Elmer F. 1947. Notes on tagging walleyes on the Wolf River, Wis.
Cons. Bull., 12 (4) :7-9.

Hubley, Raymond C., Jr., and Glenn D. Jergens. 1959, Walleye and sauger
tagging investigations on the Upper Mississippi River, West Central Area
Invest. Memo No, 1, Wis. Cons. Dept., 9 pp.

Olson, Donald E., and Warren J. Scidmore. 1962. Homing behavior of spawn¬
ing walleyes, Trans, Amer. Fish, Soc., 91:355-361,

Patterson, Donald L. 1953. The walleye population in Escanaba Lake, Vilas
County, Wisconsin. Trans. Amer. Fish. Soc., 82:34-41.

Rawson, D. S. 1957. The life history and ecology of the yellow walleye, Sti¬
zostedion vitreum, in Lac la Ronge, Saskatchewan. Trans. Amer. Fish.
Soc., 86:15-37.

Rose, Earl T. 1949. The population of yellow pike-perch (Stizostedion v. vit¬
reum), in Spirit Lake, Iowa. Trans. Amer. Fish. Soc., 77:32-41.

Shetter, David S. 1936. The jaw-tag method of marking fish. Pap. Mich. Acad.
Sci., 21:651-653.

Smith, Lloyd L., Jr., Laurits W. Krefting, and Robert L. Butler. 1952.
Movements of marked walleyes Stizostedion vitreum vitreum (Mitchill)
in the fisherey of the Red Lakes, Minnesota, Trans. Amer. Fish. Soc., 81 :
179-196.

Stoudt, Jerome H. 1939. A study of the migration of the walleyed pike (Sti¬
zostedion vitreum) in water of the Chippewa National Forest, Minnesota.
Trans. Amer. Fish. Soc., 68:163-169.

— — — , and Samuel Eddy. 1939. Walleye pike tagging study, 1937; 1938, Chip¬
pewa National Forest. Trans. 4th N. Am. Wildl. Conf, :305-310.
Yamashita, Daniel T., and Kenneth D. Waldron. 1958. An all-plastic dart-
type fish tag. Calif. Fish and Game, 44:311-317.

THE TAXONOMY AND ECOLOGY OF LEECHES
(HIRUDINEA) OF LAKE MENDOTA, WISCONSIN

J. A. Sapka/rev
University vo Skopje
Skopje, Yugoslavia

Introduction

Wisconsin, with over 8000 lakes, mainly located in the northern
half of the state, ranks second only to Minnesota in number of
lakes in the United States. Although many studies have been con¬
cerned with the taxonomy and ecology of their biota, there are
certain groups of animals in these lakes on which little research
has been done. Among these are the leeches, which from the sys¬
tematic point of view have been studied for only a few lakes (Mutt-
kowski, 1918; Baker, 1924; Pearse, 1924; Bere, 1931). Questions
related to their spatial and temporal distribution, population densi¬
ties, and life histories have remained almost completely un¬
answered. Such is the case with the leeches found in Lake Mendota.

Lake Mendota is situated in the central portion of southern Wis¬
consin and is the largest of a chain of four lakes in the Yahara
Basin, all of which were formed by modification of the river valley
by glacial activity. Among the morphometric characteristics of
Lake Mendota cited by Birge and Juday (1914) are: maximum
length, 9.5 km ; maximum breadth, 7.5 km ; maximum depth, 25.6 m ;
mean depth, 12.1 m; circumference, 23.4 km; and total surface
area, 39.4 sq. km.

The first to mention the leeches of Lake Mendota was Muttkow-
ski (1918) who noted the following species in his study of the
fauna of the lake:

Erpohdella punctata

Nephelopsis ohscura

Glossiphonia sp.

Another early study which included observations on the leeches
of Lake Mendota was done by Pearse (1924). He mentioned
Piscicola punctate as a parasite of carp, bluegill and large-mouth
black bass and Placobdella parasitica of the bluegill.

225

226 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Methods

From September, 1964 through August, 1965, I had the oppor¬
tunity to study the taxonomy and ecology of the leeches in Lake
Mendota.

A quantitative analysis of Lake Mendota leeches to determine
their vertical and horizontal distributions and seasonal variations
in population density was accomplished by collecting material with
the aid of an Ekman dredge (15 x 15 cm) and a sheet metal frame
(50 X 50 cm) . A transect from Bascom Woods to Governor’s Island
was sampled every month (usually between the 20th and 22nd day)
from September, 1964 through August, 1965. In May, 1965 ten
other transects were sampled also (Figure 1). Samples were taken
from the bottom of the following isobaths of each transect: 0, 1,
2, 3, 4, 6, 8, 10, 12, 14, 16, 18, 20, 22, and 24 meters. Wet and dry
weights measurements were made on a single-pan analytical
balance.

Results

The following 16 species of leeches were identified during the
study :

Glossiphonia complanata
Glossiphonia heteroclita
Batracohdella phalera
Batracohdella picta
Helobdella stagnalis
Helobdella punctata — Lineata
Helobdella lineata
Helobdella elongata
Placob della ornata
Placobdella parasitica
Placobdella montifera
Placobdella sp.

Dina parva
Erpohdella punctata
Nephelopsis obscura
Haemopis marmorata

My list of species of Lake Mendota leeches includes all those
mentioned by previous authors except for Piscicola punctata, which
I did not find. Thus, the fauna of leeches in Lake Mendota could
be represented by seventeen species which belong to four families ;
namely Piscicolidae (1 genus and 1 species), Glossiphonidae (4
genera and 12 species), Erpobdellidae (3 genera and 3 species),
and Hirudinidae (1 genus and 1 species).

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

227

Figure 1. Lake Mendota. Position of transects,

228 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Family Glossiphonidae
1. Glossiphonia complanata (Linnaeus) 1758

This species is known over the whole of North America (Verrill,
1874; Moore, 1901, 1906, 1924, 1952; Nachtrieb, Hemingway and
Moore, 1912; Ryerson, 1915; Baker, 1924; Miller, 1929, 1937;
Bere, 1931; Rawson, 1953; Meyer and Moore, 1954). It is also
found over the .whole of Europe, in Asia (India — Japan — Bering
Islands), and Africa (for example in the Congo). G. complanata
occurs in many habitats but appears to show a preference for lakes
and running water and especially stone bottoms, while it is rarely
found in vegetation. It occurs in oligotrophic lakes, but more fre¬
quently in eutrophic lakes. This study is the first to report this
species in Lake Mendota.

In Lake Mendota G. complanata may be found at almost all times
on the northern and southern shores of the lake, where the bottom
(0-1 meter depth) is covered with boulders, stones and pebbles,
with a mixture of gravel and sand. In these areas it has a popula¬
tion density of 44.4-266.4 individuals per square meter. As may
be seen from Table I, G. complanata may be encountered in some
other habitats in Lake Mendota, such as sand, sand and mud, mud
with detritus, or mud and detritus with vegetation. In such habi¬
tats, it occurs with a smaller population density.

The vertical distribution of G. complanata in this lake is slight
and during the year 1964-65 it was not found at depths of over
one meter (Figure 2) . Such is the case also in the markedly eutro¬
phic Dojran Lake, Macedonia, Yugoslavia (Sapkarev, 1964), eutro¬
phic Fures, Denmark (Bennike, 1943), but not in the markedly
oligotrophic Ohrid Lake or the Prespa Lake, Macedonia, Yugoslavia
(Sapkarev, 1963). In the latter two lakes, G. complanata is a
eurybathic form found at all depths of the lake.

The average, seasonal changes of the population density and
biomass of G. complanata, calculated for the whole littoral zone of
Lake Mendota, are given in Figure 3. From this figure it may be
seen that the population tends to have the lowest density in the
winter and spring months (11.1 individuals per square meter)
and the highest density in the summer and fall months (maximum
in August — 99.9 individuals per square meter). The maximum
population density was associated with the appearance of young
individuals after a reproductive period from the end of May to
the beginning of July. Concurrently, the biomass, in wet and dry
weight, attains its relative maximum in fall (0.93 gr. wet .weight
or 0.20 gr. dry weight per square meter) two or three months
after the appearance of the new generation.

Table I. The Population Density (Individuals Per Square Meter) of Leeches
IN Different Habitats of Lake Mendota

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

229

230 Wisconsin Academy of Science, Arts and Letters [Vol. 56

DEPTH IN METERS

I 3 5 7 9 II 13 15 17 19 21 23 25

Placobdella parasitica
Placobdella montifera
Placobdella ornata
Placobdella sp.

Batracobdel la picta
Glossiphonia complanata
Dina parva

Haemopus nnarmoratis
Nephelopsis obscura
Erpobdella punctata
Helobdella punctata -I ineata
Helobdella lineata
Batracobdella phalera
Glossiphonia heteroclita
Helobdella elongata
Helobdella stagnalis

Figure 2. Bathymetrical distribution of the leeches in Lake Mendota.

Figure 3. Average seasonal changes of the population density and biomass of
Glossiphonia complanata in the littoral zone of Lake Mendota during 1964-65.

Two specimens of G. complanata with eggs were found in the
period between the end of May and the beginning of June. Another
specimen with eggs was found at the beginning of July and still
another specimen with young ones in July. Young individuals were
encountered during all of July and August. A similar reproductive
period for G, complanata was described by Bennike (1943) and
Sapkarev (1946).

I had an occasion to observe G. complanata preying upon Physa
gyrina and Planorhis parvus and on the oligochaeta Limnodrilus.
Muttkowski (1918), on several occasions, found small leeches of
the genus Glossiphonia (I think that these are individuals of G.

1967-68] Sapkarev — Taxonomy and Ecology of Leeches 231

complanata) attached to the underside of the beetle larva,

Paephenus lecontei.

2. Glossiphonia heteroclita (Linnaeus) 1758

This species, like the previous one is quite common in North
America. In addition to other habitats, it has also been found in
lakes such as Nipigon (Moore, 1924), Georgian Bay (Ryerson,
1915), Allequash, Man and Trout lakes (Bere, 1931). It has been
reported in many European lakes also (Pawlowski, 1936; Bennike,
1943 ; Sapkarev, 1963, 1964; Okland, 1964).

G. heteroclita is also identified for the first time in Lake Mendota.
It occurs most frequently in a habitat of mud with detritus and
overgrown with vegetation (11.1-88.8 individuals per square
meter), but is found also in sand with mud, as well as on stones
(11.1-44.4 individuals per square meter). Its vertical distribution
does not extend below a depth of three meters (Figure 4). It is
found frequently at a depth of two meters, and less often in shal¬
lower parts of the littoral zone.

In the course of this study the maximal density of population
occurred in August, as described previously for G. complanata,

I found only one specimen of this leech, on July 10, 1965, with
eggs attached. During August, 1965, 1 found several specimens with
both eggs and young attached to the ventral surface. Bennike
(1943) has found G. heteroclita with eggs and with young from
June to October in Denmark. Thus the period of reproduction ex¬
tends slightly over four months. In central Europe the period of
reproduction appears to last from April to September.

3. Batracohdella phalera (Graf) 1899

Batracobdella phalera is known only throughout the United
States (Graf, 1899; Baker, 1924; Miller, 1929; Bere, 1931) and
Canada (Moore, 1906; Ryerson, 1915). In Wisconsin it can be
found in many of the northeastern lakes (Bere, 1931), in the Lake
Winnebago region (Baker, 1924), and I have now found it in Lake
Mendota. Its distribution in the lake is limited in depth from 0 to 2
meters. With the greatest density of population being 88.8 indi¬
viduals per square meter, it is rather rare in this lake. I found it
most frequently in the Second Point and B'ascom Woods areas. It
occurs in various habitats, but has the greatest density of popula¬
tion in areas having a bottom of mud with detritus and covered
with vegetation.

4. Batracohdella picta (Verrill) 1872

Like the previous species of this genus, B. picta is known in both
the United States and Canada (Verrill, 1872; Moore, 1906, 1952;

232 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Miller, 1929; Bere, 1931). In northeastern Wisconsin it has been
found in many lakes. I have found only one specimen of this leech
in Lake Mendota, between University Bay and Picnic Point at
about a depth of 0.20 meters. Pearse (1924) mentions it as a para¬
site on the sucker and perch in Lake Michigan.

5. Helohdella stagnalis (Linnaeus) 1758

This species is very common and cosmopolitan, being found in
both North and South America, over all of Europe, and in Asia
and North Africa.

One of the reasons for the wide distribution of this leech lies
in its ability to exist in numerous habitats. Bennike (1943) in his
paper, '‘Contributions to the Ecology and Biology of the Danish
Freshwater Leeches'’, writes : "There is only one type of fresh¬
water, in which it has not been found, i.e., the sphagnum bogs;
it has even been found in an extremely dystrophic lake, but in one
locality only (Store okosso).” Thus in Lake Mendota, as in many
other lakes, it may be found in various habitats as can be seen
from Table 1. This means H. stagnalis is an eurjTopic form, but
also an eurybathic form, because its vertical distribution is rather
great — from the shore line to a depth of 12-15 meters. Esrom Lake
(Berg, 1938), Fures Lake (Bennike, 1943) and Prespa Lake (Sap-
karev, 1963) offer similar cases. It is unique in being the only leech
whose range extends throughout all zones in Lake Mendota.

From Figure 5 it is possible to see that H, stagnalis has the
greatest population density in the littoral zone (an average density
of 421.8 individuals per square meter at the 0 meter depth), a
much smaller population density in the sublittoral zone (an aver¬
age of 14.8 individuals per square meter) and the smallest density
in the profondal zone (an average of 3.7 individuals per square
meter at a depth of 12 meters). This is also the case in Esrom
Lake (Berg, 1938), Fures Lake (Bennike, 1943), Dojran Lake
(Sapkarev, 1964) and Borrewant Lake (Okland, 1964).

The vertical distribution of H. stagnalis in the lake as related to
the seasonal change in the density of population may be seen in
Figure 6. From this, one can see that the population density is
greatest in the littoral zone, especially during the summer and
early fall.

The average seasonal change in number and biomass for the
littoral zone only is presented in Figure 7. The maximum popula¬
tion density and biomass of H. stagnalis is found in July and
August (e.g., 1527.4 individuals, 0.83 gr. wet weight or 0.15 gr.
dry weight per square meter in month of July). The minimum
occurs in the winter months (e.g., in February, 22.2 individuals,
0.06 gr. wet weight or 0.01 gr. dry weight per square meter) . The

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

233

10 ind. /m.^
I - 1

200 ind./m.^

I - 1

Figure 4. Average verti¬
cal distribution in the
density of population of
Glossiphonia complanata
in Lake Mendota.

Figure 5. Average verti¬
cal distribution in the
density of population of
Helobdella stagnalis in
Lake Mendota.

occurrence of the maximal population density in the summer, es¬
pecially in the month of July, may be explained by the recruitment
of a new generation. The earliest date that I observed a specimen
with eggs attached to the ventral surface was May 10. In my
collections during the period from May 20 to May 30, I found about
80 percent of the individuals with eggs, 10 percent with young, and
about 10 percent with neither eggs nor young. Individuals captured
during the period from June 1 to June 9 were found in the follow¬
ing state : about 60 percent with young, 30 percent with eggs, and
approximately 10 percent with neither young nor eggs. During the
whole month of July (especially beginning with July 5) I found
free, young individuals of H. stagnalis, while the number of the
old ones was reduced to a minimum. In August samples, all cap¬
tured individuals were of the new generation. This fact may indi¬
cate that a great number of the old individuals die off after the
reproduction period.

234 Wisconsin Academy of Science, Arts and Letters [Vol. 56

500 ind./m. -

Figure 6. Vertical distribution and seasonal change in the density of popula¬
tion of Helobdella stagnalis during 1964-65 in Lake Mendota.

Individuals of H. stagnalis with eggs on their ventral surface
which I captured on May 10, were kept in an aquarium having a
water temperature between 16 and 19 °C. Observations showed
that between May 19 and May 22 some had young on their ventral
surface and by the end of May and the beginning of June all were
with young.

Thus, on the basis of the above, the reproductive period of H.
stagnalis in Lake Mendota appears to cover less than two months
(May and June). In eutrophic Do j ran Lake I have found individ¬
uals of H, stagnalis with young during the months of August and
September (Sapkarev, 1964). Bennike (1943) has found that the
reproductive period of H. stagnalis in freshwater habitats of Den¬
mark extends over more than four months (from May to Septem¬
ber) . An even longer season, from April to September, has been
reported from North America (Castle, 1900) Central Europe (Her-
ter, 1937) and Iceland (Bruun, 1938). Even in the northwestern
part of Iran, the reproductive period has been found to last from
February to June (Bennike, 1940). On the other hand, in the Alps
the reproductive period is much shorter from late July to early
August (Zshokke, 1900).

To determine the horizontal distribution of this leech in the lake,
I took quantitative samples from ten different transects during
May. The results are shown in Figure 8. Except for Morris Park
and Governor’s Island, it was found in all transects. A somewhat
larger population density was found in the vicinities of Bascom
Woods, Mendota Beach, Fox Bluff, and Maple Bluff. The greatest
vertical distribution occurred at Second Point and Yahara Canal.

I was able to observe that H. stagnalis uses Chironomid larvae,
tubificids and Hyalella azteca for food. This leech has been reported
to feed on Chironomid larvae and other small freshwater inverte¬
brates by Pawlowski (1936), Bennike (1943), Mann (1955, 1957)
and Hilsenhoff (1963).

1250

1200'

1 150

I 100-

1050-

1000-

950-

900-

850-

800-

750-

700-

650-

600-

550-

500'

450-

400-

350-

300-

250-

200-

150-

loo¬

se-

0

okarev — Taxonomy and Ecology of Leeches

INDIVIDUALS
WET WEIGHT
DRY WEIGHT

1 — — 1 - j - 1 - ^

II III IV V VI VII VIII

TIME OF YEAR

5000
4800
4600
4400
4200
-4000
-3800
3600
[-3400
3200
3000
2800
2600
h2400
2200
h2000
1800
h 1600
1400
[-1200
1000
800

600

400

200

0

7. Average seasonal changes of the population density
omass of Helobdella stagnalis in the littoral zone of
dendota during 1964-65.

235

MILLIGRAMS PER SQUARE METER

236 Wisconsin Academy of Science, Arts and Letters [Vol. 56

DEPTH IN METERS
ro o 00 o) 4^ ro o

BASCOM WOOD

SECOND POINT

MENDOTA BEACH

fox's bluff

oi

o

MORRIS PARK jO
CATFISH RIVER -L \

• * ro

governor's is.

MAPLE BLUFF

YAHARA CANAL

CAPITOL BLDG.

Figure 8. Horizontal distribution of the density of population of Helobdella
stagnalis in Lake Mendota on the month of May, 1965.

6. Helobdella punctata — lineata (Moore) 1939

This species was described for the first time by J. P. Moore
(1939) on the basis of material from Puerto Rico. My collection
of leeches from Lake Mendota included only a single individual of
this species. It was caught at a depth of two meters, where the lake
bottom was composed of mud and sand with detritus and covered
with vegetation. Because I found just one individual, I was able to
recognize it only by its external morphological characteristics. It
would be useful if more individuals could be found and studied
in greater detail in order to establish whether it is in fact H. punc¬
tata — lineata.

1967-68] Sapkarev — Taxonomy and Ecology of Leeches 237

7. Helohdella lineata (Verrill) 1874

This North American leech is encountered in Lake Mendota very
infrequently. I was able to collect several specimens at a depth of
one to two meters (Figure 2), where the bottom was composed of
stones and gravel, mud and sand, mud with detritus, or mud and
detritus with a cover of vegetation (Table 1). One specimen .with
eggs and two specimens with young were found on August 12.

Baker (1924) writes of it in another Wisconsin lake. Lake Win¬
nebago, referring to it as Helohdella fusca variety lineata.

8. Helohdella elongata (Castle) 1900

Helohdella elongata is found in Canada and the United States
(Castle, 1900; Moore, 1906, 1912, 1924; Ryerson, 1915; Miller,
1929; Rawson, 1930; Bere, 1931). It is mentioned here for the
first time as occurring in Lake Mendota. It was found with greater
frequency than the previous two species of this genus. I found it
together with H. Stagnalis, especially in the Bascom Woods and Pic¬
nic Point transects. Excluding H. stagnalis, this leech has the great¬
est vertical distribution of all leeches in Lake Mendota (Figure 2).
Its range in depth extends from the shore line to four meters, which
would mean that it like all others except H. stagnalis, occurs only
in the littoral zone.

The maximal population density is found along the shore line
on the stoney bottom (177.6 individuals per square meter) with
a biomass of 2.13 gr. wet weight or 0.36 gr. dry weight per square
meter. The population density decreases with increasing depth. It
settles in all kinds of habitats except in the deep lake muds (See
Table 1).

Hilsenhoff (1964) is of the opinion that H. elongata feeds on
tendipeded larvae or small mollusks.

