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Vol. LIX— 1971
Cover Design by Arthur Thrall, Lawrence University
TRANSACTIONS OF THE
WISCONSIN ACADEMY
OF SCIENCES, ARTS
AND LETTERS
LIX — 1971
TRANSACTIONS OF THE
WISCONSIN ACADEMY
Established 1870
Volume LIX
OPPORTUNITIES FOR THE FUTURE OF THE WISCONSIN
ACADEMY OF SCIENCES, ARTS AND LETTERS 1
William B. Sarles
FREDERICK JACKSON TURNER:
NON-WESTERN HISTORIAN 7
Ray Allen Billington
THE RHETORICAL HERITAGE OF
FREDERICK JACKSON TURNER 23
Goodwin F. Berquist, Jr.
THE GOSPEL OF POVERTY: THE MESSAGE OF
CONSERVATIVE PROTESTANTISM TO THE POOR
AT THE TURN OF THE CENTURY 33
Walter F. Peterson
THE BROWN RAT IN EARLY WISCONSIN 45
A. W. Schorger
THE INFLUENCE OF DRUMLIN TOPOGRAPHY ON FIELD
PATTERNS IN DODGE COUNTY, WISCONSIN 55
Charles W. Collins
SEDIMENTOLOGICAL AND CHEMICAL PARAMETERS OF THE
LAKE SUPERIOR NERITIC ZONE, SOUTH SHORE, WISCONSIN 67
J. W. Horton, R. C. Brown, D. W. Davidson, A. B. Dickas,
W. Lunking, and R. K. Roubal
VEGETATIONAL PATTERNS AND ORDINATION IN
CEDARBURG BOG, WISCONSIN 79
Thomas Foster Grittinger
THE LITTORAL MACROPHYTE VEGETATION OF LAKE WINGRA 107
Stanley A. Nichols and Scott Mori
VARIABILITY IN WISCONSIN IN TR1ENTALIS BOREALIS RAF 121
Roger C. Anderson
THE INSECT PARASITES OF THE INTRODUCED PINE SAWFLY,
DIPRION SIMILIS (HARTIG) (HYMENOPTERA: DIPRIONIDAE),
IN WISCONSIN, WITH KEYS TO THE ADULTS AND
MATURE LARVAL REMAINS 127
James W. Mertins and Harry C. Coppel
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% 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 should be typed on a separate sheet
entitled “Footnotes” and appended to the section entitled “References Cited.”
Contributors should avoid unnecessary documentation wherever possible. Other
matters of style should be in harmony with current practice in the subject
matter area.
Galley proofs and manuscript copy will be forwarded to the author for
proofreading prior to publication; both should be returned to the Editor within
two weeks . Papers received on or before July 1 5 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:
Doctor Walter F. Peterson
Editor, Transactions of the Wisconsin Academy
University of Dubuque
Dubuque, Iowa 52001
i9th President of the
WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS
OPPORTUNITIES FOR THE FUTURE OF THE
WISCONSIN ACADEMY OF SCIENCES, ARTS AND LETTERS
William B. Sarles , President 1969-70
May 7, 1970
The “Theme" of the Academy's Centennial year is : “Preserving
the Past— Planning the Future." Our meetings in Madison and
Milwaukee, and our publications during this, our 101st year, will
emphasize and amplify the theme, and will illustrate service to
the present.
My excursion into the past will be brief,. but hopefully significant
because it is designed to serve as at least part of the foundation
for that to be said about opportunities for the future.
Members and guests of the Academy will receive a reprinted
copy of Bulletin No, 1, dated April, 1870, of the Wisconsin Academy
of Sciences, Arts and Letters. This valuable publication tells how
the Academy was organized, and by whom; presents its Constitu¬
tion, By-Laws, and its March 16, 1870 Charter from the Wisconsin
Legislature; and finally, states its Plan of Operations.
The Plan of Operations starts as follows : “Having thus a legal
existence, and being provided by the State with secure and con¬
venient apartments in the Capitol for its office, library and collec¬
tions, the Academy is ready to commence the work for which it was
established." This description of early housing and facilities, makes
us envious but at the same time serves as a reminder that we should
strive more diligently and effectively to recapture that which was
provided 106 years ago.
After describing work to be done and studies to be made in the
sciences,, arts and letters, the Plan, written by the Academy's Presi¬
dent, J. W. Hoyt, presents a highly significant point : “The measure
of accomplishment, in other words the efficiency and degree of use¬
fulness of the Academy, will, of course, be determined by the com¬
petency and zeal of its members, the wisdom, energy, and devotion
of its officers, and the cordiality and liberality with which their
plans and efforts for the public good are seconded and sustained
by the people and the State."
President Hoyt was right ; the members and officers of the Acad¬
emy have served it well. But at some time during the century we
failed to accomplish our objectives with sufficient success to justify
strong, continuing financial support from “the people and the
1
2 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
State.” Perhaps we can regain the needed recognition and support
from the State ; we may if we can prove our worthiness.
President Hoyt continues in his “Plan of Operations” to make
recommendations that have come to be of special significance today.
“No institution of this or any other kind can be efficiently main¬
tained without the means to employ and fairly compensate one or
more competent and efficient officers, so that their whole time and
energies may be consecrated to its work. . . . For the liberal support
of these there must be a permanent fund. . . . Wisconsin may not
yet hope to vie with some of the older States in the number and
munificence of . . . private benefactions . . . but she may justly
boast of men, not a few, who, by favor of the rare opportunities
she has given them, have acquired so large a measure of wealth
that the donation of an amount sufficient to place this Academy at
once upon a sure foundation . . . would advance the public interest
by a signal act of noble generosity.” This was a hopeful plea that
has turned out to be a prophetic statement. Our gathering this
evening is an inadequate but sincere attempt to recognize the gen¬
erosity of Professor Harry Steenbock who, “by a signal act of noble
generosity” bequeathed a large share of the residue of his estate
to the Wisconsin Academy of Sciences, Arts and Letters. We are
grateful to Professor Steenbock to a degree that makes impossible
an adequate expression of our gratitude. At the same time, we wish
to honor him because he was a great and accomplished gentleman
who contributed so much to the advancement of scientific knowledge
as a research worker and teacher, and who appreciated and sup¬
ported the arts and letters. Professor Steenbock loved Wisconsin
and made it his life. All of Wisconsin — its people and its institu¬
tions— benefited by his life.
The Academy now has opportunities for the future that were
envisaged 100 years ago by its President and his fellow charter
members.
It is not my desire to present a detailed, step-by-step plan for the
future of the Academy, but rather to emphasize some of the princi¬
pal opportunities that we, as a truly interdisciplinary organization,
can develop. Another way to do this is to speak of opportunities as
challenges to the Academy, and this is what I propose to do.
The first opportunity, or challenge, involves communication of
facts, concepts, and ideas.
There is an ever increasing number of scientists, technologists,
scholars, and artists at work in the world. Their discoveries and
proposals must be published or by some other means made known
to all who have the intelligence to understand. Specialization of the
discoverers and the innovators in any field of endeavor adds to the
complexity of communication and the difficulty of comprehension.
1971] S arles— Opportunities for the Future of the Academy 3
Use of different languages and alphabets contributes to the magni¬
tude of the problems of communication faced by even highly
trained, competent scientists and scholars. Consider, for example,
the difficulties encountered by an accomplished microbial geneticist
who must try to translate intelligently a significant article pub¬
lished in Japanese by one of the growing number of microbiologists
working in his special field in Japan! Microbial geneticists whose
native language is English have enough trouble trying to under¬
stand the writings of their English-speaking colleagues who delight
in the introduction of new terminology. Communication becomes
even more difficult when ideas, interpretations, explanations, con¬
cepts, or descriptions must be elucidated and understood.
The question might be asked : Why is communication a challenge
to the Academy? The Academy is concerned with support and de¬
velopment of sciences, arts and letters. The Academy, by means of
its meetings and publications, is involved in the dissemination of
information ; in communication of knowledge to those competent to
understand. The Academy is committed to stimulation of learning
and to the awakening of interests. The Academy is concerned with
bridging the gaps that exist between specialists and those who need
to know significant facts, concepts, interpretations, and ideas.
What can the Academy do toward improvement of its efforts to
achieve its objectives? The diversity of interests and of competence
represented by its membership is valuable. It makes it possible for
members and guests to gather in meetings and to publish volumes
devoted to interdisciplinary communication of facts, concepts, and
ideas. The specialists have their own journals and their own meet¬
ings that can be devoted entirely to their specialties. The Academy,
because of its diversity, provides an opportunity for development
of awareness of the accomplishments of others and the possibilities
for improvement of communication.
In his book, “Scene of Change,” published by Charles Scribner’s
Sons in 1970, Dr. Warren Weaver speaks of diversity and of unity
of knowledge and concepts when he says : . science should have
no quarrel with the humane arts or with contemplative fields of
thought, nor they with science. They are all, each using its charac¬
teristic methods, seeking to perceive order and unity in diversity.
They are all based on faith, they are all creations of imaginative
minds; they are all alive, growing, changing; they all are limited
by what our linguistic apparatus and our cultural concepts per¬
mit. . . .” Dr. Weaver goes on to quote the late physicist, J. Robert
Oppenheimer : “The artist and scientist both live always at the edge
of mystery, surrounded by it. Both struggle to make partial order
in total chaos. They can, in their work and in their lives, help them¬
selves, help one another, and help all men.”
4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
There is still something missing from that which I’ve said and
quoted about the opportunity — or the challenge — of communication
and of diversity, and the missing part concerns communication
with young men and young women. This is a special problem and a
very special opportunity for the Academy. Dr. Weaver said that
the sciences and arts are limited by what our linguistic apparatus
and our cultural concepts permit. In his book, he describes the
problems of trying to communicate facts, concepts, and ideas to the
Hopi Indians, who do not share our grammar, our ideas, or our
ways of dealing with experience. He points out that the Hopi In¬
dians, even if they could understand English, could not comprehend
our desires to study the sciences,, arts and letters because “they
have a metaphysics of their own, and deal with experience and
reality in ways that are quite unlike ours, but which nevertheless
work entirely satisfactorily for them.”
No doubt I've gone a bit far in relating the challenges of com¬
munication with the young to those encountered in attempts to
communicate with Hopi Indians. Our children and our young
men and women have been exposed to English, mathematics,
sciences, the humanities, and the arts. They have had the oppor¬
tunity to grasp the basics of the facts, concepts, and ideas that
we consider to be important. The principal opportunity— and the
challenge — is to arouse their interests and enthusiasms for learn¬
ing. Isn’t there some way in which we can reach them more effec¬
tively, and help them to generate the curiosity, the thirst for
knowledge, the practices of thinking, and the skills of doing that
can be theirs?
Our Junior Academy of Science, now celebrating its 25th year
of accomplishment and service should, I believe, be expanded to
become the Junior Academy of Sciences, Arts and Letters. The
time to appreciate diversity and unity of thought and endeavor
might be earlier in a young person’s life than we’ve thought. A
broadened Junior Academy could, hopefully, provide the same
opportunities now made available by the Academy only to adults.
Another opportunity for the Academy’s development exists, I
believe, in working with the young men and women who are
undergraduate students in colleges and universities. We have at
present a Junior Academy for high school students; we make no
provisions for undergraduates who are beginning the serious
business of finding themselves and deciding what they really
want to do. These students, perhaps even more than those in high
school, need the stimulants and the services that the Academy can
provide.
My proposals of opportunities— and challenges — for the future
of the Academy may be too general; too much concerned with
1971] S cirles— Opportunities for the Future of the Academy 5
strategy rather than tactics. But I have attempted to generate
some thinking along lines that may lead to the improvement and
development of the Academy; to make it achieve more effectively
the objectives set forth in its Charter in March of 1870. Professor
Steenbock’s bequest has given us a chance to make progress. The
Academy can become a more significant, dynamic force in the
life of the State.
Frederick Jackson Turner: Non-Western Historian
Ray Allen Billing ton
Ladies and Gentlemen: I am indeed honored to participate in
such a momentous occasion as this. For a fruitful century the Wis¬
consin Academy of Sciences, Arts and Letters has stimulated the
intellectual currents that elevated the state and its universities to an
enviable spot in the hierarchy of the nation’s harbingers of civiliza¬
tion. It has done so by refusing to succumb to the forces of spe¬
cialization that increasingly departmentalize all knowledge today.
As I have read over the fat volumes of Transactions of a generation
ago, I have been struck by two things : the incomprehensible forti¬
tude of those who endured hour after hour of learned discourse, and
their good fortune in doing so. As they listened, half dozing per¬
haps, to papers on subjects far from their principal interest, they
may well have been startled into rapt attention by a hypothesis, or
a technique, or an idea that applied to their own research interest
and that illuminated a hitherto dark corner. The Academy is to be
commended to keeping alive the spirit of interdisciplinary inves¬
tigation in a day when the knowledge explosion threatens to com¬
partmentalize all learning.
It is for this reason that I want to talk with you tonight about
Frederick Jackson Turner, a long-time member of the Academy,
who more than any other historian of his day sought to popularize
its ideals and utilize the approaches that it advocated. This, admit¬
tedly, is not the image of Turner in the popular mind. He is remem¬
bered, in Wisconsin, as the state’s most distinguished contribution
to the historical profession who trained a legion of students at
Madison between 1889 and 1910 when he reluctantly left for Har¬
vard because he believed that his resignation would awaken the
regents to the dangers of their attacks on pure research. He is
remembered nationally as the intellectual father of two theories.
One, advanced in 1893 in his famed paper on “The Significance of
the Frontier in American History,” held that certain distinctive
features of the American character could be traced to the three-
centuries-long process that settled the continent ; as opportunity in
the form of free land and untapped natural resources altered
behavioral patterns, men became more optimistic, inventive, mate-
* Doctor Billington delivered this address at the banquet of the fall gathering- of the
Academy held in Milwaukee, October 3, 1970.
7
8 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
rialistic, nationalistic, and democratic than their fellows with no
frontier experience.1 The other, the sectional hypothesis, occupied
Turner for most of his life. As the frontier moved westward, he
held, pioneers encountered and overran vast physiographic regions
where differing natural conditions created differing modes of life.
These ‘sections’ corresponded to European nations, and the history
of the United States could be understood only by comprehending
the manner in which national policy was decided by sectional con¬
flict, interplay, and compromise.2 Turner devoted most of his life
to a vain attempt to prove that hypothesis.
Turner’s stress on these two theories, linked with the immense
popularity of the frontier thesis during the early years of the cen¬
tury, have created a popular impression of Turner as a propa¬
gandist who preached that one giant causal force shaped the
nation’s past. Critics rose by the score during the depression-
oriented 1930s and 1940s to brand him as a monocausationist who
ignored more vital causal forces by his stress on the frontier, who
glorified nationalism at a time when internationalism was necessary
to prevent world chaos, who blinded the people to the problems bred
of industrialization and urbanization by his stress on the rural past.3
My purpose tonight is to challenge that distorted image and
reveal Frederick Jackson Turner in his true light: as a historian
whose views were so modern, whose techniques were so in advance
of his times, whose conception of history was so broad, that he
would feel as much at home in a meeting of the American Historical
Association of today as he did in 1910. Above all I hope to demon¬
strate that he was a pioneer in the interdisciplinary approach to
research, and that those who today explore those borderland areas
in the social sciences could learn from his example.
Before plunging into that discussion, let me answer one necessary
question : if Turner’s views were so modern, why has his reputation
1 This general essay was first presented to the American Historical Association on
July 12, 1893, at a meeting held in Chicago in connection with the World’s Columbian
Exposition. It was repeated before the State Historical Society of Wisconsin on Decem¬
ber 14, 1893. Its first printing was in the Proceedings of the Forty-First Annual Meet¬
ing of the State Historical Society of Wisconsin (Madison, 1894), 79-112; it was
reprinted later that year in American Historical Association, Annual Report for the
Year 1893 (Washington, 1894), 199—227. The essay has appeared dozens of times since
then, in whole or in part, in every form from historical anthologies to expensively
designed special books. It is perhaps most readily available in Frederick Jackson
Turner, The Frontier in American History (New York, 1920), 1—38.
2 Turner published a number of essays embodying his theories on sectionalism. Most
important among these are those entitled “Sections and Nation,” Yale Review, XII
(October, 1922), 1—21, and “The Significance of the Section in American History,”
Wisconsin Magazine of History , VIII (March, 1925), 255—280. Both were reprinted
in Turner, The Significance of Sections in American History (New York, 1932).
3 For a summary of the views of Turner’s critics see Ray A. Billington, The Amer¬
ican Frontier (Washington, 1965. 2nd edn. ), and America’s Frontier Heritage (New
York, 1966).
1971] Billing ton— Turner : Non-Western Historian
9
been so tarnished since his death in 1932? The answer, I suspect,
can be found in the popularity of his frontier thesis. Repeatedly
through his lifetime he was called upon to speak or write about the
frontier; this was the subject expected of Turner when he was
demanded for a commencement address, enlisted for the annual Phi
Beta Kappa banquet, or seduced into writing a lucrative article for
the Atlantic Monthly . Yet he had, after the turn of the century,
abandoned research on the frontier to begin his sectional studies or
to venture briefly into the field of diplomatic history. The lectures
or essays composed under those circumstances simply expanded
ideas that he had already voiced, substituting extravagant rhetoric
for proof. When unshackled by factual information, as he was on
such occasions, he was inclined to succumb to oversimplification and
overstatement, substituting the bludgeon for the rapier, and writing
with the unrestrained prose of the poet rather than the exactly
defined words of the historian. These are the essays, collected in
two published volumes, that aroused the ire of his critics and gave
them evidence of his traitorism to the standards of his profession.4
Let me give you an example of the way in which he dug his own
grave by these practices. Turner was to teach on the west coast
during the summer of 1914, and had agreed to give the commence¬
ment address at the University of Washington on “The West and
American Ideals” — a topic broad enough to cover anything he
might possibly want to say when he began composing his remarks.
Unhappily, he had little time for that composition. He intended to
prepare a polished oration before leaving Cambridge, but Turner
was a natural procrastinator, and the round of oral graduate
examinations, the ocean of end-of-term blue books, the need of a
hurried trip to Washington for a committee meeting, and the chaos
involved in moving their residence from one house to another
allowed the spring to pass with nothing done.5 He left Boston on
June 4, intending to make some progress during the week allotted
to Madison, but that was ill-advised, for his daughter Dorothy was
marrying John Main that week, and between wedding plans and
old friends no work was done. Turner arrived in Seattle on June 15
or 16 with his commencement address still to be prepared — an
4 Turner’s two published volumes of essays, referred to in the footnotes above, were
his Frontier in American History (1920), and The Significance of Sections in American
History (1932), the latter published after his death.
5 Turner to Edmond S. Meany, May 26, 1914, in Roy Lokken, ed., “Frederick Jack-
son Turner’s Letters to Edmond S. Meany,’’ Pacific Northwest Quarterly , XLIV (Jan¬
uary, 1953), 35; Turner to Mrs. William Hooper, May 26, 1914, Frederick Jackson
Turner Papers, Henry E. Huntington Library and Art Gallery, TU-H, Box 2. Here¬
after referred to as : “HEH.”
10 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
address scheduled for delivery on June 17. 6 To compound his prob¬
lems the packet of notes that he had hastily assembled had disap¬
peared, so that he must depend on his memory even for the
quotations.7
The result was predictible— an address that marvelously paraded
most of the cliches of frontier history. The pioneer “knew not
where he was going but he was on his way, cheerful, optimistic,
busy and buoyant/’ He was “an opportunist rather than a dealer
in general ideas/’ but possessed “a courageous determination to
break new paths, indifference to the dogma that because an institu¬
tion or a condition exists, it must remain.” American democracy,
Turner told his audience, “was born of no theorist’s dream ; it was
not carried in the Sarah Constant to Virginia nor in the Mayflotver
to Plymouth. It came out of the American forest, and it gained new
strength each time it touched a new frontier.”8 In one hastily pre¬
pared essay Turner had provided fuel that was to keep his critics
happy for two decades.
In his own behalf, it must be said that he was less than proud
of the fantasies that he invented on that occasion. “I am still mov¬
ing by reflex action after my poor commencement address,” he
wrote a friend two days later,9 and added that he felt as a man
might who relaxes in the electric chair after the first shock.10 His
one solace was that the crying babies in the audience of 2,500
drowned out most of his words. “Always,” he advised a friend,
“take along a supply of babies when you preach.”11 These bantering
words hid a genuine humiliation. At first he refused to allow the
lecture to be published — “it was written to be spoken,” he
explained12— but when the editor of the Washington Historical
Quarterly persisted, he succumbed. Only when he saw it in print
did he recognize the many errors of fact and theory of which he
was guilty. Preserved among his papers is a printed copy filled
with corrections, adorned with frequent marginal notes, and bear-
6 Turner, “Memorandum,” in Harvard Commission on Western History Correspond¬
ence, Harvard University Archives, Widener Library, Harvard University, Box 5,
Folder: Turner, F. J. Hereafter referred to as “HC on WH Corr.” The story of the
Commission, which was financed by Mrs. Hooper to purchase books and manuscripts
in western history for the Harvard University Libraries, is told in the introduction to
Ray A. Billington, ed., “Dear Lady The Letters of Frederick Jackson Turner and
Alice Fortes Perkins Hooper (San Marino, Calif., 1970).
7 Turner to Max Farrand, October 26, 1914. HEH TU Box 22.
8 Turner, “The West and American Ideals,” in The Frontier in American History,
290-310.
9 Turner to Roger Pierce, June 19, 1914. HC on WH Corr., Box 9, Folder: In Re
A. B. Hulbert.
10 Turner to Charles Homer Haskins, June 18, 1914. Charles Homer Haskins Papers,
Firestone Library, Princeton University.
11 Ibid. A warm, sunny day attracted a larger crowd than Turner had anticipated —
or wanted. Seattle Sun , June 17, 1914.
12 Turner to Edmond S. Meany, July 22, 1914, in Lokken, ed., “Frederick Jackson
Turner’s Letters to Edmond S. Meany,” loc. cit 35-36.
1971] Billing ton- — Turner : Non-Western Historian
11
ing beside the more extravagant statements the underlined words :
“Too strong/’13
I do not want to suggest that all of Turner’s later papers on the
frontier were as inadequately prepared as this ; I do maintain that
the true Turner can be found not in warmed-over versions of a
theory that no longer concerned him but in the research papers and
letters to fellow historians where he expressed his real views on the
nature and meaning of historical research. If we focus on this evi¬
dence we reveal a scholar who stood head and shoulders above his
own generation in concepts, purpose, and methodology. Above all,
Turner emerges as a pioneer in the interdisciplinary approach for
which the Academy stands, and which is proving so valuable among
the social sciences today.
This viewpoint was a product of his training, his intellectual
environment, and his whole concept of history. His training under
Professor William Francis Allen at the University of Wisconsin
pointed him in the right direction. Allen, a mediaevalist, was inter¬
ested to an unusual degree in the impact of social conflicts, eco¬
nomic forces, and cultural factors in shaping the civilization of the
Middle Ages; he taught his young disciple that society was an
evolving organism responding to a variety of pressures that must
be investigated for complete understanding.14 As Turner turned
his own attention to studies of the American frontier he recognized
the wisdom of his master. Alterations in the traits of intruding
ethnic groups under the impact of frontier opportunity could be
appraised only after the physical environment was properly under¬
stood. This meant mastering geography and geology ; in 1898, when
a young professor at Madison, Turner enrolled in a course on the
physiography of the United States given by his friend and neigh¬
bor, Charles H. Van Hise.15 This proved immensely valuable in his
studies; what was more logical than to assume that the study of
economics, or government, or sociology, would be equally revealing.
13 Turner listed some of his errors in writing- to his good friend Max Farrand (in¬
cluding the statement that George Washington was born in South Carolina), then
added: “There are other reasons why it lacks the perfection which is the dream of the
wise and the good.” Turner to Farrand, October 26, 1914. HEH TU Box 22. A printed
copy of the address, with Turner’s marginal comments, is in HEH TU File Drawer
15B, Folder: Commencement Address. University of Washington.
14 The only adequate biography of Professor Allen is Owen P. Stearns, “William
Francis Allen : Wisconsin’s First Historian,” Unpublished Masters Thesis, University
of Wisconsin, 1955. Mr. Stearns has kindly allowed me to use the well-researched
dissertation. Turner frequently commented on his debt to Allen. For a typical example
see Turner to Merle Curti, August 8, 1928. HEH TU Box 29.
15 The University of Wisconsin undergraduate newspaper. The Daily Cardinal,
October 4, 1898, described the lecture course that attracted Turner. The careful lecture
notes that he took during the course, all dated October and November, 1898, are scat¬
tered through his papers at the Huntington Library. They may be found in HEH TU
File Drawer 12C, Folder: Van Hise Course; File Drawer 14D, Folder: Lecture.
Physical Geography ; and File Drawer 15A, Folder: Notes on Van Hise’s Lectures.
12 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
This early training inclined Turner toward the interdisciplinary
approach to history, but I like to believe that his resolutions were
strengthened by the intellectual environment of Wisconsin provided
by the Academy of Sciences, Arts, and Letters. He was elected a
member in 1887, when still a graduate student, probably under the
sponsorship of Professor Allen who was president that year.16 From
that day until his death in 1932 he served actively, presenting two
of his important research papers for the first time at meetings in
1895 and 1896, commenting frequently at other sessions, persuading
his students to present their findings in a series of papers at the
annual gatherings, and acting as vice-president for the letters sec¬
tion in 1896.17 We will never know the ideas planted in his mind
or the flashes of inspiration that occurred as he listened to endless
papers on the whole range of scholarly inquiry. Did Humphrey J.
Desmond, a political scientist, turn Turner’s interests to sectionalism
when in 1888 he reported on “The Sectional Feature of American
Politics?”18 Did T. C. Chamberlin and Charles H. Van Hise stir the
young historian’s curiosity as they gave their numerous reports on
geology, and turn him toward the geographic investigation that
underlay his frontier and sectional concepts?19 Did he suddenly
awaken when listening in 1892 to E. A. Birge explain “Weissman’s
[sic] Theory of Heredity,” to the knowledge that nearly all biol¬
ogists believed in the inheritance of acquired characteristics, and
thus see how traits stemming from the frontier environment could
be passed on to future generations?20 Those are unanswerable ques¬
tions, but there seems no question that Turner’s immersion in the
interdisciplinary atmosphere of the Wisconsin Academy helped
broaden his interests to the everlasting benefit of historical
scholarship.
Finally, and probably most important, Turner was nudged toward
an interdisciplinary approach by his own concept of the nature of
16 Transactions of the Wisconsin Academy of Sciences , Arts and Letters, VII, 1883 -
1887 (Madison, 1889), 267.
17 Turner read his paper on "State Making- in the West” before the Academy in
1885, and another on ‘‘The Projected French Expedition of George Rogers Clark
Against Louisiana in 1793” in 1896. Ibid., XI, 1896-189 7 (Madison, 1898), 549. The
first of his student’s papers before the society was delivered in 1891 when Turner
himself read an essay by Kate A. Everest on ‘‘Early Lutheran Immigration to Wis¬
consin,” Ibid., VIII, 1888-1891 (Madison, 1892), 415. Others were presented at irregular
intervals during the remainder of Turner’s stay at Wisconsin. Notice of his activities
on committees and as vice-president are in Ibid., VIII, 1888—1891 (Madison, 1892),
413, and XI, 1896-1897 (Madison, 1898), 256. The Daily Cardinal, January 6, 1897,
also noted his elevation to the vice-presidency of the letters section.
18 Ibid., VIII, 1888-1891 (Madison 1892), 1-10. A printed copy of this essay, heavily
underlined and annotated by Turner, is in HEH TU File Drawer 14B, Folder: Sec¬
tional Feature in American Politics.
19 Charles H. Van Hise’s evening address in 1893 on ‘‘The Evolution of the North
American Continent” would certainly attract Turner’s interest. T. C. Chamberlin was
a constant reader of geological papers before the Academy at this time. Ibid., X,
1894-1895 (Madison, 1895), 582.
20 Ibid., IX, 1893 (Madison, 1893), viii.
1971] Billington— Turner : Non-Western Historian
13
history. He was, above all else, no monocausationist, as his critics
have branded him. More than most men of his generation— or today
—he recognized the complexity of human behavior and the variety
of forces motivating every action. “In truth,” he told an audience
early in his career, “there is no single key to American history. In
history, as in science, we are learning that a complex result is the
outcome of the interplay of many forces. Simple explanations fail
to meet the case.”21 This remained Turner’s credo ; through his life¬
time his quest was not for the force underlying an event, but for all
forces. Early in his academic career he read the germinal essay by
T. C. Chamberlin on “The Method of the Multiple Working Hypoth¬
esis,” first published in 1897, and from that day on applied to his¬
torical studies the techniques Chamberlin used in geology.22 Tur¬
ner’s method was to postulate every possible explanation for any
happening, then test each successively with all available evidence.
Only in this way, he believed, could the historian escape what he
called “the warping influence of partiality for a simple theory.”23
Before showing you how Turner applied that belief, let me
digress to ask one obvious question. Should a historian wedded to
the concept of the multiple hypothesis and aware that the complex¬
ity of human behavior precluded simple explanation, have devoted
his life to proving the influence of the frontier and section on past
behavior? Did he consider them more important than other forces:
the class struggle, slavery, constitutional interpretation, or many
more? Of course he did not. “I do not,” he explained to Carl Becker
in 1925, “think of myself as primarily either a western historian,
or a human geographer. I have stressed those two factors, because
it seemed to me that they had been neglected, but fundamentally
I have been interested in the inter-relations of economics, politics,
sociology, culture in general, with the geographic factors, in
21 Turner, “The Development of American Society,” The [ Illinois ] Alumni Quarterly,
II (July, 1908), 120-121. He used almost these words in a Phi Beta Kappa lecture
at the University of Nebraska in 1907. The manuscript of this lecture is in HEH,
TU Box 55. Edward E. Dale, “Memories of Frederick Jackson Turner,” Mississippi
Valley Historical Review , XXX (December, 1943), 355-356 remembered that he em¬
ployed virtually the same phrase when lecturing- to a class some years later.
22 Thomas C. Chamberlin, “The Method of Multiple Working Hypotheses,” Journal
of Geology, V (November-December, 1897), 837-848. Chamberlin first read the paper
at a meeting of the Society of Western Naturalists in 1889, and Turner may well
have learned of it at that time. Many years later he wrote one of his students : "I, as
you perhaps recall, valued Chamberlin’s paper on the Multiple Hypothesis, which I
have aimed to apply to history as he did to geology.” Turner to Merle Curti, August
8, 1928. HEH TU Box 39. Wilbur It. Jacobs, “Turner’s Methodology: Multiple Working
Hypotheses or Ruling Theory?,” Journal of American History, LIV (March, 1968),
853-863, argues that Turner never correctly applied the methodology advocated by
Chamberlin, but that he treated his frontier and sectional hypotheses as ruling
theories that were true instead of postulates to be tested.
23 Turner used this phrase in his presidential address before the American Historical
Association in 1910. Turner, “Social Forces in American History,” in The Frontier in
American History, 333.
14 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
explaining- the United States of to-day.”24 This was common sense.
Each man must specialize, and the mere fact of specialization does
not mean that he assigns greater influence to his field of special
interest than any other.
Turner, then, was committed to the use of the multiple hypoth¬
esis, and to the belief that no single key unlocked the secret of the
past. Equally important was his realization that history was more
than the “past politics” current in the 1890s ; it was more too than
the colorful narratives spun from the pens of the Francis Park-
mans and the William H. Prescotts of that day. History, to Turner,
was “a study of the social forces which caused and modified the
political events, institutions, and ideas of the period.”25 Enough
time had been wasted by nineteenth century scholars tracing the
surface manifestations of political behavior. The time had come to
probe below the surface, and analyze the underlying forces-— eco¬
nomic, social, and cultural — that found ultimate expression in
politics. “Behind institutions,” he wrote in his famous 1893 essay,
“behind constitutional forms, and modifications, lie the vital forces
that call these organs into life and shape them to meet changing
conditions.”26 This was his declaration of independence. His inves¬
tigations would be focused on “the changes in the economic and
social life of the people,” as he put it, “that . . . ultimately create
and modify organs of public action.” No mere narrative, no super¬
ficial survey of political behavior would do.27 His area of study was
society as a whole, not its leaders. “The important point,” he told
the well-known sociologist Albion W. Small in 1904, “is to get more
sociology into history and more history into sociology.”28
This view of history doomed Turner to a lifetime of perpetual
learning — and shamefully little writing. He had marked out for
himself an impossible assignment; to understand human behavior
in all its manifestations he must become a master of many dis¬
ciplines, and a slave to none. He must use the tools of the economist,
the sociologist, the demographer, the geographer, and the political
scientist. At the same time he must remain true to his own dis-
24 Turner to Carl Becker, October 3, 1925. Carl Becker Papers, Cornell University
Library. Hereafter referred to as : “Becker Papers. Cornell.”
25 Turner, Review of J. W. Burgess, The Middle Period, 1817—1858 (New York,
1897), in the Educational Review, XIV (November, 1897), 390-391.
26 Turner, “The Significance of the Frontier in American History,” in The Frontier
in American History, 2.
27 In 1921 Turner wrote a publisher who had tried vainly for a generation to per¬
suade him to write a textbook : “It is in narrative history that I am least experienced
or (I fear) competent . . . My strength, or weakness, lies in interpretation, correlation,
elucidation of large tendencies to bring out new points of view ... I am not a good
saga-man.” Turner to Lincoln MacVeagh of Henry Holt & Company, April 5, 1921.
Henry Holt & Company Archives, Firestone Library, Princeton University.
28 Turner to Albion W. Small, November 4, 1904. College of Letters and Science,
Department of History, Turner Correspondence, 1901-1905, Box 5, Folder: S, Uni¬
versity of Wisconsin Archives, University of Wisconsin Library. Hereafter referred
to as: “Li & S, Dept, of Hist. Turner Corr., 1901-5. Wis. Archives.”
1971] Billing ton— Turner: Non-Western Historian
15
cipline and avoid the traps of other social scientists who were all
too ready to lay down rules of human conduct. “The human soul/'
he wrote in 1904, “is too complex, human society too full of vital
energy and incessant change, to enable us to pluck out the heart
of its mystery— to reduce it to the lines of an exact science or to
state human development in terms of an equation/'29 Historians
had made that mistake in the past, and their chronicles were strewn
with the wrecks of “known and acknowledged truths."30 Now they
must utilize their breadth of experience, and with it temper their
findings as they used the tools of other social scientists. To fail to
do so would be to abandon the field to others less well equipped to
ferret out the truth. “History," he wrote in 1914, “had a right to
deal with large mass statistics, tendencies, etc., as well as the event
and the individual psychology. I dislike to yield good territory to
sociologists, political scientists, etc., on which the historian may
raise good crops/'31
More than any other historian of his generation, Turner suc¬
ceeded in preserving that territory for his fellow craftsmen. At
times he was a lone voice crying in the wilderness, but always he
was an evangelist preaching a vital message with the fervor of con¬
viction. As early as 1901 he urged on the students in the Division of
Economics, Political Science and History the belief that “the vari¬
ous branches of our work are related/' and cautioned them against
letting the branches of social science drift apart unduly.32 He drove
home the message even more strongly when addressing a historical
gathering at the St. Louis World's Fair three years later, their most
serious problem was “how to apportion the field of American his¬
tory itself among the social sciences.” This meant calling into coop¬
eration sciences and methods hitherto little used. “Data drawn from
the studies of literature and art, politics, economics, sociology,
psychology, biology, and physiography all must be used,” he told
his listeners. “The method of the statistician as well as that of the
critic of evidence is absolutely essential. There has been too little
cooperation of these sciences, and the result is that great fields have
been neglected."33 Once more in 1910 Turner made this plea the
central theme of his presidential address before the American His¬
torical Association. Just as geologists were using chemistry, mathe-
29 Turner, “The Historical Library in the University/’ in Brown University, John
Curler Brown Library. The Dedication of the Library Building , May the Seventeenth,
A.D. MDCdGC'IIII . With the Addresses by William Vail Kellen , LL.D., and Frederick
Jackson Turner , Ph.D. (Providence, R.X., 1905), 52-53.
30 Turner, “Social Forces in American History,” in The Frontier in American History,
333.
31 Turner to George Lincoln Burr, September 5, 1914. George Lincoln Burr Papers,
Cornell University Library.
32 Notice sent to the members of the School of Economics, Political Science and His¬
tory, October 5, 1901, by Turner and Richard T. Ely. A copy is in L & S, Dept, of Hist.,
Turner Corr, 1901—5. Wis. Archives.
33 Turner, “Problems in American History,” in The Frontier in American History , 20.
16 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
matics, botany and zoology to understand the dynamics of the inor¬
ganic earth, he warned, so must historians master the essential
tools of the social sciences to unravel the skeins of the past.34 This
was a unique plea ; Turner was the first president of this influential
body to urge interdisciplinary techniques.
Moreover, Turner practiced what he preached, and used a variety
of disciplines so successfully that his students and friends some¬
times wondered if he had deserted his own cause. One of his Har¬
vard undergraduates mirrored this confusion when he was heard
to say that what Turner was doing in the classroom was all right,
“but it wasn’t history.”35 When a colleague accused him of being
more of a sociologist than historian, Turner answered: “It is the
subject that I am interested in, and I don’t particularly care what
name I bear.”36 Nor did he, so long as his unquenchable thirst for
knowledge could be satisfied. The quest for the truth about the past
mattered to him; the tools used in unearthing that truth did not.
“I sometimes wonder,” he told Carl Becker late in his life, “if after
all I have not been simply, rather blindly, trying to explain America
to myself instead of writing history! or writing agriculture, or
geography, or diplomacy, or economics, land transportation, etc.,
or literature, or religion.”37 So he had, but he had still been study¬
ing history. For history to Turner was, as he once put it, “a com¬
plex of the social sciences.” It could be made understandable only
if the watertight compartments in which they had been divided
could be broken down, and the One-ness of the subject brought
home to investigators.38
He labored to break down those compartments, just as he tried
to persuade other historians to do so. When, early in his career, he
joined the editorial board of the American Historical Review his
first step was to urge the acceptance of more articles that lay “on
the border-land between history and its neighbors.”39 A lifetime
later when in his twilight years he was asked to advise the Hunting-
ton Library on its book-buying policies, he pointed out that history
was no longer a placer-mining form of narrative collecting, “but
involves the use of wide agencies more like the chemical labora¬
tories, dredging process, geological experts, quartz crushers, etc.
The library extended into economic, social, literary, political, relig¬
ious sources becomes necessary to the modern type of historian of
3i Turner, “Social Forces in American History,” in ibid., 331. For an excellent dis¬
cussion of the originality of Turner’s views in relation to other presidents of the asso¬
ciation see Herman Ausubel, Historians and Their Craft: A Study of Presidential
Addresses of the American Historical Association , 188^—19^5 (New York, 1950), 326-
329.
35 Turner to Merle Curti, August 8, 1928. HEH TU Box 39.
36 Turner to Luther L. Bernard, November 24, 1928. HEH TU Box 40.
37 Turner to Carl Becker, February 13, 1926. Becker Papers, Cornell.
38 Turner to Carl Becker, December 1, 1925. Becker Papers. Cornell.
33 Turner to J. Franklin Jameson, January 26, 1910. HEH TU Box 14.
1971] Billington — - Turner : Non-Western Historian 17
the evolution of civilization/’40 Few men have clung so tenaciously
to a belief as did Turner in the virtues of interdisciplinary studies,
and few have both preached and practiced a cause so devotedly.
His own contribution to the breaking of line-fences between dis¬
ciplines led him principally into the field of geography, for this was
the allied subject that promised the greatest help in understanding
American sectionalism. This was a complex problem; to what
degree did varying physiographic features in each region — South¬
east, Southwest, New England, North Central States, and the like
— alter the behavioral patterns of the intruding stocks? Were the
differences in economic, social, political, and cultural behavior that
distinguished New England from the North Central States, or the
Southeast from the Southwest due more to environmental forces
or to the persistence of ethnic behavioral patterns among the
settlers? This was the basic problem in the study of sectionalism,
and that to which Turner devoted most of his research between
1905 and his death in 1932.
It could be solved only by devising a completely new research
technique, build on methods of both geographers and statisticians.
This involved the drawing of maps that would correlate physi¬
ographic features with political, social, economic, and cultural
behavior. Some that he devised revealed the physiographic basis
of society; they showed on a county-by-county basis the nature of
soils, land values, agricultural yields, value of farm produce, and
extent of improvements. Others depicted patterns of social behav¬
ior, illuminating degrees of illiteracy, levels of education, church
preferences, and various cultural attainments. A third set of maps
illustrated voting patterns, showing which counties were consist¬
ently Whig or Democratic, and which showed slight political alle¬
giance. By comparing these maps, Turner was able to show to his own
satisfaction that there was a direct relationship between geographic
conditions and political or economic behavior. Thus he could demon¬
strate that areas of poor soil, high illiteracy rate, and evangelical
sects in the pre-Civil War era were inclined to vote Democratic;
that western counties changed their attitude on the tariff from free
trade to protectionism with a change in crop emphasis from corn
to wool growing ; and that counties with agricultural surpluses for
export were those principally supporting government-financed
internal improvements.41
40 Turner to Max Farrand, March 8, 1927. HEH TU Box 36.
41 Dozens of Turner’s manuscript maps are preserved in the Turner Papers at the
Huntington Library. A few of the most important were published in his last book,
completed by friends after his death, The United States 1830— 1850 : The Nation and
Its Sections (New York, 1935). Turner was convinced of the value of his maps, and
of their use in various social-study disciplines. “I really think,” he wrote in 1925,
“the maps which exhibit the correlation between physical geography (especially
topography and soils), land values in 1850, illiteracy, party politics, and culture have
a real merit in the line of showing the interdependence of the social studies.” Turner
to Carl Becker, October 3, 1925. Becker Papers. Cornell.
m
18 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Today’s student of statistical cartography or statistics is inclined
to look at the dozens of maps so laboriously produced by Turner
with a mixture of amused tolerance and contempt, for we now know
that they woefully failed to accomplish their purpose. His misfor¬
tune was twofold: he lived before statisticians devised the sophis¬
ticated techniques that would endow his masses of data with sig¬
nificant meaning, and he was so wedded to the use of maps that he
failed to take advantage of more useable means of achieving the
correlations that he sought. Neither he nor his students made any
serious effort to master the devices that might have solved their
problems : mathematical correlations, sampling, time-series trend¬
ing, and the use of punched cards for storing and sorting data.
Instead Turner’s faith was in a device that did not lend itself to
the complex correlations that he required. As he crowded on the
necessary information the result was an undecipheral maze of col¬
ors, symbols, lines, and numbers, so jumbled and crowded that they
meant nothing. Even when kept simple the visual comparison of
two or more maps to identify correlations was almost impossible
unless the correlation was so high that it could not be missed.
Today’s statistical techniques allow the type of analysis that Turner
sought, but not through the maps on which he pinned his faith.42
This latter-day criticism should not blind us to the undisputed
fact that in his own time Turner’s methods were accepted as
irreproachable and his findings hailed as revolutionary by both
geographers and statisticians. His services were sought as speaker
at professional associations of geographers,43 and when he did con¬
sent in 1914 to deliver a paper at a joint meeting of the Association
of American Geographers and the American Geographical Society
on “Geographical Influences in American Political History,” his
talk was hailed as one of the most stimulating ever given and by
far the hit of the program. “Your work,” he was assured by a lead¬
ing geographer, Isaiah Bowman, “is so sympathetic with respect
to geographic factors that it is a pity that we do not see more of
you and hear a paper every year.”44 The high regard in which
Turner was held by practitioners of this discipline was shown when
42 Richard Jensen of Washington University has graciously allowed me to use his
unpublished paper on “The Development of Quantitative Historiography in America.”
This excellent essay, together with the same author’s study of “American Election
Analysis : A Case History of Methodological Innovation and Diffusion,” in Seymour M.
Lipset, ed., Politics and the Social Sciences (New York, 1969), provide the best
criticism of Turner’s mapping techniques, yet assign him his proper place as a pioneer
in the use of statistical methodology.
43 Ralph S. Tarr to Turner, May 25, 1911. HEH TU Box 16. Turner has written
“NO” at the top of this letter. He had, however, participated in two conferences at
meetings of the American Historical Association in 1907 and 1908 on “The Relation
of Geography to History.” Reports of these meetings are in the Annual Report of the
American Historical Association for 190 7 (Washington, 1908), I, 45-48, and the Annual
Report of the American Historical Association for 1908 (Washington, 1909, I, 61.
44 Isaiah Bowman to Turner, April 7, 1914. HEH TU Box 21.
1971] Billing ton— Turner : Non-Western Historian 19
he was elected a member of the Association of American Geograph¬
ers — an honor restricted to one hundred scholars who made
the most significant contributions— and chosen a fellow of the
American Geographical Society.45 His passing in 1932 was mourned
as sincerely by geographers as by historians. “Professor Turner,”
wrote the editor of the Geographical Review, “was a rare combina¬
tion of historical originality with geographical insight. His death
is a loss no less severe to American geography than to the study of
American history.”46 Clearly Turner had broken the line-fence
between two of the social-science disciplines as no one had in
the past.
Despite the criticism of modern sceptics who scorn his statistical
techniques, he was also viewed by statisticians of its own genera¬
tion as the outstanding pioneeer in applying their discipline in the
field of history. When the American Statistical Association cele¬
brated its seventy-fifth birthday in 1913, the one historian invited
to participate was Turner, who was asked to give a paper on “The
Importance and Service of Statistics to History.” “No one,” the
president of the association assured him, “is better qualified to
speak on this subject from the view of the historian than you are.”47
A decade later when a team of scholars began preparation of a book
on the inter-relationship of the social sciences, they turned nat¬
urally to Turner for the chapter on the relationship between history
and statistics.48 Turner declined both of these invitations, for he
was too deeply involved in his sectional studies to be interrupted,
but the point remains : his own generation viewed him as the out¬
standing disciple of the use of statistical methods in historical
research.
That Turner pioneered in blending historical, geographic, and
statistical techniques even his enemies must concede; they insist,
however, that despite his lip service to interdisciplinary studies he
"virtually ignored the other social sciences. Economists in particular
have castigated him for distorting the past by refusing to recog¬
nize the significance of class conflicts and economic growth. If
Turner is to be judged solely on the basis of his published works
this charge seems warranted, for he wrote little and largely of a
rural America in which economic forces played a lesser role than
in the industrialized-urbanized twentieth century. But if we ex¬
amine his unpublished essays, and read his letters of advice to
friends, it becomes clear that he was no less aware of their impor-
45 Isaiah Bowman to Turner, March 1, 1915. HEH TU Box 25. Turner’s certificates
•of membership in the two associations are in HEH TU Box 53.
46 Geographical Review, XXII (July, 1932), 499
47 John Koren to Turner, November 18, 1913. HEH TU Box 20A.
48 William F. Ogburn to Turner, December 15, 1924. HEH TU Box 33. Turner sent
Jiis “regrets” to both of these proposals.
20 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
tance than later economic historians who built their whole inter¬
pretation of the American past on an economic foundation.
Turner’s interest in economic forces and recognition of their
importance in shaping human behavior can be traced to the very
beginning of his career. As early as 1892 he urged that historians
cease their preoccupation with politics and “interpret events from
the economic point of view.”49 Until this was done, history would
be superficial, and scholars would have no real understanding of
the basic subsurface social and economic tides that determined
political action. In adopting this point of view, Turner aligned him¬
self with the little group of social scientists that included Lester
Ward, Richard T. Ely, and John R. Commons who first recognized
that modern society rested on an economic base, and that this must
be understood before political actions could be understood. He saw
too, as did John R. Commons, that they were linked with social and
cultural forces, and that all operated together in human
motivation.50
With this as a basic assumption, Turner would never ignore the
role of economics in shaping human behavior, even though his focus
on agrarian America left him little to say on the subject. When he
concentrated, as he occasionally did, on a modern period, he recog¬
nized that even a Marxian interpretation had some validity. A final
and very important phase of the nation’s growth, he told an audi¬
ence in 1908, had been “the steady stratification of our society by
the development of contesting social classes. It is hard to realize
how recently it has become possible to use the words proletariat
and capitalistic classes in reference to American conditions.”51 In
his seminars, too, he elaborated the idea of a continuous conflict
between “the capitalist and the democratic pioneer” persisting from,
colonial times to the present.52 Even such a latter-day critic as
Charles A. Beard paid tribute to Turner as a pioneer in exploring
the field that he was to make his own. “Almost the only work in.
economic interpretation which has been done in the United States,”
he wrote in 1913, “seems to have been inspired at the University of
Wisconsin by Professor Turner.”53
At one point Turner did break with Beard and his school, and
we know today that Beard was wrong and Turner right. He saw,
49 Turner, “The School of Economics, Political Science and History,” The Aegis, VI
(April 8, 1892), 448.
50 For an intelligent appraisal of Turner’s influence in this group of thinkers see
George D. Blackwood, “Frederick Jackson Turner and John Rogers Commons — Com¬
plementary Thinkers,” Mississippi Valley Historical Review, XLI (December, 1954),.
471-489.
51 Turner, “Development of American Society,” loc. cit ., 132-133.
52 Merle Curti to Turner, August 13, 1928. HEH TU Box 39.
53 Charles A. Beard, An Economic Interpretation of the Constitution of the United-
States (New York, 1913), 5-6.
1971] Billing ton— Turner : Non-Western Historian
21
as less perceptive scholars did not, that idealism played a role in
human motivation, and that those tracing behavioral patterns solely
to material desires were blind to the truth. Over and over again
he protested to his friends, “I am not an economic determinist.”54
This broad view led to a more sophisticated— and more complex-
view of history than that of Beard and his followers, but no mes¬
sage that he preached was more important in leading to a correct
understanding of the past. “I tried,” he told a friend in 1928, “to
keep the relations steadily in mind; but it isn’t an easy job, and
the effort is sometimes conducive to unwritten books!”55
Frederick Jackson Turner, then, more than any historian of his
generation, succeeded in merging history with geography, history
with statistics, history with economics. Nor did his assault on com-
partmentalization end even there, for he was a welcome guest on
programs of the American Sociological Association and a pioneer
in introducing sociological methods into historical research. Frank¬
lin H. Giddings, one of the most eminent practitioners of that sub¬
ject, in 1928 judged Turner to be “a sound sociologist, and a
ground-breaking one of first rate importance.”56 In the judgment
of his peers, Turner stood head and shoulders above his fellow-
historians in urging interdisciplinary studies, and in proving them
essential in unraveling the secrets of the past. Avery Craven, a
student of Turner and himself an eminent historian, summed up
the judgment of his generation in his remarks at the funeral of
his master. “He is claimed by the historians,” Craven said, “and
the sociologists and the geographers and yet he was more than any
of these. He was a student of the whole field of the social sciences,
and more than any other man . . . saw the field as one and was able
to integrate it.”57 No historian could deserve a more appropriate
epitaph than that.
54 Turner to Andrew C. McLaughlin, May 29, 1915, HEH TU Box 24; Turner to
Edgar E. Robinson, December 12, 1924, HEH TU Box 33 ; Turner to Archibald Hen¬
derson, January 29, 1930, HEH TU Box 43. On the margin of a term paper written
by one of his students, Herman K. Murphy, Turner wrote : “History not all economic
■determinism.” HEH TU Students’ Papers (1).
55 Turner to Merle Curti, August 15, 1928. HEH TU Box 39.
56 Franklin H. Giddings to Merle Curti, August 20, 1928. HEH TU Box 39.
57 Avery Craven, “An Appreciation of F.J.T.,” HEH TU Box 57.
THE RHETORICAL HERITAGE
OF FREDERICK JACKSON TURNER
Goodwin F. Berquist, Jr.
Ninety -three years ago a young man named Frederick Jackson
Turner became a member of the Wisconsin Academy of Sciences,
Arts and Letters. Turner was at the time an instructor in rhetoric
and oratory at the University of Wisconsin. He would later be¬
come Vice-President of the Academy for Letters and one of Amer¬
ica's truly great historians.1 2 Turner’s fame began with a speech
he delivered in Chicago in 1893 entitled “The Significance of the
Frontier in American History.” The purpose of this paper is to
identify and explore Turner’s rhetorical heritage prior to that time.
The Rhetorical Environment at Portage
Portage, Wisconsin where Fred Turner grew up a hundred
years ago was a rough and ready frontier community. Indians with
their ponies and dogs came to town frequently to trade furs for
paint and trinkets. Lumberjacks from the pineries of central Wis¬
consin “took over” the town on Saturday nights. The Bierhall of
Carl Haertel was a haven for local Germans as well as the town
club house. Pomeranian immigrants, Irish, Scots, Welsh, Norwe¬
gians and Swiss lived about Portage, along with a few Southerners
and Negroes, some Englishmen and one or two Italians. Turner’s
parents were among the Yankee settlers from upstate New York
who came in the 1850’s.3
In such a melting pot, not everyone could read or write English.
But all could listen and speak when they had a mind to. The spoken
1 Paper read at the Centennial Meeting of the Wisconsin Academy of Sciences, Arts
and Letters, Milwaukee, Wisconsin, October 3, 1970. The author is former editor of the
Academy Transactions and Professor of Speech, Ohio State University. Research for
this paper was undertaken at the Henry E. Huntington Library and Art Gallery, San
Marino, California under a grant from the Department of Speech, Ohio State University.
2 Authority for the above statements comes from the following sources : Transactions
of the Wisconsin Academy of Sciences, Arts and Letters, 7(1883-87), 267 ; biographical
data dictated by Turner to his secretary, TU Box 57, Frederick Jackson Turner Papers,
Henry E. Huntington Library ; Merle Curti, “Frederick Jackson Turner” in Wisconsin
Witness to Frederick Jackson Turner, O. Lawrence Burnette, Jr. comp. (Madison: State
Historical Society of Wisconsin, 1961), p. 175. Turner served as an Academy Vice-
President from 1896 to 1899 and remained a member of the organization until he left
Wisconsin in 1910. The Council of the American Historical Association chose him as one
of the two most eminent historians America has produced.
3 The above description of Portage is based upon an account Turner himself wrote
to C. L. Skinner, March 15, 1922, reprinted in Burnette, Wisconsin Witness, pp. 65-6.
and in p. xi introducing the same work.
23
24 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
word was not only the most common mode of communication; it
was also the necessary instrument for education, progress and
enlightenment. Indeed, probably no event of importance took place
in Portage without some spokesman or other urging the popu¬
lace to action or stronger belief.
Fred Turner was raised in a literate household. His mother
had been a school teacher and came from a long line of preachers.
His father was editor of the Wisconsin State Register and a pillar
of the community who gave talks on local history and published
several manuscripts in this field.4 Fred Turner early learned the
importance of the printed word. As he wrote his future wife in
later years, “I thank heaven that I have an imagination and a
love of books, two things that have lifted me out of my surround¬
ings at Portage. Life, in any case, is more or less coarse, and re¬
quires imagination to idealize it a bit, but this is especially so in
such an environment as that in which my early life has been
spent.”5
Andrew Jackson Turner was much more than a literate parent.
As editor of a small town newspaper, it was his duty to know what
was going on in town, across the state, and throughout the nation.
Contemporary affairs were part of the regular fare at mealtime
in the Turner household. The elder Turner was also leader of
the Republican party in Portage; he “went as delegate to state
and national Republican conventions, assigned the candidates of
his party to the varied nativities and towns of the county, as chair¬
man of the Board of Supervisors, harmonized the rival tongues
and interests of the various towns of the county, and helped to
shepherd a very composite flock.”6 An expert hunter and fisher¬
man, A. J. Turner often took Fred with him on his forays into
the wilderness. As soon as he was allowed, Fred began to frequent
the newspaper office and in time, his father let him set type for
local news and edit an exchange column.7 As a high school student,
he kept a scrapbook of newspaper clippings including texts of
speeches and quotations from such prominent spokesmen as Emer¬
son, Disraeli, Ingersoll, Webster and Schuyler Colfax. Fred was
4 Burnette, Wisconsin Witness , p. 67 ; p. xi.
5 Turner to Caroline Mae Sherwood, August 24, 1887, TU Box B.
6 Burnette, Wisconsin Witness, “Turner’s Autobiographic Letter”, p. 66.
7 Burnette, Wisconsin Witness, “Turner’s Autobiographic Letter”, p. 66. Turner’s
early - experience working in a newspaper office was later to prove of great value
in his work as a historian : “My practical experience in newspaper work, and in
contact with politics through my father probably gave me a sense of realities
which affected my work and my influence. I had to see the connections of many
factors with the purely political. I couldn’t view things in the purely ‘academic’
way ...” Turner to Merle Curti, August 8, 1928, reprinted extract, Wilbur R.
Jacobs, The Historical World of Frederick Jackson Turner With Selections From
His Correspondence (New Haven: Yale University Press, 1968), p. 6.
1971] Berquist — The Rhetorical Heritage of Turner
25
also among a half-dozen students to declaim on Memorial Day in
1877 and 1878.8
Not much is known about Turner’s day-to-day activities in these
early years but we do have a document which reveals a good deal
about his rhetorical development. In his senior year, Fred partici¬
pated along with eighteen others in an oratory contest, held on
graduation day. Fred won first place and his oration 'The Power
of the Press” was published soon after in his father’s newspaper.9
The first rule in public speaking is to choose a speech topic that
interests you, and this Fred obviously did.
A second rule is to speak in such a way as to interest your
listeners. Fred Turner talked of native democracy, of the role of
the press in educating everyone, not just a powerful elite as the
Athenians did in ancient Greece. The want of education by the
masses is shown by the recent rise in communism, Fred declared.
Orators will continue to persuade their audiences but the press
provides a new dimension, a larger audience. "The Press is an
instrument of unspeakable good in the diffusion of education ....
as the freedom of the Press increases, so does the freedom of the
people” — sentiments dear to the heart of Andrew Jackson Turner
as well as his son. The speech also includes timely references
to the Union Army, the Christian religion, Uncle Tom’s Cabin, and
the Declaration of Independence. The people of Portage could
easily identify with the young orator.
By today’s standards, Fred’s oration would rank "above average”
but not "superior.” What makes this speech significant is not its
rhetorical excellence but its insight into a young mind. Journal¬
ism was Fred’s intended career. His faith in American democracy,
Portage style, would be lifelong. And he would be an avid reader
of books to his dying day.10 But above all else, Fred Turner ex¬
celled as a speaker in an age when oratory was usually a prompt
and sure road to recognition and power.
Rhetorical Experiences at Madison
In the fall of 1878, Fred Turner matriculated at Madison. There
he was exposed to two men who were to have a marked influence
upon his life: President John Bascom, "a versatile scholar, and,
for his day, a progressive thinker on social and economic issues,”
and David B. Frankenburger, newly appointed professor of rhe¬
toric and oratory. President Bascom preached the duty of Uni¬
versity students to improve the state which made their higher edu-
8 Wisconsin State Register , May 26, 1877 ; June 1, 1878.
9 Wisconsin State Register, July 6, 1878.
10 Fred’s conclusion is a flowery peroration to books : “the true Elysian field? where
spirits of the dead converse, and into these fields a mortal may venture unappalled.”
26 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
cation possible, by using the information thus acquired for the
public good.11 Professor Frankenburger was the kind of instructor
who taught individuals, not classes.12 Turner, who spent four highly
successful years under his tutelage, described his rhetoric teacher
as a man with a rare, questioning smile, glad expectancy, appealing
sympathy, an instructor who sought only the best in his students.
Frankenburger encouraged and inspired, made helpful suggestions,
and gave unstintingly of his time and energy. He was himself an
unusually popular reader, lecturer and speaker, a lawyer, prom¬
inent Unitarian and staunch admirer of Emerson.13 In May of
1883, Fred Turner’s oration entitled “The Poet of the Future”
won the coveted Burrows Prize at the Junior Exihibition. A year
later, at graduation, Turner’s “Architecture Through Oppression”
was chosen the best of eighteen orations delivered before a dis¬
tinguished audience including the Governor, President Bascom, and
members of the Board of Regents.14 In a day when skill in ora¬
tory and debate were widely admired, young Fred Turner, the
shy, quiet youth from Portage, had found the key to success and
self-confidence.
Turner did not debate at Wisconsin, probably for two reasons.
In the summer of 1879, Fred became seriously ill and spent the
following academic year recuperating at home. In those days at
Madison, being on the debate squad was a four year proposition
which included an intensive period of research and apprentice¬
ship for the underclassmen.15 In so popular and competitive an
activity, a student who missed a whole year usually missed out
entirely. Secondly, Turner simply did not have a “logical mind” ;16
while he would have no trouble tackling the research on a question,
his forte was imagination. Avery Craven put it this way: “There
was something of the poet and much of the philosopher about Tur¬
ner. He had the ability to see deep into the meaning of things and
the power to catch the universals. This did not weaken his capacity
for scientific research nor lessen his interest in details, but it
did cause him to emphasize trends and flavors, to attempt to deal
11 Authority for the statements in this paragraph comes from the following sources :
University of Wisconsin photostat L 18 14, Turner Papers ; Burnette, Wisconsin
Witness, “Frederick Jackson Turner”, p. 178 ; Merle Curti and Vernon Carstensen,
The University of Wisconsin : A History 1848-1925 (Madison: University of Wisconsin
Press, 1949), 1, 344.
12 The portrait of Frankenburger which follows is one Turner painted himself in
a funeral eulogy to one of his favorite professors, TU Box 55.
13 Curti and Carstensen, University of Wisconsin, 1, 344.
14 Copies of both college orations appear in TU Box 54.
15 A revealing account of the seriousness with which students took their debate
responsibilities in Turner’s day appears in Curti and Carstensen, University of Wis¬
consin, 1, 433-8.
16 “Inviting us one day to consider the problem of sovereignty, he (Turner) quoted
Austin’s definition ; said he couldn’t understand it ; admitted he wasn’t blessed with
the logical mind.” Quoted in Carl Becker, “Frederick Jackson Turner”, American
Masters of Social Science, ed. by Howard Odum (New York: Henry Holt, 1927), p. 278.
1971] Ber quiet— The Rhetorical Heritage of Turner
27
with intangibles, to sweep over minor things in the effort to get
at larger truths.”17
That Turner respected the work done in debate is clear enough.
An entry in his commonplace book, written apparently in Septem¬
ber, 1887, includes notes for a speech to the debate society.18 Debate
work is important, Turner told the undergraduates, because it
helps one to be a ready speaker and exposes the student to much
information he might otherwise miss. There is a danger, Turner
told the squad, to acquiring glibness and getting into careless
habits of enunciation and gesture. "Do not let the violence of de¬
bate carry you away from rules of graceful oratory— -when form
becomes more prominent than the idea— the effect is lost.” Don't be
prejudiced in important questions by reading on only one side
and hearing the opposition in a spirit of antagonism. 'Tight your
side for all it is worth but hold your judgment in reservation.”
"Young speakers are too apt to make their point as a dry resume.”
"Use as few notes as possible.” Excellent advice today as when
it was given over eighty years ago.19
The thrills and excitement attending the return of a winning
college orator in Turner's day is nowhere more clearly docu¬
mented than by Turner himself. Writing from Madison on May
3, 1879, Fred vividly describes the reception given Robert La
Follette upon winning the state title at Beloit— -an account Turner's
father later published.20
Fred Turner's own rhetorical talents were substantial. Each year
his grades improved in Professor Frankenburger's classes until
at last he received a 98 in his senior year.21 Turner's junior ora¬
tion, "The Poet of the Future”, is a philosophical eulogy of modern
science and modern democracy, replete with literary allusions.
In his senior year he chose a simpler theme: that great architec¬
ture is built at the cost of great toil and evil. This final col¬
lege effort is a more professional composition, the best rhetorical
production of his three prize-winning orations.
Undoubtedly Turner's sustained exposure to classical authors
and his continuing work in journalism also contributed to his rhe-
37 Burnette, Wisconsin Witness , “Frederick Jackson Turner, Historian”, p. 110.
is tu VoL III (1-3), commonplace books, 1881—87, n.p.
39 In 1892 when Turner, then a professor of history at Madison, was trying- to per¬
suade Richard Ely to leave Johns Hopkins for Wisconsin, he noted that “vigorous
and deeply interested” students in the debating societies paid more attention to the
department of economics than any other. Turner to Richard Ely, January 25, 1892,
Box E, Ely correspondence (State Historical Society Library, Madison, Wisconsin).
Turner later commended C. H. Haskins to Woodrow Wilson in these terms : “He
is one of the strongest reasoners and effective debaters in our faculty meetings.”
Turner to W. Wilson, December S, 1896, extract reprinted in Jacobs, Historical World,
p. 200.
20 Wisconsin State Register , May 10, 1879.
21 Turner Papers, University of Wisconsin photostat L 18 14, uncatalogued ma¬
terial — Turner’s undergraduate records.
28 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
torical development. At Madison, “he received a thorough classical
training” and did honors work there; quite probably, Turner’s
sensitivity to literary style was cultivated in this way.22 For three
of his four undergraduate years, Turner was associated with The
Badger, the student newspaper.23 As a senior he intended to
make journalism his lifetime career.
Turner’s commonplace books reveal his growing awareness of
style, metaphor, balance and epithet. Begun April 9, 1881, these
student notebooks present a remarkable insight into the mind of
a developing scholar. The materials included are varied indeed:
quotations from books, essays, lectures and speeches; Greek and
Latin translations; drafts of orations;24 lists of books Turner
read, concerts, operas and lectures he attended ; poetry and
songs Turner composed; and random asides about philosophy
and religion, ideas for essays and topics for orations. Turner’s
favorite author was Ralph Waldo Emerson; his picture appears on
the back cover of the first commonplace book. While an extended
analysis of these notebooks is not appropriate here, a listing of
some of the speech-related themes may suffice to suggest the whole :
here are quotations about talk, the spoken word, silence, the human
voice, being understood, the role of character in men of intellect,
elocution, modes of style and originality of thought, rhythm and
meaning, figures of speech and eloquence. Here, indeed, is docu¬
mentary proof of Turner’s rhetorical heritage!
Tutor in Rhetoric and Oratory
The year following graduation Fred Turner turned his attention
to journalism full-time, serving as correspondent for newspapers
in Chicago and Milwaukee.25 He appears to have gotten on well
enough, but something was missing; the career he dreamed of
since boyhood was somehow unsatisfactory. At first, Turner
thought the problem was one of locale and he toyed with the idea
of joining Reuben Gold Thwaites in starting a paper out West.26
But actually the problem lay elsewhere. In his junior year at Madi¬
son Turner had his first exposure to college history with Professor
22 Burnette, Wisconsin Witness , “Frederick Jackson Turner”, p. 178. In 1880 Turner
was asked to tutor a college preparatory student in Greek by his former high school
principal. W. G. Clough to Turner, September 7, 1880, TU Box 1.
23 Turner served at various times as exchange editor, secretary-treasurer, and presi¬
dent of the Badger Association. When Turner first joined, the paper was called
The Campus; it received its new name in 1882. University of Wisconsin, University
Archives, The Badger, Vols. 1—4.
24 According to Wilbur Jacobs, “the drafts of Turner’s college orations sometimes
read like preliminary versions of an essay on the frontier theory, for they deal with
the rise of the common man, freedom, and social evolution.” Historical World, pp. 8-9.
25 Wilbur R. Jacobs, Frederick, Jackson Turner’s Legacy (San Marino: Huntington
Library, 1965), p. 6.
28 Frederick Jackson Turner, “Reuben Gold Thwaites: A Memorial Address”, p. 38,
printed copy, Huntington Library.
1971] Berqui§t—The Rhetorical Heritage of Turner
29
William Francis Allen; he was simply never the same thereafter.
A revealing entry in Turner's commonplace book at this time prob¬
ably holds the key to his decision to make history his. life work:
“Science has revolutionized Zoology, Botany, etc. It must now take
up recorded history and do the same by it. This I would like to
do my little to aid, but find not the time. It is a very egotistical idea
that haunts me that if I were to drop my detestable dishing up of
newspaper flippancy, I could . . ff The sentence ends here.
But how could Turner earn a living while he pursued graduate
work? There simply were no assistantships available in history.
Rhetoric solved his financial problem now as it would frequently
In the years to come.27 Professor Frankenburger arranged to have
Fred appointed instructor in rhetoric and oratory. From the fall
of 1885 to the end of the winter term, 1888, Fred Turner tutored
over a thousand students in the communication skills he had him¬
self so recently mastered.28
As a speech teacher, Turner was. conscientious and content-ori¬
ented. He taught twelve hours of formal speech classes each week
and listened to “rehearsals” an additional hour and a half daily.
Sometimes his work was made easier when the majority of a new
crop of students had earlier training; sometimes, harder because,
as Fred put it, “to tell the truth none of the contestants are natu¬
rally very good dec I aimers/1' When judges were overly long in
rendering a decision in a speech contest on a hot day, Fred lec¬
tured them on their lack of consideration and thereby expedited
matters. Turner preferred oratory to declamation “not being” he
said “much of an elocutionary alchemist myself!”29
Like many another teacher of public speaking, Turner soon
became a highly sensitive critic, reacting to speech communication
wherever he found it. Whether it was poor preaching or the
polished oratory Professor Allen took him to hear at a Harvard
commencement, unenlightening papers read by fellow historians,
President Bascom’s farewell address, or merely social conversation
at a Madison soiree, the critical faculties of the speech teacher
came into play.30
27 Turner had “champagne” tastes impossible to satisfy on existing income. Conse¬
quently he accepted dozens of speaking engagements, before and after 1893, to
supplement Ms salary.
28 Turner Papers, University of Wisconsin photostat L 18 14, uncatalogued ma¬
terial — Turner’s instructional reports.
29 References to Turner’s experiences as a speech teacher appear in the following
letters: Turner to his parents, September 23, 1885, TU Box A; Turner to Caroline
Mae Sherwood, June 6, 1887 ; June 15, 1887, TU Box B.
30 References to Turner as speech critic appear in the following letters : Turner
to Caroline Mae Sherwood, December 12, 1886, TU Box A; June 12, 1887, TU Box
B; Turner to Mary O Turner, June 30, 1887, TU Box B; Turner to William Francis
Allen, December 31, 1888, TU Box 1 ; Turner to Caroline Mae Sherwood, May 15,
1887, TU Box B ; May 5, 1887, TU Box A.
30 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Not surprisingly, Turner was frequently called upon to speak
himself. In 1885, he agreed to replace Professor Allen temporarily
as “recent history” section leader for the Contemporary Club of
the Unitarian Church. Later he lectured to this body on the
fisheries question and with Professor Allen, on the history of
the northwest. In 1887 Turner was chosen to give the address
of welcome at his class reunion. On still another occasion he
served as toastmaster at the annual dinner-dance of his social
fraternity. As a doctoral student at Johns Hopkins in 1888-89
Turner was active in both chautauqua and extension work. Soon
after his return from Baltimore, he was simply unable to satisfy
all the requests he received for extension lectures throughout
the state. Because Turner believed in the need for competent
history teachers in the public schools, he found himself addressing
teachers’ conventions from time to time. Frederick Jackson Turner
mastered the platform as he would later master his profession.31
Turner in the Classroom
Despite his many appearances off campus, it was in the class¬
room that Turner exerted his lasting impact. Turner’s rhetorical
training is clearly a part of his approach to the teaching of
history. In March of 1889, for example, he complained to Professor
Allen that his freshman history students experienced difficulty in
reporting the results of their research in class.32 In class, Turner
himself dropped the role of polished lecturer to take up that of
Socratic inquirer. As Sidney Packard, a former student recalls,
“Turner was a poor lecturer in his classroom almost never getting
very far from a small box of notes. When he did manage to get a
few feet from those notes, or while he was on the way back to
them after explaining a diagram or map, he was another man
entirely and spoke with real eloquence and charm and addressed
himself to large topics in an easy and relaxed manner. He seemed
almost apologetic for so doing as soon as he got back to the
notes.”33
31 References to Turner’s speaking- appear in the following sources : Turner to his
parents, September 23, 1885, TU Box A; Turner to Caroline Mae Sherwood, May 11,
1887, TU Box A; Curti and Carstensen, University of Wisconsin, 1, 723; Turner to
Caroline Mae Sherwood, May 27, 1887, TU Box A; Ray A. Billington, “Frederick
Jackson Turner: Biography of a College Teacher” (An unpublished manuscript Dr.
Billington kindly permitted the author to examine in March, 1970) ; Turner to
William Francis Allen, October 31, 1888, TU Box 1 ; March 14, 1889, TU Box 1 ;
Turner to Herbert Baxter Adams, December 9, 1891, TU Box 1 ; January 18, 1892,
TU Box 1 ; Curti and Carstensen, University of Wisconsin, 1, 642. In one rural Wis¬
consin community of six hundred inhabitants. Turner’s extension lectures attracted
an audience of over two hundred people.
32 Turner to William Francis Allen, March 14; 1889, TU Box 1; Turner to Merle
Curti, August 8, 1928, TU Box 39.
33 Turner Papers, student reminiscences, August 20, 1968, L 18 14, uncatalogued
material ; see also Jacobs, Turner’s Legacy, p. 14.
1971] Berquist—The Rhetorical Heritage of Turner
31
One should not assume that Turner went to class unprepared.
Indeed, just the opposite was true. According to Ray A, Billington,
Turner was never “guilty of using the yellowing notes of yester¬
year; every assignment was freshly prepared in endless hours of
labor. This was illustrated in his last year of teaching in 1923-24.
The lectures for his 'history of the West1 were carefully recast,
even though they had been given dozens of times and would
never be given again. . . . Little wonder that a Harvard under¬
graduate told his tutor that the students learned more from
Turner than from any other instructor, and on being asked why
replied 'Turner gives all his time to us, instead of .spending it
writing books and articles like others !’ ”34 What made Turner
one of the great teachers of his time, Merle Curti maintains, is
the fact that “he possessed the rare gift of inspiring students, of
imbuing them with a deep love of his subject and a belief in its
great importance.”35
Along with many others, Carl Becker believed that a key
ingredient in Turner’s remarkable hold on others was his voice.
Describing his first encounter with Turner as a college sophomore,
Becker wrote “Haltingly I asked my foolish question, and was
answered. The answer was nothing, the words were nothing, but
the voice— the voice was everything: a voice not deep but full, rich,
vibrant and musically cadenced; such a voice as you would never
grow weary of, so warm and intimate and human it was. I cannot
describe the voice. I know only that it laid on me a kind of
magic spell which I could never break, and have never wanted to.”36
As Allyn Young, a prominent economist at Wisconsin and Harvard,
wrote “My wife who is nearly blind and upon whom voice, there¬
fore, makes a deep impression, has said that Turner has the most
pleasantly modulated voice and the most winning manner of speech
that she has ever heard.”37
In his writing as well as his speech, Frederick Jackson Turner
revealed his rhetorical heritage for all to see. Some commentators
were fascinated by the “compelling power” of his metaphor; others
with his poetic touch. A Harvard graduate student fresh in from
southwestern Oklahoma was intrigued by Turner’s ability to
capture the spirit of the frontier West he had recently left: “I
had lived so close to all these things that they were conditions
34 "Why Some Historians Rarely Write History : A Case Study of Frederick Jack-
son Turner,” Mississippi Valley Historical Review 50, No. 1 (June, 1963), 18-19.
See also Jacobs, Turner’s Legacy , p. 14.
35 Burnette, Wisconsin Witness , “Frederick Jackson Turner,” p. 201.
38 Becker, “Frederick Jackson Turner”, p. 276; see also George C. Sellery, “The
Spirit of Wisconsin”, baccalaureate address, June 20, 1937, TU File Drawer 15— E ;
and Richard Hofstadter, The Progressive Historians: Turner , Beard and Farrington
(New York: Alfred A. Knopf, 1968), pp. 81-2.
37 Jacobs, Historical World , pp. 14-15.
32 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
to be accepted as a matter of course. It had been impossible for
me to see the forest for the trees, or the city for the houses.
But Turner’s lectures changed all this.” Still others commented
on Turner’s philosophical insight and broad humanitarianism.
Joseph Schafer summed it all up best when he wrote “Other
things being equal, men and women prefer to work under a
leader instinctively recognized as a supreme gentleman. But when,
to that character, is added not only the profound historian and
philosopher, but also the artist in speech and the poet in imagina¬
tive conception, even the unseeing feel the resulting blend as
something strictly unique, to be enjoyed as a 'gift of the gods.’ ”3S
Three historians clearly perceive the rhetorical spirit of Turner’s
work. Ray A. Billington reports that Turner “enlisted an army
of propagandists who preached the gospel of the frontier in class¬
rooms and seminars throughout the land.”39 John D. Hicks
observed of Turner that “In his essays, most of which were
written to be read aloud, he could rise to the heights of eloquence.”40
A final observation comes from one who knew Professor Turner
better than anyone else: “My mother’s ancestors were preachers!
Is it strange that I preached of the frontier?”41
38 The various facets of Turner’s rhetoric cited above are identified in the following
sources : Frederick Jackson Turner, The Significance of the Frontier in American
History, ed. by Harold P. Simonson (New York: Frederick Unger, 1963), pp. 9-10;
Turner Papers, reminiscences of Merrill Crissey, TU Box 57 ; Hofstadter, The
Progressive Historians ; p. 73. Edward Everett Dale, “Memoirs of Frederick Jackson
Turner”, Mississippi Valley Historical Review, XXX, no. 3 (December 1943), 340,
342 ; Frederick Jackson Turner, The Frontier in American History, foreword by Ray
A. Billington (New York: Holt, Rinehart and Winston, 1962), p. xviii ; Joseph
Schafer, “Editorial Comment — Death of Professor Turner”, Wisconsin Magazine of
History, XV, no. 4 (June, 1932), 497. From the time he composed his junior oration,
“The Poet of the Future”, it was apparent that Turner was philosophically oriented,
that he yearned to grasp the significance of events and periods rather than merely
capture their details. Fulmer Mood, “Turner’s Formative Period” in Everett E.
Edwards, The Early Writings of Frederick Jackson Turner (Madison: University
of Wisconsin Press, 1938), p. 6; Becker, “Frederick Jackson Turner”, p. 296. Such
a mind could be strongly prophetic on occasion. Thus one of Turner’s speeches in
the 1920’s “gloomily forecast what portended for America: population pressures;
diminishing food supply ; the exhaustion of forest, oil, and coal reserves ; the threat
of war ; the horror of a dreaded ‘chemist’s bomb’. He also lamented in his addresses
the increasing tendency toward conformity with an accompanying ‘decline in self-
confidence’ in America.” Jacobs, Turner’s Legacy, p. 42.
39 Turner, The Frontier in American History, foreword by R. A. Billington, p. xiii.
40 Review of Wilbur Jacobs, The Historical World of Frederick Jackson Turner
With Selections From His Correspondence, by John D. Hicks, Journal of American
History, LVI, no. 2, (September, 1969), 413. In 1920 for example Turner was per¬
suaded to publish a collection of his essays under the title The Frontier in American
History. Ray Billington wrote of this work in 1967, “no one volume has done more
to reshape the writing of American history or to recast the popularly held image
of the American past” ; of the thirteen essays included, nine were originally presented
as speeches. Turner, The Frontier in American History, foreword by R. A. Billington.
41 Burnette, Wisconsin Witness , “Turner’s Autobiographic Letter,” p. 67.
THE GOSPEL OF POVERTY: THE MESSAGE OF CONSERVATIVE
PROTESTANTISM TO THE POOR AT THE TURN OF THE CENTURY
Walter F. Peterson
An article simply entitled “Wealth” appeared in the North Amer¬
ican Review of June 1889. The editor declared that this article
written by the articulate industrialist Andrew Carnegie was “the
finest article I have ever published in the Review”1 The former
bobbin boy detailed three “modes” by which the wealthy man could
dispose of his surplus : “It can be left to the families of the descend-
ents ; or it can be bequeathed for public purposes ; or, finally it can
be administered during their lives by its possessors.” Carnegie held
that the first was “the most injudicious.” The failure of man to
dispose of his accumulated wealth during his lifetime marked him
as selfish and lacking in foresight.
The rich man is thus almost restricted to following the examples of Peter
Cooper, Enoch Pratt of Baltimore, Mr. Pratt of Brooklyn, Senator Stanford,
and others, who know that the best means of benefiting the community is
to place within its reach the ladders upon which the aspiring can rise — •
parks, and means of recreation, by which men are helped in body and mind;
works of art, certain to give pleasure and improve the public taste, and
public institutions of various kinds, which will improve the general condi¬
tion of the people; in this manner returning their surplus wealth to the
mass of their fellows in the forms best calculated to do them lasting good.
The solution to the problem of wealth and poverty, the rich and the
poor had been discovered. “Such, in my opinion,” said Carnegie,
“is the true Gospel concerning Wealth . . .”2
Few literate Americans at the turn of the century escaped ex¬
posure to Carnegie's apologia for the accumulation of wealth. The
spirit of acquisitiveness abroad in the land had assumed enormous
prestige and support. From 1890 to the present the upholding of
this doctrine, or attack upon it, the examination and re-examination
of Carnegie's and succeeding positions have led to the development
of a sizable body of literature on the Gospel according to “Saint
Andrew.”
Studies of the Gospel of Wealth normally cite the writings of
President James McCosh of Princeton and Noah Porter of Yale,
Russell Conwell, whose “Acres of Diamonds” was supposed to have
1 Quoted in B. J. Hendrick. The Life of Andrew Carnegie (New York, 1932), I, p. 330.
3 Andrew Carnegie, “Wealth,” North American Review , 148 (June, 1889), pp. 653-664.
33
34 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
been repeated six thousand times throughout the East and Mid¬
west, and conclude with the benediction of the Right Reverend Wil¬
liam Lawrence, Episcopal Bishop of Massachusetts.3 But lesser
men, ministers of all the major Protestant denominations, who
were often closer to the people, endorsed the same general prin¬
ciples with equal and sometimes greater fervor. They held that
wealth and civilization went together and provided man with the
possibility of rest and reflection. The worshipers at Plymouth
Congregational Church in Indianapolis heard that “Wealth is the
rich soil in which a human soul-root unfolds its powers and becomes
its possibility. God meant we should flee poverty/’4 The question of
accumulation was not one that the Christian could view dispassion¬
ately. The chancellor of Nebraska Wesleyan University pointed out
that it was no sin to get money provided it was done through honest
methods : “Indeed to gain in this way is a Christian duty.”5
Through the assiduous application of their Christian duty men
such as George Peabody and A. T. Stewart made millions of dollars.
Charles E. Bronson, pastor of the First Presbyterian Church,
Saginaw, Michigan, thanked God for the noble examples of John
Wanamaker and Chauncey Depew, who proved to the young men
of America that it was possible to reach the highest posts in busi¬
ness and at the same time preserve the Christian character.6 Of
course they did not do these things alone, for God put the twelve
millions of dollars in Peabody’s hands. “Everybody honors him
because he saved his country from financial ruin, set up institutions
of learning, and because he was always looking for an opportunity
to make some one happy,” S. P. Long told the First Lutheran
Church of Mansfield, Ohio.7 To accomplish these good works a man
had to be determined, sometimes even as harsh as A. T. Stewart
when he sent a servant out of his home because she burned the two
ends of a match when matches were very dear. When the girl’s
father found fault with the great merchant for giving a large sum
to the church the next week, Stewart replied, “ ‘If I had not saved
3 See the brilliant chapter entitled “The Gospel of Wealth of the Gilded Age,” by
Ralph Henry Gabriel, The Course of American Democratic Thought (New York, 1956),
pp. 151-169.
4 Oscar C. McCulloch, The Open Door: Sermons and Prayers (Indianapolis, 1892),
p. 145. McCulloch who was minister of Plymouth Congregational Church, Indianapolis,
Indiana, made this statement in a sermon entitled “The New Vow of Poverty.” John
Sweet, pastor of the First Methodist Episcopal Church, Owosso, Michigan said much
the same thing : “Money is power, and all power is good in the hands of those who
know its proper place and use,” in C. S. Eastman, ed., The Methodist Episcopal
Pulpit (Monroe, Michigan, 1897), p. 140.
5 D. W. C. Huntington, Half Century Messages to Pastors and People (Cincinnati,
1905), p. 164.
6 Charles E. Bronson. The Presbyterian Pulpit ; a Volume of Sermons by Ministers
of the Synod of Michigan (Monroe, Michigan, 1898), p. 213.
7 S. P. Long, The Eternal Epistle: Sermons on the Epistles for the Church Year
(Columbus, 1908), p. 399.
1971]
Peterson— The Gospel of Poverty
35
the two ends of the match, I could not have given this large sum
for this benevolent cause/ ”8
The Christian benefactor was a selfless man who gave mostly to
religious institutions and did so with some self-effacement. On this
count even the great and generous Carnegie did not escape the
chastisement of the irascible S. P. Long. When the wealthy iron¬
monger refused to support Wesleyan University and the Protestant
Hospital of Columbus, Long concluded that he was no Christian.
Long also resented the Carnegie library program. “I do not blame
a man even like Carnegie for putting up a library in every city if
he can get the dumb public to pay for half of his monument.” He
held that the difference between a man of God and a man of the
world is that the man of the world wants all the glory himself while
the man of God wants the glory to go where it belongs, to the God
who gave him the money.9
While righteousness and business success normally went hand
in hand, all affairs of men were seen as guided by God’s unerring
wisdom. uHe knows when to give and when to withhold, when to
check and when to impel, when to enrich and when to impoverish,
when to create and when to destroy,” wrote Lutheran theologian
Luther Gotwald in Joy in the Divine Government , In short, God
cannot be guilty of the slightest mistake. The students of Matthias
Loy, Professor of Theology in the Evangelical Lutheran Seminary
at Columbus, Ohio, were told that this same God assigns men to
their place in society. One man was a merchant prince, another but
a servant in his house or a mechanic in his shops. To complain,
therefore, was to show a thankless heart and behave in a manner
unworthy of a Christian who knows that God bestows different
gifts. If his place in society was lower than that of another, he
should not be envious of the man whose station was higher. Since
each man receives only what is right and 'fair in the eyes of the
Lord, each individual should be content with his lot in life.10
But it was not altogether likely that the poor, content in the
knowledge that it was God’s will, would accept a perpetual state
of poverty. Many Protestant ministers at the turn of the century
recognized this possibility. The revolt of the Grangers and Popu¬
lists, the railway strikes and the Haymarket Riot were convincing
evidence that this situation was unfortunately true. As a conse-
8 S. P. Long, The Great Gospel (Columbus, 1904), pp. 571-572.
9 S. P. Long, The Eternal Epistle , p. 400.
10 Luther A. Gotwald, Joy in the Divine Government ; and Other Sermons (New
York, 1901), p. 8. Matthias Loy, Sermons on the Epistles for the Sundays and Chief
Festivals of the Church Year (Columbus, 1900), pp. 140-141. These statements have
the same tone as those of Henry Ward Beecher a half century earlier in Life
Thoughts (Boston, 1858), p. 181. “We have aristocrats, but God made them. ... It
was designed that some should be high, some intermediate, and some low, as trees
are some forty, some a hundred, and some, the giant pines, (how solitary their tops
must be!) three hundred feet in height/'
36 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59>
quence, to provide for those to whom the “Gospel of Wealth” did
not apply, they built an elaborate rationale that might be called
the “Gospel of Poverty.” This statement did not appear as one
simple and consistent declaration of good news to the poor but
rather as a variety of assertions which can be seen as variations-
on the basic theme.
Perhaps the least acceptable answer to the question was the one
proposed by the Lutheran minister, Olaf Lysnes. If you have ac¬
cepted Jesus then you are independently rich even though you may
be poor in this world’s goods. Suddenly you belong to the nobility
of the household of faith.11 In Minneapolis the members of the First
Baptist Church heard much the same message. “Talk about Pov¬
erty ! He is not the poorest man who is out of money, whose build¬
ings have gone up in smoke, whose deposit account is swallowed
up in bankruptcy. The poorest man is the man who is without God,
and without hope, the man from whom the Scriptures have been
snatched away.” Thus did the Reverend Doctor William B. Riley
admonish the city’s poor in Messages For the Metropolis.12
If the poor still chafed under the burden of their worldly troubles
they were counselled to keep in mind that their troubles were really
of no consequence. Samuel Smith Harris, Episcopal Bishop of Mich¬
igan declared that he did have a gospel for the poor which they
would find soothing, for it was full of the good news of hope, love
and life. This gospel proclaimed that the wants and needs, the
poverty of the unfortunate man, belonged to a world that was pass¬
ing away. God had another world, a real world, in store for him
where all inequalities of the present life would be redressed, for
God had reserved all of eternity merely to console the poor. Heaven
would be
An eternity of peace for the troubled, of rest for the weary, of joy for the
afflicted, when men and women and children shall hunger no more, and
thirst no more; where there shall be no more pain, neither sorrow nor cry¬
ing, for God shall wipe away all tears from their eyes. Ah, yes, this begins
to sound like good news indeed, like a real gospel to the poor.13
S. P. Long, who preached to as many as three thousand people
11 Olaf Lysnes. “The purpose of Jesus’ Poverty,” in Pastors of the United Norwegian
Lutheran Church of America , a Free Text Church Postil (Minneapolis, 1913), pp.
36-37.
12 William B. Riley. Messages for the Metropolis (Chicago, 1906), pp. 185-186.
S. P. Long in The Great Gospel, pp. 568-569, wrote, “Who are the poor people in
this city of Mansfield? The very people who are giving nothing to God.”
13 Samuel Smith Harris. The Dignity of Man ; Select Sermons (Chicago, 1889),
p. 252. Bishop Matthew Simpson of the Methodist Episcopal Church preached much
the same doctrine when he said, “If I have but little treasure on earth, I can have
treasure laid up in heaven. There are rich men on earth who will not be rich in
eternity ; and there are poor men who will be rich in the day of the Lord Jesus ; and
if the mind can be thrown forward thus, how this view of the future will compensate
for the privations of the present!” Sermons by Bishop Matthew Simpson (New York,
1885), p. 232.
1971]
Peterson-— The Gospel of Poverty
37
each Sunday in Mansfield, Ohio, addressed himself to the topic of
“Plain Philosophy for Poor People” on Christmas morning, 1903.
He called the attention of his congregation to “those homes in the
tenement houses, dark, black, filthy rooms, drunken husbands,
sometimes no clothing to wear, no bread to eat, no decent meal
for the children, and it almost makes our hearts bleed.” Yet there
was another side to this question. Christ the Savior of the world
was born not even in a tenement, but as the poorest child on earth
never was born, in a low, common stable in order that the poor
might have comfort.
There is comfort for the poorest people in the world; they can live until
they die, and that is all the rich can do. We are here, my friends, to live,
and it does not take a great deal to exist, and when life is done there is just
as much in store for the poorest man that ever lived as there is for the
wealthiest. When life is over, then comes death, and let me say to all the
poor this morning, that that is all the rich have; they simply live and at
the end of life comes death; they take nothing with them, and how much
better off are they than the poorest?
The poorest people can live and die saved. “This my friend is
the true philosophy of a Christian — Plain Philosophy for Poor
People.”14
The Presbyterian leader J. G. K. McClure, President of McCor¬
mick Theological Seminary, held that people could gain an insight
into the entire question of poverty by seeing how Jesus dealt with
it when he told the rich young ruler to sell all that he had and
give to the poor. Actually, Christ did not mean that man should
take the injunction literally, for the young ruler would become
forever a poor man himself and such a step would be most unwise.
Christ felt so deeply for the poor that he would never do anything
to weaken or hurt them. The outright gift of money would be the
worst possible use of the young ruler’s goods because it would dis¬
courage efforts for self-support and thus most certainly injure the
poor. “What the poor always need — whether they are poor in
money, or poor in strength, or poor in comfort, is stimulus and
encouragement to rise above their circumstances, to struggle be¬
yond them, to have a larger spirit, and to put forth an earnest
effort.”15 This treatment of the question, an evasion of the real
issue, seemed to be quite standard in Protestant circles at the turn
of the century.
If man needed a stimulus to rise, as McClure suggested, no
better one could be found than in poverty itself assisted by calamity.
14 S. P. Long. The Great Gospel, pp. 8 9-9 0.
15 James G. K. McClure. Loyalty ; The Soul of Religion (Chicago, 1905), pp. 94-95.
S. P. Long in Prophetic Pearls (Columbus, 1913), p. 195, wrote, ‘We never can help
this world back to God till we mingle with the poor and extend them a warm, helping
hand.”
38 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
In fact, it would be disastrous to do away with poverty, for it was
a most valuable condition. Just as gold and silver are dug out of
rugged mountains, so out of the hardships of life come some of
the richest treasures of human experience and of human character.
Many of the greatest men in history have come from the most
humble surroundings, and had they not been faced by this chal¬
lenge of poverty they might never have reached the heights of
fame. The struggle with poverty and hardship developed a strength
of character that could not be found in any other way. The best
thing in life was to struggle with these difficulties and overcome
them.16 It was much better that a man struggle with poverty, than
with wealth, for “through the liftlock prosperity , one man passes
upward while ninety-nine pass downward. Through adversity one
man passes downward while ninety-nine pass upward/’ Obviously,
our society must preserve and cherish poverty to develop men of
character in the future.17
The man who could find no consolation in adversity as a stimulus
to building character and who, though striving mightily, still failed
to rise above his poverty, was counselled in another fashion. As
a mere “drudge” in an industrial society he must cultivate a spirit
of love “ere he is perfectly at home and happy in his task.” His
honor was at stake to do his humble part in God’s plan.18 “The
vision of someone who is helped will be his final motive to the
humble and complete and glad offering up of himself.” Through
his faithfulness others were made rich. “The factory hand, the
miner under ground, the seamstress in her chamber, the laborer
by the roadside, wear out their lives for others’ comfort. In the
accomplishment of great enterprises how many die for the honor
and the progress of mankind!”19 Congregationalists in Wilmette,
Illinois, and Presbyterians in West Bay City, Michigan, were thus
admonished to give their all in the name of progress.
The new industrial society continued to widen the gap between
the poor, who seemed only to become poorer, and the rich whose
wealth seemed continually to accumulate. Because of the growing
disparity a very definite effort was made to prove to the poor how
16 John Woods, Pastor of the First Presbyterian Church, Ludington, Michigan,
The Presbyterian Pulpit, pp. 115-116.
17 P. E. Holp, Pastor of the Congregational Church, Sioux Falls, South Dakota, in
The Golden Age and Other Sermons (Sioux Falls, 1887), p. 177. Precisely the same
point of view was expressed by the Disciples of Christ minister J. Z. Tyler in Talks
to Young People (Cincinnati, 1896), pp. 15-16: the Presbyterian minister David
Edwards Beach in Sermons and Addresses (Marietta, Ohio, 1890), pp. 199, 272, 276;
and the Lutheran minister Luther A. Gotwald in Joy in the Divine Government,
pp. 12, 58. These expressions were in the same vein as those of Henry Ward Beecher
in Life Thoughts published in 1858, p. 73, when he said “How blessed, then, is the
stroke of disaster which sets the children free, and gives them over to the hard but
kind bosom of Poverty, who says to them, ‘Work!’ and, working, makes them men!’’
18 Roy Edwin Bowers. The Wholesome Life (Chicago, 1911), p. 67.
19 E. K. Strong. “Life in Death,” in The Presbyterian Pulpit , pp. 367—368.
1971]
Peterson - — The Gospel of Poverty
39
very much they actually had to be thankful for. B. L. McElroy of
the First Methodist Episcopal Church, Ann Arbor, Michigan,
stressed a “doctrine of compensation,” which, he said, “is calcu¬
lated to be of service to all who have not made themselves utterly
‘reason-proof/ ” This doctrine proposed some measure of mediation
between the rich and the poor. McElroy hoped that the gap between
these two groups would be narrowed if men would just remember
that “for everything they have missed they have gained something
else.”20 Possession became only another name for relinquishment. If
the poor would only open their eyes to that fact they would look
without envy on the holdings of the rich. By way of illustrating this
principle, the Episcopal rector in Muscatine, Iowa, notes that most
wealthy men had ruined their health in the process of acquiring
their riches and then would have given all their accumulated wealth
if only they could have had in return the good health of the com¬
mon laboring man.21
In the kitchen of the General Otto H. Falk mansion as it over¬
looks Lake Michigan from Milwaukee is a plaque hung at the turn
of the century by the General for the edification of his servants.
The central portion of the plaque shows a happy, care-free servant
giving counsel to a troubled king. Beneath the figures an inscription
reads: “Riches are always restless; ’tis only to poverty the gods
give content.” Falk, a good Episcopalian, and one-time President
of the Allis-Chalmers Manufacturing Company, felt that this mes¬
sage was salutary. This argument was carried into the pulpit by
Louis Buchheimer, a Lutheran pastor in St. Louis, and his state¬
ment constitutes the ultimate exposition of the Gospel of Poverty
as it carried the good news concerning their condition to the poor.
Buchheimer pointed out that there was really no very great dis¬
tinction between the gifts of God to various men. The rich man had
his park, but the poor man could look at it and enjoy it without the
expense of maintaining it. Although others lived in stately man¬
sions, they had to pay very heavily for the privilege. While the rich
man may have had a valuable picture gallery, the poor man could
see in the sunrise and sunset a splendor that no artist could ever
capture. While the poor man did not possess some of the con¬
veniences and delights of the more favored, in return he was free
from many embarrassments to which the wealthy were subject.
By the very simplicity and uniformity of his life the poor man was
mercifully delivered from the great variety of cares that continu¬
ally plagued his wealthy brother. Surely the plain meal eaten with
20 B. L. McElroy. The Methodist Episcopal Pulpit, pp. 256-258.
21 Edward Clarence Paget, Silence: With Other Sermons (New York, 1896), pp.
116-117.
40 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
relish and appetite by a poor man was more delicious than the
most luxurious banquet.22
Andrew Carnegie, addressing the Nineteenth Century Club at the
close of 1887, exclaimed, “I defy any man to show that there is
pauperism.”23 Those Protestant ministers who preached the Gospel
of Poverty agreed completely with the author of the Gospel of
Wealth. In Minneapolis, William B. Riley said that he knew for a
fact that the inhabitants of “ninety-nine houses out of a hundred
are in perfect comfort.”24 S. P. Long told his congregation that
every man is supposed to own his own home.25 Even evangelist Billy
Sunday, the ordained Presbyterian minister who worked with the
poor and downtrodden more than did most of his brethren, said
that “I do not believe that there are any people beneath the sun
who are better fed, better paid, better clothed, better housed, or
any happier than we are beneath the stars and stripes— no nation
on earth.”26
The working man in the United States may have been better off
than his counterpart elsewhere, but how prosperous was he in the
decade of the nineties? Industrialization was making the United
States an increasingly wealthy nation. But hours of work were
long and wages were low. According to one estimate, weekly hours
of work in manufacturing industries averaged sixty in 1890 and
fifty-nine in 1900, and annual earnings of all wage earners except
farm laborers averaged only $486 in 1890 and $490 in 1900.27
According to the census of 1900, two-thirds of the male workers
over sixteen years of age earned less than $12.50 per week.28 It is
true that prices were not high. In 1898 sugar sold for 5% cents
a pound, coffee for 28 cents a pound, roasting beef for 14%. cents
a pound, and milk for less than 6 cents a quart. Coal cost $6 a ton,
men’s heavy shoes $2 a pair, and the best ready-made suits sold
for $20 or less.29 But even with prices so low, the real income of a
great many American families was very meager indeed. Without
question, the condition of vast numbers of Americans at the turn
of the century bordered on poverty.
Because they failed to recognize the facts of urban American
industrial life as they applied to the working man, most of the
Protestant clergy consistently misread the causes of poverty. The
22 Louis Buchheimer, pastor of the Evangelical Lutheran Church of Our .Redeemer,
St. Louis, Missouri, in Faith and Duty ; Sermons on Free Texts with Reference to the
Church Year (St. Louis, 1913), pp. 24, 81-82.
23 Quoted in Social Science Review (December 14, 1887), p. 9.
24 William B. Riley. Messages for the Metropolis, pp. 10—11.
25 S. P. Long. The Eternal Epistle, p. 148.
26 William T. Ellis. Billy Sunday: The Man and His Message ( , 1936), p. 363.
27 Paul H. Douglas. Real Wages in the United States, 1890-1926 (New York, 1930),
Table 147 opposite p. 392.
28 Bureau of the Census, Employees and Wages (Special Report, 1900), pp. ci-civ.
29 Bureau of the Department of Labor, No. 18 (September, 1898), p. 696.
1971]
Peterson ■ — -The Gospel of Poverty
41
catalog of reasons given for the existence of poverty included sloth
and indifference, the failure to anticipate expenses through saving,
and the use of liquor and tobacco.30 Very considerable emphasis
was placed on the lack of cleanliness, thrift and economy. “If they
knew those things,” T. G. Soares, a Baptist minister in Oak Park,
Illinois, wrote, “they would not be needy. It is foolish to be im¬
patient with the poor, because they have not the methods and
virtues of the successful. If they had them they would not be
poor.”31 Grinding poverty was also seen as the warning hand of
God. In 1908, S. P. Long said that there were families without
bread in every city in the land and the reason for it was very
simple. “The good Lord cannot bear it any longer to see families
go to hell with full stomachs and no knowledge of Him. So He now
pulls the bell of famine and rings into our ears: ‘Prepare to meet
thy God !’ Oh, what an unthankful world! You unthankful souls, I
am surprised that God has not starved you long ago!”32 A far
easier solution was not to resort to reason or explanation but
simply write off the poor as did J. G. K. McClure. “People are not
alike. Those who grow up in slums and are foul with evil from
their youth are different from those who grew up clean and whole¬
some in religious homes.”33
But this was the period also of the rise of the Social Gospel. At
the turn of the century Walter Rauschenbusch, Washington Glad¬
den, Shailer Mathews, Harry F. Ward and Charles Stelzle were
issuing a clarion call to the churches to accept their social responsi¬
bilities and reorient the pattern of Protestant thought and action
to meet the challenge of a rapidly developing urban, industrial
society. The impetus within the churches for a Social Gospel was
provided, however, by only a small but articulate group. These
liberal Protestant leaders represented the religious part of the
Progressive period in American history. The Social Creed of the
Churches adopted by the Federal Council of the Churches of Christ
in America in 1912 was the high point of the Social Gospel.34 But
it had little or no impact on the average Protestant minister in
Beaver Dam, Wisconsin, or Mansfield, Ohio, largely because the
Protestant church had, by the turn of the century, been captured
30 B. L. McElroy, “Compensation, A Law of Life,” in The Methodist Episcopal
Pulpit, p. 358. S. P. Long-, Prophetic Pearls, p. 102. Billy Sunday, Billy Sunday’s
Sermons in Omaha (Omaha, 1915), p. 114.
31 Theodore Gerald Soares, The Supreme Miracle and Other Sermons (Chicago,
1904), pp. 35-36.
32 S. P. Long, Prepare to Meet Thy God (Columbus, 1908), p. 62.
33 James G. K. McClure, Loyalty ; The Soul of Religion, pp. 98-99.
34 See Harry F. Ward, ed., Social Creed of the Churches (New York, 1912), for not
only the Creed itself but also the official definition of each of the articles. Article 16
was the most far-reaching, “For a new emphasis upon the application of Christian
principles to the acquisition and use of property, and for the most equitable division
of the product of industry that can ultimately be devised.”
42 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
by the middle class and because most Protestant ministers simply
did not understand the age in which they lived.
The clergy were not prepared to meet the new problems of the
city and industry directly, for they had no contact with them. Only
very rarely did ministerial candidates come from those classes pro¬
duced by modern industrial society — the very rich and the urban,
industrial poor. Charles Stelzle once observed that the average
Protestant minister believed that having come from a poor home
he was in a position to sympathize with the unfortunate. At the
same time these ministers completely failed to comprehend the
problems of industrial and urban poverty and unemployment.
Stelzle made a survey of ministers to study this matter and found
that fully ninety per cent of the men that he interviewed in city
churches had been born and reared in what might be classed as
rural areas. The poverty to which they referred was the simple
life of the small town or of the farm which was quite a different
thing from the pangs of cold and hunger which he had experienced
in city tenements. Leaving their rural areas for college and the
seminary, they had no occasion to encounter the realities of city
life as they affected the urban poor. The result was an ever-
widening rift between the American worker and Protestantism.35
To an increasing extent, the great American middle class came to
sustain the churches of the major Protestant denominations as the
nineteenth century progressed. Clearly the upper and lower strata
of society, both from an economic and educational point of view,
had by the turn of the century ceased to actively support these
denominations. Now the “solid” and “responsible” middle class,
including the employers, salaried persons, small tradesmen and
farmers, came to be identified with the major Protestant denomina¬
tions. Among them were also included those who had a personal or
sympathetic attachment to this class or were engaged in personal
service. Some ministers candidly admitted that their churches min¬
istered primarily to the middle class.36
What was the answer of Protestantism to the new age that was
emerging at the turn of the century ? The great mass of the Protes¬
tant clergy, conservative in temper and philosophy, beholden to
35 Charles Stelzle, A Son of the Bowery ; The Life of an East Side American (New
York, 1926), pp. 82-83. In a sample of 1800 ministers taken in 1930 it was found that
only 12 per cent were reared in cities over 100,000 population and 48 per cent came
from communities of less than 1000. In this same sample, less than 1 per cent reported
the economic status of their parents as wealthy, about 4 percent said their parents
were poor. Well over half stated that their fathers were farmers or small tradesmen.
Mark A. May, “Theological Education,” in Samuel McCrea Cavert and Henry Pitney
Van Dusen, eds., The Church Through Half a Century: Essays in Honor of William
Adams Brown (New York, 1936), p. 257.
36 Andreas Bard, The Dawn of To-Morrow and Other Sermons (Burlington, Iowa,
1911), p. 30. Lewis O. Brastow, The Modern Pulpit: A Study of Homiletic Sources
and Characteristics (New York, 1906), p. 321.
1971]
Peterson — The Gospel of Poverty
43
their middle class congregations, beat a dismal retreat in the face
of growing problems. This trend can be seen in Manhattan where,
in the period before 1900, 40 Protestant churches left the district
below 20th Street while 300,000 immigrants and workers moved
into the same area.37 Most ministers were not conscious of society
as a whole except to resent the intrusion of new problems on their
peace of mind. The course of least resistance was to maintain the
old individualistic approach and respond to the new questions with
the same old answers. The Gospel of Poverty was the message of
conservative Protestantism to the poor at the turn of the century.
Perhaps it was just as well that few of the class for which this
gospel was intended ever heard the sermons preached ostensibly for
their benefit to middle class congregations. However, the develop¬
ment of this message was an interesting intellectual exercise for
many a conservative Protestant clergyman and provided peace of
mind for the middle class parishioner.
37 Charles Howard Hopkins, The Rise of the Social Gospel in American Protestantism,
1865-1915 (New Haven, 1940), p. 250.
(
THE BROWN RAT IN EARLY WISCONSIN
A. W. Schorger
Rats have been the greatest plague of all animals to mankind.
On account of their destruction of provisions, property, and as
carriers of disease, they have been universally detested. The two
most common rats in the United States are the brown or Norway
rat (Rattus norvegicus) and the black rat (Rattus rattus), the
former being by far the most abundant.
It is commonly stated on the authority of Pallas that the brown
rat entered Europe in 1727 at which time vast numbers swam the
Volga. According to Baumann (1949) this species arrived in
Europe in the 15th century and not in the 18th. The dates of its
arrival in several European countries are given by Barrett-
Hamilton (1910) : Denmark, 1716; Norway, 1762; Sweden, 1790;
France, 1750; England, 1728 or 1729 (Pennant, 1761-66).
An attempt to determine the dates of arrival of the two species
in North America is made extremely difficult since usually only
“rats” are mentioned without specific information for identifica¬
tion. The black rat was the first to arrive. Both species were brought
by ships. Port Royal, Nova Scotia was founded in 1605. Lescarbot
(1914 :226) was present from July, 1606 to July, 1607. In the mean¬
time rats had increased greatly and had traveled a distance of
over 400 paces to the Indian lodges. At Jamestown, Virginia, in
April, 1609, the corn imported for the colonists was almost totally
destroyed by decay and the “many thousand rats” (Smith, 1910).
To have reached this abundance, rats must have arrived on the
ships of 1607. Lawson (1907:129) came to North Carolina in
1700, and nine years later wrote that they had the same house rat
as in Europe. When Kalm (1772 :348) was in New Jersey in Janu¬
ary, 1749, he observed that the rats were of the same size as those
in Sweden but differed in color, being “grey or blue-grey.” Both
brown and black rats may have been present. The black rat was
frequently called the blue rat in Pennsylvania at the beginning
of the present century (Rhoads, 1903). Pennant (1761-66) was
assured by several natives of Scandinavia that the Norway rat
was unknown there and stated that Linnaeus failed to take notice
of it ; hence Kalm may have referred only to the black rat.
45
46 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Methods of Distribution
Rats were distributed mainly by boats such as sailing vessels,
steamers, and the large barges and scows of traders. They were
carried inland, from waterways by freighters, and in the wagons
of emigrants which usually carried seeds for planting and goods
protected from breakage by packing in straw. Cook stoves were
brought to Fort Wayne, Indiana, in 1836. The parts, wrapped in
straw, were packed in crates. When these were opened several
rats escaped to form the first colony (Griswold, 1917). At this
time nearly all freight arrived by water. The railroad did not come
until much later. About 1811, Audubon (1851) observed several
Norway rats escape from a barge from New Orleans which was
unloading freight at Henderson, Kentucky. The Norway rat ap¬
peared at Brookville, on the White River, Franklin County, Indiana,
the summer of 1827. The black rat, which was numerous at the
time, disappeared in a year or two (Hammond, 1869). Both species
undoubtedly arrived by way of the Ohio River.
A few migrations of the brown rat in large numbers have been
recorded (Lantz, 1909). Since it is mainly nocturnal, the move¬
ments of individuals in frequency and distances overland are diffi¬
cult to determine. Migration appears to be induced by a shortage
of food or an unfavorable environment. Rats usually remain in
the vicinity of buildings but in summer and fall sometimes move
away from them.
When Maximilian (1906, 2:235) was at Fort Clark in 1833 the
Norway rat had not yet reached the Indian villages but the Indians
killed on the prairie seven that were on their way from the fort.
The Minitari villages were at the mouth of Knife River, Mercer
County, North Dakota and Fort Clark was on the Missouri eight
miles below them. Lyon (1936) mentions that he had taken this
rat on a wild island in the Potomac. It could be reached only by a
swim of at least 100 yards. In September, 1899, Snyder (1902)
observed one running along the beach at Fox Lake, Dodge County,
Wisconsin, fully a mile from any building. Some of the marked
rats released in New Orleans were subsequently trapped four miles
distant (Nelson, 1917).
The fall movement is usually to cornfields. In Rock County Jack-
son (1908) found them in these fields, the only localities at a dis¬
tance from buildings. Errington (1935) in the early winter of
1930-31 found them living in holes in the banks of Lake Kegonsa
where they subsisted on dead fish, ducks, and other animal matter.
For the most part they attempted to winter in corn shocks but
few if any survived the season. When the old sandpit near Uni¬
versity Bay was used as a refuse dump, it was infested with rats.
1971] Schorger—The Brown Rat in Early Wisconsin 47
In winter I saw much sign of them in the corn shocks in an adja¬
cent field. The great horned owl is one of the most important
predators on wintering rats (Errington, 1932). On May 29, 1914,
a rat was found in the nest of a bank swallow, a considerable dis¬
tance from any building, on the Yahara River near Upper Mud
Lake (Betts, 1914; Schorger, 1937).
Spread of the Brown Rat
It is impossible to state definitely when the brown rat first ar¬
rived in North America. Harlan (1825) was informed by an eye
witness that the Norway rat did not appear in what is now the
United States until shortly before 1775. The commander of the
Spanish Fort Panmure (Rosalie), on the site of present Natchez,
Mississippi, complained bitterly in 1779 of the damage done to
grain by the enormous number of rats (Delavillebeuve, 1932). The
species is unknown. Michaux (1805) mentioned that the “grey
European rats” had not yet reached Cumberland (the western two-
thirds of Tennessee) but were common at the white settlements
elsewhere in the country. I do not know when rats arrived in
Chicago, but their presence in 1854 is graphically described by
Carl Schurz (Schafer, 1928) : “Chicago has 'wooden sidewalks’
under which live millions of rats. These rats regard the streets
at night as their domain, and in my presence made great use of
their freedom. Rats of all sizes and colors, old and young, white
and gray, played charmingly about my feet. And when I stepped
on one and it squeaked, it seemed to me as if I ought to beg pardon.”
The presence of the white phase is to be noted. The black rat is
not mentioned for Cook County by Kennicott (1855).
Strangely, Nuttall (1950) reported that there were no rats in or
around Detroit in 1810. The earliest references to rats in the Upper
Great Lakes region are to be found in the papers of the American
Fur Company. On April 16, 1827, Robert Stuart wrote from Macki¬
nac to J. J. Astor that he did not consider it advisable to keep furs
over winter on account of the destruction by rats. Damage by
rats is also mentioned in a letter of June 2, 1827.
Rats were present in the states of the upper Missouri far earlier
than Silver (1927) indicates. They infested the trading posts and
forts and were highly destructive to the stored corn. The earliest
reference found to rats at St. Louis is 1846. At that time Pancoast
(1930) was operating a boat on the Missouri and when docked at
St. Louis several rats went ashore. Rats must have been present
much earlier for the boats which plied the Missouri started from
St. Louis. When Luttig (1920) was ascending the Missouri in the
fall of 1812 he mentioned the abundance of mice on his boat but
does not mention rats. The first steamboat to ascend the Missouri
48 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
beyond Council Bluffs went to Fort Union in 1832. However, in
1833, Maximilian (1906, 3:13) found Norway rats abundant at
Fort Clark. At this fort, which was not built until 1831, Chardon
(1932), between August, 1834 and May, 1839, killed nearly 5000
rats by tally. In 1858 rats were not only a scourge to the fort but
also to the Indian villages (Boiler, 1868).
Norway rats were so numerous in 1833 at Fort Union in extreme
eastern Montana that the daily loss of corn was estimated at 250
pounds (Maximilian, 1906, 2:235). Oddly, Kurz (1937) wrote at
Fort Union in 1851 that “there are no rats in the fort any more
than at Fort Berthold.” The ambiguity may be due to translation.
Sackett (1866) who subsequently visited Fort Berthold found it
free of rats. He also inspected forts Randall, Sully and Rice where
rats swarmed. At Fort Rice 90,000 pounds of the 237,000 pounds
of stored corn had been destroyed, and the remainder so defiled
that only the Indians would eat it. The rats at Fort William,
located at the junction of the Laramie and the Platte, in 1837 had
cut to pieces the old epichimores (Russell, 1914). No large boats
could ascend the Platte.
The Black Rat
The black rat does not appear to have entered Wisconsin.
Lapham (1853) mentions it as having occurred at Racine, and
Kessinger (1888) as “very frequent’' in Buffalo County. These
were without doubt melanistic brown rats. Smith (1958) found
that 19 percent of the Norway rats in southeastern Georgia were
melanistic, and Rohe (1961) captured five in the black phase at
Pasadena, California. J. L. Diedrich of the Milwaukee Public
Museum has informed me that the Museum does not have a speci¬
men of the black rat; however they do have several melanistic
brown rats which were obtained in the late 1930’s at the old Wash¬
ington Park zoo site where there was a quite large population.
Arrival in Wisconsin
It is logical to assume that the brown rat arrived in eastern
Wisconsin from vessels on Lake Michigan and in the western part
of the state from boats on the Mississippi. The dates when rats
are mentioned for some of the counties in the state are shown in
Fig. 1.
Rats were probably at Prairie du Chien by 1823 for on Septem¬
ber 10 of that year R. Stuart wrote to Joseph Rolette at Prairie du
Chien that the deer skins shipped by the Upper Mississippi Outfit
were badly damaged by moths, rats, and dampness (Am. Fur Co.).
The headquarters of the Upper Mississippi Outfit were at St. An-
1971] Schorger — The Brown Rat in Early Wisconsin
49
thony Falls near St. Paul. According to Atwater (1893) rats and
mice were unknown at Minneapolis until about 1852. They certainly
were present much earlier. The invoices inward of the American
Fur Company show that of the “House Rat Traps” supplied in
1833, 18 traps remained with the Upper Mississippi Outfit and 7
at Prairie du Chien.
Grant County had rats at least by 1844. In June of this year
about 30 were killed in demolishing an old log stable (Lancaster,
1844). On April 5, 1847, 225 rats were killed by a Mr. Nordyke
and sons living near Lancaster (Lancaster, 1847). Jones (1857),
50 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
residing near Platteville, entered in his journal on January 29,
1857 : “I cleaned wheat and killed rats.” Two dogs killed 63 large
rats at Boscobel in June, 1873 (Boscobel, 1873).
The first rat ever seen at Taylor, Jackson County was killed in
September, 1878. It was supposed to have arrived in a freight car
(Merrillan, 1878). The first rat ever seen, supposedly, at Merrillan
was found on July 21, 1882 (Merrillan, 1882). A correspondent at
nearby Alma Center promptly wrote that “we count ours by such
numbers that we should not miss a small colony'5 (Merrillan
1882.1). After a rat was caught in 1880 in Black River Falls, it
was stated that this pest had yet to become a permanent resident
of the village (Black River Falls, 1880). The first rat for Bloomer,
Chippewa County, was caught November 9, 1899 (Bloomer, 1899).
Rats were seen daily in Galesville, Trempealeau County, in
August, 1873 (Galesville, 1873). Durand, Pepin County, was over¬
run by rats in 1878 (Durand, 1878) ; and in 1885 they were multi¬
plying rapidly at New Richmond, St. Croix County (New Rich¬
mond, 1885). In September, 1908, 42 rats were killed under one
stack of grain while threshing on a farm near Barron, Barron
County. They had been around only a year or two but were “mak¬
ing excellent progress’5 (Barron, 1906).
The brown rat arrived in central Wisconsin later apparently than
along Lake Michigan and the Mississippi. Jackson (1961:255) was
informed that it was present at Milton, Rock County, prior to the
arrival of the railroad in 1852. Chase (1858) spent the winter of
1839-40 in an old log cabin three miles from Watertown, Jefferson
County, in which were rats which he looked upon as better than
no company at all They lived under the floor. Welch (1881) under¬
stood that rats were unknown in the interior of Dane County until
the arrival of the railroad in 1854. By 1860 Madison (1860) had
rats in abundance and contests were scheduled to see which dog
could kill ten rats in the shortest time. An advertisement was run
offering ten cents apiece for rats. One contest did not materialize
as the rats escaped (Madison, 1860.1). The Capital House had
been nearly freed of rats by “Costar’s celebrated vermin extermina¬
tor55 (Madison, 1862). While at DeForest Stoner (1938) wrote in
his diary on August 27, 1862, that he was disturbed most of the
night by a large rat. The following night he caught one. In 1858
the first rat ever seen in Baraboo, Sauk County, was killed by boys
(Western Hist. Co. 1880), Rats were observed in Reedsburg in
1873 (Reedsburg, 1873).
While at Menasha, Winnebago County in 1851, Mackinnon
(1852) intended to dine with some Indians. The squaw put in the
kettle several species of squirrels and undressed fish, then “to cap
the climax, some rats were pulled from a heap of rubbish and
1971] Schorger — The Brown Rat in Early Wisconsin 51
actually added to the stew. My stomach began to mutiny, and I
was peremptorily compelled to run off.” At Neenah in 1857, rats
and mice were a great nuisance (Neenah, 1857). The Peters House
in Oshkosh was overrun by rats in 1869 (Oshkosh, 1869). It was
announced in April, 1872, that rats had at last arrived in Stevens
Point, Portage County (Stevens Point, 1872). They did not appear
in Colby, Marathon County until the spring of 1881 (Colby, 1881).
While at Keshena, Menominee County, in 1859, Father Gachet
wrote: “In order to make ourselves a little soup, the brother fer¬
reted out the chapel and produced some ends of old tallow candles
which the teeth of rats had spared.” In 1874 it was stated that it
was about seventeen years since rats had become “acclimated” to
Lake Superior (Ashland, 1874) ; however, when six rats were
killed in Ashland the following year, they were pronounced to be
the first ever found in the village (Ashland, 1875).
The first wheat raised near Racine was purchased by Charles
Wright the fall of 1840. While in storage a large amount of it was
destroyed by rats and mice (Western Hist. Co., 1879) . Subsequently
Lapham (1853) listed the brown rat as a resident of Racine. The
fall of 1839, H. Vail of the town of Vernon, Waukesha County,
stored his corn in the loft of the house of Almon Welch only to
have it purloined by rats, mice, and squirrels (Western Hist. Co.
1880.1). Between October, 1868 and the end of April, 1869, the
watchman at Rock Mills, Sheboygan, killed between 400 and 500
rats (Sheboygan, 1869). By 1859 rats had become an intolerable
nuisance at Manitowoc (Manitowoc, 1859). Mrs. A. B. Williams
came to DePere, Brown County in 1847. The following year she
moved into the Frontier House. It had been occupied so long by
“rats, bats, swallows and pigeons” that it took three men a week
to make it livable (French, 1876). Florence, Florence County,
could say in 1901 that there was neither a rat nor a mouse in the
city (Florence, 1901). On the night of October 4 of the following
year the first rat was killed at the pumping station (Florence,
1902).
Eradication
There is no hope of eliminating the brown rat. The most that
can be expected is to keep the population at a low level. Shortly
after the discovery by Karl P. Link of Warfarin as a rodenticide,
it was used in an eradication campaign in the village of Middleton
in the fall of 1950. Bait, consisting of a mixture of corn meal and
Warfarin, was widely distributed on November 4 (Madison, 1950).
Dr. Link has informed me that in the spring of 1951 it was diffi¬
cult to find a rat but it was not exterminated.
52 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
References
American Fur Company papers. Wis. Hist. Soc.
Ashland. 1874. Press Oct. 17.
• - . 1875. Press Sept. 18.
Atwater, I. 1893. History of the city of Minneapolis, Minnesota. New York,
I: 76.
Audubon, J. J. and J. Bachman. 1851. The quadrupeds of North America.
New York. II: 26.
Barrett— Hamilton, G. E. H. 1910. A history of British mammals. London..
pp. 608-609.
Barron. 1906. Shield Sept. 21.
Baumann, F. 1949. Die freileben Saugtiere der Schweiz. Bern. p. 215.
Betts, N. D. 1914. A rat in a swallow’s nest. Bird-Lore 16: 283.
Black River Falls. 1880. Banner March 12.
Bloomer. 1899. Advance Nov. 9.
Roller, H. A. 1868. Among the Indians. Eight years in the far west: 1858-
1866. Philadelphia, p. 33.
Boscobel. 1873. Dial; Madison State Journal June 5.
Chardon, F. A. 1932. Chardon’s Journal at Fort Clark, 1834-1839. Pierre-
458 pp.
Chase, Warren. 1858. The lifeline of the lone one. Boston, pp. 93, 94.
Colby. 1881. Enterprise May 25.
Delavillebeuvre, J. 1932. Fort Panmure, 1779. Miss. Vail. Hist. Rev. 18: 543-
Durand. 1878. Courier Sept. 28.
Errington, P. L. 1932. The food habits of southern Wisconsin raptors. Part I.
Owls. Condor 34: 181.
- . 1935. Wintering of field-living Norway rats in southcentral Wiscon¬
sin. Ecology 16: 122-23.
Florence. 1901. Mining News Nov. 2.
- . 1902. Mining News Oct. 11.
French, Bella. 1876. History of Brown County. Green Bay. p. 134.
Gachet, Father A. M. 1934. Journal, September, 1859, Keshena, Wis. Wis.
Mag. Hist. 18:204.
Galesville. 1873. Journal and Record Aug. 15.
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Harlan, R. 1825. Fauna americana. Philadelphia, p. 149.
Haymond. 1869. Mammals found at the present time in Franklin County. First
Ann. Rept. Geol. Surv. Indiana, p. 207.
Jackson, H. H. T. 1908. A preliminary list of Wisconsin mammals. Bull. Wis.
Nat. Hist. Soc. 6: 20.
- . 1961. Mammals of Wisconsin. Madison.
Jones, Orlando S. 1857. Diary. Jan. 29. Wis. Hist. Soc.
Kalm, P. 1772. Travels in North America. London, p. 348.
Kennicott, R. 1855. Catalogue of the animals observed in Cook County, Illi¬
nois. Trans. Ill. State Agr. Soc. for 1853-54. 1: 579.
Kes singer, L. 1848. History of Buffalo County, Wisconsin. Alma. p. 42.
Kurz, R. F. 1937. Journal of Rudolph Friedrich Kurz. Bur. Am. Ethn. Bull.
115: 187.
Lancaster. 1844. Herald June 8.
- . 1847. Herald April 10.
Lantz, D. E. 1909. The brown rat in the United States. Biol. Surv. Bull. 33 : 17.
Lapham, I. A. 1853. A systematic catalogue of the animals of Wisconsin.
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1971] Schorger—The Brown Rat in Early Wisconsin
53
Lawson, John. 1937. Lawson’s history of North Carolina (1709). Richmond,
p. 129.
Lescarbot, M. 1914. The history of New France. Toronto. 3: 226.
Luttig, J. C. 1920. Journal of a fur-trading expedition on the upper Missouri,
1312-1813. St. Louis, pp. 62-63.
Lyon, M. W. 1936. Mammals of Indiana. Notre Dame. p. 271.
Mackinnon, L. B. 1852. Atlantic and transatlantic sketches. London. I: 248.
Madison. 1860. Daily Argus and Democrat April 28, 30.
- — — . 1860.1. Wis. Daily Patriot May 26, 28.
— - — — — . 1862. Wis. Daily Patriot July 31,
- — . 1950. State Journal Nov. 3, 19.
Manitowoc. 1859. Tribune July 27.
Maximilian, Prince. 1906. Travels in the interior of North America, 1832-
1834. Cleveland. II: 235; III: 13
Merrillan. 1373. Leader Sept. 27.
— ■ — - 1882. Leader July 28,
- •• - . 1882.1. Leader Aug. 4
Michaux, F, A. 1805. Travels to the west of the Allegheny Mountains. Lon¬
don. p. 293.
Neena.Ii and Menasha. 1857. Conservator Sept. 17.
Nelson, E. W. 1917. The rat pest. Nat. Geogr. Mag. 32: 5.
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Botanica 14: 61.
Oshkosh. 1869. Journal Aug. 29.
Pancoast, C. E. 1930. A Quaker forty-niner. Philadelphia, p. 130.
Pennant, T. 1761-66. British zoology. London.
Reedsburg. 1873. Free Press Sept. 5.
Rhoads, S. N. 1903. The mammals of Pennsylvania and New Jersey. Philadel¬
phia. pp. 18-19.
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42: 268.
Russell, O. 1914. Journal of a trapper. Boise, p. 60.
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Sess., House Ex. Doc. 23: 24, 30, 38, 44.
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244.
Sheboygan. 1869. Herald; Madison State Journal April 27.
Silver, J. 1927. The introduction and spread of house rats _ in the United
States. Jour. Mam. 8: 58-60,
Smith, Capt. John. 1910. Travels and works. Edinburgh, pp. 154-55, 471.
Smith, W. W. 1958. Melanistic Rattus norvegicus in southwestern Georgia.
Jour. Mam. 39: 304-306.
Snyder, W. E. 1902. A list with brief notes, of the mammals of Dodge Co.,
Wise. Bull. Wis, Nat. Hist. Soc. 2: 117.
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Stoner, G. W. 1938. Diary. Wis, Mag. Hist. 21 : 424.
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— — — . 1880. The history of Sauk County, Wisconsin. Chicago, p. 553.
— — — . 1880.1. The history of Waukesha County, Wisconsin. Chicago, p. 476.
THE INFLUENCE OF DRUMLIN TOPOGRAPHY
ON FIELD PATTERNS IN DODGE COUNTY, WISCONSIN
Charles W. Collins
The influence exerted by drumlin topography on field patterns is
readily apparent to those persons who have the opportunity to fly
over the drumlin fields of southeastern Wisconsin. Marsehner, 1 in
his classic work on land use and patterns, observed that land use
in Dodge County, Wisconsin, is controlled by both the rectangular
land divisions and the orientation of the drumlins. It was observed
by this writer that the relationship between field patterns and
drumlins is not only complex, but also systematic. It is the purpose
of this paper to demonstrate that a combination of factors, includ¬
ing drumlin trends and height as well as the township and range
grid, are responsible for systematically varying sizes, shapes and
orientations of the field patterns in Dodge County.
Dodge County lies in the southeastern portion of Wisconsin, ap¬
proximately 30 miles northwest of Milwaukee. (See Figure 1.) The
county is nearly homogeneous insofar as farm type, farm size
and crop combinations are concerned. In general, the farms are
between 130 and 150 acres in size.2 Although the majority are
dairy operations with hay, corn and oats occupying a large percent
of the cropland, some specialized operations such as beef, chicken
and turkey ranches are occasionally encountered. Woodlands and
unimproved pasture generally are found on the steeper slopes.
Pasture, farmsteads, farm gardens and a few specialty crops, such
as green peas and sweet corn, account for most of the remaining
cropland. In several instances wetlands are found in the lowland
areas between drumlins. Occasionally these wetlands are used as
unimproved pasture, although more often they would fall into the
category of wasteland.
Soils vary considerably throughout the drumlin area with sandy
loams and silt loams commonly found on drumlin sides and tops.3
Peat and muck soils are commonly found in the depression between
drumlins even though actual swamp or marsh conditions may be
absent due to the implacement of drain tile.
Drumlins are present in approximately % of the country. These
elongated hills, with their major axes orientated in the direction
of the flow of the ice sheet that created them, vary in their axial
orientations from almost true north-south in south central Dodge
55
56 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Figure 1. Index Map.
County to northeast-southwest at the western border. The major
non-drumlin area is located in the north-central section, where
typical swell and swale topography is found. Horicon Marsh, one
of Wisconsin’s largest wetland areas, is also located in this north-
central area.
Method
In order to study in detail the relationships between drumlin
topography and field size, shape and orientation, a method by
1971] Collins— Influence of Drumlin Topography 57
which these phenomena could be measured and evaluated had to be
developed. Because the large number of fields presented a problem
in handling the data, four areas of nine square miles each were
selected as sample areas. These areas were chosen by visual exam¬
ination of the topographic maps of Dodge County. They were
selected because they represent four types of topography character¬
istic of the non-wetland areas of the county. (See Figure 2.) These
areas include the following types of topography :
Elba Area —High drumlins oriented northeast-southwest.
Lowell Area —Low drumlins oriented north-northeast south-
southwest.
Lebanon Area— High drumlins oriented nearly true north-south.
Trenton Area — Non-drumlin area with local relief generally less
than 40 feet.
Field patterns were studied through the use of large scale aerial
photographs (eight inches to the mile) provided by the Dodge
County Office of the Agricultural Stabilization and Conservation
Service. Maps of the field patterns, roads and farmsteads were
constructed by tracing these elements directly from the aerial pho¬
tographs. (See Figure 3.) As in a recent study by Hart dealing with
field patterns, changes in the tone or texture of the photos were
considered to be an indication of a boundary.4 Field checking in
the study areas corroborated Hart’s conclusion that this method is
indeed accurate. For the purposes of this study, farmsteads were
not considered to be fields, and minor drainage lines were not con¬
sidered to be field boundaries unless they coincided with a tone or
texture change on the photo. True wetlands were observed in only
a few instances. They were delineated, but for the purposes of the
study were not considered to be fields, Woodlots, which occupied
less than 10% of the study areas, were considered to be fields for
the purposes of this study. In the case of strip cropping, each strip
was treated as a separate field.
It was desirable, for the purposes of this study, to measure
the degree of topographic orientation in such a way so that a
statistical, yet visual, presentation could be made. A grid with
four lines to the inch was constructed and randomly placed over
the topographic maps of each square mile of the nine-square-mile
units. Each intersection of a contour line and the grid was then
noted and the bearing of the contour line was measured. In some
instances contour lines were observed to bend toward a grid line,
touching it, but not crossing. These cases were disregarded due to
the difficulty of obtaining an accurate angular measurement. A fre¬
quency diagram of the angles was then constructed. The angles
58 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Trenton Lebanon
Figure 2. Topography in the four study areas.
were grouped for convenience into ten-degree units. The results
were plotted on the compass roses shown in Figure 4.
In the analysis of field shape, all fields were considered. Each
was put into one of the following nine categories: squares, rec¬
tangles, parallelograms, trapezoids, clipped figures, triangles, com¬
plex polygons, trapeziums and irregulars. Of these nine categories,
1971]
Collins— Influence of Drumlin Topography
59
perhaps clipped figures, complex polygons, trapeziums and irregu¬
lars are in need of further explanation. The clipped figure is a
five-sided field which has two parallel sides which were, in turn,
generally parallel to drumlin trends. The other three sides were
usually coincident with some elements of the township and range
grid. Two of these three sides were generally parallel The complex
polygon category included fields having five or more sides, all of
60 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Figure 4. Topographic and Field Orientation. Lengths of the shafts are
proportional to the percent of orientation in that direction. Data are
grouped into ten-degree units. The dark portions of the semi-circles are
proportional to the percent of fields having no definite orientation.
1971] Collins— Influence of Dmmlin Topography 61
which were straight lines. Trapeziums included four-sided fields
where all sides were straight, but where no two sides were parallel.
Fields where one or more sides were not straight lines were placed
in the irregular category.
The fields to be used in determining field orientation and size
were selected with the use of a random dot sample. The size of each
sample field was measured with a planimeter. Some subjectivity
was involved in the measurement of field orientations since the
determination of orientation was entirely visual. This visual analy¬
sis was chosen because of difficulties encountered in attempting to
develop a non-sub jective method. The method employed herein in¬
volved the simple placing of a line parallel to the general direction
of field orientation as computed by the operator. Although this
method involves some degree of subjectivity, it should be noted that
in a test six geography graduate students at the University of
Wisconsin-Milwaukee agreed as to the orientation of test fields
within two degrees, ninety-eight percent of the time. Therefore,
although some subjectivity is certainly involved in this measure¬
ment it is the belief of the writer that it is minimal. Field orienta¬
tion data, like topographic orientation, was grouped into ten degree
units and plotted on compass roses. (See Figure 4.)
Field Patterns
Several interesting relationships come to light when topographic
orientation and field orientation data are compared. For instance, if
Figure 4 is examined, it is evident that the two roses representing
the nine square miles in Elba Township display a strong and nearly
identical northeast-southwest orientation. Thus, drumlins and field
orientation are nearly identical in the Elba Area. In the Lowell
Area, however, although drumlins have a similar orientation to
that of the Elba Area, the majority of the fields display a north-
south or east-west orientation. This leads one to believe that topo¬
graphic orientation alone will not produce the angular fields seen
in Elba, and that the lower relief of Lowell is most certainly a fac¬
tor. Note also that in the Trenton area, where low relief is coupled
with little or no general topographic orientation, most of the fields
are oriented with the township and range grid. Lebanon with its
nearly north-south drumlins, displays an exceptionally high degree
of north-south field orientation.
When field shape was compared to the topographic orientation
of the four study areas, it was discovered that Elba and Lowell,
where drumlins were oriented north-east to south-west, displayed
the greatest variety of field shapes. It was also in these areas where
the greatest number of fields with acute and obtuse angles were
62 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
observed. Field shapes, such as trapezoids, clipped figures, parallelo¬
grams and triangles were common. In Elba, for example, over 36%
of the fields are trapezoids. If we add to this, 7 % parallelograms,
3% clipped figures, and 8% triangles, we include 54% of the sample
fields. (See Figure 5.) Compare this to less than 10% of these four
shapes in Trenton where there is little topographic orientation,
12% in Lebanon and 24% in Lowell.
Elba
FrJ
mnpfm
Lowell
CD
£ g
3 P
■s. Z
s s JS 1 S !
C ii; ^ w JS
§c e « &.-=
C a 4- .2 r .=- 2
Z ru £ u > — &
- H — H r* u —
3
S£
O
Figure 5. Field shapes, percents.
1971] Collins— Influence of Drumlin Topography 68
In Lebanon, where high drumlins give a strong north-south
orientation to the topography, 56% of the fields were found to be
rectangular. Compare this figure to that of Elba with 6%, Lowell
with 34% and Trenton with 49% rectangles. Given these data
only, one might be led to believe that the Trenton and Lebanon
areas are similar. It is field orientation, however, that is the
important difference, since Lebanon displays a high degree of
north-south field orientation which is parallel to the drumlin
trends, while Trenton has as many fields oriented east-west as
north-south. A further difference in the nature of the rectangular
fields of these two areas can be seen when the field pattern maps
of sections in Trenton and Lebanon (Figure 3) are compared.
In Lebanon the closely spaced high north-south oriented drumlins
impose a restriction on field width, the ratio between the length
and width of fields is considerably greater than in Trenton.
In Elba the low figure of 6% rectangles is related to both the
high degree of topographic orientation in a direction not parallel
to elements of the township and range grid, and the relatively
high relief. The Lowell Area, with the intermediate figure of 34%
rectangles again emphasizes the importance of lower relief in
governing field shapes. Here, although topographic orientation is
strongly northeast-southwest, fields are preferentially oriented
north-south or east-west, resulting in the larger percentage of
rectangular fields for Lowell.
In the case of Lebanon, two other factors regarding field shapes
stand out. They are, the almost complete absence of square and
the relatively lower number of irregular fields. The latter can be
perhaps explained by the absence of streams and swamps in the
Lebanon area, since these hydrographic features are responsible
for many of the irregular field boundaries. The absence of streams
and swamps can be attributed in part to the implacement of drain
tiles in many of the lower areas between drumlins. Consequently,
straight line drainage ditches frequently serve as field boundaries
in the lower intra-drumlin areas. The near absence of square fields
can be related to the parallelism of the high north-south drumlins.
This parallelism seems to make the elongated rectangle so practical
for cultivation as to exclude squares.
In Hart's study of field patterns in Indiana,5 field shapes were
related to differences in farm types. This, however, does not seem
to be the case in Dodge County where the scale of the study is
larger and the study areas are much closer together. Furthermore,
field study revealed that the only significant difference in farm type
or farming practices throughout the four study areas was that strip
cropping was practiced more in those areas of higher relief.
64 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Field Size
Field size differs considerably throughout the four study areas.
The smallest fields were located in Lebanon where 8.33 acres was
the mean size of those fields sampled. The smaller field size in
Lebanon is undoubtedly partly attributable to the greater use of
strip cropping in this area of relatively higher relief. The largest
fields were found in Trenton with a mean of 15.0 acres. Here,
lower relief and the absence of topographic orientation have re¬
sulted in fewer restrictions on field size than in the other three
regions. An examination of the bar charts in Figure 6, further
demonstrates the field size differences between the study areas. For
example, in Lebanon 86.5% of the fields sampled were below 13
acres, which was the median field size in Trenton. This compares
to 65% below 13 acres in Lowell and 54.5% in Elba. Insofar as
large fields, Trenton and Elba both had approximately 20% of
their fields over 20 acres in size, while the Lowell area had 15%
over 20 acres, and the Lebanon area only 3% in this larger size
category. It is further interesting to note that the “forty-acre
field” is missing in all four of the study areas.
Conclusions
It is the opinion of the writer that this study bears out the
stated hypothesis that drumlin trends and heights as well as the
rectangular land division are largely responsible for the varying
field sizes, shapes and orientations in Dodge County. Four signifi¬
cant facts regarding drumlin and fields can be drawn from the
study. First, the presence of drumlins in portions of Dodge County
appears to be an important limiting factor with regard to field
size. Second, the relief in the drumlin areas is an important factor
in determining field orientations, in that the fields in the regions
of higher drumlins tend to show a greater degree of parallelism
to the topographic orientation. Third, more complex field shapes
are found in areas where drumlins cross elements of the rectangular
land division at acute and obtuse angles. Fourth, in the area where
high drumlins are parallel to elements of the rectangular land
division fields were found to have the highest degree of topographic
orientation and the smallest size.
Although the conclusions listed above only apply to the four
nine-square-mile study areas in Dodge County, Wisconsin, rela- ,
tionships uncovered in this study may apply to other drumlin areas.
Obviously this paper represents only a preliminary study and more
work is needed before the relationships between drumlins and
field patterns are completely understood. It is the hope of this
1971]
Collins— Influence of Drumlin Topography
65
Figure 6. Field sizes in the four study areas.