9. Placohdella ornata (Verrill) 1872

For the genus Helohdella as well as for the genus Placohdella,
I have found four species, all of them very rare. This species is
found throughout the rivers, bogs, ponds, and lakes of North Amer¬
ica (Verrill, 1872; Moore, 1901, 1906, 1912; Andrews, 1915; Ryer¬
son, 1915; Miller, 1929; Rawson, 1930; Bere, 1931) and Japan
(Oka, 1917) . In my collection of leeches from Lake Mendota, I have
collected a few specimens along the shore line in University Bay
and Catfish Bay.

10. Placohdella parasitica (Say) 1824

Placohdella parasitica has been found in North America mainly
in bogs, but also in lakes, occurring as a parasite of various ani-

238 Wisconsin Academy of Science, Arts and Letters [Vol. 56

mals. It was observed in Lake Mendota by Pearse (1924) as a
parasite on bluegills, and elsewhere in the vicinity of Madison
(Wingra springs Region) by Cahn (1915). I found just one speci¬
men from the marshy part of University Bay of Lake Mendota.

11. Placohdella montifera (Moore) 1912

Placohdella montifera is distributed throughout the United States
and Canada. It is found in many lakes in these countries; for ex¬
ample in Lake Nipigon (Moore, 1924) , Georgian Bay of Lake Huron
(Ryerson, 1915) , in many of the lakes (oligotrophic and eutrophic)
of northeastern Wisconsin (Bere, 1931). Pearse (1924) mentions
it as parasite on smallmouth black bass in Lake Geneva and on
carp and hackleback sturgeon in Lake Pepin. I collected just one
specimen in Lake Mendota in the area between Catfish Bay and
Governor’s Island.

12. Placohdella sp,

I observed two young specimens of a leech near the Yahara Canal
along the shore line on the stones. Both of them belong to the same
species of the genus Pkicobdella, but I was not able to determine
the species to which they belonged.

Family Erpobdellidae

1. Dina parva (Moore) 1912

The family Erpobdellidae is represented in Lake Mendota by
three species belonging to three different genera. Dina parva is
distributed throughout the United States and Canada (Moore, 1912,
1924), including many Wisconsin lakes (Baker, 1924; Bere, 1931).
In Lake Mendota I encountered it most frequently in the area from
Picnic Point to Mendota Beach together with Erpobdella punctata
and Nephelopsis obscura. The latter two are more abundant than
the former. I found it primarily on a rocky bottom and less fre¬
quently on a bottom of vegetation or sand (Table 1) . It has a very
limited vertical range, which extends in depth from 0 to 1 meters
(Figure 2). The same distribution, density of population, and habi¬
tat for Dina lineata was observed in the macedonian lakes, Prespa
and Dojran (Sapkarev, 1963, 1964).

2. Erpobdella punctata (Leidy) 1870

Erpobdella punctata is widely distributed in North America and
occurs in various habitats including lakes (Moore, 1901, 1924;
Ryerson, 1915; Muttkowski, 1918; Baker, 1924; Bere, 1931; Raw-

1967-68] Sapkarev — Taxonomy and Ecology of Leeches 239

son,, 1953 and others). This species was found in Lake Mendota by
Muttkowski (1918).

In my research, it was the most abundant species among the
Erpobdellidae and .almost always was found together with Nephe-
lopsis obscura.

As is stressed by Muttkowski (1918), optimal conditions for this
species are on the shore margin where coarse gravel and stones
are intermixed. This appears to account for the great abundance
of the species on the northern and southern shores of Lake Mem
dota. From Table 1, one can see that the density of population is
greatest on a stony bottom. Here the population density can attain
a magnitude of 1332.0 individuals per square meter. In July and
August, the period when the young generation appears, the density
can be considerably greater in certain .areas.

As shown in Figure 2, E. pwictata was found at a depth of about
0-1,50 meters. The largest number of individuals occurs at a depth
of 0-0.50 meters with an average of 85,1 individuals per square
meter at a depth of 0.20 meters.

In winter .and early spring E, punctata occurs in small numbers.
In some months during this period, no individuals were found.
From July to October and especially from July to August, a marked
maximum occurred (Figure 9). This maximum density of popu¬
lation is associated with the appearance of a new generation.

Figure 9. Average .seasonal changes of the population density and biomass of
Erpobdella punctata in the littoral zone of Lake Mendota during 19'64-65.

240 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Egg-cocoons have been found from May to August. After that
time only empty cocoons have been found. The length of these
cocoons varies from about two to eight millimeters. The number
of eggs per cocoon is variable, but the largest number of cocoons
were found with two to six eggs. When the young first appear in
the cocoon, their length is about three millimeters, and when they
leave the cocoon they are about five millimeters in length. The
young in the same cocoon were frequently of very different sizes.
Most of the young leave the cocoons in July. The egg-cocoons are
common on the stones of rock and gravel shores to a depth of 0-1
meter. The number of egg-cocoons varies in the different habitats
and localities. I was able to calculate their number per square meter
as 888.0-2220.0. However, in some places the number can be either
much larger or much smaller. The common European species,
namely E, octoculata, has the same period of reproduction (Paw-
lowski, 1936; Berg, 1938; Bennike, 1943; Sapkarev, 1964).

I have observed that Erpobdella punctata uses Oligochaeta and
larvae of Chironomidae for food. Muttkowski (1918) has found
may-fly larvae (Hexagenia, Caenis) and Trichoptera larvae in
Erpobdella punctata.

3, Nephelopsis obscura (Verrill) 1872

Like Erpobdella punctata, this species of the Erpobdellidae is
widely distributed in North America and is found in different
habitats, especially in the stony shore line of lakes (Verrill, 1872;
Ryerson, 1915; Moore, 1924). N. obscura has been reported in
many Wisconsin lakes (Muttkowski, 1819; Baker, 1924; and Bere,
1931) . It was noted for the first time in Lake Mendota by Muttkow¬
ski (1918).

As is shown in Figure 2, the species was found at depths of
0-1.50 meters. The greatest density was found at about 0.20 meters
with an average of 66.6 individuals and a biomass of 1.29 gr. wet
weight or 0.21 gr. dry weight per square meter.

The species was found on almost all dredging days throughout
the year. It is numerous in the summer samples (July and August) ,
where an average number of individuals for the littoral zone was
97.7 per square meter. The corresponding biomass was 0.95 gr. wet
weight or 0.14 gr. dry weight per square meter (Figure 10). The
maximal density of population in July and August was due to the
appearance of a new generation in that period. It seems that this
species has a reproductive period similar to Erpobdella punctata.

Family Hirudinidae

1. Haemopis marmorata (Say) 1824

This single species of the family Hirudinidae in Lake Mendota

1967“68] Sapkarev — Taxonomy and Ecology of Leeches

241

TIME OF YEAR

Figure 10» Average seasonal changes of the population density and biomass of
Nephelopsis obscura in the littoral zone of Lake Mendota during 1964-65.

has been found in the Picnic Point and Pheasant Branch areas.
It is frequently discovered under stones in the shore line or im¬
bedded in the ,wet soil. Near Picnic Point it is found to a depth of
1 meter (Figure 2) .

It has been encountered in some lakes of northeastern Wisconsin
(Bere, 1931) as well as in other parts of the United States (Moore,
1901; Cahn, 1915) and Canada (Ryerson, 1915; Moore, 1924; Raw-
son, 1953).

Discussion

A biocenotic analysis of the leeches in Lake Mendota shows that
all species inhabit the littoral zone (Table 2). For each family, the
number of species decreases with an increase in depth. Only one
species of Glossiphonidae, namely Helobdella stagnalis, was present
in the sublittoral and the upper region of the profundal zone.

It is readily seen from Table 3 that the fauna of Hirudinea of
Lake Mendota is composed mainly of two families of leeches,
namely Glossophonidae and Erpobdellidae, Of the total number of
individuals, 85.7% are in the family Glossophonidae , whereas
17.6% belong to Erpobdellidae, Therefore, Glossiphonidae is the
dominant family from the standpoint of numbers of species and
individuals. During the year (1964-65), Helobdella stagnalis (Glos¬
siphonidae) was dominant in numbers of individuals (75.6%),
while Glossiphonia complanata (Glossophonidae) ranked second
(7.7 % ) . Very few individuals of Helobdella elongata, Batracobdella
phalera and other species of Glossiphonidae were encountered ; less
than 0.9% for each.

242

Wisconsin Academy of Science, Arts and Letters [Vol. 56

i I

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W [ij

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5 g

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Profundal

25

1

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Zones

Depth in meters

Family

Glossiphonidae .

Erpobdellidae .

Hirudinidae .

Total .

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

243

Table III. Relative Dominance of the Species of Leeches in Lake
Mendota Based on the Total Number of Individuals
Encountered During 1964-65

Species

Percent

Family Glossiphonidae . . . .

1. Glossiphonia complanata . .

2. Glossiphonia heteroclita. . . .

3. Batracobdella phalera .

4. Batracobdella picta .

5. Helobdella stagnalis .

6. Helobdella puctata — lineata

7. Helobdella lineata .

8. Helobdella elongata .

9. Placobdella ornata .

10. Placobdella parasitica .

11. Placobdella montifera .

12. Placobdella sp .

8H68
7.66
0.94
0.36
0.01
75.58
0.01
0. 10
0.88
0.10
0.01
0.01
0.02

Family Erpobdellidae

1. Dina parva .

2. Erpobdella pinctata. .

3. Nephelopsis obscura . .

13.77

1.01

7.05

5.71

Family Hirudinidae . .
1. Haemopsis marmorata

0.77

0.55

The average total number and biomass of all Glossophonidae at
various depths in the lake is presented in Figure 11, As may be
seen from this figure, the maximum density of population and
biomass occurs in the littoral zone at a depth of 0 to 3 meters (the
average of a one year period at the 0 meter depth is 558.7 individ¬
uals, corresponding to a biomass 2.13 gr. wet weight or 0.43 gr.
dry weight per square meter).

Figure 12 shows average seasonal changes in population density
and biomass of Glossiphonidae in the littoral zone during 1964-65.
The minimal density of population and biomass occurs in the winter
months (December-February) . The maximum number of individ¬
uals occurs in the summer months (July-August) and is associated
mainly with the appearance of new generations of Helobdella stag¬
nalis and Glossiphonia complanata.

The average vertical distribution in numbers and biomass of
Erpobdellidae in Lake Mendota is presented in Figure 13. Erpobdel¬
lidae has little vertical distribution, occurring only to a depth of
1.50 meters. Maximum density occurs along the shore line (0-0.50
meters) .

244 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 11. Average verti¬
cal distribution in the
density of population of
Glossiphonidae in Lake
Mendota.

The most abundant species of Erpobdellidae are Erpohdella punc¬
tata with 7.0% and Nephelopsis obscura with 5.7% of the total
number of individuals of leeches in Lake Mendota. This means that
the former comes as the third and the latter as the fourth in num-

INDIVIDUALS PER SQUARE METER

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

245

TIME OF YEAR

Figure 12. Average seasonal changes of the population density and biomass of
Glossiphonidae in the littoral zone of Lake Mendota during 1964-65.

ber of individuals of leeches in the lake. The third species of Erpoh-
dellidae, Dina parva, has a relative density of just about one
percent.

The average seasonal changes in population density and biomass
(wet and dry weight) of the Erpohdellidae during the study are
represented in Figure 14.

MILLIGRAMS PER SQUARE METER

246

Wisconsin Academy of Science, Arts and Letters [Vol. 56

100 ind. /m.^

I - 1

Figure 13. Average verti¬
cal distribution in the
density of population of
Erpobdellidae in Lake
Mendota.

The minimal population density and biomass of Erpobdellidae
occurs in the winter months ( January-March) and is probably
associated with the severity of the environment at that season of
the year. The maximal density of population occurs in the summer
months (July-August) and corresponds with the appearance of a
new generation of Erpohdella punctata and NepJielopsis obscura.

Finally, Figure 15 shows the average, vertical distribution of
all leeches in Lake Mendota. It may be seen that: 1) the greatest
density of population and biomass (710.4 individuals, 5.16 gr. wet
weight, or 0.92 gr. dry weight per square meter at a depth of 0

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

247

Figure 14. Average seasonal changes of the population density and biomass of
Erpohdellidae in the littoral zone of Lake Mendota during 1964-65,

meter) is found in the littoral zone; 2) an insignificant number of
individuals (14.8 individuals, 0,05 gr. wet weight, or 0.01 gr. dry
weight per square meter) occurs in the sublittoral zone; 3) in only
a few cases during the study period were several individuals (of
Helobdella stagnalis) found along the rim of the profundal zone
(3.7 individuals, 0.0014 gr. wet weight or 0.0003 gr. dry weight
per square meter) at a depth of 12 meters.

Table 4 shows the average population density and biomass of
the leeches per square meter in the three zones of Lake Mendota,
as well as the average figures calculated for all zones.

Figure 16 shows the vertical distribution of the population den¬
sity of the leeches during the study period. It is possible to see that
the greatest vertical distribution occurs at the end of winter and
the beginning of spring, but that the greatest density of population
occurs in the summer (July-August).

248 Wisconsin Academy of Science, Arts and Letters [Vol. 56

500 ind. / m. ^

I - 1

Figure 15. Average verti¬
cal distribution in the
density of population
of Hiriidinea in Lake
Mendota.

As in the case of Glossiphonidae and Erpobdellidae, it is natu¬
ral that the maximum density of population and biomass of all
Hirudinea fauna should occur in the summer months (July-
August). At this time the young generation of several species
appears, particularly of the most abundant leeches, such as Helob-

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

249

Table IV. The Average Population Density and Biomass of Leeches
(Hirudinea) in Different Zones of Lake Mendota

Zones

Littoral

SUBLIT-

TORAL

Profundal

Average
FOR All
Zones

Per Square Meter

Individuals .

355.2

13.6

0.7

90.3

Wet weight in gr .

1.8657

0 . 0443

0.0003

0 . 4648

Dry weight in gr .

0.3455

0.0092

0.00006

0.0888

della stagnalis, Erpohdella punctata, N ephelopsis ohscura and Glos-
siphonia complanata. It is also natural that the minimum density of
population and biomass should occur in the winter months (De-
cember-February) , probably as the result of very severe conditions
during- that period of the year. The average seasonal changes in
population density and biomass of all leeches in the littoral zone
of Lake Mendota during 1964-65 are represented in Figure 18.

The horizontal distribution in the density of the leeches in Lake
Mendota in May 1965 is represented in Figure 17. From this one
can see that the leeches having a larger density of population occur
in the Maple Bluff, Fox’s Bluff, Bascom Woods, and Second Point
areas. However, the greatest vertical distribution is found in the
Second Point, Yahara Canal, Maple Bluff and Fox’s Bluff regions.

Acknowledgments

I would like to take this opportunity to thank the University of
Wisconsin for supporting my research and sponsoring the Ameri-
Skopje. I wish to extend my particular appreciation to Professor

1000 ind. /m.^

Figure 16. Vertical distribution in the density of population of the leeches
during 1964-65 in Lake Mendota.

250 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1000 ind./m.^

I - 1

X

LU

O

10-

12 - - -

Figure 17. Horizontal distribution in the density of population of the leeches
in Lake Mendota on the month of May, 1965.

can- Yugoslav Exchange for staff members of the University of
Arthur D. Hasler, Director of the Laboratory of Limnology, for
the many suggestions which he made concerning my work; to Dr.
Marvin C. Meyer, Professor of Zoology, University of Maine, for
verifying my identifications of Lake Mendota leeches ; to Mr. Gary
L. Hergenrader, graduate student in the Laboratory of Limnol¬
ogy, with whose help I was able to take the bottom samples during
the winter period; and to Mr. Paul E. Sager, graduate student at
the Laboratory of Limnology, for his help in reviewing the manu¬
script. Finally, I should also like to thank all others, especially the
members of the staff of the Laboratory of Limnology, Department
of Zoology of the University of Wisconsin, for the many ways in
which they have contributed to this project.

INDIVIDUALS PER SQUARE METER

1967-68] Sapkarev — Taxonomy and Ecology of Leeches

251

Figure 18. Average seasonal change of the population density of biomass of
Hirudinea in the littoral zone of Lake Mendota during 1964-65.

MILLIGRAMS PER SQUARE METER

252 Wisconsin Academy of Science, Arts and Letters [Vol. 56

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Andrews, 0. V. 1915. An ecological survey of Lake Butte des Morts Bog,
Oshkosh, Wisconsin. Bull. Wis. Nat. Hist. Soc., 13:196-211.

Autrum, H. 1936. Hirudineen. In: Broons Klassen und Ordnungen des Tier-
reichs. Band IV, Abt. Ill, Buch 4, Teil 1:1-96.

Baker, F. C. 1924. The fauna of the Lake Winnebago region (a quantitative
and qualitative survey with special reference to the Mollusca) . Trans. Wis.
Acad. Sci. Arts & Lett., 21:109-146.

Bennike, S. a. Boisen. 1940. On some Iranian Freshwater Hirundinea. Danish
Scientific Investigations in Iran. 2:1-10, Copenhagen.

- . 1943. Contributions to the ecology and biology of the Danish fresh¬
water leeches (Hirudinea). Folia Limnologica Scandinavica. No. 2:1-109.
Berg, K. 1938. Studies on the bottom animals of Esrom Lake. D. Kgl. Danske
Vid. Selsk. Skrift., Afd. Naturv. Math., 9. Baekke, VIII: 1-255.

Bere, R. 1931. Leeches from the lakes of northeastern Wisconsin. Trans. Wis.
Acad. Sci. Arts & Lett., 26:437-440.

Birge, E. a. and C. Juday. 1914. The inland lakes of Wisconsin. Wis. Geol. Nat.
Hist. Survey, No. 27.

Bruun, F. 1938. Freshwater Hirudinea. The Zoology of Iceland, 2, Part 22.
Cahn, a. R. 1915. An ecological survey of the Wingra Springs region, near
Madison, Wisconsin, with special reference to its ornithology. Bull. Wis.
Nat. Hist. Soc., Vol. 13, No. 3:123.

Castle, W. E. 1900. Some North American freshwater Rhynchobdellidae and
their parasites. Bull. Mus. Comp. Zook, 36:15-64.

Herter, K. 1937. Die Okologie der Hirudineen. Bronns Klassen und Ordnungen
des Tierreichs. Bd. 4, Abt. Ill, Buch 4, Teil 2, Lief. 3.

Hilsenhofp, W. L. 1963. Predation by the leech, Helobdella stagnalis, on
Tendipes plumosus (Diptera :Tendipedidae) . Ann. Entomol. Soc. Amer.,
Vol. 56:252.

- . 1964. Predation by the leech, Helobdella nepheloidea on larvae of

Tendipes plumosus (Diptera :Tendipedidae) . Ann, Entomol. Soc. Amer.,
Vol. 57:139.

Mann, K, H. 1955. Some factors influencing the distribution of freshwater
leeches in Britain. Proc. Internatl. Assoc. Theoret. and Appl. Limnol.,
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- . 1957. The breeding, growth, and age structure of a population of the

leech, Helobdella stagnalis. J. Anim. Ecol., 26:171-177.

Meyer, M. C. and J. P. Moore. 1954. Notes on Canadian leeches. Wasmann J.
Biol., 12:63-96.

Miller, J. A. 1929. The leeches of Ohio. Contrib. Franz Theodore Stone Lab.,
2:1-38.

Miller, J. A. 1937. A study of the leeches of Michigan with key to orders,
suborders and species. Ohio J. Sci., 37:85-90.

Moore, J. P. 1901. The Hirudinea of Illinois. Bull. Ill. State Lab. Nat. Hist.,
5:479-547.

- . 1906. Hirudinea and Oligochaeta collected in the Great Lakes region.

Bull. U. S. Bur. Fish., 21:153-171.

- . 1924. The leeches (Hirudinea) of Lake Nipigon. Univ. Toronto Studies.

No. 23:17-31.

- . 1939. Helobdella punctata — lineata, a new leech from Puerto Rico.

Puerto Rico J. Public Health Trop. Med., Vol. 14:422-429.

- . 1952. Professor Verrilks freshwater leeches. Notulae Naturae Acad.

Nat. Sci. Phila., No. 245:1-15.

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1967-68] Sapkarev — Taxonomy and Ecology of Leeches 253

Muttkowski, R. a. 1918. The fauna of Lake Mendota, A quantitative and
qualitative survey with special reference to the insects. Trans. Wis. Acad.
Sci. Arts & Lett., 19:374-482.

Oka, a. 1917. Zoological results of a tour in the Far East. Hirudinea. Mem.
Asiat. Soc. Bengal, 6:157-176.

OKLAND, J. 1964. The eutrophic Lake Borrevann, Norway. An ecological study
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Pawlowski, L. K. 1936. Zur Okologie der Hirudineen fauna der Wigryseen.
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der Systematik und Okologie der Hirudinea des Dojran-Sees. Folia
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Verrill, a. E. 1872. Synopsis of American freshwater leeches. Report of Com-
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Nachtrieb, H. F., E. E. Hemingway and J. P. Moore. 1912. The leeches of
Minnesota. Geol. Nat, Hist. Survey, Zool. Ser. No, 5.

ZSHOKKE, F. 1900. Die Tierwelt der Hochgebirgseen. N. Denschr. Schweiz.
Ges., 37.