66 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
writer that more studies of held patterns in all types of topography
will be undertaken by geographers and other scholars so that we
may one day more fully understand the fields upon which our very
existence depends.
References Cited
1. Marschner, F. M., Land Use and Its Patterns in the United States, Agri¬
culture Handbook No. 153, United States Department of Agriculture,
Washington, 1959, reference on p. 150.
2. Finley, R. W., Geography of Wisconsin: A Content Outline, College Print¬
ing and Typing Co. Inc., Madison, 1965, reference on pp. 116-117.
3. Beatty, M. T., et al, 196U Wisconsin Blue Book, pp. 149-170, reference on
pp. 166-167.
4. Hakt, John Fraser, Field Patterns in Indiana, The Geographical Review,
Vol. LVIII, No. 3, July 1968, pp. 450-471, reference on p. 462.
5. Ibid, p. 471.
6. I would like to thank Professor Canute Vander Meer of the Department of
Geography, University of Wisconsin-Milwaukee for the encouragement
necessary to continue this study. Professors Dale Fatzinger and Lloyd
Flem of the Department of Geography at Wisconsin State University-
Platteville read the first draft and offered valuable comments. The final
paper remains, of course, my responsibility.
SEDIMENTOLOGICAL AND CHEMICAL PARAMETERS OF THE
LAKE SUPERIOR NERITIC ZONE, SOUTH SHORE, WISCONSIN
J. W. Horton R. C. Brown,2 D. W. Davidson ,3
A. B. Dickas ,4 W. Banking ,4 and R. K. Roubal1
Wisconsin State University , Superior
Introduction
In October, 1967, a group of faculty members at the Wisconsin
State University, Superior, began to analyze the opportunities
oceanography offered by their location on the shore of Lake Supe¬
rior. An area was selected for examination for the purpose of
establishing baseline studies. Parameters selected for study in¬
cluded; sediment sieving analysis, turbidity, pH, electrical con¬
ductivity, dissolved oxygen, and total dissolved solids. Development
of techniques and procedures was a primary objective of the pilot
study and, as anticipated, the exploratory traverses exposed several
errors.
While Lake Erie has been classified by investigators as either
“dead” or in a isemi-permanent state of contamination, Lake Supe¬
rior is usually classified as in a near-pristine condition. In an effort
to avoid such generalized terminology it is necessary to conduct
analyses of Lake Superior in a manner to establish bases from
which the degree of contamination may be determined, particularly
along the Wisconsin portion of the south shoreline where there is
definitely a problem of shoreline erosion (Red Clay Interagency
Report, 1967). With these baseline studies, studies of water quality
may then be compared in qualitative as well as quantitative terms.
These baseline studies are particularly important because there
are significant quantities of non-natural materials being discharged
into the lake at the present time. From these baseline studies it
is hoped that the study of the physical-chemical factors of the
environment will lead to studies of the producer-consumer dy¬
namics of the area discussed in the present paper. These baseline
studies are in addition to other baseline studies which have been
and are being conducted in the western Lake Superior region by
the National Water Quality Standards Laboratory (Duluth) and
by the University of Minnesota (Duluth and Minneapolis).
1 Department of Chemistry.
2 Department of Geography.
3 Department of Biology.
4 Department of Geology.
67
68 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
In October of 1967 an area was selected for the purpose of
establishing baseline studies of the Lake Superior neritic zone,
south shore, Wisconsin. This research work was the beginning of
studies which have culminated in the establishment of the Center
for Lake Superior Environmental Studies at Wisconsin State Uni¬
versity, Superior.
The area selected for study extends for two miles along the south
shoreline from the eastern end of Wisconsin Point immediately
east of the City of Superior in Section 35, Township 49 North,
Range 13 West (figure 1). The eastern end of the study area is
500 yards east of the mouth of Morrison Creek in Section 36,
Township 49 North, Range 13 West.
Factors considered in selection of the study area included accessi¬
bility by land for convenience in establishing triangulation stations,
proximity to the University, an open or unprotected coastal area,
and a variety of onshore topography ranging from marsh to wooded
and actively undercut clay banks. Two streams flow into the lake
within the study area, Dutchman Creek and Morrison Creek, with
their mouths periodically blocked with sediment.
Figure 1. Area of Study in Western Lake Superior.
1971] Horton et al. — Parameters , L. Superior Neritic Zone 69
Five data collecting stations were established at equal intervals
along the shore as indicated in figure 2. Triangulation points for
use by the boat party in positioning along the traverse lines were
located using the 1954 1:24,000 Superior Topographic Quadrangle
Sheet, Aerial Photographs, and Hydrographic Charts.
The five traverse lines were oriented at right angles to a line
parallel to the shoreline at the appropriate station. Standard
plane table triangulation methods were utilized to map the location
of the offshore stations along the traverse lines. The traverse lines
were spaced at half-mile intervals and extended offshore to a water
depth of thirty feet, the operational limit of the equipment.
Four data collecting stations were located along each traverse
line: at the shore, and where the depth of the bottom was ten,
twenty, and thirty feet. Samples were taken along the traverse
lines at water depth intervals of ten feet, from surface to bottom,
to a maximum depth of thirty feet. A set of ten samples of water
and four sediment samples were obtained per traverse. Maximum
distance offshore at the thirty foot depth was 0.8 miles while
minimum distance was 0.4 miles.
70 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Methods
A seventeen foot boat with ten horsepower motor was available.
The boat crew maintained position along the traverse lines by
visual alignment of markers previously erected on shore and
oriented using a Brunton Pocket Transit. The positions of the
sample collecting stations were then plotted on the base maps.
Water samples were taken at the surface, at depths of ten foot
intervals, and at the bottom. For example, at the thirty-foot depth
station along each traverse, one bottom sediment sample and two
sets of water samples were collected. Separate samples were ob¬
tained for dissolved oxygen analysis. Each suite of water samples
thus consisted of four aliquots collected; from the surface, at a
depth of ten feet, twenty feet, and at the bottom.
The size distribution of sands along the South Shore of Lake
Superior was examined. Significant qualitative changes in the
lithology were noted. Sediment samples were obtained at each
traverse station with a Peterson Dredge. These bottom samples
were oven dried and 100 gram aliquots were retrieved from a
Jones Sediment Splitter. This sample was passed through a nest
of ten U. S. Standard Sieves and Ro-Tapped for a period of five
minutes. The various size fractions were regarded as percentages
of the total weights of each aliquot.
A grain-size frequency distribution of material from each sample
station was then plotted as a comparative curve of cumulative
frequency. From these curves the significant parameters of median
diameters, coefficient of sorting, skewness and kurtosis (the latter
two representing the first and second moments of dispersion) were
calculated.
Water turbidity was measured at each offshore station by use
of a Secchi Disk. All observations were made under conditions of
direct sunlight and within a three-week period in October 1967.
Sunlight was of approximately the same intensity during all
observations.
The pH of the water samples was obtained by means of a Beck¬
man Model G pH Meter with Sargent No. S-30072-15 combination
electrode. Freas type conductivity cells with cell constants of
approximately 0.3 reciprocal ohms were used in conjunction with
a Model RC Conductivity Bridge from Industrial Instruments
Incorporated operating at 1000 Hz to determine the specific
conductance.
Dissolved oxygen in parts per million was determined by the
Iodometric Method, not using the azide modification (Standard
Methods, 1965). Values for total dissolved solids were determined
by evaporation of a 25.00 milliliter water sample at 105° C. in a
platinum crucible.
1971] Horton et al. — Parameters, L. Superior Neritic Zone 71
Results
The sediment groups identified by median size analysis are
derived from three sources. The first is continental clastic glacial
till of Wisconsin age (minimum age of 10,000 ± 2,000 years)
which forms the surface stratigraphic unit throughout much of
the Great Lakes Region. The second source consists of the rather
extensive Keweenawan Sandstones (age greater than 500 million
years) found adjacent to the shoreline in the Apostle Island area.
The third source is the Valders Red Clay which outcrops along the
South Shore of Lake Superior from the Duluth-Superior area east¬
ward to the Apostle Islands. Loy (1962) reported that while this
material appears to be pure clay, sieve analysis reveals a consistency
of approximately five-percent boulder to sand-sized particles. Val¬
ders Clay was deposited as a result of sedimentation in glacial
Lake Duluth and is of post-Wisconsin age.
Median grain size of the lake sediments ranged from a nega¬
tive 2.70 </> (small pebbles) to a positive 3.05 <f> (very fine sand)
value (figure 4.). Along the typical traverse, medium-sized sand
particles were found high on the beach and in the littoral zone.
Fine sand particles were recorded at the ten foot and twenty foot
depths and very fine sand particles were dredged at the thirty
foot depth.
Using Trask statistics (Trask, 1932), the quartile deviation
(coefficient of sorting) varied from 1.51 </> to 0.69 <f> indicating
well sorted elastics with all calculations based on phi scale (-log2)
(see figure 3). The skewness (symmetry) ranged from a negative
2.21 to a positive 1.09, while the kurtosis (peakedness) ranged
from a negative 0.42 to a negative 0.2.
On this basis bottom contours were mapped and the areal dis¬
tribution pattern of the median diameter size grades of the lake
bottom sediment indicated (see figure 4). Translated into the
Wentworth Grade Scale (1922) lake bottom material in the study
area ranged from granules to very fine sand.
This wide range in particle size reflects the active erosion
presently existing in the area studied. As figure 4 indicates, the
coarse sediment is located adjacent to the shoreline and sediment
size decreases with an increase in distance offshore. The division
between fine and very fine sand occurs approximately one-half
mile offshore.
Isolated pockets of pebbles are found occasionally along the
South Shoreline. These pockets are residuals resulting from the
reworking of banks of Valders Red Clay which as indicated above
contains a small percentage, by weight, of boulders. Large volumes
of this clay are eroded and reworked during periods of lake storms
72 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
LAKE SUPERIOR PILOT OCEANOGRAPHIC PROJECT
AREAL DISTRIBUTION OF SORTING
□ TRAVERSE POSITION
© SAMPLING STATIONS
AtRIANGULATION POINTS
.0.9 _ CALCULATIONS BASED ON PHI SCALE C-L062 )
Figure 3. Areal Distribution of Sedimentary Particles (calculations based on
Phi Scale, -Logo).
and frost heaving. After the fine clay is transported offshore by
lake currents the boulders are concentrated in pockets and slowly
transported parallel to the shoreline by longshore currents.
Dispersion of a fraction of these gravels occurs by a process i
described by Dickas and Lunking (1968). Blocks of clay eroded
from the banks and carried into the lake are reworked into tri-
axial ellipsoids. The tri-axial “mudball” acquires a layer of pebble
armour. The armoured mudballs are then carried by the current
and cast upon the beach where the armour plate of gravel and
sand spalls off as the hydrous clay mud-ball core desiccates. The
result is a small residue of pebbles on an otherwise finely-sorted |
beach sand. It should be emphasized that these small pebble residu¬
als are not the same as those of much greater volume caused by
longshore current sorting along the beach.
Anticipated exceptions to the offshore median diameter particle
size decrease occur adjacent to the mouth of Morrison Creek (see
figure 4) . At this location a submerged sand bar trends 55 degrees
west of north. The highest portion of this bar is composed of
granules and pebbles and is centered approximately 1800 feet
1971] Horton et dl.—. Parameters, L. Superior Neritic Zone 73
Figure 4. Areal Distribution of Median Diameter Size Grades of Sands
Particles.
offshore. The shallowing water over the bar permits westerly-
moving longshore currents to winnow out the finer clastic elements
leaving the coarse sediment. This winnowing activity is also related
to the increase in turbidity noted in the vicinity of the bar (fig¬
ure 5).
The orientation of this bar cannot, at this time, be attributed
to any particular type of lake currents. A more complete under¬
standing of surface current activity requires a thorough study
of the variable wind patterns in the immediate vicinity of the
Lake Superior study area.
The average depth of visual extinction of the Secchi Disk at
all stations was 2.95 meters (8' 3") and ranged from a minimum
value of 1.67 meters (5' 6") to a maximum of 3.81 meters (12' 6")
(figure 5). These data indicate that wave action and longshore
currents hold enough sediment in suspension to decrease the water
transparency below that determined in mid-lake where Loy (1962)
reported values of 14 meters. Figure 5 indicates an apparent
increase of water transparency to the northwest which may be
74 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
due to the lessening influence of the sand bar which is crossed
by traverses one and two.
pH ranged from a low value of 7.49 to a high of 8.40 (see table 1) .
The average value was 7.89. These appear to be in agreement with
the Silver Bay study from which results of 7.8 pH units were
reported (Swain and Prokopovich, 1957.) The higher values as
compared with samples obtained at Silver Bay, Minnesota, may
be due to increased leaching of alkaline earths from the red clays
along the South Shore of Lake Superior. Average pH unit values
for all sample stations by depth were 7.91 at the surface, 7.88 at
ten feet, 7.88 at twenty feet, and 7.85 at thirty feet.
Specific conductance (k x 106 ohm -1 Cm _1 at 25 °C.) had
values ranging from 96.5 to 126.3 (see table 2). The average was
102.3. Homogeneity was within reasonable limits and indicated that
samples from depths to thirty feet were within the active mixing
zone. Averaging values by depth were 105.1 at the surface, 100.9 at
ten feet, 100.9 at twenty feet, and 98.5 at thirty feet.
Dissolved oxygen values ranged from an average of 10.8 parts
per million at the surface layer to 10.5 parts per million at the
thirty foot level. Thus, there was no significant variation in parts
Figure 5. Water Transparency by Secchi Disk (values in meters from surface).
1971] Horton et al.— Parameters , L. Superior Neritic Zone 75
Table 1. pH of Lake Water.
xThe number of the sample station is the maximum depth at the station. The
letter ‘a’ is the surface sample; ‘b’ is at 10' depth; ‘c’ is 20' depth; etc.
per million from one depth to another and apparently the samples
were all obtained from the mixing zono. However, the slightly
higher surface readings in parts per million of dissolved oxygen
concurs with the findings of Putnam and Olson (1960), that parts
per million of dissolved oxygen and oxygen saturation (%) were
both higher at the surface. With one exception shore surface read¬
ings were higher than from offshore samples.
Putnam and Olson's observations further indicate that during
a sampling period at Knife River Station (L-16) , Lake Superior
on September 16, 1959, the parts per million of dissolved oxygen
were approximately comparable with results obtained in the present
study ranging from sampling station averages of 10.6 to 11.3.
Table 2. Specific Conductance of Lake Water.
(Values in K x 106 ohm-1 cm-1)
76 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Putnam and Olson also found oxygen saturation for September
10, 1959, to be near or greater than 100% for the zone covered by
the present study.
Beeton (1966) found the dissolved oxygen content of Lake
Superior, except in certain bays, to be near saturation at all depths.
It is therefore presumed that dissolved oxygen is near or greater
than 100% saturation in the study area.
Total solids determined by evaporation of a 25.00 milliliter water
sample at approximately 105° C. in a platinum crucible averaged
71.5 parts per million.
Conclusions
Data indicates a standard distribution of the Lake Superior
elastics within the area studied which is not unusual considering
the high energy conditions of the lake shoreward of the six-fathom
depth. The few anomalous zones recorded are attributed to small
lake floor surface pockets of clay being transported to lower depths.
Derived from gravity slumping along the South Shoreline, such
clays were initially deposited in the area as lacustrine sediments
during the Wisconsin sub-epoch in Lake Duluth, the ancestral
glacial high-watermark of Lake Superior.
Generally the specific conductance average value decreases
slightly from shoreline to the thirty foot area and also decreases
from surface to bottom. The specific conductance values are ap¬
proximately equivalent to the specific conductance of a 7 x 10~4
M KC1 solution.
pH unit values obtained were quite uniform except for a few
anomalies which may have some significance. The highest and
lowest values occur at the shoreline which may be indicative of
leaching. An average of pH unit values at each sample station
indicates that the area is quite homogeneous.
Dissolved oxygen values were nearly constant with average
values ranging from 10.5 to 10.8 parts per million. There was
no significant variation in parts per million from one depth to
another and apparently the samples were all obtained from the
mixing zone. Highest values were obtained from shoreline stations
which are the sites of greatest water agitation.
The average of random samples for total dissolved solids was
determined to be 71.5 parts per million.
Ultimate conclusions are difficult to assess. It was not possible
to locate the mixing zone utilizing the data obtained. While much
more information should be gathered, the data collected are as¬
sumed to be normal considering the near-pristine condition of Lake
Superior.
1971] Horton et at,—- Parameters , L. Superior Neritic Zone 77
The project has served as an initiation into the field of water
study for an interdisciplinary group and has established a base
line for further investigation.
The findings presented apply only to the study area and are not
intended to indicate an interpretation of the entire dynamic proc¬
esses operating in Lake Superior. Since, however, the study was
along and adjacent to open lake shoreline, it is expected that the
findings may validly be employed in a comparative study of other
unprotected shoreline sectors of Lake Superior.
Acknowledgments
The authors are grateful to the late Mr. Edward Moyer, who was
dedicated to the protection and conservation of Lake Superior, and
who provided the boat and motor for use in this study.
Financial support was supplied by a Research Grant to the
authors, from the Wisconsin State University-Superior, Research
Committee.
Literature Cited
Beeton, A. M. 1966. Indices of Great Lakes Eutrophication. Great Lakes Re¬
search Division Publ. No. 15, The University of Michigan, Ann Arbor,
pp. 1-8.
Dickas, A. and W. Lunking. 1968. The Origin and Destruction of Armoured
Mud Balls in a Fresh-water Lacustrine Environment. Journal of Sedi¬
mentary Petrology. Dec. pp. 1366-1370.
Loy, W. 1962. The Coastal Geomorphology of Western Lake Superior. M.S.
Thesis. Department of Geography, University of Chicago.
Putnam, H. and T. Olson. 1960. An Investigation of Nutrients in Western
Lake Superior, School of Public Health, University of Minnesota, Minne¬
apolis.
Red Clay Interagency Report. 1967. Prepared by Wisconsin Red Clay Inter¬
agency Committee.
Standard Methods for the Examination of Water and Waste Water. 1965.
American Public Health Association. 12th Edition. New York.
Swain, F. and Prokopovich. 1957. Stratigraphy of Upper Part of Sediments
of Silver Bay Area, Lake Superior. Geological Society of America. Bulle¬
tin No. 68, pp. 527-542.
Trask, P. 1932. Origin and Environment of Source Sediments in Petroleum.
Gulf Publishing Company, Houston, pp. 71-72.
Wentworth, C. A. 1922. Scale of Grade and Class Terms for Clastic Sedi¬
ments. Journal of Geology, Vol. 30, pp. 377-392.
VEGETATIONAL PATTERNS AND ORDINATION
IN CEDARBURG BOG, WISCONSIN
Thomas Foster Grittinger 1
Introduction
Cedarburg Bog, located only 25 miles north of Milwaukee, is
one of the largest bog areas in southern Wisconsin. This bog,
occupying the basin of a postglacial lake, lies in sections 20, 21,
28, 29, 31, 32, and 33, in the township of Saukville (TUN, R21E)
in Ozaukee County (Figure 1). An aerial photograph of this 2,000
acre bog reveals numerous patterns throughout its surface (Fig¬
ure 2), which reflect differences in the vegetation. The peripheral
portions of the bog generally appear dark in tone and of coarse
texture, whereas around the margins of the lakes the tone is lighter
and the texture finer. Large areas near the center of the bog appear
to be composed of alternating dark and light bands. Other parts
are sharply distinct from nearby areas, often giving a rectilinear
effect, and still other regions have less well defined patterns. Since
preliminary observations on the ground confirmed these patterns,
and the only previous research in Cedarburg Bog concerned plant
species and associations encountered (Cutler, 1935) and water
level changes (Cutler, 1936), further study was indicated.
The first objective concerned a description of the vegetation
types which form these patterns and the mapping of the major
cover types present within the bog. The second objective involved
the quantitative analysis of the vegetation and the communities
present by utilizing an ordination of these communities within
the bog.
Methods
Pattern Analysis and Mapping of Major Cover Types
To map vegetational patterns within Cedarburg Bog, several
aerial observations were made during the summer and fall of 1966.
1 Contribution No. 2 from The University of Wisconsin-Milwaukee Field Station.
This paper represents a portion of a thesis submitted in partial fulfillment of the Doc¬
tor of Philosophy Degree, University of Wisconsin-Milwaukee, 1969, and it represents
a portion of a paper presented at the 99th annual meeting' of the Wisconsin Academy
of Sciences, Arts and Letters. The author wishes to express appreciation to P. B.
Whitford- for advice and counsel throughout the course of this work, to M. D. Heinsel-
man for positive identification of the string bog, to P. J. Salamun and A. L. Throne
for taxonomic assistance, to G. W. Argus for identification of willows, to W. H. Ellis
for identification of red and silver maple hybrids, and to J. H. Zimmerman for identi¬
fication of sedges and grasses. Financial aid was provided by a National Science
Foundation Summer Fellowship for Graduate Teaching Assistants, a University of
Wisconsin-Milwaukee Graduate Fellowship in Botany and The University of Wiscon¬
sin, University Center System. The author is Assistant Professor of Botany and Zool¬
ogy at the University of Wisconsin, Sheboygan County Campus.
79
Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
scale in feet
I - - 1
1 667
Figure 1. Section numbers and lakes of Cedarburg Bog.
1971] Gritting er—V eg etational Patterns in Cedar burg Bog 81
Figure 2.
Also, numerous ground reconnaissance trips were carried out dur¬
ing the winters of 1966-67 and 1967-68. References were constantly
made to aerial photographs and to topographic maps to corroborate
these observations.
Under stereoscopic examination, two aerial photographs taken
from slightly different angles reveal both the topography and
vegetational characteristics (Losee, 1942). The characteristics of
the vegetation, other than height, were analyzed by examination
of photographic tone, texture, shadow, and the shape and size of
crowns as suggested by Spurr and Brown (1946).
Vegetational Sampling
After the bog area had been mapped, the general vegetation
types were studied quantitatively, using the transect method for
82 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
cover values and the quarter method for trees and saplings. Sam¬
pling lines were laid out independently of the aerial map. Thus
they served to check on the accuracy of the mapping and to amplify
and refine the description of the vegetation types.
Cover values for trees of all sizes, shrubs, woody vines, and
some of the more conspicuous herbaceous material, e.g. the cattails
and sedges, were measured by the line intercept method (Can-
field, 1941; Buell and Cantlon, 1950). Such methods are useful
where differences in the vegetation are evident and are to be com¬
pared with one or several factors that change between two points
(Costing, 1956). The transect method has been used to study
vegetation and some environmental factors by Bauer (1936),
Penfound and Hathaway (1938), and Pierce (1953).
East-west lines (lines 2, 3, 5, and 6) were placed every half
mile to insure thorough coverage of the bog (Figure 3). A north-
south line near Mud Lake (line 1) was added to supplement
coverage in this area and terminated at the south shore of the
lake. The short east-west line (line 4) was used to gain additional
data since it was most accessible and had water level sampling
devices for other work (Grittinger, 1969) ; this line ended at the
creek north of Mud Lake. All of these lines were laid out by
compass.
The six lines were divided into 100 foot segments, resulting
in a total of 236 segments. Cover in each segment was measured
by placing a tape on the ground and estimating the vertical
projection of the canopy. Since the segment was 100 feet long, the
total intercept of any given species could easily be expressed in
per cent cover. This was converted to relative cover by dividing
the total intercept of a species within the 100 feet by the total
intercept of all species and multiplying by 100.
Relative density, relative frequency, and relative dominance
of trees and saplings were determined by the quarter method. The
merits of the quarter method are discussed by Cottam and Curtis
(1956). This method had been employed by Curtis (1950) based
on a system of recording “witness trees” used by the Federal Land
Survey in the last century (Cottam, 1949; Stearns, 1949). A point
was placed at the center of every 100 foot segment.
Ordination and Compositional Index
Ordination has been defined as the “arrangement of data of
a continuous and non-discrete sort into an orderly spatial pattern”
(Curtis, 1959). This method, unlike classification techniques, relies
less on subjective judgement for placing vegetational types in a
successional pattern. While the general classification and mapping
permits an introduction to the problem, the ordination of data
1971] Grittinger — Vegetational Patterns in Cedarburg Bog 83
Figure 3. Line intercepts.
84 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
will, it is hoped, permit a more thorough and objective study of
the variation within Cedarburg Bog.
To use an ordination technique, it was necessary to establish
adaptation numbers for the woody and the major herbaceous
species in the bog. Curtis and McIntosh (1951) used a similar
method with upland forest trees. An index of joint occurrence
was used in ranking all species according to their ecological re¬
sponses. This index, used by Swindale and Curtis (1957), was
calculated by finding the number of quadrats in which two species
occur together as a percentage of the number of quadrats of occur¬
rence of the less common species of this pair. In the present study,
100 foot segments of the line intercepts were used rather than
quadrats. The species were then placed in an order based on
similarity of behavior (Guinochet, 1955). As in Swindale and
Curtis (1957), the species with high indices of joint occurrence
are placed close together and those with low indices are placed far
apart in the order.
To facilitate ordination, additional information gathered from
field experience and the literature was used. This involved the
knowledge that some species have fairly predictable ecological
requirements, i.e. a given species will be more important on the
mesic end rather than the wet end of a continuum ranging from
emergent aquatics to a mesic situation, or vice versa. Reference
species were chosen as follows: white birch ( Betula papyrifera
Marsh.)* was arbitrarily used to represent a more mesic condition
than bog birch (Betula pumila L. var. glandulifera Regel) or
the emergent aquatic sedge (Car ex spp.) mat, and the emergent
sedges were used to represent the most wet situation since
they were abundant next to the edge of the water. The white
birch is typical of the dense forest nearer the outer edges
of the bog. Bog birch lies midway between the two extremes.
All other species were compared to these three on the basis
of indices of joint occurrence. For the sake of clarity, three
different lines were used with one of these three species as a
reference point on each line. All of the other species were placed
along these lines in an order determined by their joint occurrence
indices. For example, one line had white birch as a reference point,
another had bog birch, and the third had emergent sedges. Each
species was compared to each of these three ispecies, and then
placed an appropriate distance from the reference point. A linear
scale was computed by subtracting the index from one and multi¬
plying by 20, an arbitrary number. The results were then measured
out in centimeters along the line. Thus two species with an index
The nomenclature used here is according- to Fernald (1950).
1971] Grittinger — Vegetational Patterns in Cedar burg Bog 85
of 0 (never occurring together) would be placed 20 centimeters
apart, and two species with an index of 1 (always together) would
be placed on the same spot on the line.
A scale was devised, similar to that of Curtis and McIntosh
(1951), but with emergent aquatics at one end and the mesic
upland hardwoods at the other. A scale from 1 to 10 was used
with the low value to represent those species most likely to be
found at the edge of the water with the emergents and 10 to
represent those species likely to be present in an upland forest. In
addition to the three original species, yellow birch ( Betula lutea
Michx.) , cedar ( Thuja occidentals L.) , tamarack (Larix laricina
(DuRoi) K. Koch), and a flat, open sedge mat area referred to as
the Rhynchospora-Carex mat (dominated by Rynchospora alba
(L.) Vahl and Carex lasiocarpa Ehrh. var. americana Fern.)
were found to lie in a similar sequence on all three of the ordination
lines. An adaptation number was then assigned to each of the five
species and two associations based on their distance from each
other and from the ends of the 20 centimeter lines. The emergent
aquatic mat was given a value of 1.0, the Rhynchospora-Carex
mat 2.0, bog birch 3.5, tamarack 5.0, cedar 6.0, white birch 7.0,
and yellow birch 8.0. Sugar maple ( Acer saccharum Marsh.),
an infrequent species in the bog, was assigned a value of 10.0.
The rest of the species were then assigned adaptation numbers
on the basis of comparisons with the species and groups designated
above. For example, winterberry (Ilex verticillata (L.) Gray)
has a joint occurrence of 0.062 with respect to the emergents,
0.825 compared to Rhynchospora-Carex, 0.698 compared to bog
birch, 0.875 compared to tamarack, 0.830 compared to cedar,
0.610 compared to white birch, and 0.470 compared to yellow
birch. Thus, the winterberry was given an adaptation number
which would place it between tamarack and cedar, i.e. 5.5. In a few
cases the indices were high over a wide scale, and an adaptation
number was assigned on the basis of field experience and literature.
A compositional index for each segment was calculated by
summing the products of the adaptation number and the relative
cover value for each species in the segment. For example, a 100
foot segment with 60% cedar, 30% tamarack, and 10% dog¬
wood would have these relative cover values multiplied by 6.0,
5.0, and 5.0 respectively to produce a compositional index after
summation of 560. The possible range of these index numbers is
from 100 for a segment of pure cattail or emergent sedges to 1,000
for a segment of pure sugar maple. Thus each segment with its
accompanying point had a value that was determined by the rela¬
tive cover and the ecological response of the individual species
found along that particular segment. According to Curtis (1959),
86 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
this method of summing1 weighted values, which combines a meas¬
ured value of plant quantity with an estimate of plant response,
has been used by a number of ecologists (Whittaker, 1952; Ellen-
berg, 1952) and has been discussed by Whittaker (1954).
The ecological response of the individual species was analyzed
by plotting the relative cover of a single species against the
ordination after arranging all segments by numerical order of the
compositional index. These figures were made by plotting the
moving averages of the averages of the relative cover within each
80 units of the ordination. The moving average was calculated by
the formula:
B _ nia + 2n2b + n3c
ni + 2n2 + n3
where B is the smoothed average of b, and m, n2, and n3 are the
number of segments included in the averages a, b, and c respec¬
tively. This formula has been used in upland ordination by Brown
and Curtis (1952). This procedure was carried out for many of
the woody species and some of the more important herbaceous
species.
The data from the relative cover analysis of each 100 foot line
segment were combined for each 100 unit interval of the compo¬
sitional index scale, so that all of the 100 foot line segments falling
within each 100 unit interval were used to obtain the mean relative
cover values for each species of tree, shrub, and some of the
herbs. One hundred unit intervals were arbitrarily used for
convenience.
The tree and sapling data obtained by the quarter method were
computed on the basis of the same 100 unit intervals of the compo¬
sitional index.
Comparison of Ordination with Mapped Patterns
Each transect segment was plotted on an overlay of an aerial
photograph by its compositional index. All of the segments with
compositional index values ranging from 100 to 200 were plotted
with one color, those with values from 200 to 300 were plotted
with another color, and so on until all of the segments were rep¬
resented on the overlay. These plotted segments were then compared
to the vegetational patterns and cover types. Since an extremely
large aerial photograph and overlay (40" by 40") would be re¬
quired to adequately show the mapped indices, this could not be
included. However, all of the segments within each pattern given
on the vegetational pattern map were summed and the mean and
range of compositional index values for each type were calculated.
1971] Gritting er—V eg etational Patterns in Cedarburg Bog 87
Results
Pattern Analysis and Mapping of Major Cover Types
On the basis of the aerial photographs and ground recon¬
naissance, eight major vegetational cover types were recognized
within Cedarburg Bog:
1. Emergent aquatics
2 . String bog or Strangmoor
3. Bog birch-leatherleaf shrub area
4. Dogwood-willow shrub area
5. Dead tamarack area
6. Conifer forest
7. Conifer-swamp hardwood forest.
8. Upland hardwood forest
These zones outlined on the map (Figure 4) are necessarily
somewhat arbitrary since cover types blend one into another
throughout the bog. In addition, within any larger area there
are frequent cover types too small to be shown on a map of the
scale given here (1 inch = 1667 feet). Where there is an obvious
and intimate mixing of cover types over a given area two or more
numbers are used to indicate the nature of the mixture, e.g. 4,6
indicates a dogwood-willow shrub and conifer forest mixture.
Vegetational Sampling, Ordination, and Compositional Index
The adaptation numbers assigned to the woody and the major
herbaceous species of the bog are listed in Table 1.
The compositional index numbers for the 236 segments of the
six transect lines in Cedarburg Bog range from 99.9 to 799.9.
There are 6 segments in the 100-200 range, 16 in the 200-300
range, 28 in the 300-400 range, 53 in the 400-500 range, 117 in
the 500-600 range, 13 in the 600-700 range, and only 3 in the
700-800 range, indicating that the major part of the bog is in
shrub or early swamp forest stages of succession.
The ecological responses of 13 of the species are illustrated on
Figures 5-7, and some of the more important ones are discussed.
The results of the relative cover analysis are included on Tables
2-4 and those of the mean importance values for trees and saplings
are found on Tables 5 and 6.
Comparison of Ordination with Mapped Patterns
The correspondence between the areas mapped within the bog
and the compositional index values are listed in Table 7. For each
cover type there is a mean compositional index value and a range
of values.
88 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Figure 4. Vegetational patterns of Cedarburg Bog.
1971] Gritting er—V eg etational Patterns in Cedarhurg Bog 89
Table 1. Adaptation Numbers of Species in Cedarburg Bog.
Carex aquatilis Wahlenb. . . . . . . . 1.0’
Sagittaria sp . 1.0
Scirpus validus Vahl var. creber Fern . 1.0'
Typha spp . . . 1.0
Andromeda glaucophylla Link . . 2.0
Phragmites communis Trin. var. Berlandieri (Fourn.) Fern . . . 2.0
Rhynchospora alba (L.) Vahl . 2.0
Sarracenia purpurea L. . . . . . 2.5
Betula pumila L. var. glandulifera Regel . . . 3.5
Chamaedaphne calyculata (L.) Moench var. angustifolia (Ait.) Rehd . 3.5
Salix pedicellaris Pursh . 3.5
Salix Candida Fliigge . . 4.0'
Salix gracilis Anderss . 4.0
Salix serissima (Bailey) Fern . 4.0
Cornus racemosa Lam. . . . . . 4.5
Gaylussacia baccata (Wang.) K. Koch . 4.5
Rosa palustris Marsh . 4.5
Salix discolor Muhl . . . 4.5
Spiraea alba Du Roi . . . 4.5
Alnus rugosa (Du Roi) Spreng. var. americana (Regel) Fern . 5.0
Amelanchier spp . 5.0'
Cornus obliqua Raf . 5.0
Cornus stolonifera Michx . 5.0
Larix laricina (Du Roi) K. Koch . 5.0
Lonicera dioica L . 5.0
Lonicera oblongifolia (Goldie) Hook . 5.0
Lonicera villosa (Michx.) R. & S . 5.0
Nemopanthus mucronata (L.) Trel . 5.0
Pyrus melanocarpa (Michx.) Willd . 5.0
Rhus Vernix L . 5.0
Vaccinium myrtilloides Michx . 5.0
Viburnum trilobum Marsh . 5.0'
Ilex verticillata (L.) Gray. . . 5.5
Juniperus communis L. var. depressa Pursh . 5.5
Lonicera canadensis Bartr . 5.5
Parthenocissus sp . 5.5
Picea mariana (Mill.) BSP . 5.5
Rhamnus alnifolia L’Her . 5.5
Ribes sp . 5.5
Salix Bebbiana Sarg . . . 5.5
Solanum Dulcamara L . 5.5
Viburnum Lentago L . 5.5
Vitis riparia Michx . 5.5
Populus tremuloides Michx . 6.0
Rhamnus cathartica L . 6.0
Rhamnus Frangula L . 6.0
Rubus sp . 6.0
Thuja occidentalis L. ...... . . 6.0
Acer rubrum L . 7.0
Acer rubrum L. x A. saccharinum L . 7.0
Betula papyrifera Marsh . 7.0
Fraxinus nigra Marsh . 7.0
Fraxinus pennsylvanica Marsh, var. subintegerrima (Vahl) Fern . 7.0
Ulmus americana L . 7.0
Betula lutea Michx . 8.0
Fraxinus americana L . 8.0
Taxus canadensis Marsh. . . 8.5
Prunus virginiana L . 9.0
Tilia americana L. . . . . 9.0
Acer saccharum Marsh. . . 10.0
Ostrya virginiana (Mill.) K. Koch. . 10.0
Viburnum acerifolium L . 10.0
90 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Compositional Index
Figure 5. Behavior of the major bog trees along compositional gradient. 1,
tamarack; 2, cedar; 3, white birch; 4, black ash; 5, elm.
Compositional Index
Figure 6. Behavior of the major bog shrubs along compositional gradient. 1,
leatherleaf ; 2, bog birch; 3, red osier dogwood; 4, poison sumac; 5, winterberry.
1971] Gritting er — -V eg etational Patterns in Cedarburg Bog 91
__i i - _ _ i :T , —.1 _ _ _ j —
200 300 liOO $00 600 700
Compositional Index
Figure 7. Behavior of selected bog herbs along compositional gradient. 1, cat¬
tail; 2, reed grass; 3, pitcher-plant.
Table 2. Relative Cover Means of Trees in 100 Unit Compositional
Index Intervals.
*Subtotal includes trees with less than 1.0% relative cover in any interval:
Acer rubrurn
Acer rubrurn x Acer saccharinum
Fraxinus pennsylvanica var. subinteger rima
Populus tremuloides
92 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Table 3. Relative Cover Means of Shrubs in 100 Unit Compositional
Index Intervals.
*Subtotal includes shrubs with less
Amelanchier spp.
Andromeda glaucophylla
Cornus racemosa
Gaylussacia baccata
Lonicera canadensis
Lonicera dioica
Lonicera oblongifolia
Lonicera villosa
Nemopanthus mucronata
Parthenocissus sp.
Pyrus melanocarpa
Rhamnus alnifolia
Rhamnus cathartica
1.0% relative cover in any interval:
Ribes sp.
Rosa palustris
Salix babylonica
Salix Candida
Salix humilis
Salix pedicellaris
Sambucus canadensis
Spiraea alba
Vaccinium myrtilloides
Viburnum Lentago
Viburnum trilobum
Vitis riparia
than
1971] Gritting er- — V eg etational Patterns in Cedarburg Bog 93
Table 4. Relative Cover Means of Selected Herbs in 100 Unit
Compositional Index Intervals.
*Subtotal includes herbs with less than 1.0% relative cover in any interval:
Hypericum sp.
Table 5. Mean Importance Values of Trees by Quarter Method for
100 Unit Compositional Index Intervals.
Species with importance values less than 5 in any interval:
Acer rubrum x Acer saccharinum
Populus Iremuloides
Salix Bebbiana
*It should be noted that the Importance Value is based on relative values among
trees only so that, where tree cover is less than 10% of the total cover, importance
value has little meaning.
94 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Table 6. Mean Importance Values of Saplings by Quarter Method for
100 Unit Compositional Index Intervals.
Species with importance values less than 5 in any interval:
Acer rubrum x Acer saccharinum
Cornus obliqua
Fraxirxus pennsylvanica var. subintegerrima
Ilex verticillata
Populus tremuloides
*It should be noted that the Importance Value is based on relative values among
saplings only so that, where sapling cover is less than 10% of the total cover, impor¬
tance value has little meaning.
1971] Gritting er—V eg etational Patterns in Cedarhurg Bog 95
Table 7. Correspondence Between the Mapped Areas Within the Bog and
the Compositional Index Values,
Discussion
Pattern Analysis and Mapping of Major Cover Types
The patterns and zonation of the vegetation types of Cedarburg
Bog form a mosaic (Figure 2), Some of these types, such as the
upland hardwoods, dead tamarack area, string bog, and, to some
extent, bog birch-leatherleaf shrub zone and emergent aquatic
zone are usually distinct from each other. Others, such as the
conifer forest, the conifer-swamp hardwood forest, and the dog-
wood-willow shrub areas, are often indistinct, blending into one
another. This blending seems greatest where disturbance such as
lumbering or wind-throw of trees has occurred. Clearly the classi¬
fication of vegetation by general types, while useful in mapping the
general vegetational cover, is not sufficiently discriminating.
Although sequential, concentric zones representing different stages
of successional development are present throughout much of the
bog, i.e. the northeast shore of Mud Lake, the mosaic effect
demonstrated by aerial photograph suggests more complex relation¬
ships.
The first zone to be considered on the vegetation map is that
of the emergent aquatics (Figure 4, Number 1). It is finely
textured and very light in tone on the aerial photograph (Figure 2) ,
This zone is found at the margins of the lakes, especially northern
and eastern Mud Lake, northern and southern Long Lake, and
northern Donut Lake. It can be found along the several creeks
between the lakes as well. It lies between the open water and the
successional shrub zone (Number 3). The emergent aquatic zone
96 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
is composed mostly of bulrushes (Scirpus validus Vahl var. creber
Fern.), sedges (chiefly Car ex aquatilis Wahlenb.), and cattails
( Typha spp.), while locally water willow ( Decodon verticillatus
(L.) Ell.) and arrowhead ( Sagittaria sp.) may dominate at the
water’s edge. The appearance of this zone is very open — -almost
marsh like. This is referred to as the emergent aquatic mat or
association.
The center portion of the bog (Figure 4, Number 2) is distinct
from all other zones in that it appears as dark gray, more or less
parallel bands on a finer, lighter background on aerial photographs.
There are larger islands of dark gray vegetation, similar to the
narrower bands, scattered in this zone too. Ground studies, obser¬
vations from an airplane, and stereoscopic analysis reveal this to
be string bog or “Strangmoor” composed of ridges and islands of
tamarack and cedar, usually very stunted on the smaller ridges,
lying on an open, flat sedge mat. When seen from the ground or
at lower elevations, this area has the appearance of open meadows
separated from one another by ridges or hedgerows of conifers.
Sometimes the ridges are small with little woody vegetation. The
larger, darker islands are stands of fairly dense conifer forest.
This part of the bog is of special interest since it is the southern¬
most location of “patterned organic terrain” of the northern type.
The previous known limit for such patterns is about 200 miles
northward, near Seney, Upper Michigan (Heinselman, 1965). Dr.
M. L. Heinselman states that the areas “west and southwest of
Long Lake clearly constitute a 'string bog’ or 'strangmoor’ complex”
(personal communication). His identification was made on the
basis of an examination of an aerial photograph and a description
of the area.
The Rhynchospora-Carex mat (called “flarks” by Swedish
workers) forms the relatively flat, open mat; the area is domi¬
nated by Rhynchospora alba (L.) Vahl, a ishort, fine sedge, with
lesser amounts of Carex lasiocarpa Ehrh. var. americana Fern.,
pitcher-plants ( Sarracenia purpurea L.), bogbean ( Menyanthes
trifoliata L. var. minor Raf.) , water horsetail ( Equisetum fluviatile
L.) , bladderwort ( Utricularia spp.), arrowgrass ( Triglochin
maritima L.), dense clones of reed ( Phragmites communis Trin.
var. Berlandieri (Fourn.) Fern.), sundew {Bros era linearis
Goldie and D. rotundifolia L.), some orchids ( Pogonia ophioglos-
soides (L.) Ker. and Calopogon pulchellus (Salisb.) R. Br.) , and
some cattails ( Typha latifolia L. and T. angustifolia L.) . This
mat is very soft and treacherous to tread upon, and though it
appears to float, seems to lack some of the resiliency of the emergent
aquatic mat.
1971] Gritting er—V eg etational Patterns in Cedar burg Bog 97
The ridges of the string bog are elevated from a few inches to a
foot or more above the surrounding mat, and range in width from
a foot or so to twenty or more feet. The ridges may be more than
one hundred feet long before attenuating into the mat or fusing
with another ridge, thus forming a net like structure. Within each
ridge or island there is a miniature ecotone, grading from low
Rhynchosporcu-Carex mat to bog birch, leatherleaf ( Chamaedaphne
calyculata (L.) Moench var. angustifolia (Ait.) Rehd.) , and some
bog-rosemary ( Andromeda glaucophylla Link) , along with Sphag¬
num spp., cranberry ( Vaccinium Oxycoccos L.) , and sundew
( Drosera rotundifolia L.) , to cedar and tamarack with numerous
shrubs beneath. The shrubs are especially frequent on the edges of
the larger ridges and islands and in openings within ; included here
are bog birch, leatherleaf, poison sumac ( Rhus Vernix L.) , red
osier dogwood (Cornus stolonifera Michx.) , black chokeberry
( Pyrus melanocarpa (Michx.) Willd.) , juniper ( Juniperus com¬
munis L. var. depressa Pursh), winterberry, and dwarf alder
(Rhamnus alnifolia L’Her.). Beneath the dense cedar and tamarack
of the larger ridges and islands, velvet-leaf bilberry ( Vaccinium
myrtilloides Michx.) and winterberry may be found.
The bog birch-leatherleaf or successional shrub zone (Figure 4,
Number 3) appears quite fine in texture and a medium gray in
tone on the aerial photograph. This zone is found near lakes,
streams, and the edges of the low islands within the lakes. It lies
shoreward of the emergent aquatics. Near the edge of Mud Lake
it is up to several hundred feet wide and ranges from almost pure
bog birch to a bog birch-tamarack mixture. A dense cover of
leatherleaf is present on some of the low islands within the lakes
where stunted tamaracks are found.
The dogwood-willow shrub areas are very numerous in the bog
(Figure 4, Number 4). Their appearance on the aerial photographs
is very similar to that of the previous zone, but ground observations
reveal some differences. Near the west shore of Mud Lake it has
the characteristics of a shrub carr, with a dense, almost pure and
impenetrable stand of slender willow ( Salix gracilis Anderss.) . In
other areas red osier dogwood and several species of willow are
important with lesser amounts of winterberry. Another form of this
cover type occupies large areas near Mud Lake, where there are
many dead or dying elm ( TJlmus americana L.) , black ash ( Fraxi -
nus nigra Marsh.), and tamarack. Here red osier dogwood and
cattail are abundant. These shrub areas are often mixed with other
types, especially where the trees were cut or wind thrown.
The zone of dead tamaracks (Figure 4, Number 5) occupies
principally two disjunct, but very similar areas. On aerial photo¬
graphs it is similar to the previous two types, and again can be
98 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
best distinguished by its location within the bog and by ground
observations. This zone is characterized by dead tamaracks, many
of which are still standing, though free of bark and most branches.
It is an open area with some young cedar, large amounts of bog j
birch, some red osier dogwood, and lesser amounts of poison sumac,
willows, winterberry, and juniper. Old stumps are found scattered
here and there.
The sixth cover type, that of the conifers (Figure 4, Number 6),
has a coarse texture and a mottled gray tone. Throughout most
of the bog this cover type is composed of both cedar and tamarack,
but white birch is often found near the peripheral portions of
this cover type. Near the bog birch-leatherleaf shrub zone almost
pure stands of tamarack may occur and between Mud Lake and the
east side of the bog black spruce ( Picea mariana (Mill.) BSP.)
occurs. In still other locations, small thickets of cedar may exclude
almost all other species.
The conifer-swamp hardwood forest (Figure 4, Number 7) is i
more heterogeneous than the conifer forest due to the increased
number of hardwood trees. This forest type tends to be found on
the peripheral portions of the bog, especially the northwestern and
western parts of the bog, and around the southern parts of Mud
Lake. The latter example has been subjected to disturbance, how¬
ever. This forest is composed of cedar, tamarack, black ash, elm,
white birch, with lesser amounts of red maple (Acer rub rum L.),
and red-silver maple hybrids (A rubrum L. x A. saccharinum L.).
The latter hybrid was corroborated by Dr. W. H. Ellis of Austin
Peay State University. This cover type, as represented by the >
conifer and white birch mixture blends freely with the pure conifer
forest. There are areas near the edge of the bog, e.g. south of Mud
Lake, where there is a preponderance of swamp hardwoods. How¬
ever, since these areas are relatively small and blend freely with
the conifer-swamp hardwood mixture, they are included in the
conifer-swamp hardwood forest.
The most mesic cover type in the bog is the upland hardwood
forest (Figure 4, Number 8). On the aerial photographs, the vege¬
tation appears to be of a very coarse texture, mottled gray in tone,
and composed of large flat tree crowns. This type of cover occupies
the upland mineral soils of the islands within the bog. Stereoscopic
examination indicated that these tree tops are at a higher elevation
than any others within the bog. One island is found in Mud Lake,
three are north of Mud Lake, and another is surrounded by bog
northwest of Long Lake. Upland hardwood forest also covers the
high ridge that is near the western edge of the bog. These upland
areas appear to be formed of glacial debris, mostly limestone boul¬
ders and gravel and as such are elevated above the surface of the
1971] Gritting er — Vegetational Patterns in Cedarburg Bog 99
surrounding peats. The upland hardwood forest is composed mainly
of sugar maple, white ash ( Fraxinus americana L.) , basswood
(Tilia americana L.), ironwood (Ostrya virginiana (Mill.) (K.
Koch), and other mesic species.
Vegetational Sampling, Ordination, and Compositional Index
The ecological responses of members of a plant community have
been widely applied for ordination in Wisconsin. Adaptation num¬
bers (Curtis and McIntosh, 1951) have been used for forest trees
(Curtis and McIntosh, 1951; Brown and Curtis, 1952; Ware, 1955;
and Christensen, Clausen, and Curtis, 1959). In these cases actual
joint occurrences of species served as a guide in setting up a Classi¬
fication system based on ecological responses of trees (Curtis,
1959). They all found sugar maple to be the most mesic species,
with other tree species at the opposite extremes of the adaptational
series (Curtis, 1959). In all cases a scale of adaptation numbers
was set up ranging from 1 to 10. The most mesic species, sugar
maple, was assigned a value of 10.0, and the least mesic in either
wet or dry sites, a value of 1.0.
Bray (1955) assigned adaptation numbers to southern Wisconsin
savanna trees on the basis of mutual occurrences. According to
Curtis (1959), the results gave a compositional gradient based on
moisture rather than moisture and light as was the case with the
forest trees.
Vegetation other than trees has been similarly treated in order
to set up a classification based on ecological response. Curtis (1955)
evolved five indicator groups for Wisconsin prairie species based
primarily on the moisture gradient. The scale of numbers ran from
1 for the wet indicators to 3 for the mesic and finally on to 5 for
dry indicators.
In this study, adaptation numbers (Table 1) based on an index
of joint occurrence were obtained for all of the woody and some
of the important herbaceous species present. A compositional index
for trees alone was inadequate since many areas in the bog are tree¬
less and, in other areas, the relatively uniform tree composition
precluded an ordination based on joint occurrence of trees.
The series of adaptation numbers represents a range in vegeta¬
tion from cattail to sugar maple. The higher the number of any
given species, the greater the likelihood that that species will be
found on mesic areas, dominated by sugar maple. As might be
expected for a wet area with little topographic variation, many
species encountered in Cedarburg Bog have adaptation numbers
below the middle of the scale. The species with higher numbers
are found mostly on upland areas; sugar maple and ironwood do
not grow in the bog proper.
100 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
The adaptation numbers assigned in this bog continuum may
differ considerably from those in the upland continuum for several
reasons. First, the scale utilized here is wider than that of the
upland ordination since it ranges from emergent aquatics to mesic
upland hardwoods rather than only from pioneer trees to mesic
upland hardwoods. Second, a given species may exhibit a somewhat
different response in a bog as compared to the uplands.
The string bog area presents a unique problem in ordination.
Since the string bog is composed of two distinct cover types, i.e.
islands and ridges with conifers and shrubs on a matrix of Rhyn-
chospora and Carex, the data from the line intercepts represents
a “hybrid” cover type. Since 100 foot segments in the string bog
include both cover types, the Rhynchospora-C arex mat will have a
very high joint occurrence index with any of these woody species.
For example, the joint occurrence index value between the Rhyn-
chosporar-Carex mat and tamarack or bog birch is 1.00, and be¬
tween the mat and cedar it is 0.925. Thus, it is difficult to sub¬
stantiate the assignment of an adaptation number of 2.0 to
Rhynchospora alba on one hand, and 3.5 to bog birch, 5.0 to tama¬
rack, and 6.0 to cedar on the other, on the basis of the above com¬
parisons. Of course when tamarack is compared to cedar or to bog
birch, a more meaningful relationship is seen : 0.842 and 0.862 are
the respective joint occurrence indices. Although this suggests that
the Rhynchosporcu-C arex mat does not fit into the ordination, the
desirability of including this extensive cover type on a standard
ordination precluded its elimination, while the sampling method
using 100 foot segments precluded treating the ridges and open
mat separately.
The ecological response curves for most of the species (Figures
5-7) show an intergrading similar to that found in upland forests
by Curtis and McIntosh (1951). No species is distributed entirely j
at random. Each species is present in more than one section of
the continuum but showed a tendency to a normal curve of relative
cover values, with a peak in one section of the continuum. This :
would be expected from the general knowledge of species’ responses
to environmental factors. It is unfortunate that there are not
enough samples of segments with higher values present to enable
a complete study including examples from the 800 to 1000 range, i
but the paucity of mineral islands in the bog and their abrupt
nature, precluding a gradual transition from island to bog, made
a complete study impossible.
White birch proved relatively mesic, reaching a maximum cover i
value in segments between 600 and 800 (Figure 5 and Table 2).
White birch was found in situations ranging from conifer-swamp i
hardwood forest to the lower slopes of the mineral soil islands
1971] Gritting er — Vegetational Patterns in Cedarburg Bog 101
where it appeared to reach maximum size. Mixed with cedar and
tamarack in the bog forest,, it often occurs in areas where stumps
show evidence of cutting. This pattern is corroborated by Curtis
(1959), who mentions that this species is often found as a gap-
phase tree in small openings in the forest, and shows intermediate
shade tolerance.
Elm and black ash have high relative cover values in the seg¬
ments from 600 to 700 (Figure 5 and Table 2). Both species were
once abundant near Mud Lake, but now only a few isolated trees
survive. Although elm (Eyre and Zillgitt, 1953) and ash (Curtis,
1959) appear able to sprout from the bases of nearly dead trees,
the black ash seems to recover better after flooding. This is prob¬
ably due to the additional effect of the Dutch elm disease on the
elm population. The increase in relative cover for black ash in the
segments near 200 is due to the survival of a few isolated trees on
hummocks near Mud Lake.
Cedar has a high relative cover value over a wide range on the
compositional index scale (Figure 5 and Table 2), with highest
values in the 500 to 700 range. There is a wide range in size and
growth-form in the bog, with stunted and gnarled trees in the
string bog and larger trees on slopes of the mineral soil islands.
Curtis (1959) reported cedar stumps over 4 feet in diameter on
an island in Cedarburg Bog. The only areas that do not contain
this species are the emergent aquatic, some of the dogwood-willow,
and the bog birch-leatherleaf areas near the lakes. Cedar is con¬
sidered tolerant by Baker (1949) and Zon and Graves (1911), but
only intermediate by Curtis (1959). It is sometimes seen in almost
pure thickets in the conifer forest area, as a result of layering
following wind throwing of this shallow rooted species. This means
of reproduction is common in swamps (Curtis, 1959; Curtis, 1944;
and Rudolf, 1949). Cedar has been considered as a climax species
in boggy areas (Gates, 1942). Clausen (1957) found cedar to be
important in the wet-mesic conifer swamps, where its optimum
presence is attained (Curtis, 1959).
Tamarack, though less mesic than cedar, also has a wide ecologi¬
cal response (Figure 5 and Table 2), ranging far and wide in the
bog. Unlike cedar, it sometimes occurs very near the emergent
aquatic areas. It is more frequent than cedar in the bog birch-
leatherleaf shrub, and less frequent than cedar in the more mesic
conifer-swamp hardwood forest. According to Conway (1949),
Cooper (1913), and Gates (1942), tamarack is usually the first
forest tree to invade the hydrosphere in bogs. As an intolerant tree
(Curtis, 1959), it is replaced by black spruce, cedar, balsam fir,
and swamp hardwoods, in about that order, on better drained less
acid swamps (Soc. of American Foresters, 1954). Tamarack is a
102 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
major species in the wet conifer forests of Wisconsin (Clausen,
1957),, and attains its optimum presence in this forest type (Curtis,
1959).
Winterberry has quite mesic requirements (Figure 6 and Table
3). It frequently forms small, dense thickets in the conifer forest
areas; it is especially common on the conifer islands and larger
ridges in the string bog where it may be found beneath cedar and
tamarack.
Red osier dogwood and poison sumac are typical of disturbed
areas in the bog (Figure 6 and Table 3). They often occur in the
dogwood-willow areas in the bog (Figure 3, Number 4). Red osier
dogwood can assume high relative cover values in the segments in
the 400 to 600 interval, but is really very common elsewhere, too.
It is probably the most frequent woody species in this bog. It may
be found in the smallest openings, but develops best in large, open
areas such as under dead elm and black ash near Mud Lake. Poison
sumac has almost a bimodal curve since it is frequent in small
open but otherwise more mesic areas resulting from disturbance
as well as on some ridges and islands in the string bog.
Bog birch and leatherleaf both reach maximum relative cover
values in the 300 to 400 interval of the compositional index scale
(Figure 6 and Table 3) . Gates (1942) places bog birch in the “High
Bog Shrub Association” which succeeds the “Chamaedaphne Asso¬
ciation.” On the other hand, Conway (1949) considers both species
part of the “Moss-heath Association.” In Cedarburg Bog, both
species often occur together, showing similar ecological require¬
ments. As mentioned previously, these species, especially the bog
birch, are dominant over large portions of the bog as the bog
birch-leatherleaf shrub areas (Figure 3, Number 3), Leatherleaf
and the more common bog birch appear in an apparent successional
sequence between the emergent aquatics and the tamaracks, in the
string bog areas, and in the dead tamarack areas.
Pitcher-plant is common in the Rhynchospora-Carex portions of
the string bog (Figure 7 and Table 4). However, it is also found
on the ridges under a light canopy of cedar and tamarack; its
greatest cover is in the 200 to 300 range.
Reed grass finds optimum ecological conditions in the low-indexed
segments in the string bog (Figure 7 and Table 4). Here it is
usually found in the open, very wet flarks of the Rhy ncho sporar-
Carex mat, where it may form dense clones.
Cattail is typically a member of the emergent aquatic commu¬
nity, but is often found in wet depressions caused by wind-thrown
trees in the 300 to 700 range (Figure 7 and Table 4). Most of this
cattail is Typha latifolia L., though some T. angustifolia L. is
present.
1971] Grittinger — -V eg etational Patterns in Cedarburg Bog 108
The ecological responses of other bog species are graphed and
discussed elsewhere (Grittinger, 1969).
The mean importance values of the trees and saplings generally
parallel the results seen with the relative cover treatment, i.e. those
species with low adaptation numbers usually have high importance
values in the lower compositional index intervals and those species
with high adaptation numbers have high importance values in the
higher intervals (Tables 5 and 6). Some exceptions should be
noted, however. In the tree class (Table 5), black ash attains the
importance value of 300 in the compositional index intervals of
100 to 200. In the interval of 200 to 300, tamarack and cedar
have importance values of 134.5 and 165.5 respectively. Likewise
in the sapling class (Table 6), black ash is important in the 100
to 200 range (201.6) as are cedar and elm in the 200 to 300 range
(140.5 and 12.3 respectively). In all of these cases the small sample
is probably a factor, as there are only 6 segments in the 100 to 200
range, and 16 in the 200 to 300 range. Furthermore, the impor¬
tance value is based on relative values among tree species only,
so that in relatively treeless areas, where shrubs and herbs make
up most of the cover, the importance values of the few scattered
trees have little meaning. In the 100 to 200 range the sum of the
relative cover of the trees is only 1.2 and in the 200 to 300 range,
it is 8.9. In such situations one accidental tree on a hummock among
cattails would give that species an importance value of 300 if no
other tree species were present, even though its contribution to the
total cover is insignificant.
Comparison of Ordination With Mapped Patterns
The plotted segments matched the original patterns on the aerial
photographs. There is a progressive increase in mean composi¬
tional index, with a few slight exceptions, in going from the mapped
emergent aquatic areas to the upland hardwood forests. The areas
where large amounts of dogwood-willow are found mixed with
other cover types also fit into this progression. For example, the
mixture of dogwood-willow and conifer forest give a mean that is
higher than the “pure” dogwood-willow but lower than “pure”
conifer forest.
The uniformity with which the compositional indices of the tran¬
sect segments correspond to their expected localities on the aerial
photograph seems to indicate that the adaptation numbers assigned
to the plant species are valid for such ordination.
The range of compositional indices within a given pattern type
is indicative of several factors. First, as in the case of the conifer
forest with a wide range of indices, it is probably due to the mix¬
ture of stunted conifer forest within the string bog at one extreme
104 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
and the existence of dense stands of tall cedar and tamarack with
a few scattered white birch trees at the other. Another situation of
note is the wide range of the mixture (4, 6) of dogwood-willow
shrub and conifer-swamp hardwood forest. Here the segments with
lower compositional indices are found in disturbed areas, especially
near Mud Lake, and in some cases approach being a shrub-carr or
even an aquatic emergent area, while the segments with higher
compositional indices are represented by conifer-swamp hardwood
forest and shrubs near the edges of the bog.
Conclusions
Cedarburg Bog is composed of a mosaic of vegetational patterns
that may be arbitrarily classified and mapped into eight major
cover types based on aerial and ground studies. The major cover
types are : emergent aquatic, string bog or strangmoor, bog birch-
leatherleaf shrub, dogwood-willow shrub, dead tamarack, conifer
forest, conifer-swamp hardwood forest, and upland hardwood for¬
est areas. The string bog is an area of particular interest since it
represents the southernmost reported example of this type of pat¬
terned terrain. In addition to the major cover types, mixtures of
some of these types exist, especially where disturbance occurred.
Adaptation numbers were assigned to the plant species on the
basis of an index of joint occurrence, and these numbers were used
to weight relative cover values to obtain a compositional index for
each 100 foot segment of the line intercepts. The compositional in¬
dices strongly correspond to the vegetation patterns or cover types
suggesting that the adaptation numbers assigned are valid for this
ordination. The ordination of the communities within the bog was
related to the ecological responses of the individual species as shown
by the graphed curves of relative cover on the compositional index
scale. Of the eight major cover types,, only the string bog does not
lend itself to this ordination.
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THE LITTORAL MACROPHYTE VEGETATION OF LAKE WINGRA
An example of a Myriophyllum spicotum invasion in a southern Wisconsin lake.
Stanley A. Nichols and Scott Mori
Abstract
Over the past century the aquatic vegetation of Lake Wingra,
Dane Co., Wisconsin has changed from a vegetation type which
was probably dominated by Vallisneria americana and Potamogeton
spp. to one which is 68% Myriophyllum spicatum . M. spicatum is
a European species that appears to be a very aggressive invader.
The value of M. spicatum to the aquatic plant community is low.
The time or mechanism of the invasion could not be precisely
determined.
To provide a basis for assessing the role of M. spicatum in the
present community, and to provide a basis for continued research
on the lake, the submerged aquatic plants were sampled by the line
intercept method. Forty lines, approximately perpendicular to the
shoreline and extending to the depth at which growth of submerged
aquatic plants ceased, were sampled. All plants intercepting the
line were recorded within consecutive 5 m segments of the line.
Depth data, shore vegetation and soils data were taken relative to
the transect lines. For descriptive and mapping purposes the vege¬
tation was divided into five communities. The data are summarized
in map and tabular form.
Introduction
Over the past century a striking change has occurred in the
vegetation of Lake Wingra. Up to this time no quantitative work
has been done on the aquatic plants of the lake. The purpose of
this study was to assess the change in the vegetation during the
past hundred years, and to describe and map the present vegetation
for current research and future vegetational studies.
In the 1870’s Lake Wingra, or Dead Lake as it was known then,
was shallower and larger. Rowley (in Sachse, 1965) described the
lake as marshy on all sides with a good bit of wild rice (Zizania
aquatica) and wild celery (Vallisneria americana). “The shores
of the lake were shallow and one had to push a boat through a
hundred yards or more of weeds and cattails before reaching open
water.”
107
108 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
A hydrographic survey in 1904 revealed a maximum depth of
14 feet. Beginning in 1905 and continuing through 1920 consider¬
able filling and dredging of the lake took place. Today the majority
of the shoreline is controlled by the University of Wisconsin
Arboretum.
The shore vegetation of the lake today is considerably different
than the broad Typha marshes with Z. aquatica as described by
Rowley. A more striking change occurred in the submerged vege¬
tation, with the disappearance of V. americana and the almost
complete dominance by Myriophyllum spicatum.
Description of the Area
Lake Wingra is a natural, shallow basin on an outwash terrace
overlying a feeder stream of the preglacial Yahara River. The Lake
lies in central Dane Co. (T-7-N, R-9-E), within the city limits of
Madison. Poff and Threinen (1962) report the surface as 139.6 ha
and the maximum depth as 6.4 m.
Two small streams and numerous springs are the chief source
of water supply. The outlet, at the northeastern corner of the lake,
drains into Lake Monona via Murphy’s Creek. A dam on Murphy’s
Creek maintains a 0.6 m head of water.
!
Methods
Field Methods
Lake Wingra was sampled using the line intercept method as
described by Lind and Cottam (1969). A plastic coated rope,
marked in five meter intervals, was used as the intercept line.
The intercept lines (Fig. 1) ran from shore toward the center of
the lake at approximately 100 m intervals. Because of dense vegeta¬
tion and unsure bottom it was found easiest to run the lines with
a canoe. On the first trip the rope was taken out and anchored to
a float in deep water. The line was run a second time with one
paddler calling off the plant species intercepting the line, one
paddler taking depth measurements, and a third investigator acting
as a recorder. By using different symbols on the data sheet it was ,
noted whether the plant species was present (less than one plant
per meter), scattered (non-continuous) , or continuous (a solid, .
unbroken stand of plants) in each 5 m segment of line. Depth was
recorded in 0.5 m intervals until the end of the vegetation. A sound¬
ing rod was used for depths up to 2.5 m and a weighted line after
this depth. For mapping purposes the azimuth of each line was
taken with a Brunton Compass.
In the case that a community obviously ended between intercept
lines, the community was sketched. Notes referring to its position
1971] Nichols and Mori— Vegetation of Lake Wingra
109
by distance and azimuth from the enclosing intercept lines were
taken.
The soil was examined 5 m from shore at the beginning of each
transect. Shore vegetation for each transect was noted.
From the line intercept data, five macrophyte communities were
recognized. Using SCUBA apparatus, 20, one meter square quadrats
were harvested from each community. The number of stems of
each species in each quadrat was tallied for density data. In every
fourth quadrat the species were dried and weighed to give an esti¬
mate of the standing crop.
Two areas along the north shore of the lake were not considered
in this survey. An area between transects 31 and 32 and transects
39 and 40 were deleted because of a great amount of disturbance
to the plant community due to mechanical harvesting and chemical
poisoning with Aquathol. The ponds and lagoons entering the lake
were also disregarded.
Plant species were collected during the summers of 1968 and
1969. Voucher specimens are on file at the University of Wisconsin
Herbarium.
Computations
Each five meter interval of every transect was used as a basic
sampling unit (BSU). The percent occurrence of each community
type and the frequency of each species was calculated on the basis
of 872 BSU containing vegetation.
110 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
The outer limit of the littoral zone was calculated as the mean
± 2<r of the end point depth of the 40 transects. The open area in
the littoral zone was computed as the percentage of BSU under
2.3 m (x — 2 o-) containing no vegetation.
Swindale and Curtis (1957) devised an ordination of aquatic
communities based on the relative frequency and joint occurrence
of each species. Only nine species in the Wingra study were given
joint occurrence numbers by Swindale and Curtis. Therefore, the
following modification was devised to calculate a compositional in¬
dex (Cl) value.
Cl =
5 relative frequency of joint occurrence #4 spp. X 100 + 300
2 relative frequency of joint occurrence #3 + joint occur. # 4 spp.
The modification can be used because the greatest percentage of
the relative frequency is concentrated in species with joint occur¬
rence numbers, therefore only minor species were eliminated. The
Swindale and Curtis (1957) scale goes from 100-400. The low end
of the scale indicates oligiotrophic conditions and the high end
represents eutrophic conditions. Lake Wingra is a highly eutrophic
lake, therefore few, if any, species with a joint occurrence number
of less than 3 would be expected.
Results
Depth, Lacustrine Soils , and Shore Vegetation
A hydrographic map of the littoral zone is presented in Fig. 2.
The outer limit of the vegetation is controlled by depth. The mean
depth of the end of the littoral zone is 2.7 ± 0.4 m. Because of the
extreme turbidity of the water, it was often hard to ascertain the
absolute end of the vegetation, even with SCUBA gear. It is as¬
sumed that one could find vegetation beyond the limits indicated
on the vegetation map (Fig. 3). If so it would be scattered and
highly stunted plants of M. spicatum. For more complete hydro-
graphic data, one should consult Noland (1951).
Marl composes the major part of the littoral zone soil. 68% of
the 40 transects had a marl bottom 5 m from shore, 20% had sand
and 12% had an organic bottom.
The organic bottom is found on the north, south, and west sides
of the lake. It is found extensively in areas of water lily vegetation
and areas where the shore vegetation is cattail marsh.
Sand beach is found along a road fill on the south shore of the
lake, and in an area centrally located along the north shore line.
The remainder of the areas have a marl bottom. All the beach soil
types grade into a black, organic “ooze” before the limit of the
littoral zone.
1971] Nichols and Mori— Vegetation of Lake Wingra
111
Figure 2. Map of Lake Wingra showing shore vegetation and depth in meters.
Shore vegetation: 1-developed land, 2-lowland shrub, 3-lowland forest,, 4-cattail
marsh.
Figure 3. Map of Lake Wingra showing the location of plant communities.
Plant communities : 1-Nymphaea community, 2-shallow water Myriophyllum
community, 3-deep water Myriophyllum community, A-Nuphar community, 5-
P otamogeton-Myriophyllum community, 6-Scirpus beds.
112 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
No obvious correlations could be made between shore vegetation
and aquatic macrophyte communities. Figures 2 and 4A depict the
vegetation types bordering the lake.
Flora of Lake Wingra
Fassett (1930) divided aquatic macrophytes into four different
life forms: forms with long lax stems and flexous leaves; forms
with basal rosettes and unbranched stems; forms with vegetative
stems and horizontal floating leaves; and forms that are rooted
underwater but have photosynthetic parts emergent. In considering
Lake Wingra a fifth category should be added to include small, free
floating plants.
Naming of species, except for M . spicatum follows Fassett, 1960,
For a history of the naming of M. spicatum one should consult
Fernald, 1919 and Fernald, 1945.
Four species, Lemna minor L., L. trisulca L., Spirodela polyrhiza
(L.) Schleid, and Wolfia Columbiana Karst., of the small, free float¬
ing plants were collected.
Nymphaea tuberosa Paine and Nuphar variegatum Engelm. rep¬
resent forms with vegetative stems and horizontal floating leaves.
Three species, Typlna latifolia L., T. angustifolia L. and Scirpus
Validus Vahl. represents forms that are rooted underwater but
have photosynthetic parts emergent.
No species with a basal rosette and unbranched stems were
found.
The remainder of the species, Potamogeton pectinatus L., P.
nodosus Poir, P. Richardsonii (Benn.) Rydb., P. zosterif ormis
Fern., P. crispus L., P. foliosus Raf., P. natans L., Zanichellia
palustris L., Najas flexilis (Willd.) Rostk., Schmidt, Anacharis
canadensis (Michx.) Planchon . Peter anther a dubia (Jacq.) MacM.,
Ceratophyllum demersum L., Utricularia vulgaris L., Ranunculus
longirostris Godron., and Myriophyllum spicatum L. are plants with
long lax stems and flexous leaves.
Table 1 shows the relative frequency each major species occupies
in the flora of the lake. M. spicatum is the most dominant member
with a relative frequency of 68.4%. No other species exceeds a
relative frequency of 10%.
An examination of herbarium sheets in the University of Wis¬
consin Herbarium has revealed specimens of Potamogeton freisii
Rupr., P. illinoinensis Morong., P. amplifolius Tuckern., P. prae-
longus L., and V. americana Michx. which were not recollected by
the authors in this study. None of these species have been collected
later than 1929.
M. spicatum and P. crispus are European introductions.
Table 1. Species Composition and Compositional Index of the Plant Communities in Lake Wingra.
1971]
Nichols and Mori — Vegetation of Lake Wingra
113
114 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Vegetation of Lake Wingra
For mapping and descriptive purposes five communities were
recognized in Lake Wingra. The shallow water Myriophyllum com¬
munity constituted 68.4% of the total vegetation. The Potamoge-
ton-Myriophyllum community constituted 16.5% of the total vege¬
tation. The deep water Myriophyllum community constituted 13.3%
of the total vegetation. The Nuphar community constituted 5.2%
of the vegetation and the Nymphaea community constituted 1.6%
of the vegetation. Figure 3 shows the location of each of these
communities in the lake.
The shallow water Myriophyllum community (Fig. 4B, 4F) is a
solid stand of M. spicatum with other species scattered or present,
and not more than one species of Potamogeton scattered in any
BSU. The stand is characterized by an average of 253 stems/m2
and a standing crop of 385 g/m2. All species combined, except for
M. spicatum, constitute less than 1 % of the density or the standing
crop. The frequency, relative frequency and Cl value are given in
Table 1. With a Cl value of 398, the shallow water Myriophyllum
community is an example of a community in a stage of advanced
eutrophication.
At a point (about 2.4 m) with increasing depth the Myriophyl¬
lum community changes from a solid stand to scattered plants.
This is the dividing line between the deep water and shallow water
Myriophyllum communities. The deep water community is more
monotypic; P. crispus and C. demersum (Table 1) are the only
species occurring with M. spicatum. With a scattering of plants
the density and standing crop drops considerably. The average
density was 70 stems/m2 and the average standing crop was 142
g/m2. Virtually 100% of the density and standing crop was com¬
posed of M. spicatum. M. spicatum colonizes the deepest portions
of the littoral zone. It has enough of a competitive advantage to
exclude virtually all other species.
The Potamogeton-Myriophyllum (Figure 4C) community is diffi¬
cult to define. For mapping purposes it was defined as an area
having one Potamogeton sp. continuous through the sampling unit
or at least two species scattered or present. The difficulty arises
in the fact that there is tremendous variety in this area. There
were many small associations of one or two species of Potamogeton
spatially located very near one another, but mutually exclusive.
A very intensive mapping technique would be required to separate
them, but the physiognomic difference was obvious in the field.
On the other hand, much of the Potamogeton-Myriophyllum com¬
munity appeared to be an ecotone between a narrow, shallow water
Potamogeton community and the somewhat deeper, shallow water
Myriophyllum community. The community is characterized by an
1971]
Nichols and Mori — Vegetation of Lake Wingra
115
Figure 4. Aquatic communities of Lake Wingra: A. Nymphaea tuberosa com¬
munity with Typha shore vegetation, B. Shallow water Myriophyllum com¬
munity, C. P o tamo geton-Myriophy Hum community, D. Nymphaea tuberosa
community, E. Nuphar variegatum community, F. Foreground: Nymphaea and
Nuphar communities showing close proximity. Background: Shallow water
Myriophyllum community showing broad, dense expanse of plant material.
average density of 192 stems/m2 and a standing crop of 196
g/m2. P. natans, M. spicatum and P. pectinatus were the three
most common species (Table 1). Monotypic stands of N. flexilis,
P. natans, P. nodosus, and P. pectinatus could be found. C. demer-
sum , although not wholly a member of the Potamogeton-
Myriophyllum community, was also observed in pure stands. These
stands never exceeded 10m2 and were not mapped as a separate
community. The Cl value of 374 indicates a lower level of eutrophi-
116 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
cation in this community. This is a direct reflection on the impor¬
tance of M. spicatum in the community.
Usually Nymphaea tuherosa and Nuphar variegatum are lumped
into a floating leaved community (Fassett, 1980). Again from field
observations, it was obvious that these two species occur spatially
close together, but mutually exclusive of one another (Fig. 4F).
An attempt was made, therefore, to separate the two communities.
With analysis the difference between the communities is striking.
The average density in the Nymphaea community (Fig. 4D) was
13 stems/m2 compared to 114 stems/m2 in the Nuphar (Fig. 4E)
community. The average standing crop was 49 g/m2 in the
Nymphaea community compared with 183 g/m2 in the Nuphar
community. M. spicatum was the only other species to occur com¬
monly in the Nymphaea community. It occurred only where
N. tuherosa became widely scattered. N. variegatum had a variety
of species (Table 1) associated with it. The species difference
reflects the Cl differences in Table 1.
The emergents, S. validus, T. latifolia , and T. angustifolia,
because of their minor importance, were not analyzed as a sepa¬
rate community. S. validus beds are indicated on the vegetation
map (Fig. 3).
Discussion
Over the last century, and more particularly the last few decades,
the submerged aquatic vegetation of Lake Wingra has changed
drastically from a community that was probably dominated by
Potamogeton spp. and V. americana to a type strongly dominated
by an European invader, M. spicatum. This change has not been
documented, but has most likely been paralleled in other areas.
M. spicatum was first collected in the Chesapeake Bay Region
in 1902 (Steenis et al, 1961). By 1962 the plant covered 200,000
acres and was considered a serious aquatic pest. Currituck Sound,
North Carolina illustrates the rapidity of the M. spicatum invasion.
First reports of the plant from here were received in 1965. At this
time approximately 100 acres were in the infestation stage and
500-1,000 additional acres showed initial establishment. By the
summer of 1966 these figures had risen to 8,000 and 67,000 acres
respectively (Crowel et al, 1967).
By 1967 M. spicatum was established in the Northeast in Ver¬
mont, New York, Pennsylvania, New Jersey and Delaware. In the
Midwest it had been reported in Ohio, Indiana, Illinois and Wis¬
consin. Its presence has been noted in the states of Maryland,
Virginia, North Carolina, Georgia, Florida, Alabama, Louisiana,
Texas and California.
1971] Nichols and Mori — Vegetation of Lake Wingra 117
Lake Mendota (also within the city of Madison) has been studied
and can be used for comparison. Denniston (1921) listed Vallisneria
spiralis L., Najas flexilis, P . Richardsonii , P. zosterif ormis and
P. pectinatus as the most abundant species in the order given.
Today even the most uncritical observer would have to rank
M. spicatum as the most abundant Lake Mendota aquatic vascular
plant. In a study by Rickett (1921) the average dry weight of
Myrophyllum per square meter in University Bay of Lake
Mendota was given as 12.6 g. This differs sharply with the value
of 172 g/m2 reported by Lind and Cottam (1969).
Andrews (1946) states, “By late June or early July, the long¬
stemmed pondweeds have risen above it (M. exalbescens Fern.),
and it is no longer conspicuous.” By contrast, in 1967, the peak
growth of Myriophyllum was in June and July (Lind and Cottam,
1969). Within the last fifty years there has been a startling
change in the abundance of Myriophyllum in Lake Mendota.
The invasion of M. spicatum appears to be a very recent occur¬
rence in Lake Mendota. Voucher specimens of M. spicatum were
identified for the authors by John Steenis in the summer of 1969.
The bulk of the plant material in 1969 was M. spicatum. Andrews
(1946) found no M. spicatum. Voucher specimens of M. exalbescens
were collected as late as 1962 on Lake Mendota. The earliest
recorded specimen of M. spicatum was collected in the state in
1936 by J. J. Davis in the Tomahawk region.
Like many other pests M. spicatum ranks high in uselessness.
It is of greatest public concern because it restricts man's recrea¬
tional and aesthetic activities. The concern in this report is di¬
rected toward its effect on the biological system.
M. spicatum is a very aggressive plant with a wide ecological
amplitude. Initially it appears beneficial to fish and wildlife. These
appearances are deceiving, and, as in the case of Lake Wingra, the
plant soon crowds more desirable aquatic plants. How stable the
Myriophyllum communities are, is yet to be determined. A mono-
typic community is ecologically unhealthy. It is felt that M. spicatum
has reached the maximum extent of its growth in Lake Wingra. The
Myriophyllum communities have also reached the maximum extent
of eutrophication as described by Swindale and Curtis (1957).
Although M. spicatum has a competitive edge in the deep water,
soft bottom areas of the lake, the effect is less intensive in the
shallow water, sandy bottom areas. There, other species constitute
a more substantial portion of the vegetation.
Curtis (1959) defines terrestrial weeds as plants of open or
disturbed habitats. The last major disturbance took place in Lake
Wingra in 1920. John H. Steenis (personal communication) re-
118 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
ported no M. spicatum in Lake Wingra from a date in the 193 0’s,
at least ten years after the last disturbance.
Much research is being expended on control measures for
M. spicatum by means of chemical poisoning and mechanical
harvesting. Little has been discovered about natural control, the
role the plant plays in the natural community, or the invasion
mechanism the plant uses to become established in the community.
Lake Wingra is the ideal location for continued research on
the community relations of M. spicatum. Up to this time only a
small portion of the lake has been disturbed by man’s control efforts.
By continued resampling of the lake and by using this report as
a base, questions on the role of M. spicatum in the community,
and questions on the course of community succession after maxi¬
mum eutrophication as described by Swindale and Curtis (1957)
might be answered.
Acknowledgements
The authors wish to express their thanks to Dr. Grant Cottam
for critical reading of the manuscript, to Mr. Robert Read and
Mr. David Nelson for aid in the field, and to the University of
Wisconsin Arboretum. Arboretum Journal Series Paper No. 77.
The work upon which this publication is based was supported
in part by funds provided by the United States Department of the
Interior as authorized under the Water Resources Research Act
of 1964, Public Law 88-379, OWRR JB-019-Wis, Agreement No.
14-01-0001-1968.
References Cited
Andrews, J. D. 1946. The macroscopic invertebrate population of the larger
aquatic plants in Lake Mendota. Unpublished Ph.D. Thesis, University
of Wisconsin.
Crowell, T. E., J. H. Steenis, and J. L. Sincock. 1967. Recent observations
of Eurasian watermilfoil in Currituck Sound, North Carolina, and other
coastal Southeastern states. Mimeo. 8 pp.
Curtis, J. T. 1959. The vegetation of Wisconsin. The University of Wisconsin
Press, Madison.
Denniston, R. H. 1921. A survey of the larger aquatic plants of Lake Men¬
dota. Wisconsin Academy of Sciences 20: 494-500.
Fassett, N. C. 1930. The plants of some northeastern Wisconsin Lakes. Wis¬
consin Academy of Sciences 25 : 157.
Fassett, N. C. 1960. A manual of aquatic plants. The University of Wisconsin
Press, Madison.
Fernald, M. L. 1919. Two new Myriophyllum and a species new to the United
States. Rhodora 21(247) : 120-124.
Fernald, M. L. 1945. Incomplete flora of Illinois. Rhodora 47 : 204-219.
Lind, C. T. and G. Cottam. 1969. The submerged aquatics of University Bay:
A study in eutrophication. The American Midland Naturalist 81(2) : 353-
369.
1971] Nichols and Mori— Vegetation of Lake Wingra
119
Noland, W. E. 1951. The hydrography, fish and turtle population of Lake
Wingra. Wisconsin Academy of Sciences 40(2) : 5-57.
Poff, R, J. and C. W. Threinen. 1962. Surface water resources of Dane
County. Wisconsin Conservation Department, Madison.
Rxckett, H. W. 1921. A quantitative study of the larger aquatic plants of Lake
Mendota, Wisconsin. Wisconsin Academy of Sciences 20: 501-527.
Saghse, N. D. 1965. A thousand ages. Regents of the University of Wisconsin,
Madison.
Steenis, J. H., V. D. Stotts, and C. R. Gillette. 1961. Observations on dis¬
tribution and control of Eurasian watermilfoil in Chesapeake Bay, 1961
Mimeo 7 pp.
Swindale, D. N. and J. T. Curtis. 1957. Phytosociology of the larger sub¬
merged plants in Wisconsin lakes. Ecology 38(3) : 397-407.
VARIABILITY IN WISCONSIN
IN TRIENTALIS BOREALIS RAF.
Roger C. Anderson 1
In North America Trientalis borealis extends from the Great
Slave Lake to the east coast. Its distribution approximates that of
the boreal forest in Canada and the northern conifer hardwoods
in the United States. Its range extends into the Appalachian Moun¬
tains and on to the New Jersey coastal plain (Anderson, 1968).
During an autecological study of Trientalis borealis in Wisconsin,
the great morphological plasticity of this species became apparent.
Studies were undertaken to examine the range of variability of
Trientalis in Wisconsin and compare it with the variability reported
for the European species of Trientalis ( Trientalis europaea L.).
A further objective was to determine if morphological differences
existed between different sectors of the state, specifically between
the area north of the tension zone (Curtis, 1959) and the southern
portion of the state. The tension zone divides the state into two
floristie provinces, the prairie forest and the northern hardwoods.
Trientalis has been reported for all but 6 of 72 counties in Wiscon¬
sin; however, it is most abundant north of the tension zone (Ander¬
son, 1968).
The morphological variation of T. europaea in Great Britain has
been discussed by Matthews and Roger (1941) and in Poland,
Norway, and Finland by Medwecka-Kornas (1963). In Canada,
Lepage (1946) delineated three forms of T. borealis based largely
on leaf shape and plant growth form, and Curtis (1959) comments
on the variability of T. borealis in Wisconsin.
Methods and Materials
Pressed specimens from the University of Wisconsin herbarium
were used to obtain the variability data across the state. Data per¬
taining to the fruits and seeds was obtained from material
collected at a single site, the University of Wisconsin Arboretum
Finnerud Forest, in Oneida County. The information collected
from the herbarium specimens included the stem length in centi¬
meters (from the point on the stem where it would be out of the
soil to the major leaf whorl), the number of leaves in the major
whorl, the number of flowers per plant, and the number of floral
1 University of Wisconsin Arboretum, 1207 Seminole Highway, Madison, Wis. 53711.
121
122 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
parts. For any one flower, the number of stamens, sepals and
petals was constant. However, multi-flowered plants occasionally
had flowers with different numbers of floral parts so that each
flower had to be counted separately. For some specimens it was
not possible to obtain all the information due to lack or damage
of stem, leaves or flowers. For each specimen, collection north
or south of the northern tension zone boundary was noted. Chi-
square tests were used to determine whether there were significant
morphological differences between northern and southern speci¬
mens. Fruit and seed size were determined by measuring the
longest dimension with the aid of a microscope ocular micrometer.
Results
Figure 1 shows the results of several measures of variability
on Wisconsin Trientalis . Plant height is the only measured feature
that does not differ significantly between northern and southern
specimens. In Figure la plant height has been divided into five
stem height classes (4-8 cm, 9-10, 11-12, 13-14, 15-19). Northern
plants tend to be slightly taller, whereas the southern specimens
have more individuals that are in the shortest group and propor¬
tionately fewer tall plants, Figure la. The average height of the
northern specimens was 11.9 cm compared to 11.1 cm for the
A North
O South
b. No. Loovo* /Whorl
c. No. Flowor
Port*
No. Flowor*
FINNERUD
40-
Figure 1. Some aspects of the variability of Trientalis borealis.
1971] Anderson— Variability in Trientalis Borealis Raf. 123
southern plants. Considering all the specimens, plant height ranges
from 4 cm to 19 cm with an average of 11.6 cm.
The number of leaves on the major whorl varied from 4 to 10
leaves with an average for the complete sample of 6.8 and a mode
of 7 leaves. Figure lb shows that for the northern specimens six
leaves was as common as seven. The average number of leaves
for the northern specimens was 6.5 leaves compared to 7.1 for the
southern. A chi-square test indicates that there is a significant
difference (.01 level) in the number of leaves between locations of
collection. For the test statistics, specimens having five leaves or
fewer per whorl were combined. Similarly those with nine or more
leaves were grouped, because of the smaller number of specimens
in these groups.
A significant difference (.005 level) in the number of flowers
and the number of floral parts (.025 level) was found between
northern and southern specimens. Figure Id shows that northern
plants commonly produced only a single flower (75 per cent of the
specimens) and the usual number of flowering parts was seven
(77.6 per cent), Figure 1c. In the south, specimens with 6 flower
parts were more numerous, accounting for 22.9 per cent of all the
southern specimens, compared to 4.0 per cent for the northern
plants.
The capsules of Trientalis vary in size from 1.66 mm to 2.66 mm
with an average of 2.2 mm. The capsules contained from 2 to 14
seeds, Figure le, with a mode of 10 and an average of 9.4 seeds.
Seed length, Figure If, ranged from .56 mm to 1.43 mm with an
average of 1.18 mm. The weight of 233 air dried seeds was 111
mg with an average weight of .476 mg.
Discussion
The variability described by Matthews and Roger (1941) for
T. europaea in Great Britain was largely variation in plant height
and the number of leaves per whorl. They report that plants with
6 leaves are most frequent, accounting for 42.4 per cent of all
the specimens examined. For Wisconsin specimens, 20.1 per cent
of the plants had 6 leaves per whorl, while plants with seven
leaves per whorl were the most common, 30.6 per cent of the
specimens. Table 1 compares the number of leaves per plant for
the Wisconsin specimens with T. europaea as reported by Matthews
and Roger (1941).
The table shows that all three groups differ in the number of
leaves per whorl, with the southern Wisconsin T. borealis plants
having the most leaves. However, the variation between the two
T. borealis groups is less than that between the species. Medwecka-
Kornas (1963) reports that in northern Europe the number of
124 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Table 1.
Number of Plants
T. europaea .
North Wis. T. borealis .
South Wis. T. borealis .
Combined Wisconsin .
Average North — 6.5 leaves.
Average South — 7. 1 leaves.
Average combined — 6.8 leaves
(Wisconsin).
No. Leaves Per Whorl .
277
27
11
38
488
29
13
42
212
29
35
64
115
19
20
39
9 10
leaves per whorl varies from 4 to 11, but Hegi (1927) gives a
range of 5 to 12 leaves for T. europaea. In Wisconsin, plants of
T. borealis with 12 leaves have been observed. Thus, the variation
in number of leaves between the two species is about the same.
For Trientalis europaea in Great Britain the variation in stem
length was 3 to 20 cm (Matthews and Roger, 1941), and for
northern Europe .6 to 27.5 cm (Medwecka-Kornas, 1963) com¬
pared with 4 to 19 cm for the Wisconsin plants. The height of the
European species is influenced by environment; plants exposed to
severe environments are smaller. In Wisconsin, plant height is
also related to environment, with shorter plants generally asso¬
ciated with drier habitats.
litis and Shaughnessy (1960) indicate that the star-shaped
flowers of T. borealis, 12-20 mm in diameter, are usually 7-merous.
In this study the southern specimens were more variable than the
northern. The 7-merous flowers were found to be the most com¬
mon throughout the state, but the 6-merous flowers were more fre¬
quent south of the tension zone.
For the Eurasian species, the number of seeds per capsule ranged
from one to eighteen with an average of eight seeds per capsule
(Matthews and Roger, 1941). Capsules of Trientalis borealis con¬
tain two to fourteen seeds with an average of 9.4. Matthews and
Roger (1941) report the average seed weight to be .680 mg, 30
per cent heavier than T. borealis seeds, .476 mg.
Another variation in Trientalis is the “ramose” form that is
characterized by having an additional verticel of leaves above the
main whorl. The earliest published record of this growth form is
by Hegi (1927). Lepage (1946) recognized three forms of T.
borealis in Canada, one of which is the “ramose” form. Three
morphological forms were delineated by Medwecka-Kornas (1963)
in northern Europe: a normal form distributed south of the tree
1971] Anderson— Variability in Trientalis Borealis Raf. 125
line, a diminutive form with a northern range beyond the limits of
the boreal forest of the fjeld field of Lapland, and a “ramose”
form growing on the peripheries of the distribution of the normal
form in disturbed or open habitats. The morphological forms inter¬
grade, and she suggests that the forms are the results of environ¬
mental modifications. Individuals of the normal form were found
to develop additional verticels of leaves after being transplanted
to an open garden (Hiirsalmi, 1960). In Wisconsin, plants with
this form were occasionally encountered in the held, but they are
more common among plants grown under greenhouse conditions.
Because of the morphological variability of Trientalis in Wiscon¬
sin and the diverse habitats in which it grows, Curtis (1959) sug¬
gested that there may be ecotypes within the population of
Trientalis borealis in Wisconsin. Some of this variation may be due
to environmental differences. However, near the edge of a species
range it seems likely that there may be selection for genetic com¬
binations that are better adapted to conditions that differ from
those in the main part of the range. In Wisconsin the tension zone
delineates areas that have climatic and edaphic differences. When
plants collected north and south of the tension zone are examined,
significant morphological variations between the two are found.
However, the differences are quantitative rather than qualitative,
and nearly all the variability found in Wisconsin Trientalis occurs
both north and south of the tension zone. The Wisconsin specimens
of Trientalis appear to be a single taxon displaying the same kind
of variation from north to south within the state.
Literature Cited
Anderson, Roger C. 1968. The autecology of Trientalis borealis. Ph.D. Thesis,
University of Wisconsin, Madison.
Curtis, John T. 1959. The vegetation of Wisconsin. University of Wisconsin
Press, Madison. 657 p.
Hegi, Gustav. 1927. Flora of middle Europe. 5(3) : 1816-1864. J. F. Lehmans,
Munich.
Hiirsalmi, H. 19'61. Studies on the reproduction, biology, variation, and ecol¬
ogy of Finnish populations of Trientalis europaea L. Msc. Thesis. Dept,
of Botany. University Turku, Finland. (In Medwecka-Kornas’, 1963).
Iltis, Hugh H. and Winslow M. Shaughnessy. 1960. Preliminary reports on
the flora of Wisconsin No. 43. Primulaceae. Wis. Acad. Sci. Arts Lett.,
Trans. 49: 113-135.
Matthews, J. R. and J. G. Rogers. 1941. Variation in Trientalis europaea L
J. Bot. 79: 80-83.
Medwecka-Kornas, Anna. 1963. Observations on the variability of Trientalis
europaea L. in Finland, Norway, and Poland. In Botanical Institutes
Berichte, Des Geobotanischen Institutes Der Eidg. Tech. Hochshule, Stif-
tung Rubel, 34 Heft, Bericht uber das Jahr 1962, Redaktion Heinz Ellen-
berg, Zurich, 1963.
Lepage, Ernest. 1946. Variation taxonomique de trois especes Laurentiennes.
Nat. Canad. 73: 5-16.
;
li
THE INSECT PARASITES OF THE INTRODUCED PINE SAWFLY,
DIPRION SIM I LIS (HARTIG) (HYMENOPTERA:
DIPRIONIDAE), IN WISCONSIN, WITH KEYS TO
THE ADULTS AND MATURE LARVAL REMAINS
James W. Merlins and Harry C. Coppel
The introduced pine sawfly, Diprion similis (Hartig), was first
discovered in North America in 1914, in a nursery at New Haven,
Connecticut (Britton, 1915). The larvae collected may have been
the first progeny arising from foreign stock which probably entered
this country on nursery material, Kapid spread of the insect
resulted in reports of infestations throughout the northeastern
states and as far south as Virginia by 1923 (Middleton, 1923). In
1931, D. similis appeared in Canada at Oakville, Ontario (Munro,
1935), and was reported from Wisconsin near Menomonie by
1944 (Coppel, 1962),
The hosts of the sawfly include species of the genus Finns. In
North America the favored host is eastern white pine, Finns strobus
L. Other species of pines commonly attacked in Wisconsin are
Scotch pine, P. sylvestris L., and red pine, P. resinosa Ait. Trees
of all ages are defoliated, but feeding is particularly severe in the
most exposed locations and in the overstory, where stripping of
foliage may result in branch mortality. Tree mortality occasionally
results when a large second generation of larvae destroys the
buds set for the following year. Although the sawfly is primarily
a pest of plantations, nurseries, and ornamentals, its habits in
Wisconsin make it a serious threat to the production of eastern
white pine forests.
The data, accumulated since 1957 on the parasite complex of
the introduced pine sawfly in Wisconsin, were organized in 1965.
This information was compiled from both field collections and labo¬
ratory studies of material collected during the summers of 1957-68
in northwestern Wisconsin ; principally in Polk, Burnett, and Wash¬
burn Counties.
This paper, the third in a continuing but unnumbered series con¬
cerning the insect parasites of Wisconsin forest insect pests, deals
with the known parasites of D. similis in Wisconsin. Keys are pre¬
sented both for the separation of the adult parasites, and for the
remains left in the host cocoon after the parasite has emerged.
Descriptions of the adults, final larval instar cephalic structures
127
128 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
and spiracles of Hymenoptera, and buccopharyngeal apparatuses
and stigmal plates of Diptera are given. Notes on the biology of the
parasites are also included.
Methods
The keys are based on the adult parasites, and the host and
parasite remains positively associated with them. The information
used for preparation of the key to parasite remains consisted of
the host cocoon itself, especially the size, position, and type of
emergence hole made by the parasite; the absence or presence,
and appearance of the parasite cocoon or puparium ; the cast larval
skin of hymenopterous parasites and the buccopharyngeal arma¬
ture of dipterous parasites. Cocoons from which parasites had
emerged were cut open with a razor blade to observe the location
and appearance of the contents. Cast larval skins of hymenopterous
parasites were removed, softened in warmed 10% KOH for 20-80
minutes, and rinsed in distilled water. While rinsing, the skins
were spread under the dissecting microscope with two insect pins,
and then mounted on microscope slides in non-resinous mounting
medium (Turtox CMC-10). The final instar buccopharyngeal arma¬
ture of each dipterous parasite was removed from the inside of the
puparial case, softened in KOH, cleaned of extraneous membranes
in distilled water, and mounted. Posterior spiracular plates of
Diptera were taken from the puparial cases, softened, and mounted.
Illustrations of adults and gross characters of larval remains
were made with the aid of Bausch and Lomb binocular dissecting
microscope fitted with lOx eyepieces and a 2x enlarger lens. Fine
details of immature remains were added through the use of an
Ernst Leitz binocular compound microscope. Measurements were
made with a Reichert ocular micrometer calibrated with a stage
micrometer (American Optical Co.) on each microscope.
Terminology used for the parts of the cephalic structures and
spiracles of final instar hymenopterous larvae and the bucco¬
pharyngeal armature of the Diptera is the same as that assembled
by Finlayson (1960) from various authors.
Parasites Obtained
The following 21 species of Hymenoptera and 4 of Diptera were
reared from D. similis cocoons in Wisconsin :
Hymenoptera
Ichneumonidae : Scambus hispae (Harris), Delomerista japonica
(Cushman), D. novita (Cresson), Itoplectis conquisitor (Say),
Exenterus amictorius (Panzer), E. canadensis Provancher,
Gelis tenellus (Say), Agrothereutes lophyri (Norton).
1971] Merlins and Coppel — Parasites of the Pine Sawfly 129
Eulophidae: Dahlbominus fuscipennis (Zetterstedt) , Tetrastichus
coerulescens Ashmead, Elasmus apanteli Gahan.
Eupelmidae: Eupelmus spongipartus Foerster, Eupelmella (—Mac-
roneum of Peck, 1963) vesicularis (Retzius) .
Torymidae: Monodontomerus dentipes (Dalman).
Pteromalidae : Amblymerus verditer (Norton), Tritneptis scutel -
lata (Muesebeck), Dibrachys cavus (Walker), Catolaccus
cyanoideus Burks, Habrocytus thyridopterigis Howard.
Eurytomidae : Eurytoma pini Bugbee.
Chalcidae: Spilochalcis albifrons (Walsh).
Diptera
Tachinidae: Spathimeigenia spinigera Townsend, Bessa harveyi
(Townsend), Diplostichus lophyri (Townsend), Euphorocera
edwardsii (Williston).
Two keys have been prepared for the separation of the afore¬
mentioned species. The first allows separation of the parasite species
on the basis of the remains left in the host cocoon after adult
emergence. The second separates the adult parasites.
Key to the Parasites of D. similis Based on Parasite Remains
1. Host cocoon containing parasite puparium or cocoon _ 2
Host cocoon not containing parasite puparium or cocoon_14
2. (1) Host cocoon containing puparium, exit hole at tip of
cocoon _ 3
Host cocoon containing parasite cocoon _ 6
3. (2) Exit hole usually with edges of tapering thickness, ragged
and bent outward in appearance; no lid attached (Fig.
144) _ Spathimeigenia spinigera Tnsd.
Exit hole sharply cut, with or without lid attached _ 4
4. (3) Exit hole usually with hinged lid (Fig. 146) ; mandibular
hooks fused with intermediate sclerite (Figs. 99, 100) __5
Exit hole without lid (Fig. 145) ; mandibular hooks sepa¬
rate from intermediate sclerite (Fig. 98) _
- Bessa harveyi (Tnsd.)
5. (4) Length of mandibular hooks plus intermediate sclerite less
than one-half the total length of buccopharyngeal appa¬
ratus (Fig. 99) - Diplostichus lophyri (Tnsd).
Length of mandibular hooks plus intermediate sclerite
greater than one-half the total length of buccopharyngeal
apparatus (Fig. 100) _ Euphorocera edwardsii (Will.)
6. (2) Parasite cocoon abortive, seldom more than a crude tan¬
gled covering of silk over host remains _ 7
Parasite cocoon complete _ 9
130 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
7. (6)
8. (7)
9. (6)
10. (9)
11. (10)
12.(11)
13. (9)
14. (1)
15. (14)
16. (15)
Epistoma complete, labial sclerite open dorsally, mandibles
without teeth (Fig. 80) _ Itoplectis conquisitor (Say)
Epistoma incomplete, labial sclerite closed dorsally, mandi¬
bles each with two rows of fine teeth on blade and a large
posteromedial tooth (Figs. 76, 77) _ 8
Labial sclerite angular dorsally, resembling either an acute
or truncate peak (Fig. 76) _ Delomerista japonica Cush.
Labial sclerite rounded dorsally (Fig. 77) _
- Delomerista novita (Cress.)
Labral sclerite present ; epistomal arch incomplete or very
lightly sclerotized so as to appear incomplete (Figs. 75,
81, 82) _ 10
Labral sclerite absent ; epistoma complete (Figs. 78, 79) _ 13
Labial sclerite with a group of dome-like protuberances
ventrally (Fig. 75) Scambus (Scambus) hispae (Harris)
Labial sclerite with no such protuberances _ 11
Labial sclerite open dorsally _ 12
Labial sclerite closed dorsally _ 8
Spiracle short and broad, closing apparatus constituting
about one-half of its total length (Fig. 112) _
_ Agrothereutes lophyri (Nort.)
Spiracle longer, funnel shaped, the closing apparatus con¬
stituting only about one-third of its total length (Fig. Ill)
- Gelis tenellus (Say)
Atrium of spiracle wider than deep, usually elliptical in
outline (Fig. 110) _ Exenterus canadensis Prov.
Atrium of spiracle almost as deep as wide, widest at top
and tapering toward its stalk (Fig. 109) _
_ Exenterus amictorius (Panz.)
Exit hole very nearly at tip of host cocoon, edge slightly
ragged or irregular; diameter 1. 3-2.0 mm _ 15
Exit hole in various positions, often on side of cocoon;
edges smoothly cut or slightly scalloped; diameter 0.4-
1.2 mm _ 17
Exit hole large, 1. 5-2.0 mm in diameter (Fig. 128) _
_ _ _ Itoplectis conquisitor (Say)
Exit hole smaller, about 1.3 mm in diameter _ 16
Remains of parasite consisting of exuviae and bucco¬
pharyngeal apparatuses of the first two instars of dip¬
terous larva within host cadaver ; exit hole as in Fig. 145
_ Bessa harveyi (Tnsd.)
Parasite remains other than within host cadaver contained
within the host cocoon; cephalic structure as in Fig. 95
_ Spilochalcis albifrons (Walsh)
1971] Mertins and Coppel — Parasites of the Pine Sawfly 131
17. (14) Cast skin of last larval instar with long hairs or setae _ 18
Cast skin of last larval instar without conspicuous long
hairs or setae _ 22
18.(17) Skin densely covered with long (0.4 mm) setae; antennae
dome-like; blades of mandibles straight (Fig. 88) _
- - - Monodontomerus dentipes (Dalm.)
Skin sparsely covered with setae ; antennae peg-like ; blades
of mandibles curved _ 19
19. (18) Mandibles each with a large tooth (Fig. 94) _
- - - Enrytoma pini Bugbee
Mandibles without large tooth _ 20
20. (19) Clypeus absent _ _ _ _ Catolaccus cyanoideus Burks
Crescent-shaped, toothed clypeus present (Figs. 86, 87) _21
21. (20) Clypeus with 7 or 8 well-formed teeth (Fig. 86) _
- - Eupelmus spongipartus Foerst.
Clypeus with 3-5 well-formed teeth (Fig. 87) _
- - Eupelmella vesicularis (Retz.)
22. (17) Atrium of spiracle with at least 10 chambers (Figs. 113,
114, 115) _ __23
Atrium of spiracle with 4-8 chambers; antennae cone¬
like _ 25
23. (22) Cephalic structure of last larval instar showing mandibles,
pleurostoma, hypostoma, superior and inferior mandibular
processes (Fig. 84) _ Tetrastichus coerulescens Ashm.
Cephalic structure of final larval instar with only man¬
dibles visible _ 24
24. (23) Cast pupal skins dark brown; ventral process of mandible
much longer than dorsal process; teeth on blade of man¬
dible prominent (Fig. 85) _ _ Elasmus apanteli Gah.
Cast pupal skins golden-yellow; ventral process of man¬
dible but little longer than dorsal process; teeth on blade
of mandible obscure (Fig. 83) _
_ Dahlbominus fuscipennis (Zett.)
25. (22) Cephalic structure of final larval instar with mandibles,
epistoma, pleurostoma, hypostoma, superior and inferior
mandibular processes (Figs. 90, 91) ; atrium of spiracle
with 4 or 5 chambers (Figs. 120, 121) _ _ _ 26
Cephalic structure of final larval instar with only man¬
dibles and sometimes a lightly sclerotized articulation vis¬
ible (Figs. 89, 93) ; atrium of spiracle with 5-8 chambers
(some may appear sub-divided) (Figs. 119, 123) _ 27
26. (25) Ratio of mandibular width: length fg0.8 (Fig. 90) ; exit
hole diameter 0.5-0. 6 mm; usually secondary parasite
_ _ _ Tritneptis scutellata (Mues.)
132 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
Ratio of mandibular width: length >0.8 (Fig. 91) ; exit
hole diameter 0.7-0. 8 mm ; usually primary parasite _
- Dibrachys cavus (Walk.)
27.(25) Antennae usually with sockets visible (Fig. 93); total
length of spiracle 2. 8-3.0 times length of closing apparatus
(Fig. 123) _ Habrocytus thyridopterigis How.
Antennae with no visible antennal sockets (Fig. 89) ; total
length of spiracle 2.3-2. 5 times length of closing apparatus
(Fig. 119) _ Amblymerus verditer (Nort.)
Key to Adult Parasites of D. similis
1. Antennae 3 segmented, the third bearing an arista; body
bristly (Diptera) _ 2
Antennae with at least 8 apparent segments ; body without
bristles (Hymenoptera) _ 5
2. (1) Vein Mi+2 rounded distally, not angular (Fig. 68) ; only 2
bristles at base of vein R4+5; posterior tip of abdomen red-
brown _ Spathimeigenia spinigera Tnsd.
Vein Mi+2 distinctly angular distally (Figs. 70, 72, 74) ;
4-8 bristles at base of vein R4+5; posterior tip of abdomen
black or gray _ 3
3. (2) With a pair of small, usually cruciate bristles on apex of
mesoscutellum (Figs. 72, 74) ; palpi yellow _ 4
Without small pair of bristles at apex of mesoscutellum;
palpi dark brown (Fig. 70) _ Bessa harveyi (Tnsd.)
4. (3) With a pair of bristle-like hairs on the mesoscutellar disc
larger than the others; vein M4+2 pronouncedly recurved
distally (Fig. 72) _ Diplostichus lophyri (Tnsd.)
All the hairs on the mesoscutellar disc of equal size; vein
M4+2 forming an obtuse angle, with little or no tendency
towards recurvation (Fig. 74) _
_ Euphorocera edwardsii (Will.)
5. (1) Antennae geniculate (Chalcidoidea) _ 13
Antennae filiform (Ichneumonidae) _ 6
6. (5) Clypeus with a median apical notch (Figs. 1, 5, 9) _ 7
Clypeus rounded or angular apically, never with a median
notch _ _ _ _ 9
7. (6) Tarsal claws of female with a large basal tooth (Fig. 4) ;
front below antennal sockets black in male _
_ Scambus (Scambus) hispae (Harris)
Tarsal claws of female without basal tooth; front below
antennal sockets creamy white in male _ 8
8. (7) Ovipositor with a strong dorsal curve apically (Fig. 8) ;
hind tibia of male not as long as the femur and trochanters
combined _ _ _ Delomerista japonica Cush.
1971] Merlins and Coppel — Parasites of the Pine Saw fly 133
Ovipositor without a strong dorsal curve apically (Fig.
12) ; hind tibia of male as long as femur and tronchanters
combined _ Delomerista novita (Cress.)
9.(6) Ovipositor short and inconspicuous, mostly concealed
within the abdomen (Figs. 17, 20) ; mesoscutellum yellow
_ 10
Ovipositor longer, extending beyond the tip of the abdomen
and at least one-third as long; mesoscutellum white, black
or brown _ 11
10. (9) With a large yellow spot on either side of propodeum; all
yellow margins of abdominal tergites less than one-third
the length of the tergite _ Exenterus canadensis Prov.
No yellow spot on sides of propodeum; yellow margins of
tergites I and II broad, one-third to four-fifths the length
of the tergite _ Exenterus amictorius (Panz.)
11. (9) Eyes prominently emarginate at the level of antennal
sockets (Fig. 13) _ Itoplectis conquisitor (Say)
Eyes not emarginate _ 12
12. (11) Forewings with two broad, transverse dark bands (Fig.
23) _ Gelis tenellus (Say)
Forewings evenly dusky hyaline _
_ Agrothereutes lophyri (Nort.)
13. (5) Mesofemur dorsoventrally flattened; mesotarsus short,
broadened and spinous beneath (Figs. 37, 39) _ 20
Mesothoracic legs not so modified _ 14
14. (13) Metafemora greatly enlarged and toothed ventrally (Fig.
64) _ Spilochalcis albifrons (Walsh)
Metafemora not so modified _ 15
15. (14) Metacoxae at least 3 times as large as procoxae (Figs. 33,
41) _ 16
Metacoxae only slightly larger than procoxae _ 17
16. (15) Tarsi 4 segmented, extremely long; radius attached only
proximally (Fig. 33) _ Elasmus apanteli Gah.
Tarsi 5 segmented, normal length; radial vein apparently
forming a complete semicircle attached to front margin of
wing at both ends (Fig. 41) _
- Monodontomerus dentipes (Dalm.)
17. (15) Pronotum large and rectangular when viewed dorsally ;
antennae with 10 segments, last 3 fused (Figs. 60, 62)
- - - - - Eurytoma pini Bugbee
Pronotum narrowed, at least dorsomedially ; antennae with
8 or 13 segments _ 18
18.(17) Antennae with 8 segments; tarsi 4 segmented _ 19
Antennae with 13 segments, the last 3 of which may be
more or less fused; tarsi 5 segmented _ 21
134 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
19. (18) Forewings infumate centrally (Fig. 28); male antennae
with 3 branch-like projections (Fig. 29) _
_ Dahlhominus fuscipennis (Zett.)
Forewings hyaline _ Tetrastichus coerulescens Ashm.
20. (13) Wings reduced to non-functional vestiges (Fig. 39) _
- Eupelmella vesicularis (Retz.)
Wings normally developed _
- - - Eupelmus spongipartus Foerst.
21. (18) Antennal sockets about one-half way up the front (Fig.
56) _ Habrocytus thyridopterigis How.
Antennal sockets lower, usually about even with lower
margin of eyes (Figs. 43, 47, 50, 53) _ 22
22.(21) Forewings infumate centrally (Fig. 48) _
_ Tritneptis scutellata (Mues.)
Forewings hyaline _ 23
23. (22) Head disproportionately large; cheeks deeply grooved or
excavated (Figs. 53, 54) _ Catolaccus cyanoideus Burks
Head of normal proportions ; cheeks not hollowed out _ 24
24. (23) Postmarginal vein about as long as radius (Fig. 51) ; body
very dark green _ Dibrachys cavus (Wlkr.)
Postmarginal vein distinctly longer than radius (Fig. 44) ;
body bright metallic green --Amblymerus verditer (Nort.)
Notes on Biology and Descriptions of Parasites
Hymenoptera
Ichneumonidae
Scambus (Scambus) hispae (Harris)
Figs. 1, 2, 3, 4, 75, 105, 126
S. hispae was reared sporadically from D. similis during the
present study. It has not been reported previously from this host.
It is a solitary, usually internal feeder on D. similis, attacking and
emerging from the cocoon, and is usually a primary parasite.
The round, sharply cut emergence hole (Fig. 126) is slightly to
the side of one end of the host cocoon and is 1.5-1. 9 mm in diameter.
The parasite cocoon is semi-opaque, thick, and is made of tough
tan silk. It is nearly the size and shape of the host cocoon. The host
remains are appressed to the inside of the host cocoon but are out¬
side of the parasite cocoon. In the end of the parasite cocoon op¬
posite the emergence hole are a number of hard, shiny black pel¬
lets of larval meconium. These are flattened-oval in shape and are
usually fused together in a mass. The translucent pale white pupal
skin of the parasite is loose within the parasite cocoon. The skin
of the female shows a prominent and characteristically long ovi-
1971] Mertins and Copy el— -Parasites of the Pine Sawfly 135
Figures 1-8. Adult hymenopterous parasites of D.
similis. 1-4, Seambus (Scambus) hispae ; 1, head
capsule, frontal view; 2, female, lateral view; 3,
male abdomen, lateral view; 4, tarsal claw of fe¬
male. 5-8, Delomerista japonica; 5, head capsule,
frontal view; 6, female, lateral view; 7, male ab¬
domen, lateral view; 8, tip of ovipositor, lateral view.
positor sheath. White pellets of adult parasite meconium often are
found within the cocoon.
The final larval skin is light brown and is heavily pigmented
about the head capsule. It usually does not shrink completely, but
remains distended, at least lengthwise. Under magnification it is
densely covered with blunt conical papillae and scattered short
setae. The cephalic structure (Fig. 75) has a weakly sclerotized
epistoma which is usually broken in molting, and prominent labral
and suspensorial sclerites. The mandibles each have 2 rows of fine
teeth. The stipital sclerites are elongate and straight. The most
easily identifiable feature of the structure is the group of dome¬
like protuberances on the ventral portion of the labial sclerite.
136 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Although usually a primary parasite, S. hispae was reared once
as a secondary through I. conquisitor and once through E. amic-
torius. In one case, both S. hispae and G. tenellus emerged from
the same cocoon, and in another instance, a female I. conquisitor
shared a sawfly cocoon with a female S. hispae; both reached ma¬
turity, but only S. hispae emerged successfully. The sex ratio of
specimens reared was 48 females:? males. Males differ from fe¬
males in their narrower abdomen without an ovipositor (Fig. 3),
and in the absence of a basal tooth on the tarsal claw.
Delomerista japonica Cushman
Figs. 5, 6, 7, 8, 76, 106, 127
D. japonica is a fairly common and occasionally abundant para¬
site of D. similis , with some being reared every year. The insect
attacks and emerges from the host cocoon. It is a solitary primary
parasite.
The exit hole (Fig. 127) is slightly off the tip of the host cocoon,
round, but irregularly cut, and 1.8-2. 2 mm in diameter. The host
cocoon contains the flattened sawfly remains against a side wall,
and usually walled off with a layer of coarse tan or white silk.
Sometimes the silk is extended to form a nearly complete inner
lining of the host cocoon. The hardened, deep red-brown to black
larval meconium is in the end of the cocoon opposite the exit hole
and may be in the form of pellets. The shrunken, shiny pale yellow
pupal skin is loose within the cocoon. Several chalky white pellets
of adult meconium usually are also present.
The skin of the Anal instar is loose in the cocoon and is light
yellow with brown, strongly sclerotized cephalic structures. It is
covered densely with minute conical papillae and intermittently
with setae 0.13 mm long. The epistoma of the cephalic structure is
incomplete (Fig. 76). A prominent bow-shaped labral sclerite is
present. The mandibles each have 2 rows of fine teeth on the blades,
and 1 large posterior medial tooth. The stipital sclerites are short
and broad.
The sex ratio of reared specimens favored females 37 :28. Males
can be differentiated from females by the absence of an ovipositor,
and the presence of a white face and white pro- and mesothoracic
legs.
Delomerista novita (Cresson)
Figs. 9, 10, 11, 12, 77, 107, 127
D. novita is a rare parasite of D. similis, only 4 females and
possibly 1 male having been reared, and it has not been reported
previously from this sawfly. Its life history is similar to D. japon¬
ica, and all the life stages are comparable to that species. The adult
1971] Mertins and Coppel — Parasites of the Pine Sawfly 137
Figures 9-15. Adult hymenopterous parasites of D.
similis. 9-12, Delomerista novita; 9, head capsule,
frontal view; 10, female, lateral view; 11, male ab¬
domen, lateral view; 12, tip of ovipositor,, lateral
view. 13-15, Itoplectis conquisitor; 13, head capsule,
frontal view; 14, female, lateral view; 15, male ab¬
domen, lateral view.
female may be separated readily from D. japonica by the gentler
dorsal curve in the tip of the ovipositor (Fig. 12). According to
Townes and Townes (1960) the males of the two species may be
separated by the relative lengths of the metatibia (couplet 8 of the
key to adult parasites), but the single specimen at hand is cer¬
tainly borderline. The immature stages of the two species are as
difficult to separate. In the specimens examined, the cephalic struc¬
ture of D. novita was larger and more heavily sclerotized than that
of D. japonica, but this could be due to individual variation. The
best differentiating character seems to be the difference in the
shape of the labial sclerite (Fig. 77). That of D. japonica is some¬
what angular, especially dor sally, whereas in D. novita it is
138 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
smoothly rounded. In addition, the ratio of width to length of the
spiracle of D. novita is 0.7 :1, whereas that of D. japonica is about
0.6:1, although a number of D. japonica specimens have spiracles
similar in shape to those of D. novita (Figs. 106, 107).
Itoplectis conquisitor (Say)
Figs. 13, 14, 15, 80, 108, 128
I. conquisitor was a common and sometimes abundant parasite
of D. similis. It nearly always functioned as a primary, solitary,
internal feeder in the cocoon.
The emergence hole (Fig. 128) is close to the tip of the host
cocoon, 1.55-2.0 mm in diameter, usually round, but with ragged
edges. The host remains consist either of larval integument, or
occasionally a pupal skin. The remains are flattened solidly against
the cocoon wall and usually partitioned off with at least a crude
mat of rough brown silk, which may be expanded to a semi-complete
lining of the host cocoon. A tan to red-brown pellet mass of larval
meconium adheres to the side of the cocoon opposite the exit hole,
and occasionally one to several elongate white pellets of adult
meconium are present within the cocoon. The last larval and pupal
skins are free in the end opposite the exit hole. The pupal skin is
pale, translucent and often fragmented.
The final instar skin is yellow to light brown with a prominent,
strongly sclerotized cephalic structure. It is covered with minute
spines, and a few inconspicuous setae. The cephalic structure
(Fig. 80) cannot be confused with that of any other parasite of
D. similis. It appears as a complete circle broken only by the labial
sclerite ; the hypostomal arms are lacking ; the mandibles are large
and usually show 15 or 20 rounded protuberances medially; the
area above the silk press is densely covered with sharp spines. The
disc-like antennae are small and inconspicuous. The spiracle (Fig.
108) is short and oval-shaped.
The adult is easily recognized by the prominent light and dark
bands on the posterior tibiae and tarsi. The sex ratio of reared
specimens favored females by 4.5:1. Males are separated easily
from females by the absence of an ovipositor (Fig. 15). The single j
instance of I. conquisitor as a hyperparasite found it as a secondary
through E. amictorius, and in one case, both 7. conquisitor and G.
tenellus emerged from the same host cocoon.
Exenterus amictorius (Panzer)
Figs. 16, 17, 18, 78, 109, 129
This European species was not recorded from D. similis in Wis¬
consin until 1961, and has increased steadily in abundance since
that time to rival Monodontomerus dentipes as the most important
1971] Merlins and Coppel — Parasites of the Pine Saw fly 139
Figures 16-21. Adult hymenopterous parasites of D.
similis. 16-18, Exenterus amictorius ; 16, head cap¬
sule, frontal view; 17, female, lateral view; 18, male
abdomen, lateral view. 19-21,, Exenterus canadensis ;
19, head capsule, frontal view; 20, female, lateral
view; 21, male abdomen, lateral view.
species in the parasite complex (Mertins and Coppel, 1968). It is
a solitary, primary parasite of the sawfly larva, and emerges from
the cocoon.
The exit hole (Fig. 129) is near the end of the host cocoon, round
and jaggedly cut. Except for E. canadensis, which makes an equally
large hole, the exit hole is the largest of the parasites emerging
from D. similis cocoons, being 2. 1-2. 5 mm in diameter. The parasite
cocoon is about the size and shape of the host cocoon, thin, trans¬
lucent, shiny, and white to pink in color. The host remains adhere
to the lateral wall of its cocoon and outside that of the parasite.
The partially embedded, empty chorion of the egg from which the
parasite larva emerged can frequently be seen along its dorsum,
usually in the thoracic region. At the end of the cocoon opposite
the exit hole are found several flattened, oval brown pellets of larval
140 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
meconium ; the shrivelled yellow to tan larval skin ; the transparent,
whitish cast pupal skin; usually a large mass of white adult
meconium; and numerous pieces of cocoon cut from the exit hole.
To separate the larvae of the species of Exenterus a perfect
mount is necessary, and even then identification is not always cer¬
tain (Finlayson, 1960). The cast skin of the final instar is covered
with minute needle-like spines and a few short setae. The narrow
epistoma of the cephalic structure (Fig. 78) is complete, but lightly
sclerotized in most specimens. The stipital sclerites are large and
bend dorsally along the margins of the U-shaped labial sclerite. The
blade of the mandible is long, slender and pointed. The spiracular
atrium (Fig. 109) is about as deep as it is wide, and tapers towards
its base.
Adults are black with yellow markings, and a greenish sheen to
the eyes in life. Living males can be separated readily from fe¬
males. Females have a single large hypopygial plate and males have
two, the anterior of which is margined with yellow giving the ap¬
pearance of a narrow yellow line across the black ventral apex of
the abdomen. In pinned specimens, collapse of the abdominal
sternites makes microscopic examination of the genitalia necessary
for proper sex determination. The sex ratio of reared specimens
was 6 females: 5 males.
Since becoming an important parasite of D. similis, E. amictorius
has become the focal host of the complex of hyperparasites found
associated with the sawfly, being attacked by no fewer than 10
species. The appearance of several native species of hyperparasites
not previously reared from D. similis cocoons may be the result
of the increased abundance of E. amictorius, the species with which
they have become commonly associated. Functioning as a multi¬
parasite, E. amictorius emerged from the same host cocoon with
M. dentipes, G. tenellus, D. fuscipennis, D. cavus, or T. scutellata.
Exenterus canadensis Provancher
Figs. 19, 20, 21, 79,110, 129
A rare parasite of D. similis in Wisconsin, E. canadensis has
been reared about a dozen times, and in recent years has been
encountered only once by dissection. The appearance and life his¬
tory are, in general, similar to E. amictorius.
The exit hole (Fig. 129) is similar to E. amictorius, as are the
remains left in the host cocoon, except that the silk of the parasite
cocoon is never pink. The larval skin is best separated from E. amic¬
torius by the shape of the spiracular atrium which is oval and
wider than deep (Fig. 110). In addition, well mounted cephalic
structures show both a lower epistomal arch, and shorter, more
robust lacinial sclerites (Figs. 78, 79).
1971] Mertins and Compel— Parasites of the Pine Saw fly 141
Figures 22-26. Adult hymenopterous parasites of D.
similis. 22, 23, Gelis tenellus; 22, head capsule,
frontal view; 23, female, lateral view. 24-26, Agro-
ther eutes lophyri; 24, head capsule, frontal view;
25, female, lateral view; 26, male abdomen, lateral
view.
Gelis tenellus (Say)
Figs. 22, 23, 81, 111, 130
G. tenellus is a common though not abundant parasite of D.
similis. It is a solitary, external feeder, and functions as either a
primary or secondary parasite. In three cases two individuals
emerged from the same host cocoon.
The exit hole (Fig. 130) is just to one side of the tip of the host
cocoon, round, smoothly cut, and 1.0-1. 3 mm in diameter. The
parasite cocoon adheres tightly to the inside of the host cocoon
along its length on the side where the exit hole occurs. It is about
one-fifth to one-fourth the volume of the host cocoon, thin, and
loosely woven of light gray silk which is shiny on both sides. Host
remains, which are excluded from the parasite cocoon, indicate that
frequently the host pupa is attacked, as well as the prepupa. A
142 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
fused clump of hard brown pellets of larval meconium is found
opposite the exit hole within the parasite cocoon, and lying loose
nearby is the pale, transparent pupal skin with its long ovipositor
sheath.
The pale, transparent final larval exuvium lies in close proximity
to, and often is tangled with, the pupal skin. It is covered with
minute bump-like protuberances and scattered setae. The cephalic
structure (Fig. 81) has a complete, unsclerotized epistoma; the
stipital sclerites are long, extending almost to the hypostomal arms ;
the blades of the mandibles are straight with two rows of fine
teeth; a widely arched labral sclerite turns down over the man¬
dibles; a U-shaped suspensorial sclerite occurs dorsally and poste¬
rior to the mandibles. The funnel-shaped spiracle has 10-12 annu-
lations (Fig. 111).
In the years before E. amictorius became prevalent, G. tenellus
was most always a primary parasite. Recently, however, it has
been found frequently as a secondary through E. amictorius. It
also parasitizes through /. conquisitor. In addition, six instances in
which both Gelis and E. amictorius emerged from the same cocoon
were encountered, and it also exhibited a similar multiparasitic
relationship with M. dentipes, S. hispae, I. conquisitor, T. scutellata,
and H. thyridopterigis. G. tenellus is the only parasite with two
dark bands on each forewing. Of nearly 600 specimens reared from
the sawfly, no males have been encountered.
Agrothereutes lophyri (Norton)
Figs. 24, 25, 26, 82, 112, 131
A. lophyri was reared consistently, although not in great numbers
from D. similis. It was always a primary, solitary, external parasite
of the sawfly within its cocoon.
The emergence hole (Fig. 131) is slightly off the tip of the host
cocoon, round with slightly ragged edges, and 1. 5-2.0 mm in diame¬
ter. The parasite cocoon is complete and nearly the size and shape
of the sawfly cocoon. It is white, papery, and brittle, and made up
of two to several layers. Its outside layer is rough and sometimes
has a brownish tinge, whereas the inside is smooth and shiny. The j
end of the parasite cocoon opposite the emergence hole contains a
layer of hardened red-brown larval meconium, in or on which lie
the pale whitish, shrivelled pupal skin, the yellowish shrunken
larval skin, and small particles of cocoon from the exit hole.
The head capsule of the larval skin is brown, and the skin is
covered with minute, sharp, conical spines and scattered setae.
The epistoma of the cephalic structure is unsclerotized but appears
complete (Fig. 82). The structure is similar to G. tenellus, but is
1971] Mertins and Coppel — Parasites of the Pine Saw fly 143
Figures 27-35. Adult hymenopterous parasites of
D. similis. 27-29, Dahlbominus fuscipennis ; 27, head
capsule, frontal view; 28, female, lateral view; 29,
male antenna, lateral view. 30, 31, Tetrastichus
coerulescens ; 30, head capsule, frontal view; 31, fe¬
male, lateral view. 32-35, Elasmus apanteli; 32, head
capsule, frontal view; 33, female, lateral view; 34,
male abdomen, lateral view; 35, male antenna,
lateral view.
larger, has stronger teeth on the mandibles, a sclerotized prelabium,
and widened dorsal arms on the labial sclerite. The spiracle has a
tapering reticulate atrium, a short stalk of two or three annuli,
and a large closing apparatus (Fig. 112).
The adults are readily identified by the wide red-brown to orange
band around the abdomen. Males differ from females in the lack
of an ovipositor (Fig. 26), and the absence of a white band around
each antenna. The sex ratio of 48 reared specimens was 3:1 in
favor of females.
144 Wisconsin Academy of Sciences , Arts and Letters ,[Vol. 59
Eulophidae
Dahlbominus fuscipennis (Zetterstedt)
Figs. 27, 28, 29, 88, 118, 132
The European D. fuscipennis is established on D. similis in Wis¬
consin, is reared regularly, but is never very numerous. In all
examined instances but one, the insect was a primary, gregarious,
external parasite of the sawfly prepupa. It attacks and emerges
from the host cocoon.
Emergence is most often via a hole (Fig. 132) slightly off one
end of the host cocoon, but the hole may be anywhere along the
side, and occasionally two holes are cut at opposite ends. The hole
is irregularly round and about 0.6-0. 7 mm in diameter. No parasite
cocoons are present. The host remains are shrivelled and frequently
unrecognizable. Careful dissection usually reveals that the host
remains are located in the center of the cocoon, surrounded on all
sides by the parasite remains. This arrangement is seen most easily
before emergence, when the parasite larvae are lined up against
the walls facing in one direction before pupation. After emergence
the inside of the cocoon is a dishevelled mass of shiny, golden-
yellow broken pupal skins, and dead larvae. The larval meconium
is a hard dark-brown to gray mass occasionally made up of small
ovoid pellets.
The minute last larval skin is often found adhering to the
meconial mass, or sometimes to the posterior end of the pupal skin.
Because of their small size it is sometimes easier to use a dead
larva for identification purposes rather than a skin. One or two
dead larvae are commonly found in the cocoon, and may be recog¬
nized by their shiny brown color, and the fact that broken pieces
of pupal skins frequently adhere to them. The larvae are softened
in KOH, punctured with a needle, and the contents squeezed out
before mounting in the normal manner. Mounted skins are smooth
and without structure other than mandibles, antennae and spiracles.
The mandibles are triangular with a long, straight blade (Fig. 83).
A row of minute teeth along the blade are usually very difficult to
discern. The antennae are dome-like with two sensoria each. The
spiracle is funnel-shaped, with at least ten chambers and a long
stalk (Fig. 113).
D. fuscipennis is usually a primary parasite, but in one instance
it apparently developed hyperparasitically through an A. lophyri
larva. In one other case both D. fuscipennis and E. amictorius
emerged from the same cocoon. The average number of individuals
emerging per cocoon was 28 (range 3-59), and the overall sex
ratio of insects reared was 4.3 females:! male. Males differ from
females in the shape of the antennae (Fig. 29), and their nearly
hyaline forewings.
1971] Merlins and Coppel — Parasites of the Pine Sawfly 145
Figures 36-39. Adult hymenopterous parasites of
D. similis. 36, 37, Eupelmus spongipartus ; 36, head
capsule, frontal view; 37, female, lateral view. 38,
39, Eupelmella vesicularis ; 38, head capsule, frontal
view; 39, female, lateral view.
Tetrastichus coerulescens Ashmead
Figs. 30, 31, 84, 114, 133
The hyperparasite, T. coerulescens, was encountered only twice
during the study, and has not been reported previously from D.
similis.
In the single example of emergence available, 2 exit holes were
cut in the host cocoon on the side near one end. Probably 1 hole is
cut normally, as this is the usual case in other parasites which
develop gregariously on the sawfly. The hole (Fig. 133) is oblong
(.47 x .40 mm). The chalcid pupae parasitized by T. coerulescens
turned dark brown as opposed to the orange color of unparasitized
pupae. Emergence from the host pupa is characteristically via a
large irregular round hole cut in the ventral portion of the abdo¬
men, usually completely destroying the abdominal tergites. The
host pupa is hollow except for a dark, shiny gray or brown deposit
of larval meconium in the head capsule, and the yellow parasite
larval skin and yellow-brown pupal skin which lie on the meconium.
The skin of the final instar is extremely small and thread-like.
Under magnification the only visible features are the spiracles and
146 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
the cephalic structure (Fig. 84), which has an incomplete epistoma,
large and prominent superior mandibular processes, and a ventral
sclerotized bridge apparently formed from the fused inferior man¬
dibular processes. The mandibles are bulbous ventrally and lack
teeth on the blades. The spiracle (Fig. 114) has 15-18 chambers and
a small closing apparatus.
When this insect emerged from a D. similis cocoon it was a sec¬
ondary parasite through A. verditer. Three female parasites
emerged from the cocoon, each having matured as an internal soli¬
tary parasite of a single pupa of A. verditer. The second occurrence
of T. coerulescens was revealed by dissection, and was part of the
most complex association of insects found in a single sawfly cocoon.
Dissection of a cocoon which had produced two adult female M.
dentipes, revealed in addition to the M. dentipes and sawfly re¬
mains, those of an E. amictorius larva, one adult female H. thyri-
dopterigis, one dead H. thyridopterigis pupa, and one adult female
T. coerulescens. In this instance T. coerulescens was a tertiary para¬
site through H. thyridopterigis on E. amictorius. T. coerulescens
was the smallest parasite reared from D. similis and can be identi¬
fied easily by the blunt-tipped shape of the wings (Fig. 31). No
males were encountered, but according to Burks (1943) they differ
from the female both in having long setae on the antennal funicle
segments, and in having the first segment of the funicle only three-
fourths as long as the second.
Elasmus apanteli Gahan
Figs. 32, 33, 34, 35, 85, 115, 134
E. apanteli is a rare parasite of D. similis cocoons. It was en¬
countered only twice, during the late summer of 1968, once by
emergence and once by dissection. It is a gregarious secondary
parasite through E. amictorius. This is a new host record.
The nearly circular exit hole (Fig. 134) is located at the tip of
the host cocoon. It is about 0.6 mm in diameter and the edges are
slightly scalloped. No parasite cocoons are present. The contents
of the sawfly cocoon are the same as those described for the pri¬
mary parasite, E. amictorius. The E. amictorius pupa is shrivelled
and dark brown. The Elasmus remains are within the Exenterus
cocoon, and include several dark brown pupal cases, which are vir¬
tually intact, save the head and pro- to mesothoracic region; sev¬
eral dead larvae or pupae; clusters of small, brown ovoid larval
meconium pellets ; and the pale yellow, thread-like larval exuvium,
which is usually attached to the posterior tip of the pupal case
abdomen.
The mounted exuviae (Fig. 85) closely resemble those of D. fusci-
pennis. In general, the row of fine teeth along the blade of the man-
1971] Merlins and Copp el— Parasites of the Pine Sawfly 147
Figures 40-46. Adult hymenopterous parasites of
D. similis. 40-42, Monodontomerus dentipes; 40,
head capsule, frontal view; 41, female, lateral view;
42, male abdomen, lateral view. 43-46, Amblymerus
verditer; 43, head capsule, frontal view; 44, female,
lateral view; 45, male abdomen, lateral view; 46,
male antenna, lateral view.
dible is more distinct in E. apanteli, and in addition, the postero-
ventral process of the mandible is much longer than the dorsal
process as compared to those of D. fuscipennis which are about
equal in length. Perhaps the easiest separation may be made on the
basis of the color of the cast pupal skins which are brown in
Elasmus and golden-yellow in Dahlbominus.
The recent rearings of E. apanteli from D. similis cocoons may
be the result of the increasing importance of its primary host,
E. amictorius, in recent years. In addition to E. apanteli, Catolaccus
cyanoideus and Tritneptis scutellata have also been found recently,
and nearly exclusively, associated with E. amictorius. The adults
of E. apanteli are distinctive (Fig. 33) . The hind coxa is enormously
148 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
enlarged and flattened and the tarsus is elongate and four seg¬
mented. Males may be distinguished by the shape of the antennae
(Fig. 35). The sex ratio of reared adults was 5 females :1 male.
Eupelmidae
Eupelmus spongipartus Foerster
Figs. 36, 37, 86, 116, 135
One female E. spongipartus has been reared from D. similis in
Wisconsin, and it was a solitary, secondary cocoon parasite through
D. cavus. It is undoubtedly an insignificant member of the parasite
complex.
The exit hole (Fig. 135) is on the side of the host cocoon near
the tip, round, smooth, and 1.0 mm in diameter. No cocoon is spun,
but the small, white, fibrous egg mat noted by Muesebeck and
Dohanian (1927) is attached to the inside of the host cocoon. The
parasite remains are loose within the cocoon and consist of a group
of dark red-brown ovoid pellets of larval meconium; the small,
pale yellow larval skin which is covered with long setae; and the
fragile, golden-yellow pupal skin, whose abdomen retains its shape.
The cast skin of the last instar is smooth except for a sparse
covering of long setae. The most prominent feature of the cephalic
structure is the large, crescent-shaped, 8-toothed clypeus (Fig. 86).
The inferior mandibular articulations occur as depressions on a
lightly sclerotized, transverse ventral bar, and the superior articu¬
lations (one of which is hidden in Fig. 86) are separate and lightly
sclerotized as well. The mandible is large and sclerotized, bulbous
basally, and has a curved toothless blade. The spiracle (Fig. 116)
is funnel-shaped with 14-19 chambers, a prominent closing appara¬
tus, and an annulated stalk. An irregular line occurs around the
first chamber.
Eupelmella vesicularis (Retzius)
Figs. 38, 39, 87, 117, 136
Reared in small numbers every year, this insect can act as
either a primary or secondary parasite of D. similis cocoons. It
is a solitary, external feeder.
The exit hole (Fig. 136) is on the side of the host cocoon near
the tip, is smooth and oblong (0.8 x 1.0 mm) . As in E. spongipartus
no cocoon is spun, but one to several white fibrous mats (0.6 x
0.3 mm) are found on the inside of the host cocoon. Clausen
(1940) believed these fibrous mats which cover the eggs serve as
protection from primary parasites upon which the developing
Eupelmella larvae will prey. The parasite remains are similar to
those of E. spongipartus except that they are slightly smaller and
the last larval exuvium is darker yellow.
1971] Mertins and Coppel — Parasites of the Pine Sawfly 149
Figures 47-52. Adult hymenopterous parasites of
D. similis. 47-49, Tritneptis scutellata; 47, head cap¬
sule, frontal view; 48, female, lateral view; 49, male
antenna, lateral view. 50-52, Dibrachys cavus; 50,
head capsule, frontal view; 51, female, lateral view;
52, male abdomen, lateral view.
The sparsely setate skin of the final instar presents features
nearly identical to E. spongipartus under magnification. However,
the clypeus usually has only five well-developed teeth (Fig. 87)
with two or three imperfect ones laterally ; the inferior mandibular
articulations are less sclerotized and more distinctly separated
from each other, and the superior articulations are not discernible.
Contrary to the findings of Finlayson (1962), in our material it
was not possible to distinguish E. spongipartus from E. vesicularis
by the number of chambers in the spiracular atrium. In both species
the number ranged from 14-19, although the prominent wavy line
around the first chamber in spongipartus is not as evident in vesicu¬
laris (Fig. 117). In any case, because of the rarity of E. spongipar¬
tus , one can be fairly certain that a parasite exhibiting a cephalic
structure such as that in Figs. 86 or 87 is E. vesicularis.
E. vesicularis is most commonly secondary, especially through
D. fuscipennis , as noted by Morris (1938), but also matured on a
150 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
number of other parasite species, and in a number of instances
was a primary parasite. In 2 instances 2 individuals of this species
emerged from a host cocoon, both utilizing the same exit hole, and
in 3 cases an E, vesicularis and several M. dentipes emerged from
the same sawfly cocoon. The E. vesicularis had functioned as a
secondary parasite, but had apparently failed to destroy all of the
primary parasites within the cocoon before completing its develop¬
ment. No males of this species are known (Gahan, 1933) .
Torymidae
Monodontomerus dentipes (Dalman)
Figs. 40, 41, 42, 88, 118, 137
M. dentipes is one of the two commonest parasites of D. similis
in Wisconsin. It is nearly always reared from every collection of
cocoons made in the field, and is frequently observed in the process
of oviposition. It is a gregarious, external parasite of D. similis,
attacking and emerging from the cocoon.
The exit hole is usually located slightly to one side of the tip of
the host cocoon, and is round with jagged edges (Fig. 137). It is
normally 1.1 mm in diameter, but may be as large as 1.3 mm, or as
small as 0.6 mm if only small males are produced. In about 3%
of the cocoons examined, 2 emergence holes were cut, and occasion¬
ally the hole is cut on the side of the cocoon. The arrangement of
the inside of the cocoon is similar to that left by D. fuscipennis,
with the host remains and meconia of the parasites in the center,
and the parasite pupal and larval skins layered against the walls.
This is a result of the tendency of the mature parasite larvae to
line up in a single layer around the host cocoon walls all facing in
one direction before pupation. There are no cocoons. The larval
meconium consists of numerous single or massed brown or black
ellipsoid pellets located usually at the end opposite the exit hole.