RABIES AND RABIES CONTROL IN WISCONSIN

Daniel O. Trainer^'

A prerequisite to any consideration of rabies control in Wisconsin
is a review of the status of the disease, a resume of some of the
factors that have contributed to the situation, and a summary of
the specific control approaches utilized.

Status of Rabies

The prevalence and relative distribution of rabies in wild and
domestic animals in Wisconsin (Table 1) is not unlike that reported
in the United States (Scholtens and Tierkel, 1963). Despite annual
fluctuations in the total number of rabies cases in the state, the
disease in domestic animals has not varied significantly since 1952.
The major change has occurred in wild animal rabies, especially the
skunk (Mephitis mephitis).

* Department of Veterinary Science and Wildlife Ecology, University of Wisconsin-
Madison.

Table 1. A Summary of Animal Rabies in Wisconsin, ( 1952-1966 )

Year

Total

Number

OF Labop

LATORY (

2ases

Domes¬

tic

Wild

Skunk

Fox

Ba t

Rac¬

coon

1952 .

56

27

29

27

1

0

1

1953 .

49

21

28

25

2

0

1

1954 .

90

47

43

36

6

0

1

1955 .

39

17

22

19

3

0

0

1956 .

41

23

18

13

2

0

2

1957 .

74

23

51

37

5

6

2

1958 .

227

33

194

184

7

3

0

1959* .

92

23

69

64

4

1

0

1960 .

24

15

9

5

1

0

1

1961 .

30

21

9

5

1

2

0

1962 .

42

24

18

12

1

5

0

1963 .

62

33

28

17

7

5

1

1964 .

95

62

33

12

11

6

3

1965 .

64

39

25

21

2

1

1

1966 .

68

41

27

22

4

1

0

*Since 1959, wild animals were tested only if there was human exposure.

255

256 Wisconsin Academy of Science, Arts and Letters [Voi 56

In 1958, there were 184 laboratory confirmed cases of rabies in
skunks, more than 10 times the number recorded two years earlier.
Due to the volume of wild animals submitted for rabies examina¬
tion and the fact that at least 85 per cent of all submitted skunks
were rabid, the State Laboratory of Hygiene, beginning in 1959,
examined wild animals only if there was human exposure. All of
the rabies laboratory examinations in Wisconsin are conducted by
the State Laboratory of Hygiene. Wild animal rabies figures sub¬
sequent to 1959 are therefore not directly comparable with those of
prior years. Since the establishment of this new laboratory policy,
the number of confirmed skunk rabies cases, all involving human
exposure, has persisted and even risen moderately indicating the
existance and a possible increase of rabies in this species.

Despite a sizable population, the fox (Vulpes fulva and Urocyon
cinereoargenteus) has not been an important rabies target in Wis¬
consin (Table 1) . Rabies in insectivorous bats was initially detected
in Wisconsin in 1957, and since then has been detected annually in
low numbers (Table 1). Despite a large and increasing raccoon
(Procyon lotor) population, rabies persists in this species at a low
level (Table 1).

Until 1960 the principle domestic animal target of rabies was the
dog. Since then, the cow and the dog have fluctuated as the leading
domestic animal victim. Sporadic rabies cases have also occurred in
cats, swine and horses.

Wildlife Populations

Since the reservoir of rabies in Wisconsin exists among wild
populations, a review of some population trends of involved species
is appropriate. To census any wild population on a state-wide basis
is difficult, however related data can sometimes be utilized to project

Table 2. Some Wildlife Harvest Figures in Wisconsin. (1920-1965)

Year

Species (thousands)

Skunk*

Fox**

Raccoon*

1920 .

56.3

4.6

1930 .

56.7

3.4

6.4

1940 .

50.7

11.0

13.6

1950 .

11.6

28.5

34.3

1960 .

0.8

57.0

50.0

1965 .

0.4

52.8

63.2

*Harvest figures from WCD trapping and hunting records.
**Harvest figures from WCD bounty records.

1967-68]

Trainer — Rabies and Rabies Control

257

trends of these wild populations. For example, bounty payment fig¬
ures provide a kill figure which can be utilized to project population
trends. A summary of fox bounty records (Table 2) suggests a
fluctuating population iwhich has tended to increase since 1930.
Despite this apparent increase of fox numbers there has been no
conspicuous alteration in rabies prevalence (Table 1), During this
35 year span, fox pelt prices have ranged from $12 to 40^ with the
highest prices paid at either end of the period (Wisconsin Conser¬
vation Department, 1965^).

The raccoon is not a bountied animal in Wisconsin, but it is
utilized as fur, meat and sport. Harvest figures for the raccoon
(Table 2) have increased from 4,600 in 1920 to 63,200 in 1965.
Raccoon fur prices have varied during this period ($4.35 in 1920;
65f^ in 1948; $2.50 in 1965) and undoubtedly influence the harvest.
Low fur prices result in less trapping effort and a larger raccoon
population. Since 1945 the raccoon has become an important sports
animal and harvest figures now include a larger proportion of
animals taken by hunting than by trapping. Increasing harvest
figures and reports of crop depredation, vandalism, etc. indicate an
extensive and growing raccoon population. This apparent rise in
raccoon numbers has not been accompanied by a parallel increase
in rabies.

The skunk, Wisconsin’s major rabies target, was once an im¬
portant fur animal. In fact it was the second most important fur
bearer in the state in 1918 when 74,300 skunks brought Wisconsin
trappers almost one-third of a million dollars (Scott, 1940). Be¬
cause of its fur value the skunk was protected by prescribed trap¬
ping seasons until 1930 when the Conservation Commission was
asked by the Department of Agriculture to withdraw protection of
the skunk because it was a reservoir of rabies. As late as 1945,
more than 58,000 skunks were harvested. Fur prices have declined
steadily since the mid-1940’s and today a prime skunk pelt is worth
less than a dollar.

Trapping in general and for skunks in particular has declined
drastically since 1945, and in recent years the number of skunks
harvested is less than a thousand. Table 3 depicts this decline in
trapping interest despite a marked increase in other outdoor activi¬
ties. The drop in fur value accompanied by the decline in trapping
has resulted in a decreased harvest of skunks and an apparent in¬
crease in their numbers.

Land Use Changes

There are 36 million acres in Wisconsin of which 34% in crop
land and 40% is forests. Several important land-use changes in
Wisconsin have undoubtedly contributed to changes in wildlife

258 Wisconsin Academy of Science^ Arts and Letters [Vol. 56

numbers. Similar to the national trend of farming and land use,
the number of farms, farmers, and acres in farms has declined in
recent years (Table 4). In addition to the reduction of acres in
farms (from 23.5 to 21.2 million) more than 770,000 acres have
been retired in various Conservation Reserve programs (Buse and
Brown, 1965).

The purchase and development of land for wildlife purposes is
a major program in Wisconsin. The Game Division of the Wiscon¬
sin Conservation Department in 1927 initiated a land acquisition
program. Since that time they have 208 individual projects under¬
way or completed in which they own 273,000 acres and lease
another 291,000 acres for public hunting. In addition there are 4.5
million acres of national, state or county forest land, and private
forest croplands available for public recreation. In 1961 the Wis¬
consin Outdoor Recreation Act established a one cent per pack tax
on cigarettes. These funds (approximately 5 million dollars
annually) are earmarked for land acquisition to protect and pro¬
mote natural resources in the state.

Another growing industry in Wisconsin is wildlife farming.
There are 27 beaver farms (6,600 acres), 156 deer farms (100,582

Table 3. Wisconsin Conservation Department License Sales. (1920-1965)

Year

Type of

License (thousands)

Trapping

Hunting

Deer

Fishing

Sportsman*

1920 .

20.0

nr**

50.2

NR

NR

1930 .

18.9

NR

77.3

NR

NR

1940 .

15.3

295.7

102.3

233.1

2.8

1950 .

10.4

455.8

289.4

716.7

20.0

1960 .

4.4

278.3

269.8

612.9

65.3

1965 .

2.1

378.8

382.6

490.9

218.5

*A sportsman license allows hunting, fishing and trapping.
**Not required.

Table 4. Farm Trends in Wisconsin. (1935-1965)

1935

1945

1955

1965

Total Number Farms .

(Thousands)

200

178

155

124

Total Farm Acreage .

(Millions)

23.5

23.7

23.2

21.2

Total Land in Farms .

(%)

66

66

64

60

1967-68]

Trainer — Rabies and Rabies Control

259

acres), 1,012 game bird farms (7,085 acres), 350 muskrat farms
(45,717 acres) and 128 shooting preserves (43,775 acres). In addi¬
tion the federal government has more than 150,000 acres in wildlife
refuges.

Reforestation provides another example of land alterations which
can result in wildlife habitat improvement. In 1959 state nurseries
alone distributed 43 million trees for reforestation (Wisconsin Con¬
servation 'Department 1965^) . Another 1.9 million game food shrubs
were sold to private land owners by state nurseries.

Various combinations of the aforementioned changes could have
an important impact on wildlife populations. Accompanying this
increase in the number of potential rabies vectors, is the increased
opportunity for human contact with wildlife, Wisconsin is in step
with the nation concerning increased outdoor recreation. It has 71
state recreation areas with camp sites, 39 federal areas, 169 county
or city areas and 270 private camping establishments (Wisconsin
Conservation Department, 1965^) , In state parks alone during 1964
there were six million visitors and over 700,000 camper days spent.
The steady increase in hunting and fishing license sales (Table 3)
further attests to this outdoor trend.

Rabies Control Program

Despite these increased opportunities for human contact with
rabies, the disease has not been a major human health problem in
Wisconsin. There has been one human rabies death, the result of a
bat bite, in the state during the last decade.

Although wild animal bite records are not maintained in Wiscon¬
sin, the majority of wild animal rabies suspects submitted for
diagnosis involve human exposure ; therefore, the threat of human
rabies is ever present and real. To combat this potential rabies
problem, various agencies and organizations acting independently
and on occasion together have produced a variety of rabies control
programs. Basically the approach has been one of education involv¬
ing the public, physicians, veterinarians and wildlife professionals
as well as pet vaccination programs and the control of local wildlife
populations.

The Wisconsin Department of Agriculture through its Animal
Health Division promotes an educational program on rabies for
Department employees and veterinarians. A monthly computation
of animal rabies cases by county is issued to all concerned individ¬
uals. Its monthly newsletter “Animal Health” is supplied free to
state veterinarians and reports the status of rabies in wild and
domestic animals, the location of recent cases, current rules and
regulations concerning the disease, and other significant rabies in-

260 Wisconsin Academy of Science, Arts and Letters [Vol. 56

formation. On several occasions a geographic section of the state
has been quarantined due to the threat of rabies.

Local Veterinary Associations with the aid of University of
Wisconsin extension personnel have established county vaccination
clinics. Almost half of the 72 counties in Wisconsin have sponsored
local rabies vaccination clinics which have varied in size, procedure
and success. Most of these programs were initiated in 1958 when
rabies was very prevalent in the state. Some programs were dis¬
continued after several years, others exist today, and new clinics
are being added annually — especially in recreation areas.

The State Health Department conducts an educational program
similar to the Department of Agriculture, but directed toward local
health agencies and physicians. Data on the status of rabies, appro¬
priate therapeutic procedures, and recommended laboratory pro¬
tocol are stressed. Their newsletter as well as conventional news
media are utilized. The State Laboratory of Hygiene of the State
Health Department conducts all of the diagnostic rabies work in
Wisconsin, a free service available to physicians and veterinarians.

Since the major rabies problem involved wildlife, the Wisconsin
Conservation Department is concerned and sponsors an active pro¬
gram of information and education. Department personnel are in¬
formed on the status of the disease in wild and domestic animals
via periodic administrative directives stressing signs of disease,
procedure for handling rabies suspects, and the appropriate pro¬
tocol following human exposure.

Campers are an important high risk group ; therefore, an educa¬
tional program on rabies in wildlife involving the press, television,
and radio is periodically directed at this group as well as other
outdoor sportsmen. Rabies warnings are posted at appropriate state
camp grounds and dogs are restrained in all parks. The vaccination
of hunting dogs is promoted.

If rabies problems exist in local skunk, fox or raccoon popula¬
tions adjacent to a camp site, control by shooting and trapping is
initiated. In addition an extension predator trapping program has
been initiated to teach farmers or other interested sportsmen to
trap wild animals.

The combination of the aforementioned programs, education of
the public and involved professionals, has proven elfective in con¬
taining this important disease problem in Wisconsin. The continu¬
ation of this educational approach with rabies surveillance, vaccina¬
tion, and control of local wild populations is anticipated and
essential.

Summary

Rabies has been present in Wisconsin for many years in both
wild and domestic animal populations. Despite annual fluctuations

1967-68]

Trainer— Rabies and Rabies Control

261

since 1952, no significant change in the prevalence of rabies in
domestic animals has occurred. Wildlife rabies, specifically in the
skunk, has varied considerably during this period. Some of the
environmental alterations that have contributed to the wildlife
rabies picture are new agricultural patterns, reforestation and in¬
creased recreation activities.

Various agencies including the State Departments of Agricul¬
ture, Health, and Conservation pursue an informational and educa¬
tional rabies program aimed at the public, physicians, veterinarians
and wildlife professionals. This in combination with vaccination
clinics and control of local wildlife populations contains the rabies
problem in Wisconsin.

Literature Cited

Buse, R, C. and R. N. Brown, 1965. The Conservation Reserve in Wisconsin.

Agr, Exp, Sta. Bull,, University of Wisconsin, Madison. 18,

Scholtens, R. G, and E. S. Tierkel, 1963. Incidence of animal rabies in the
United States. J.A.V.M.A. 143:52,

Scott, W, 1940. Early Wisconsin game and fur harvest summary. Wisconsin
Conservation Department, Madison. 12.

Wisconsin Conservation Department 1965"^, Wisconsin Conservation Depart¬
ment game and fur reports. Wisconsin Conservation Department, Madison.
Mimeograph.

Wisconsin Conservation Department 1965''. Twenty-ninth biennial report of the
Wisconsin Conservation Department. Wisconsin Conservation Department,
Madison. 148.

Wisconsin Conservation Department 1965®. Wisconsin camp ground directory.
Vacation and Travel Service, Wisconsin Conservation Department, Madi¬
son, Pub, 110:67.

NOTES ON WISCONSIN PARASITIC FUNGI. XXXIII

H, C. Greene

Department of Botany, University of Wisconsin, Madison

This series of notes is, unless stated otherwise, based on collec¬
tions made during the season of 1966.

General Observations

A high incidence of powdery mildews in Wisconsin is noted by
Dr, Koji Hirata who has recently (1966) published a book entitled
^'Host Range and Geographical Distribution of the Pov/dery Mil¬
dews''. Through 1962, according to his tabulation, 380 different
powdery mildew-host combinations had been reported for this
state— more than for any other state in the Union, This despite
the fact that Wisconsin is not exceptionally rich in total number of
higher plant species. Hirata ascribes this profusion, no doubt cor¬
rectly, to the intensive collecting efforts of the late J. J. Davis
and the writer, carried on almost continuously for the past eighty-
odd years.

Mycosphaerella sp. on stems of Eleocharis acicularis (L.) R.
& S. collected June 26 at Madison appears to have developed para-
sitically. The clear grayish-black perithecia are closely gregarious,
deeply sunken, subglobose, about 115-140 diam. ; asci hyaline,
narrowly clavate, 60-65 x 9-11 p; ascospores hyaline, subfalcate,
approx. 16-17 x 5 /a. The host plants were on rapidly drying soil,
but were in the main still green and healthy.

Mycosphaerella sarraceniae (Schw.) House on the brown
upper portions of otherwise still green “pitchers" of Sarracenia
purpurea L,, collected June 11 at Hope Lake Bog near Lake Mills,
Jefferson Co., may possibly, it would seem, have initiated its devel¬
opment parasitically,

Mycosphaerella sp. is amphigenous on suborbicular to irregu¬
larly elongate brownish-cinerous, mostly marginal areas on leaves
of Heliopsis helianthoides (L.) Sweet, collected in Iowa Co,, near
Mazomanie, August 5. The perithecia are scattered to gregarious,
black, thick-walled, subglobose, about 175-225 /a diam. ; asci hyaline,
short-pedicellate, subcylindric or slightly obclavate, 37-42 x 5-7.5
p; ascospores hyaline, fusoid, 10-11 x 2. 5-2.7 ,/a. Possibly parasitic.

263

264 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Sphaerulina sp., probably parasitic in its early development,
has been noted at different times in several localities on still
attached and also on fallen leaves of Quercus ellipsoidalis Hill. The
scattered, epiphyllous perithecia do not mature in the fall but can
be brought to maturity in the following spring by holding in a moist
chamber for several days, as was done with a collection made Octo¬
ber 13, 1965 at Tower Hill State Park, Iowa Co. The black, globose,
thick-walled perithecia are approx. 135-150 /x diam. ; asci hyaline,
curved and clavate, 40-45 x 8-9 y; ascospores hyaline, straight,
narrowly cylindric, 3-septate, about 15 x 3 /x. The dimensions of
asci and spores do not correspond with any of the several species
of Sphaerulina described as occurring on oak leaves.

Pringsheimia ( ?) sp., possibly parasitic, occurs on dead white
areas on leaves of jet-bead, Rhodotypos tetrapetala Makine (cult.),
collected at Madison, July 27. This is a coarser form than Pring-
shemia sepincola (Fr.) Hoehn. which develops on rose twigs. In
the specimen on Rhodotypos the fruiting structures (perithecia?)
are epiphyllous, black, globose, erumpent, gregarious, widely osti-
olate, about 135-150 /x diam. ; asci hyaline, 8-spored, broadly clavate,
the wall appearing quite thick, overall about 60 x 35 /x; ascospores
hyaline, broadly fusoid, 4-septate with usually one or two cross
septa, about 26-28 x 10-11 /x.

Phyllostictae, appearing parasitic, but undetermined as to spe¬
cies, continue to be found. Descriptive notes on some of these fol¬
low mention of the names of the host plants on which they
occurred :

1) On Sorghastrum nutans (L.) Nash, near Leland, Sauk Co.,
June 14. The spots are 1.5-3 mm. long, narrowly ellipsoid with
tan centers and purplish borders, the pycnidia scattered, black, sub-
globose, small, about 85-115 /x diam., the conidia hyaline, subcylin-
dric to subfusoid, biguttulate, approx. 12-16 x 3.5-5 /x. No septa
were seen, but it seems possible this may be a poorly developed
Ascochyta or Stagonospora. 2) On Aquilegia canadensis L., near
Leland, Sauk Co., July 8. The lesions are subcircular, sordid brown,
sinuous, with pale brown inner portion and relatively wide blackish-
purple margins, the pycnidia epiphyllous, gregarious or scattered,
black, subglobose, widely ostiolate, about 100 /x diam., the conidia
hyaline, biguttulate, subcylindric, broadly ellipsoid, or more rarely
broadly subfusoid, 7.5-11 x (2.5) 3.5-4 /x. 3) On Pyrus ionensis
(Wood) Bailey, at Madison, August 30. The lesions are orbicular,
reddish-brown, subzonate, about 1-2 cm. diam., the pycnidia epiph¬
yllous, scattered, flattened, quite superficial, light brown, thin-
walled, rather widely ostiolate, about 75-100 /x diam., the conidia
hyaline, short-cylindric or ellipsoid, shall, approx. 4-5 x 1.7-2 /x.