The yellow larval skins are covered with long hairs, and are loose
within the cocoon. The ovipositor sheath of female pupal skins
is long and conspicuously curved antero-dorsally. Many times the
flattened, banana-shaped egg chorions adhere to the host remains.
The skin of the last instar is smooth, but densely covered with
long (0.4 mm) setae. The cephalic structure (Fig. 88) has an
incomplete epistoma, short hypostomal arms, large superior man¬
dibular processes and greatly elongate inferior processes upon
which the mandibles rest. The mandibles have a straight blade
without teeth. The antennae are prominent dome-like structures
each bearing two sensoria apically. The spiracle (Fig. 118) is
large, reticulate, funnel-shaped with thick walls, and has 8-10
chambers.
1971] Mertins and Coppel — Parasites of the Pine Saw fly 151
Figures 53-58. Adult hymenopterous parasites of
D. similis. 53-55, Catolaccus cyanoideus; 53, head
capsule, frontal view; 54, female, lateral view; 55,
male abdomen, lateral view. 56-58, Habrocytus thy -
ridopterigis ; 56, head capsule, frontal view; 57, fe¬
male, lateral view; 58, male abdomen, lateral view.
The adults of M. dentipes are large and robust. They are dark
green with red eyes. Males are distinguished easily by the absence
of an ovipositor (Fig. 42) . The average number of adults emerging
from a host cocoon was 8 (range 1-82), and the sex ratio was 1.1
females : 1 male.
The host relations of M. dentipes are complex. Dissections indi¬
cate that the insect is most often a primary parasite, but it was
found as a secondary through I. conquisitor, E. amictorius , D. fusci-
pennis, and D. lophyri. This relationship is probably facultative.
In other instances it was the successful competitor in a multipara-
sitic situation, although it apparently did not act as a secondary.
Instances of secondary and multiparasitism by this species are most
common in second generation sawfly cocoons which are most heavily
attacked by the large numbers of M. dentipes present in the fall.
152 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
The insect seems to be a successful multiparasite more often than
it is unsuccessful, for it is seldom found dead in cocoons attacked
by other species. M . dentipes has emerged from the same cocoons
with the following species : E. amictorius, E. vesicularis, G. tenellus,
H. thyridopterigis, or A. verditer.
Pteromalidae
Amhlymerus verditer (Norton)
Figs. 43, 44, 45, 46, 89, 119, 138
A. verditer was reared sporadically, but not in great numbers
from D. similis. It is a gregarious, external feeder in the cocoon,
and in nearly every case acted as a primary parasite.
The exit hole is on the side of the host cocoon, often near the tip,
but sometimes in the middle. Occasionally more than 1 hole is
cut. The hole (Fig. 138) is small, round and regular, and 0.8 mm
in diameter. No parasite cocoons are present. The parasite remains
include numerous small shiny brown to black pellets of larval
meconium, numerous tiny white larval skins resembling twisted
threads, and fragmented shiny golden-yellow pupal skins, all loose
within the cocoon. Dead and shrivelled parasite larvae and pupae
are sometimes present. All 5 members of the Pteromalidae which
parasitize D. similis leave remains similar to those described above,
and therefore cannot be separated from each other without making
slides.
The last larval skin is nearly featureless, except for a few
inconspicuous widely scattered setae, the spiracles, and the cephalic
structure which consists only of the mandibles and lightly sclero-
tized articulations (Fig. 89). The antennae are nearly as large as
the blades of the mandibles, though not as pointed, and are without
antennal sockets. The spiracle (Fig. 119) is funnel-shaped with
5-8 chambers, some of which appear subdivided, and has a narrow
closing apparatus.
Although commonly a primary parasite, A. verditer occasionally
was found to develop at the expense of E. amictorius. In one
instance, individuals of both A. verditer and M. dentipes emerged
from the same host cocoon. The adult is recognized most easily by
its coloration which is brilliant metallic green in the female, and
metallic yellow-green with a yellow band around the abdomen in
the male. In addition, the male abdomen (Fig. 45) is shorter and
narrower than that of the female. The scape of the male antenna
(Fig. 46) is thicker than that of the female, and shows an anterior
depression. An average of 22 individuals emerged per cocoon
(range 6-40), and the sex ratio of reared individuals was 2.6
males: 1 female.
1971] Merlins and Coppel — Parasites of the Pine Sawfly 153
Figures 59-66. Adult hymenopterous parasites of
D. similis. 59-62, Eurytoma pini; 59, head capsule,
frontal view; 60, female, lateral view; 61, male ab¬
domen, lateral view; 62, male antenna,, lateral view.
63-66, Spilochalcis alhifrons; 63, head capsule, fron¬
tal view; 64, female, lateral view; 65, male abdomen,
lateral view; 66, male antenna, lateral view.
Tritneptis scutellata (Muesebeck)
Figs. 47, 48, 49, 90, 120, 132
T. scutellata was reared from 16 host cocoons collected during
the summers of 1967 and 1968. D. similis has not been previously
reported as its host. It is usually a secondary gregarious parasite,
attacking and emerging from the sawfly cocoon.
The emergence hole is on the side of the host cocoon near the
tip, round, and has slightly irregular edges (Fig. 132). It is 0.5-0. 6
mm in diameter. In 2 cases more than 1 hole was used in emerging.
No parasite cocoons are formed, and the other remains are similar
to those described for A. verditer.
The exuvium of the last instar also resembles that of A. verditer,
but may be separated by the cephalic structure and the spiracles.
154 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
The cephalic structure (Fig. 90) has, in addition to the mandibles,
an incomplete epistoma, pleurostoma, hypostoma, superior and
inferior mandibular processes, and shorter antennae. The atrium
of the spiracle has but 4-5 chambers (Fig. 120). It is especially
difficult to separate the parasite remains of T. scutellata from those
of D. cavus, but the characters used in the key should allow reason¬
able certainty in most cases.
T. scutellata is usually a secondary parasite, and in most cases
was found to be associated with E. amictorius. In 2 cases both
T. scutellata and E. amictorius adults emerged from the same
cocoon ; in 10 cocoons T. scutellata developed as a secondary para¬
site through E. amictorius, and from 1 of these an adult G. tenellus
also emerged. In 4 other cocoons, T. scutellata was secondary
through I. conquisitor in 1, shared a sawfly host with A. lophyri
in another, and was a primary parasite in the others.
The average number of individuals emerging per cocoon was 8
(range 3-15), and the sex ratio of reared specimens favored fe¬
males 3:1. Males differ from females in their smaller abdomen
and broader antennal scape (Fig. 49).
Dihrachys cavus (Walker)
Figs. 50, 51, 52, 91, 121, 139
D. cavus is an occasional parasite of D. similis, with only a few
individuals reared each year. It is a gregarious, external parasite
in the cocoon, and most often functions as a primary.
The exit hole is as in A. verditer, 0.7-0. 8 mm in diameter, and
may have scalloped edges (Fig. 139). More than 1 hole is sometimes
cut. The contents of the cocoon are similar to those described for
A. verditer . The skin of the last instar is similar to A. verditer
and the cephalic structure (Fig. 91) and spiracle (Fig. 121) re¬
semble those of T. scutellata. The characters used in the key are
sufficient to separate these species in the majority of cases.
D. cavus is usually a primary parasite, but in several cases it was
found to develop at the expense of E . amictorius . In 1 instance both
D. cavus and E. amictorius emerged from the same cocoon. An
average of 12 individuals emerged per cocoon (range 1-24) , and
the sex ratio of reared insects was 17.3 females : 1 male. The adult
is dark green, approaching black, and a light yellow band encircles
the abdomen of the male. The male abdomen (Fig. 52) is shorter
and narrower than that of the female.
Catolaccus cyanoideus Burks
Figs. 53, 54, 55, 92, 122, 140
C. cyanoideus was encountered 12 times, all from cocoons col¬
lected during the summer of 1967. It was always a solitary sec-
1971] Merlins and Coppel — Parasites of the Pine Saw fly 155
Figures 67-70. Adult dipterous parasites of D. simi-
lis. 67, 68, Spathimeigenia spinigera; 67, head of
male, dorsal view; 68, female,, dorsal view. 69, 70,
Bessa harveyi ; 69, head of male, dorsal view; 70,
female, dorsal view.
ondary parasite through E. amictorius, a previously unreported
host.
The exit hole (Fig. 140) is on the side of the host cocoon
usually near the tip, but in one case was located in the middle of
the side. It is round with slightly scalloped edges, and for the
females was 0. 9-1.0 mm in diameter. The exit hole of males was
0.7 mm in diameter. No parasite cocoon is formed. The parasite
remains include the dark brown, shiny pupal skin which remains
with little breakage, usually only the head and part of the thorax
being destroyed. The last larval exuvium is often still attached to
the dorso-posterior end of the pupal skin at one end, and to the
hard, dark-brown, pellet-like meconial mass of the parasite larva
at the other.
156 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
The last exuvium is smooth and bears some scattered setae
0.02-0.05 mm long. The cephalic structure (Fig. 92) is similar to
A. verditer, and the spiracle (Fig. 122) resembles that of H. thy -
ridopterigis except for the lack of prominent lines on the first
chamber of the atrium.
The disproportionately large head is an obvious characteristic
of the metallic blue-green adult (Fig. 54). The cheek area on
either side of the head is excavated. The sex ratio of the specimens
reared was 5 females: 1 male. Males are about one-half the size
of females and have a proportionately smaller abdomen (Fig. 55).
Hahrocytus thyridopterigis Howard
Figs. 56, 57, 58, 93, 123, 141
H. thyridopterigis is an occasional parasite of D. similis and has
never been reared more than 4 or 5 times in a season. It is a
gregarious external parasite in the host cocoon, and is usually
primary.
The exit hole (Fig. 141) is similar to A. verditer, but slightly
larger, and is 0. 7-0.9 mm in diameter. The contents left in parasi¬
tized cocoons are like those left by Amhlymerus, Dihrachys, and
Tritneptis, except that the average sizes of the parasite remains
are somewhat larger, and the pupal skins are yellow-brown.
The cast skin of the last instar may be differentiated from the
aforementioned species by the lack of the partially developed
cephalic structures found in D. cavus and T. scutellata, and the
presence of large antennae set in usually distinct antennal sockets
(Fig. 93). In addition, the row of fine teeth on the blade of the
mandible is usually more distinct than in the other species of
pteromalids. The spiracle (Fig. 123) is usually reticulate apically,
and the closing apparatus is usually shorter and broader than in
Amhlymerus, Dihrachys, and Tritneptis.
Although normally primary, H. thyridopterigis was reared sev¬
eral times as a secondary parasite through I. conquisitor. In one
instance, both it and G. tenellus emerged from the same cocoon.
Other cases of successful multiparasitism involving H. thyridop¬
terigis included G. tenellus (with both species as primary para¬
sites), M. dentipes, and E. vesicularis. The hyperparasitic rela¬
tionship of H. thyridopterigis to I. conquisitor is apparently com¬
mon (Cushman, 1927; Langston, 1957; Balduf, 1937).
The adults are metallic green, and slightly larger than the other
pteromalids, except the equally large C. cyanoideus. An average
of 5 adults emerged per cocoon (range 1-22), and the sex ratio
of reared adults was 10.6 females: 1 male. In its role as a sec¬
ondary parasite only 1 or 2 adults were produced per cocoon, but
as a primary, the average was 6 per cocoon.
1971] Merlins and Coppel — Parasites of the Pine Sawfly 157
Figures 71-74. Adult dipterous parasites of D. simi-
lis. 71, 72, Diplostichus lophyri; 71, head of male,
dorsal view; 72, female, dorsal view. 73, 74, Eupho-
rocera edwardsii; 73, head of male, dorsal view; 74,
female, dorsal view.
Eurytomidae
Eurytoma pini Bugbee
Figs. 59, 60, 61, 62, 94, 124, 142
E. pini is a rare parasite of D. similis in Wisconsin, only 6 hav¬
ing been reared from a single study plot during 1958 and 1959.
It is a primary, solitary parasite of the prepupa in the cocoon.
The emergence hole is cut in the side of the host cocoon, fre¬
quently near one end (Fig. 142). It is round, smoothly cut, and
0.9-1. 2 mm in diameter. There is no parasite cocoon. The parasite
remains consist of a mass of hard brown globular meconial pellets,
a yellow larval skin with a sparse covering of long hairs, and a
shiny golden-yellow pupal skin, all loose within the host cocoon.
158 Wisconsin Academy of Sciences, Arts and Letters [Vol. 59
The setae of the last larval skin are about 0.2 mm long, and are
scattered sparsely over its surface. The cephalic structure (Fig. 94)
has an incomplete epistoma, large superior mandibular processes,
long inferior mandibular processes, and narrow trailing hypostomal
arms. The mandibles have curved blades and a large medial tooth.
The antennae are conspicuously sclerotized. The large funnel-
shaped spiracle (Fig. 124) has about 15 chambers and a distinctive
closing apparatus. The apical chamber is reticulate.
The adult (Fig. 60) is robust and shiny black, and is the only
parasite of D. similis with 10 segmented antennae. Males differ
from females by their pediculate abdomen (Fig. 61) and their
unusual setate antennae (Fig. 62). The sex ratio favored females
2:1.
Chalcidae
Spilochalcis albifrons (Walsh)
Figs. 63, 64, 65, 66, 95, 125, 143
S. albifrons is a rare, and apparently not very successful para¬
site of D. similis; it has not been previously reported from the saw-
fly. In all, 11 adults and 1 larva have been obtained from D. similis
cocoons : 10 by dissection and 2 by emergence. All adults were fe¬
males, and all were solitary, primary parasites. The larva was a
secondary through E. amictorious.
The emergence hole (Fig. 143) is slightly off the tip of the host
cocoon, round with irregular edges, and is about 1.3 mm in diame¬
ter. There is no parasite cocoon. The parasite remains consist
of 1 large sub-spherical brown meconial deposit opposite the exit
hole, the tan or light brown larval skin which is almost always
attached to the posterior end of the dark brown, and shiny pupal
skin, and numerous fragments of cocoon cut from the opening.
The cast skin of the final larval instar is smooth except for 2 or 3
short inconspicuous setae. The cephalic structure (Fig. 95) has
an incomplete epistoma, well-developed superior mandibular proc¬
esses, long narrow inferior mandibular processes, and very long
hypostomal arms. An indistinct silk press is present, and the
lightly sclerotized labral sclerite is vacuolate at the ends. The long
sharp blades of the mandibles show faint indications of a row of
small teeth. Antennae could not be located. The spiracle (Fig.
125) has about 7 annulations, and is widest at the second and
narrowest at the fifth annulation. The closing apparatus is large
and marked with a dark ring.
It appears that the adult parasites have little success in cutting
through the tough silk of the host cocoon for emergence, as only
2 of the 12 individuals encountered managed to escape normally.
1971] Mertins and Coppel — Parasites of the Pine Sawfly 159
Figures 75-82. Cephalic structures of final instar
ichneumonids ; frontal view. 75, Scambus (Scambus)
hispae; 76, Delomerista japonica; 77, Delomerista
novita; 78, Exenterus amictorius; 79, Exenterus
canadensis; 80, Itoplectis conquisitor; 81, Gelis tenel-
lus; 82, Agrothereutes lophyri.
The black and yellow marked adults are easily recognized by the
greatly enlarged, toothed metafemur. Two males of this species
from another host were examined, and can be easily distinguished
by their small abdomen (Fig. 65) which is about the size of the
metafemur, and their swollen antennal scape (Fig. 66).
160 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Diptera
Tachinidae
Spathimeigenia spinigera Townsend
Figs. 67, 68, 96, 101, 144
S. spinigera is a rare parasite of D. similis in Wisconsin, having
been reared only twice. It has not been previously reported from
this sawfly. It is a primary, solitary parasite, attacking the larva
and emerging from the cocoon.
The emergence hole is on the tip of the host cocoon (Fig. 144),
is nearly round, and about 2 mm in diameter. The thin, tapered
edges are ragged and pushed outward. The host remains consist
of little more than a hollow integument appressed to the lateral
wall of the cocoon. Next to the host is the puparium of the parasite,
about 6 mm in length, red-brown, semi-transparent, and brittle.
The end of the puparium near the exit hole is broken into two semi¬
circles and the inside is lined with a thin whitish membrane.
The mouth-hooks of the third instar larva are attached securely
to one of the broken semi-circles under the membranous lining.
The buccopharyngeal armature (Fig. 96) has heavily sclerotized
mandibular hooks with a strong antero-dorsal curve, and postero-
dorsal and ventral points. The intermediate sclerite is distinct and
heavily sclerotized. The basal sclerite is heavily sclerotized ante¬
riorly, but its dorsal and ventral wings are less so, the lower trailing
portion of the ventral wings becoming membranous. The posterior
spiracular plates of the puparium (Fig. 101) are sharply elevated,
wedge-shaped, and each is surrounded by a heavily sclerotized band.
The adult insect is most readily identified by the reddish mark¬
ings on the tip of the abdomen and on the legs. The pulvilli and
tarsal claws are decidedly longer than the fifth tarsal segment in
the male, but of equal length in the female. Orbital bristles are
lacking (Fig. 67) in the male, and present (Fig. 68) in the female.
Bessa harveyi (Townsend)
Figs. 69, 70, 97, 98, 102, 145
Bessa harveyi is an uncommon parasite of D. similis in Wiscon¬
sin, having been obtained only 16 times. D. similis is apparently one
of the species “occasionally” attacked by B. harveyi (Turnock and
Melvin, 1963), but has not been reported previously as a host in
the literature. It is a solitary, primary parasite, attacking the
host larva and emerging from the cocoon. In about half of the
instances observed, the mature parasite larva emerged from the
host cocoon before forming its puparium.
The oval emergence hole (Fig. 145) is on the tip of the host
cocoon and about 1.3 mm in diameter. Some of the edges may
1971] Mertins and Copp el— Parasites of the Pine Sawfly 161
Figures 83-95. Cephalic structures of final instar
chalcidoids; frontal views. 83, Dahlbominus fusci-
pennis; 84, T etrastichus coerulescens ; 85, Elasmus
apanteli; 86, Eupelmus spongipartus ; 87, Eupel -
mella vesicularis; 88,, Monodontomerus dentipes ; 89,
Amblymerus verditer; 90, Tritneptis scutellata; 91,
Dibrachys cavus ; 92, Catolaccus cyanoideus; 93,
Habrocytus thyridopterigis ; 94, Eurytoma pini; 95,
Spilochalcis albifrons.
appear slightly pushed out. When the parasite larva emerges from
the cocoon to pupate, the only remains are the mouth-hooks and
exuvia of the first and second instars. The buccopharyngeal appa¬
ratus of the second instar (Fig. 97) is 0.5 mm long and is fused
into a single structure. The mandibular hooks are long, narrow and
sharp, the dorsal wings are elongate and less sclerotized posteriorly,
and the ventral wings are short and simple. A small salivary
sclerite occurs below the intermediate region.
162 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Figures 96-104. Remains of immature dipterous
parasites. 96-100, Buccopharyngeal apparatuses,
lateral views; 96, Spathimeigenia spinigera, third
instar; 97, Bessa harveyi, second instar; 98, Bessa
harveyi, third instar; 99, Diplostichus lophyri, third
instar; 100, Euphorocera edwardsii, third instar.
101-104, Right posterior spiracular plates of third
instar dipterous larvae; 101, Spathimeigenia spini¬
gera; 102, Bessa harveyi; 103, Diplostichus lophyri;
104, Euphorocera edwardsii.
When the parasite pupates within the host cocoon, the remains
include the puparium in addition to those remains described previ¬
ously. The puparium is formed with its anterior end toward the
exit hole, and lies side by side with the host remains. It is similar
in appearance to that of S. spinigera , but is only 5 mm in length.
A creamy-white meconium is usually present in its posterior end.
The mouth-hooks are attached to the open end of the puparium
under the membranous lining-.
The buccopharyngeal armature of the third instar (Fig. 98) is
separated into three distinct parts. The mandibular hooks are
1971] Mertins and Coppel — Parasites of the Pine Saw fly 163
Figures 105-125. Spiracles of final instar Hymenop-
tera. 105, Scambus (Scambus) hispae; 106, Delome-
rista japonica ; 107, Delomerista novita; 108, Itoplec -
tis conquisitor ; 109, Exenterus amictorius ; 110, Ex -
enterus canadensis; 111, Gelis tenellus; 112, Agro-
thereutes lophyri; 113, Dahlbominus fuscipennis;
114, Tetrastichus coerulescens; 115, Elasmus apan-
teli; 116, Eupelmus spongipartus ; 117, Eupelmella
vesicularis ; 118, Monodontomerus dentipes; 119,
Ambly menus verditer; 120, Tritneptis scutellata;
121, Dibrachys cavus; 122, Catolaccus cyanoideus;
123, Habrocytus thyridopterigis ; 124, Eurytoma
pini; 125, Spilochalcis albifrons.
heavily sclerotized, curved on the antero-dorsal margin, and are
equipped with postero-dorsal and ventral processes. The intermedi¬
ate sclerite has a large posterior lobe. The basal sclerite is heavily
sclerotized anteriorly and lightly so posteriorly on both the dorsal
and ventral wings. Two small sclerites are present below the
164 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
intermediate sclerite. The posterior spiracular plates (Fig. 102)
are slightly elevated and each is completely surrounded by a black
ring of roughly circular form.
The sex ratio of adults reared was 1.7 females: 1 male. Adult
B. harveyi are the smallest dipterous parasites of D. similis, and
are quickly recognized by their size, dark-brown palpi, and the
lack of a pair of cruciate bristles on the apex of the scutellum.
Males are separated most easily from females by the absence of
orbital bristles (Fig. 69).
Diplostichus lophyri (Townsend)
Figs. 71, 72, 99, 103, 146
D. lophyri was by far the commonest dipterous parasite of
D. similis in Wisconsin. It was reared in small numbers every year.
The insect attacks late instar larvae and is a primary, solitary para¬
site. It emerges from the cocoon.
The distinctive emergence hole is at the tip of the host cocoon
(Fig. 146). It is round with smoothly cut edges, and a flap that
remains attached by a small hinge of uncut silk. The hole is 2.4
mm in diameter. Within the cocoon the host remains lie next to
the puparium which faces the exit hole. The puparium is 6 mm
long and similar to those described for S. spinigera and B. harveyi.
The buccopharyngeal armature (Fig. 99) of the third instar is
attached to the inner open end of the puparium. It consists of the
heavily sclerotized mandibles which are completely fused to the
intermediate sclerite, and the basal sclerite which is closely articu¬
lated with, but separate from, the intermediate sclerite. The
mandibular hooks are broad and have a postero-ventral projection.
The heavily sclerotized intermediate region also has a ventral
process. The basal sclerite is strongly sclerotized centrally, but
becomes less so posteriorly on its dorsal and ventral wings. The
ventral wings show an antero-dorsal projection. The posterior
spiracular plates (Fig. 103) are large, with only the areas immedi¬
ately surrounding the spiracular openings being elevated. The
spiracular openings are long and narrow, and the black ring sur¬
rounding each plate is incomplete medially.
Probably any housefly-sized Diptera reared from D. similis can
be safely called D. lophyri because the other Diptera are so uncom¬
mon. Specifically the insect may be identified by the pronounced
distal recurvation of vein Mi+2 (Fig. 72) and the pair of enlarged
hairs on the mesoscutellar disc. Males lack the orbital bristles of
females, and also have greatly elongate tarsal claws and pulvilli.
In one instance, a puparium of D. lophyri served as a food source
for several individuals of M. dentipes.
1971] Mertins and Copp el— Parasites of the Pine Saw fly 165
Figures 126-149. D. similis cocoons showing sawfly
and parasite emergence holes, and holes made by
predators. 126, Scambus (Scambus) hispae; 127,
Delomerista japonica or D. novita ; 128, Itoplectis
conquisitor; 129, Exenterus amictorius or E. cana¬
densis ; 130, Gelis tenellus; 131, Agr other eubes
lophyri; 132, Dahlbominus fuscipennis or Tritneptis
scutellata; 133, Tetrastichus coerulescens ; 134,
Elasmus apanteli; 135, Eupelmus spongipartus ;
136, Eupelmella vesicularis ; 137, Monodontomerus
dentipes; 138, Amblymerus verditer; 139, Dibrachys
cavus; 140, Catolaccus cyanoideus; 141, Habrocytus
thyridopterigis ; 142, Eurytoma pini; 143, Spilochal-
cis albifrons ; 144, Spathimeigenia spinigera; 145,
Bessa harveyi; 146, Diplostichus lophyri; 147,
Diprion similis; 148, bird predation; 149, rodent
predation.
Euphorocera edwardsii (Williston)
Figs. 73, 74, 100, 104
E. edwardsii is a rare, and apparently unsuccessful parasite of
D. similis. It was encountered 13 times in Wisconsin (8 of these
in 1967), and then only by dissection. No adults have been reared.
166 Wisconsin Academy of Sciences , Arts and Letters [Vol. 59
Since no adults of the species emerged, the position and appear¬
ance of the exit hole are unknown. It is probably large, and because
of the near relationship of the species to D. lophyri, it may have
a hinged cap. Pupation probably occurs within the host cocoon as
with D. lophyri.
The buccopharyngeal armature of the third instar is large and
heavily sclerotized (Fig. 100). The mandibular hooks are long,
slender, and show a definite constriction at their juncture with
the intermediate sclerite. They have both dorsal and ventral proc¬
esses posteriorly. A small sclerite projects anteriorly below the
intermediate region. The basal sclerite is freely articulated with the
intermediate sclerite, and is heavily sclerotized anteriorly. The
posterior portions of both the dorsal and ventral wings are lightly
sclerotized. The ventral wings have an anterior projection. The
posterior spiracular plates of the third instar larva (Fig. 104)
are large, elevated, and somewhat irregular in outline. Each is
surrounded incompletely by a black ring. The spiracular openings,
especially the two outside ones, are long and slender.
The adult specimens borrowed from the U. S. National Museum
for the purposes of illustration ranged to 8.9 mm in length, which
is larger than any of the other dipterous parasite of the sawfly,
and almost equal to the length of a large sawfly cocoon. The mouth-
hooks of the third instar are one-half again as large as any of the
other dipterous parasites. It may be that eggs deposited on
D. similis larvae are incapable of maturing on the sawfly because
it provides an insufficient amount of food. Only dead third instars
and several puparia with completely devoured hosts have been
encountered in cocoons. In any case, E. edwardsii is successful inso¬
far as it kills a small number of sawflies, but it is rare and
insignificant in its overall effects.
The size of the adults will set them apart immediately if they
should ever be reared. In addition, the presence of apical cruciate
bristles on the mesoscutellum, but absence of a pair on the disc,
is characteristic. Males (Fig. 73) lack the orbital bristles of fe¬
males, and also have pulvilli and tarsal claws twice the lengths
of those from the females.
Summary
Twenty-five species of parasites, 21 Hymenoptera and 4 Diptera,
have been reared from Diprion similis (Hartig) in Wisconsin.
Nine of these species are reported here for the first time as para¬
sites of this sawfly. Two illustrated keys have been prepared to
aid in the separation of these parasites. The first is designed to
separate the species on the basis of remains left in the host cocoon,
1971] Mertins and Coppel — Parasites of the Pine Sawfly 167
while the second will allow identification of the adults. Brief notes
on the biology of each species, and descriptions of their final instar
cephalic structures are also presented.
Acknowledgements
The authors wish to express their appreciation to the staff of
the Insect Identification and Parasite Introduction Branch of the
U. S. Department of Agriculture, especially Miss L. M. Walkley,
and Messrs. B. D. Burks and C. W. Sabrosky, and to R. E. Bugbee,
Allegheny College, Meadville, Pennsylvania, for the identification
of parasites.
Approved for publication by the Director of the Research Divi¬
sion, College of Agricultural and Life Sciences. This project was
supported in part by the Wisconsin Department of Natural Re¬
sources through the School of Natural Resources, and in part by
the University of Wisconsin Research Committee of the Graduate
School with funds supplied by the Wisconsin Alumni Research
Foundation. The authors are Research Assistant and Professor of
Entomology, respectively, University of Wisconsin.
References Cited
Balduf, W. V. 1937. Bionomic notes on the common bag worm, Thyridopteryx
ephemerae formis Haw., (Lepidoptera, Psychidae) and its insect enemies
(Hymenoptera, Lepidoptera). Proc. Entomol. Soc. Wash. 39: 169-184.
Britton, W. E. 1915. A destructive European sawfly in Connecticut. Conn.
Agr. Expt. Sta. Rept. 39: 118-125.
Burks, B. D. 1943. The North American parasitic wasps of the genus
Tetrastiches — a contribution to the biological control of insect pests. Proc.
U. S. Nat. Mus. 93: 505-506.
Clausen, C. P. 1940. Entomophagous Insects. McGraw-Hill Book Co., N. Y.
and London. 688 p.
Coppel, H. C. 1962. The introduced pine sawfly in Wisconsin. Wis. Conserv.
Dept. For. Pest Leafl. 4, 4 p.
Cushman, R. A. 1927. The parasites of the pine tip moth, Rhyacionia frustrana
(Comstock). J. Agr. Res. 34: 615-622.
Finlayson, T. 1960. Taxonomy of cocoons and puparia, and their contents, of
Canadian parasites of Neodiprion sertifer (Geoff.). Can. Entomol. 92:
20-47.
Finlayson, T. 1962. Taxonomy of cocoons and puparia, and their contents, of
Canadian parasites of Diprion similis (Htg.) . (Hymenoptera: Diprioni-
dae). Can. Entomol. 94:271-282.
Gahan, A. B. 1933. The serphoid and chalcidoid parasites of the hessian fly.
U. S. Dept. Agr. Misc. Coll. 174, 148 p.
Langston, R. L. 1957. A synopsis of hymenopterous parasites of Malacosoma
in California. Univ. Cal. Pub. Entomol. 14: 1-49.
Mertins, J. W., and H. C. Coppel. 1968. The changing role of Exenterus amic -
torius (Panzer), a parasite of Diprion similis (Hartig) in Wisconsin.
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Middleton, W. 1923. The imported pine sawfly. U.S.D.A. Bull. 1182, 22 p.
Morris, K. R. S. 1938. Eupelmella vesicularis Retz. (Chalcididae) as a predator
of another chalcid, Microplectron fwscipennis Zett. Parasitology 30 : 20-32.
Muesebeck, C. F. W., and S. M. Dohanian. 1927. A study in hyperparasitism,
with particular reference to the parasites of Apanteles melanoscelus
(Ratzeburg). U. S. Dept. Agr. Bull. 1487, 36 p.
Munro, H. A. U. 1935. The ecology of the pine sawfly, Diprion similis Htg.
(Unpub.) M. S. Thesis, McGill Univ., Montreal, 72 p.
Townes, H., and M. Townes. 1960. Ichneumon-flies of America North of Mex¬
ico. U. S. Nat. Mus. Bull. 216(2), 676 p.
Turnock, W. J., and J. C. E. Melvin. 1963. The status of Bessa harveyi
(Townsend) (DipteraiTachinidae). Can. Entomol. 95: 646-654.
WISCONSIN ACADEMY OF SCIENCES, ARTS & LETTERS
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L. G. Monthey
University of Wisconsin — Madison
Chairman — Junior Academy of Science
LoRoy Lee
W.A.S.A.L. Office, Madison
ACADEMY COUNCIL
The Academy Council includes the above named officers and officials and
the following past presidents:
Robert J. Dicke
Henry A. Meyer
Merritt Y. Hughes
Carl Welty
J. Martin Klotsche
Aaron J. Ihde
Walter E. Scott
Harry Hayden Clark
David J. Behling
John W. Thomson
Paul W. Boutwell
A. W. Schorger
Henry A. Schuette
Lowell E. Noland
Otto L. Kowalke
Katherine G. Nelson
Ralph N. Buckstaff
Joseph G. Baier
Stephen F. Darling
A. A. Suppan
W. B. Sarles
MAY 3 1971
/
Cover Design by Arthur Thrall, Lawrence University
TRANSACTIONS OF THE
WISCONSIN ACADEMY
OF SCIENCES, ARTS
AND LETTERS
LX— 1972
Editor
W ALTER F. PETERSON
TRANSACTIONS OF THE
WISCONSIN ACADEMY
Established 1870
Volume LX
THE AESTHETIC EDGE 1
Norman Olson
THE NATURAL SCIENCE OF AN AMERICAN PIONEER:
A CASE STUDY 7
Donald Zochert
THE EVOLUTION OF FACULTY GOVERNMENT OF THE
UNIVERSITY OF WISCONSIN— MILWAUKEE 17
Ted J. McLaughlin
THE EFFECT OF RESTAURANT SERVICES ON THE
SURVIVAL RATE OF TOURIST-LODGING
ESTABLISHMENTS IN WISCONSIN 33
L. G. Monthey and R. A. Ricketts
THE REMAKING OF “AMERICAN LITERATURE” 45
Donald Emerson
DISCONTINUITIES IN DEMOCRATIC SYSTEMS
AND MASS SOCIETIES 53
Charles Redenius
THE FIFTH PAN AMERICAN CONFERENCE: PROVING
GROUND FOR WARREN G. HARDING’S
LATIN AMERICAN POLICY 69
Kenneth J. Grieb
FOOD HABITS OF THE COHO SALMON,
ONCORHYNCHUS KISUTCH,
IN LAKE MICHIGAN 79
Margaret A. Harney and Carroll R. Norden
LIMNOLOGY OF SOME MADISON LAKES:
ANNUAL CYCLES 87
Kenton M. Stewart and Arthur D. Hasler
THE ALGAE OF THE WINNEBAGO POOL
AND SOME TRIBUTARY WATERS 125
William E. Sloey and John L. Blum
KINETICS OF ORTHOPHOSPHATE UPTAKE
BY PHYTOPLANKTON POPULATIONS
IN LAKE WINNEBAGO 147
Steven Bartell and Sumner Richman
A RECORD OF THE FRESHWATER NEMERTEAN,
PROSTOMA RUBRUM , IN WISCONSIN 179
Robert F. Browning
A RECORD OF CRASPEDACUSTA SOWERBYl
IN WISCONSIN 181
Richard P. Howmiller and G. M. Ludwig
PEDOLOGY OF THE TWO CREEKS SECTION,
MANITOWOC COUNTY, WISCONSIN 183
Gerhard B. Lee and M. E. Horn
LOWER WISCONSIN RIVER VALLEY SOIL
RESOURCES AND USE POTENTIALS 201
G. E. Musolf and F. D. Hole
THE PITUITARY GLAND OF THE ALE WIFE IN LAKE
MICHIGAN: CYCLICAL CHANGES IN THREE
ADENOHYPOPHYSEAL CELL TYPES 211
Alexander H. H. Li and Eldon D. Warnei*
OVIPOSITIONAL SITE PREFERENCES OF THE OAK
DEFOLIATING GRASSHOPPER, DENDROTETT1X
QUERCUS , IN WISCONSIN 225
Douglas A. Valek and Harry C. Coppel
WILD RIVERS OF NORTHEASTERN WISCONSIN
(WILD RIVERS COOPERATIVE
RESEARCH PROJECT) 233
George Becker
CANOEING THE WILD RIVERS PINE AND POPPLE 239
Joe Mills
AN ARCHAEOLOGICAL SURVEY OF THE PINE,
PIKE AND POPPLE RIVERS 265
Robert J. Salzer
THE MAMMALS OF THE PINE AND POPPLE RIVER AREA 275
Robert A. McCabe
THE AVIFAUNA OF THE PINE-POPPLE WATERSHED 291
Howard Young
THE AMPHIBIANS AND REPTILES OF FOREST, FLORENCE
AND MARINETTE COUNTIES WITH SPECIAL REFERENCE
TO THE PINE, POPPLE AND PIKE WATERSHEDS 303
William E. Dickinson
ANNOTATED LIST OF THE FISHES
OF THE PINE-POPPLE BASIN 309
George C. Becker
BIOGRAPHIES
331
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NORMAN OLSON
50 th President of the
WISCONSIN ACADEMY OF SCIENCE, ARTS AND LETTERS
THE AESTHETIC EDGE
Norman Olson, President 1970-71
May, 1971
Over the past several years I have developed a concern that the
critical faculties of a great number of people have become im¬
paired in the ability to understand and evaluate The Arts as they
exist in our present culture. It is my sometimes worry that the
Post-World War II period has seen the destruction in many, both
artist and layman, of what may be termed the aesthetic edge — the
artistic sensitivity that exists within a person. For, just as this
aesthetic edge can be sharpened and fine-honed with proper study,
self-discipline and training, it can be dulled and destroyed by con¬
stant exposure to the corroding effects of the vulgar, the false,
and the inept in The Arts.
A problem for every man lies in recognizing those qualities of
a piece of music, or a painting, or a play, or a poem, that make it
a work of art or make it a fraud. For now, I will be content to
define the problem and break it down into what to me are three
of its major aspects. It will be your task to weigh my arguments
with the indulgence that you would wish for if you were in my
shoes. I will, for simplicity’s sake, concentrate mainly on the art
of painting.
It is a frustrating experience for many of us these days to
visit an art gallery and discover that it is filled from ceiling to
floor — and probably from wall to wall — with what is collectively
called modern art. It is not unusual to travel the length of the
gallery and find nothing whatever in it that produces a pleasur¬
able response in us. We may sometimes recognize the object that
the artist had in mind to create in paint or sculpture, but why he
bothered to make it, or dared to show it, remains a mystery.
But do we dare reveal our inability to appreciate and respond
to many creations of contemporary artists? Those who are skilled
and schooled in such matters, or pretend to be, may scoff at us.
They are proficient — we are deficient. They feel clever — we feel
stupid. But why are we unable to appreciate so much of today’s
artistic production? I must warn you that I intend to explore the
matter with the obvious bias of one who has lived in reverent
appreciation of the rational in all of the arts. By “rational” I
really mean the incorporation of the quality into the work of art
that early Renaissance humanists called “right reason.”
1
2 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
There is a vast field of scientific study aimed at analyzing human
response in terms of psychological cause and effect. I do not reject
the data pouring forth from this source, but merely acknowledge
it in moving on to a more primitive body of data ; i.e,, the aesthetic
theories, and critical observations, of some of the philosophers
and artists whose reputations and writings have survived the test
of time. For as Andre Malraux has pointed out in his brilliant
book, The Metamorphosis of the Gods, just as the artist tries to
immunize his creative work against time, “A work . . . becomes
a work of art in virtue of being outside time.”1 And of course if
this test of time applies to painting, it also pertains to endeavors
of art in literature and other fields as well.
Plato in his Timaeus dialogue explains that, “God devised the
gift of sight for us so that we might observe the movements which
have been described by reason in the heavens and apply them to
the motions of our own mind which are akin to them, so far as
what is troubled can claim kinship with what is serene.”2 A bit
farther along he observes that rhythm was given to us, “to help
us in dealing with what is unmeasured and chaotic in the minds
of most of us.”3
Writer upon writer points out the common chords of rhythm,
order and balance in all branches of the arts. Poetry, music, and
the dance clearly depend upon these. So do sculpture and painting
and when they are missing from a work by choice, chance, or the
ineptness of the artist, the lack is conveyed inside us by our own
powers of perception. Chaos in anything is disturbing, even fright¬
ening, and we tend to reject it when we see it on the canvas or
in the plastic arts merely because it is contrary to our natural
instincts.
How do we know that so much of modern art is without rhythm,
order and balance? Is it there but we cannot comprehend through
ignorance? Does the fault, dear Brutus, lie within ourselves? And
one last question: what is abstract expressionism? The last ques¬
tion is necessary because that is the school or style arising from
what Katherine Kuh describes as the break-up of traditional art
forms.
Miss Kuh, in commenting upon the painting of Jackson Pollock
says, “Very different in motivation from surrealism, abstract ex¬
pressionism was not concerned with symbols of the unconscious,
but with the artist’s spontaneous feelings at the moment of paint¬
ing. Abstract expressionism and especially Jackson Pollock were
the final denial of all that Renaissance and Classical Art stood for.”4
1 Andre Malraux, The Metamorphosis of the Gods, p. 32.
2 Plato, Timaeus, in E. F. Carritt, Philosophies of Beauty, p. 28.
sIhid, p. 28.
4 Katherine Kuh, Break-Up : The core of m.odern art, p. 105.
1972]
Olson — The Aesthetic Edge
3
She added that Pollock “. . . created a new kind of fury to echo
the fury within himself.”5 One can’t help but wonder at the dura¬
bility of such a great passion, and ponder whether the evidence
of it via paintbrush on canvas is of interest as art or merely as
a psychic efflux. In her comment on a painting by William DeKoon-
ing entitled “Excavation” Miss Kuh observes that the picture . .
with its all-over pattern and warm loam-like color, has to do with
the sensations of digging into the earth, and possibly into one¬
self.”6 After reading that, I had the impression that the author
had reached very far into the realm of speculation.
The artist Frank Kline confesses, “I don’t paint a given object—
a figure or a table; I paint an organization that becomes a paint¬
ing.”7 Miss Kuh in talking about a painting of Pierre Soulages
that has no name, but only a date, explains, “For both artist and
viewer, meaning relies on the gratification of rich pigment deftly
manipulated. In other words, the paint itself becomes at once the
raison d’etre and the image.”8 We have just been informed that
for this painting at least, the medium is the message!
So, now we have one reason for our dilemma as viewers in not
understanding a great many of the paintings that are included in
exhibits today. The problem is that in abstract expressionism the
artist often has nothing to convey to us. If we do think we have
found a meaning, quite probably it is one evoked entirely within
ourselves. Perhaps the artists, relying entirely upon the subjec¬
tive creativity of the viewers for effect, are taking Oscar Wilde
seriously in his statement that, “It is the spectator, and not life,
that art really mirrors.”9
Another kind of painting that confronts us these days is that
which admittedly does contain the representation of a recognizable
object. It may be the enlarged picture of a tin can, complete with
label, or a character from a comic strip. The artist has used metic¬
ulous care— the rendition is perfect, and the reaction is that here
is the work of someone who should get “A” in mechanical draw¬
ing, but who is woefully lacking in imagination.
The subject is prosaic; the message is that printed on the can’s
label or found innocently scrawled within the picture’s confines.
As Aristotle points out in the poetics, “The ludicrous is only one
species of the ugly.”10
Another painting hanging close by may be concerned with human
anatomy. The chances are excellent that an entire figure will not
6 Ibid .
6 Ibid. p. 103.
7 Ibid. p. 101.
8 Ibid. p. 102.
9Hesketh Pearson, Oscar Wilde , His Life and Wit, p. 132.
10 Aristotle, <c Poetics” in Carritt, p. 32.
4 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
be there, however, but just a selected portion rendered with pho¬
tographic precision. As counterpoint to pictures of this kind we
find enormous plastic sculptured pieces placed strategically about
the gallery. These are, seemingly, reconstructions of the entrails
of fabulous monsters, or giant earthworms, or both. Yes, the or¬
ganic element is definitely included in contemporary art.
Still another type of painting is that which depicts a subject
that is sordid, mean or ugly. Satire with its sharp teeth has its
place in art as witness the drawings of Hogarth, Grosz or Daumier.
But when there is no purpose revealed in the picture except to
move the viewer to feel disgust and perhaps admit that ugliness
exists, what credit can we give to the piece as art?
The second aspect of the problem then, appears also to focus
upon the artist. It is that the artist must utilize his genius within
the framework of what is aesthetically acceptable and understand¬
able to the sensitive viewer, and it is not a sufficient excuse on
the part of artists or gallery directors to say that it is the view¬
ers’ limitations that make a painting misunderstood — or not under¬
stood at all. Tolstoy has commented upon this by observing, “To
say that a work of art is good, but incomprehensible to the major¬
ity of men, is the same as saying of some kind of food that it is
very good but that most people can’t eat it.”11
The third and last aspect of the problem, I feel, is that which
concerns the taste — or aesthetic edge — of the gallery visitor. To
consider this we must swing the mirror around until we see our¬
selves squarely in it. We have contended that there is good art and
bad art; and that there are paintings and sculpture with a great
message and some with a poor message; and very many with no
message at all. But when we reach a conclusion as to the merits
of a particular painting, that conclusion is the end result of our
entire lifetime of conditioning. Thomas Hobbes who is emphatic
in his philosophical observations and conclusions says in Leviathan,
“Whatsoever is the object of any man’s appetite or desire; that is
it, which he for his part calleth good: and the object of his hate,
and aversion evill, and of his contempt, vile and inconsiderable .”12
If Hobbes is right, and we see the evidence of this particular
truth all around us, then we must realize that we can destroy our
natural abilities to appreciate what is good and reject what is bad.
The woodland Indian of Wisconsin Territory days, who could track
game with uncanny skill because of his highly developed and un¬
shattered sensitivity toward what he saw, heard, smelled, touched
or tasted, is in great contrast to the Modern American who over¬
eats three times a day and falls asleep at the symphony. We can-
11 Lyov Nikolayevitch Tolstoy, “What is Art?” in Carritt, p. 192.
12 Thomas Hobbes, Leviathan, quoted in E. F. Carritt, Philosophies of Beauty, p. 56.
1972]
Olson — The Aesthetic Edge
5
not as a steady diet read books in the Last Exit to Brooklyn genre
and then appreciate the delicate characterization in James’ The
Wings of the Dove . We cannot drown ourselves daily in rock music,
and hope to experience the thrill that lurks for the responsive
listener in a Brahms concerto. We cannot live and work surrounded
by psychedelic posters without becoming color-numb.
Milton believed that for a man to become a great poet, he must
rigidly follow rules of self-discipline and education. He must be¬
come in effect a priest in the purity of the conditioning he applies
to his natural talents. That kind of life is, of course, beyond the
reach or resolve of most of us. But we can resolve to avoid con¬
stant exposure to those things that would certainly dull our aes¬
thetic edge.
What we do as individuals will not change the world of art.
Painters and sculptors will continue to pour forth a never-ending
line of questionable artifacts. And where these probable frauds
would normally perish under the dead weight of their inherent
lack of artistic merit, the directors of many art centers and some
affluent collectors march forth in the role of dens ex machina, pur¬
chase the doomed pieces, and preserve them forever in our muse¬
ums and galleries.
We must fight the good fight to defend and support the artists
who produce honest art. To do this we have an obligation to know
what is good in art and have faith in our knowledge. We may
lose the battle to the frauds and their sponsors, but as Ajax said,
“Oh Zeus ... if so be that we must die, let us die in the light!”13
Lists of Works Consulted
Carritt, E. F., ed. Philosophies of Beauty. Oxford, The Clarendon Press, 1962.
Kuh, Katherine. Break-TJp: The core of modern art. London, Cory, Adams &
MacKay, Ltd, 1965.
Malraux, Andre. The Metamorphosis of the Gods. Garden City, Doubleday &
Company, Inc., 1960.
Pearson, Hesketh. Osccur Wilde, His Life and Wit. New York, Harper &
Brothers Publishers, 1946.
13 Longinus, “On the Sublime,” Carritt, p. 38.
i
I
I
THE NATURAL SCIENCE OF AN AMERICAN PIONEER:
A CASE STUDY
Donald Zo chert
How much did the common man of the frontier know about
natural science? What grasp did he have of theoretical concepts?
What evidence did he leave of scientific or philosophic specula¬
tion? The answers to these questions appear locked in an inarticu¬
late past. Indeed, when students of science have turned their
attention to the frontier it has generally been to trace the passage
of great naturalists, or to demonstrate the growth of scientific
interests in isolated academic contexts, or to detail the accom¬
plishments of government-sponsored surveys. The nature and ex¬
tent of popular scientific knowledge on the frontier have been all
but ignored.
It may be argued that the common man rarely left a record
of his mind at work, and that his intellectual stores must there¬
fore remain inaccessible and uncounted. This, however, is to over¬
look the production on the frontier of a substantial literature—
journals, letters, diaries— which has gone largely unexamined
from the perspective of science. It is the purpose of this paper to
examine from that perspective the journals of James Clyman
(1792-1881), an early pioneer of Milwaukee and a man who spent
his life on the successive edge of settlement.1
James Clyman is representative of the common man on the
frontier in several respects. He had little or no formal education.
He followed no specific trade, being by turn a farmer, a sur¬
veyor's assistant, a trapper, a storekeeper, a soldier, a miller, and
ultimately the owner of a modest California ranch. His life, in
common with others on the frontier, was restless and westering.2
If he be considered atypical for having produced a journal, the
thoughts and information he recorded give no evidence of having
1 Clyman’s journals have been published in Charles L. Camp, ed., James Clyman,
Frontiersman : The Adventures of a Trapper and Covered-Wagon Emigrant as Told
in His Own Reminiscences and Diaries , “definitive” edition (Portland, Ore., 1960).
The present writer is preparing a biography of Clyman.
2 Born in Virginia, east of the Blue Ridge, Clyman moved to Ohio at the age of
sixteen. He left home for southern Indiana in 1818, spent the winter, and moved on
to Illinois. Prom 1823 to 1827 he was engaged in the Rocky Mountain fur trade, rang¬
ing the Plains and Central Rockies north of Colorado. He settled at Danville, Ill.,
upon his return from the mountains, but in 1832, under the impetus of the Black Hawk
War, resumed a wandering — and this time military — life, serving in the Illinois Vol¬
unteers, U.S. Mounted Rangers, and First U.S. Dragoons. Three years later he settled
at Milwaukee.
7
8 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
been inspired by anything but the common experience of the
frontier.
Clyman emigrated to Milwaukee in 1835 and made that frontier
village his home until 1844. In the latter year, for what he said
were reasons of health, he joined an emigrant train to Oregon.
From Oregon, Clyman went to California in 1845 and the following
year returned to Milwaukee; in 1848 he left Wisconsin to make
his home in California.
During the early months of 1840 in Milwaukee, and through his
entire western tour from 1844 to 1846, Clyman maintained a
series of journals which indicate again that the conditions of the
frontier, and more specifically of agrarian or even village life,
imposed an intimacy with the natural world that is almost wholly
dissipated today. Here, in the wildlife, the weather and the topogra¬
phy of the frontier must be the starting point for any survey of
popular scientific knowledge. Clyman’s journals contain a great
number of elementary descriptions which derive from the direct¬
ness of this experience.
His identification of trees and birds, for instance, suggests the
energy with which the frontier environment acted upon the mind
of the pioneer. Among trees he refers to the alder, aspen, birch,
boxwood, cedar, cherry, cottonwood, dogwood, elder, elm, red fir,
white fir, white balsam fir, hackberry, hazel, hemlock, hickory, man-
zanita, maple, sugar maple, black oak. white oak, red pine, spruce
pine, white pine, yellow pine, redwood, sycamore, spruce, walnut,
willow and yew. He identifies the following birds : the raven, moun¬
tain grouse, duck, pheasant, swan, teal, plover, heron, brant, shagg,
mocking bird, goose, crane, blackbird, woodcock, fir grouse, quail,
meadowlark, redbreasted woodpecker, sparrow, condor, buzzard,
crow, hawk, pigeon, vulture, royal vulture, bald eagle, turkey buz¬
zard and snipe. In addition, his journals contain references to
fifty-one different kinds of plants and flowers, thirty-two species
of mammals, twenty-one varieties of fruit, seven reptiles and in¬
sects, and four species of fish.3
None of these botanical or zoological references demonstrates a
technical, systematic or theoretical understanding of the objects
Clyman described; he does not suggest relationships between dif¬
ferent plants or animals — other than to indicate the similarity
between eastern and western species — nor does he draw analogies
between plants and animals, as was common among popularizers
of science. Despite the great naming and counting compulsion of
the times, and perhaps because of the confusion generated by
contending taxonomic systems, Clyman, like other laymen on the
frontier practiced only the most rudimentary sort of classification,
Camp, passim. Specific references are available from the present writer.
1972] Zochert — Natural Science of American Pioneer 9
categorizing in such indistinct terms as “animal kingdom” or
“plant kingdom” or “feathery tribe.” This naming of the natural
world forms the most elementary level of Clyman’s knowledge of
science.
Clyman’s description of topographical features and meteorologi¬
cal phenomena, with one exception, likewise carries no indication
of technical or theoretical competency. The exception is an observa¬
tion made in September 1844, when Clyman, on his way to Ore¬
gon, noted that the emigrant trail had for several days been
obscured by smoke from Indian fires. “The day verry Smoky,”
he wrote on September 26, “& I Begin to daubt Mr. Espys theory
of produceeing rain by any phisical means as the whole country
has been on fire for a month past & no rain yet.”4 The allusion
is to a theory of James Pollard Espy, proposed in 1838 and given
wide circulation in popular and scientific periodicals through 1843,
that a rainfall of great duration would occur “if masses of timber
. . . should be prepared and fired simultaneously every seven days
in the summer, on the west of the United States, in a line of six
or seven hundred miles long . . .”5
With his geological references, however, Clyman indicates a
deeper and more sustained interest in the theoretical aspects of
science. Many of his references, to be sure, remain simply descrip¬
tive. He notes the presence, for instance, of slate, sandstone, lime¬
stone, shell rock, granite, flint, pebble rock, basalt, chalk, obsidian,
marble, slag and scoria, and attributes to them such qualities as
“saturated,” “vitreous,” “decomposed,” “indurated” and “porous.”
He identifies deposits of salt, lime, saleratus, coal, soda, potash,
sulphur, lead, lye, mercury, gold, silver, iron and quartz.6 But in
several other observations he demonstrates a grasp of geological
concepts and a recognition of classification.
In his 1840 diary at Milwaukee, Clyman summarizes the rock
formations he had encountered in northeastern Wyoming seven¬
teen years earlier: “I do not recolect that I saw any primitive
rock in this place except some granite Boulders all the rock that
I saw being secondary Lime rock although all the petrifactions
and even pebble stone are verry hard and flinty and in fact all
the rock formation in this region is Trasition [ transition ] and
secondary . . .”7 Clyman here followed an anglicized version of
the system of classifying rock formations developed by the great
Austrian mineralogist Abraham Gottlob Werner. Werner placed
4 Camp, 107.
6 See James Pollard Espy, The Philosophy of Storms (Boston, 1841), 492-500. The
passage quoted, however, is from W. E. Knowles Middleton, A History of the Theories
of Rain and Other Forms of Precipitation (New York, 1966), 160.
6 Camp, passim. See Note 3.
7 Ibid., 51-52. My italics.
10 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
rock formations in three classes — primitive, transition and floetz;
his American followers substituted the more easily understood
term “secondary” for the germanic “floetz.”8
Clyman’s periodic reference to the stratification of rocks also
suggests a recognition of rock classification. The inclination of
rock strata is a characteristic once thought to distinguish primi¬
tive from secondary rock formations.9 Clyman’s references in¬
clude the terms strata, substrata, “regular stratification,” “rocks
without strata,” as well as observations of the inclination of strata,
e.g. “perpendicular strata.”
Additional evidence of geological conceptualization can be found
in Clyman’s frequent allusions to the volcanic process; these in¬
clude the terms volcanic eruption, volcanic mud, vitrification, con¬
vulsion, crater, globule and fusion. His description of a western
valley as being bounded “by a range of Bald mountains shewing
in a peculiar manner their volcanic origin by their standing in
the form of wavse of the ocean” suggests a recognition of the
molten nature of lava. On the same subject, Clyman attributes
the turbidity of the Missouri River to airborne volcanic ash.10
The origin of erratic boulders or “lost rocks” posed a different
sort of problem for scientist and layman alike. Benjamin Silliman,
who had established a wide influence as professor at Yale College
and editor of the American Journal of Science, pronounced it in
1821 “among the most interesting of geological occurances,” and
it continued to exercise a fascination for the next fifty years.11
Considering the extent of his travels in the old Northwest Terri¬
tory, Clyman could hardly avoid comment on the glacial drift.
Every person that has ever passed through the western country [he wrote
in his 1840 diary at Milwaukee] must have observed the Quanty of
granite Boulders that lay scattered all over the vast extent of country . . .
and which seem to grow larger and more plenty in allmost Regular
progression as you traverse the Region northward ... as none of this
rock is found in regular strata it has been a matter of much speculation
to know how they came situate whare they are as likewise whare they
8 Werner’s “primitive” rock formations were of chemical origin, “transition” were
partly mechanical and partly chemical in origin, and “floetz” were formed chiefly
by mechanical deposition. For the influence of Werner upon such American scientists
as Benjamin Silliman, Amos Eaton and Thomas Cooper, see William Martin Small¬
wood and Mabel Sarah Coon Smallwood, Natural History and the American Mind
(New York, 1941), 243-244, 267-268, 296.
9 John Playfair, Illustrations of the Huttonian Theory of the Earth (1802; facsimile
reprint, Urbana, Ill., 1956), 12. This view was being challenged in the late 1820s,
however ; see Charles Schuchert, “The Progress of Historical Geology in North Amer¬
ica,” in Edward Salisbury Dana, et al., A Century of Science in America with Special
Reference to the American Journal of Science, 1S18-191S (New Haven, 1918), 78.
10 Camp, 221, 49.
11 See Herbert E. Gregory, “Steps of Progress in the Interpretation of Land Forms,”
in Dana, A Century of Science, 131-139. Silliman’s observation is quoted on p. 132.
J. D. Dana, writing in 1871, concluded (p. 138) it “is still a mooted question in
American geology whether the events of the Glacial era were due to glaciers or ice¬
bergs. ”
1972] Zochert — Natural Science of American Pioneer
11
came from and as all Speculation on this head must be mere conjecture
my oppinion is that [in a] remote period the whole of the Missisippi valy
was covered with water at which time those rocks were brought from
the base of the Rocky mountains in the ice and carried southward wore
let loos as they progressed by the thaws and sunk whare they are now
found12
Clyman’s explanation of erratic boulders clearly involved both
ice and water as agents of transportation, a popular and persistent
stand which comfortably reconciled the Mosaic account of the
Deluge with the new science of glaciology. More importantly, how¬
ever, Clyman’s discussion demonstrates a speculative, as well as
observational, approach to science.
Popular science obviously could not be restricted to quanti¬
tatively describing the natural environment. The same impulse
which led to the description of natural objects led as well to
speculation concerning their origin and meaning. While scholastics
calculated such imponderables as the depth of Hell,13 more modest
men were brought face to face with concepts and abstractions
almost too great to bear. James Clyman, in a reflective mood at
Milwaukee, grappled with no less an abstraction than the nature
of infinity and the dimensions of the universe.
He begins by asserting that time and space are infinite, but this
leads him rather quickly to a quandary:
Two things Infinite Time and space Two things more appear to be
attached to the above infinity (wiz) Matter and number matter appears
to prevade the infinity of space and number attempts to define the
Quantity of matter as well as to give bounds to Space — which continually
Expands before matter and number — and all human speculation is here
bounded in matter and number leaveing space at least allmost completely
untouched. . .14
Infinity, Clyman is saying, defies measurement; therefore how
can we define it? We are restrained from a definition of infinity
by our dependence upon number. There is, however, another way
to approach the problem. If one could find an instantaneous
phenomenon— -that is, one beyond measurement— then the universe
might be inferred to be infinite. But if there are no phenomena
except which can be measured, then the universe is ultimately
finite— even if beyond our capacity to measure it.
The velocity of light seems to be the greatest of all Known principals
unless Electricity should be greater Some have thought that Electricity
12 Camp, 49-50.
13 1,832,308,363 miles, as computed by Josiah Meigs. Dirk J. Struik, Yankee Science
in the Making, new revised edition (New York, 1962), 463.
14 James Clyman, MS. memorandum and diary, Everett D. Graff Collection, New¬
berry Library, Chicago. My transcription differs slightly from Camp’s in spelling and
punctuation, although notes will be to Camp. Camp, 49.
12 Wisconsin Academy of Sciences, Arts and Letters [VoL 60
is insantaneous throughout all universal space I can hardly think this to
be the fact but if it should it puts to rest the difficult Question of infinity
of space For if any Known and palpable principle is instantaneous
through eternal space then Eternal space may be infinite as to bounds
and duration and mater may likewise be infinite as to Quanty and
duration But on the contrary if no Known palpable or impalpable princi¬
ple or matter can be found but what is limited by size or time then
infinity means nothing more than such an imense mass of space matter
or time as becomes immeasureable and incomprehensable to all means
of comparison for instance
We may comprehend the globe we inhabit pretty fully and even the
sollar System but a million of such systems becomes incomprehensible
allthough even a million such Systems may fall verry short of the
Quantity of matter in existence throughout the universal Kingdom
But notwithstanding the immense Quantity still Finity becomes a part
of infinity and the globee being Finnite or mesureable so by comparison
of one mesurable part or particle of infinite matter occupying a speck of
space may we geathur some crude Idea of infinity itself allthough this
Idea may ammount to nothing more than to say all things have thier
Bounds and limits space has its bounds and time has its limmits mater
occupies all space and time wears out all things15
What has Clyman accomplished thus far? He has begun by
asserting time and space to be infinite, but has quickly found it
difficult to define infinity. Then he has subtly changed the grounds
of the problem, from a consideration of the nature of infinity to a
consideration of whether the universe — space — is indeed infinite.
To this question he provides two possible answers. One, the uni¬
verse may be considered infinite if there exists an instantaneous
phenomenon — the speed of light, for instance, or of electricity.
Clearly he does not consider either of these to be instantaneous or
beyond measurement. This leads him to the second possibility:
that the universe can be finite even though “incomprehensible.”
Our perceptions, crude and limited, cannot finally settle the ques¬
tion, although by comprehending a part of the universe we might
get a “crude Idea” of the whole. But to suggest, as Clyman does,
that this crude idea would be “of infinity itself,” and might mean
no more than that all things have limits, appears to collapse the
argument in a contradiction of terms. He must go back and once
again try to find a definition of infinity:
Some seem to think that infinity means something than can never Even
have a beginning nor an end and that if it were posible to move with
the velocity of Light for millions and millions of years and even time
without limit that you then have not more than set out But admit all this
and say that after you have flown with the velocity of light for as many
yares as there is particle of sand included in the whole Solar System
even at that immense time and immense swiftness if you have advanced
a Quarter of an inch comparatively you at once give imaginary Bounds
15 Ibid., 53-54.
1972] Zochert — Natural Science of American Pioneer 13
to space although it may not be posible to measure or comprehend but
a verry small Quantity of Space or matter10
The definition of infinity as something without beginning or
end carries no endorsement from Clyman; he uses it, in fact, as
a device with which to reconsider the dimensions of the universe.
I would suggest that Clyman was content with the position that
infinity appears to be beyond definition, as he indicated at the
beginning of the passage. As for the extent of the universe, he
appears to rest on the assumption that the definition of a part in
effect sets limits on the whole, even if the limits of the whole are
beyond perception.
I have quoted these reflections on infinity and the universe at
such length chiefly to illustrate the strength and quality of Cly-
man’s thought. Crude, untrained, repetitious, without discipline—
it nevertheless testifies to a forcing of the intellect, and to thoughts
being carried forward and at least momentarily sustained, not on
the basis of formal knowledge but purely on the speculative
impulse.
James Clyman’s interest in natural science found expression
at three different levels— one of simple observation, another of
classification, and a third of speculation or theory. In the fields
of botany, zoology, geography, and to some extent meteorology
and geology, he clearly operated at the most elementary, observa¬
tional level, without attempt at classification or explanation. His
concern with geological phenomena, however, frequently did ex¬
tend to the level of classification ; his reference to the classification
of rock formations, and the terminology of stratification (imply¬
ing classification) provide evidence of this. At the level of specu¬
lation and theory may be placed his refutation of Espy’s theory
of the artificial production of rainfall; his explanation of the
turbidity of the Missouri River; the conceptualization which lay
behind the terminology of vulcanism; and his reflections on the
nature of infinity and the dimensions of the universe.
If there is a common element to these levels of scientific interest
and capability, it is a willingness to confront and define the natural
world. Even though the physical world may ultimately surpass
man’s ability to perceive or comprehend it, Clyman was intent
upon preserving the possibility of limits — quite clearly a rational
sentiment, and the expression of a desire to perceive and com¬
prehend.
16 Ibid. From here, Clyman goes on to a brief exegesis of Genesis 1:1 in an effort
to determine what limits are placed upon time and matter by the creation account.
This passage is noteworthy in two respects : it represents Clyman’s only appeal to
authority — in this case biblical — and by referring to “the first revolution of the first
globe of matter,” he indicates an exposure to the nebular theory of the origin of the
solar system.
14 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
The extent to which other men on the frontier shared this desire
is beyond the province of a case study. It may be useful, however,
to suggest some coeval expressions of scientific interest as clues
to the commonality of Clyman’s interest and competency.
Our knowledge of how frequently, and with what success, the
common man engaged in metaphysical speculation is sharply lim¬
ited by the infrequency with which such thought was transcribed ;
it is much easier, for instance, to make a journal of botanical
and zoological notations than it is to wrestle with the devil of
infinity— with pen in hand. We are fortunate, then, to have the
reflections of Osborne Russell, a fur trapper who worked the
Rocky Mountains a decade after James Clyman. Russell writes of
being “almost lost in contemplation’’ while standing on a moun¬
tain peak: “In viewing scenes like this the imagination of one
unskilled in Science wanders to the days of the Patriarchs and
after numerous conjecturings returns without any final decision
wonder is put to the test but having no proof for its argument a
doubt still remains but supposition steps forward and taking the
place of Knowledge in a few words solves the mysteries of ages
Centuries and Eras . . .”17 Here is the same pell-mell rush of
thought, the same undisciplined cession to speculation that one
finds in Clyman.
Clyman’s description of topographical features was matched in
similar language by many other frontier journalists, as was his
interest in geological phenomena.18
What truly determined an inclination toward popular science
seems to be related less to education and station in life than to a
specific frontier experience. In Clyman’s case, service in the state
land surveys probably sharpened his awareness of many scientific
objects,19 and the presence in Milwaukee of Increase A. Lapham
and other men of serious scientific interest must have provided
17 Osborne Russell, Journal of a Trapper, Aubrey L. Haines, ed. (Lincoln, Neb.,
1967), 63.
18 Compare Clyman’s topographical terms with those employed in the journals of
Nicholas Carriger and James Mathers, in Dale Morgan, ed., Overland in 18^6: Diaries
and Letters of the California-Oregon Trail (Georgetown, Calif., 1963), 1:150-158,
225-236, and in the journal of the Mormon immigrant Appleton Milo Harmon, in
Maybelle Harmon Anderson, ed., Appleton Milo Harmon Goes West (Berkeley, Calif.,
1946). Harmon, who had no more education than Clyman, is much weaker in geology.
However, Thomas J. Farnham, whose training was in law, reported during his western
journey on alluvium, basalt, scoria, stratification and “regular” stratification, and
“primary formations” ; see his Travels in the Great Western Prairies, the Anahuac
and Rocky Mountains (1843) reprinted in Reuben Gold Thwaites, ed., Early Western
Travels 17^8-18^6 (Cleveland, 1906), vols. 28-29.
10 The formal instructions given to surveyors at this time required them to take
notice of “all rivers, creeks, springs . . . the kinds of timber and undergrowth . . .
all swamps, ponds, stone quarries, coal beds, peat or turf grounds, uncommon natural
or artificial productions, such as mounds, precipices, caves ... all rapids, cascades
or falls of water . . . minerals, ores, fossils . . . the quality of the soil . . . the true
situation of all mines, salt springs and mill seats . . .” Lowell O. Stewart, Public
Land Surveys: History, Instructions, Methods (Ames, Iowa, 1935), 146.
1972] Zochert — Natural Science of American Pioneer 15
some stimulus toward scientific thinking on the part of many lay¬
men.20 But as was suggested earlier, the natural science of James
Clyman gives no evidence of having been generated by anything
but the common experience and inspiration of the frontier.
This is not to say that events beyond the frontier did not on
occasion stimulate thoughts about science. Cly man’s comment on
Espy, and those portions of his science in which he demonstrates
an exposure to concepts of classification suggest the importance,
for instance, of popular scientific literature.
The interest in popular science was fed more strongly by the
very closeness of the natural world, and the familiarity it bred.
As the “aesthetic contemplation of a perfected universe,” it gained
impetus from the desire to escape from provinciality, or from the
crudeness of frontier life. Natural science itself — especially on a
popular level — was still an accessible enterprise, as open to the
dilettante as to the devotee.
As James Clyman demonstrated, one did not need formal train¬
ing to pursue the pleasures of science. All that was needed — and
they were there in abundance — was a rich and vital environment,
and an aggressive, Whiggish spirit that sought to improve not only
property but the mind.
Note
The author wishes to acknowledge the kindness of George H.
Daniels, Department of History, Northwestern University, in
criticizing an earlier draft of this article.
20 Lapham witnesses Clyman’s signature to a letter of attorney dated July 20, 1836.
Camp, 301. There is no additional evidence of a relationship between them however.
THE EVOLUTION OF FACULTY GOVERNMENT OF
THE UNIVERSITY OF WISCONSIN— MILWAUKEE*
Ted J. McLaughlin
Outsiders frequently are surprised to discover that life within
an academic community is no more tranquil or stable than it is
among citizens who live and work off campus. In a time of social
unrest and revolt, internal campus confusion leads to public con¬
cern over the operation of state supported universities. No more
pressing priority faces the state university than does the achieve¬
ment of understanding of its structure and conduct. This paper is
a response to the critical need for clarification of the historical de¬
velopment, contemporary status, and probable future of faculty
government of The University of Wisconsin — Milwaukee (UWM).
Scope and Rationale of the Study
The Concept of University Academic Organization. Like all
human enterprises, an institution of higher learning must operate
under some rational system characterized by identifiable roles,
predictable continuity, and group goals. Unlike a military unit
with the “command'' implications of concentrated authority or a
business enterprise with the ‘‘management team” functions of
authority, responsibility, and accountability, a university is or¬
ganized as a polity. By traditional practice and by legal sanction,
a state university operates under a faculty government, with
varying degrees of democracy.
The jurisdiction and powers of a university faculty are seldom
definitive, attempts at legal specificities notwithstanding. Does
the faculty “control” or merely “participate in” decisions con¬
cerning the academic program? Such extreme disclaimers and
claims are useless rhetorical exercises. Ultimately, “. . . an effec¬
tive system of campus governance should be built on the concept
of ‘shared authority' between the faculty and the administration.”1
A democratic government depends on mutual checks and balances —
especially a balance of authority (“effective influence”)2 among
its major branches. At least in the ideal university academic or-
* This paper was presented in condensed form at the Centennial Meeting of the Wis¬
consin Academy, October 3, 1970.