1967-68] Greene — Wisconsin Parasitic Fungi. XXXIII

265

It seems possible that the spots are due at least in part to Fusi-
cladium dendriticum action. 4) On Geum canadense Jacq., near
Leland, Sauk Co., July 8. The lesions are subcircular, sordid brown,
immarginate, slightly sunken, small, about 1.5-3. 5 mm. diam., the
pycnidia epiphyllous, loosely gregarious, black, subglobose, thick-
walled, rather widely ostiolate, about 125-175 diam., the conidia
hyaline, fusoid, subfusoid, or narrowly ellipsoid, frequently biguttu-
late, (5~)6.5-9.5 x 2.3-2. 6 jx. Suggestive of Phomopsis, but no sco-
lecospores were observed, 5) On Prunus virginiana L., at Madison,
June 30. The lesions are orbicular, bright reddish-brown, subzo-
nate, circumscissle and dehiscent, about .3-1 cm. diam., the pyc¬
nidia epiphyllous, pallid brown, flattened and imperfect below, tiny,
about 40-60 /x diam., the conidia subhyaline with a faintly sooty
tinge, variable in shape, oblong, ovoid, broadly ellipsoid or broadly
subfusoid, 4.5-8 x 2.7-4.2 /x. 6) On Gaultheria procumbens L., near
Leland, Sauk Co., June 22. The lesions are white, rounded, up to
5 mm. diam., the pycnidia epiphyllous, scattered, shining black,
rather deeply immersed, globose, about 100-150 /x diam., the conidia
pallid brownish in mass, but individually appearing hyaline,
straight, rod-shaped, approx. 5. 5-8. 5 x 1.32-2 /x. The conidia of
Phyllosticta gaultheriae Ell. & Ev. are described as running 5-7 x
4-5 /X. 7) On Solidago riddellii Frank, at Madison, September 16.
The lesions are dark-bordered with gray central portion, narrow
elongate, ranged along the leaf midrib, the pycnidia subcuticular,
epiphyllous, gregarious, black, thick- walled, somewhat flattened,
about 150 /X diam., the conidia hyaline, ellipsoid or sufusoid, approx.
10-13 X 5-7 /X. Except for the subcuticular habits this is much like
Leptothyrium similisporum (Ell. & Davis) Davis which occurs on
several species of Solidago in Wisconsin. 8) On Solidago patula
Muhl., near Leland, Sauk Co., August 12. The lesions are grayish,
immarginate, orbicular, 1 cm or more in diam., the pycnidia scat¬
tered, pallid brownish, thin-walled, subglobose, about 200 /x diam.,
the conidia hyaline, slender-cylindric, contents granular, approx.
7.5-10 X 1.2-1. 7 /X, non-septate. Approaches Septoria in conidial
dimensions. 9) On Erigeron annus (L.) Pers., near Leland, Sauk
Co., July 8. The spots are tan with narrow, darker brown margin,
orbicular or semi-orbs impinging on the leaf margin, .6-1.2 cm.
diam., the pycnidia epiphyllous, gregarious, pallid brownish, thin-
walled, subglobose, about 90-150 /x diam., the conidia hyaline,
broadly ellipsoid, ovoid, occasionally subfusoid, approx. 5-7.5 x 2.5-
3.5 /X, very numerous. 10) On Silphium perfoliatum L., near Leland,
Sauk Co., July 20. The spots are few and scattered, usually only one
or two per leaf, rounded, with cinereous centers and darker bord¬
ers, about 2-6 mm. diam., the pycnidia appearing amphigenous,
gregarious, pallid brownish, thin-walled and translucent, subglo-

266 Wisconsin Academy of Science, Arts and Letters [VoL 56

bose, approx, 100-150 p. diam., the conidia hyaline, subcylindric or
subfusoid, approx. 4.5-7 x 2.3-2.7 y.

Phomopsis sp., which appears to have originated parasitically,
infects the stems and capsules of Aquilegia canadensis L. collected
near Leland, Sauk Co., August 3. The pycnidia are black, thick-
walled, subglobose, about 125-150 p diam. Both types of conidia are
present in abundance, the scolecospores flexuous, thread-like, hya¬
line, (15-) 17-21 X ,7-1 p the others fusoid, hyaline, approx. 7,5 x
(1.8-)2.2-2.7(-3) /X.

Phomopsis (?) spp. have been studied on several hosts. 1) On
Veronica arvensis L., at Tower Hill State Park, Iowa Co., June 1.
Superficially this collection closely resembles examples of Septoria
veronicas Desm., but the latter all have scolecospores about 1 p
thick and mostly quite long and flexuous. The specimen in question,
however, has two classes of spores: a) robust, obtuse, mostly
slightly curved, hyaline, continuous, about 22-25 x 2.5-3 p, and
b) less numerous ordinary scolecospores, approx. 25-40 x 1-1,30 p.
2) On Aster sagittifolius Wedem. near Leland, Sauk Co., August 3.
The spots are large, up to 3 cm. diam., suborbicular, light reddish-
brown with cinereous zonate banding, the pycnidia epiphyllous,
zonately arranged, black, thick-walled, subglobose, approx. 150-200
p diam., the conidia hyaline, slender-fusoid, straight to slightly
curved, (8-) 9-12 (-15) x (2.3-) 2. 5-2. 7 (-3) p. No scolecospores
observed. 3) On Tragopogon pratensis L., near Edgerton, Eock Co.,
September 18, 1965. On the flower peduncles. The pycnidia are gre¬
garious, sometimes even confluent, black, subapplanate, widely osti-
olate, approx. 150-250 p diam., the conidia hyaline, fusoid or sub¬
fusoid, (7.5-)8.5-10(-12) x (2-) 2.5-3 /X. No scolecospores observed.

Ascochytae which occur on the leaves of higher plants are gen¬
erally strong and obvious parasites, characterized by conspicuous,
often extensive, but nevertheless sharply defined lesions, with the
pycnidia being rather large and flesh-colored or light brownish,
and often in a zonate arrangement. Most investigators of these
fungi, the writer included, have assumed strong host specificity,
and in many cases this is almost certainly so. However, in a series
of collections made during the past several years, particularly in
1966, in the vicinity of Leland, Sauk Co., host specificity seems
open to question. On June 21, 1966 three Ascochytas were collected
on Asarum canadense L., Mitella diphylla L. and Sanguinaria
canadensis L. in a very limited area of not more than 150 feet
square. Those on Mitella and Sanguinaria appear to be identical
with previous collections made in the same general vicinity and
reported on in my Notes 30 and 31. The following day, June 22,

1967-68] Greene — Wisconsin Parasitic Fungi. XXXIII 267

at a point less than two miles distant, the Ascochyta on Sanguinaria
was found again and close by there occurred a similar form on
Symplocarpus foetidus (L.) Nutt. The following tabulation of
pycnidial and conidial measurements would seem to suggest forms
closely related, if not identical. The measurements of conidia are
based on those which are septate and presumably mature :

Asarum—Fycnidisi 125-200 ju, diam. ; Conidia 7.5-10 (-12) x
(2.5-)2.8-3.5 /X

Mitella — ^Pycnidia 90-150 /x diam.; Conidia 6.5-8 (-10) x 2.5-
3.3 ja

Sanguinaria — Pycnidia 150-180 /x diam.; Conidia (7-)7.5-
10 (-12) X 2.5-3.8 /X

Symplocarpus — Pycnidia 100-200 /x diam. ; Conidia 7.5-11 x 3-
3.5 /X

The only report I have found of Ascochyta on any of these genera
is that of A. versicolor Bubak, said to occur on Asarum caudatum
Lindl. in Idaho, but this species has conidia 10-25 x 4-6.5 /x, out
of the range of the Wisconsin specimens. Whether one or more
species is involved in the Wisconsin material is a question unfor¬
tunately not likely to be resolved in the foreseeable future, but the
need for caution in the description of these fungi is plain.

Ascochyta spp., undetermined but obviously parasitic, including
two mentioned in the preceding note, show the following character¬
istics: 1) On Symplocarpus foetidus (L.) Nutt., near Leland, Sauk
Co., June 22. One large lesion is narrowly oval and elongate, about
3 cm. X .8 cm., with narrow blackish-brown border and sordid
greenish internal portion, the flesh-colored pycnidia amphigenous,
thin-walled, subglobose, approx. 100-200 /x diam., the conidia
hyaline, subcylindric or subfusoid, occasionally curved, often
slightly constricted at septum, about 7.5-11 x 3-3.5 /x, A very few
conidia are somewhat longer and 2-septate. 2) On Asarum
canadense L., near Leland, Sauk Co., June 21. The lesions are cir¬
cular or broadly oval, .5-1 cm. diam., with greenish or grayish
centers, the pycnidia gregarious, flesh-colored, thin-walled, sub-
globose, approx. 125-200 /x diam., the conidia hyaline, cylindric or
subcylindric, 7.5-10(12) x (2.5-) 2.8-3. 5 /x. Only about 20% of the
conidia are septate, the continuous ones being smaller and presum¬
ably immature. 3) On Anemone canadensis L., near Leland, Sauk
Co., July 8. The lesions are orbicular and grayish-brown, with
darker borders, up to 1 cm. diam., the pycnidia epiphyllous, gregari¬
ous to crowded, pallid brownish, subglobose, about 125-150 /x diam.,
the conidia hyaline, cylindric, almost uniformly septate, approx.
7.5-9 X 2.5-3 ,/x. 4) On Decondon verticillatus (L.) Ell., near Trout
Lake, Vilas Co., July 1, 1914. Collected by J. J. Davis who placed

268 Wisconsin Academy of Science, Arts and Letters [Vol. 56

the specimen in the herbarium as ''Ascochyta decodontis ined/',
but did not report on it in his Notes. The pycnidia are pallid
yellowish-brown, thin-walled, rather widely ostiolate, mostly about
100 fx diam. In a note enclosed with the specimen Davis states the
conidia are “cylindrical to subfusiform and acute at one end,
straight or slightly curved, 15-22 x 3 In a mount examined by
the writer the conidia were mostly shorter and the majority with¬
out a septum. Phyllosticta neseae Peck on this host is said to have
conidia approx. 7-10 x 2.5 /x, not too different from the Wisconsin
specimen. 5) On Hieracium aurantiacum L. at Madison, June 24.
The lesions are oval to orbicular, dull brownish with narrow darker
margin, zonate or subzonate, about .5-1.7 cm. diam., the pycnidia
epiphyllous, scattered to gregarious, pallid brownish, subglobose,
approx. 150-250 /x diam., the conidia hyaline, uniformly septate,
cylindric or subcylindric, straight or slightly curved, (6.5-) 7.5-11
X (2.6-)2.8-3.7 /X.

Hendersonia crastophila Sacc. occurs in seriate arrangement
on elongate, narrow, white lesions on the adaxial surfaces of leaves
of Panicum virgatum L. collected at Madison September 11, 1964.
The Hendersonia appears to be in association with Puccinia panici
which occurs on the reverse side of the leaves. Presumably the rust
infection preceded the development of the Hendersonia, so the latter
is probably only weakly parasitic. On the other hand, H. crastophila
is present on leaves of Sporobolus asper (Michx.) Kunth., collected
the same day at a location in Iowa Co. near Blue Mounds, in a prob¬
able parasitic relationship, as there is no other fungus present.

“Septoria"' sigmoidea Ell. & Ev., occasionally found on Panicum
virgatum L. in Wisconsin, is plainly not a Septoria in the usual
conception of that genues. It has been suggested that it may be
identical with Hendersonia crastophila Sacc., but the conidia with
their strong curvature, pronounced taper at both ends, and length
of about 60-75 /x do not seem to fit in this species. It is more likely
the fungus should be referred to Phaeoseptoria Speg. It is to be
regretted that S. sigmoidea seems to have escaped the attention of
the late Roderick Sprague, the outstanding authority on
Phaeoseptoria.

Septoria, undetermined as to species, occurs on several hosts,
with descriptive notes as follows: 1) On Eriophorum virginicum
L. at Hope Lake Bog near Lake Mills, Jefferson Co., June 11. The
pycnidia, which occur on the dead tips of otherwise living leaves,
are shining black, globose, about 150 /x diam., the conidia hyaline,
mostly somewhat lax and slightly curved, with granular contents
but appearing continuous, approx. 30-50 x 1.7-2. 5 /x, Septoria

1967-68] Greene — Wisconsin Parasitic Fungi. XXXIII 269

eriophori Oud. has pycnidia only about 75 u diam., while S'.
eriophoricola Hollos has pycnidia 110-150 /x diam., with filiform
conidia 30-40 x 1 /x. 2) On Decodon verticillatus (L.) Ell., near
Trout Lake, Vilas Co., July 1, 1914. Coll. J. J. Davis. The pycnidia
are globose, brownish, thin- walled, about 125-175 /x diam., the
conidia hyaline, indistinctly several-septate, laxly curved, wider and
obtuse at one end, tapering at the other, approx. 2-3.5 (-4) x 35-
45 fjL. Not Septoria lythrina Peck which has filiform conidia. There
appear to be no previous reports of Septoria on Decodon. 3) On
Eupatorium sessili folium L. collected October 13, 1965 at Tower
Hill State Park, Iowa Co., and overwintered out-of-doors at Madi¬
son until mid-May 1966. The specimen as collected had well-marked,
rather small, orbicular or angled, fuscous spots, on which
epiphyllous pycnidium-like structures with poorly defined contents
were scattered. The spring of 1966 was in the main cold and dry,
but when examined the pycnidia were found to be filled with well-
developed conidia. The pycnidia are black, rather thick-walled,
ostiolate, subglobose, approx. 125-150 /x diam., the conidia hyaline,
straight or slightly curved, continuous or obscurely septate,
(10-) 12-17 (-22) X 1,4-1. 7 /x. Perhaps a cold-weather development
of Septoria eupatorii Rob. & Desm. which normally has conidia
about 25-35 x 1.5 /x. 4) On Ambrosia psilostachya DC., near Mazo-
manie, Dane Co., July 20, 1935. Coll. J. J. Davis. This seems close
to, perhaps identical, with Septoria ambrosicola Speg. The Wis¬
consin specimen has small, orbicular, light brownish spots, with
pycnidia approx. 50-100 /x diam., the conidia hyaline, mostly
strongly curved, tapering toward the apices, continuous or ob¬
scurely multiseptate, about 40-85 x 1-2 /x. S, ambrosicola is de¬
scribed as having the spots amphigenous, orbicular, determinate,
1-3 mm. diam., whitish, pycnidia 90-100 /x diam., the conidia hya¬
line, attentuate at both ends, subarcuate or merely fiexuous, con¬
tinuous, 50-100 X 1.5-2 /X.

Leptothyrium similisporum (Ell. & Davis) Davis normally
occurs as a definite parasite on the leaves of Solidago rigida L. and
other goldenrods. However, in a specimen on S. rigida, r^.ollected at
Madison, September 1, the fungus is confined to, but still profusely
present on, the outer margins of applanate, rounded, yellowish
galls produced by a midge (cecidomyiid) on the leaves. The writer
has, over the years, seen hundreds of such gall-infested leaves, but
never before have any of the galls been seen to bear a fruiting
fungus.

Melasmia ulmicola B. & C. has long been reported as occurring
on elms in Wisconsin and has been represented by a number of
specimens from Wisconsin and elsewhere in the Wisconsin Her-

270 Wisconsin Academy of Science, Arts and Letters [Vol. 56

barium. Recent collections of the conidial stage of Gnomonia ulmea
(Schw.) Thum., Cylindrosporella ulmea (Miles) v. Arx, have led
to a re-examination of the ‘‘Melasmia ulmicola'' specimens and it
appears highly likely that most, if not all, are really more or less
well-defined C. ulmea. The conidia of Melasmia ulmicola are said to
be minute and oblong-botuliform, but the overall description is in¬
adequate and the identity of the fungus remains somewhat
uncertain.

Monoceras kriegerianum (Bres.) Guba is the type species of
Monochaetia and Pestalotia” , p. 290 (1961). This fungus causes a
conspicuous leaf of fireweed, Epilobium angustifolium L. It has
been variously referred to Pestalotia, Monochaetia and Hyaloceras.
Cuba’s description of Monoceras and its type species is based in
part on a specimen collected by J, J. Davis near Luck, Polk Co.,
Wis., August 25, 1916.

Pestalotia (?) sp. is epiphyllous on small, sharply defined,
rounded, reddish-brown spots on Hypericum kalmianum L. collected
at Madison, July 29. The conidia are variously ciliate, from almost
straight to moderately curved, clear grayish, 3-septate, about 15-
18 X 5 ja. Except for the tiny (1-2 mm.) spots, the leaves are green
and vigorous appearing, so parasitism seems a possibility.

The Monilia stage of Monilinia fructicola (Wint.) Honey or¬
dinarily develops on the flowers and fruits of species of Prunus, but
what appears to be this stage is hypophyllous or large, orbicular,
purplish-brown lesions, up to 3 cm. diam., on shoot leaves of Prunus
americana Marsh. Collected at Madison, June 30.

Botrytis spp. have been noted on several additional hosts. These
are but more of a considerable number of so far undetermined,
parasitic-seeming members of this genus observed over many years
on a variety of plants: 1) On Rumex acetosella L., near Verona,
Dane Co., September 9. The fungus appears definitely parasitic on
pallid streaks of varying width which originate near the apices
of otherwise still green leaves. 2) On Ranunculus ahortivus L., near
Albany, Green Co., May 26. Botrytis has occasionally been found
on the rounded basal leaves of this host, as has been reported in
these notes. However, in a specimen collected May 26 near Albany,
Green Co., the bract-like leaves encircling the bases of the flower¬
ing pedicels were involved and the infection had spread to the
pedicels causing a conspicuous drooping of otherwise still healthy
appearing flowering parts of many plants. 3) On Geranium
maculatum L., near Leland, Sauk Co., July 1, 1965. On rounded,
sharply defined, brown lesions, about .7-1 cm. diam. 4) On Helian-
thus strumosus L., New Glarus Woods Roadside Park, Green Co.,

1967-68] Greene — Wisconsin Parasitic Fungi. XXXIII 271

June 16. All the plants in a large clone had the leaves blighted just
below the growing point. It did not appear frost damage could have
been responsible,

Acremonium sp, has overgrown sori of Puccinia heucherae
(Schw.) Diet, on Heuchera richardsonii R. Br. collected July 1 at
Faville Prairie near Lake Mills, Jefferson Co. In my Notes 29
(Trans. Wis, Acad. Sci. Arts Lett. 52: 239. 1963) there is mention
of a very similar, if not identical, fungus overrunning Kuehneola
uredinis and Pucciniastrum agrimoniae.

Cladosporium ( ?) sp. is epiphyllous on small, 1.5-3 mm. diam.,
rounded, very sharply defined reddish-brown spots on Acer penn-
sylvanicum L, (cult.) collected at Madison August 11, 1965. The
cylindric, grayish-olivaceous conidia are rather strongly echinulate,
about 11-16 X 6.5”6.5 ,/x, mostly uniseptate, but some with two septa,
borne on loosely fasciculate, often tortuously geniculate, brownish-
olivaceous, several-septate conidiophores, approx. 60-85 x 4-5 ft.
Perhaps parasitic, since there is no other obvious causal agent and
since, aside from the small spots, the leaves are in thriving
condition.

Cercospora camptosori J. J. Davis on Camptosorus rhizophyllus
(L.) Link was collected June 17 ,at Wyalusing State Park, Grant
Co., where Davis first found it in 1914 — a. few years later he col¬
lected an additional specimen at Werley, also in northwestern Grant
Co. An examination of numerous Wisconsin and extra-Wisconsin
specimens of this fern in the Wisconsin Herbarium shows no speci¬
mens bearing this fungus. Since the walking fern attracts much
attention and is frequently collected, and since according to Chupp
C. camptosori is known only from Grant Co., Wis., it would seem to
indicate that this fungus does indeed have a restricted range. In
view of the notable sporadicity of many parasites, it is interesting
that C. camptosori should have persisted for more than 50 years
within these narrow boundaries.

Cercospora (?) sp. occurs in profusion on rounded, reddish-
brown lesions on leaves of Steironema lanceolatum (Walt.) Gray
collected at Madison, September 1. The grayish-olivaceous conidio¬
phores are simple, with only a suggestion of geniculation, in more
or less dense fascicles, when they may be up to 80 /x in length, or
they arise from definite, subglobose stromata, in which cause the
measurable phore length is considerably less. Only a very few
mature Cercospora-type conidia have been observed. These are ob-
clavate or narrowly obclavate, subhyaline, 1-3 septate, about 25-35
X 2. 5-3. 5 with a conspicuous basal scar. Also present are con¬
siderable numbers of hyaline, narrowly cylindric or fusoid conidia,

272 Wisconsin Academy of Science, Arts and Letters [Vol. 56

mostly continuous, a few uniseptate, occasionally in short chains.
No conidia were seen attached in a large number of mounts
examined.

Cercospora sp. is epiphyllous on cultivated Syringa (“Blue
Hyacinth” — not in the S. vulgaris group) collected in the Univer¬
sity of Wisconsin Arboretum at Madison, October 4. The spots are
rounded, mottled brownish-cinereous, up to about 1 cm. diam., the
conidiophores sinuous, subgeniculate, clear brownish-olivaceous,
several-septate, about 45-75 x 4-5 y, and fascicled on small
stromata, the few conidia seen pallid olivaceous, rather broadly
obclavate, truncate at base, tip obtuse, about 60 x 5 y, 3-septate.
The only species with conidia of this type mentioned by Chupp as
occurring on Syringa is Cercospora amurensis Zilling, but it does
not otherwise resemble the Wisconsin specimen.

Cercospora (?) sp. is hypophyllous on small, rounded, immar-
ginate, pallid yellowish areas on Kuhnia eupatorioides L. collected
near Leland, Sauk Co., August 3. The conidiophores are scattered,
approx. 20-40 x 3.5-5 y, 1-2 septate, from slightly sinuous to sub-
geniculate, sometimes denticulate, clear light grayish-brown, the
conidia subhyaline, subfusoid to subcylindric, approx. 12-18 x 2.5-3
y, continuous or uniseptate. Scarcely typical Cercospora conidia,
but the specimen is perhaps somewhat immature and it seems likely
that longer conidia might be produced in time. Chupp does not
report any Cercospora on Kuhnia.

Solidago uliginosa Nutt., collected in northern Forest Co., Oc¬
tober 2, 1965, has the upper stem and smaller leaves profusely
covered with a Sarcinella-\We fungus which, while largely super¬
ficial, appears to have invaded the trichomes and is thus perhaps
weakly parasitic.