1 American Association for Higher Education, Faculty Participation in Academic
Governance (Washington, D. C., 1967), p. 1.
2 Faculty Participation , p. 14.
17
18 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
ganization, the faculty legislates academic policy proposals, while
the administration exercises the executive functions of review,
approval, and implementation. Curiously, the judicial function
of democratic governance appears to operate as a de facto rather
than a de jure phenomenon in contemporary university practice.
Method, Data, and Limitations of the Study. This study reports
the results of a descriptive evaluation of UWM faculty government
as an effective instrument in the operation of the institution. The
period covered begins with the initial academic year of 1956-57
and extends to the conclusion of the 1969-70 academic year. The
writer has compared the theoretical assumptions underlying fac¬
ulty government with the actual decision-making processes of the
faculty. In arriving at the conclusions and predictions stated at
the end of this report, the author has used two major categories
of information. Archival records of faculty meeting agendas,
minutes, documents, and reports provide the primary source basis
for developmental assessment, in addition to statutory laws and
other codified regulations. Critical observations of faculty meet¬
ings and practices constitute the other body of source material.
Except as faculty-administrative relationships impinge directly
on the subject of faculty governance, no attempt is made to study
administration, per se. And although discrete faculties of the
several colleges, schools, and departments are responsible for aca¬
demic policy within their restricted jurisdictions, the operation
of these subsidiary UWM units is outside the limits of this gen¬
eral study.
Historical and Legal Basis of UWM Faculty Government
In a landmark action, the 1955 special session of the Wisconsin
Legislature created a Coordinating Committee (later Council)
for Higher Education, directed the CCHE to merge the competing
state college and university extension center in Milwaukee into
a single institution of higher learning as an integral part of the
University of Wisconsin under the governance of its Board of
Regents, and placed administrative authority for the new institu¬
tion in a Provost (later Chancellor) reporting directly to the
President. For purposes of this study, however, the most signifi¬
cant legislative provision was that “. . . this unit of the univer¬
sity . . .” (shall have) “. . . the same degree of self-government
by its own faculty as is vested in other units of the university/’3
The political solution in Madison to the Milwaukee problem
of educational consolidation was a pragmatic decision, as are
most political acts. The Executive Committee of the “Committee
of Thirty” (composed of representatives of the three institutions
Wisconsin Statutes, 39.024 (3) (h).
1972] McLaughlin — Faculty Government of University 19
involved in the merger) commented in its final report of imple¬
mentation recommendations to the Board of Regents that the
Legislature’s language had been . . probably fortunately far
from precise.”4 The Committee noted its difficult preoccupation in
arriving at decisions regarding organization of UWM to empha¬
size autonomy while insuring the integrity of the total University
of Wisconsin. Developments in succeeding years were to demon¬
strate continuing shifts in this delicate balance of academic gov¬
ernment which the basic legislation and initial implementation
had attempted. Changing powers, structure, and external relation¬
ships of the Milwaukee campus faculty seem to have been inevitable.
Because there was no other unit of the University which was
comparable to the Milwaukee institution and because of the psy¬
chological and physical separation of the Milwaukee faculty from
the rest of the University faculty in Madison, the implementation
document suggested that the UWM faculty would have “. . . a
smaller degree of participation in affairs considered by the total
University faculty, and (2) a larger degree of self-government
than “. . . existed in other units of the University.”5 Although
notices of the Madison-based general faculty meetings were to be
sent to Milwaukee faculty members,6 it was clear that they were
effectively disenfranchised. This was especially irksome to UWM
faculty members in search of their own unique identity, in view
of the Regents’ definition :
The faculty of the Milwaukee unit, operating within policies and standards
governing the University as a whole, and its several units, shall hold
meetings at regular intervals (1) to discuss matters which require action
by the [general] University faculty and to make recommendations
thereon; and (2) to take actions on matters which are within established
University policy but which relate to the Milwaukee campus only.7
The Regents did make one concession to the Milwaukee institu¬
tion’s different character and tradition inherited from its state
college predecessor institution : Faculty membership was extended
to those holding the rank of instructor. (The general University
faculty and its co-terminous Madison campus faculty excluded
academic staff members below the rank of assistant professor.)8
Milwaukee faculty members anxious to assert their own self-
governing identity may have overlooked a major result of the act
of merger. As interpreted in an official opinion by the state attorney
general, a three-way merger had been effected.9 The two Milwaukee
4 Summary Report of the Actions Leading to the Establishment of The University
of Wisconsin — Milwaukee (Madison, Wisconsin, 1957), p. 1.
5 Summary Report, p. 57.
6 Summary Report, p. 48.
7 Summary Report, p. 29.
8 Laws and Regulations Governing the University of Wisconsin (Madison, Wisconsin,
1951), 4.112.
20 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
institutions had become one with the University of Wisconsin. A
university system had been created which would eventually op¬
erate as a federal academic government. Of greater immediate
importance, Milwaukee faculty members had become instant heirs
of a long and strong tradition of faculty authority over educational
affairs. In over a century, University faculty members had achieved
a remarkable degree of self-determination over courses, degree
programs, and personnel matters subject to usually only nominal
administrative and regent approval. By legislative enactment and
bylaws of the Regents,10 the immediate government of the Univer¬
sity had become the province of the faculty.
The contemporary structure and scope of faculty authority in
the University is a product of intensive self-examination around
the turn of this century. University historians Merle Curti and
Vernon Carstensen point out that in the waning years of the nine¬
teenth century “The faculty was not only a legislative body but
a judicial and, to some degree, an administrative agency as well/’11
Following a critical controversy in 1910 over the respective roles
of the faculty and the Regents in educational policy decisions, the
faculty adopted in 1916 a committee recommendation which
“. . . went a long way toward solving the problem of maintaining
democratic faculty control over educational policy and of relieving
the teaching staff from routine matters”12 of administrative imple¬
mentation. In essence, the plan recognized the faculty’s direct legis¬
lative interest in policy formation based on investigation and
recommendations of a new faculty standing committee, called the
University Committee. Under the new rationale, an Adminis¬
trative Committee composed of the President, other University
administrators and the Secretary of the Faculty would super¬
vise the execution of routine matters. To complete the separation
of powers in academic government, the plan classified other fac¬
ulty authorized committees according to whether their chief func¬
tions were policy determining or administrative. The basic ration¬
ale of the 1916 faculty reorganization has continued to underlie
the philosophy and practice of faculty government.
But if the new faculty government at UWM was an heir, it
was also a parent of change. The merger of 1956 precipitated a
series of academic government revisions in structure and relation¬
ships which affected the total University system, the Madison
9 Attorney General, Wisconsin Statutes , 39.024 (3) (h), cited in West’s Wisconsin
Statutes Annotated (St. Paul, Minnesota, 1966), Vol. 5 and in Wisconsin Annotations
(Madison, Wisconsin, 1960).
10 Wisconsin Statutes, 36.02 (1), 36.06 (1), and 36.12.
u Merle Curti and Vernon Carstensen, The University of Wisconsin, A History:
1848-1925 (Madison, Wisconsin, 1949), Vol. I, pp. 608-609.
12 For a succinct account of this action, see Curti and Carstensen, Vol. II, pp.
105-107.
1972] McLaughlin — Faculty Government of University 21
campus operation,, and other units of the University. In a 1958
special report to the UWM faculty, its University Committee ac¬
knowledged successful operation of UWM self-governance within
its legally required framework as an integral part of the total
University. But the report called for a reaffirmation of the fac¬
ulty's traditional prerogative to have charge of academic affairs
and for appropriate safeguards to insure the faculty role in policy¬
making. At the same time, the University Committee insisted that
. . faculty committees should reverse the trend toward greater
concern with administrative detail and non-policy matters by in¬
sisting that these functions be carried out by administrative per¬
sonnel/'13 in keeping with the traditional role of the faculty of the
University. Viewed as a major faculty committee statement on
the future of the University of Wisconsin — Milwaukee, the 1958
report is perhaps surprising in its mild and scant mention of de¬
sirable modifications in academic government relationships within
the University.
Despite the initial guarantee of faculty self-government, the
operation of academic affairs at the Milwaukee campus for the
first few years was essentially a branch or satellite activity of
the Madison-based faculty. An elaborate system of inter-campus
conference committees to coordinate curricular and personnel ques¬
tions began to break down in complexity. Requirements for review
and approval of Milwaukee campus faculty policy decisions be¬
came steadily more irksome. Finally in 1963, the UWM faculty
set the stage for the University system-wide reform in academic
government which has continued to the present. It adopted pro¬
posals which would (1) make the UWM faculty the final faculty
approval body for curricular programs, (2) authorize the UWM
faculty to inaugurate Its own campus committees to review course
proposals, (3) establish a discrete Madison campus faculty, (4) es¬
tablish a University of Wisconsin faculty for consideration of gen¬
eral policy matters which affect all campuses of the University,
and (5) recognize the UWM faculty as having final faculty juris¬
diction over Milwaukee campus matters. Approval by the general
University faculty and by the Board of Regents contributed to a
comprehensive overhaul and codification of University rules.
Today, the Laws and Regulations of the University of Wiscon¬
sin provide for the academic government of a University federal
system. General “constitutional” provisions relating to the whole
University set forth basic statutory laws and Regent bylaws,
describe the operation and jurisdiction of the system-wide Uni¬
versity Faculty Assembly, state system-wide rules, and prescribe
13 Special Report of the University Committee — Milwaukee on the Future of the Uni¬
versity of Wisconsin — Milwaukee , UWM Faculty Document 55, May 26, 1958.
22 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
a minimal legal framework for legislation by unit faculties, such
as the UWM faculty. Legislation adopted by the UWM faculty,
following approval by the Regents, is embodied in a set of chapters
reserved for the Milwaukee campus government.
Operation of Faculty Government
Structural Elements of the Polity. The faculty government of
The University of Wisconsin — Milwaukee operates as a deliberative
body and as a cluster of subordinate committees. Meeting as a
body, the faculty legislates academic policy decisions usually based
on investigations conducted by subject matter committees. Begin¬
ning with the 1969-70 academic year, the faculty delegated its
powers and jurisdiction to a representative senate, between meet¬
ings of the faculty. An examination of meeting practices of both
bodies reveals to a large degree the identifiable roles, predictable
continuity, and group goals of UWM faculty government.
Sessions of the Body : Faculty Meetings . The initial regulations
governing UWM faculty meetings14 were a brief adaptation of
the existing rules of the parent University of Wisconsin faculty.
With the preoccupation of integrating the two diverse faculties
and academic programs into the University system, UWM faculty
members apparently were satisfied with minimal and non-original
rules of procedure in the early years of the institution. Succeeding
years brought piecemeal changes, culminating in a codified exposi¬
tion in 1967 and a major experimental revision of faculty govern¬
ment processes in 1969.15
According to current faculty adopted and regent approved legis¬
lation, the faculty meeting is parliamentary, democratic, systematic,
and definitive. But in the light of empirical observation of recorded
experience, the faculty meeting is sometimes licentious, authori¬
tarian, anarchic, and indecisive. Viewed from the perspective of
fourteen years, a meeting of the UWM faculty is remarkably simi¬
lar to other public deliberative bodies. Like other legislative polities,
the faculty places increasing faith in an increasing corpus of com¬
plex rules directed toward simplistic ends. To some extent, it shares
the common communication mystique which assumes that Truth
and Understanding are inevitable products of free and unlimited
verbal confrontation. A summary look at faculty procedures im¬
plemented in practice illustrates its strengths and weaknesses.
14 Proposed Regulations Governing University of Wisconsin — Milwaukee Faculty
Meetings by University Committee — Milwaukee, UWM Faculty Document 1, February
6, 1957.
15 The University Faculty — Milwaukee, UWM Faculty Document 384 (revised),
March 9, 1967 and The University Faculty — Milwaukee and its Senate, UWM Faculty
Document 485 (revised), March 20, 1969.
1972] McLaughlin — Faculty Government of University 23
From its first organizational meeting on September 14, 1956
through the last session of the 1969-70 academic year, the UWM
faculty was convened as a body 123 times. Ninety nine of these
sessions were regularly scheduled monthly meetings; twenty four
were additional special meetings. (With the inauguration of the
senate, the full faculty held only two prescribed regular meetings
during 1969-70, with four additional special meetings.) Legal
membership of the faculty exactly doubled during the fourteen
years of this study: from 310 to 620. (In 1964-65, except for
those who enjoyed “grandfather’s rights,” instructors were dis¬
enfranchised by system-wide rules and the net membership de¬
clined by eight from the previous year.)
Because the faculty convenes as a “town meeting” with no
regular quorum requirement, attendance is affected by the urgency
of issues, the intensity of feeling of special interest groups, or
ceremonial obligations. The percentage of those attending ranged
from 25% to 37% during the first year, and the range has not
varied significantly in the succeeding thirteen years. The all time
low of 7% was recorded at a meeting which debated important
basic portions of the University code concerning personnel, UWM
faculty government, and the system-wide University Faculty As¬
sembly.16 Two special meetings tied for the second low percentage
of participating attendance : One adopted the UFA legislation, and
the other passed a resolution in opposition to the Vietnam war.17
The record high attendance figures were achieved at two special
faculty meetings in response to emotional campus issues. Recruit¬
ing policies and attendant student protests concerning the Vietnam
conflict brought out 69% of the faculty, plus 70 student visitors.
At the height of campus disruption associated with the national
student strike of May, 1970, 64% of the eligible faculty debated
and passed a series of resolutions greeted by the jeers and cheers
of an estimated 250 students in a standing room only auditorium.
In neither of these two most highly attended faculty meetings in
fourteen years were the stated consensus or adopted resolutions
binding or effective on University policy and action.18
Although the official minutes of faculty meetings record only
fragmentary or sporadic excerpts of meeting dialogue, veteran
faculty meeting-goers conclude that a small number of members
dominate discussion and debate. Categories of frequent vocal par¬
ticipants include spokesmen for the “safe” positions of the faculty
“establishment,” apologists for militantly activist causes of a
16 Minutes,. Regular Meeting of the UWM Faculty, March 9, 1967.
17 Minutes , Special Meeting of the UWM Faculty, March 21, 1967 and Minutes,
Special Meeting of the UWM Faculty, November 14, 1969.
18 Minutes, Special Meeting of the UWM Faculty, November 27, 1967 and Minutes,
Special Meeting of the UWM Faculty, May 14, 1970.
24 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
para-educational nature, self-appointed defenders of real and
imagined “oppressed” minorities of faculty opinion, and chronic
participants indulging a need for public recognition. Explanations
for non-attendance and non-participation by the overwhelming
silent majority probably range from passive apathy to active dis¬
gust. Regardless of motivations, faculty meeting government obvi¬
ously is a minority exercise.
Although faculty meeting attendance and participation are pre¬
dictably unpredictable, the content of agenda items is relatively
certain. Faculty rules provide that, except by unanimous consent,
business at a faculty meeting is limited to written proposals in
proper form which have been included in the prepared calendar
(agenda) distributed in advance. The original mechanism desig¬
nated the Administrative Committee to prepare the calendar, in¬
cluding only those matters under the jurisdiction of the faculty.
Although that criterion is still implicit, a later codification dropped
the stated requirement. The latest rule assigns the task of calendar
preparation to a faculty committee elected by the senate. As a
further indication of faculty preoccupation with democratic due
process, regulations assure that any matter omitted from a cal¬
endar shall be included in the calendar of the next regular meeting
by affirmative vote of those present.19 Calendar regulations were
intended to insure advance familiarity with issues, parliamentary
efficiency, and protection of individual rights. But periodic com¬
plaints of arbitrary and capricious actions by the former Adminis¬
trative Committee and the current Calendar Committee pose a
continuing and unresolved question for the future of faculty gov¬
ernment.
The problem of appropriateness of faculty meeting business is
substantive as well as procedural. Controversy about access to
faculty meeting deliberation is matched by controversy about the
appropriateness of the issues themselves. The question is not
merely academic; it is at once political and social. UWM faculty
members are not merely state employees; they are also public
officers. The transacted business of a faculty meeting is not merely
University educational policy; it is also public policy. The provi¬
sions of the Wisconsin Anti-Secrecy Law, as interpreted by the
attorney general, are applicable to meetings of the UWM faculty
as to those of other public bodies concerned with the transaction
of governmental business.20
19 Chapter 31, The University Faculty — Milwaukee and its Senate, Laws and Regu¬
lations of the University of Wisconsin. 31.04 (4) (c).
20 Attorney General Bronson C. La Follette, letter to University of Wisconsin Presi¬
dent Fred Harvey Harrington, December 23, 1968.
1972] McLaughlin — Faculty Government of University 25
A review of the calendars and minutes of the meetings of the
UWM faculty during its early years shows only rare recourse to
special meetings devoted to issues of dubious or uncertain faculty
jurisdiction. Until 1959-60, the faculty deviated from its preoccu¬
pation with academic programs only once; it passed a resolution
to the Regents protesting the inauguration of fee parking facilities
on campus and claiming “faculty control” of University affairs as
a traditional right.21 Only two of the five special meetings of the
1959-60 academic year22 involved strongly controversial matters,
and both concerned academic policy: ROTC and a discussion of
the loyalty oath provisions of the National Defense Education Act.
In general, through the first eleven years, the UWM faculty’s
handling of controversy was parliamentary, democratic, systematic,
and definitive. Appropriate issues of scholarship, campus planning,
grading systems, and related academic subjects were deliberated
with little public notice. But in 1967-68 and ensuing years, a
public mood of division and unrest was equally apparent on the
campus and in meetings of the faculty. Public issues became Uni¬
versity issues, and faculty controversies became public contro¬
versies.
However historians of the future may judge the conduct and
results of faculty meetings of our recent past, the pragmatic con¬
clusion of many faculty members from their limited perspective
must have been that the general faculty meeting as an academic
government device was tried and found to be wanting. Impatience
with reliance on reasoned discourse, the raising of peripheral and
emotional issues, or aborted attempts to inject such subjects into
faculty deliberations often seemed to arise in frustration and to
end in frustration. The crisis in faculty government perhaps was
inevitable. The growth in numbers and complexity of the institu¬
tion had increased the probability that the faculty meeting as a
“system of communication”23 would fail. George Reed Field’s
doctoral study had found that “. . . only slightly over 50% of
the Milwaukee faculty members reported that they had substantial
authority to participate in academic policy determinations” and
that in other critical areas of institutional relationships Milwau¬
kee faculty members had a greater degree of dissatisfaction than
did their well established counterparts on the Madison campus.24
UWM academic government also seemed to have become a classic
21 Minutes, Special Meeting- of the UWM Faculty, December 3, 1958.
22 Minutes , Special Meeting- of the UWM Faculty, September 24, 1959 and Minutes,
Special Meeting of the UWM Faculty, December 17, 1959.
23 George Reed Field, Satisfaction and Dissatisfaction of University of Wisconsin
Faculty Members by Campus Location, Ph. D. Thesis (University of Wisconsin, Madi¬
son, 1965), p. 11.
24 Field, p. 128.
26 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
case of “executive-legislative conflict/’25 In this period of strife,
the conflict was only partly the traditional campus administration-
faculty clash over authority. In a larger sphere, the authority of
the established faculty parliamentary system and leadership
seemed to be as much in question as the authority of the admin¬
istration. In the waning years of a decade in which most American
college and university presidents reported increased faculty in¬
fluence as the most important campus change,26 the UWM faculty’s
perception of its role was confused and divided.
The testing and at least tentative resolution of the role of the
faculty meeting in governance began in November of 1967. A
special meeting of the faculty was called to consider a resolution
attacking administrative policy on demonstrations by students
against job recruiting agencies associated with the Vietnam con¬
flict. In a ruling subsequently supported by the faculty Codifica¬
tion Committee,27 the presiding officer ruled that the issue was
limited to discussion without formal action. The angry response
of proponents was the adoption of a resolution which distilled the
essence of faculty division. In a regular meeting attended by 13%
of the eligible members, the faculty directed that the Chancellor
establish a student-faculty committee to inquire into matters of
university autonomy, academic freedom, and decision-making. The
“Committee of 32” was to be concerned specifically with the rela¬
tionship of the University to critical public issues and political
action on campus.28 Without official sanction as a formal faculty
committee,29 the vague alliance of faculty members and students
went about its investigation with no definite conclusion.
The underlying issues raised in the “Committee of 32” resolu¬
tion erupted again in a special meeting in March of 1969. Approxi¬
mately 18% of the faculty and an uncounted number of students
listened to an emotional debate which ended with the adoption
of three resolutions and adjournment fifty minutes after the regu¬
lation time. Again the faculty motions reaffirmed the “validity of
the Wisconsin tradition of shared faculty power,” called on the
administration to consult with faculty committees prior to acting
contrary to recommendations, and requested administrative and
regent permission to enable the admission of black students who
had been expelled from Wisconsin State University— Oshkosh fol-
25 A. Clarke Hagensick, ‘‘A Propositional Inventory of Executive-Legislative Con¬
flict,” Transactions of the Wisconsin Academy of Sciences , Arts and Letters , Vol. LVI i
(1967-68), 81-92.
28 Carnegie Commission on Higher Education, report quoted in New York Times
News Service dispatch, The Milwaukee Journal, July 19, 1970, p. 4.
27 Minutes, UWM Codification Committee, February 5, 1968.
28 Minutes, Regular Meeting of the UWM Faculty, December 14, 1967.
20 Minutes, UWM Codification Committee, January 23, 1968.
1972] McLaughlin — Faculty Government of University 27
lowing a disturbance at that institution.30 One week later, another
special meeting adopted motions which would develop a degree¬
granting Center for Afro-American Culture. Counter-proposals,
charges of “institutionalized racism,”31 and a dramatic walkout of
dissident faculty members and students were observed in silence
by a majority of the faculty.
The minutes of the regular meeting of March 20, 1969 consti¬
tute an understated record of a peak of strong feeling which ac¬
companied the almost continuous intrusion of faculty authority
issues. The main scheduled business of the meeting was considera¬
tion of a recommendation of the Codification Committee for amend¬
ment of the charter chapter of UWM faculty government to
provide for the creation of a faculty senate. Consideration of the
proposal previously had been delayed at the last regular meeting
by discussion of the Oshkosh students case. Before the senate
proposal was referred finally to the full faculty (and subsequently
approved by mail ballot), it was subjected to a barrage of sub¬
stantive objections and extraneous verbal maneuvers. Interrup¬
tions included attempts to permit students to speak on a petition
circulated earlier, challenges to rulings by the chair, frivolous
amendments, a premature motion to adjourn, a call for a vote
recount, and requests for parliamentary rulings.32 The conduct
and atmosphere of this and previous meetings probably contributed
to final approval of the senate as a partial replacement for general
faculty meetings.
The controversy over controversy continued into the final year
covered by this study, in spite of and because of the creation of
a faculty senate. In two regular and four special meetings, the
faculty debated Indo-China resolutions, response to a student strike,
sustained senate action on the academic year calendar, and pro¬
hibited the senate from amending provisions of the basic charter
of faculty government.
Sessions of the Body : Senate Meetings. A summary of the steps
leading to the creation of the UWM faculty senate33 reveals a
lengthy, deliberate, and democratic process in the achievement of
faculty government goals. Because of political realities, the senate
represented an evolution in faculty governance rather than an
abrupt departure. The persuasive case rested on a compromise
solution to schedule two regular meetings of the general faculty
each year and to delegate interim legislation to the smaller repre-
30 Minutes, Special Meeting- of the UWM Faculty, March 6, 1969.
31 Calendar and Minutes, Special Meeting- of the UWM Faculty, March 13, 1969.
32 Minutes, Regular Meeting of the UWM Faculty, March 20, 1969.
33 Documents Leading to the Establishment of the Faculty Senate at the University
of Wisconsin — Milwaukee, undated compilation, Office of the Secretary of the Faculty,
UWM.
28 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
sentative body. In response to faculty fears of loss of participatory
rights, the power of review of senate actions was retained by the
faculty, senate meetings were opened to non-voting participation
by non-senators, the right to convene special meetings of the
faculty was retained, and senators would be bound to conduct their
sessions according to general faculty rules. Finally, the senate
meeting mechanism was frankly proposed as experimental legisla¬
tion, subject to future faculty review and modification. The limited
experience of the first year of operation can only suggest some
tentative characteristics.
With a quorum requirement of a majority of its membership of
47 and a roll call provision, the freshman year senate maintained
an attendance ranging from a low of 68 to a high of 82 per cent.
In another evidence of increasing concern with “pure” faculty
governance, the senate elected its own president pro tem to preside
in the absence of the Chancellor. Meetings generally adhered
closely to established parliamentary rules and reasoned debate,
after some initial uncertainty over jurisdiction and procedures.
Reflecting in part its composition of faculty members of higher
academic rank and greater seniority, however, the initial senate
spent little time expressing doubt about its authority in academic
policy determinations.34 Legislative actions involved internal mat¬
ters of faculty concern, except for the adoption of a motion provid¬
ing for committee study of academic cooperation among the several
University campus units in southeastern Wisconsin.35 This action
was taken despite the contrary advice of a visiting officer of the
Milwaukee campus administration who asked for delay.
The Faculty Committee System. One of the first acts of the UWM
faculty was to define a system of faculty committees. The process
of definition has continued at an increasing pace through the period
of this study. Certain general themes and trends are readily ap¬
parent; they include standardization of codified committee descrip¬
tions, consolidation of outmoded and overlapping committee func¬
tions, recognition of distinctive Milwaukee campus problems,
procedures, and goals, emphasis on committee membership by
election rather than by administrative appointment, fluctuating
interest in providing for student involvement, and identification
of committee authority, responsibility, and accountability. The
basic charter of the UWM faculty declares that it “. . . may delegate
functional authority and responsibility to committees . . . ; however,
such bodies . . . are accountable to the University Faculty —
Milwaukee which retains final jurisdiction over all educational
matters. . .”.36 A separate Milwaukee unit chapter of University
34 Field, p. 128.
35 Minutes, Regular Meeting of the UWM Faculty Senate, January 8, 1970.
33 Chapter 31, 31.02 (4).
1972] McLaughlin— Faculty Government of University
29
laws and regulations specifies provisions for the establishment and
regulation of both standing (permanent) and special (ad hoc)
faculty authorized committees.37 This chapter also prescribes the
membership and functions of the current roster of standing
committees.
Committees exercise an important and sometimes confused role
in academic government. Most business transacted in meetings of
the faculty or its senate is based on informational reports of
committee activities or specific recommendations to the faculty
legislative body. But much committee activity is in separate imple¬
mentation of faculty delegated duties. Although the Secretary of
the Faculty is administratively responsible for mechanical details
of the committee structure and operation, each committee reports
directly to the faculty or its senate. Problems of interpretation of
functions, conflicts of jurisdiction, charges of usurpation of powers,
failures to carry out specific assigned responsibilities, and disagree¬
ments between committees are either ignored by neglect, resolved
by private negotiation, or subjected to a faculty body vote. In the
absence of any real judicial mechanism, there is no alternative to
these options. For example, when a joint meeting of the University
Committee and the Codification Committee failed to resolve the
question of applicability of general committee regulations to the
University Committee,88 the issue was adjudicated by a vote of the
senate.39
The existing committee structure is a product of a flurry of
codification activity which began in 196640 and extended through
the 1969-70 academic year. A significant part of this activity was
due to the intervention of the University Board of Regents in re¬
quiring or requesting faculty legislation affecting or involving
students. Faculty committees on student conduct were defined in
accordance with regent instructions. In 1969, the Regents requested
the various unit University Committees to investigate and to de¬
velop procedures for greater student involvement in broad educa¬
tional matters. A subsequent survey by the UWM University and
Codification Committees led to greatly increased student member¬
ship on a number of faculty standing committees.41 Earlier efforts
to provide for student representation in the committee element of
faculty government were essentially ineffective, with the notable
exception of the active Student Life and Interests Committee.
37 Chapter 34, Milwaukee Campus Committees.
38 Minutes, Joint Special Meeting of the UWM Codification Committee and the
University Committee — Milwaukee, September 18, 1969.
39 Minutes, Regular Meeting of the UWM Faculty Senate, October 9, 1969.
40 On December 5, 1966, the Secretary of the Faculty issued a memorandum in¬
forming the chairmen of all faculty committees that faculty legislation required a
self-study report of functions and membership to the Codification Committee by Feb¬
ruary 1, 1967.
41 Action by the University Board of Regents, July 25, 1969.
30 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
“Meaningful participation in . . . university government is not guar¬
anteed merely by the presence of students on committees, . . .”42
but the experience of another state university system with unit
campus governance suggests that “the development of increased
student participation must . . . grow naturally. . .”.43 After four¬
teen years, the starting point for student participation in UWM
academic government appeared to have been committed to
committees.
Conclusions and Predictions
To describe the shape and nature of the past and present is
difficult enough; to describe the precise pattern of the future is
absurd. We do not know what conditions will affect The University
of Wisconsin — Milwaukee in 1984 — nor in 1983, for that matter.
We cannot do more than enjoy the good-humored warning of
McGeorge Bundy’s prophetic report from an “Academic Utopia”
of 1975 in a pseudo-retrospective look at our shortcomings of the
1960s.44 But any future UWM faculty government will be derived
from its past. Here, then, are some major conclusive assessments
with their likely predictive corollaries.
1. The judicial functions of a democratic academic government
are not being served as an identifiable activity or are being exer¬
cised on a sporadic and non-systematic basis. These normal judicial
functions include: (a) determination of compliance with prescrip¬
tive and proscriptive rules, (b) interpretation of the meaning/intent
of discrete rules and provisions, (c) reconciliation of apparent con¬
flicts or inconsistencies among rules, and (d) judicial remedy for in¬
dividual grievances in cases of illegal acts, usurpation of powers in
the practice of faculty government or failures to act as required.
This study suggests that judicial concerns will continue to be faced
in something less than a comprehensive rationale.
2. The “doctrinal anti-administrative attitude”45 as a faculty
characteristic may be expected to persist in the muted form of an
“arm-length” communicative relationship between faculty govern¬
ment representatives and members of the campus administration.
This study suggests that traditional anti-administrative feeling
is becoming more translated into anti-faculty establishment au¬
thority bias by individuals and groups of faculty members who are
alienated from the silent majority which supports gradual change.
42 “Draft Statement on Student Participation in College and University Govern¬
ment,” American Association of University Professors Bulletin (March 1970), 35.
43 Executive Vice President John W. Oswald, University of California, letter to
Professor Kirk R. Petshek, Chairman, UWM Codification Committee, March 23, 1970.
44 McGeorge Bundy, “A Report from an Academic Utopia,” Harper’s Magazine (Jan¬
uary 1962), 10-15.
43 T. R. McConnell, “Faculty Interests in Value Change and Power Conflicts,”
American Association of University Professors Bulletin (September 1969), 346.
1972] McLaughlin — Faculty Government of University 31
3. The modest but increasing* trend to increasing exercise of
legal power by administrative and regent levels to cope with im¬
mediate and potential campus problems may be expected to persist,
“lacking faculty action”46 of specific appropriateness and acceptable
speed. The future may see a reversal of the traditional “faculty
proposal — administratiop/regent disposal” process unless faculty
anticipatory behavior involves more than the adoption of resolu¬
tions on public issues which spill over into campus controversy and
disruption.
4. During the period covered by this study, UWM faculty gov¬
ernment became increasingly codified into a systematic rationale
of fixed and delegated authority. But the academic tradition of
free and extensive dialogue is so strong that action oriented faculty
government leaders and campus administrators will continue to find
it difficult to heed Bobert M. Hutchins’ conclusion that “durable
action” in university governance requires “patience.”47
5. The structure and operation of UWM faculty government
has changed gradually into its present form and practice. Con¬
fronted with the impact of persistent or recurring social disruption,
it will continue to evolve. The major faculty goal of self-
determination of institutional uniqueness through parliamentary
democracy will continue to dominate its individual and collective
behavior.
40 “The Harrington Resignation,” Wisconsin Alumnus (June 1970), 9.
47 Robert M. Hutchins, “The Administrator Reconsidered,” reprinted in Adminis¬
trative Control and Executive Action, edited by B. C. Lemke and James Don Ed¬
wards (Columbus, Ohio, 1961), p. 65,
THE EFFECT OF RESTAURANT SERVICES ON THE SURVIVAL
RATE OF TOURIST-LODGING ESTABLISHMENTS IN WISCONSIN
L. G. Monthey and R. A. Ricketts
Introduction
The number of tourist-lodging- (T-L) businesses in Wisconsin
has been decreasing steadily since the mid-1950s, when the total
was in excess of 8,000 establishments. However, the greatest de¬
crease has occurred in the years since 1964, when the rate of de¬
cline accelerated to about 3% a year. As Figure I shows, every
major region of the State lost T-L businesses between 1964 and
1968; and the total loss during the period was about 870.
Previous studies have shown that the largest declines during
the Sixties were in the small, seasonal lodging businesses. On the
other hand, year-round establishments — particularly of the motel
or motor-hotel type — -have gained both in numbers and size.
Meanwhile, the State's total capacity in bedroom units (B.U.) has
remained near 80,000 for at least 10 years.
The apparent reasons for these general trends in the T-L indus¬
try have been discussed elsewhere in some detail. However, one
of the most frequent observations is that the small lodging opera¬
tion doesn't provide enough service to the traveler or vacationist
and hence finds it increasingly difficult to compete in the lodging
market. Or, to put it another way, it has been postulated that the
T-L establishments which offer the most services are the ones
most likely to stay in business longer.
This study is an attempt to test the latter supposition by se¬
lecting a single important factor, i.e. food service, and determining
its relationships to the survival of lodging establishments over a
recent 5-year period 1964-68. This factor was selected because it
could be readily identified through the issuance of restaurant
permits by the Wisconsin State Board of Health. During each of
the years concerned, the presence (or absence) of such permits
was cross-referenced on the records of all lodging-business inspec¬
tions made by the Board of Health. Thus, reliable counts of T-L
businesses holding restaurant permits were available for each year
of this study, and the type, size and seasonality of each establish¬
ment having restaurant services (RS) was readily determined.
Incidentally, it is our belief that those T-L establishments that
offer food service are more likely to provide various additional
33
34 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
LODGING ESTABLISHMENTS (1964 - 1968)
Figure I. Tourist-Lodging Establishments in Wisconsin by Regions.
(Comparing September 1964 with September 1968)
services as well. If this is true, then the restaurant permits may
serve to identify those establishments which offer, in general, a
greater array of guest services to their special clientele and other
travelers too.
Figure II shows the regional distribution of restaurant permits
in Wisconsin, comparing 1964 and 1968 figures. These data include
all types of food-service businesses with the exception of temporary
operations, such as hot-dog stands at county fairs, which obtain a
special short-term permit. It should be noted that the total number
1972] Monthey and Ricketts — Effect of Restaurant Services 35
RESTAURANT PERMITS (1964 - 1968)
January 1964 with January 1968. (Temporary Restaurant Permits are not
included.)
of restaurant operations, unlike the T . L establishments, remained
fairly constant during the 5-year period. All regions of the State
showed modest gains, with the exception of the southeast district
(including Milwaukee) which lost 200 food establishments, a de¬
cline of 8.3%. However, the over-all increase, statewide, was only
0.9%. Of the 13,330 restaurant permits in 1964, 11.1% (1,484)
were issued to T-L establishments. In 1968, when the total was
13,447 permits, 10.5% of them (1,414) were associated with
lodging business.
36 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
In order to obtain basic data on the relationship between restau¬
rant services and the number, type and size of T-L establishments
involved, Wisconsin Board of Health inspection records and mailing
lists for the years 1964 through 1968 were used. The appropriate
data were then coded for each establishment, transferred to IBM
cards, and the results compiled by data-processing techniques.
Five major categories were used in classifying T-L businesses:
(1) Hotel type; (2) Motel type; (3) Resort type; (4) Cottage type;
(5) Other. The distribution of restaurant permits among the vari¬
ous T-L businesses was then determined for all types and sizes of
establishments. The data are presented in a series of tables, which
follow.
General Findings
Table I summarizes the findings of this 5-year study relative
to the restaurant permits issued to all T-L establishments. In 1964,
19.3% of all T-L businesses had food services, whereas in 1968 the
corresponding proportion was 20.7% However, this small increase
of 1.4% may not be of much significance, since the sum total of
lodging establishments had declined from 7,690 in 1964 to 6,821
in 1968, a net decrease of 11.3%.
In view of this general decline in the number of T-L businesses,
a more definitive measure of the effect of restaurant services on
the survival of lodging establishments would be the comparative
rates of decline for establishments with and without food service.
For example, if the net decrease during the 5 years is 11% for all
T-L businesses, but is only 4.8% for those establishments with
restaurants (as in Table I), we can assume that food service is
associated with a lower business-mortality rate in the case of such
establishments. Similarly, Table I shows that the net decrease in
Table I. Total of Lodging Establishments with and without
Restaurant Operations (1964-68).
(RS = Restaurant Services)
1972] Monthey and Ricketts — Effect of Restaurant Services 87
T-L businesses without restaurant facilities (using 1964 as the
base year) was 12.9% — a drop that is somewhat greater than the
general rate of decline for the industry.
Table II illustrates the rates of increase or decrease in the
number of establishments, grouped into size categories by bedroom
units (B.U.), with and without restaurant facilities. Within the
1-4 bedroom unit grouping, both establishments with and with¬
out restaurant facilities have decreased. However, using the num¬
ber of 1964 establishments as a base, there appears to be a greater
rate of decrease in those establishments without restaurants
(—19.1 percent) in 1968 compared to those which provided restau¬
rant facilities (—13.5 percent) in 1968. In the 5-19 room grouping
this same relationship is also apparent. Those establishments with¬
out restaurant facilities had, by 1968, decreased to 89.6 percent
of the number of lodging operations existing in the base year of
1964. Meanwhile, those establishments with restaurant operations
declined only slightly and still comprised 97 percent of the base
year.
The next size grouping (20-29 rooms) indicates a reversal of
the relationship illustrated in the first two size groupings. Although
there was a decrease in establishments both with and without
restaurant facilities, the greatest decrease, using the 1964 base,
had occurred in those establishments which provided restaurant
facilities. Establishments with restaurant facilities decreased to
91.4 percent of the 1964 base, while 1968 establishments without
restaurants represented 98.9 percent of the base-year total. This
reversed relationship becomes even more apparent in the next size
grouping of 30-99 rooms. Here establishments which provided
restaurant facilities have showed a slight decrease to 98.9 percent
of the base year, while those without restaurant facilities increased
to 128.8 percent of the 1964 total.
Finally, the last size grouping (100 B.U. or more) showed no
change in establishments without restaurants, while there was an
increase of 17.3 percent in establishments with restaurant facilities.
In looking at all classes of lodging establishments, it appears
that as one moves from smaller size categories to the larger, the
trend in number of establishments without restaurant facilities
changes from a negative to a positive relationship relative to the
base year. However, the trends in those establishments which pro¬
vided restaurant facilities become less clear as one moves from the
smaller to the larger establishments. It appears that restaurant
facilities (in the house) may play a less significant role in medium-
to-large establishments than they do in the smaller establishments
(if we omit the 100-plus category).
38 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Table II. Trend in Lodging Establishment Numbers, with and without
Restaurant Services, by Various Size Groups (1964-68).
(RS = Restaurant Services)
1. SMALL ESTABLISHMENTS:
2. MEDIUM-SIZE ESTABLISHMENTS:
3. LARGE ESTABLISHMENTS:
1972] Monthey and Ricketts — Effect of Restaurant Services 39
The Relationships Vary
What possible conclusions might be drawn from the above data?
First, it appears that within the smaller size categories (1-19 bed¬
room units), the decrease in lodging establishments and the lack
of restaurant facilities is directly related. It could be hypothesized
that the lack of such facilities has resulted in the greater rate of
decrease for establishments without restaurants versus those with
restaurants. An alternative hypothesis might be that those estab¬
lishments which do not have restaurant facilities are economically
worse off than those which do provide restaurant services. One
could say that the greater decrease in the case of smaller estab¬
lishments without restaurant facilities is more likely to be a result
of economic considerations than the fact that they do not provide
such facilities.
Conversely, those smaller establishments which do provide res¬
taurant facilities might do so only because of better economic suc¬
cess with their lodging operation, which may account for the lower
rate of decrease in the number of these establishments. To deter¬
mine if the restaurant operation in smaller establishments has
enhanced their survival, or if the success of the lodging operation
has resulted in expanding into restaurant services, still remains
open to consideration.
Secondly, with establishments of 30 to 99 bedroom units, there
seems to be a trend toward constructing establishments without
restaurant facilities in the house. At any rate, there has been no
net increase in establishments with restaurants (in fact a slight
decrease), while there has been a substantial net increase (within
this size grouping) of establishments without restaurants. There
has been a tendency for new motels in this medium-size range to
rely on “outside” restaurants near the premises.
The trend in large establishments with more than 100 B.U.
showed no change in number of lodging operations without res¬
taurants. However, a net increase of eight establishments having
restaurant facilities was recorded over the 5 years, a gain of 17
percent.
At this point, it might be concluded that the net increase in
operations containing restaurant facilities (1.4 percent of total
establishments) can be attributed to the large net decrease in
smaller establishments without restaurants. The effect of such a
net decrease has been to increase the proportion of existing
establishments which provide restaurant facilities.
Hotel-Motel Businesses
Table III illustrates the relationship of restaurant facilities to
types of establishments. Again, using the base year of 1964 as a
40 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
standard of comparison, it appears that provision of restaurant
facilities is related to both increases and decreases within the
several types of lodging: establishments.
The hotel category shows a significant difference between the net
decrease rates for establishments which do and do not provide res¬
taurant facilities. Hotels without restaurants showed a greater net
rate of decrease ( — 16.9 percent) than those which do provide
restaurant facilities (—4.9 percent). Again, the cause-and-effect
relationship is unknown and can only be speculated.
Table III illustrates that motels with and without restaurants
have increased over the base year 1964. The rate of increase,
however,, has been greater for motels without restaurant facilities
( + 9.6 percent) than those which provided restaurant facilities
( + 3.9 percent). This seems in line with the findings in Table II
in that there has been a net increase in establishments without
restaurants within the 30-99 bedroom unit size group. Many of
the new motel entries fall within this size grouping, thus account¬
ing for the larger net increase of establishments without restau¬
rants in the motel category. This still does not answer the question
why motels are not emphasizing restaurant facilities, at least in
this size category. The most probable explanation appears to be
that these medium to medium-large motels are locating in areas
where there is an independent restaurant operation available in the
immediate area , there being no necessity to provide a restaurant
facility for the guest's convenience. Other considerations might
include past experience, space allocation and available capital.
Table III. Trend in Numbers of Lodging Establishments with and
without Restaurant Services, Grouped by Type of
Operation, 1964-68.
1972] Monthey and Ricketts — Effect of Restaurant Services 41
Other Establishments
There appears to be no significant relationship in the decrease
of establishments with and without restaurants for the resort
category, and only a slight relationship is found within establish¬
ments classified as “cottages.” However, there appears to be a
pronounced relationship between restaurant facilities and the
survival of establishments classified as “Other.” In this case, inter¬
pretation should be approached with caution. Those establishments
(classified as “Other”) without restaurants have decreased 19
percent and those with restaurant facilities have increased 1.9
percent. The important fact to recognize is that this category is
composed of a great number of heterogeneous types of establish¬
ments which could not be classified in the above categories. It would
be unwise to make comparisons because of the widely differing
characteristics of the various establishments.
Table IV illustrates the change in average size, measured in
bedroom units, of lodging operations which do and do not provide
restaurant facilities. The data indicated that lodging operations
with restaurant facilities increased in size an average of 2.0 bed¬
room units per establishment from 1964 to 1968, while those
without restaurant facilities have increased an average of only
0.7 bedroom units per establishment. Table IV also indicates that
T-L establishments with restaurant operations are more sizable
businesses (21.9 bedroom unit average) than those without res¬
taurant facilities (9.1 bedroom unit average).
Table V provides a more detailed analysis of the average size
of operations. Both seasonal and year-around operations with and
without restaurant facilities are considered. Between 1964 and 1968
there was no significant difference between the increase in size
of seasonal establishments with restaurants (0.1 bedroom units)
and those without (0.2 bedroom units). However, note the differ¬
ence in the average size of seasonal businesses with restaurants
for 1968 (14.5 bedroom units) and seasonal operations which did
not provide restaurant facilities (7.4 bedroom units).
Table IV. Changes in the Average Size of Lodging Establishments
WITH AND WITHOUT RESTAURANT SERVICES, 1964-68.
(RS £= Restaurant Services)
42 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Table V. Average Size Change for Seasonal and Year- Around Lodging
Establishments, with and without Restaurant Services, 1964-68.
Year-around operations show a more pronounced difference rela¬
tive to the change in average size of establishments with and
without restaurants for the five year period. Although the differ¬
ence is slight, operations with restaurants have increased 2.9
bedroom units while those without restaurant facilities have in¬
creased only 2.1 bedroom units. As with seasonal operations, the
average size of year-around operations with restaurant facilities
is much larger (30.5 bedroom units) than those establishments
without restaurants (15.1 bedroom units).
Highlights
Slightly over 10% of the 13,400 restaurant permits issued by
the State of Wisconsin are issued to tourist-lodging (T-L) estab¬
lishments.
Slightly over 20% of Wisconsin’s 6,800 T-L establishments
obtain a restaurant permit and provide a food service for their
guests. This study included all of the establishments that were in¬
spected by the Wisconsin State Board of Health during the years
1964 to 1968, inclusive. It attempts to determine relationships
between the availability of food service at T-L establishments and
the subsequent increase or decrease in the numbers of such
establishments.
There was a net decrease of 11.3% in the total number of lodging
businesses between 1964 and 1968. However, the State’s total
housing capacity for travelers, in terms of bedroom units (B.U.),
remained substantially the same over the period. Those establish¬
ments which provided restaurant services (RS) declined 4.8%
during the 5 years, while those without RS decreased 12.9%.
Small T-L establishments (1 to 19 B.U.), as a category, showed
a much higher survival rate where restaurant facilities were pro¬
vided. It was 94.4% after 5 years, compared to only 85.7% where
no food services were offered.
1972] Monthey and Ricketts — Effect of Restaurant Services 43
Medium-sized T-L establishments (30 to 99 B.U.) seem to be
placing less emphasis on restaurant services “in the house.” During
the 5-year period the number of properties with food service
remained virtually unchanged, while those without restaurants
increased from 104 to 134, about 29%.
Lodging establishments with 100 or more B.U. showed no decline
in the small number that did not provide RS between 1964 and
1968. However, the total for those that had restaurants increased
by 17%.
The number of hotels without restaurant facilities declined 17 %
during the 5 years, while those with food service dropped only
5%. Motels, on the other hand, gained in both categories; those
with restaurants gained 4% in number, and those without such
facilities increased by 10%.
The biggest declines in numbers occurred in the case of cottage-
type establishments. Those with food service showed a net de¬
crease of 13.5%, while those without restaurant permits dropped
20% between 1964 and 1968.
Both seasonal and year-round establishments which provided
restaurant services were considerably larger than those which did
not have them, averaging about 100 percent bigger in both groups.
References
Fine, I. V. and R. E. Tuttle. (1963) The Tourist Overnight Accommodations
Industry in Wisconsin. Wis. Dept, of Resource Development and the
University of Wisconsin, Madison.
Monthey, L. G. (1964) The Resort Industry of Wisconsin . Wisconsin Academy
of Sciences, Arts and Letters Transactions, Vol. 53 (Part A), pp. 79-94.
Monthey, L. G. (1969) Trends in Wisconsin’s Tourist-Lodging Industry.
Wisconsin Academy of Sciences, Arts and Letters Transactions, Vol. 58
(1970), pp. 71-99.
THE REMAKING OF “AMERICAN LITERATURE”
Donald Emerson
''American Literature” in quotation marks, for one must dis¬
tinguish between the raw bulk of what may be included by gen¬
erous definition, and the more or less official "American Literature”
of which I wish to speak* Generous definition has come to include
many things not always accorded status as contributory to the
literature— for example, the types of fiction, verse, and orally-
transmitted tales and anecdotes these days designated as The
Literature of Popular Culture. This is partly a matter of intellectual
class distinctions, but if New England once had a Brahmin caste,
the general levelling tendency has brought it low, to the point where
articles are now written on the decline of WASP culture in the
United States. And in much the fashion of anti-discrimination
statutes and compensatory programs for the disadvantaged in the
social sphere, there is these days an effective attitude of intellec¬
tual anti-discrimination, accompanied by active attention to the
literature of ethnic minorities. Both tendencies lead today to an
active redefinition of "American Literature.”
It may be foolish to speak of a more or less official American
Literature when in fact no such thing exists. There is no pres¬
tigious Academy-of-Something-or-Other to sanctify American lit¬
erary gospels, no Federal-Office-of-This-or-That to proscribe un¬
worthy works. The literary historian who should attempt to de¬
fine the scope of American literature of the past half-century by
reference to the most distinguished literary awards would make
himself laughable. Even the international Nobel Prize guarantees
Mrs. Pearl S. Buck no standing whatever. As is customary in the
United States, a vague consensus is reached in quite unsystematic,
even chaotic,, fashion. This shifting consensus loosely defines
"American Literature” at any given moment. The active parties
include the writers, scholars, and critics whose judgments are
effective with the mass of relatively passive culture-customers. The
Dun and Bradstreet volumes of this "American Literature” are
the anthologies, which reflect the decline of once successful ven¬
tures and the establishment and consolidation of new enterprises.
Anthologies exist in great variety to meet differing demands
and expectations. I refer particularly to the anthologies imposed
on students who wish to understand something of the growth and
development of literature in the United States. In the context
45
46 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
of this discussion “American Literature” (note the continuing
marks of quotation) refers to that minimum of authors and se¬
lections the anthologist feels he must include to keep himself
honest. Or to put it another way, “American Literature” for this
discussion includes whatever the anthologist feels he can’t afford
to leave out for the students who are to have only an overview
of the field, with little expectation of more intensive studies. I am
talking about the “American Literature” of college sophomores,
not of doctoral candidates, and I note that instructors further
anthologize. They select for assignment only part of what must
seem, to those sophomores, God’s plenty in excess of all reason¬
able appetite. In this they contribute to the consensus which pro¬
duces for each year’s class an “American Literature” which has
undergone an annual model change. The students make their con¬
tribution by responses on the scale from apathy to enthusiasm,
and not many instructors willingly repeat assignments which even¬
tually bore even themselves in the absence of student response. And
there comes a point where every anthology is beyond hope of
revision and must simply be abandoned, for “American Literature”
has become something other, and the addition of new selections
will not modernize it.
Two principal approaches to the problem have been made
through literary history and esthetic judgment, neither of which
ever operates apart from the other, and both of which have greater
and more continuing importance than the temporary pressures
of the times. Of political history Carl Becker remarked that “His¬
tory is not what happened but what we think about what hap¬
pened,” and I suggest that the dictum applies to literary as well
as to political history. Both are efforts at understanding from
the vantage point of distance. But greater distance alters the
landscape of the past, and as Becker further observed, each gen¬
eration must rewrite its histories, and, in the literary context,
also remake its anthologies as the result of remaking its histories.
In the process, “American Literature” is effectively changed. I
obviously now mean by “American Literature” what each genera¬
tion, or, if you will, each historian understands American litera¬
ture to be. Content and emphasis both vary. As Professors Henry
Pochmann and Gay Wilson Allen point out in the Introduction to
their Masters of American Literature, the history has been seen
from various points of view : Vernon Louis Parrington read Ameri¬
can literature as an essentially native phenomenon to be under¬
stood in the continuing clash of the forces of liberalism and con¬
servatism. Howard Mumford Jones related the literature to three
animating forces in American culture: the cosmopolitan spirit ;
the frontier spirit; and the middle-class spirit. Oscar Cargill dis-
1972] Emerson — Remaking of “ American Literature i
47
cussed the literature in terms of “ideas on the march,” while Nor¬
man Foerster examined the interplay of foreign importations
and native conditions in terms of four broad factors : Puritanism ;
romanticism ; realism ; and the frontier spirit.
Pochmann and Allen themselves emphasize a diversity which has
received even greater emphasis in the twenty-some years since they
wrote :
For upwards of two centuries Boston and Cambridge held a position
of primacy, and the conventional history treated American literature as
the peculiar province of New England, with the result that these earlier
studies read strikingly like histories of Harvard College .... Scant
attention was given to the cavalier tradition of the South . . . , to the
literary coteries in Baltimore, Richmond, and Charleston during the
nineteenth [century], or, for that matter to the dissenting groups in
Puritan New England herself; while the evaluation of so-called “foreign”
(that is to say, non-English) strains or influences in American literary
culture was entirely neglected until recently.
I shall wish to pursue later that matter of cultural diversity in
emphasizing trends which have become clearer since Pochmann
and Allen wrote, but for the moment I wish to cite the evidence
of a variety of approaches to the presentation of “American Litera¬
ture” to the consuming public. I shall have to cite, at possibly
tedious length, the statements of anthologists in the prefaces to
their awkwardly-heavy volumes. And I shall cite dates, for they
seem to me significant. Mr. Pochmann and Mr. Allen made their
very sensible statement in 1949, after World War II. Three state¬
ments of principle published shortly before it may illustrate the
diversity of possible approaches, the ways in which, late in the
decade of the Depression, two respectable editors could insist on
the sole validity of their estheticism while others clung to their
sense of a social and historical approach, and yet a third composed
his excellent anthology in anticipation of a coming war against
totalitarianism.
1938 ; William Rose Benet and Norman Holmes Pearson, editors
of The Oxford Anthology of American Literature:
A man may look at writing as he chooses. We have regarded it as litera¬
ture. Undoubtedly by the introduction of a social approach, an interest
in the history of American letters has been enormously stimulated. This
has been occasioned partly by a general concern with social matters
and social history; but it has been mostly seized on with a defensive
enthusiasm for one quality when the presence of another, the purely
literary was not certain. While the endowment of a novel with proletarian
significance, or the identification of an essay with the deistic movement,
or the recognition of the spirit of democracy in a poem may form the
basis of useful estimates, they leave unanswered the stubborn questions
of literary values.
The editors themselves answer stubborn questions of literary
48 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
values only through their selections, and at this point in time some
of their discriminations seem, at the least, curious.
1939; Milton Ellis, Louise Pound, and George Weida Spohn in
A College Book of American Literature:
All the phases of our literary development have had careful considera¬
tion and an effort has been made to combine selections which embody
reflections of the political and social history of the age with those which
embody their authors’ best literary art, without allowing either tendency
to go to an extreme.
This is hedging one’s bets, but it is at least ninety degrees away
from the avowed estheticism of Mr. Benet and Mr. Pearson. A more
forthright declaration was made by Bernard Smith in The Demo¬
cratic Spirit, 1941. He was anthologizing for a nation about to go
to the wars:
The motive for the creation of this volume should be obvious. In these
days, when we are one of the few peoples that can still cherish democracy,
it behooves us to recollect our peculiar tradition and to review its growth
so that we may know exactly what it is and not be led into believing it
is something else. Here is a noble heritage. We must know it well if we
are not to lose it. There are always forces at work to deprive us of it.
By a later shift of interest which Mr. Smith could not have an¬
ticipated, his thirty-year-old anthology is more “modern” in the
world of 1971 than any which followed it for the next thirty years.
Because he worked from the tradition of American idealism he
included the work of such black writers as Frederick Douglass,
W. E. B. DuBois, Langston Hughes, Countee Cullen, James Weldon
Johnson, Claude McKay, and Richard Wright; and he also included
Sacco and Vanzetti, Michael Gold, and Albert Maltz, “minority
writers,” as they are all termed in the very latest anthology,
writers almost entirely ignored in other, and very successful
anthologies.
By 1956 Sculley Bradley, Richmond Groom Beatty, and E. Hud¬
son Long had produced The American Tradition in Literature,
probably the most successful anthology of the decade.
Our effort has been to represent major authors in the fullness of their
stature and variety. Besides the titans, we have included writers of
lesser stature whose works endure; but no author was introduced pri¬
marily for the purpose of illustrating literary or social history. In the
same way, works of popular literature and humor have been admitted as
literature, not as social or cultural documents.
By 1967, there have been further recognitions for the third
edition :
Since time has brought changes in the prevailing evaluations of American
literature, we endeavor to reflect these here, preserving at the same time
that organization and editorial attitude which has best served the ever-
1972] Emerson— Remaking of u American Literature J
49
evolving American tradition in literature. The most significant revaluation
is, perhaps, the desire to study in greater depth certain nineteenth-
century masters — Hawthorne, Melville, Thoreau, Emerson, and Henry
James. Another revaluation results in the greater attention paid to the
best writers of our own century.
So it goes, and here is the latest statement, from Literature in
America, 1971, under the general editorship of Robert C. Albrecht:
This anthology is not a social history despite an organization of topics
that could be considered extra-literary. For example, slavery and theology
were important to authors of the early period, and they wrote about them
in works of indisputable worth. On such issues, we have carefully chosen
selections that reflect both the concern of the authors and the quality of
their work. We accept a broad definition of literature, yet insist upon
the substantial differences between a social-problems reader and a litera¬
ture anthology.
The publisher is somewhat more forthright in proclaiming the
virtues of the anthology, in a letter to Professors of American
Literature :
Now especially, at a time of heightened racial and ethnic awareness, these
volumes can serve you remarkably well. The Modern Age, in the section
titled “Minority Reports,” contains the largest selection of works by
minority writers of any introductory literature text; included are works
by Black, Indian, Jewish, and Hispano- American authors. The Founding
of a Nation offers a number of works by and about Indians and slaves.
A Century of Expansion, also rich in material that bears heavily on
previously “hidden” aspects of our history, contains works by Black
writers, abolitionists, and fragments from the untapped riches of our
Indian heritage.
There is an obvious state of tension. The modern anthology is to
preserve literary values ; it is also to reflect the spirit of the age by
presenting the work of minority groups, and this is touted as one
of the claims of virtue. It may be virtue, but it is virtue lately
revealed.
Besides the changes in “American Literature” which are the
product of a changing sense of history — one’s way of understand¬
ing the past — -there are more purely literary considerations which
ought to be mentioned, for they, too, change “American Litera¬
ture.” It is not altogether a matter of that changing sense of liter¬
ary history which I mentioned earlier. For one, there is discovery.
The great example for our age is the discovery of the Dead Sea
Scrolls, but here at home there was an interesting development
in Puritan poetry. No anthology before 1937 includes the work of
Edward Taylor, for his manuscript volume passed unregarded
through generations of his family until Thomas H. Johnson found
it in the Yale library and transcribed the works. Every sophomore
now knows that Edward Taylor was the principal Puritan poet,
50 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
even if Taylor’s contemporaries and the scholars of two and a half
centuries didn’t.
Emily Dickinson is another interesting example. At her death in
1886 she had published few verses, but from manuscripts dis¬
covered after her death six hundred and sixty-eight poems were
brought forth in successive volumes between 1890 and 1945; and
in 1955 Thomas H. Johnson prepared the three-volume variorum
text which is the definitive edition of her work. That sophomore,
secure in his knowledge of the place of Edward Taylor, also knows
that Emily Dickinson was, with Walt Whitman, one of the great
poets of the second half of the nineteenth century, although her
reputation was made in the twentieth.
There is the other process of rediscovery or revival of interest.
Herman Melville suffered neglect until 1924, when Raymond
Weaver produced a biography and in the course of research dis¬
covered the manuscript of “Billy Budd.” Henry James was com¬
paratively unregarded from the second decade of the century
until 1934, when a Henry James issue of Hound and Horn marked
the beginning of an interest that created a scholarly industry.
When Malcolm Cowley set out to anthologize The Portable Faulk¬
ner in 1945, none of William Faulkner’s novels were in print,
although he had completed the bulk of his best work. The com¬
bination of the Cowley volume and the award of the Nobel Prize
in 1950 stimulated an interest, and it shows no signs of diminish¬
ing. Faulkner, like Taylor, Melville, Dickinson, and James, is
safely placed in “American Literature,” although the places of
all of them have been lately defined.
Conversely, there is the redefinition of the literature by elimina¬
tion. Everyone understands the disappearance of very popular but
admittedly poor work after its day has passed, but I refer now
to the destruction of great and lasting reputations when the intel¬
lectual and spiritual climate in which they flourished has passed
away. Henry Wadsworth Longfellow was the universally known
and admired poet of his time, and his reputation abroad was
immense; he is the only American honored in the Poet’s Corner
of Westminster Abbey. Yet it may not be too many years before
The New Yorker can carry a cartoon of one American tourist
looking up at the bust and asking another, “Who the hell was
Henry Wadsworth Longfellow?” unless, of course, the editors fear
the point will be lost on their readers. Longfellow is not men¬
tioned in the 1963 anthology Poetry in English of Warren Taylor
and Donald Hall which includes the work of twenty-eight Ameri¬
can poets beginning with Philip Freneau. Nor does the name of
James Russell Lowell appear. Yet the likenesses of both were
familiar to me from one of those framed holy pictures of Ameri-
1972] Emerson — Remaking of u American Literature ” 51
can middle-class culture heroes which adorned the parlor of my
childhood. My mother taught me their verses from memory. They
have been eliminated by one of those great changes of climate
which destroy whole species.
These discoveries and revaluations are part of what I consider
the perennial process of remaking “American Literature.” There
are conspicuous forces which act over shorter periods of time. In
the Thirties, the general concern with social problems led to great
emphasis on the literature of social action. A novelist like Thorn¬
ton Wilder was berated for writing of pre-Christian Greece when
Michael Gold demanded to know, “Mr. Wilder, are you a Greek
or an American?” and indicted him for frivolity in not writing
of the labor struggles of Depression America. The Seventies al¬
ready oppress me with a sense of deja vu, even if the terms have
somewhat changed, for the spokesmen of the New Left sound
remarkably like the Marxists of the Thirties, like Michael Gold,
or Granville Hicks before he left the Communist party. Again
there is the equation of literary value and social usefulness in a
cause. The New Leftists have made T. S. Eliot their test case.
Eliot is open to censure on several grounds — his snobbishness,
his anti-Semitism, his neo-fascist attitudes. A teacher of literature
who includes Eliot, it is now claimed, is hopelessly reactionary;
he should be teaching the literature of protest, or better, not
teaching literature at all, but the gospels of Herbert Marcuse or
Norman 0. Brown. Departments of English should teach sociology
or be abolished.
But this seems to me possibly as transitory an emphasis as that
of the Thirties. Far more noteworthy is the way in which, travel¬
ling down the Mississippi of our literature, we passed the Ohio
in the mid-Forties, and the Missouri in the early Sixties. I refer
thus metaphorically to recognition of the great contribution of
American Jewish writers and American Negro writers. I name
the Jewish writers first, for although their claims date only from
the turn of the century, they were earlier recognized; Black
Americans have a claim going much further back, but only since
the activist years of the Sixties have they passed that anthology
test. Now there is great eagerness to recognize the claims of other
minority groups. And this is good, but over the next decade we
shall see the general revaluation process operating, as it always
does. The “American Literature” of 1981 will require a new
anthology.
DISCONTINUITIES IN DEMOCRATIC SYSTEMS
AND MASS SOCIETIES
Charles Redenius
Introduction
The linkages between democratic political systems and mass
societies are explored in this paper. Specifically, an attempt is made
to show that there are serious discontinuities between the political
system and the social system by examining certain interactions of
these two systems. The paper itself is divided into three parts. The
first section is largely analytic. It attempts to elucidate the environ¬
mental and cultural features that led to the presence of a demo¬
cratic political system within the context of a mass society. The
driving force behind this development has been, and remains, eco¬
nomic modernization. The result of this movement into an indus¬
trial culture has been a serious dislocation between the political and
social spheres. Although it is widely recognized that certain condi¬
tions must be met before it is possible for a democratic political
system to emerge, it is not recognized to the same extent that these
conditions are not the same as the conditions that sustain a mature
democratic political system. Both sets of conditions are set forth
and explored in this section.
The second section of the paper describes the discontinuities
between the democratic system and mass society. It tries to do this
in three principal ways. First, it is argued that democratic theory
has remained very stable and has undergone little evolution since
the Industrial Revolution. This is not the case with the structure
of society. Indeed, it would not be incorrect to say that there have
been fundamental changes in the social system. The second method
used to describe the discontinuities between the political and social
system examines the internal structure of both a traditional society
and a mass society. Finally, the discontinuities are described by
noting the violations of the laws of social change. The conclusion
that is reached is that the dislocations occurred as the result of the
failure of democratic theory to evolve, rapid social change, and the
fact of inadequate response to the demands generated by the
industrial culture.
The third, and final, section is largely prescriptive. This part
of the paper tries to show what steps need to be taken in order to
bring about a realignment between the political and social systems.
53
54 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
The underlying assumption here is that the political system, or
more precisely, the government can bring about the necessary
changes.
Since this section is prescriptive, the values behind it should be
spelled out. These values are only made explicit here. Neither here
nor in the body of the paper are there any arguments to sustain
these values. They are simply posited. The values are : that abund¬
ance is better than want ; that ecological balance is better that eco¬
logical imbalance; that fraternity is better than prejudice; and
that peace is better than war.
The discussion in this part of the paper centers around four
primary areas; the need for new myths to replace the exploded
ones ; the strengthening of voluntary associations, the basic instru¬
ment of the democratic system; the need to stabilize our growing
population which aggravates myriad other problems ; and a neces¬
sary change in our foreign policy from militarism to economic
assistance. In short, we must more successfully address ourselves
to the changes, and the consequent problems, wrought by the
Industrial Revolution.
This research is cross-disciplinary or inter-disciplinary in nature.
The footnotes will provide ample evidence of that. With few excep¬
tions, however, the bulk of the research was done in either the
behavioral sciences, or those fields with a behavioral orientation.
Survey data from opinion polls conducted by both academic and
commercial pollsters were an important source that were used in
this analysis. The data from these polls were used in two ways:
first, as evidence supporting the arguments presented in the body
of the paper; and secondly, as an empirical “screen” for assump¬
tions made by authors who either did not utilize such data, or did
not consider such data germane to their subjects.
I
Democratic political systems have been in existence for roughly
three hundred and fifty years. We can date the emergence of demo¬
cratic political systems from the seventeenth century where the
conflict between absolutism and liberalism was first resolved in
favor of the latter.1 These democratic systems were linked with
traditional societies until the advent of the Industrial Revolution.
To describe the conjunction of the political system and the social
system in this pre-industrial era such terms as classical democracy,
aristocratic democracy, and even traditional democracy have been
1 John H. Hallowell, Main Currents in Modern Political Thought (New York: Holt,
Rinehart and Winston, 1950), p. 71.
1972] Redenius — Democratic Systems and Societies 55
used.2 Nevertheless, the root meaning of democratic theory re¬
mained the same regardless of the social system qualifier used. The
qualifiers though did introduce some confusion. This confusion cen¬
tered around the failure to distinguish clearly between what con¬
stituted the political system and what constituted the social sys¬
tem. Indeed, this failure to distinguish between the political and
the social has persisted to this day.3 Although not a part of this
paper, the same confusion and failure of discrimination exists in
regard to the linkages between the political system and the eco¬
nomic system in the minds of many people, and not all of them
laymen.
Thus, by keeping the concepts political system and social
system distinct in our minds, we can see that it is both possible
and probable that democratic political systems can be and will be
linked with social systems having radically different internal struc¬
tures. This does not mean that such conjunctions will necessarily
be harmonious ones. Indeed, the opposite is true in certain cases
where dislocations are bound to exist.
The Industrial Revolution can be viewed as the watershed in
terms of the internal structure of social systems. Prior to the Indus¬
trial Revolution there was only one type of society, the traditional
society, although many variations of this type existed. The advent
of industrial modernization destroyed this single form of social
organization. There now exists three primary types of social sys¬
tems.4 First, there still remains the traditional society which is
pre-industrial and has not yet begun the move toward economic
modernization. Next, there is the transitional society which is
gripped by internal conflict between the traditional elite and the
industrial managers over the issue of the modernization of the
economy. Finally, there is the modern, or the mass, or the industrial
society. The terms are used interchangeably in this paper. In this
form of social organization economic modernization has been, or is
virtually completed. There are, of course, many variations on each
of these major types. Indeed, in any organization as complex as a
social system it is unlikely that a “pure” type exists.
Thus, the Industrial Revolution created an environment that led
to the emergence of mass societies linked with democratic political
systems. This development has been accompanied by serious dis¬
locations between the political and social systems. The structure
2 Joseph Schumpeter, Capitalism, Socialism and Democracy (New York and Lon¬
don: Harper and Bros., 1947), Chapters xxi and xxii ; A. D. Lindsay, The Modern
Democratic State (New York: Oxford University Press, 1962), p. 12.
3 Sheldon Wolin, Politics and Vision (Boston: Little, Brown and Company, 1960),
pp. 286-94 ; David Easton, “An Approach to the Analysis of Political Systems,” World
Politics, IX (April, 1957), p. 388.
4 William A. Faunce, Problems of An Industrial Society (New York: McGraw-Hill,
1968), pp. 27-29.
56 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
of the social system linked with a democratic political system
changed radically while the structure of democratic theory under¬
went little change. It should occasion no surprise that the discon¬
tinuities in the linkages between the two systems stand in dire need
of realignment.
Despite the fact that industrialization led to the transformation
of the structure of social systems, it did not immediately work the
same transformation on the structure of political systems. Indeed,
the effect of industralization on the structure of political systems
was a conservative one. The structure of both democratic and non-
democratic systems remained relatively stable. Overall, industrial
modernization had two primary effects on political systems. First,
modernization did not lead to the decline of non-democratic political
systems. Secondly, the transformation of the social system that led
to discontinuities between the social system and the political system
in democratic systems produced the same results in non-democratic
systems.
Thus, the societal conditions that act as part of the linkage
between the political and social system, and that make a democratic
political system possible can be considered only as necessary con¬
ditions. They are not the necessary and sufficient conditions.
Indeed, the possibility of the emergence of a democratic system
does not mean the necessity nor does it indicate the degree of
probability of such emergence. These societal conditions, then, per¬
mit development of radically different political systems.5 The
impact of the Industrial Revolution, which transforms the struc¬
ture of society, merely continues the further development of these
different political systems. Historically, the political systems that
have continued their development since the drive toward economic
modernization began range from syncratic politics on the right of
the political spectrum to stalinist politics on the left.6 This develop¬
ment has included the linkage of democratic systems with economic
systems as different as capitalism, socialism, and the welfare state.