K. T. Harper, in the years 1960-63, made an intensive ecological
study of the climax maple-basswood forest of Wisconsin. As an
adjunct of this study he checked 114 more or less characteristic
higher plant species of the maple-basswood community in connec¬
tion with the fungus parasites reported over the years as occurring
on them in Wisconsin. He found that 104 of these higher plant
species were known to be attacked by one or more fungal parasites,
that a total of 339 species of parasitic fungi were concerned, and
that there was an average of more than 4 parasites for each of the
114 higher plants considered.

Additional Hosts

The following hosts have not been previously recorded as bear¬
ing the fungi mentioned in Wisconsin.

1967-68] Greene — Wisconsin Parasitic Fungi. XXX 11 1 273

Albugo tragopogonis (DC.) S. F. Gray on Ambrosia trifida L.
Iowa Co., Ridgeway, June 24, 1921. Coll. J. J. Davis & A. B. Sey¬
mour. Davis failed to report A. tragopogonis on this host and the
single specimen was overlooked until recently.

Peronospora ficariae Tul, on Ranunculus acris L. Manitowoc
Co., Point Beach State Forest, June 20, 1960. Coll. J. A. Reed.

Peronospora parasitica (Pers.) Fr. on Arabis canadensis L.
Sauk Co., near Leland, August 12.

Peronospora arthuri Farl. on Oenothera strigosa (Rydb.)
Mack. & Bush. Pierce Co., Spring Valley, July 20, 1925. Coll. J. J.
Davis,

Basidiophora entospora Roze & Cornu on Solidago gigantea Ait.
Dane Co., Madison, October 12, 1920. Coll. J. J. Davis. Davis did
not report this, but a scraping shows the sporangiophores so char¬
acteristic of this species.

Erysiphe CICHORACEARUM DC. on stems of Monarda punctata
L. Sauk Co., near Leland, September 12. Powdery mildew conidia
are regularly present on the leaves of this host, but this is the first
Wisconsin collection of cleistothecia. Also on Artemisia biennis
Willd. Dane Co., Madison, October 2.

Sphaerotheca humuli (DC.) Burr. vslt. fuliginea (Schl.) Salm.
on Erigeron strigosus MuhL, Sauk Co., near Leland, August 17.

Phyllactinia CORYLEa (Pers.) Karst, on Cornus obliqua Raf.
Dane Co., Madison, September 30. Coll. D, P. Mahoney.

Powdery mildews indet. (conidia only) have been noted 1) On
Trifolium arvense L. Barron Co., near Prairie Farm, September 19,
1965, and 2) On Salvia haematodes L. (cult.). Dane Co., Madison,
September 24.

Peckiella lateritia (Fr.) Maire on Lactarius indigo (Schw.)
Fr. Milwaukee Co., Milwaukee, October 1903. Coll. C. Thot.

Mycosphaerella spleniata (C. & P.) House on overwintered
leaves of Quercus alba L. Sauk Co., near Leland, May 13. The micro-
conidial stage, Phyllosticta livida Ell. & Ev., without doubt devel¬
oped parasitically in the previous season.

Coleosporium ASTERUM (Diet.) Syd. II on Solidago caesia L.
Milwaukee Co., Cudahy, October 24, 1965. Coll. C. T. Lind.

PUCCINIA ANDROPOGONIS Schw. I on Pentstemon pallidus Small.
Oneida Co., near Tripoli, June 19, 1965. Coll. & host det. F. S.
Crosswhite.

274 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Pellicularia filamentosa (Pat.) Rogers on Prenanthes
racemosa Michx. Dane Co., Madison, July 19.

Phyllosticta eminens H. C. Greene on Salix discolor Muhl.
Dane Co., Madison, September 5. The fungus is in rather small
amount and does not produce the conspicuous lesions so character¬
istic of the type specimen, but microscopic correspondence is close.

Phyllosticta virginiana (Ell. & Halst.) Seaver on Prunus
pumila L. Bayfield Co., Barnes, September 12, 1956. The specimen
was included in a large collection of Tranzschelia on this host and
overlooked until recently.

Phyllosticta decidua Ell. & Kell, on Agastache scrophulariae-
folia (Willd.) Ktze. Iowa Co., Gov. Dodge State Park, July 15.

Phyllosticta cacaliae H. C. Greene on Prenanthes alba L. Sauk
Co., near Leland, September 8.

Neottiospora umbelliferarum H. C. Greene on Cicuta maculata
L. Dane Co., Madison, July 6. Both pycnidia and conidia are some¬
what smaller on the average than in the type on Oxypolis (see
Trans. Wis. Acad. Sci. Arts Lett. 47 :113. 1958) , but the dimensional
differences would seem to be within the acceptable range.

Ascochyta equiseti (Desm.) Grove on Equisetum laevigatum
A. Br, Iowa Co., near Arena, August 5.

Ascochyta graminicola Sacc. on Setaria viridis (L.) Beauv.
Burnett Co., Roosevelt Twp., October 2, 1965. Coll. G. Patz.

Ascochyta nepetae J. J. Davis on Glecoma (Nepeta) hederacea
(L.) Trev. Dane Co., Madison, August 30. Referred here with some
doubt. Accompanying the typical uniseptate conidia are much more
numerous rod-shaped microconidia about 3-5.5 x 1-1.7 y, and some
pycnidia contain microconidia only.

Ascochyta compositarum J. J. Davis on Aster cordifolius L.
Sauk Co., near Leland, August 3. Also on Helianthus tuberosus L.,
Sauk Co., near Leland, August 17.

Darluca filum (Biv.) Cast, on Cronartium quercuum (Berk.)
Miyabe on Quercus ellipsoidalis Hill. Burnett Co., Webster, August
31, 1916. Coll. J. J. Davis.

Stagonospora albescens j. j. Davis on Carex trichocarpa Muhl.
Sauk Co., near Leland, September 8.

Septoria pentstemonicola Ell. & Ev. on Pentstemon pallidus
Small. Marathon Co., near Knowlton, June 19, 1965. Coll. & host
det. F. S. Crosswhite.

1967-68] Greene — Wisconsin Parasitic Fungi. XXXIII 275

Septoria rudbeckiae Eli & Halst. on Rudbeckia serotina Nutt.
Marathon Co., near Rothschild, August 4, 1965. Coll, M. Torin. On
a phanerogamic specimen in the University of Wisconsin
Herbarium.

Phaeoseptoria festucae R. Sprague var. muhlenbergiae
Sprague on Muhlenhergia tenuiflora (Willd.) BSP. Sauk Co,, near
Leland, September 12. On narrow, rather elongate, cinereous,
brown-bordered lesions. Sprague considered the species of
Phaeoseptoria to be essentially saprophytes, but the present speci¬
men suggests parasitic development.

PiROSTOMA CIRCINANS Fr. On Andropogon scoparius Michx.
Green Co., near Albany, September 23.

Hainesia lythri (Desm.) Hoehn. on Geum triflorum Pursh f.
pallidum Fassett. Dane Co., Madison, August 10. Although this
white-flowered form is not generally recognized as distinct it is
very constant, certain plants having been observed annually by the
writer over a 23 year period, and is evidently much less susceptible
to fungus attack, with the present instance being the first noted in
the entire period, in contrast to the species proper which is fre¬
quently infected. Also on Cary a cordiformis (Wang.) K. Koch. Sauk
Co., near Leland, September 7. The Sclerotiopsis stage of the fungus
has been earlier noted on this host in Wisconsin.

Sclerotiopsis concava (Desm,) Shear & Dodge on Vitis riparia
Michx. Dane Co., Madison, September 14, 1965. The Hainesia stage
has already been noted on this host in Wisconsin.

COLLETOTRICHUM GRAMINICOLA (Ces.) Wils. on Cenchrus pauci-
florus Benth. Dane Co., near Middleton, September 24, 1965. Coll.
G. Patz.

Phleospora anemones Ell. & Kell, on Anemone riparia Fern.
Trempeleau Co., near Dodge, October 2, 1965. Coll. J. Graham.

Monilia stage of Monilinia fructicola (Wint.) Honey on
flowers of Prunus cerasus L. Door Co., Sturgeon Bay, June 7. Coll.
& det. D. A, Biris.

Cladosporium ASTERICOLA J. J. Davis on Aster macrophyllus L.
Sauk Co., near Leland, August 17.

Cercospora nigricans Cooke on Cassia fasciculata Michx. Dane
Co., Madison, September 16,

Cercospora perfoliata Ell. & Ev. on Eupatorum sessilifolium
L. Sauk Co., near Leland, July 30. The spots here are more sharply

276 Wisconsin Academy of Science^ Arts and Letters [Vol. 56

defined than is usual with this species, but microscopically it cor¬
responds well.

Tuberculin A persicina (Ditm.) Sacc. on Puccinia podophylli
Schw. I on Podophyllum peltatum L. Green Co., near Albany, May

26.

Additional Species

The fungi mentioned, with several exceptions, have not been
previously reported as occurring in Wisconsin. Some name revisions
of earlier reported entities are listed here as it is felt they are
more likely to come to the attention of the reader under this
heading.

'Dimeriella erysiphoides (Ell. & Ev.) Farr (Venturia erysi-
phoides Ell. & Ev.) appears to be the correct name of the fungus
on Andropogon scoparius Michx., reported by me as Venturia
sporoboli Greene.

Strom ATINIA gladioli (Drayton) Whetzel on Gladiolus hor-
tulanus Bailey. Columbia Co., Cambria, September 16. Coll. & det.
R. Pinney.

Coleroa sporoboli (Greene) Barr, replaces Venturia sporoboli
H. C. Greene according to M. E. Barr who has made a revisionary
study of the Venturiaceae. She states (personal communication)
‘The fungus in its subcuticular development, superficial appear¬
ance, and other characters, agrees with modern concepts of Coleroa
rather than VenturiaP This fungus occurs on Sporoholus cryptan-
drus (T.) Gray, S\ heterolepis Gray and Oryzopsis asperifolia
Michx, in Wisconsin.

Coleroa rubicola (Ell. & Ev.) Muller is the name applied by
M. E. Barr to Coccochora rubi J. J, Davis which occurs on Rubus
canadensis L. and R. hispidus L, in Wisconsin. In a personal com¬
munication she states that the latter is but a stromatic form of
Coleroa rubicola,

Seynesiella juniperi (Desm.) Am. is, according to M. E. Barr,
the name properly applied to the fungus previously reported as
Asterina ciopressina Cooke which occurs on Juniperus communis
L. var. depressa Pursh in Wisconsin.

Puccinia brachypodiuphoenicoidis Guyot & Malencon var.
DAViSii Cummins & Greene on Oryzopsis asperifolia Michx, Various
stations, mostly in northern Wisconsin. The rust on this particular
host was reported in former Wisconsin lists as Puccinia pygmaea
Erikss., but Cummins & Greene (Mycologia 58:719. 1966), in a

1967-68] Greene — Wisconsin Parasitic Fungi. XXXIII

277

paper entitled review of the grass rust fungi that have uredial
paraphyses .and aecia on Berberis-Mahonia”, have erected this
segregate and selected as the type a specimen collected by J. J.
Davis at Agenda, Ashland Co., October 13, 1911.

Gymnosporangium asisticum Miyabe ex Yamada III on
Juniperus virginiana L. Columbia Co., Gibraltar Rock County Park
near Okee, M,ay 13. Coll & det, J. L, Cunningham. Confirmed by
F. D, Kern. Probably a new host worldwide, according to Dr.
Cunningham.

Phyllosticta MALI Prill & Delacr. on Pyrus malus L. Dane Co.,
near Pine Bluff, August 4, 1960. The Wisconsin specimen corre^
sponds closely with No. 1676 “Herbarium Mycologicum Romani-
cum^\ distributed as this species. The conidia are oval, about 6 x 3 /x,
and of a greenish tint. In the Saccardian description conidial dimen-
sions are given as 6.5-8 x 4-4.5 ,/x, but in other particulars the speci¬
mens correspond fairly well with the description.

Phyllosticta primulicola Desm. on Primula ohconica Hance
(cult.). Dane Co., Madison, July 7, The small, black pycnidia are
about 100-125 /a diam., the hyaline conidia ellipsoid to subfusoid,
approx. 3.5-5 x 1.5-2. 5 Grove in “British Stem and Leaf Fungi’'
p. 34, states concerning this species “Very common, but the
pycnidia are nearly always empty”. This seems to be the case with
the Madison specimen as well, since only one of the pycnidia ex¬
amined had conidia, but in that one they were numerous and
well-developed,

Coniothyrum wisconsinensis sp. nov.

Maculis distinctis, plerumque tantum uno vel duo in frondibus,
obscuro-cinereis vel brunneis vel rufo-brunneis, marginibus
angustis fuscis, subcirculis, ca. 2-4 mm. diam, ; pycnidiis epiphyllis,
gregariis, nigris, muris crassis, ostiolatis, subglobosis, ca.
(75-) 100-125 (-150) ,/A diam.; conidiis levibus, Claris, cano-
olivaceis, late ellipticis vel subcylindraceis vel subglobosis interdum,
(3.5-) 4-5.5 (-6.5) x 2.5-3.5 ,/a.

Spots sharply defined, usually only one or two per leaf, dull
cinereous to brown or reddish-brown with narrow darker border,
rounded, about 2-4 mm, diam.; pycnidia epiphyllous, gregarious,
black, rather thick-walled, ostiolate, approx. (75-) 100-125 (-150)
/A diam. ; conidia smooth, clear grayish-olivaceous, broadly elliptic or
subcylindric, or occasionally subglobose, (3.5-) 4-5.5 (-6.5) x 2.5-
3.5 /A.

On living leaves of Ulmus americana L. University of Wisconsin
Arboretum, Madison, Dane County, Wisconsin, U. S. A., July 19,

278 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1966. Two other specimens were collected in the Arboretum on the
same date within a quarter of a mile of the type. There are further
collections from Dane, Iowa and Sauk counties.

Coniothyrium ulmi Tharp (Mycologia 9:116. 1917) had whitish,
angular spots ,5-3 mm. diam., with the conidia brown and ovate,
4-6 X 2-2.5 juL. It occurred on cultivated Ulmus campestris L.

Septoria minuta Schroet. on Luzula multiflora (Ehrh.) Le-
Jeune, Sauk Co., near Leland, May 28. The small black pycnidia,
50-75 p. diam., are subseriate to crowded on conspicuous, dark
reddish-brown spots. The conidia are hyaline, appear continuous,
and are mostly slightly curved, (1 5-) 18-23 (-28) x 1.7-2. 3 p.

Septoria clementsii sp. nov.

Maculis obscuro-viridulis, angulatis, magnitudinibus variis, saepe
confluentibus, in foliis languidis; pycnidiis hypophyllis, gregariis,
numerosis, nigris, muris crassis modice, subglobosis vel subap-
planatis, ca. (75-) 85-125 (-150) p diam.; conidiis hyalinis, con-
tinuis vel multiseptatis obscuris, sinuosis vel curvis varie, saepe
attenuatis in apicibus unicis, ca. (45-) 55-75 (-90) x (1-)1.5-
2(2.5) p.

Spots dull greenish, angled, variable in size and often confluent
on otherwise faded leaves; pycnidia hypophyllous, gregarious,
numerous, black, moderately thick-walled, subglobose or somewhat
flattened, approx. (75-) 85-125 (-150) p diam,; conidia hyaline,
continuous or appearing faintly multiseptate, sinuous to variously
curved, often tapered strongly at one end, approx. (45-) 55-
75 (-90) X (1-) 1.5-2 (-2.5) p.

On languishing current season’s leaves of Aralia nudicaulis L.
Gov. Dodge State Park, Iowa County, Wisconsin, U. S. A., Septem¬
ber 21, 1966.

Septoria clementsii was first collected in Wisconsin in 1962 and
was discussed at some length in my Notes 30 (Trans. Wis. Acad.
Sci. Arts Lett. 53 :183. 1964) . As pointed out there, it is considered
to be the same as Septoria macrostoma Clements (No. 55 in
Clements’ “Cryptogamae Formationum Coloradensium”, issued in
1906) and apparently represented only by the type specimen in the
U. S. National Fungus Collections. As stated ''According to Dr.
C. R. Benjamin . . . publication was effected through distribution
of the exsiccati, but it appears that in the case of Septoria
macrostoma the Latin indication of host appearing on the label is
insufficient to be considered an adequate description and therefore
the name is not valid.” The accompanying description is offered to
remedy this and at the same time indicate Clements’ connection
with the species. The reason for his choice of the specific epithet

1967-68] Greene- — Wisconsin Parasitic Fungi. XXXIIl

279

^^macrostoma^^ is not clear for there is no well-defined ostiole in
this species. When the pycnidia are viewed by transmitted light the
greater depth of wall looked through at the periphery does give the
effect of a dark ring surrounding a lighter area, so perhaps the
name was applied because of this.

Dothistroma PINI Hulbary on Pinus nigra Arn. var. austriaca
Aschers. & Graebn. Jackson Co., near Hixton, June 6. Coll. & det.
A. J. Pray. It appears that the previous report of the similar Leca-
nosticta acicola (Thum.) Syd. on this host is erroneous and prop¬
erly referable to Dothistroma, Lecanosticta does, however, occur
in Wisconsin on Scotch pine, Pinus sylvestris L.

Cylindrosporella caricina sp. nov.

Maculis conspicuis, castanets, centris pallidioribus saepe, orbicu-
laribus vel ovatis, saepe confluentibus, ca. 1-2.5 cm. diam. ; acer-
vulis epiphyllis, subcuticularibus, planis, inconspicuis, sparsis vel
gregariis, ca. 100-175 /x diam.; conidiophoris hyalinis, exilibus,
saepe curvis, confertis, ca. 12-18 x 1-1.5 ,/x; conidiis hyalinis, rectis,
angusto-cylindraceis, ca. 7-9 x 1 /x.

Spots conspicuous, chestnut-brown, often with lighter-colored
centers, orbicular to ovate, often confluent, about 1-2.6 cm. diam. ;
acervuli epiphyllous, subcuticular, flattened, inconspicuous, scat¬
tered to gregarious, about 100-175 jx diam.; conidiophores hyaline,
slender, often curved, closely ranked and compacted approx. 12-18 x
1-1.5 /t; conidia hyaline, straight, narrow-cylindric, approx. 7-9
X 1

On living leaves of Carex lacustris Willd. Tower Hill State Park,
Iowa County, Wisconsin, U. S. A., August 24, 1966.

This fungus was collected in small amount at the same station
in 1957 and was mentioned in my Notes 24 (Trans. Wis. Acad. Sci.
Arts Lett. 47 : 108. 1958) . In 1966 the organism occurred in great
profusion over an area of an acre or more. As noted in 1958, the
large air spaces of the host leaves are well filled with a coarse, rami¬
fying mycelium which is assumed to have been produced by the
just described fungus,

Gloeosporidiella cercidis sp. nov.

Maculis magnis, conspicuis, suborbicularibus vel elongatis varie,
plerumque ca. 3-5 cm, diam., centris rufo-brunneis splendidis, mar-
ginibus nigris, obscuris ; acervulis hypophyllis, gregariis, immersis,
ca. 75-150 /i diam. ; conidiophoris inconspicuis, fere obsoletis ; conid¬
iis in cirrhis, hyalinis, formis variabilibus, subcylindraceis, ovoideis,
allantoideis late, vel subfusoideis, etiam in magnitudinibus varie,
ca, 6-12 X 2.5-5

280 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Lesions large, conspicuous, suborbicular to variously elongate,
mostly about 3-5 cm. diam., with bright reddish-brown central por¬
tions and rather poorly defined blackish margins ; acervuli hypoph-
yllous, clustered in groups, deeply imbedded in the mesophyll,
approx. 75-150 p. diam.; conidiophores inconspicuous, almost obso¬
lete; conidia extruded in short cirrhi, hyaline, variable in shape,
subcylindric, ovoid, broadly allantoid, or subfusoid, and also vari¬
able in size, approx. 6-12 x 2.5-5 p.

On living leaves of Cercis canadensis L. University of Wisconsin
Campus, Madison, Dane County, Wisconsin, U. S. A., August 16,
1966.

The lesions dry out quickly and the central portions often shred
and break away. The fungus appears highly parasitic and I have
found no report of any similar organism on Cercis. On some leaves
the acervuli are strongly developed on portions of the principal
veins.

Ramularia ANGELICAS Hoehn. on Angelica atropurpurea L. Sauk
Co., near Leland, September 30.

Cercospora murina Ell. & Kell, on Viola pennsylvanica Michx.
Sauk Co., near Leland, September 7. In my Notes 24 I mentioned
the occurrence of what was probably this species on Viola canaden¬
sis L. from Sawyer Co., but the specimen was rather poor and no
formal record was made at that time.

Beniowskia sphaeroidea (Kalchbr. & Cke.) Mason is the name
applied by S. J. Hughes (Can. Jour. Bot. 36: 742. 1958) to the
fungus, parasitizing Panicum virgatum L., misnamed Botrytis
uredinicola by Peck, and discussed at length in my Notes 28 (Trans.
Wis. Acad Sci. Arts Lett. 51 : 77. 1962).

PRELIMINARY REPORTS ON THE FLORA OF WISCONSIN
NO. 59. PLANTAGINACEAE— PLANTAIN FAMILY'

Melvern F. Tessene
Department of Botany and Botanical Gardens
University of Michigan, Ann Arbor

The Plantaginaceae is best known for its weedy members, al¬
though these constitute a very small percent of the total number
of species. Two of the three genera in the family are native in
Wisconsin. Littorella, a specialized aquatic, is a rare member of
the northern flora. The genus Plantago is represented by five na¬
tive and four introduced species, one or more of which can readily
be found almost anywhere in the state. Plantago cordata Lam. is a
rare plant of small streams in the southern half of the state and
should be actively looked for by anyone who ventures into the field.

Fortunately, for those who are unfamiliar with the family, the
Wisconsin species are fairly distinct once one learns which charac¬
ters are important. The perennial confusion of P. major L. and
F. rugelii Dene., especially of sterile or depauperate specimens, can
only be resolved by close examination of the material. It would be
pretentious to assume the keys in this report will “work’' for every
specimen that may be collected in the state, but I hope they will
prove useful in the identification of most specimens. To facilitate
correct determinations, I have illustrated the habit and important
morphological features of each species. These drawings are ideal¬
ized representations drawn from many different specimens and are
to be thought of as representing only the “average” condition. All
species are plastic and highly subject to environmental variation.
The range of variation, as shown by the specimens at hand, is
given in the description of each species. I have tried to restrict
these discussions to include only those data which I believe accurate
either through personal observations or experiments. In a few
cases, I have relied on other authors for supplementary informa¬
tion. Chromosome counts, except where indicated, have been made
by me. All localities are represented by specimens which I have
seen. Reports in the literature, where no vouchers were available

1 Research for this paper has been supported in part by NSF Grant GB-3366 (Sys¬
tematic and Evolutionary Biology Prog-ram) and by the University of Michig-an Gradu¬
ate Student Research Fund.

281

282 Wisconsin Academy of Science, Arts and Letters [Vol. 56

to me, have been omitted. Except for P. cordata, localities known
only by county are not given.

I wish to express my sincere gratitude to the directors and cura¬
tors of the following herbaria who so generously made their speci¬
mens available to me: Illinois Natural History Survey, Iowa State
University-Ames, Michigan State University, Milwaukee Public
Museum, Ohio State University, University of Illinois-Urbana,
University of Michigan, University of Minnesota, University of
Wisconsin, University of Wisconsin-Milwaukee, Wisconsin State
University-Oshkosh and Wisconsin State University-Whitewater.
My special thanks are given to Dr. Warren H. Wagner, Jr., for his
continuing advice and encouragement in this and other projects
and for critically reading the manuscript ; Dr. Hugh H. litis with¬
out whose assistance this paper would have been impossible; Dr.
Edward G. Voss for assisting in the identification of the Littorella
specimens and for reading the manuscript; Dr. Neil Harriman who
made several collecting trips in the poorly collected Lake Winne¬
bago area; and Dr. Ronald L. Stuckey with whom I frequently
discussed the material in this paper.

Key to Fertile Specimens of the Plantaginaceae of Wisconsin

1. Flowers unisexual; pistillate flowers in a group in the axil of a
single bract, each flower having 1-4 sepals; staminate flowers
solitary, terminal, sepals four; fruit an achene; leaves terete
or subterete, linear; stolons present --Littorella americana (1)
1. Flowers bisexual (except in gynomonoecious species) each
flower subtended by a bract; all flowers having four sepals (two
connate in P. lanceolata) ; fruit a circumscissile capsule; leaves

flattened, linear to cordate; stolons absent _ Plantago

2. Leaves cordate to broadly elliptic, definite blade and petiolar
portions present; sepals and bracts glabrous.

3. Major roots fleshy, 0.5-1. 3 cm thick; peduncle 3-5 mm
wide, central cavity % or more of total diameter; seeds
2 per capsule; major veins of leaf not parallel to margin,

appearing to arise from a “midrib”, leaves leathery _

_ Plantago cordata (2)

3. Major roots filamentous, 0. 1-0.3 cm thick; peduncle 1-2
mm wide, central cavity less than % of total diameter;
seeds 4-12 per capsule; major veins of leaf parallel to
margin, leaves thin.

4. Capsules oblong, dehiscence near base; seeds 4-9 per
capsule; bracts and sepals lanceolate-attenuate; leaves
broadly elliptic, margins bearing 3-7 small teeth,
glabrous _ P. rugelii (3)

1967-68] Tessene — Reports on Flora in Wisconsin

283

4. Capsules rhombic-ovate, dehiscence medial; seeds 8-12
per capsule ; bracts and sepals ovate-obtuse ; leaves
cordate-ovate, mangius entire, glabrate or hirsutulous
_ P. major (4)

2. Leaves linear to spatulate, definite blade and petiolar portions
absent or inconspicuous; sepals and/or bracts usually
pubescent.

5. Stem freely branching, internodes 0.5-2 cm long; leaves
opposite; seeds 2.8-4 mm long; spikes often appearing

capitate _ _ _ P. indica (5)

5. Stem rarely branching, internodes absent or inconspicu¬
ous; leaves rosulate; seeds less than 3 mm long; spikes
elongate.

6. Leaves linear, pilose, major veins 3; flowers conspicu¬
ously zygomorphic; stamens and style usually shorter
than posterior corolla lobe; lobes variously pigmented
at corolla throat; seeds lightly sculptured, medially
constricted.

7. Bract 1.2-4 times as long as sepals ; flowers spiraled
on peduncle; tips of petals rounded; seeds 2.2-3 mm
long _ P. aristata (6)

7. Bract usually shorter than or just equaling sepals;

flowers 4-ranked on peduncle; tips of petals acute;
seeds 1.6-2 mm long _ P. patagonica (7)

6. Leaves spatulate or lanceolate, hirsute or glabrous, ma¬
jor veins 3,5,7 ; flowers actinomorphic or only slightly
zygomorphic ; stamens and style usually exceeding
posterior corolla lobe (unless flowers are cleistoga-
mous) ; lobes translucent or pigmented only at tip;
seeds rarely sculptured, not constricted.

8. Leaves lanceolate; 2 sepals adjacent to bract con¬
nate ; pigmented areas of bracts and sepals a narrow

stripe; seeds 2 per capsule, hirsutulous _

_ P. lanceolata (8)

8. Leaves spatulate; all 4 sepals free; pigmented areas
of bracts and sepals % of total area; seeds 3 (2-4)
per capsule, glabrous.

9. Major veins 5,7 ; leaves 2.5-4 cm wide; bracts and
sepals glabrous; flowers contiguous along spike;
stamens well developed; root system extensive

_ P. media (9)

9. Major veins 3; leaves 0.6-1. 5 cm wide; bracts and
sepals hirsute; flowers separated by internodes

284 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 1. Habit drawings of some Wisconsin species in the Plantaginaceae :
A) Plantago media; B) P. aristata; C) P. patagonica; D) P. indica; E) P.
lanceolata; F) P. virginica; G) Littorella americana.

1967-68] Tessene — Reports on Flora in Wisconsin

285

along spike ; stamens frequently aborted ; root

system sparce and poorly developed _

_ P. virginica (10)

Key to Sterile Specimens of the Plantaginaceae of Wisconsin

1. Leaves terete or slightly flattened on adaxial side, linear ca.
1 mm wide, fleshy, glabrous, major vein 1, inconspicuous;

stolons present ; plant aquatic _ Littorella americana ( 1 )

1. Leaves flattened, linear to cordate, 0.5-19 cm wide, thin to
leathery, never fleshy, glabrous to pilose, major veins 3-9 ;

stolons not present; plant usually terrestrial _ Plantago

2. Leaves opposite, cauline; internodes 0.5-2 cm long; stem
freely branching _ P. indica (5)

2, Leaves alternate, rosulate; internodes usually absent or less
than 0.5 cm long; stem a corm or caudex, rarely branching.

3. Leaves cordate, ovate or broadly elliptic with definite
blade and petiole; blades 2-19 cm wide; major roots
fibrous, or branched and fleshy.

4. Major roots fleshy, 0.5-1. 3 cm. thick; leaves leathery,
spatulate to cordate-ovate; major veins appearing to
arise from a ‘‘midrib’’ i.e. pinnate, not parallel to mar¬
gin _ P. cordata (2)

4. Major roots filamentous, 0. 1-0.3 cm thick; leaves thin
to chartaceous, cordate to broadly elliptic; major veins
parallel to leaf margin.

5. Leaves broadly elliptic, never cordate; major veins
5,7, margin bearing 3-7 small teeth, base of petiole

usually purple; plant glabrous or glabrate _

_ P. rugelii (3)

5. Leaves ovate-cordate, major veins 3,5, (7), margin
entire, undulate, base of petiole usually green ; plant

hirsutulous or glabrate _ P. major (4)

3. Leaves linear, lanceolate spatulate without definite blade
and petiole; blades 0.5-4 cm wide; major roots in a tap
system or (in old P. lanceolata) fibrous and arising from
a stout caudex.

6. Leaves spatulate, hirsute, margins entire or bearing
3-7 pectinate teeth.

7. Major veins 5,7 ; leaf 2.5-4 cm wide, often revolute,
appressed to ground ; plant perennial, often forming

caudex to 1.5 cm thick _ P. media (9)

7. Major veins 3; leaf 0.6-1. 5 cm wide, erect or spread¬
ing; plant a short-lived annual with poorly devel¬
oped root system, never a caudex__P. virginica (10)

286 Wisconsin Academy of Science, Arts and Letters [Vol. 56

6. Leaves linear or lanceolate, pilose or glabrous, margins
entire or bearing 5-7 small teeth.

8. Leaves lanceolate, glabrous, 0.5-4 cm wide, dark
green, margins bearing 5-7 small teeth, major veins
5 (3,7) ; plant perennial often forming caudex 4-6
cm long, internodes absent ; secondary branches fre¬
quent on large specimens _ P. lanceolata (8)

8. Leaves linear, pilose, 0.2-0. 6 cm wide, light green
or silvery, margins entire, major veins 3; plant an-
annual, occasionally forming short aerial stem with
or without internodes; branches absent.

9. Plants light green, thinly pilose; leaves of juve¬
nile plants erect or slightly spreading; internodes
short (2-5 mm) when present; secondary root

system well developed _ P. aristata (6)

9. Plants silvery-green, thickly pilose; leaves of
juvenile plants broadly spreading or appressed
to ground ; internodes absent ; secondary root sys¬
tem poorly developed _ P. patagonica (7)

Key to Flowers and Fruits of the Plantaginaceae
OF Wisconsin

1. Flowers unisexual, spike interrupted, i.e. bearing cluster of pis¬
tillate flowers at base and one terminal staminate flower sepa¬
rated 0.8-2 cm along floral axis; fruit an achene _

_ Littorella americana (1)

1. Flowers bisexual, gynomonoecious or apomictic, flowers evenly
distributed along spike; fruit a circumscissile capsule. _Pte^apo

2. Seeds 4-16 per capsule, testa more or less deeply sculptured;
anthers conspicuously horned; bract and sepals glabrous;
stigmatic hairs scattered, never in two rows, absent at corolla
level.

3. Bract and sepals attenuate; capsule oblong, 6.5-8 mm
long, dehiscence near base; lower valve below sepals after
dehiscence; 4-9 seeds per capsule; seeds 1. 8-2.3 mm long;

floral outline with conspicuous narrowing near base _

_ P. rugelii (3)

3. Bract and sepals obtuse-acute; capsule rhombic-ovate, 4-5
mm long, dehiscence medial ; lower valve above sepals after
dehiscence; 8-16 seeds per capsule; seeds 0.6-1 mm long;

floral outline ovate _ P. major (4)

2. Seeds 1-4 per capsule, rarely 5, testa smooth or very lightly
sculptured; anthers cordate or horned (P. indica) ; bract and
sepals glabrous-pilose; stigmatic hairs scattered or in two
parallel rows, usually present at corolla level.

1967-68] Tessene — Reports on Flora in Wisconsin

287

4. Stamen and style length 0.5 mm or less ; sepals and bracts
pilose; petals with dark pigmentation at corolla throat;
capsules with constriction above line of dehiscence; seeds
slightly sculptured; face of seeds white with one or more
dark bands.

5. Bract 1.2-4 times as long as sepals; 4-8 pigmented
areas on each petal near corolla throat ; seeds 2.2-3 mm
long, face with 2 concentric bands _ P. aristata (6)

5. Brace shorter than or equaling sepals; 1 pigmented

spot on each petal near corolla throat; seeds 1.6-2 mm
long, face with one dark band _ P. patagonica (7)

4. Stamen and style length 1 mm or more or aborted ; sepals
and bracts glabrous or hirsute; petals without dark pig¬
mentation at corolla throat or only near tip ; capsules not
constricted; seeds smooth; face of seeds usually lacking
dark bands.

6. Two sepals next to bract connate; pigmented areas on
bracts and sepals a narrow stripe less than % total
width ; petals, bract and the fused sepals brown at tip ;

capsule dehiscence basal ; seeds hirsutulous _

_ P. lanceolata (8)

6. All sepals free ; pigmented areas on bract and sepals at
least % of total width; petals, bract and sepals green
or translucent throughout; capsule dehiscence medial;
seeds glabrous.

7. Bract and sepals glabrous; corolla tube extending
beyond calyx; planar area of seed inconspicuous or
only partially developed.

8. Style exceeding stamens ; bract smooth, ovate, tip
, rounded; seeds 2 per capsule, 2.8-3 mm long;
planar area visible only at one end ; flowers odor¬
less _ P. cordata (2)

8. Style ca. as long as stamens; bract keeled,

lanceolate, tip acute; seeds 3 (2-4) per capsule,
1. 8-2.1 mm long, planar area not well defined;
flowers fragrant _ P. media (9)

7. Bract and sepals hirsute; corolla tube equalling or
shorter than calyx; planar area of seed %-% of
face.

9. Stigmatic hairs evenly distributed; petals with

distinct midrib ; bract keeled, chartaceous margin
extending to tip; seeds 2.8-4 mm long, 2 per
capsule _ P. indica (5)

9. Stigmatic hairs in two rows (or absent in
apomictic flowers) ; petals membranaceous

288 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Figure 2. Habit drawing's of some Wisconsin Plantagos: A) Plantago major;
B) P. rugelii; C) P. Cordata; a,b,c) Leaves of respective species.

1967-68] Tessene — Reports on Flora in Wisconsin

289

throughout, i.e. without a midrib; bract rounded,
chartaceous margin not reaching tip; seeds 1.6-
1.9 mm long, 3 per capsule _ P. virginica (10)

Littorella bergius

1. Littorella Americana Fern. Rhodora 20 :62 1918.

Map 2, Figs. IG and 3.
L. uniflora (L.) Aschers. sensu Gleason (1962) ; and Gleason
and Cronquist (1963).

L. uniflora v. americana (Fern.) Gl.

Plant a perennial herb. ROOTS : Fibrous ca. 1 mm thick. STEM :
compact, may form a short caudex, frequently producing stolons
0.5-4 cm long. LEAVES: Rosulate, linear, terete or slightly flat¬
tened on adaxial side, truncate, with only one major vein, this only
rarely visible on lower % of leaf; submerged leaves 1.3-3. 7 cm
long, erect; emersed leaves 2-8 cm long, arching. INFLORES¬
CENCE: Total length 0.5-3 cm, flowers unisexual. STAMINATE
FLOWERS : Terminal, only one per inflorescence ; bract rounded,
transparent; sepals 4, fleshy, often tinted light pink-brown, pig¬
mented area of total area, glabrous, 2. 8-3. 5 mm long, slightly
keeled; corolla 4-lobed, erect at anthesis, transparent, projecting
1-1.7 mm beyond calyx; filaments 9-12 mm long; anthers ca. 2.5
mm long, tip rounded, base cordate, versatile, purple and/or yellow.
PISTILLATE FLOWERS: In groups of 2-5 at base of inflores¬
cence, hidden except for stigmas by leaves, entire cluster subtended
by a single large, obtuse, transparent bract; sepals 1-4, linear,
acuminate, transparent ; corolla 3-4 toothed, teeth erect, attenuate ;
style filamentous 0.8-2. 5 cm long ; stigmatic hairs in one spiralling
row, FRUIT: Achene, oblong, ca. 2 mm long; apparently rarely
sets fruit. PHENOLOGY : Flowering June-August.

Fernald (1918) refers to Littorella americana as ‘‘One of the
rarest plants of the North American flora . . It is surely un¬
common but may often be overlooked or confused with other
aquatics as it rarely blooms (For keys to vegetative specimens
and morphologically similar aquatic plants, see Voss 1967.) Fas-
sett (1934) referred to it as “. . . a characteristic and abundant
plant in many lakes of northern Wisconsin.’’ Voss (1965) dis¬
cusses the species and gives a description of material from the
Upper Peninsula of Michigan. Unfortunately, I have not seen the
plant in the field but I do have a large collection of living plants
collected at Cusino Lake (Michigan) by Ronald L. Stuckey. The
data on this species, as given below, are based upon these
collections.

290 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Littorella americana

Figure 3. Flowers of Littorella americana.

Littorella is known from lakes in Maine, Michigan (Voss 1965), .

Minnesota (Lakela 1958), Newfoundland, New York (Muenscher |
1934), Nova Scotia, Ontario, Vermont and Wisconsin (Map 2). ij

1967-68] Tessene — Reports on Flora in Wisconsin 291

The name, Littorella, referring to the shore, is very appropriate
as it is only in the littoral zone that it is found in bloom. Voss
(pers. com.) reports that the Cusino Lake locality, where flower¬
ing specimens were collected in 1964, was under 2-3 feet of water
in 1965 and 1966. The plants were still there, but without flowers.
Experiments at the University of Michigan Botanical Gardens
have shown that the plants require cool temperatures when
emersed and cool or cold water when submersed. In cultivation,
flowering can only be induced in emersed plants and under long-
day conditions. The cultivated specimens were considerably larger
than any collected in the field. Although I had many plants bloom¬
ing simultaneously (all from the same clone), they did not set
fruit. This indicates, perhaps, a genetically controlled self¬
incompatibility. This may also explain the rarity of fruit-set in
the field. Reproduction is mainly vegetative by means of stolons.
New plants are readily obtained by dividing old individuals or by
cuttings from stolons.

PLANTAGO L.

SECT. PALEOPSYLLIUM Pilger

2. PLANTAGO CORDATA Lam. Illustr. Genr. 1:338. 1791.

Heart-leaved Plantain Map 1, Figs. 2C, 4, 6A and 7A.

P. kentuckensis Michx.

P. canadensis Hort.

Plant a perennial herb. ROOTS: Primary massive and fleshy,
0. 5-1.3 cm thick, branching several times ; secondary roots fibrous.
LEAVES : Spatulate in winter rosette, 1-3 cm wide ; cordate-ovate
but never truly cordate in late spring and summer rosette, 8-19
cm wide, 12-23 (50) cm long, lamina with a leathery texture,
petiole base often purple; major veins (3) 5,7 (9), not parallel to
margin but appearing to arise from a '‘midrib” I/2 to l/s the dis¬
tance up from the base of the expanded blade; leaf margin entire
or undulate. INFLORESCENCE: Total length including peduncle
12-40 cm, glabrous, central cavity % total diameter of peduncle,
2-4 flowers per cm. FLOWERS: (Fig. 6A) Obovate in outline;
sepals 2-2.3 mm long; bract 5/6 the length of sepals, obovate, tip
truncate, pigmented area bordered on sides by very narrow hyaline
margin, glabrous; petals elongate-deltoid, slightly revolute, trans¬
lucent, spreading at anthesis; style length 1.5-2 times the stamen
length, stigmatic hairs present at corolla level ; anthers cordate at
base, yellow. CAPSULE: (Fig. 7A) Ovate, 9-11 mm long, de¬
hiscent below the middle, lower valve of capsule equalling sepals
after dehiscence. SEEDS : 2 per capsule, tan — brown, testa smooth,
mucilaginous coat heavy, 3-3.8 mm long, planar area conspicuous

292 Wisconsin Academy of Science, Arts and Letters [Vol. 56

on one end of seed, obscured on other end, hilum and micropyle
separated. (N = 12) PHENOLOGY : Flowering mid-April to early
June (rarely in late fall). Fruit matures 1-3 weeks after anthesis.

Plantago cordata is a rare and fascinating plant. It has been
collected so infrequently in the past 50 years that whenever a new
locality is discovered or an old one rediscovered, it is either pub¬
lished (Svenson 1935, Chute 1942, Harper 1944, 1945) or cher¬
ished as a well-kept secret. So few botanists have seen it alive
that I have heard respected workers question whether it still
exists and whether it is even a distinct species.