The linking conditions that make a democratic system possible
are not the same conditions that sustain a mature democratic
political system, that is, a democratic system linked with an indus¬
trial mass society. By examining first the conditions that make a
democracy possible, and then, the conditions that sustain a mature
democracy, it will be possible to observe the discontinuities in part
of the linkages between the political and social systems. The con¬
ditions that make a democracy possible are four in number. First,
5 Kenneth Boulding, The Meaning of The Twentieth Century (New York: Harper
and Row, 1964), pp. 175-76.
8 A. F. K. Organski, The Stages of Political Development (New York: Alfred A.
Knopf, 1967), pp. 7-16. The terms are broadened to include all totalitarian systems
regardless of their stage of economic development.
1972] Redenius — Democratic Systems and Societies 57
there must exist some measure of widespread economic security.7
In other words, the biological requisites of food, clothing, and
shelter must be met in a relatively satisfactory manner. Wide¬
spread poverty and want is not conducive to political freedoms.
The biological drives of survival and hunger are the two strongest
instinctual drives in human beings. If these drives occupy a posi¬
tion uppermost in a man's mind, he will not have time for, or be
concerned with, such luxuries as freedom of speech and freedom
of association. There exists a hierarchy of needs for human beings,
and economic needs must be satisfied before humans concern them¬
selves with political ideals.
Second, the society as a whole must have a relatively high
literacy rate.8 There are no precise parameters to indicate how
high this literacy rate must be, but it is safe to assume that it must
be well over fifty percent. Literacy is an essential element in the
socialization process that must occur if a society is to view itself
as a political unit, and as fit to govern that political unit. Without
a relatively high literacy rate, this socialization process cannot
occur among the entire society, it will be restricted to an elite.
Literacy makes possible the communication and interchange of
political ideas throughout the entire structure of society.
Third, an acceptance of the dignity of human life is necessary
if a democracy is to succeed.9 The first two conditions seem to be
necessary if this one is to be recognized. However, the notion of
an intrinsic moral worth of every individual is so central to demo¬
cratic theory that it must be stated separately. This condition pro¬
vides the framework for the ideal of political equality — that all men
are equal. More than this is meant, however. Men are more than
equal to one another, they are brothers. The relationship between
democratic citizens is one of fraternity. Non-democratic systems
are always paternalistic in some way. The relationship between
rulers and ruled in a democracy is one of equals. The same relation¬
ship is a non-democratic system is one of superordinate and
subordinate.
The final condition that makes a democratic system possible is
that there must be a widespread acceptance of the exchange sys¬
tem.10 This involves a consequent rejection of the threat system.
The key difference between a democratic system and a non-
7 Marian D. Irish and James W. Prothro, The Politics of American Democracy
(Englewood Cliffs, N. J. : Prentice-Hall, 1965), p. 80.
8 Lucian Pye, “Transitional Asia and the Dynamics of Nation Building,” in Marian
D. Irish (ed. ), World Pressures on American Foreign Policy (Englewood Cliffs, N. J. :
Prentice-Hall, 1964), pp. 154-72.
9 Weston La Barre, The Human Animal (Chicago: The University of Chicago Press,
1954), pp. 315-18.
10 Kenneth Boulding, The Impact of the Social Sciences (New Brunswick, N. J. :
Rutgers University Press, 1966), pp. 57—58.
58 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
democratic system in regard to this condition is that use of the
threat system is viewed as normal in a non-democratic system
whereas in a democratic system threats, force, and the use of
violence are considered extraordinary, and thus, illegitimate. The
basic instruments of democracy are the bargain and the compro¬
mise, which are the heart of the exchange system.
These, then, are the societal conditions that make a democratic
system possible. The democratic systems that emerged were linked
with a traditional society. With the coming of the Industrial Revo¬
lution the internal structure of society was radically altered.
Following industrialization, democratic systems were linked with
mass societies. The conditions that sustain a mature democratic
system, as noted above, are not the same as the conditions that
make a democratic system possible. By contrasting the sustaining
conditions with the necessary conditions, the dislocations between
the democratic system and the mass society will be readily appar¬
ent. The sustaining conditions are also four in number.11 First,
symbols and forms that have continuity and that speak men’s lan¬
guage, that is, excite their imagination, must exist. Economic mod¬
ernization is a traumatic experience. Without the appropriate
symbols, that shock may be more than the political system can
absorb. The process of modernization necessarily involves the
debunking of myths found in the traditional society. The modern
democratic system has failed to replace the symbols that have been
displaced.
Second, a mature democracy is sustained by a modernized econ¬
omy and culture. Modern democracies have faltered here because
industrialization is an uneven process. Certain parts of the econ¬
omy are left relatively unaffected by modernization. It is the task
of democratic government to direct the forces of industrialization
to those parts of the economy. A modernized culture is a by-product
of the industrial process. By removing economic backwardness,
democratic government insures that the culture does not remain
backward.
Third, there must be a reasonable distribution of wealth and
power in the community to sustain a modern democratic system.
The necessary conditions for a democracy call for a widespread
measure of economic security. In a mature democracy the concern
shifts from economic security to the distribution of wealth and
power. The concentration of wealth and power in the hands of
a few violates the pluralism that is an essential part of a modern
democracy.12 The power to redistribute the resources of a modern
11 Charles Frankel, The Democratic Prospect (New York: Harper and Row, 1962),
p. 24.
12 Ibid.
1972] Redenius — Democratic Systems and Societies 59
society rests only with the government. The mature democratic
systems have failed to act on this responsibility. By not acting in a
resolute manner democratic governments have actually encouraged
the drift toward greater inequalities in wealth and power. This
drift will not correct itself; governmental action is necessary to
redress the inequalities.
Finally, mature democratic systems are sustained by civil liber¬
ties and a framework of vigorous private groups and associations.
The basic civil liberties are effective instruments for maintaining
men's loyalty to a political system even when they disapprove of
many of the actions of that system. The erosion of civil liberties,
and the stifling of dissent, destroys that loyalty and undermines
the very basis of democratic government— government by consent.
The basic instrument of the democratic citizen, the instrument
through which he exercises his civil liberties, is the voluntary asso¬
ciation, Democratic governments have been vigorous in protecting
civil liberties. The same is not true for voluntary associations.
Thus, the exercise of one's civil liberties is often an exercise in
futility. Freedom of speech is ineffective without freedom of
association.
The role of the government, then, marks the greatest contrast
between the democratic system linked with a traditional society
and the democratic system linked with a mass society.13 In the
traditional democracy, the role of government is essentially a nega¬
tive one. It allows, or permits, and in rare cases, promotes, the
transition from a traditional society to an industrial society. In
the mass democracy, the role of government is essentially a posi¬
tive one. In addition to promotion, it must also regulate and super¬
vise many of the activities that occur in an industrial society. The
contrast between the necessary conditions and the sustaining con¬
ditions reveals this change in role very vividly. A large share of
the discontinuities between the democratic system and the mass
society can be directly attributed to the extreme reluctance of
both political leaders and followers in accepting this necessary
change in the role of democratic government. These vestiges of
cultural lag can be eradicated only by a social learning process
that will reeducate both the political leaders and the masses as to
the proper role of government in a mass democracy. However, be¬
fore this social learning process can occur, the seeming stability
of democratic theory in the face of extensive structural changes in
society must be broken down. That is the task of the second section
of this paper. But before turning to that task the figure presented
on next page will summarize the arguments of the paper to
this point.
13 Lindsay, op. cit p. 116, and pp. 245—48.
60 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
m - >-
Industrialization
Figure 1.
II
The evolution of the democratic political system has been re¬
tarded by the lack of fundamental change in democratic theory.
Despite the widespread changes occurring in the economic and so¬
cial systems, democratic theory has remained very stable. The
proof of this stability can be demonstrated by examining four
different facets, all of which concern democratic theory. First, the
authors writing about democratic theory today are merely restat¬
ing what authors since the seventeenth century have been saying.14
In some cases it is refinement of the basic tenets of democracy,
and in other cases, it is a working out again of the logical conse¬
quences of those basic tenets. In either case, however, the basic
tenets of democratic theory — popular sovereignty, limited govern¬
ment, and political equality — have remained virtually unchanged.
Next, democratic political systems have manifested different
institutional arrangements, yet the same theory is used regardless
of the institutions in the system.15 Indeed, a particular institutional
arrangement takes on a definitional aspect when it is defined as
something a democracy is not, such as, democracy is not separation
of powers, checks and balances, federalism, and judicial review.
This diversity of institutional arrangement in the political system
and the unitary quality of democratic theory seems to suggest that
the linkages between systems and theory are in need of further
exploration. If such an exploration were successful, it might be
found that different democratic systems are supported by different
democratic theories. On the other hand, the results might show
why a single theory permits diverse institutional arrangements to
develop.
Third, democratic systems have been linked with economic sys-
14 Frankel, ap. cit., pp. 33-48.
15 Schumpeter, op. cit., chap. xxi.
1972] Redenius — Democratic Systems and Societies 61
terns that are radically different.16 Thus, the same argument that
was used in the second point would seem to be valid here. Without
satisfactory knowledge of the linkages between the political system
and the economic system, it is impossible to say why democratic
systems are linked with different economic systems. The traditional
response to the problem posed here, and in the second point, is to
assume that democratic theory remains the same, that is, stable,
while the economic system evolves independently of the political
system. The fact of the matter might be that democratic theory
only appears to be stable, it is quite possible that it evolves right
along with the social and the economic systems.
Finally, the greatest revolution in democratic systems since the
Industrial Revolution, it seems, has concerned the role of the mass
citizenry. This revolution has often been confused with a revolution
in the tenets of democratic theory. However, that is not the case.
The tenets have remained the same. In the traditional democracy,
it was assumed that the citizen was highly interested in, and in¬
formed about, political issues. This atomistic individual made ra¬
tional decisions in pursuit of his self interest. The voting behavior
studies have shown that the mass citizenry do not conform to this
model.17 Despite these findings, the voting behavior studies have
not brought about dramatic changes in democratic theory. The
response has been a call for participatory democracy. The thrust
of this movement has been an attempt to remove the obstacles
blocking participation in the system . Through it all, again, demo¬
cratic theory appeared to remain unchanged.
Thus, the stability of democratic theory has contributed to the
discontinuities between the democratic system and mass society.
By examining first, the structure of a traditional society, and then
the structure of a mass society, the dislocations between the po¬
litical and social systems can be described in a second way.
A traditional society is characterized by extended family ties,
a face-to-face society, and mechanical solidarity.18 Social cohesion
and integration is the result of the sharing of a common culture
or way of life. In this type of society there is very little mobility
either in terms of geography or social status. Given this limited
mobility there are strong ties to, and a strong sense of, community.
Control of behavior is not normally a problem because of the
reliance of the internalization of the community’s norms. This is
16 Schumpeter, op. cit., chap. xxii.
17 Paul F. Lazarsfeld, et al., The People’s Choice (New York: Duell, Sloan and
Pierce, 1944) ; Bernard Berelson, et al., Voting (Chicago: University of Chicago Press,
1947) ; Angus Campbell, et al., The American Voter (New York: John Wiley and Sons,
Inc., 1960) ; Angus Campbell, et al., Elections and the Political Order (New York:
John Wiley and Sons, 1966).
18 Faunce, op. cit., pp. 28 and 170.
62 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
the result of a socialization process that stresses internal, or pri¬
mary, controls rather than external, or secondary controls.
One of the ideals of the traditional society was that of crafts¬
manship, or imitation. Thus, the pace of change was very slow.
Innovation was not a major virtue. Indeed, it occupied a very lov
place in the hierarchy of communal values, and in some cases, il
was even suspect. Thus, continuity, rather than change, was insti¬
tutionalized in the social system. The coming of the Industrial
Revolution transformed the structure of society. The process of
economic modernization destroyed the traditional society, and in
its place, a mass society arose.
Mass society means more than simply density of population. It
means a society where the traditional controls have been shat¬
tered.19 Control of behavior becomes a problem, and increasing
reliance is placed on external controls. The specific form of these
external controls is the modern bureaucracy. These controls are
necessary because there is confusion over the values of the society.
More particularly, a mass society means the breakdown of ex¬
tended family ties, and the loss of the face-to-face mechanical
society. These are replaced by the nuclear family, and the “face¬
less”, organic society. This type of society, for the most part, rests
on the fact of interdependence and not on a sense of community.
Social cohesion is imperfect and social integration is never fully
accomplished. The result is the loss of a sense of community. The
effects of this loss are compounded by the shift of the population
from small rural towns to large urban centers. Thus, the industrial
society is characterized by a high degree of mobility both in terms
of geography and social status. Social change is very rapid due to
the institutionalization of change at all levels of the society. Inno¬
vation replaces continuity as the leading virtue.
The democratic system fails to keep pace with the changes that
are occurring, and have occurred, in the social system. The result
is serious dislocations between the two systems. Up to this point
in time, the democratic systems have still failed to adequately re¬
spond to the challenges of industrialization.
The causes for the divergence between democratic systems and
mass societies can be described by a third method. That method
is to examine the violations of the laws of social change.20 First,
industrial societies, and non-industrial societies that have been
affected by economic modernization, have failed miserably to check
population growth. Even the most successful countries have only
slowed such growth. No country has stabilized its population nor
has any country been able to reduce its population.
10 Ibid.
20 Kenneth Boulding, The Organizational Revolution (Chicago: Quadrangle Books,
1968), pp. 77-80.
1972] Redenius — Democratic Systems and Societies 63
Second, although it is widely known today that the culture
patterns of a society are transmitted mainly through the instru¬
ment of the family, the transition from a traditional to a mass so¬
ciety altered the basic structure of the family almost without
notice. This alteration in basic structure left the family unable
to adequately carry out its historic function of acculturation. Due
to the fact of rapid social change, society also seems incapable
of developing the necessary adjunct to the family that would work
together with the family to carry out this socialization process.
Third, organizations of all kinds have an optimum size. In the
social system today, the most acute problem is how to organize
large masses of people without sacrificing liberty and even decency.
The political system is faced with the breakdown of its hierarchy.
Large democratic organizations with an elaborate hierarchy are
faced with a dual problem: the failure of communications from
ruled to ruler (and vice versa) ; and the obvious fact that hierarchy
violates the democratic principle of political equality.
Next, there is the social law of oligopoly. This law states that if
the number of independent interacting organizations is few a
situation of acute instability and conflict will be created. The eco¬
nomic systems of most mass democracies are dominated by
oligopolistic firms. The international political system is dominated
by three or four great powers. The instability in both of these
systems is too obvious to need further comment.
On the other hand, the social law of instability states that the
uncontrolled interaction of a large number of organizations pro¬
duces unacceptable consequences. The Great Depression, for ex¬
ample, was one of those unacceptable consequences. Thus, the
political system, or more precisely the government, must steer a
middle path between the instability caused by oligopoly and the
instability caused by unregulated interaction.
Finally, there is the social law of the persistence of role. Despite
the transformation of the social system, the political system has
persisted in its role developed for an earlier time. This problem is
essentially the same one that has been discussed from two points
of view; the stability of democratic theory, and the changes from
traditional society to mass society. Therefore, any further discus¬
sion of this problem will be unnecessarily redundant.
To say that the causes of the divergence between the political
and social systems are clear does not mean that the remedies are
going to be easy to implement. Indeed, here is the crux of the
problem. We have not developed any instruments that can exercise
effective control over these causes. The laws of social change are
known to us, but are not controlled by us. The techniques and
innovations that are necessary to insure the continuance of demo-
64 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
cratic systems within the context of a mass society have not yet
been discovered, or if discovered, have not been fully utilized and
implemented. We have for too long* a period of time worked at cross
purposes with our biological and social natures. The challenge of
our times is to see that mass democratic systems meet and master
the problems generated by the industrial culture. If the mass
democracies fail, the world will surely turn to the obvious alterna¬
tive, the non-democratic systems, to view their response to this set
of problems.
To summarize the conclusions up to this point, we can state
that the discontinuities between the democratic system and the
mass society occurred as the result of the failure of democratic
theory to evolve, the transformation of the structure of society,
and the violation of the laws of social change. Here is a classic
example of cultural lag. The ideas of society have not kept pace
with technical innovation. This is a consequence of social inertia,
that is, a tendency to persist in past modes of thought, and vested
interests, a conscious attempt on the part of a few to block change
in order to preserve the status quo. It is the task of the final section
of this paper to point out some ways which will reduce cultural
lag and bring about a better alignment between the political and
social systems.
Ill
The relationship between the democratic system and the mass
society as this paper has shown is not one of isomorphic models.
Societal changes have led to discontinuities between the two sys¬
tems. The problem, simply stated, is to remove those discontinui¬
ties. The solution to this problem will not be a final one. The best
that can be hoped for is that we can bring our problems under ra¬
tional control and then keep them manageable.21 It seems rather
senseless to waste our time searching for permanent solutions. In
a dynamic system change is inevitable. But, given the fact that
change is inevitable, we can learn to cope with the dislocations
that will occur. Our first step must be to bring the industrial
culture under rational control.
An optimum strategy would seem to call for at least the imple¬
mentation of the following four points. All of these points call for
vigorous leadership by the government. The need for such positive
government was demonstrated earlier. As these points are opera¬
tionalized other features may be called to our attention that need
amelioration. But the time is past for continued speculation, there
is a need for action now.
-1 Charles Lindblom, “The Science of ‘Muddling Through’,” in Raymond Wolfinger
(ed. ), Readings in American Political Behavior (Englewood Cliffs, N. J. : Prentice-
Hall, 1966), pp. 211-226.
1972] Redenius — Democratic Systems and Societies 65
First, there should be a deliberate attempt made to foster new
symbols and ideologies for our democratic system.22 We need new
myths to replace the ones exploded by the process of industrializa¬
tion and by the transformation of society from a traditional base
to a mass base. This problem is an especially acute one since politics
has replaced religion as the central concern of our time.23
It is useless to try to reconstruct the solidarity of the face-to-face
society. The unit upon which society relied for the transmission of
cultural patterns, the extended family, has been destroyed for the
most part by the Industrial Revolution. Even if the extended family
had remained intact throughout the transition it would still be im¬
possible to retain the solidarity of the face-to-face society because
the values transmitted by such a family structure are the ones that
were broken down in order that the process of economic moderniza¬
tion might occur. There is no going back, new values must be fos¬
tered, not old ones recaptured. The family unit, now the nuclear
family, will still be the primary transmitter of values, but it will be
transmitting new values.
What must these new values be like? First, it must be accepted
that these values stand somewhere between science and religion,
but not contradicting either. Second, these new values must be
viewed as relative values and not absolute ones. There is no set
hierarchy involved, nor is there a definite content to these values.
We must learn to live with openness (uncertainty) and forsake
comprehensiveness. Finally, we must also forsake some of the logi¬
cal constraints of ideology. Democratic values necessarily involve
a tension between competing values such as liberty and order, or
excellence and equality. There can be no final resolution of these
tensions. The incomplete nature of these values allows for their
continued evolution. This is a process that is unending if the les¬
sons of history remain true.
Second, the basic instrument of the democratic citizen, the volun¬
tary association must be strengthened.24 The failure of the demo¬
cratic system to provide mechanisms that will enhance private
groups is one of the discontinuities between the political and social
system that is felt most strongly. As mentioned earlier, one of the
key differences between traditional society and mass society is the
transition from the face-to-face society to the “faceless” society.
In this transition there is lost what is perhaps the heart of the
traditional society— a sense of community. A sense of community
is built on voluntary rather than economic relationships. In the
mass society, economic relationships dominate interactions between
22Frankel, op. cit., pp. 20-22.
23 Ernst Cassirer, The Myth of the Stale (New Haven: Yale University Press, 1961).
2i Frankel, op. cit., pp. 64-71.
66 Wisconsin Academy of Sciences, Arts and Letters [VoL 60
citizens with only one clear exception, the nuclear family. Economic
relationships do not seem capable of providing the milieu in which
a sense of community can develop. Part of the inadequacy of demo¬
cratic theory is its failure to recognize the need for a sense of com¬
munity, that is, vigorous private groups and associations. The prin¬
cipal tenets of democratic theory are clearly negative in character.
They have allowed or permitted the rise of a mass democracy but
they have not fostered nor encouraged the voluntary associations
that would make for a healthy mass democracy.
Specifically, how do we strengthen voluntary associations ? First,
we must counter non-democratic forms of organizations in our pri¬
vate groups. These groups are the training grounds for democracy,
and if they are allowed to deteriorate the socialization process nec¬
essary for democratic participation is sure to deteriorate also.
Second, participation in a voluntary association must foster more
than a sense of involvement, it must also foster a sense of efficacy.
Democratic governments can encourage this development by recog¬
nizing the vital role voluntary associations play. Private associa¬
tions are effective intermediaries between the democratic citizen
and democratic government when the government is influenced by
the opinions of the group. Refusal by the government to listen to
the petitions of private groups is sure to lead to feelings of aliena¬
tion and helplessness. Finally, democratic governments can encour¬
age voluntary associations by providing easier access to the mass
media. The technical facilities for strengthening the organizations
of private groups are available. We are not making full use of our
technical capabilities at a time when it is most important that we
do so.
Third, there is a crying need to stabilize the population of mass
democracies.25 As noted earlier, the problem of controlling the size
of the population is basically unsolved ; no country has a stationary
population. Without a stable population it will be nearly impossible
for a democratic (or a non-democratic) government to cope with
the institutional problems of an industrial society. Such ecological
problems as air, land, and water pollution cannot be brought under
control when confronted with ever greater and greater demands
upon these resources. Many of the urban problems of our society
such as waste disposal, education, crime, traffic, and housing are
aggravated by population pressure.
The problems of population pressure go beyond national bounda¬
ries. Population control is perhaps mankind’s most serious long-
run problem. We have been practicing death control in both devel¬
oped and under-developed countries. Indeed, one of the first
23 Boulding, The Meaning of the Twentieth Century , pp. 121-37.
1972] Redenius — Democratic Systems and Societies 67
“benefits” of a developed society that is introduced into an under¬
developed country is death control. Unfortunately, neither society
is practicing birth control. One of the effects of unrestricted popu¬
lation growth is the exacerbation of international conflicts. Who
can forget Hitler’s cry for “Lebensraum.” Factions who oppose
birth control must face the fact that they are engaged in danger¬
ous warmongering.26 One author goes so far as to say that the
only sound biological solution to the problem of war is massive
depopulation.27
It would seem that population equilibrium will be achieved only
if we can accomplish some of the following : first, we must reform
those factions who oppose birth control. This will mean, for ex¬
ample, that the Catholic Church must be convinced of the necessity
of adopting a more realistic attitude. Next, we must create social
institutions to control population growth. It is social institutions
which are dominant in determining population growth and not
mere individuals with knowledge of the physiology of reproduction.
Thus, the “pill”, or other methods of contraception, alone are not
enough. Such groups as Planned Parenthood must be fostered, and
others created, to insure efficacious population control. Finally, a
vast reeducation on the role of marriage and the family in society
must be undertaken. Marriage and the family must be viewed now
and in the future as primarily an institution of companionship and
not procreation. For this reason, the Women’s Liberation Move¬
ment ought to be supported.
The fourth and final step that is proposed here as necessary to
realign the democratic system and mass society calls for a shift
from a militaristic foreign policy to one based on economic assist¬
ance.28 It seems a stark fact of life that huge defense expenditures
have not increased our security. Increasing defense expenditures
beyond what they are today would merely continue a counter¬
productive policy. Internally, outsize defense expenditures have se¬
riously distorted our domestic priorities. Social and economic in¬
equalities persist, and in some cases, are exacerbated by lack of
funding for programs designed to eliminate or reduce these inequal¬
ities. In terms of international relations, the “traditional ’ policies
have been miserable failures. This is especially the case with “Third
World” countries. Racism and poverty are at least as important
a factor as ideology, whether left or right.29 The white-have world
cannot continue to confront the non-white, have-not world with
a policy designed to contain communism. That policy does not fit
20 Ibid. ; Desmond Morris, The Naked Ape (New York: McGraw-Hill, 1967), p. 178.
27 Morris, op. cit., p. 177; E. E. Schattschneider, In Search of a Government (New
York: Holt, Rinehart and Winston, 1969), p. 14.
28 Edward H. Carr, The New Society (Boston: Beacon Hill, 1951), p. 92.
29 Ibid.
68 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
the facts very well. We desperately need to return to a foreign
policy similar to that carried out under the Marshall plan. Simply
stated, instead of continuing arms shipments we should shift to
capital investment and technical aid in a scheme of planned inter¬
national trade. This would enable underdeveloped countries to
combat totalitarianism by meeting the requisites of democracy.
The surest defense against communism is economic security and
not military security. This does not mean we should neglect military
security but rather that we should recognize that it is of a lower
priority than economic security. The limitations of, and after a
point, the counter-productive nature of military security should
have alerted us a long time ago to the need for a more viable de¬
fense policy.
Conclusion
The crisis of our times is not, as commonly depicted, one of ideo¬
logical confrontation. It goes far deeper than that and actually
dwarfs ideological clashes. Indeed, the crisis is more “revolution¬
ary” than the struggles that are occurring. The crisis of our times
is the fact of inadequate response on the part of mass democracies
to the challenges generated by the industrial culture.30 This paper
has made an attempt to analyze the way in which the discontinui¬
ties between the political and social system developed. These dis¬
continuities in the linkages between the two systems were described
and the reasons for their persistence, and in some cases, for their
exacerbation, were examined. The results of this examination re¬
vealed the causes for mass democracy’s insufficient reaction to the
problems of industrialism. Finally, some of the steps that will be
necessary to bring about a realignment of the democratic system
and mass society were spelled out. Unless the dislocations between
the two systems are removed, it will be impossible for a mass
democracy to successfully meet the challenges wrought by the
Industrial Revolution. The resources which are necessary to launch
this challenge already exist. This paper has demonstrated there is
information enough for action. All that is apparently lacking is
political will.
30 Paul Meadows, The Culture of Industrial Man (Lincoln, Nebr. : University of
Nebraska Press, 1950), p. 2.
THE FIFTH PAN AMERICAN CONFERENCE:
PROVING GROUND FOR WARREN G. HARDING’S
LATIN AMERICAN POLICY
Kenneth J. Grieb
President Warren G. Harding and Secretary of State Charles
Evans Hughes sought to promote friendship with Latin America,
and endeavored to reverse the long trend of interventions which
had characterized United States relations with that area of the
globe through the administrations of Theodore Roosevelt, William
Howard Taft and Woodrow Wilson. Harding and Hughes sought
stability in Latin America, but proposed to attain this by peaceful
means. The United States would continue to act as “big brother”
to the Latins, but would rely on diplomatic persuasion and calm
counseling instead of force. In this way, with the exercise of a
little patience, the desired objectives could be achieved without the
expense and ill feeling involved in armed intervention. Harding
and Hughes perceived that stability would facilitate American
financial penetration far more effectively than military occupation,
and hopefully would also terminate the frequent disputes that in¬
evitably entangled the United States. The Harding administration
avoided sending troops to Latin America, and by the time the Fifth
Pan American Conference assembled in Santiago, Chile in March,
1923, had completed arrangements for the withdrawal of the
troops from the Dominican Republic, recalled the detachment from
Cuba, and turned most police duties in Haiti over to a native
constabulary.1
The President went to considerable lengths to demonstrate his
cordial feelings toward Latin America. For example, in May, 1921,
he attended a Pan American Union reception honoring the Foreign
Minister of Venezuela, a gesture which impressed Latin diplomats.
Harding also opened a personal correspondence with President
Arturo Alessandri of Chile, after Alessandri, in congratulating
Harding on his inauguration, commented upon their common affil¬
iation with the Masonic order. Immediately recognizing the oppor-
xFor the Dominican Republic, see Kenneth J. Grieb, “Warren G. Harding and the
United States Withdrawal from the Dominican Republic,” Journal of Inter- American
Studies, XI, 3 (July, 1969), pp. 425-440. For the Cuban withdrawal see, Russell H.
Fitzglbbon, Cuba and the United States: 1900-1935, (Menasha, Wisconsin, 1935), p.
161, and New York Times, January 27, 1922. The instances of Cuba and Haiti, will
be treated more fully in a volume dealing with Harding’s Latin American Policy, which
the author is presently preparing.
69
70 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
tunity, Harding responded in a letter addressed to “My dear and
worthy brother Alessandri,” stressing their union in “fraternal
friendship.” Although the succeeding missives seldom touched on
important questions, they did promote understanding, especially
in view of Latin personali-smo.2 This correspondence may well have
influenced Chile’s decision to accept arbitration of the Tacna-
Arica dispute by Harding. To further demonstrate United States’
friendship, the President dispatched Hughes to the inaugural ses¬
sion of the Brazilian Centennial exposition of 1922 in Rio de
Janeiro. Harding and Hughes went beyond declamations, and en¬
deavored to demonstrate their sincerity through actions. Instead
of dispatching troops to Cuba under the Platt Amendment they
relied on General Enoch Crowder, using political rather than
military intervention.3 In addition to the troop withdrawals pre¬
viously cited, Harding devoted considerable effort to securing Sen¬
ate approval of the treaty returning the Isle of Pines to Cuba,
and also persuaded the Senate to give its advice and consent to
the accord compensating Colombia for the seizure of Panama.4
Both of these agreements had been previously rejected by the
Legislature, and their approval removed longstanding grievances
that had served as festering sores stimulating resentment in Latin
America.
Harding also wished the United States to assume an active peace¬
making role in settling disputes between Latin American states.
His offer to arbitrate the Tacna-Arica controversy is the best
known example, but was not his only effort.5 In 1922, during an
informal visit to the White House, President-elect Pedro Nel Ospina
of Colombia requested American aid in reopening diplomatic rela¬
tions with Panama. Harding enthusiastically wrote Hughes: “I
realize very well that this is none of our affair. . . . However, so
long as we play the role of big brother, I suppose we shall have
errands of this sort to perform.” The President suggested: “per¬
haps our Minister in Panama might make informal and wholly
discreet inquiry as to diplomatic representation in Colombia. Please
2 Dr. Leo S. Rowe to Harding-, May 12, 1921, Papers of Warren G. Harding-, Ohio
State Historical Society, Box 141 ; and Arturo Alessandri to Harding-, March 21, 1921,
and Harding- to Alessandri, May 2, 1921, Harding Papers, Box 693. Other letters from
the exchanges may also be found in the Harding Papers.
3 See Charles E. Chapman, A History of the Cuban Republic , (New York, 1927), pp.
413-449, and David A. Lockmiller, Enoch H. Crowder: Soldier , Lawyer , Statesman,
(Columbia, Missouri, 1955), pp. 230-246.
4 For the Colombian Treaty see E. Taylor Parks, Colombia and the United States:
1765-1934, (2 ed. New York, 1968), pp. 440-460. There are no satisfactory treatments
of the Isle of Pines Treaty available in secondary literature.
5 The United States was also active in several other disputes, such as the Panama-
Costa Rican war, and the attempt to form a Central American Federation. See for
example, Kenneth J. Grieb, “The United States and the Central American Federation,”
The Americas, XXIV, 2 (October, 1967), pp. 107-121. For the Panamian-Costa
Rican Conflict, see William D. McCain, The United States and the Republic of Pan¬
ama, (Durham, North Carolina, 1937), pp. 206-221.
1972] Grieb — Fifth Pan American Conference 71
note that I make no suggestion of formal proceedings. If we can
discreetly and helpfully broach the subject ... it would be a wholly
becoming thing to do.”6 Hughes proved reticent, but eventually
mediation was undertaken. This incident provided a clear illus¬
tration of Harding’s technique. As a practical politician, he re¬
alized the value of informal and friendly exchange, and was ac¬
customed to working tactfully and discreetly, without a formal
commitment. While the President was a neophyte in the field of
foreign policy, and was unacquainted with the diplomatic back¬
ground of the questions, his political training had provided him
with a thorough understanding of the methods involved. Harding
was well aware of his limitation, and constantly badgered the State
Department for position papers and background memoranda.7
When presented with the full particulars of a question, he had
no difficulty perceiving the proper course, and recognized sound
advice when it was provided. To facilitate their policies, Harding
and Hughes sought able diplomats to represent the United States
in Latin America. They carefully selected career envoys for the
most sensitive posts, and the Chief Executive found other positions
for political nominees, contrary to the image normally presented
in standard texts.8
The Fifth Pan American Conference took place, therefore, dur¬
ing a period of improving relations between Latin America and
the United States. Inevitably, such an assemblage of diplomats
from throughout the hemisphere would provide both a barometer
to measure the success of previous administration policy innova¬
tions, and a vehicle useful in advancing the administration’s prin¬
cipal objective in the region — improving the diplomatic atmosphere
by dissipating mistrust of the Colossus of the North. Harding and
Hughes shrewdly perceived that such a conference could effectively
contribute to a transformation of the milieu of inter-American
relations, rather than serving as a forum for political settlement.
Selection of the American representatives was crucial. Hughes
was “a little anxious about commitments that may be made at the
White House,” for the President was under “very great pressure”
6. Harding- to Hughes, May 5, 1922, Papers of Charles Evans Hug-hes, Manuscript
Division, Library of Congress, Box 24.
7 There are many items in the Harding papers and State Department papers indi¬
cating that the President requested such memoranda. See for example Hughes to
Harding, November 17, 1921, United States Department Papers, National Archives,
RG 59, 813.00/1145a, in which Hughes transmitted a memorandum to the President
“in accordance with your request.’’ Hereinafter State Department Papers will be cited
by slash number only.
8 The Harding image has been considerably distorted in standard historical texts,
and considerable revisionism is presently underway, using the information revealed
by his papers. The outstanding example to date which indicates the erroneousness of
the standard Harding image, and reveals prudence in the appointments to many
positions, is Robert K. Murray, The Harding Era: Warren G. Harding and his Admin¬
istration, (Minneapolis, 1969).
72 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
from Senators “anxious to have constituents appointed,”9 but the
delegation was carefully chosen. The President named Henry P.
Fletcher as chairman. Although Fletcher was then serving as am¬
bassador to Belgium, he possessed extensive experience in and
was widely known in Latin America, and had formerly served in
the host country. A close friend of the President, Fletcher had
served as Undersecretary of State throughout the first year of
the administration, during which he was “a full member of the
golf cabinet.”10 Fletcher’s appointment was acclaimed by the Amer¬
ican and Latin American press, and the President of Paraguay
told the American minister in Asuncion that: “He was particu¬
larly pleased with the selection of Mr. Fletcher, who from long
experience with Latin temperament, will be able to accomplish
more than any other person.”11 The selection of Dr. Leo S. Rowe,
Director of the Pan American Union, provided the American dele¬
gation with another individual knowledgeable about and well known
in Latin America. George E. Vincent, head of the Rockefeller
Foundation, and Frank C. Partridge, a former minister to Ven¬
ezuela, were also chosen. Contrary to the usual image of Harding,
the Chief Executive side-stepped pressures from legislators in
behalf of constituents, by selecting two Senators, Frank B. Kel¬
logg of Minnesota, a Republican, and Atlee Pomerene of Ohio, a
Democrat. Former Senator Saulsbury of Delaware and Washing¬
ton attorney William E. Fowler completed the list.12 By selecting
Senators, Harding assured support for the ratification of any
agreements. The delegation thus represented an astute compro¬
mise between ability and political expediency. Harding correctly
discerned that Fletcher, by virtue of his experience and friend¬
ship with both the Chief Executive and the Secretary, would dom¬
inate the delegation.
There was considerable speculation that the Secretary of State
would attend the conference and Harding strongly urged him to
make an appearance. The President viewed this as a means of
dramatizing American friendship for Latin America, and when
Hughes proved reluctant, Harding placed considerable pressure
upon him. As early as November 28, 1922, he wrote the Secretary:
“If the circumstances are such that you can arrange to go and
9 Dr. Leo S. Rowe to Henry P. Fletcher, November 10, 1922, Box 9, Papers of Henry
P. Fletcher, Manuscript Division, Library of Congress ; and Rowe to Fletcher, Janu¬
ary 15, 1923, Fletcher Papers, Box 10.
10Hug’hes to Fletcher, January 5, 1923, State Department Papers, 710 E. 002/2a;
Hughes to Harding, January 6, 1923, and Harding to Hughes, January 6, 1923, 710
E. 002/3 ; Fletcher to Harding, January 10, 1923, Fletcher Papers, Box 10, Fletcher
to Hughes, January 10, 1923, Hughes Papers, Box 21, and New York Time s, March 4,
1923.
11 New York Times, March 4, 1923 ; and William J. O’Toole (United States Minister
in Paraguay) to Hughes, February 12, 1923, reporting the remarks of President Ayala,
710. E. 002/85.
12 Hughes to Fletcher, January 31, 1923, 710 E. 002/161.
1972]
Grieb — Fifth Pan American Conference
78
open the conference I think it would be an exceedingly fine thing
to do. ... I think the benefits accruing from your visit to Brazil
might be duplicated by a visit to Chile. I should be happy to pro¬
mote such visits on your part as will tend to enhance our relations
with South American states/’13 The Secretary however, had no
desire to attend the conference, since he regarded Pan American
Conclaves as mere “friendship festivals.”14 While assuring the
Chief Executive that he would “do all in my power to aid our Latin
American relations,” he continued: “I confess . . . that I have
no love whatever for speechmaking trips.” Harding replied that
he would discuss the matter with Hughes personally, indicating
that he still favored an appearance by the Secretary.15 Obviously,
Harding was far more aware of the value of personal diplomacy
than was Hughes, and in this sense the President was ahead of
his time.
Hughes yielded to Harding’s admonitions, and accepted a Chil¬
ean invitation to attend the inaugural session of the conference,
but reversed himself at the last moment, citing the “press of work”
in Washington, and ignoring renewed appeals from Harding.16
The Secretary’s action appears particularly regrettable in view of
his success at the Sixth Pan American Conference in 1928, and
in this context Harding was certainly perceptive in urging Hughes
to attend the Santiago conclave. Yet, in retrospect, perhaps the
absence of Hughes benefited the United States, for the appoint¬
ment of Fletcher assured American success at Santiago. With his
knowledge of Latin temperament, Fletcher was more subtle and
less committed to rigid legalism than the Secretary, and conse¬
quently was able to secure American ends more tactfully, remain¬
ing pliant, stressing cooperation, and exuding friendship. This
was the very embodiment of the tactics Harding advocated and
hoped to make the basis of hemispheric understanding.
Problems began before the conference convened, as Mexico de¬
clined the invitation, citing the fact that its dispute with the United
States denied it representation on the governing board of the
Pan American Union, which prepared the conference agenda. That
body’s membership was limited to the representatives of the re¬
spective states in Washington, and diplomatic relations between
the United States and Mexico remained severed. While Chile in¬
vited Mexico to the conference, the Mexicans considered it beneath
13 Harding- to Hughes, November 28, 1922, Harding Papers, Box 361.
u Charles Evans Hughes, The Pathway of Peace, (New York, 1925), pp. 137 and
160, and Murlo J. Pusey, Charles Evans Hughess, (New York, 1951), p. 55.
15 Hughes to Harding, November 28, 1922, Hughes to Harding, December 1, 1922, and
Harding to Hughes, December 1, 1922. Harding Papers, Box 361.
16 Hughes to Fletcher, January 5, 1923, 710 E. 002/2a; New York Times, January
14, 1921, January 31, 1921, February 27, 1923, March 4, 1923, and William Miller
Collier (United States Ambassador in Chile) to Hughes, March 3, 1923, 710 E. 022/66.
74 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
their dignity to attend after being unable to contribute to the
agenda, and charged that the Yankees were deliberately seeking
to exclude them. When Chile and Brazil informed Washington
that they desired Mexican participation, Hughes replied that al¬
though the United States did not object to the presence of Mexico
at the Santiago conference, and despite continuing negotiations
with Mexico, “It would be quite impossible for this Government
to enter into treaties with a government which has not been rec¬
ognized/’17 Hughes’ inflexible stand forced the Latins to choose
between Mexico and the United States, but this caused little diffi¬
culty in view of Mexican intransigence.
The governments of Peru and Bolivia also abstained from par¬
ticipating in the conference. These nations feared that the conclave
might attempt to compel them to accept some settlement of the
Tacna-Arica dispute, and also objected to a meeeting in the capital
of the nation which they felt had committed a transgression.18
When the conference convened on March 25, 1923, Ambassador
Fletcher immediately became the focal point. Chilean crowds lin¬
ing the streets cheered him as he passed through the city, and
delegates applauded when he entered the inaugural session. Sen¬
ator Pomerene was awed by the “most flattering ovation” Fletcher
received, and wrote Hughes: “You have named the right man for
chairman of our delegation.” Chilean President Arturo Alessandri,
who opened the conclave, presented the original manuscript of his
keynote address to Fletcher, a gesture which clearly demonstrated
both the ambassador’s popularity with Latin diplomats19 and the
success of Harding’s policies.
The chairman of the American delegation deftly worked to secure
maximum benefits for the United States, without ruffling Latin
feelings. Fletcher needed all his diplomatic skill to blunt the vest¬
iges of suspicion of the United States, since the conference agenda
was studded with political topics, which appeared in greater pro¬
fusion than at previous Pan American conferences. Debate on any
one of these controversial questions could become anti-American,
and the chairman of the American delegation was compelled to
oppose some of the aspirations of the Latin diplomats which clashed
with American policy. Fletcher had to simultaneously promote
good will and defend his country’s interests.
Realizing that the Latins would raise the question which had
kept Mexico from the conference, Dr. Leo S. Rowe Director of
the Pan American Union, offered a proposal to broaden the pow-
17 New York Times , January 13, 1923, and Hughes to Fletcher, March 5, 1923, Hughes
Papers, Box 21.
18 New York Times, March 25, 1923.
19 Atlee Pomerene to Hughes, March 28, 1923, 710 E. 002/89 ; and Fletcher to Arturo
Alessandri, April 5, 1923, Fletcher Papers, Box 10.
1972] Grieb— -Fifth Pan American Conference 75
ers of the Union, and permit it to consider political questions. Costa
Rica proposed a board composed of delegates accredited directly
to the Union, to enable a government denied United States recog¬
nition to retain its representation. Most of the Latin American
nations supported this plan. The resulting compromise sanctioned
appointment of a special representative by any government that
so elected, accepted Rowe's proposals, and also provided for an
elected board chairman, terminating the automatic appointment
of the American Secretary of State; though the practice of elect¬
ing the Secretary continued.20
Given Latin American membership in the League of' Nations,
questions relating to the world body were implicit throughout the
conference. Agustin Edwards, a Chilean diplomat then serving as
President of the League of Nations, was elected President of the
Pan American Conference. Edwards saw no conflict in this dual
post, contending that the League had promoted Pan American
solidarity by bringing the Latin nations together in a voting block
at the world body.21 Yet such parallels raised suspicions in the
United States, due to the passionate feelings aroused by the dis¬
pute over League membership. The New York Times commented
satirically that the discussion of “League plans" at Santiago “will
compel the State Department to admit that the League of Nations
exists, or else walk out of the conference. If our delegates walk
out," the Times continued, “Pan American harmony will have
been destroyed; if they admit that there is a League of Nations,
Henry Cabot Lodge will be stultified, and Heaven knows what harm
may happen to the Republic."22
In accordance with a suggestion by Uruguayan President Dr.
Baltasar Brum some three years earlier, several delegations advo¬
cated the formation of an American League of Nations.23 The pro¬
posed Charter contained a provision continentalizing the Monroe
Doctrine that caused considerable debate. Dr. Brum advocated
Latin support for the Doctrine, in order to transform it into a
multi-lateral instrument, which would be enforceable jointly, but
the United States steadfastly insisted that the Monroe Doctrine
must remain a unilateral policy.24 Although the proposal died in
the face of determined opposition by the United States, the fact
that the American delegation had not prevented discussion of the
idea was significant. A Costa Rican suggestion for an all Ameri¬
can Court of Justice, raised another question the United States
20 New York Times , April 5, April 10, April 17, and April 19, 19 23.
21 “The Fifth Pan-American Conference,” Current History, XVII (1923), pp. 184-
188, and Augustin Edwards, “Latin America and League of Nations,” Current His¬
tory, XVII (1923), pp. 181-183.
22 New York Times , July 11, 1922.
23 New York Times, July 16, 1922.
24 New York Times , March 24, and May 2, 1923.
76 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
preferred to avoid, but the Latins themselves were by no means
agreed on the establishment of a court, and the proposal was
deferred.25
Arms limitation provided an additional perplexing issue. The
futility of seeking agreement on limitation of ground forces was
readily apparent, and discussion was restricted to naval limitation,
following the example of the Washington Disarmament Confer¬
ence. Naval limitation, in turn, hinged on agreement between
Argentina, Brazil, and Chile, which possessed the largest fleets.
But the A.B.C. countries deadlocked, and the only result was an
agreement to continued talks between them.26
The outstanding development of the Fifth Pan American Con¬
ference that resulted in a concrete agreement was the adoption
of the Treaty to Avoid or Prevent Conflicts between the American
States, also known as the Gondra Treaty, after its sponsor, Dr.
Manual Gondra of Paraguay. The signatories pledged to submit
all outstanding disputes to an International Commission of Inquiry,
which would tender a report after a six month study. While the
decision would not be binding, the respective states would be re¬
quired to refrain from initiating war preparations during the
investigation and the succeeding six months.27 Thus the treaty
functioned in the same manner as the Bryan “cooling off” treaties,
merely multilateralizing the process. The plan assumed that post¬
poning hostilities would prevent their outbreak by allowing time
for negotiations. Presumably the other American states would
apply pressure for a settlement during the interim, although the
treaty did not commit them to do so.
The delegates also codified a number of technical matters. A
convention providing for publication of passport regulations and
standardization of forms to facilitate intercourse among the sev¬
eral states was approved. Another agreement established standard
nomenclature for commercial shipping of merchandise, to stimu¬
late trade. The conference also approved a convention providing
reciprocal protection for copyrights, trademarks, and commercial
names.28
Fletcher’s ability to uphold American policy in a subtle manner
drew praise when the conference adjourned May 3, 1923. Dr. Rowe
hailed “the masterly way in which you have handled the work of
the Delegation,” while Senator Pomerene wrote Harding: “You
26 New York Times, April 6, and April 29, 1923.
28 Washington P'ost, April 7, 1923, and New York Times, April 12, April 18, and April
19, 1923. See also Current History, XVIII (1923), pp. 924-925.
27 Washington Post, April 16, 1923 ; New York Times, May 12, 1923 ; and Pan Amer¬
ican Union, Tratados y convenciones suscritos en la Quinta Conferencia Internacional
Americana, Serie sobre derecho y tratados, No. 19, (Washington, 1949).
28 New York Times, April 3, 1923, and Pan American Union, Tratados convenciones
suscr'itos en la Quinta Conferencia, passim.
1972]
Grieb — Fifth Pan American Conference
77
made a ten strike in naming Ambassador Fletcher as chairman
of this Delegation. He is very popular here among the Chileans and
all South Americans.”29
From the viewpoint of the Harding administration, the confer¬
ence was a resounding success. A change in atmosphere was the
key objective, and the United States delegation attempted to exude
good will and allow the Latins to feel free to express their views.
Despite the fact that Fletcher became the focus of attention at the
conclave, he nevertheless refrained from commenting on the ques¬
tions under consideration until the Latin delegates had stated their
positions. This change in the role of the United States delegation
constituted the pivotal factor in setting the mood of the confer¬
ence. As a result, in the words of one observer, for “the first time
in the history of these conferences . . . the Latin Americans felt
they could freely speak their minds.” So successful was
the American delegation at cultivating good will and listening
to Latin views, that domestic press criticism of the conclave was
answered from Latin America. Taking exception to a New York
Evening Post editorial criticizing the defense of the Monroe Doc¬
trine as “imperialism,” El Mercurio of Santiago replied: “The
North American delegation observed such a deferential and re¬
spectful attitude toward all the other republics that without a
doubt they succeeded in convincing all the delegations that the
United States does not follow an imperialistic policy, nor seek
to impose its policies, but seeks harmony of interests, and cor¬
diality founded on mutual respect and equality of treatment.”31
Such a comment would scarcely have been possible at earlier Pan
American conclaves, and it reflected the success of the Harding
policy.
Fletcher reported to President Harding and Secretary Hughes
that: “The frank, free, full discussions had made the conference
a success as far as the establishment of better and more friendly
relations among the American nations was concerned.”32 The Amer¬
icans did not go to Santiago seeking any spectacular agreements.
The Harding administration desired only to dissipate the residue
of ill feeling that remained from previous American actions, and
create a new spirit to facilitate mutual understanding. Withdraw¬
ing objections to the discussion of political issues was one way to
show the new policy of friendship, but this did not mean sanc¬
tioning the policies. The Americans desired only an airing of
views. That they succeeded in obtaining this .without an excessive
^Alessandri to Fletcher, May 7, 1923, Fletcher Papers, Box 10; Rowe to Fletcher,
May 3, 1923, Fletcher Papers, Box 10 ; and Pomerene to Harding, May 4, 1923, Fletcher
Papers, Box 10.
30 Current History , XVII (1923), p. 923.
31 El Mecurio (Santiago), August 22, 19 23.
32 New York Times, June 1, 1923.
78 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
amount of anti-American tirades, was certainly an accomplish¬
ment.
If the results of the conference appear somewhat limited at first
glance, this is because the item of greatest significance, the change
in atmosphere, could not be recorded in an agreement. The New
York Times noted editorially that the results would disappoint
the idealists but cheer the realists.33 Discussion of numerous polit¬
ical questions represented a significant step along the road of Pan
Americanism, and although the resulting disputes could not be
resolved, the airing of the controversial issues was an important
transition. The accomplishments of the conference were more ex¬
tensive than the treaties indicate.
The Harding administration thus secured its ends at the Fifth
Pan American Conference, and this accomplishment indicated the
skill with which it pursued its goals. Harding shrewdly focused
upon realistic objectives, and prudently selected a delegation chair¬
man whose skill matched his objectives. The combination of careful
advance preparation and able representatives brought success for
Harding’s practical common sense approach to diplomacy. Thus
the Fifth Pan American Conference both contributed to his objec¬
tives and demonstrated the success of his policies.
33 New York Times , May 12, 1923, and Samuel Guy Inman, “Pan-American Unity in
the Making-,” Current History, XVII (1923), p. 919.
FOOD HABITS OF THE COHO SALMON,
ONCORHYNCHUS KISUTCH, IN LAKE MICHIGAN
Margaret A. Harney and Carroll R. Nor den
Abstract
The stomach contents of thirty-six coho salmon, Oncorhynchus
kisutch, taken from southern Lake Michigan were examined. The
contents were identified and the numerical and volumetric meth¬
ods used to analyze the data. The results indicated that the coho
salmon in Lake Michigan is primarily piscivorous. Fish comprised
96.9 percent by volume of the coho’s diet and the alewife accounted
for 62 percent of the total stomach contents. Other items included
in the coho’s diet were smelt, stickleback and insects.
Introduction
The coho salmon, Oncorhynchus kisutch , were stocked in Lake
Michigan in the spring of 1966 by the Michigan Department of
Natural Resources (Borgeson, 1970). They were stocked in the
lake to serve as a predator on the alewife and to convert their fish
flesh into a valuable sport fishery. Since the 1950’s, the alewife
population in Lake Michigan has increased dramatically. The in¬
crease in the alewife population followed the decline and virtual
extinction of the lake trout as a result of changing lake conditions
and to the marine lamprey. As often happens, when a top predator
is lost from an ecosystem, the organism that had served as its
prey increased unchecked. It was hoped that coho stocked in the
lake would replace the lake trout as one of the top predators and
would help to control the alewife population. The objectives of
this study were to determine the food habits of the coho in Lake
Michigan and to determine whether the coho were in fact preying
upon the alewife.
Materials and Methods
The stomach contents of thirty-six coho salmon were examined.
The coho were caught by gill net in Illinois waters of Lake Michi¬
gan between March 28 and May 1, 1968. Two main collections
were made, one on April 11, 1968 and the other on April 22, 1968.
The stomachs were removed by severing the gullet and the intes¬
tine behind the plyoric caeca and then placed in 10% formalin. A
binocular microscope was used to identify the stomach contents.
79
80 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Contents rendered unrecognizable either through mastication or
digestion were classed as unidentifiable. The quantitative analysis
was done by water displacement in which the volume of each item
of food was expressed as a percentage of total volume of stomach
contents. The frequency of occurrence was recorded as the number
of samples in which each food item occurred and as a percentage
of the total number of specimens which contained food. The rela¬
tive importance of each food item was also determined using the
numerical method. The results were reported as a percentage of
the total number of organisms consumed (Lagler, 1956).
Results
The coho salmon in Lake Michigan was mainly piscivorous and
fish comprised 96.86 percent by volume of their diet (Table 1).
Alewife
Smelt
Stickleback
Skeleton
Insecta
Unidentified
Figure 1. Percent of total, by volume which each food item contributed to the
diet of the coho salmon taken from southern Lake Michigan during the spring
of 1968.
1972] Harney and Nor den — Food Habits of Coho Salmon 81
Alewives comprised 62 percent of the total volume followed by
smelt with 17 percent (Fig. 1). Stickleback and other fishes made
up an additional 18 percent whereas insects were of minor impor¬
tance, less than one percent (Fig. 1). Alewives were found in 50
percent of the stomachs which contained food (Fig. 2) and ac¬
counted for the largest amount of food by volume (Table 1). They
were not, however, the most numerous organism in the stomach
but this was due to the fact that two coho stomachs contained many
small smelt. In addition to alewife and smelt, the cohos also con¬
sumed several sticklebacks. These three species were the only spe¬
cies of identifiable fish found in the cohos’ stomachs.
Discussion
It was found that 14 stomachs or 38.9 percent were empty. This
was considered a rather high proportion of empty stomachs.
LeBrasseur (1966) caught four species of Pacific salmon in gill
nets set for 10-hour periods (overnight). He analyzed the stomach
Alewife
Smelt
Stickleback
Skeletons
Insecta
Unidentified
“555 -
Frequency
- 43”
Occurrence
“50 - W
( Percent )
Figure 2. Frequency of occurrence of each food item in the diet of the coho
salmon taken from southern Lake Michigan during the spring of 1968.
Table 1. Food of Coho Salmon Taken From Southern Lake Michigan During the Spring of 1968.
82
Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
00
s
00
xd
o
00 —
00 —
00 —
xO
C
£
<S
Stomachs with
1972] Harney and Norden — Food Habits of Coho Salmon 83
contents of the four species and found that the coho had the high¬
est incidence of empty stomachs, up to 43 percent whereas the
chum salmon had the least, only three percent. LaBrasseur (1966)
postulated that chum fed early in the evening as the nets were
being set whereas pink, coho and sockeye tended to feed in early
morning about the time the nets were being retrieved.
In Lake Michigan the principle food of the coho was fish, (96.86
percent) whereas in other studies crustaceans and insects formed
a much greater share of the coho’s diet. Stanley (1937), Hasler
(1938) and Hasler and Farner (1942) reported that the main
foods of coho in Crater Lake, Oregon was daphnia, midges and
caddis flies. The food of the coho salmon in certain Oregon streams
consisted mainly of Diptera and Ephernormoptera (Rees, 1959).
The principle food of the coho salmon in Cultus Lake, British
Columbia (Ricker, 1946, 1952) was midges, particularly in the
spring.
Most of the food studies of coho have been done on fish taken
in open waters off the coast of Oregon and British Columbia. The
diet of coho in the northeast Pacific consisted of squid, fish (includ¬
ing smelt, anchovies, herring, lantern fish, hake, whiting, other
coho, black cod and rockfish) crustaceans such as amphipods, eu-
phasids, and crab larvae, and jellyfish, (Prichard and Tester, 1943;
Fraser, 1946; Oregon Fish Commission, 1949; Van Hvning, 1951;
Prakash and Milne, 1958; Roos, 1960; Prakash, 1962; Reimers,
1964; LeBrasseur, 1966; Grinolds and Gilt, 1968; Manzer, 1968,
1969; Ueno, 1969).
The relative importance of each type of food depended on where
the fish was caught and what was available in the environment
at the time of capture. A number of the authors commented on the
wide variety of food eaten by coho and expressed the idea that the
coho is an opportunistic feeder, feeding upon what is available
at the time (Prakash, 1962; Reimers, 1964; LaBrasseur, 1965).
LeBrasseur (1965) compared the diets of pink, sockeye, chum,
coho and steelhead trout taken from different areas of the north¬
eastern Pacific Ocean. He found more correlation of stomach con¬
tents among all species from one area than among members of
one species taken from the four fishing areas. He also noted that
a small change in the availability of some organisms could pro¬
duce significant changes in the stomach contents.
In the present investigation, alewives made up a major portion
of the stomach contents of the coho probably because they are
opportunistic feeders and the alewife is the most abundant fish
in Lake Michigan (Smith, 1968). Crustacea were an insignificant
part of the diet of Lake Michigan coho as compared to Pacific
84 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
coho because the open lake lacks the numbers of Crustacea that are
present in the Pacific.
Acknowledgements
The authors wish to acknowledge Mr. Bruce Muench, Area Fish¬
ery Biologist of Illinois, who collected the coho salmon.
This work was partially supported by the University of Wiscon¬
sin Sea Grant Program.
Bibliography
Borgeson, D. P. (editor). 1970. Coho salmon status report. 1967-1968. Mich.
Dept. Nat. Res., Fish Mgt. Rept. No. 3: 1-31.
Fraser, M. C. 1946. Food of fishes. Trans. Roy. Soc. Can. Third Ser. 40 (Sect.
5) : 33-39.
Grinolds, B. and D. Gill. 1968. Feeding behavior of three oceanic fish, Oncor-
hynchus kisutch, Trachurus symetricus and Anaplopoma fimbria from
the northeast Pacific. J. Fish. Res. Bd. Can. 25 (4) : 825-827.
Hasler, D. 1938. Fish biology and limnology of Crater Lake, Oregon. J. of
Wildlife Mgt. 2 (3) : 94-103.
Hasler, D., and D. S. Farner. 1942. Fisheries investigations in Crater Lake,
Oregon. J. of Wildlife Mgt. 6 (4) : 319-327.
Lagler, K. F. 1966. Freshwater Fishery Biology. William C. Brown Co. Du¬
buque, Iowa. 421 p.
LeBrasseur, R. J. 1966. Stomach contents of salmon and steelhead trout in
the northeastern Pacific Ocean. J. Fish. Res. Bd. Can. 23 (1) : 85-100.
Manzer, J. I. 1968. Food of Pacific salmon and steelhead trout in the north¬
eastern Pacific Ocean. J. Fish. Res. Bd. Can. 25 (5) : 1085-1089.
- . 1969. Stomach contents of juvenile Pacific salmon in Chatham Sound
and adjacent waters. J. Fish. Res. Bd. Can. 26 (8) : 2219-2223.
Oregon Fish Commission. 1949. Crab larvae as a food for silver salmon at
sea. Oregon Fish. Comm. Res. Bd. Can. 2 (1) : 1-17.
Prakash, A. 1962. Seasonal changes in feeding of coho and chinook salmon
in southern British Columbia water. J. Fish. Res. Bd. Can. 19 (5) : 851-
866.
Prakash, A. and D. J. Milne. 1958. Food as a factor affecting the growth of
coho salmon off the east and west coast of Vancouver Island, British Co¬
lumbia. Fis. Res. Bd. Can. Pro. Rep. Pacific Coast Sta. No. 112: 7-9.
Pritchard, A. L. and A. L. Tester. 1943. Notes on the food of coho salmon in
British Columbia. Fish. Res. Bd. Can. Prog. Repts. Pacific Coast Sta.
No. 55: 10-11.
Rees, Wm. 1959. Effects of dredging on young silver salmon and bottom fauna.
Wash. Dept. Fish., Fish. Res. Pap. 2 (2) : 52-65.
Reimers, P. E. 1964. A modified method of analyzing stomach contents with
notes on the food habits of coho salmon in coastal waters of Oregon and
southern Washington. Oregon Fish. Comm. Res. Briefs. 10 (1) : 46-56.
Ricker, Wm. E. 1946. The food supply of sockeye salmon in Cultus Lake,
British Columbia. J. Fish. Res. Bd. Can. 3 (5) : 450-468.
- . 1952. The benthos of Cultus Lake. J. Fish. Res. Bd. Can. 9 (4) :
204-212.
1972] Harney and Norden—Food Habits of Coho Salmon
85
Roos, F. 1960. Predation of young coho salmon on sockeye salmon fry at
Chignik, Alaska. Trans. Am. Fish. 89 (4) : 377-378.
Smith, S. H. 1968. The alewife. Limnos. 1 (2) : 9 p.
Stanley, B. J. 1937. Study of the contents of fish stomachs from Crater Lake,
Oregon. Manuscript.
Ueno, M. 1969. Food and feeding behavior of Pacific salmon. The stratification
of food organisms in the stomach. Bull. Jap. Soc. Sci., Fisheries. 34 (4) :
315-318.
Van Hyning, R. and J. Van Hyning. 1951. Food of chinook and silver salmon
taken off the Oregon Coast. Oregon Fish. Comm. Res. Briefs. 3 (2) : 32-40
LIMNOLOGY OF SOME MADISON LAKES:
ANNUAL CYCLES
Kenton M. Stewart and Arthur D. Hasler
Abstract
Detailed studies of Lakes Mendota, Monona, and Waubesa at
Madison, Wisconsin, provide a comparison of their annual cycles
and the influence of climatic variations on their thermal regimes.
Lake Waubesa integrates the influence of wind velocity, air tem¬
perature, and solar radiation most rapidly whereas Lake Mendota
responds most slowly. Although Lake Monona is still dominated
generally by climatic influences, the cultural influences of thermal
discharges now causes the ice to depart in spring at a significantly
earlier date than the other two lakes. The clarity of the Madison
lakes generally decreases from Mendota to Monona to Waubesa.
Although all three lakes are basically eutrophic dimictic lakes
with anoxic conditions developing in the lower waters during sum¬
mer stratification, Lake Waubesa may have aperiodic overturns
during summer.
Comparisons of temperature, oxygen, Secchi disk, photometer
readings, and ice thickness in Lake Mendota show relatively little
change from the early studies of Birge and Juday several decades
ago.
I. Introduction and Background
The fascinating thing about lakes is that they provide their
own variety. The task of the limnologist is to measure and interpret
this variation whether it concerns physical, chemical, or biological
phenomena.
This paper is an attempt to compare the annual cycles of se¬
lected variables, measured during the same time period, in Lakes
Mendota, Monona, and Waubesa at Madison, Wisconsin.
In temperate dimictic lakes, the greatest range of variation for
any variable occurs sometimes during the annual cycle. There¬
fore, an examination of selected variables over the course of at
least one annual cycle, provides some tentative limits by which
to judge past, present, and future events in the system. As the
number of years of investigation increases, the limits and expected
changes become defined more clearly and valuable base line data
are established. In addition to the fluctuations of the annual cycle,
87
88 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
there are seasonal, daily, and some second-by-second changes that
challenge the interpretation of the investigator. A lake may serve
as a multi-ring circus in which it is impossible to observe everything
at one time, but if one attends frequently a more thorough ap¬
preciation and understanding of the whole system can be gained.
Early illustrations of annual cycles in Lake Mendota were pro¬
vided by Birge and Juday (1911), and some unpublished data of
Birge (Neess and Bunge, 1956, 1957). In the latter references,
several years data on lake temperatures were presented. Even these
data, though extensive, were accumulated mostly during the ice-free
season of the year. The general lack of winter data, when com¬
pared to those of the summer season, is common for limnological
investigations throughout the world and provides a summer bias
in our interpretation of lake events.
In addition to their studies of annual cycles of temperature and
oxygen, the contributions of Birge and Juday (1929, 1931, and
1938), concerning solar radiation and transparency in water, were
significant.
It is surprising, in light of Birge 's interest in comparative lake
studies, that investigations of other Madison lakes besides Mendota
were so slighted. How much better would we have been able to
understand lakes generally and changes in the Madison lakes
specifically if Birge had concentrated his efforts in comparative
studies there?
II. Methods and Procedures
The thermal measurements were made with a Whitney ther¬
mometer at four selected stations (Fig. 1) in the three Madison
lakes considered in this paper. The accuracy of the thermometer
was maintained at ± 0.1 °C and rechecked frequently with precision
mercury thermometers.
Measurements of transparency and light were made by several
different instruments. A Secchi disk (20 cm diameter, all white)
was used to gather general transparency information. The G. M.
Photometer with a Weston photronic cell (Model N. 8564R) and
cosine filter were used to measure the 1% level and extinction
coefficients. Measurements for microstratification were taken with
a Whitney transmissometer or turbidimeter (one meter path length,
a modified version of the earlier instrument by Whitney, 1938).
Physical and chemical data were secured during the summer
by use of a boat and during the winter by hauling the equipment
over the ice on a toboggan. During dangerous periods of winter,
e.g., at the time of initial ice formation or in the last several days
of rotting ice, a small aluminum boat with a motor was dragged
over the ice to the sampling site(s). Whenever the ice broke K. M.
1972] Stewart and Hasler — Limnology of Madison Lakes
89
Stewart got into the boat and used the powered boat as a miniature
icebreaker until the ice was firm enough to hold or until the
sampling site or shore was reached. Under certain conditions of
smooth but thin sheet ice, it is possible to sit in a boat and push
oneself along with a spiked pole. However it is advisable to have
a dependable motor along in case the boat breaks through the
ice— as it usually does. These practical expediencies allowed acqui-
90 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
sition of data at a most interesting* and little studied time in
the annual cycle of a lake. As a further bit of practical informa¬
tion, the period of risk (depending on meteorological conditions)
is usually shorter during early ice formation than when the ice
is wasting.
The water samples were acquired with a Kemmerer sampler
from preselected depth intervals in each of the lakes investigated.
The azide modification of the Winkler method was used for all
oxygen determinations (Standard Methods, 1960). Reagents were
added in the boat and the samples were titrated immediately upon
return to the laboratory.
Information concerning solar radiation, air temperature and
wind velocity (means of 24 measurements daily for 1960-1963 and
eight measurements daily for 1966), was obtained from the
published data of the U.S. Weather Bureau Station at Truax
Field in Madison, Wisconsin (Annual Climatological Data, 1959-
1966). Radiation data of the Weather Bureau were compared to
and augmented occasionally by the data of the Solar Energy
Laboratory at the University of Wisconsin when the radiometer
at the Weather Bureau malfunctioned.
III. Results and Discussion of Harmonic Time Series
A. Annual Cycle: Weather Data
In most of the graphs in this paper concerning annual cycles,
the day-to-day variation of air temperature, wind velocity, and
solar radiation has been smoothed by plotting only the moving
ten-day averages.
However, one year (1962) has been selected to illustrate the
enormous detail apparent when comparing daily readings of wind
velocity and direction, air temperature, and incoming radiation
over an annual cycle. The daily means of 24 measurements of wind
velocity and direction have been plotted (Fig. 2). Solid lines above
and below the zero value indicate winds from the south to west,
and north to east respectively, with the maximum velocity for
that date indicated by a solid dash. Winds from the east-southeast
to south-southwest and west-northwest to north-northwest are
indicated by dotted lines above and below the line respectively,
with separate dots for the fastest velocity.
The 24-hour daily means of weather data (Fig. 2), owing to
their great variability and without further smoothing, would tend
to mask the plots of lake data. Therefore computer calculations
of moving ten-day means for winds, air temperature, and solar
radiation have been plotted for easier comparison of climatological
variables and lake trends over an annual cycle.
1972] Stewart and Hasler — Limnology of Madison Lakes
91
Figure 2. Data from the U.S. Weather Bureau at Truax Field in Madison,
Wisconsin. Values of wind velocity and direction are the resultant (this figure
only) means of 24 hr with maximum velocities for each day indicated by dots.
The 24 hr mean of air temperature and 24 hr total of solar radiation both
demonstrate great variability.
92 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
B. Annual Cycles: Temperature and Oxygen, 19 60-1 9 6 If and
1966
1. Lake Mendota, Temperature and Oxygen
(a) 1960
The initial program for gathering data on temperatures began
in 1960 (Fig. 3). Less information was collected in that year
than in the succeeding ones. No measurements were made of
dissolved oxygen in 1960 and all data were gathered at station 1
(Fig. 1). The 1960 climatological data from the U.S. Weather
Bureau Station at Truax Field (^ 3.2 km from Lake Mendota) in
Madison, Wisconsin, were plotted as moving ten-day averages on
the tenth day.
The temperature of the water at 18 meters was about one to two
degrees cooler in 1960 than the temperature at the same depth in
1961-1963 and 1966. This difference was not associated simply
with a change in sampling position from station 1 (1960) to sta¬
tion 2 (1961-1963 and 1966). Both stations were compared occa¬
sionally in 1960 and only slight differences were noted. Further¬
more, the data of Birge and Juday (1911) show temperatures at
18 meters from the area of station 1 in 1906 and 1907 similar to
those of the latter years of this study. Therefore, specific episodes
of wind velocities, air temperatures, and solar radiation must have
combined during the critical vernal circulation to control the warm¬
ing of the hypolimnetic waters. This supports some of the conclu¬
sions in Birge’s (1916) paper on the work of the wind in warming
the lake.
The response of the lake to the ten-day mean of air tempera¬
tures is illustrated best by temperatures in the first six meters
(Fig. 3). The lag of water temperature, compared to that of the
air, was prominent in all years and on all lakes to a varying
degree.
(b) 1961
Temperature data were collected from Lake Mendota in 1961
(Fig. 4) more frequently than they were in 1960. Measurements
were made of dissolved oxygen also. Data on both variables were
relatively sparse during the first few months of 1961. Consequently
the data taken during the spring missed completely the dramatic
change in stratification normally associated with the disappear¬
ance of the ice.
More heat was distributed to the lower depths during spring
circulation so that the lower waters began the summer season
warmer than they were in 1960.
The depth of the lake where these data were collected was
about 18.5 meters (Fig. 1, station 2) compared to about 21.5 meters
1972] Stewart and Easier — Limnology of Madison Lakes
93
fl/li
94 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 4. Lake Mendota temperature and oxygen 1961. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day means plotted
on the tenth day. Mean wind velocities are generally lower in summer.
1972] Stewart and Hasler — Limnology of Madison Lakes 95
for station 1, 1960. The probe of the Whitney thermometer was
dropped into the mud for measuring the mud temperature. The
depth of penetration by the perforated head of the thermistor
probe varied with sediment-composition and rate of lowering. The
penetration at these depths was approximately 20 to 30 centimeters.