The plant is quite similar, morphologically, throughout its range
but varies slightly in size. The species is endemic to eastern United
States and Ontario, excepting the coastal plain and northern for¬
ests, but most known localities are in the Great Lakes Region. I
have seen P. cordata alive only in Adams Co., Ohio. Data on the
biology of the species, as given below, are from plants of this local¬
ity. To my knowledge, there are only three or four other known
extant localities. It was frequently collected in southeastern Wis¬
consin in the 1880’s and 1890’s but the last known collection is
from Milwaukee Co. dated 1938. Wadmond’s collection from Som¬
ers, Kenosha Co. (MINN, WIS) includes photographs taken in the

Figure 4. Plantago cordata and habitat in Kenosha Co., Wisconsin (Wadmond,
June 30, 1899, near Sommers: MINN).

1967-68] Tessene — Reports on Flora in Wisconsin

293

field (Fig. 4), The distribution in Wisconsin (Map 1) is, as
throughout its range, correlated with limestone areas. From my ob¬
servations, it appears that the plants can not tolerate bright sun¬
light even if they are in very wet areas. This lack of tolerance may
be a factor in their decline, as the opening of the forests together
with the draining of low areas destroyed their habitats.

Blooming starts in late April and early May when the plant is
still in the winter-rosette condition (Fig. 2C) : Later inflorescences
and infructescences are subtended by summer leaves (Fig. 2c),
The induction of flowering is apparently brought about under
short-day conditions and young inflorescences are often produced
in the fall for the following spring. These young spikes are well
protected by the imbricate petiole bases. Meiosis may occur in the
fall but usually does not occur until spring. The important factor
in inducing flowering is evidently a cold period. Under constant
conditions, the plants will eventually develop a winter-rosette but
will then remain quiescent until subjected to a cold treatment.
Individuals will occasionally bloom again in the fall after a cold
snap. There is no dormancy in the seeds at maturity. They must
germinate rapidly as they only live about a month in storage at
40 °F. They will not germinate under water. A detailed discussion
of the biology of P. cordata will be given at a later date.

SECT, PLANTAGO

3, PLANTAGO RUGELII Dcne, in DC Prodr. XIII :695. 1852.

BugeFs Plantain Map 10, Figs. 2B, 6B, 7B and 8.

Plant an annual or perennial herb. ROOTS: Fibrous from a
short stem. LEAVES : Rhombic-ovate or elliptic, glabrous, 2-10 cm
wide; blades 6-20 cm long; major veins 5 (7, 9), parallel to mar¬
gin, joining at base of blade; margin entire but bearing 5-9 small
pectinate teeth. INFLORESCENCE: Total length including pe¬
duncle 12-50 cm, glabrous, 1-8 per plant, ca. 10 flowers per cm.
FLOWERS: (Fig, 6B) Outline oblanceolate ; sepals 2.2.-2.5 mm
long; bract % the length of sepals, linear-lanceolate, tip attenuate,
pigmented area total width, glabrous; petals subulate —

deltoid, chartaceous, reflexed at anthesis; style length equal to
stamen length, stigmatic hairs not present at corolla level ; anthers
conspicuously horned, purple or yellow. CAPSULE: (Fig. 7B)
Oblong, 6.5-8 mm long, dehiscent at middle; lower valve of cap¬
sule below tips of sepals after dehiscence. SEEDS: (4)-7-(9) per
capsule, dark brown, testa slightly sculptured, 1. 8-2.3 mm long,
planar area only vaguely defined but occupying ca. i% of axial
surface, hilum and micropyle separated. (N = 12) , PHENOLOGY :

294 Wisconsin Academy of Science, Arts and Letters [VoL 56

Flowering early June to November with peak in July (Fig. 8).
Fruit maturing 2-3 weeks after anthesis.

Plantago rugelii, as its ‘Jook-alike/' P. major, is an extremely
variable and plastic species. We are fortunate that students of this
species were not as anxious to name every morphological deviant
as has been the case with P. major. Only two varieties have been
named ; alterniflora and aspera both by Farwell. I have seen types
of these and hold them to be ecological variants; alterniflora is a
depauperate dry-exposed form, aspera is a juvenile form.

Native in eastern North America and west to Texas and North
Dakota, P. rugelii has been introduced in most other states but
is common only in its native area. It is not so frequent in the
northern parts of its range (in Wisconsin north of the Tension
Zone (Map 10), cf. discussion under P. major) as farther south.
This may represent a general northward migration of the species
associated with a warming climate and surely, at least for the
weedy forms, with the disturbances of man. Hansen (1961) re¬
ports it as established in certain areas of northern Europe. The
more robust forms are associated with rich bottom-lands frequently
in shady areas. Shull (1914) showed that the seeds will germinate
after being submerged in water for up to 54 months. The robust
form, when growing along streams, could be confused with
P. cordata (see above) as both can produce very large leaves often
having purple petiole bases (a character allegedly, but not, diag¬
nostic of P. rugelii). Hamilton & Buckholtz (1956) concluded from
their experiments that P. rugelii grew better in association with
Agropyron repens than in areas where the grass was removed. In
light of the putative relationship of the Plantaginaceae and the
Scrophulariaceae, this might suggest a haustorial relationship
which they did not investigate. I repeated the experiment but ob¬
tained rather poor growth in the plantain. No haustorial connec¬
tions were found. The problems of identification and confusion
with P. major involve mainly the more or less depauperate — weedy
and/or juvenile forms of both species. The species undergo parallel
changes when subjected to unfavorable conditions, especially those
of dry, open hot areas such as roadbeds, paths, cracks in masonry,
mud-flats, talus slopes and gravel pits. Here the leaves are small
(2-3 cm wide) and often thickened; the inflorescences short (5-10
cm) and sparse ; and frequently the capsules aborted or contorted
It is almost impossible to distinguish sterile material of this type
although some clues may be obtained from the leaf margins. From
general field observations, it appears to me that of the two species,
P. major is the more common weed in Wisconsin, whereas in
Michigan, P. rugelii is the more common.

1967-68] Tessene — Reports on Flora in Wisconsin

295

Figure 5. Reproductive structures of Wisconsin Plantagos: A) Plantago lance-
olata; B) P. media; C) P. virginica cleistogamous flower; C') P. virginica
chasmogamous flower; D) P. aristata; E) P. patagonica; F) P. indica; a,b,c,
d,e,f) Seeds of respective species.

296 Wisconsin Academy of Science, Arts and Letters [VoL 56

Figure 6. Reproductive structures of Wisconsin Plantagos: A) Plantago cor-
data; B) P. rugelii; C) P. major; a,b,c) Seeds of respective species.

Details on the reproductive biology of P. rugelii are not known.
Strausbaugh (1950) reports on the occurrence of branched spikes
in the species. It is a long-day plant but may remain blooming into
early November {Fig. 8). The seeds are dormant when first shed
but germinate after a cold treatment or several months in storage
(Steinbauer & Grigsby 1957).

If. Plantago major, l. Sp. PI. ed. 1:113. 1753.

Broad-leaved Plantain, White-man’s Foot

Map 9, Figs. 2A, 6C, 7F and 8.

Plant an annual or perennial herb. ROOTS : Fibrous from a
compact stem. LEAVES : Ovate to cordate, glabrate or hirsutulous,
2-13 cm wide; blade 4-15 cm long; major veins 3, 5, (7), parallel
to margin, joining at base of blade; margin entire or slightly
undulate. INFLORESCENCE: Total length including peduncle
6-25 cm, glabrate; 1-30 spikes per plant; ca. 16 flowers per cm.
FLOWERS: {Fig. 6C) Outline ovate; sepals 1. 5-2.0 mm long;
bract 1% the length of sepals, lanceolate, tip obtuse, pigmented area
% of total width, glabrous; corolla lobes deltoid, chartaceous, re¬
flexed at anthesis; style length equal to stamen length or slightly
shorter; stigmatic hairs absent at corolla level; anthers conspicu-

1967“68] Tessene — Reports on Flora in Wisconsin 297

ously hornedj frequently purple. CAPSULE : Rhombic-ovate, 4-5
mm long, dehiscent near the middle ; lower valve of capsule usually
exceeding tips of sepals. SEEDS: 12 (8-16) per capsule, black-
brown, testa deeply sculptured, 0.6-1 mm long, planar area in¬
conspicuous, hilum and micropyle adjacent. (N = 6), PHENOL¬
OGY : Flowering mid- June to November (Fig. 8). Fruit maturing
2-3 weeks after anthesis.

Plantago major is well known for its variability and plasticity.
Two subspecies, over a dozen varieties and innumerable forms have
been credited to eastern North America. Fernald (1950) mentions
several forms of PilgePs subspecies eumajor while Gleason (1952)
classifies the plants as members of subspecies pleiosperma Pilger.
These subspecies, based on number of seeds and general capsule
shape, are questionable, as Dowling (1935) showed in intergrada¬
tion of seed number in a population of P. major in Great Britain.
Likewise, my own observations of greenhouse cultures have re¬
vealed variation in seed numbers from 4 to as high as 20 per capsule
in P. major from Costa Rica, Hawaii, Michigan, Ontario and Wis¬
consin. Young plants, especially those forced to bloom when they
have produced only a few elliptic leaves, will usually have a lower
number of seeds per capsule. Capsule shape will also vary with age
but not as greatly. The var, scopulorum Fries & Broberg has been
collected on beaches in Wisconsin but such plants, in the green¬
house, will quickly revert back to 'Typical” form. The results of
various transplant experiments are given by Marsden- Jones & Tur-
rill (1930, 1933, 1935, 1937, 1938). The species is badly in need of
revision and in view of the dubious nature of the intraspecific
taxa, I choose, here, to treat the species as P. major sensu lat.

Plantago major is found throughout the world in areas disturbed
by man, but is not abundant in the far north. With the exception
of oceanic islands, it is difficult to state where the species is native
and where it has been introduced by man. Whether or not it is
native to North America is still a matter of speculation. The plant
was supposedly called "White-man's Foot” by the Indians of east¬
ern North America. If this is true, it would indicate an unfamili¬
arity with the plant by the Indians before European colonization.
An interesting problem would be to determine whether there exist
physiological-genetic differences between the Eurasian and Ameri¬
can populations. Likewise a chronological distribution map (like
that of Rorippa sylvestris, see Stuckey (1966) might help resolve
the question. It was first collected in Wisconsin in 1887 in
Wauwatosa.

Plantago major occupies a wide range of habitats from railroad
yards and sandy hills to moist, rich soils of old fields and lawns
but is rarely found in hard-packed soils, Curtis (1959) in his

298 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Species List (p. 641) includes P. major as a member of the
“Southern wet mixed forest’' yet lists it in Table XXXI-1, “Preva¬
lent species of weed communities of southern moist nitrogen-rich
soils,” as reaching its highest presence value (87%) there. He also
lists (p. 641) P. rugelii as being a member of the “Dry mesic
prairie.” From herbarium data, the literature and my own observa¬
tions, I can not help but think these two species are switched in
the Species List. Plantago major does very poorly in shaded areas
and rarely occurs in areas that are wet during the growing season.
The opposite is true for P. rugelii (see above). The latter is ap¬
parently more abundant south of the “Tension Zone” (75 locali¬
ties vs. 23 for P. major) while P. major is more abundant north
of it (51 localities vs. 29 for P. rugelii). Hartley (1966) gives it
as “infrequent” in the “Driftless Area” where P. rugelii is “com¬
mon.” This unequal distribution of the two species could, of course,
simply be a matter of chance but, on the other hand, if P. major
were truly introduced, it would not be particularly incongruous
for P. major's weedy forms to have a slight advantage in the north,
where P. rugelii is not native and both are competing in the weed
communities. In the south, on the contrary, where P. rugelii is in
its native area, the presence of autochthonous bottomland popu¬
lations in addition to the weedy forms makes it more numerous.

Plantago major starts blooming 3-4 weeks later than P.
lanceolata (Fig. 8) and continues later into the fall. Various ab¬
normal forms of the spike, some of which are genetically controlled,
have been described (Hammarlund 1921) including apical rosettes,
leafy bracts, fasciation and branching. There is no evidence of
gynomonoecism or other reduction of the normally hermaphroditic
flowers. Sagar & Harper (1964) give seed-to-seed period as 6 weeks
under cultivation. The seeds are dormant at the time of maturity
but will germinate after several months or a chilling (Steinbauer
& Grigsby 1957). The seeds may remain viable for periods of ca.
50-60 years (Chippendale & Milton 1934) and have given 10%
germination after 40 years (Crocker 1938). Unlike P. lanceolata,
seeds of P. major never germinate in the fall but do so sporadically
the following season (Marsden-Jones & Turrill 1938).

Sect, psyllium (Juess.) Harms in Engler & Prantl

5. Plantago indica l. Syst. Nat. ed. 10. 11:896. 1759.

Psyllium, Indian Plantain Map 6, Figs. ID, 5F, 61 and 8.

Psyllium indicum Du Mount de Cours

Psyllium annuuJm Thuill.

Psyllium ramosum Gilib.

Plantago ramosa (Gilib.) Ascherson

1967-68] Tessene — Reports on Flora in Wisconsin 299

Figure 7. Capsules of Wisconsin Plantagos: A) Plantago cordata; B) P.
rugelii; C) P. media; T>) P. lanceolata; E) P. virginica; F) P. major; G) P,
aristata; H) P. patagonica ; I) P. indica.

Psyllium arenarium Mirbel

Plantago arenaria Poir.

Plantago psyllium sensu Gleason & Cronquist (1963), non L.

Plant an herbaceous or slightly woody annual. ROOTS: Major
tap root with usually poorly developed fibrous side roots: STEM:
Total length 10-50 cm, freely branching; internodes 1-4 cm long.
LEAVES : Linear, hirsute, 1-7 cm long, 0.2-0. 5 cm wide, opposite ;
major veins (1) , 3, parallel the entire length of leaf; margin entire,
ciliate. INFLORESCENCE: Total length including peduncle 2-6
cm, hirsute, 1-30 or more per plant ; spike often appearing capitate,
10 flowers per cm. FLOWERS: (Fig. 5F) Outline obovate; sepals
2-3 mm long; bract 5/6 the length of sepals, ovate, lower bracts
may be aristate, 2-2.5 times as long as sepals, bracts increasingly
shorter from base of inflorescence to tip, tip of bract rounded, pig¬
mented area 1/3 of total width and bearing conspicuously rounded
keel hispid; corolla lobes elongate-deltoid, crenulate, central vein
present, reflexed at anthesis; style length equal to stamen length,
stigmatic hairs present at corolla level; anthers conspicuously
horned, yellow. CAPSULE : Rounded-ellipsoid, 10-13 mm long, de¬
hiscent at the middle. SEEDS: 2- (3) per capsule red-brown, testa
smooth, 2.8-4 mm long; planar area occupying % of face, sunken,
causing sides to appear revolute in x-section ; micropyle and hilum
separated. (2N = 12 Rahn, 1957). PHENOLOGY: Flowering
July-September.

300 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Plantago indica is readily identified by its caulescent habit. The
only species we may confuse with it is A psyllium L., a close rela¬
tive, which is rarely introduced in eastern North America. I have
seen no specimens nor heard any reports of the latter occuring in
Wisconsin. It differs from P. indica in having leaves narrower,
hirsutulous or glabrate, more linear bracts, and a generally
“spindly” habit.

Plantago indica is native in Europe being most com.mon in the
countries bordering the Mediterranean Sea. It is probably fre¬
quently reintroduced in North America, as the seeds are still a
popular remedy for constipation (Claus 1961). Additional uses of
Plantago in medicine can be found in Shyreu (1935) . In Wisconsin,
the plants are found on sandy open areas frequently along beaches
(Map 6). Goessel collected specimens at North Point in Sheboygan
from 1919 to 1932. The plants die in the fall and over-winter as
seeds. In the greenhouse, I have been able to keep individual plants
alive for only 10 months. There is apparently no primary dor¬
mancy as the seeds germinate readily. Flowering is initiated only
under long days. The species will probably become more frequent
in the future.

Sect, leucopsylliun Dene.

6. Plantago ARiSTATA Michx. FI. Boraeli-Amer. 1:95. 1803.

Bracted Plantain, Buckhorn Map 4, Figs. IB, 5D, 7G and 8.

Plantago patagonica Jacq. v. aristata (Michx.) A. Gray

P. gnaphalioides Pursh. v. aristata (Michx.) Hook.

P. purshii Roem. et Schult. v. anstata (Michx.) Jones

P. nuttallii Rapin

P. aristata nuttallii (Rapin) Morris

P. squarrosa Nutt.

P. squamosa Nutt, ex Dene.

P. frankii Steud.

P. filiformis Dene.

Plant an annual herb. ROOTS: Major root tap; secondary roots
usually abundant. STEM: In order specimens, especially those
growing in moist areas, stem up to 1.5-6 cm long with internodes
2-5 mm long. LEAVES: Rosulate, usually erect, linear, 3-17 cm
long, 3-6 mm wide, hirsute to pilose; major veins 3, parallel the
entire length of leaf; margin entire, ciliate. INFLORESCENCE:
Total length including peduncle 3-27 cm; peduncle hirsute, 1-10
per plant, 2-4 flowers per cm. FLOWERS: (Fig. 5D) Outline
elliptic ; vertically 4-ranked on peduncel but spirally arranged ;
sepals 2-2.5 mm long; bract reflexed, tip aristate, pigmented area
1/3 of total area but discernable only at base of bract, hyaline mar-

1967-68] Tessene — Reports on Flora in Wisconsin

301

gin present only on lower 2-3 mm of bract, bract 1.2-5 times as
long as sepals, thinly pilose; corolla zygomorphic, limb cordate,
involute, tip of posterior limb rounded, 4-8 darkly pigmented
patches on each limb near throat of corolla tube ; style length equal
to stamen length or slightly longer; stigmatic hairs present at
corolla level; anthers ovate in outline, not versatile, spinose, in¬
cluded in upper corolla lobe, light yellow. CAPSULE : Acutely
elliptic, narrow constriction circumscribing upper valve immedi¬
ately above line of dehiscence; dehiscent below the middle; old
corolla always remains on capsule at maturity. SEEDS : 2 per cap¬
sule, reddish-tan, testa finely sculptured, 2,2-3 mm long, with a
shallow medial constriction crossing back; planar area occupying
almost the entire face, face with two concentric dark elliptic bands ;
micropyle and hilum separate. (2N = 20 Rahn, 1957). PHENOL¬
OGY : Flowering mainly in June but sometimes extending into
October (Fig. 8). Fruit set 2-3 weeks after anthesis.

Plantago aristata is extremely variable in size but is usually
readily identified by the presence of the aristate bract. Goodwin's
study showed a general correlation between ecologically marginal
habitat and small size. Although the mature plants ranged from
single-spiked individuals ca 3 cm high to seven-spiked specimens
up to 20 cm high, the various morphological units — bracts, capsules
etc. — remained proportional (Goodwin, 1949). In the vegetative
rosette stage, it is almost impossible to distinguish from P.
patagonica with which it often forms mixed populations. Plantago
spinulosa Dene, is a morphological intermediate between the two
species and may represent hybrids. Individuals of P. aristata may
have relatively short pilose bracts while some individuals of P.
patagonica (e.g. Thomson Sept. U, 19.37 Millston, Wis. WIS) have
reflexed spinose bracts. This probably represents a small amount of
introgression. The parental species are phenologically separated
(see below; Fig. 8) but do overlap. Alva Day (personal communi¬
cation) grew plants from seeds of P. pnrshii (P» patagonica? )
having long bracts but, in cultivation, the bract length reverted to
the normal, short condition. She explains the long bract as perhaps
a result of high moisture content of the soil.

Plantago aristata is native to the Great Plains of North America
but, with the disturbances of man has spread throughout the United
States (including Hawaii) and adjacent Canada. In Wisconsin, it
is generally restricted to the prairies (Map 4) where it occupies the
open, well-drained sites. Curtis (1959) lists it as in indicator of
dry prairies. It is sometimes found as a weed along railroad beds
and sandy beaches.

Day-length requirements are not specifically known but my ob¬
servations on greenhouse materials indicate that a short day is

302 Wisconsin Academy of Science, Arts and Letters [Vol. 56

required to initiate flowering. Once flowering has started, day-
length is no longer important as the plants will continue to produce
inflorescences under long days until the apex is eventually used
up in the production of a terminal spike. If the plants are subjected
to sudden changes in day-length, an apical rosette of vegetative
leaves often develops on the spike. This may later produce lateral
inflorescences. The plants will continue blooming for several months
during which a stem 3-5 cm long may develop. Plantago aristata
begins blooming 1-3 weeks later than P, patagonica (Fig. 8) at
which time most stigmas of the latter are no longer receptive. Al¬
though I can not detect a fragrance in the flowers of either species,
the relatively large and showy corolla and small, frequently included
stamens, do suggest entomophily. I have seen small flies and bees
visit the flowers but do not know whether they are signiflcant pollen
vectors. The plants are self-compatible. The seeds have little or
no primary dormancy (Steinbauer & Grigsby, 1957). Even so, they
rarely germinate the same year they are produced as the capsules
do not dehisce readily and remain intact on an erect spike until late
fall when the plant dies and finally falls over.

7. Plantago patagonica Jacq. Icon. P. Rar. II, Coll. Supl. 35. 1796.

Map 3, Figs. 1C, 5E, 7H and 8.

Plantago patagonica v. gnaphalioides (Nutt.) A. Gray

P. purshii sensu Fernald (1950) non Roem. et Schult.

Plant an annual herb. ROOTS: Major root tap; secondary roots
poorly developed. LEAVES : Rosulate, erect or slightly spreading,
linear, 3-12 cm long, 2-6 mm wide, often woolly-pilose ; major veins
3, only one visible on adaxial surface, parallel the entire length of
leaf; margin entire, ciliate. INFLORESCENCE: Total length in¬
cluding peduncle 4-20 cm; peduncles pilose, 1-15 per plant (usually
only 4-6 per plant in Wisconsin) ; 2-6 flowers per cm. FLOWERS :
(Fig. 5E) Outline elliptic; conspicuously vertically 4-ranked; sepals i
1. 0-2.4 mm long, pilose; bract usually equal to or shorter than
sepals but sometimes projecting slightly beyond the calyx, always |
appressed to calyx, lanceolate, tip rounded, pigmented area bor- ■
dered by very narrow chartaceous margin; corolla zygomorphic, |
limbs cordate at base, involute, posterior limb erect, tip acute, one !
pigmented patch on each limb near throat of corolla; style length i
equal to stamen length or slightly longer; stigmatic hairs present !
at corolla level; anthers elliptic in outline, not versatile, included j
in upper corolla limb, light yellow. CAPSULE : Rounded — elliptic, f
narrow constriction circumscribing upper valve immediately above ;
line of dehiscence; dehiscent below the middle; old corolla always i]
remains on capsule at maturity. SEEDS : 2 per capsule, reddish-tan,

per cent of flowering specimens ceni of flowering specimens

303

1967“68] Tessene — Reports on Flora in Wisconsin

Figure 8. Phenology of selected Wisconsin Plantago species.

testa finely sculptured, 1.6-2 mm long shall ow medial constriction
crossing back ; planar area occupying almost entire face, face with
one dark elliptic band; micropyle and hilum separate. (2N = 20

304 Wisconsin Academy of Science, Arts and Letters [Vol. 56

Rahn, 1957) PHENOLOGY: Flowering early May and June occa¬
sionally into September (Fig. 8).

Plantago patagonica, like P. aristata (see above), is quite vari¬
able in general size. The problems of classification and taxonomy
arising between P. patagonica and P. purshii are readily apparent
in the literature. I agree with Gleason and Cronquist (1963, p. 644)
that our plants of P. patagonica are . . apparently identical with
those from S. Amer. . . The interesting and perplexing problem
is the explanation of this amphitropical distribution. Although the
entire section of the genus is in need of a complete revision, I believe
P. purshii of our western states is a very different entity (and
probably a good species) from the P. patagonica of the central
plains. I have never seen the western plant from any localities east
of the Mississippi River. For additional comments (not in complete
harmony with my views) see Poe (1928) and Morris (1900).

Restricted to the well drained soils of the Great Plains, P. pata¬
gonica occurs in the dry prairies of southern and western Wiscon¬
sin (Map 3) . Unlike P. aristata, it does not form large weedy popu¬
lations in areas disturbed by man. It is only rarely found along
dry beaches and railroad beds. It also differs from P. aristata in
that it can not tolerate very moist or wet soils. Under these condi¬
tions, it rapidly turns yellow and dies.

Plantago patagonica is a well marked short-day plant: I have
kept it in the vegetative stage for a period of four months under
14 hour days but could readily induce blooming in the same plants
under 8-hour days. The floral morphology and reproductive be¬
havior are similar to P. aristata (see above). The plants begin
blooming the second or third week of May (1-3 weeks before P.
aristata). Individual plants may continue blooming for about a
month. The seeds are dormant when first shed but will germinate
after 3-4 months storage or alternating cold-warm treatments.

Sect, arnoglossum Dene.

8, Plantago lanceolata l. Sp. PI. ed. 1:113. 1753.

Ribgrass, Ripplegrass, English Plantain, Buckhorn, Narrow¬
leaved Plantain. Map 7, Figs. IE, 5A, 7D and 8.

Arnoglossum lanceolatum S. F, Gray
Plantago lanceaefolia Salisb.

P. flexmsa Guad. ex Rapin

P. capensis Bojer

P. longistipes Royale ex Barneoud

Plant an annual or perennial herb, ROOTS: Juvenile, tap; old
specimens, fibrous from short stem or caudex. STEM : Caudex 4-6

1967-68] Tessene — Reports on Flora in Wisconsin

305

cm long in old specimens, frequently branching in robust individ¬
uals. LEAVES : Lanceolate 6-32 cm long, 0.5-4 cm wide, glabrate
or hirsutulous ; major veins 5 (3, 7), parallel the entire length of
the leaf; margin usually denticulate with 5-11 small teeth. IN¬
FLORESCENCE: Total length including peduncle 10-40 cm,
hirsutulous, peduncle ribbed, 1-10 per plant; floral group globose
or elongate, 12-15 flowers per cm. FLOWERS: (Fig. 5A) Outline
elongate-rhombic with construction between bract and sepals;
sepals arising from a short pedicle i.e, not directly in the axil of
bract, sepals 2-2.5 mm long, the two sepals next to bract fused into
one doubly tipped, doubly ribbed unit; bract 4/5 the length of
sepals, acuminate, ovate, the pigmented area reduced to a narrow
rib, glabrous ; petals elongate-cordate, slightly pigmented near tip,
reflexed at anthesis; style length ^2 of stamen length; stigmatic
hairs present at corolla level; anthers cordate at base, yellow;
the plants may be gynomonecious or gynodioecious. CAPSULE :
Elongate, dehiscent at the base. SEEDS: (1) 2 per capsule, dark
brown, testa lightly sculptured, often hirsutulous, 1.6-1. 9 mm long;
planar area 1/3 of face, slightly sunken; micropyle and hilum
adjacent. (2N = 12 Rahn, 1957). PHENOLOGY: Flowering mid-
May through October. Fruit maturing 3-4 weeks after anthesis.

Plantago lanceolata, like the other weedy plantains, is an exceed¬
ingly variable species throughout its range. Pilger (1937) divided
it into two varieties and seven subvarieties. Other authors (e.g.
Druce, 1928) recognize up to 12 varieties. Of these, only two,
sphaerostachya Mert, & Koch (having a globose inflorescence and
often pilose leaves) and angustifolia Poir. (having short, narrow
leaves) have been cited as occurring in our area (Gleason 1952).
Griffiths (1922) showed that the pubescence of sphaerostachya is
dependent on environmental conditions : The dry-open habitat pro¬
ducing the small pubescent plant while a more mesic or wet habitat
yielded the typical form. He also produced the sphaerostachya habit
by placing the typical form in the dry-open habitat. Leaf shape and
length are determined by the height of surrounding vegetation
( Jenkin, 1925) . My held observations and greenhouse cultures show
that pubescence, leaf size (and to some extent shape) and general
inflorescence size are apparently strongly influenced by the environ¬
ment. I am therefore recognizing for Wisconsin only the typical,
although polymorphic, variety.

A native of Eurasia but widely established throughout the world
except in subartic and low-lying tropical areas, Plantago lanceolata
is probably the best known member of the genus. The species has
had a long association with man : Pollen deposits have often been
used as an indicator of prehistoric man’s migrations (Iversen,
1941, Godwin, 1944, 1956). There is no record of when it was first

806 Wisconsin Academy of Science, Arts and Letters [Vol. 56

introduced into North America but because it is so abundant in
Europe and especially in the British Isles (Sagar & Harper, 1964)
it was probably brought to the early settlements.

Plantago lanceolata was first collected in Wisconsin in Marquette
Co. in 1861 and represents the first Plantago known to be collected
in the state. The distribution in Wisconsin (Map 7) is misleading
in that the plant is much more abundant than the number of locali¬
ties would indicate : One could probably find it in all counties. The
problem here is probably like that of other weedy species which,
unfortunately, are not collected because they are so ubiquitous. It is
a common weed of door-yards, roadsides, paths and open fields.
The plants will live under a broad range of ecological conditions
but seem to prefer hard-packed soils (Harper et at., 1965), alka¬
line conditions (Zeiner, 1946) and minimal competition for sun¬
light (Sagar & Harper, 1964).

Plantago lanceolata is a long-day plant (Snyder, 1948, Biinning
& Kodon, 1954) blooming in our area from late May to early Octo¬
ber with a peak around the second week in July (Fig. 8). The
flowers may be pistillate, staminate or hermaphroditic and show a
complete morphological spectrum of sexuality (Stout, 1919, Blar-
inghem, 1923, Hope-Simpson, 1939, Baker, 1963). The pollen is an
important cause of hayfever in early summer (Wodehouse, 1945).
Although normally wind-pollinated, P. lanceolata has been reported
to be frequented by insects (Clifford, 1962). The seeds may germi¬
nate in the late summer or fall of the year they are produced but
rarely produce flowers until the following spring. Buried seeds
may remain viable for up to 40 years (Beal, 1905, Darlington,
1922, Kjaer, 1948) which may help to explain the rapid appear¬
ance of large populations in recently disturbed areas. If favorable
conditions prevail, the seeds will germinate readily (Steinbauer &
Grigsby, 1957).

Sect, lamprosantha Dene.

9. Plantago media l. Sp. PI. ed. 1:113. 1753.

Hoary Plantain, Dwarf Plantain Map 5, Figs. lA, 5B, 7C and 8.

Plantago concinna Salisb.

P. incana Stokes

P. bertolonii Godr.

Arnoglossum incanum S.F. Gray.

Plant an annual or perennial herb. ROOTS : Major root tap with
abundant secondary fibrous rootlets. STEM : Compact, but in older
specimens a caudex 1-2 cm long, rarely branching. LEAVES:
Spatulate-obovate, 4-10 cm long, 2.4-4 cm wide, appressed, hirsute;
major veins 5, 7 (9), parallel the entire length of the leaf; margin

1967-68] Tessene— Reports on Flora in Wisconsin 307

entire, slightly revolute. INFLORESCENCE: Total length includ¬
ing peduncle 12-35 cm; peduncles smooth, 1-5 per plant, 12-17
flowers per cm. FLOWERS: (Fig. 5B) Outline obovate, fragrant;
sepals 2.0-2.4 mm long, all sepals distinct and separate; bract %
as long as sepals, broadly lanceolate, tip acute, pigmented area %
of total width, glabrous; corolla limbs lanceolate, chartaceous,
slightly revolute, posterior lobe erect, all others reflexed; style
length % of stamen length; stigmatic haris present at corolla
level; anthers broadly fusiform in outline, white or pink. CAP¬
SULE: Rhombic, dehiscent at the middle. SEEDS: 3 (2-4) per
capsule, dark brown, testa smooth, 1. 8-2.1 mm long; planar area
poorly defined, represented by slight depression; micropyle and
hilum separate. (2N = 12, 24 Rahn, 1957). PHENOLOGY : Flow¬
ering mid- June to September.

Plantago media, a widely distributed weed from northern Europe
to central Russia, is rare in our range. Pilger (1937) lists five vari¬
eties and four forms for the Old World material but does not cite
any North American specimens. In Wisconsin, the species is known
from only two localities (Map 5) and is represented by the typical
variety. The plant is apparently re-introduced frequently as there
is no record of its becoming well established anywhere in North
America, It is not even listed in Steyermark (1940), Small (1933)
or Rydberg (1932). Fernald (1950) refers to it as '^occasionaF^ as
do Gleason (1952) and Gleason & Cronquist (1963). Hartley
(1966) in his Flora of the “Driftless Area'' calls it "‘rare’" and ‘'A
weed on a golf course at La Crosse.” — where he collected it once in
1956.

The species may be confused with P. lanceolata superficially but
the higher seed number and the absence of fused sepals readily
distinguish it. Rademacker (1940) gives the habitat as confined to
base-rich areas of old, relatively undisturbed areas where cultiva¬
tion is at a minimum. Steele (1955) notes it as “an exacting cal-
cicole,” i.e. requiring both high pH and calcium level. The plant is
apparently very drought resistant (Sagar & Harper, 1964).

Plantago media is a long-day plant. The seeds are dormant at the
time of capsule dehiscence but germinate after a cold treatment
(Sagar & Harper, 1964).

Sect, novorbis Dene.

10. Plantago virginica l. Sp. PL ed. 1:113. 1753.

Pale-seed Plantain, Sand Plantain, Dwarf Plantain.

Map 8, Figs. IF, 5C&C', 7E and 8.

Plantago caroliniana Walt.

P. ludoviciana Raf.

308 Wisconsin Academy of Science, Arts and Letters [VoL 56

F. accendens Raf.

P, connivens Moench.

P, purpurascens Nutt, ex Rapin

P. missouriensis Steud.

Plant a short-lived annual herb. ROOTS : Weakly developed tap
system attached to a short compact stem. LEAVES: Spatulate,
2-8 cm long, 0.6-1. 5 cm wide, hirsute or, growing under moist con¬
ditions, hirsutulous; major veins 3, parallel entire length of leaf;
margin enitre or pectinate with 3-5 small teeth. INFLORES¬
CENCE: Total length including peduncle 3-15 cm, hirsute, 1-5
per plant, 2-4 flowers per cm. FLOWERS: (Fig. 5C & C') Outline
obovate; sepals 1.4-2. 7 mm long; bract as long as or slightly shorter
than sepals, bract ovate, tip rounded, pigmented area % of total
width, hispid ; cleistogamous and chasmogamous flowers sometimes
produced on one inflorescence, plants usually apomictic and bear¬
ing only cleistogamous flowers; cleistogamous flowers having
aborted anthers and styles, corolla lobes always erect, bract and
sepals more pointed than those of chasmogamous flowers; chas¬
mogamous flowers having well developed anthers and styles, style
shorter than stamens, stigmatic hairs in two rows down narrow
edges of flattened style; style glabrous at corolla level; anthers
inconspicuously horned, yellow. CAPSULE: (Fig. 7E) Rhombic-
ovate, dehiscent at the middle ; old corolla always present on mature
fruit. SEEDS : 3 per capsule, light brown, testa smooth, 1. 6-1.9 mm
long; planar area occuping % of face; micropyle and hilum adja¬
cent. (N=10 Chandler, 1954). PHENOLOGY: Flowering early
May to July.

Plantago virginica is a short-lived (2 months maximum) spring
ephemeral of pioneer habitats. Fernald’s variety viridescens, based
on shorter bracts and sepals than the typical variety and on bright
green, more or less glabrous leaves, can easily be produced in the
lab by growing the typical variety under moist conditions. The pres¬
ence or absence of denticulations on the leaf margin is apparently
under genetic control. The size of individuals varies from 3 to 20
cm in height (and proportional diameter of the rosette) and is
highly correlated with population density and available moisture;
the smaller specimens being from dense populations in drier areas.

Occurring natively from the East Coast west to Kansas and Ari¬
zona, P. virginica is becoming established as a weed in our western
states. The species is probably more abundant in Wisconsin than
the number of localities (Map 8) indicates. Because of its short
growing season and usually very small size, it is easily overlooked.
It ‘‘blooms’’ mainly in early or middle May (Fig. 8) when the dry
prairies, sand hills, rocky slopes, old gravel pits, borrow-pits and
similar well-drained areas are still moist from spring thaws.

1967-68] Tessene — Reports on Flora in Wisconsin

309

Plantago virginica is a day-n,eutral plant according to my experi¬
ments. I have induced flowering under both short and long day con¬
ditions, It is apparently a facultative apomict : Cleistogamous flow¬
ers with .abortive stamens and very short styles are the typical
form but often occur mixed with morphologically sexual, chasmoga-
mous flowers; either on the same inflorescence or separate inflo¬
rescence on the same plant. The mechanism controlling this phe-

Piantago cordata

Littorello americana

PIcHitogo patagonica

Plantago aristata

Fi

Fr.

M

1

3

j

A

1

i

t

0

H

FI.

Fr.

M

J

1

j

A

S

0

K

F|

Fr.

H

5

3

31

9

3

1#

Vi

A

5

S

0.

u

FI.

Fr.

M

J

18

J

14

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Maps 1-4. Triangles represent county record only. Map 1) Shaded area un¬
derlaid with limestone; Maps 3 & 4) Shaded area savannas and prairie (Curtis,
1959).

310 Wisconsin Academy of Science, Arts and Letters [Vol. 56

1967-68] Tessene — Reports on Flora in Wisconsin

311

nomenon is not known and individuals growing under identical
conditions may display the entire morphological spectrum. Whether
or not the seeds produced by the morphologically sexal flowers are
produced sexually or apomictically has yet to be determined. The
seeds are dormant when mature, but will germinate after a cold
treatment or several months in storage. I have observed a second
generation late the same year in a population west of Ann Arbor,
Michigan, but this is apparently not typical.

Anderson (1959) gives P. virginica as an important source of
the Ring-spot of pepper.

Bibliography

Anderson, C. W. 1959. A study of field sources and spread of five viruses of
pepper in central Florida. Phytopathology 49:97-101.

Baker, H. G. 1963. Evolutionary mechanisms in pollination biology. Science
139:877-883.

Beal, W. J. 1905. The vitality of seeds. Proc. Soc. Promotion Agric. Sci. 26:
89-93.

Blaringhem, L. 1923. Etudes sur de polymorphisme florale IV. Sexualite et
metamorphose des epis de Plantago lanceolata. Bull. Soc. Bot. Fr. 70:
717-725.

Bunning, E. and M. Kodor. 1954. Day-length, foliar growth and flower forma¬
tion. Planta 44:9-17.

Chandler, C. 1954. Improvement of Plantago for mucilage production and
growth in the U. S. Contrib. Boyce Thompson Inst. 17 : 495-565.
Chippendale, H. G. and W. Milton. 1934. On the viable seeds beneath pasture.
J. Ecol. 44:383-390.

Chute, W. N. 1942. Plantago cordata in Indiana. Am. Bot. 48:95.

Claus, E. 1961. Pharmacognosy. Lea & Febiger, Philadelphia.

Clifford, H. T. 1962. Insect pollination of Plantago lanceolata. Nature 196:
196.

Crocker, W. 1938. Life span of seeds. Bot. Rev. 4:235-274.

Curtis, J. T. 1959. The Vegetation of Wisconsin. The University of Wisconsin
Press, Madison.

Darlington, H. T. 1922. Dr. W, J. Beal’s seed-viability experiment. Amer. J.
Bot. 9:266-269.

Dowling, R. E. 1935. The ovary in the genus Plantago. I. The British species.

J. Linn. Soc. (Bot.) 50:329-336.

Druce, G. C. 1928. British plant list. Published by the author.

Fassett, N. C. 1934. Notes from the herbarium of the University of Wisconsin
— XI. Extensions of ranges of aquatic plants. Rhodora 36:349-352.
Fernald, M. L. 1918. The North American Littorella. Rhodora 20:61-62.

- . 1950. Gary’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 New York
Botanical Garden, New York.

- and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern

United States and Adjacent Canada. D. Van Nostrand Co., Princeton,
N. J.

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312 Wisconsin Academy of Science, Arts and Letters [Vol. 56

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Griffiths, B. M. 1922. Growth experiments on Spergula and Plantago. J. Bot.
Bond. 60:228-230.

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Hammarlund, C. 1921. Tiber de vererbung abnorm ahren bei Plantago major.
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Hansen, A. 1961. Plantaginaceernes og Lentibulariaceerner udbredelse i
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WISCONSIN ACADEMY OF SCIENCES, ARTS & LETTERS

Madison, Wisconsin

OFFICERS 1967-68

President

John W. Thomson
Department of Botany
University of Wisconsin —
Madison

Vice-President (Sciences)
Rezneat M. Darnell
Department of Biology
Marquette University,
Milwaukee

Vice-President (Arts)

Mary Ellen Pagel

1111 North Astor Street

Milwaukee

Vice-President (Letters)

Miller Upton
Beloit College
Beloit

President-Elect

Adolph A. Suppan
School of Fine Arts
University of Wisconsin —
Milwaukee

Secretary

Euniee R. Bonow
Department of Pharmaey
University of Wisconsin —
Milwaukee

Treasurer

Jack R. Arndt
University Extension
University of Wisconsin —
Madison

Librarian

Jack A. Clarke

Department of Library Science
University of Wisconsin —
Madison

APPOINTED OFFICIALS

Editor — T ransactions
Walter F. Peterson
Department of History
Lawrence University, Appleton

Editor — Wisconsin Academy Review
Ruth L. Hine

Wisconsin Conservation Division
Madison

Chairman — Junior Academy of Science
Jack R. Arndt
University Extension
University of Wisconsin — Madison

ACADEMY COUNCIL

The Academy Council includes the above named officers and the follow¬

ing past presidents of the

A. W. Schorger
H. A. Schuette
L. E. Noland
Otto L. Kowalke
Katherine G. Nelson
Ralph W. Buckstaff

Academy.

Joseph G. Baier
Stephen F. Darling
Robert J. Dicke
Henry Meyer
Merrit Y. Hughes

Carl Welty
J. Martin Klotsche
Aaron J. Ihde
Walter E. Scott
Harry Hayden Clark
David J. Behling

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