In soft bottoms, the mud-water interface felt poorly defined from
a cable.
On 15 December 1961 the entire surface of Lake Mendota froze.
The temperature of the water on the preceding day was 1.4 °C.
No measurement was made on the day of freezing. Note the im¬
mediate stratification of temperature following freezing. The mud
and lower waters are warmer than the surface waters owing to
the density anomaly.
There is a rapid stratification of oxygen after the spring “over¬
turn”. “Occasional blooms” of algae may cause super saturation
as is indicated in early June. The dominant causative organism
of that bloom was the blue-green alga, Aphanizomenon. The dis¬
solved oxygen of the hypolimnion is utilized rapidly through de¬
composition of organic and planktonic material. A continuous
“rain” of decomposing organisms from the epilimnion, contributes
to an anaerobic condition in the hypolimnion of many lakes, as has
been indicated by previous studies (Birge and Juday, 1911 ; Ruttner,
1966; Hutchinson, 1957).
(c) 1962
Thermal data were gathered from Lake Mendota on all but three
days in 1962 (Fig. 5). The data were collected at station 2
(Fig. 1). The average time of sampling during the day for the
entire year was 1418 hours with a standard deviation of one hour
and 34 minutes.
Following the rapid freeze in December of 1961, the lake gradu¬
ally increased its content of heat as noted by the slight rise in
temperatures at most depths.
In the spring, two days before the ice melted completely, there
were large areas of open water at the sampling station. However,
a big sheet of ice moved across the lake the following day and
arrived over the sampling station when the measurement was
taken. Thus, there was no apparent refreezing on 10-11 April as
Figure 5 might suggest. The remainder of the ice vanished during
the night. Following the departure of the ice, vernal circulation
occurred and the lake took that big inspiration of air, as Birge
(1908) described, until the density differences overcame the in¬
fluence of the wind and stratification began.
In 1962 and 1963 the calm clear days stand out in the form of
sharp peaks (Figs. 5 and 6), as much heat is absorbed at the sur-
96 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Figure 5. Lake Mendota temperature and oxygen 1962. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day means plotted
on the tenth day.
1972] Stewart and Hasler— Limnology of Madison Lakes 97
Figure 6* Lake Mendota temperature and oxygen 1963. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day means plotted
on the tenth day.
98 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
face. This sharp stratification at the surface is destroyed easily by
winds. Temperatures on these calm days were taken within one
centimeter of the surface with the bead thermistor of the Whitney
thermometer. Diurnal variations at the surface are significant on
calm days, particularly days associated with clear nights, owing
to heat loss by long wave radiation. On windy days, the surface
temperature was measured within the first 10 to 20 centimeters
of the surface when a similar displacement either way made no
difference in the value.
Thermal changes at the surface appear to be more rapid and
dramatic in 1962 than in 1960 and 1961. However, these fluctua¬
tions reflect the plot of daily measurements instead of weekly or
scattered data, as was the case in 1960 and 1961. Below six meters
of depth, there is rarely any indication of diurnal variation. The
temperature at 12 meters fluctuates greatly, but this fluctuation
is a function of internal waves and not diurnal heat flux.
The rise of air temperature in the fall, during the period com¬
monly known as “Indian Summer/’ does little to raise the tem¬
perature of Lake Mendota. Although the temperature of the air
during the day may exceed considerably the temperature of the
water, the evenings are usually cool. Consequently the average
daily air temperature may differ but slightly from the water tem¬
perature. The integrating effect the heat capacity of water pro¬
vides, may allow the mean water temperature to remain similar
for several days. This is illustrated fairly well in 1962 but to a
lesser extent in 1960 and 1961 owing to less frequent sampling.
Lake Monona (Figs. 8-10), and Lake Waubesa (Figs. 11-13) par¬
ticularly, tend to respond more rapidly than Lake Mendota to
changes in air temperature owing to their lesser volumes and heat
budgets.
After the relatively warm period, usually in October, there is
a precipitous drop in air and water temperatures preceding the
winter months. The remaining stratification is eliminated and
complete autumnal overturn commences. Heat is then lost at a
relatively rapid rate at all levels until freezing.
About 20% of the lake surface froze on 12 December. The water
temperature was 0°C from the surface down to 10 meters on that
day. This unusually cold temperature was verified with a preci¬
sion mercury thermometer. This thickness (10 m.) of water at
such low temperatures was apparently quite unusual for as Birge
(Neess and Bunge, 1957, page 61) remarks:
“It is not impossible, theoretically, that the water of a lake should
reach, in whole or in great part, a temperature of 0 degrees C., but it is
very improbable that such a low temperature should actually occur before
the lake froze. When the temperature has fallen below 1 degree, ice forms
1972] Stewart and Hasler— Limnology of Madison Lakes 99
on the lake if the air is cool, even during considerable wind. . . Still more
easily does freezing occur if no wind blows. In either case the rate of
conduction in water is so slow that the layer at a temperature of 0
would be very thin.”
However, Birge (Neess and Bunge, 1957, pages 70-71) in the
same article provided data (29 Dec 1911, the day after freezing)
for a thermal structure which was remarkably similar to that
recorded on 12 Dec 1962. Birge stated, “It is not likely that a
lower temperature at freezing will be found than that of 1911.”
Obviously, time has at least provided an equal.
By 14 December, about 60 to 70% of the lake was frozen but
strong winds broke up the ice-cover and reduced it to about 40%
by 15 December. The lake did not freeze completely until 24 De¬
cember. The circulation from 15 to 24 December set the stage for
cooler water temperatures during the winter than the previous
1961-1962 winter.
Measurements of dissolved oxygen were made weekly at depth
intervals of three meters (Fig. 5).
The utilization of dissolved oxygen in the lower waters pro¬
ceeds more slowly in the winter owing to reduced temperatures.
However, there is a significant reduction in the concentration of
oxygen in the lower waters during the ice-cover.
Within a few days after the ice has melted, higher values of
oxygen are present than at any other time of the year. This fea¬
ture, recorded each year when timely data were available, is com¬
mon to the three Madison lakes discussed and was recorded to a
lesser degree by Birge and Juday (1911). However, the magnitude
and rapid formation of this 1962 peak was somewhat unexpected
because a winter oxygen deficit had to be removed before the
peak and concomitant supersaturation with oxygen could occur.
No similar peak occurred a few days prior to freezing even though
the water was colder, could have held more dissolved oxygen, and
the summer oxygen deficit had long since been repaid. In fact, com¬
pensation for the oxygen deficit of summer is seen at the end of
the summer stratification when upper waters are mixing to greater
depths with entrainment of hypolimnetic water and resultant
lowering of upper oxygen values. Therefore, the main reason for
the high values of oxygen after the ice goes out lies in the phyto-
planktonic production of oxygen.
(b) 1963
Measurements of temperature were made at station 2 (Fig. 2)
in Lake Mendota every day in 1963. The mean time of sampling
was 1418 hour with a standard deviation of 1 hour and 37 minutes.
The time of sampling was nearly identical to that of 1962.
100 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
After a relatively cold winter, the ice melted in a single day.
Following this striking change in the thermal structure (Fig. 6),
vernal circulation began and continued as in 1962 until the dif¬
ferences in density between the upper and lower waters required
more energy than was available to maintain complete mixing. This
period was described carefully by Birge (Neess and Bunge, 1956).
The two peaks of surface temperature on 1 July and 8 August
were 28.8 °C and 29.5 °C respectively. The four highest surface
maxima ever recorded on Lake Mendota by Birge, (Neess and
Bunge, 1957) were 34.3°, 32.1°, 29.0°, and 29.9°C on 29 July
1916, 30 June 1910, 23 June 1911, and 20 June 1913 respectively.
All of the above surface maxima were recorded between the hours
of 1300 and 1600.
The general summer curve of both air and upper water tempera¬
tures is more abrupt in 1963 than 1962. This abruptness illustrates
the correspondence between air and surface temperatures. As
Birge (Neess and Bunge, 1956) noted:
“There is far less correlation between the air and surface during autumn
than during spring and summer. In spring, and especially in summer, the
surface follows the air pretty regularly, though always with smaller
range and with a decided lag which sometimes obscures the relation.
But in autumn no such close relation is to be affected after the lake has
become homothermous, partly also to the increased evaporation of warm
periods which uses up more heat, and thus prevents a corresponding rise
of surface temperature.”
An extended warm period maintained a partial thermal stratifi¬
cation for three weeks in October. Note the step-like shape of the
graph after the autumnal circulation commenced (Fig. 6). The
lake froze initially on 17 December and finally on 20 December.
With this fairly rapid closure, the temperatures at the six and 12
meter level were somewhat higher than they were in the first three
months of 1963.
Measurements of dissolved oxygen in 1963 were initiated in
March and, with the exception of one two-week period in late
July and early August, were taken weekly until August after
which time the lake was sampled every three to five days. Con¬
sequently the total number of measurements of oxygen in 1963 ex¬
ceeded that of 1962. Again note the peak of dissolved oxygen
shortly after the ice melted.
The more frequent sampling in 1963 provides additional detail
that was not apparent from the 1962 graph of Lake Mendota
(Fig. 5). Oscillations of the standing internal wave were respon¬
sible for some of the internal variation at the 9 and 12 meter level
during August and September. The step-like variations of oxygen
during autumn correspond fairly well to those of water tempera-
1972] Stewart and Hasler — Limnology of Madison Lakes 101
ture. Immediately after the lake froze, the lake restratified rapidly
with respect to temperature and oxygen.
(e) 1966
Data were gathered weekly at station 2 (Fig. 1) in Lake Men-
dota from 19 March (three days after an early ice-out) to 30
August (Fig. 7). Cold weather followed the warming trend, which
induced an early opening of the lake, and air temperatures dropped
rapidly. Thus the lake actually lost heat in the first week after
measurements were initiated. Then the customary rise in tem¬
peratures began during vernal circulation and the summer stratifi¬
cation developed later.
The winds certainly aided the loss and gain of heat to the lake
during spring after ice-out but their reduced effect during summer
provided no particular contrast to most previous years.
The maximum surface temperature recorded was 29.4 °C on 1
July. The lack of several sharp temperature peaks, as noted in 1962
and 1963 (Fig. 5 and 6) reflects the longer interval between
sampling and not the complete absence of hot calm days.
The concentrations of dissolved oxygen did not increase in such
a striking manner shortly after ice-out as was noted in 1962 and
1963. In fact the 1966 spring curve was similar to the one re¬
corded (Birge and Juday, 1911) in 1907. However, a prominent
sub-surface maximum of oxygen was recorded at three meters on
1 July 1966. The dominant alga during mid-June and early July
was Aphanizomenon flos-aquae. Lesser amounts of Anabaena and
Staurastrum were also present during this period.
2. Lake Monona, Temperature and Oxygen
(a) 1961
All the sampling from Lake Monona was carried out in the
deepest (21 meters) area at station 3 (Fig. 1). This deep zone of
the lake is in a small area and is somewhat difficult to locate, there
being less than 0.01% of the volume below 18 meters. Fewer
measurements were made in 1961 (Fig. 8) than in either of the
following two years.
The average depth of Lake Monona (7.7 meters) is intermediate
to that of Lake Mendota (12.4 meters) and Lake Waubesa (4.6
meters). Its response to air temperature is intermediate also, i.e.,
it warms and cools more rapidly than Lake Mendota but less
rapidly than Lake Waubesa. This is also substantiated by compar¬
ing the dates of freezing and opening of Lake Monona with Lakes
Mendota and Waubesa (Bunge and Bryson, 1956, Parts I and II).
The highest surface temperature recorded in 1961 was 28.8 °C on
28 July.
mg/ litir
102 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 7. Lake Mendota temperature and oxygen 1966. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day means plotted
on the tenth day.
1972] Stewart and Hasler — Limnology of Madison Lakes 103
Figure 8. Lake Monona temperature and oxygen 1961. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day means plotted
on the tenth day.
104 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Lake Monona has apparent “plankton blooms” more frequently
than Lake Mendota but less commonly than Lake Waubesa.
The dissolved oxygen becomes depleted rapidly in the lower wa¬
ters. Furthermore, it is not uncommon to have low oxygen values
within six meters of the surface.
The early onset of anaerobic conditions in the lower waters of
Lake Monona reflects the small hypolimnetic volume and slightly
warmer temperatures when compared to Lake Mendota.
Lake Monona receives an unnatural inflow of warm water year
round. The major source of this warm water is the thermal dis¬
charge from the Madison Gas and Electric Company. Water is
withdrawn from Lake Monona at a depth of approximately 4.6
meters through two intakes located about 106 meters from shore
off Blount and Livingstone Streets.
The cooling water from Lake Monona is heated in the con¬
densers of the power plant and returned to the lake with its tem¬
perature increased about 10 °C above the ambient lake temperature.
The power plant, with a maximum capacity of 594 X 103 m3/day
(157 mgd), discharges the heated water through two surface out¬
falls, also at Blount and Livingstone Streets (Zeller, 1967).
The thermal discharge of the Madison Gas and Electric Company
appears to have an influence on the departure of ice from Lake
Monona. This influence is apparent in spring when the area of open
water, expanding outward from the thermal discharge, allows wind
and wave activity to break up the remaining ice more readily.
Although the mean opening date of Lake Monona (5 April) over
the past 115 years precedes Lake Mendota (6 April) by only one
day, the opening date in the 15 year period (1950-65) for Monona
(20 March) precedes Mendota (8 April) by 19 days (Data from
Ragotzkie, 1960; personal records of K. M. Stewart; and records
of Capital Times Newspaper, Madison, Wisconsin, 1966).
A second minor source of heated water is from a local meat pack¬
ing company which discharges some water through a viaduct into
the Yahara River, which in turn empties into Lake Monona. The
physical and biological impact of this water may be important
at the immediate site of the discharge but, owing to its lesser
volume and irregular nature, appears to have little significance for
Lake Monona with respect to thermal structure.
In fact, other than locally, the overall influence of the relatively
warm water from both of these companies to Lake Monona was
slight at the time of this study when compared to the influence of
the general climatological conditions. However, it is likely that
the thermal discharge of the Madison Gas and Electric Company
will have an increasing impact on the formation and departure
of ice on Lake Monona as the power demands of the City of Madi-
1972] Stewart and Hasler — Limnology of Madison Lakes 105
son, and consequently the volume of cooling water increases. Some
detailed information on the local biological effects is being studied
(Magnuson, 1970).
(b) 1962
Measurements of temperature and dissolved oxygen in Lake
Monona in 1962 began in April and continued until early December
(Fig. 9).
The thermocline is usually less dictinct in Lake Monona than
it is in Lake Mendota. The maximum surface temperature recorded
in 1962 was 28.4°C on 80 June.
The general profiles of dissolved oxygen were fairly similar to
those of 1961. The two highest concentrations of oxygen at the
surface were 15.5 and 15.9 mg/1 on 7 June and 30 June respec¬
tively. Water withdrawn from the hypolimnion in Lake Monona
in August smells more strongly of H2S than anaerobic water from
either Lakes Mendota or Waubesa. The stronger odor of H2S at
this time follows a longer period of anaerobiosis in Lake Monona
than the other two lakes. The quantity of dissolved oxygen in the
upper waters is lowered prior to full autumnal circulation. The
amount of lowering or raising of the oxygen content reflects the
oxygen demand of the hypolimnetic waters and sediments.
(c) 1963
Over 80 trips were made to Lakes Monona and Waubesa in
1963 (Fig. 10 and 13). Temperature and oxygen measurements
were made twice a week except for a two week period in the latter
part of July and early August, when only temperatures were
measured. To determine the daily variation, temperatures were
measured 20 out of 21 consecutive days during late June and
early July.
The highest temperatures at the surface were 29.6 °C, 29.9 °C and
28.8 °C on 1 July, 19 July, and 8 August respectively. The tempera¬
tures in the lower waters of Lake Monona were roughly 2 de¬
grees higher than those in Lake Mendota and 4 degree lower than
those in Lake Waubesa. Note the very rapid restratification of
temperature and oxygen after the freezing date on 16 December
1963.
The high value of oxygen (15.7 mg/1) on 16 April, represents
at least a portion of the early peak after complete ice-out (3 April,
1963). This supersaturated condition indicates a large production
of oxygen by phytoplankton as was noted in Lake Mendota and
as will be apparent in Lake Waubesa. The other high value of
oxygen recorded was 14.4 mg/1 and occurred on 8 September.
106 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
1972] Stewart and Hasler — Limnology of Madison Lakes 107
Figure 10. Lake Monona temperature and oxygen 1963. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day averages
plotted on the tenth day.
108 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
3. Lake Waubesa, Temperature and Oxygen
(a) 1961
The sampling position for all years in Lake Waubesa is indicated
at station 4 (Fig. 1). Some prominent limnological features of
Lake Waubesa (Fig. 11), when compared to Lakes Mendota and
Monona, are the high mean summer temperatures and the rapid
response to wind and air temperature changes as reflected by
dramatic changes in stratification.
The highest temperature recorded on the surface was 29.5 °C
on 28 July. Lake Waubesa stratifies but this stratification may
be broken down even in mid-summer. Less difference exists in Lake
Waubesa between the temperature of the surface and of the lower
waters than in Lakes Mendota or Monona. Lake Waubesa would
actually be quite stable during summer were it not for changing
air temperatures. However, as the air temperature drops the
water temperature also falls. Because a small change in tempera¬
ture is associated with a relatively large change in density at
higher water temperatures, the stability of stratification can
change quickly and the lake circulates.
Lake Waubesa begins its autumnal overturn about one month
earlier than Lakes Mendota or Monona.
Oxygen concentrations were measured at three depths during
this first year of sampling on Lake Waubesa. Frey (1940) noted a
thermal and marked oxygen stratification in Lake Waubesa earlier.
Figures 11-13 (this paper) show this even more clearly. We men¬
tion this because there is a local belief that Lake Waubesa circu¬
lates freely all summer and does not stratify.
(b) 1962
The relatively rapid response of Lake Waubesa to climatological
conditions is noted again this year (Fig. 12). For example, the
spring warming of Lake Waubesa exceeded the rate of warming
in Lakes Mendota and Monona. A practical index of this warming
is noted by the earlier swimming in Lake Waubesa. During Octo¬
ber, the temperature of Lake Waubesa rose briefly but there was
no corresponding rise in Lakes Mendota or Monona. Waubesa is
generally the first of the Madison lakes, excluding little Lake
Wingra, to freeze and open. The highest surface temperature
recorded in Lake Waubesa during 1962 was 28.7° on 6 July.
Lake Waubesa appears to be in a state of almost perpetual
algal “bloom” and the concentration of dissolved oxygen varies
considerably. The surface waters are supersaturated frequently.
Continuous measurements of dissolved oxygen were not made on
Lake Waubesa, but if they were, greater diurnal variation of
oxygen in the surface waters would be expected than in the other
1972] Stewart and Hasler — Limnology of Madison Lakes 109
Figure 11. Lake Waubesa temperature and oxygen 1961. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day averages
plotted on the tenth day.
110 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 12. Lake Waubesa temperature and oxygen 1962. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day averages
plotted on the tenth day.
1972] Stewart and Hosier— Limnology of Madison Lakes 111
two lakes. Therefore, potential programs for sampling Lake Wau-
besa should retain limits on the time of sampling to provide rela¬
tive data.
(c) 1963
The ice disappeared 3 April 1963 and the measurements of
temperature and dissolved oxygen began on 9 April (Fig. 13).
■Data were collected more intensively and over a longer period of
time than in any previous year. The highest temperatures re¬
corded at the surface were 29.8 °C and 30.4 °C on 29 June and 1
July respectively. The many sharp peaks and general variability
of lake temperatures in early July also reflect the limited period
of increased sampling.
In conjunction with changes in air temperature, it is interesting
to note how much more closely the crest of water temperatures
in Lake Waubesa follows the crest of solar radiation (Langleys)
than in Lakes Monona or Mendota.
Following the departure of the ice, high values of dissolved
oxygen are prominent. However, within the next six weeks, the
oxygen content of the lower waters plummets nearly to zero. A
few days after this, the passage of a cool front caused a
lowering of the water temperature. Then, even with relatively
low winds, the lake essentially “turned over”. This pattern, which
affects the mid-summer stratification of temperature and oxygen,
is demonstrated several times in 1963. The algal “blooms” create
supersaturated conditions quickly. For example, the concentration
of dissolved oxygen on 21 August was 21.7 mg/1, which after
correcting for water temperature (25.8°C) and altitude, meant
a saturation of 271%,
The somewhat intermittent summer stratification and mixing
in Lake Waubesa may raise serious problems, for individuals trying
to interpret deposition in sediment cores.
The lake froze completely on 14 December.
C. Ice Thickness: (Lake Mendota only)
The earliest studies of ice on Lake Mendota were conducted
by Buckley (1900) and Birge (Neess and Bunge, 1957) and his
co-workers. Buckley was concerned with fracturing and expan¬
sion of ice as well as the physical effects of ice ramparts on the
shores of lakes. Birge investigated the rates of growth and decay,
thickness of ice, temperatures within the ice-layers, and insola¬
tion beneath the ice. Measurements of the thickness of the ice
were conducted over 12 to 14 winters prior to and including the
winter of 1916-17. The data from nine of those years were plotted
(Fig. 14) by Birge* (Neess and Bunge, 1957, Fig. 45). The maxi-
*Birge continued his interest in ice beyond 1916-17 and, as Bunge and Bryson
(1956, Part I) noted, “Collected at least 27 years of ice thickness data and about
30 years of winter water temperatures from 1894 to 1930”.
q/ liter
112 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 13. Lake Waubesa temperature and oxygen 1963. Wind velocity, solar
radiation (Langleys), and air temperature are moving ten day averages
plotted on the tenth day.
1972] Stewart and Hasler — Limnology of Madison Lakes 113
mum thickness recorded by Birge in those years was 75 cm in
1899. Only 30 cm were found in 1913. The maximum thickness
recorded in this more recent study was 64 cm in 1963.
Birge (Neess and Bunge, 1957) separated winter into three
periods with respect to ice, namely, a period of increase in thick™
DEC. JAN. FEB. MARCH APRIL
LAKE MENDOTA
Figure 14. Ice thickness on Lake Mendota. Top half of figure illustrates
earlier data of Birge for the years 1898-99 (A), 1900-01 (B), 1902-03 (C),
1911-12 (D), 1912-13 (E), 1913-14 (F), 1914-15 (G), 1915-16 (H), 1916-17
(I). Lower half of figure illustrates more recent data on ice thickness,
measured every day in the winter of 1961-62, 1962-63, and the first portion
of 1963-64.
114 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
ness, a period of stationary thickness during February and part
of March, and the period of rapid decrease in thickness.
During 1912, Birge (Neess and Bunge, 1957) observed that
the melting of ice took place mainly at the surface with but a
small fraction of ice melting at the ice-water interface. In a more
recent ablation experiment, Scott and Ragotzkie (1961) found
that during approximately the last two weeks prior to ice out, the
total ice melt on the surface exceeded the bottom melt about two¬
fold. Independent of whether the ice melt from the bottom is
less than or equal to one half of the surface melt, it is apparent
that most of the calories required for melting the ice do not
come from the water itself. Juday (1940) was aware of this when
evaluating the annual energy budget of Lake Mendota.
The inverse correspondence between ice thickness and air tem¬
perature is fairly obvious during the early growth and later wasting
of ice. However, the period of “stationary thickness” appears less
subject to change from fluctuating air temperatures.
Following the rapid freeze of smooth sheet ice on 15 December
1961, measurements of the ice thickness were made daily from
20 December 1961 for the remainder of the ice cover. The thick¬
ness of the ice was also measured daily in 1962 and 1963, and
for part of the 1963-64 winter. All data were collected at or near
Station 2 (Fig. 1). The measurements were made by inserting
a meter stick, with a bar at right angles to the base of the stick,
through a hole in the ice. The thickness was that distance from
the bar, brought up against the underside of the ice, to the top of
the ice.
The curve (Fig. 14) for 1961-62 is based on an average of two
measurements each day taken approximately 200 meters apart.
The curves for 1962-63 and 1963-64 are based on one measure¬
ment each day in new holes that were chopped or drilled in an
area of fairly uniform ice thickness. Care is required during meas¬
urement because it is easy to have day-to-day differences in the
measured thickness of ice that are not indicative of climatic
changes. Scott and Ragotzkie (1961) have shown that significant
variations in ice thickness may exist a few meters apart when
clear ice (thicker) and snow-covered ice (thinner) are compared.
Birge (Neess and Bunge, 1957) would have noticed this difference
also had his ice data been collected more frequently during the
winter.
Rather dramatic increases in ice thickness are noted after a
midwinter melting of snow followed by refreezing. This “quick”
growth of new ice is not always apparent when drilling a hole
unless the lake is sampled regularly or unless a bubbly or crusty
surface remains.
1972] Stewart and Hasler — Limnology of Madison Lakes 115
A “lens” of warm water beneath the ice is a common phenom¬
enon in the last few days of ice cover. This warm “lens” is usually
over and underlain with colder water thereby giving an impression
of hydrostatic instability. For example, on 7 April 1962, five days
prior to the disappearance of ice, a water temperature of 5.7 °C
was recorded at 53 cm below the ice-water interface. The ice was
22 cm thick. The temperature at 128 cm below the ice was 3.2 °C.
Birge (Neess and Bunge, 1957) also recorded these warm layers
during late winter. Doubts as to the stability of these layers are
usually allayed by measurements of electrical conductivity, which
when converted to a common temperature, indicate an increase in
electrolytes which counters the thermal differences.
The mean dates of freezing and opening for the years from
1852 to 1965 are 19 December and 6 April respectively. (Ragotzkie,
1960; Capital Times, 1966; and personal records of senior author,
K.M.S.) The specific days at which these events take place are
influenced greatly by weather conditions in the immediate pre¬
ceding days. Thus, as mentioned earlier, Lake Mendota may
freeze partially, then reopen partially, and so on until complete
closure.
D. Aspects of Light
1. Secchi disk
Although the results of Secchi disk readings will be described
in more detail elsewhere (manuscript in preparation), it is of
interest to note that the clarity of Lake Mendota in 1961, 1962,
and 1966 was equivalent to or better than it was in 1916 from
the data of Birge (Neess and Bunge, 1957).
Comparing 45 measurements in Lake Mendota and 30 in both
Lakes Monona and Waubesa, all in 1962, the mean values of the
Secchi disk were 4.6, 2.0, and 0.92 meters respectively. Generally
there is a noticeable decrease in transparency as one travels from
Mendota to Monona to Waubesa.
The Secchi disk was viewed at 13.2 meters in Lake Mendota
on 22 March 1969 while the lake was covered with ice. This is
a new record for transparency on the Madison lakes.
2. Submarine Photometer
Separate results from the submarine photometer were also uti¬
lized to measure changes in the clarity of water over time (Fig.
15). From the variability observed, it is obvious that measure¬
ments of the extinction coefficients (computed as in Hutchinson,
1957, p. 381) (ranging from .303 to 1.227 for data between 1 to
5 meters) may be valid for only one day. “Representative” slopes
on a semi-log graph may be difficult to obtain for one lake. The
116 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
PHOTOMETER
RELATIVE LIGHT UNITS
Figure 15. Range of variation in the submarine photometer readings in
Lake Mendota (1966).
excellent early investigations by Birge and Juday (1929) on the
transmission of solar radiation would have shown this overlap
more clearly if several readings had been taken on each lake be¬
sides Mendota.
3. Transmissometer
A transmissometer (one meter path length) was lowered hori¬
zontally to measure microstratification within the three lakes, at
half meter intervals, from August through December 1963. This
period extended from times of significant thermal, chemical, and
biological stratification, through the autumnal overturn, and into
the initial stages of ice-cover. The results for Lake Mendota are
illustrated in greatest detail by isometric projection in Figure 16.
Separate standard inserts in the upper left and lower right of
the same figure provide comparisons with Lakes Monona and
Waubesa.
During late summer, Lakes Mendota and Monona were still
stratified but the water clarity improved markedly below the ther-
mocline. Although very slight thermal stratification remained in
1972] Stewart and Hosier— Limnology of Madison Lakes 117
Lake Waubesa on 14 August 1963, the microstratification, as meas¬
ured by the transmissometer was gone and the water was very
turbid from top to bottom.
The relatively turbid epilimnetic waters of Lakes Mendota and
Monona might be expected owing to the normally increased quantity
of phytoplankton in the euphotic zone. However, Whitney (1938)
using essentially the same instrument in Lake Mendota, found a
decrease in transparency below the thermocline. He attempted to
relate microstratification to bacterial populations.
There were occasions, noted by Whitney (1938) and in this
more recent study, when there was a temporary decrease in trans¬
parency within or just below the thermocline during the descent
of the thermocline. One explanation for this metalimnetic decrease
might be some greater planktonic, bacterial, or detrital densities
in that zone of rapid thermal transition. Another or combined
possibility, since the thermocline may separate anaerobic and
aerobic water, is a redox zone of dissolution and precipitation of
ferric hydroxide. Mortimer (1941) noted a turbid layer at the
upper level of the hypolimnion and did attribute it to a zone of
iron oxidation.
Figure 16. Microstratification in Lake Mendota as measured by a trans¬
missometer (one meter light path) in 1963. Separate inserts in the upper left
and lower right corners provide some comparison of Lakes Mendota, Monona,
and Waubesa.
118 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
In 1938 and 1963 the water immediately over the mud-water
interface was more turbid. The turbid water near the mud-water
interface may reflect boundary layer disturbances from oscilla¬
tions of standing waves as well as possible density or turbidity
currents along the slopes (Hutchinson, 1941).
The general pattern of light transmission for the latter half
of 1963 in Lake Mendota, as indicated by the transmissometer,
was roughly inverse to that of the thermal profile. That is, the
clarity of the water improved in the colder lower waters in sum¬
mer and throughout the lake as the lake cooled during autumn.
The Secchi disk transparency increased during the fall of the
previous year (1962) as well. The relationship between cold water
and better transmission may have been merely fortuitous for those
lakes on those years because algal blooms do occur during autumn
and under ice as well (Sawyer, 1947).
Changes in the development and disappearance of layers in
a lake can be noted by the changes in transmission of light with
time and depth. For example, the clarity of the lower water on
4 August and the clearer, more uniform condition on 25 December
are readily apparent (Fig. 16 and 17).
On 26 September (Fig. 17) there was a turbid layer at one to
one and one-half meters that was overlain with unusually clear
water in the first half meter. The clear water at the surface may
have allowed or created light inhibition of a certain algal com¬
munity there while augmenting prolific algal activity at one to one
and one-half meters.
Another unexpected feature was recorded on 8-9 October 1963.
On these dates, a turbid and coffee-colored layer extended from the
surface to about two meters (Fig. 17). This unusually dark water
was most common in the southeastern portion of Lake Mendota
near the University of Wisconsin. However, the discoloration ex¬
tended quite some distance across the lake as well. Therefore, it
is not likely that it was something “simply washed in” from rain.
More probably the dark water was an intensive surface algal bloom
although its specific composition was not determined at that time.
The turbid layer of 8-9 October disappeared by 10 October.
The general significance of these findings and those of others
(Whitney, 1938; Sauberer, 1962; Mahringer, 1963; Stewart et al.
1966; and Pinsak, 1967) is that the transmissometer can be used
to trace or monitor the development or disposition of these layers.
Furthermore rapid changes in Secchi disk and photometer values
are understood and interpreted more readily when it is realized
that such changes may reflect a microstratified zone or narrow
layer as well as a general change in water clarity from the quantity
and size of algae or inert suspensoids.
1972] Stewart and Hasler — Limnology of Madison Lakes 119
PERCENT TRANSMISSION
Figure 17. Microstratification in Lake Mendota as measured by a trans-
missometer (one meter path length) on four dates in 1963.
Had Secchi disks and submarine photometer readings been taken
in Lake Mendota on 26 Sept 1963 or 3-9 Oct 1963, their results
alone may have been puzzling to the investigator and might have
given a skewed general picture of the clarity of the water.
IV. General Significance
Lake Mendota is a dimitic eutrophic lake in the north temperate
zone. By the beginning of the year, the lake is normally frozen.
The ice gains in thickness rapidly from the time of freezing through
January. During February and early March, the rate of gain in
ice-thickness decreases. In the latter part of March and early
April there is a precipitous decrease in ice-thickness until the ice
disappears.
120 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
The mean temperature of the water column under the ice rises
gradually through the winter owing to absorption of solar radia¬
tion and heat flow from the mud. The concentrations of dissolved
oxygen in the lower waters declines during periods of ice-cover.
When the ice departs, dramatic changes occur in the lake. The
inverse thermal stratification of winter ceases and a relatively short
period of homoiothermy follows. There is rapid heating of the en¬
tire water column during vernal circulation.
An interesting feature observed in all three lakes is indicated
by the rapid rise in the content of dissolved oxygen at all levels
within a few days after the ice melts. The total column of water
has values of dissolved oxygen that are higher than at any other
time of the year. On the basis of greater solubilities at lower tem¬
peratures one would expect theoretically that the highest values
of oxygen would occur in the last few days just prior to freezing.
This sudden increase in oxygen reflects increased photosynthetic
activity of phytoplankton.
As vernal circulation continues, rapid increases in density dif¬
ferences between the developing epilimnion and the hypolimnion
establish limits on the depth of mixing. Lakes Mendota and
Monona are relatively stable, thermally, during late June and July
and August. Lake Waubesa, although primarily a dimictic lake as
are Lakes Mendota and Monona, has a shorter period of summer
stratification and may be subject to aperiodic turnovers. The
total quantity of dissolved oxygen in the lower waters of all three
lakes generally decreases rapidly after the onset of stratification.
There is usually a period during the partial autumnal overturn
during which the temperatures of Lakes Mendota and Monona
remain relatively unchanged. Lake Waubesa shows this to a lesser
degree because of its more rapid response time to climatological
influences. The physical response of Lake Monona to climatological
variables is intermediate to that of Mendota and Waubesa.
At this point in the annual cycle of the lakes, it usually happens
that a cold northerly wind cools the remaining epilimnion further,
overcomes the remaining density differences, and permits com¬
plete mixing. The oxygen deficit in the hypolimnion is repaid
quite rapidly and an extended period of complete circulation usu¬
ally follows. With decreasing water temperatures and increasing
oxygen content, the lake proceeds toward the day or days that it
freezes. Birge (1908) described appropriately the periods of
autumnal and vernal circulation as the “inspiration’’ periods dur¬
ing a respiratory process.
The day each lake freezes, usually during December, as the
day each lake opens in the spring is “critical” in the sense that
1972] Stewart and Hasler — Limnology of Madison Lakes 121
physical and chemical conditions change dramatically following
this event.
Lake Waubesa is in a state of almost perpetual algal “bloom”
and demonstrates the most rapid variability of any of the three
lakes during their annual cycles. The algal conditions generally
are reflected in the descending clarity of the lakes from Mendota
to Monona to Waubesa.
V. Summary and Conclusions
These studies on some physical (temperature, light, and ice)
and chemical (oxygen) variables in the Madison lakes provide
detailed data on the annual cycles in Lake Mendota and the rela¬
tively little studied Lakes Monona and Waubesa.
The patterns of physical change within the Madison lakes are
dictated primarily by morphometric and cyclic climatological in¬
fluences. Prominent among the climatological influences are the
temperature of the air, wind velocities and directions, and solar
radiation. Even a cursory examination of these variables illustrates
the huge changes that they can impress upon the lakes. The lakes
respond to these cyclic external changes but do so through an inte¬
grating process.
These more recent data of Lake Mendota, especially when com¬
pared to those of Birge and co-workers several decades ago, are
for the most part surprisingly similar. The thermal structure,
oxygen profiles, and light readings of the lakes resemble those re¬
corded by earlier investigators of the Madison lakes and do not
in themselves indicate a significant change in the lakes.
The role of the thermal discharge into Lake Monona from the
Madison, Gas and Electric Company may have an increasing
effect as the population of Madison grows. This will be most obvi¬
ous visually in the earlier disappearance of ice.
Owing to the useful information transmissometers provide con¬
cerning microstratification of organisms, turbid layers, and pos¬
sible chemical and thermal stratification, it is unfortunate these
instruments are not utilized more widely.
Owing to the vast scientific, recreational, aesthetic, and eco¬
nomic importance of the Madison lakes, it is advisable, as sug¬
gested (Sawyer, 1947; Stewart and Rohlich, 1967), to devote a
small fraction of the research efforts and funds of a laboratory
to a program of systematic surveillance for physical, chemical,
and biological variables. These efforts would provide basic limno¬
logical data, vital for good management considerations, as well as
knowledge of trends or changes in the lakes which may affect the
direction of future research.
122 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Acknowledgments
The helpful comments of C. H. Mortimer during his Brittingham
Fellowship at the University of Wisconsin and constructive criti¬
cism of H. H. Lettau (Meteorology Dept.) and J. C. Neess (Zoology
Dept.) at the University of Wisconsin contributed greatly.
Grants to A. D. Hasler from the National Science Foundation
(G-14530), U. S. Public Health Service (5T1-WP-2), and the Wis¬
consin Conservation Department and to K. M. Stewart from the
Monroe Co. Conservation Council (9505-3236) N. Y., supported
this research and appreciation is expressed for funds which made
it possible.
References Cited
American Public Health Assoc. 1960. Standard Methods for the Examination
of Water and Wastewater. 11th ed. New York.
Annual Climatological Data. 1959-1966. U.S. Dept, of Commerce Weather
Bureau Data for Truax Field, Madison, Wisconsin.
Birge, E. A. 1908. The respiration of an inland lake. Pop. Sci . Mo., 72:
337-351.
Birge, E. A. 1916. The work of the wind in warming a lake. Trans. Wis.
Acad. Sci. Arts Lett., 18: 214-250.
Birge, E. A., and C. Juday. 1911. The Inland Lakes of Wisconsin. The dis¬
solved gases of the water and their biological significance. Bull. Wis. geol.
nat. Hist. Surv., 22: 259 p.
Birge, E. A., and C. Juday. 1929. Transmission of solar radiation by the
waters of inland lakes. Trans. Wis. Acad. Sci. Arts. Lett., 24: 509-580.
Birge, E. A., and C. Juday. 1931. A third report on solar radiation and
inland lakes. Trans. Wis. Acad. Sci. Arts Lett., 26: 383-425.
Birge, E. A., and C. Juday. 1933. The transparency, the color, and the
specific conductance of the lake waters of northeastern Wisconsin. Trans.
Wis. Acad. Sci. Arts Lett., 28: 205-259.
Buckley, E. R. 1900. Ice ramparts. Trans. Wis. Acad. Sci. Arts Lett., 13:
141-157.
Bunge, W. W., and R. A. Bryson. 1956. Ice on Wisconsin lakes, parts 1 and
2. Report to the Univ. of Wisconsin Lakes Investigation Committee.
Univ. of Wisconsin, Madison.
Capital Times Newspaper. 1966. Section of paper, p. 4, on “History of Lakes”,
18 March, 1966, Madison, Wisconsin.
Frey, D. G. 1940. Growth and ecology of the Carp Cyprinus carpio Linnaeus
in four lakes of the Madison region, Wisconsin. Ph.D. Thesis. Univ. of
Wisconsin, Wisconsin.
Hutchinson, G. E. 1941. Limnological studies in Connecticut. IV. The mecha¬
nism of intermediary metabolism in stratified lakes. Ecol. Monogr ., 11:
21-60.
Hutchinson, G. E. 1957. A Treatise on Limnology, Vol. I. Geography,
Physics, and Chemistry. John Wiley & Sons, New York. 1015 p.
Juday, C. 1940. The annual energy budget of an inland lake. Ecology, 21:
438-450.
Magnuson, J. 1970. Personal Communication at the Laboratory of Limnology,
University of Wisconsin during July, 1970.
1972] Stewart and Hasler — Limnology of Madison Lakes 123
Mahringer, W. 1963. Einschichtung und Verteilung des zufliessenden Wassers
irn Millstattersee. Arch . Hydrohiol , 59: 272-280.
Neess, J. C., and W. W.. Bunge. 1956. An unpublished manuscript of E. A.
Birge on the temperature of Lake Mendota; Part I. Trans . Wis. Acad .
Sri. Arts Lett , 45: 193-238.
Neess, J. C., and W. W. Bunge. 1957. An unpublished manuscript of E. A.
Birge on the temperature of Lake Mendota; Part II. Trans . Wis. Acad.
Sri. Arts Lett, 46 : 31-89.
Pinsak, A. P. 1967. Water transparency in Lake Erie. Proc. 10th Conf.
Great Lakes Res., 309-321.
Ragotzkxe, R. A. 1960. Compilation of freezing and thawing dates for lakes
in North Central United States and Canada. Tech. Rep. 3, ONR Contract
No. Nonr. 1202(07). Dept, of Meteorology, Univ.. of Wisconsin, Madison.
61 p.
Ruttner, F. 1966. Fundamentals of Limnology. University of Toronto Press,
Toronto, Ontario, Canada. 295 p.
Sauberrr, F. 1962. Empfehlungen fur die Durchfiihrung von Strahlungs-
messungen an und in Gewassern. Mitt, int Verein . theor . angew. LimnoL,
No. 11, 77 p.
Sawyer, C. N. 1947. Fertilization of lakes by domestic drainage. J. New
England Water Wks . Ass., 51 : 109-127.
Scott, J. T., and R. A. Ragotzkie. 1961. The heat budget of an ice-covered
lake. Tech. Rep. 6, ONR Contract No. 1202(07). Dept, of Meteorology,
Univ. of Wisconsin, Madison. 52 p.
Stewart, K. M., K. W. Malueg, and P. E. Sager. 1966. Comparative winter
studies on dimietic and meromictic lakes. Verh . int theor . angew . LimnoL,
16: 47-57.
Stewart, K. M., and G. A. Rohlich. 1967. Eutrophication — a review. Calif .
St Wat Quah Control Bd ., Publ. No. 34: 188 p.
Whitney, L. V. 1938. Microstratification of inland lakes. Trans . Wis . Acad .
Sri: Arts Lett, 31: 155-173.
Zeller, R. W. 1967. Cooling water discharge into Lake Monona. Ph.D. Thesis.
Univ. Wisconsin, Madison.
THE ALGAE OF THE WINNEBAGO POOL AND
SOME TRIBUTARY WATERS1'2
William E. Sloey and John L. Blum
Introduction
The Winnebago Pool (Lakes Winnebago, Poygan, Winneconne
and Butte des Morts) is one of the more important water resources
in Wisconsin. Frequent algal blooms interfere with domestic, in¬
dustrial and recreational use, but there have been no comprehen¬
sive reports on the algae. Smith (1920, 1924) mentioned the rel¬
ative abundance of some species present in Lake Winnebago in his
taxonomic treatise on the Phytoplankton of the Inland Lakes of
Wisconsin; and Marsh (1903) made a study comparing the zoo¬
plankton and phytoplankton of Lake Winnebago to those of Green
Lake. Recently, Leuschow et at., (1970) studied the major groups
of net plankton in Lake Winnebago during the open water period.
They compared Winnebago to eleven other lakes for trophic level,
and found it to be the most eutrophic of the lakes studied.
During a two and one-half year period from 1966 to 1968, the
senior author made a physical, chemical and biological study of
the Winnebago Pool with particular attention to Lake Butte des
Morts. The physico-chemical limnology, C-14 primary productiv¬
ity, phytoplankton standing crops and community structures were
reported previously (Sloey, 1970). This report is limited to the
population dynamics of the predominant phytoplankton organ¬
isms. A partial species list of the algae of the Pool, of the upper
Fox and lower Wolf Rivers and of some tributaries is also included.
Methods
Sampling
Water samples for phytoplankton counts were collected at five
selected stations (Figure 1) at the surface and 0.2 m above the bot¬
tom with a horizontal sampler (Howmiller and Sloey, 1969). Ali¬
quots of 2-10 ml, the amount depending on plankton density, were
1 This study was supported in part by a research grant from the Board of Reg-ents
of the Wisconsin State Universities and in part by a graduate fellowship provided by
the Botany Department, University of Wisconsin — Milwaukee.
2 The authors would like to thank Dr. Ruth Patrick of the Philadelphia Academy
of Natural Sciences, Philadelphia for her kind assistance in confirming some of the
diatom species identifications and for the time spent in her laboratories by Dr. Sloey
during the summer of 1967.
125
126 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
delivered with a 10 ml Mohr pipette having a 4 mm bore onto 25
mm diameter HA Millipore filters and filtered. The rapid Millipore
filter counting technique of McNabb (1960) was employed and
all counting was done at 1000 X. Two slides were prepared from
each of the surface and bottom water samples. The mean of the
four counts (2 each, surface and bottom) was determined for
Figure 1. A map of the Winnebago Pool and a portion of the associated water¬
shed showing sampling sites. Location: east-central Wisconsin.
i972] Stoey and Blum- — Algae of the Winnebago Pool 127
each species or group of algae. This was done in the hope of cor¬
recting to some extent for variation .within samples, as well as
between samples (from surface and bottom, respectively) and so
as to limit the samples to a number that could be handled.
On the days of plankton sampling, a tow net haul was also made
at each station to obtain large numbers of organisms for species
identification. During the summer of 1966, a qualitative study of
the benthic algae was made outside the limits of the numbered
stations by scraping submerged rocks, vegetation and navigational
buoys, and several plankton sampling transects were made of rep¬
resentative portions of the Pool and of the Upper Fox and Wolf
Rivers and their tributaries.
Within 24 hours of sampling, water mounts were prepared from
the tow hauls and the scrapings made; all the Chlorophyta, flagel¬
lates, and Cyanophyta that could be found in 8-6 transects of the
slide were identified. The more uncommon species, were recorded
on photomicrographs. Some of this material was then cleaned in
H0SO4 and dichromate, or simply ashed (Patrick and Reimer,
1966) and mounted in Hyrax for detailed study of the diatoms.
The remainder of the sample was preserved with 5% formalin and
filed, along with the Hyrax mounts, at the Wisconsin State Uni¬
versity, Oshkosh. The location of a voucher specimen for each spe¬
cies of diatom was recorded on a 3x5 card and filed with the slides
and photomicrographs.
Frequency of sampling
Plankton samples were taken at weekly intervals during the
summer of 1966, intermittently during the winter of 1966-67, bi¬
weekly during the ice-free periods of 1967 and 1968, and monthly
during the winter of 1967-68.
References used for the identification of diatoms were : Hustedt,
1930a, 1930b; and Patrick and Reimer, 1966. Those consulted for
other groups were: Prescott, 1962; Smith, 1920 and 1924; Tiffany
and Britton, 1952. Above the generic level, the nomenclature and
taxonomic treatments are those of Prescott (1962).
Determination of cell volumes
Two slides of Millipore filter mounts were selected from each
month's set of samples. Using the basic forms of cube, sphere,
cylinder and disc, the volume of at least ten units of each species
was determined from each slide. A “unit" was specified as a natu¬
ral aggregate of cells (e.g., filament of Melosira , colony of Asteri-
onella or a clump of Microcystis). Volumes were expressed as
/X3 X 106 ml-1.
128 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Determination of generation times
In a population of cells, each dividing vegetatively into two
daughter cells, a logarithmic or exponential growth pattern is as¬
sumed. This growth rate can be expressed in terms of the time
required for the population to double. Fogg (1965) described this
doubling time as a ''generation time” on the assumption that preda¬
tion, disease and losses by flushing and settling are insignificant
in cultures and in most planktonic situations. Under optimal con¬
ditions the generation time decreases. Making a similar assump¬
tion for the phytoplankton of the Winnebago Pool, we calculated
generation times (G) for the predominant algae according to the
formula of Fogg as follows:
r _ 0.301 t
” "log (N/N0)
where :
t = time
N = number of cells at the end of the time period
N0 = number of cells at the beginning of the time period
In this report, as in the recent literature, "generation time” is
used to designate apparent generation time. Obviously, patchiness
of phytoplankton populations, dilution from runoff and other fac¬
tors may affect a natural population. It will be necessary to con¬
firm field observations with laboratory cultures before the term
"generation time” can be literally applied to ecological studies.
Results
Occurrence and periodicities of individual species in the Pool
A total of 106 algal taxa were encountered in the Pool during
this study. One can assume that each of these formed part of a
population which varied with time, probably in response to favor¬
able and unfavorable conditions.
The bloom-forming blue-greens
ANABAENA spp. — The genus Anabaena is represented in Lake
Butte des Morts by five species, A . circinalis (Kutz.) Rabh., A.
lemmermanni P. Richter, A. limnetica G. M. Smith, A. plankton -
ica Bruhn., and A. spiroides Klebahn. The first two forms (A.
planktonica and A. spiroides) were more common in late summer
and fall. A. limnetica appeared intermittently throughout the sum¬
mer and seemed to be most frequent after rainy periods. These
forms are difficult to separate unless akinetes are present; thus,
routine counts were made at the generic level.
1972] Sloey and Blum — Algae of the Winnebago Pool 129
Temperature may be the key factor determining the presence
of Anabaena. In no case did significant populations occur below
18 C, and declines commenced when temperatures dropped below
this value. While populations fluctuated considerably, especially
during the summer of 1966, the minimum generation times (N =
no. of filaments) ranged from <1.0 during the first week in
August 1966 to 23 days in late July 1967. On the basis of biweekly
sampling intervals, it appeared that generation times averaged
between 2 and 5 days. In spite of lower temperatures in 1967, the
population maxima during that year were nearly twice as great
as during 1966 or 1968. The highest numbers recorded were 1,925
filaments ml-1 at station 4 on 28 July 1967 and 2,950 at station 3
on 22 August (Figure 2).
Aphanizomenon flos-aquae (L.) Ralfs. — This species occurs
with Anabaena and appears to have similar seasonal trends. The
similarity was particularly striking during 1967 (Figure 2), except
that Aphanizomenon had a brief surge in October and early No¬
vember. A surface film of colonies was noted on 1 November at
4 C; by 6 November the temperature had dropped to 2 C and no
filaments could be found. Populations during 1968 were much lower
than in 1966 or 1967. Maximum values reached only 365 colonies
per ml at station 2 on 22 August (1.46 million /A per ml), Lake
Winnebago had 827 per ml on the same date. The earliest seasonal
appearance was 24 May 1968 (29 per ml) at station 3 when the
bottom and surface temperatures were 15.3-15.8 C, respectively.
Apparent generation times (N = no. of colonies) averaged between
2 and 5 days and ranged from <1.0 to 29 days during periods of
population growth.
Microcystis aeruginosa Kiltz. emend. Elenkin 192A
Syns: M. aeruginosa var. major
M. flos-aquae (Wittr.) Kirchner
(see Prescott, p. 456-7)
Microcystis appeared in the plankton with the two previous gen¬
era at warm temperatures. Sizable populations appeared very sud¬
denly. For example, the population increased from 0 to 96 colonies
ml-1 between 10 and 24 May 1968. Once present, populations tended
to remain fairly constant throughout the summer (not shown).
Since few definite growth periods were detected, few generation
times could be determined. It is estimated that this period is 2-6
days inasmuch as the shortest period was 1.82 days (N = no. of
colonies) in early August 1966 and the longest was 6.19 days in
early July 1966.
The time of lowest temperature (15.0, 15.6 C, bottom and sur¬
face, respectively) was on 10 May 1968 at station 2. Unlike Aph-
130 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 2. Periodicity of Anabaena spp. ( - ) and Aphanizomenon flos-aquae
( - ) in the Winnebago Pool, 1966-1968.
anizomenon, populations were larger in 1968 than in 1967. The
greatest number was 404 colonies ml-1 (10.6 X 106 p? ml-1) on
22 August 1968 at station 1.
Oscillatoria Agardhii Gomont . — This species did not appear
to be of major biological significance in the Pool, but its appear¬
ance as a minor bloom in the low light regime under the ice dur¬
ing February and March was very striking. The highest number
found was 80 filaments per ml on 27 March 1967 at station 3. This
was just prior to the spring ice breakup.
1972] Sloey and Blum — Algae of the Winnebago Pool 131
The planktonic pennate diatoms ( Araphidineae)
Asterionella Formosa Hass. var. Formosa. — Asterionella for-
mosa populations fluctuated very rapidly in Lake Butte des Morts.
The species appeared primarily in late fall (October to December)
and in late spring (May to July, Figure 3). Populations in the
upper Fox River (station 2) and in Lake Winnebago were con¬
siderably smaller than at the other stations. Particularly large pop¬
ulations occurred in the Wolf River in November 1966 and at sta¬
tions 1 and 4 in May and June 1968. During 1967, large populations
failed to materialize at any of the stations in either spring or fall.
The maximum values recorded were 6,000 cells per ml on 6 June
at station 4 and 5,300 cells per ml at station 1 on 24 May 1968.
The latter value represented 34.5% of the total phytoplankton
volume, the largest percentage found. Fall populations frequently
persisted under the ice. In fact, there were only very small popu¬
lations in Lake Winnebago during 1966 and 1967, but 390 cells
per ml appeared under the ice in February 1968. Growth periods
of the populations occurred during September to December and
March to June. Apparent generation times ranged from 3.06 days
in early May, 1968 to 40.5 days in December, 1966 and averaged
about 10 days. The average number of cells per colony increased
from 4 to 5.4 during early June; this immediately followed a pop¬
ulation maximum and a generation time minimum.
Synedra spp. — The genus Synedra is represented in the Win¬
nebago Pool by at least six species and/or varieties. Like Anabaena,
these forms are difficult to separate during routine counting on the
Millipore filters and thus were also counted at the genus level.
The following entities were present:
Synedra acus Kiitz. var. acus. Fairly common during the spring
bloom.
S. delicatissima var. angustissima Grun. Important in the plank¬
ton from April until June.
S. pulchella Ralfs. ex. Kiitz. var. pulchella. An autochthonous epi-
lithic and epipelic form which occasionally appeared in the
plankton.
S. ulna (Nitz.) Ehr. var. ulna. In spring and fall plankton.
S. ulna var. amphirhynchus (Ehr.) Grun. Fairly common in the
spring and late fall plankton.
S. ulna var. longissima (Wm. Smith) Brun. Occurred only in the
fall in small numbers.
Members of the genus Synedra were primarily spring and fall
forms, but considered together, one or more species were present
132 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
on almost every sampling date (Figure 3). The smallest and most
erratic populations were seen in Lake Winnebago. By far the great¬
est number occurred at station 2 on 24 May 1968 (5,700 cells per
ml), when Synedra delicatissima var. angustissima represented
12.2% of the total phytoplankton volume. Generation times (N =
no. of cells) at the genus level ranged from 1.93 days in early Au¬
gust 1966 when populations were low to 27.5 days in April 1967
when populations were high.
Figure 3. Periodicity of Asterionella formosa ( - ) and Synedra spy. (- - -)
in the Winnebago Pool, 1966-1968.
133
1972] Sloey and Blum — Algae of the Winnebago Pool
The centric diatoms
Stephanodiscus Hantzschii Grun. — This nannoplankter was
the dominant form during the period of ice cover and immediately
thereafter. The species was present in higher numbers during the
summer of 1968 than during 1966 or 1967 (Figure 4). The larg¬
est number of cells recorded was 14,030 per ml on 29 December
1967 when it comprised some 96.5% of the number and 75.3% of
the volume of the phytoplankton. Minima occurred during the July-
September maxima of Melosira spp., S. niagarae and the blue-
greens. Generation times (N = no. of cells) during the growth
periods ranged from 1.55 days in June 1966 to 57.1 days in De¬
cember 1967. Generation times during spring and fall increases
averaged 9 days.
While midwinter declines in the populations occurred at every
station, there is no question but that this very light and hyaline
species persisted better in the relatively stable conditions under
the ice than any other diatom observed. There was an eight-fold
apparent increase under the ice in 1968 at station 2 (Figure 4).
Stephanodiscus niagarae Ehr. — Stephanodiscus niagarae was
primarily a late summer-fall form. By far the greatest popula¬
tions occurred during October and November 1966 (Figure 4).
On 8 November of that year, 686 cells mk1 were present at sta¬
tion 2 (22.6 X 106 jjl3 ml-1) . This species was seldom found in the
plankton during the more stable ice-cover period. It is probable
that S. niagarae is also relatively heavy ; this could account for its
settling out from the plankton under the ice but other factors may
be responsible. Generation times (N = no. of cells) were quite
long, averaging 16.0 days for 21 positive growth periods. Growth
rates were more consistent over long periods of time for this spe¬
cies than for any of the others reported here. During the period
of 1 July to 24 October 1968, generation times varied only from
10.1 to 15.7 days at station 1.
Melosira ambigua ( Grun.) O. Mull. — Melosira ambigua was a
co-dominant plankter (along with M. granulata) from June until
October. This species must be considered “persistent” (Bozniak
and Kennedy, 1968) since it was present at almost every station
on every sampling date (Figure 5). During the summer period,
phenomenally high values were recorded. The Fox River at sta¬
tion 2 had populations as high as 98,700 cells per ml on 28 July
1967 and 93,600 per ml on 19 July 1968. The latter value repre¬
sented 62.64 million p3 mk1 and 73.5% of the total phytoplankton
volume. Maxima, which occurred as one or more peaks between
June and September, were normally 3 to 5 times higher in the
upper Fox River (station 2) than in the Wolf (station 3) . Minima
134 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 4. Periodicity of Stephanodiscus Hantzschii ( - ) and S . niagarae
( - ) in the Winnebago Pool, 1966-1968.
occurred under the ice, but the species never completely disap¬
peared from the plankton. Certainly if any organism could be said
to be “typical” of Lake Butte des Morts or the Winnebago pool,
it would be Melosira ambigua. Exponential phases of population
growth occurred between April and July, and a secondary growth
phase appeared in early fall when water temperatures started to
decline. Cell division rates appeared therefore to be related to
water temperatures. Optimum growth rates occurred between 10
and 13 C in early May when the mean generation time (N = no.
1972]
Sloey and Blum — Algae of the Winnebago Pool
135
Figure 5. Periodicity of Melosira ambigua ( - ) and M. granulata ( - ) in
the Winnebago Pool, 1967-1968.
of cells) was only 5.37 days. The autumn population growth rate
was somewhat slower, with a minimum mean generation time of
9.74 days during September at water temperatures of 19.6 to
20.9 C. Midsummer and late fall growth rates were even slower.
Populations decreased throughout October and November (except
for station 3).
The number of cells per filament ranged from 6.1 in January to
28.6 in mid- July
136 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Melosira granulata (Ehr.) Ralfs. — Melosira granulata was a
co-dominant with M. ambigua from June until October when the
two species usually represented more than 70% of the total phyto¬
plankton volume and values as high as 95% were recorded. To¬
gether, they dominated the phytoplankton throughout the Winne¬
bago pool and as far upstream in the Fox River as Endeavor, some
75 miles above station 2 (see below). Maxima of M. granulata
were not reached until late August or September, well after the
initial maxima of M. ambigua, and at water temperatures above
20 C. The highest numbers recorded were 115,500 and 83,211 cells
ml-1 in Lake Winnebago on 22 August and 5 September 1968,
respectively (Figure 5). At station 2 there were 65,000 cells per
ml (57.08 X 106 y? ml"1) on 22 August. Populations decreased
during late autumn and reached winter levels well before ice for¬
mation.
Generation times (N = no. of cells) for M. granulata followed
a pattern similar to that for M. ambigua. Population growth did
not commence, however, until after ice -out at the end of March,
and minimum doubling times occurred between late May and early
July, rather than April.
The number of cells per filament increased from 2.5 in February
to 32 on 22 August. The longest filaments occurred after maximum
growth rate periods, but at approximately the same time as the
population maxima. By 13 November the mean filament length
had decreased to only 5 cells.
Melosira binder ana Kiitz. M. binder ana was an occasional plank-
ter in Lake Winnebago in spring but was not found in Lake Butte
des Morts.
M. italica (Ehr.) Kiitz., subsp. subarctica O. Miill., (status
alpha) . M. italica was found only in a few later winter collections.
It was also found in the upper Fox and Wolf Rivers.
M. granulata, var. angustissima O. Mull. This variety was not
found in any sample before 10 May and was present only sporad¬
ically until September. The number suddenly increased to a peak
in October with a maximum number of 1,347 filaments and 26,900
cells per ml on 24 October 1968. Filament lengths also increased
from about 8 or 10 to 20 cells during October. By 13 November
populations had dropped to only about 100 filaments ml-1. Inas¬
much as this narrow “variety” appears after the period of auxo-
spore formation of the nominate variety (15-30 July) one must
question if this not a separate species. To determine this, culture
studies are needed.
M. varians C. A. Ag. Occasionally occurs in the plankton, but
is considered a littoral form (Hustedt, 1930sl, p. 86) and is quite
1972] Sloey and Blum — Algae of the Winnebago Pool
137
common in rock scrapings and to a lesser extent in scrapings from
macrophytes.
Other algae found in the Winnebago pool
The other algae, those of lesser importance in the phytoplank¬
ton, and the benthic forms, are listed below according to their
general taxonomic affinities.
Bacillariophyceae
Achnanthes lanceolata (Breb. in Kiitz.) Gran, in Cl. and Grun.
Observed occasionally in the plankton during late winter and
early spring.
Amphipleura pellucida Kiitz., var. pellucida.
Amphiprora ornata Bailey.
Caloneis ventricosa (Ehr.) Meist., var. truneatula (Grun.) Meist.
Cocconeis pediculus Ehr., var. pediculus. Very common epilithic
and periphytic form in lakes Butte des Morts and Winnebago.
C. placentula Ehr., var. lineata (Ehr.) V. H. Common epiphyte,
occasionally found in plankton.
Cyclotella Kiltzingiana Thwaites. Rare, mostly in the Wolf River
(station 3).
C. Meneghiniana Kiitz. Uncommon.
Gymatopleura elliptica (Breb.) W. Smith.
C. solea (Breb.) W. Smith. Common, but never numerous.
Cymbella parva (W. Smith) Cleve.
C. prostrata (Berkeley) Cleve. Common epilithic and epiphytic
form.
C. tumida (Breb.) Van Heurck. Occasional plankter.
Diatoma tenue A g., var. elongatum Lyngb. Occasional in spring
plankton.
D. vulgare Bory., var., vulgare . Common epilithic and epiphytic
form that occasionaly occurs in the plankton.
Epithemia turgida (Ehr.) Kiitz., var. capitata Fricke.
Eunotia parallela Ehr., var. parallela.
Fragilaria capucina Desmazieres. Common in spring and fall plank¬
ton.
F. construens (Ehr.) Grun., var. construens. Occasional plankter.
F. crotonensis Kitton, var. crotonensis. Common plankter, especially
in spring and early summer.
F. vaucheriae (Kiitz.) Peters, var. vaucheriae.
Gomphonema sp. Very common epilithic and epiphytic form. Speci¬
mens are similar to Hustedt’s G. tergestinum (1930a, p. 378)
but striae are much coarser (7-8/10 jx) .
Gyrosigma attenuatum (Kiitz.) Rabh., var. attenuatum. Occasional
cold water planktont.
138 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
G. scio tense (Sulliv. & Warmley) CL, var. sciotense. Occasional
spring and fall planktont.
G. spencerii (Queck.) Griff & Henfr., var. spencerii.
Meridion circular e (Grev.) A g., var. constrictum (Ralfs) V. H.
Occasionally occurs in the plankton in winter and early spring.
Navicula cuspidata (Kiitz.) Kiitz., var. major Meist.
N. exigua Greg, ex Grun., var. capitata Patr. Found only in one
sample in March. Patrick lists this as a soft water form.
N. gastrum (Ehr.) Kiitz., var. gastrum.
N. lateropunctata Wallace, Var. later opunctata.
N. menisculus Schumann, var. upsaliensis (Grun.) Grun.
N. pupula Kiitz., var. rectangidaris (Greg.) Grun.
N. salinarum Grun., var. intermedia (Grun.) Cl. Common epilithic
form.
N. scutelloides W. Smith ex. Greg., var. scutellodies. Common, but
never numerous in the plankton.
N. viridula (Kiitz.) Kiitz. emend. V.H., var. viridula.
Neidium productum (Wm. Smith) Pfitzer.
Nitzschia acuta Hantzsch.
N. amphibia Grun.
N. filiformis (Wm. Smith) Hust.
N. gracilis Hantzsch.
N. palea (Kiitz.) Wm. Smith,
N. sigmoidea (Ehr.) Wm. Smith. Fairly common in spring and fall
plankton.
N. spectabilis (Ehr.) Ralfs.
Opephora martyi Herib., var. martyi. Fairly common in late winte?
and spring plankton.
Pinnularia brebissonii (Kiitz.) Rabh., var. brebissonii.
P. gentilis (Donk.) CL, var. gentilis .
P. major (Kiitz.) Rabh., var. transversa (A.S.) Cl. striae slightly
fewer (6-7/10 p) than Patrick’s description (8-9/10 n) .
P. streptoraphe CL, var. streptoraphe.
Rhoicosphenia curvata (Kiitz.) Grun. ex. Rabh., var. curvata. Very
common benthic form.
Stephanodiscus tenuis Hustedt. Fairly common in late winter and
spring plankton.
Surirella tenera Gregory, var. nervosa. Most common in June.
Surirella, sp. 1. This species is similar to S. linearis (Hustedt,
1930a ; p. 434), but specimens are larger with wider canals.
Surirella, sp. 2. Similar to S. capronii (Hustedt, 1930a, p. 440), but
lack the well developed central costae.
Synedra acus Kiitz., var. acus.
Tabellaria fenestrata (Lyngb.) Kiitz., var. fenestrata. Rare in Lake
Butte des Morts, occurring only in the spring.
1972] Sloey and Blum— Algae of the Winnebago Pool
139
Chlorophyceae
Asterococcus limneticus G. M. Smith. Rare summer form.
Cladophora glomerata (L.) Kiitz. Extensive growth on rocks, posts,
etc., wherever these is wave action. Frequently attached to
macrophytes in protected bays. Usually encrusted with peri-
phytic diatoms.
Chlamydomonas globosa Snow. Found only on a few occasions in
the Fox River (station 2) in the summer.
Closterium aciculare T. West, var. subpronum W. and G. S. West.
C. moniliferum (Bory) Ehrenberg. Associated with macrophytes in
bays and marshes.
Dictyosphaerium pulchellum Wood. Rare, in summer plankton.
Dimorphococcus lunatus A. Braun.
Pediastrum boryanum (Turpin) Meneghini. Appears mostly in the
fall.
P. duplex Meyen, var. gracillinum W. and G. S. West.
P. duplex , var. cohaerens Bohlin.
P. simplex (Meyen) Lemmermann, var. duodenarium (Bailey)
Ragh. Common in fall plankton.
Scenedesmus quadricauda , var. WestiL
S. quadricauda (Turp.) Breb., var. longispina (Chod.) G. M. Smith.
Selenastrum Bibrainum Reinsch.
Staurastrum longiradium W. & G. S. West. Common, but not nu¬
merous in plankton during summer and fall.
Tetraedron limneticum Borge. Occasionally appears in plankton
from August until October.
Chrysophyceae
Dinohryon divergens Imhof. Found only in Wolf River in early
summer.
Dinophyceae
Ceratium hirundinella (0. F. Muehl.) Jorgensen. Present in most
samples throughout the year, but never found in bloom pro¬
portions.
Myxophyceae
Aphanocapsa delicatissima W. and G. S. West. Occurred only in
the plankton at station 3 during November (water tempera¬
ture during one collection, 2.2 C).
Chroocoecus limenticus Lemmermann, var. subsalsus Lemmer¬
mann. Rare, during mid-summer.
C. limneticus , var. elegans G. M. Smith. Occasional, during early
summer.
140 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Coelosphaerium naegelianum Unger. A very common bloom former
in early summer.
Gloeotrichia echinuiata (J. F. Smith) P. Richter. Common in Lake
Winnebago in June; rare in Lake Butte des Morts as planktont.
G. natans (Hedwig) Rabenhorst. Common epiphyte in marsh areas
during late summer.
Merismopedia punctata Meyen.
The algae of the streams tributary to the Winnebago Pool
The Upper Fox River
The upper Fox River near the headwaters at Marcellon is a clear,
shallow stream with many aquatic macrophytes. The plankton is
represented primarily by tychoplankters and benthic escapes such
as Cymatopleura solea, Melosira varians, Cosmarium moniliferum,
Cocconeis placentula, Navicula spy,, Cymbella tumida, Phormidium
sp., Synedra ulna var. spathulifera (Grun.) V. H., Stauroneis sp.,
Diatoma vulgar e, Meridion circular e, Cyclotella Meneghiniana,
and Perionella planktonica. No “nuisance” blue-greens were found,
and only a few individuals of planktonic diatom species such as
Melosira ambigua, and Fragilaria capucina.
At Portage, below Park and Swan Lakes, some of the “nuisance”
blue-greens such as Microcystis flos-aquae, Anabaena spiroides and
A. circinalis were evident, along with considerable numbers of
planktonic diatoms such as Melosira ambigua and M. granulata and
Stephanodiscus niagarae. The plankton was, however, still repre¬
sented primarily by tychoplankters and benthic escapes including
the same species of Cocconeis, Cyclotella, Diatoma, and Cymbella
as was found at Marcellon. Achnanthes hungarica (Grun), var.
hungarica, M. varians, Synedra delicatissima, and Nitzschia sp.
were also present.
At Endeavor, the Fox River was already turbid and the plankton
was dominated by Melosira ambigua and M. granulata (about
75% of the diatoms) along with M. italica, Anabaena spiroides,
A. planktonica, Ceratium hirundinella, and Eudorina sp. Many of
the benthic forms were still present, but their contribution to the
total plankton was considerably reduced.
From Montello downstream, the plankton was typical of that
found in the Winnebago Pool. The predominant forms were
Aphanizomenon, Anabaena spiroides, Microcystis aeruginosa, Pedi-
astrum sp. Stephanodiscus niagarae and especially Melosira am¬
bigua and granulata. From Berlin downstream, Selenastrum bi-
braineum Reinsch, Eudorina sp., Chlamydomonas sp. and Scenedes-
mus quadricauda were also present in small numbers.
141
1972] Sloey and Blum — Algae of the Winnebago Pool
The Mecan River
The Mecan River at Montello is a marginal trout stream having
water slightly discolored from humic acids. The plankton was
sparse and composed primarily of Pediastrum sp Closterium sp
Navicula spp., Phormidium sp . and a few filaments of Melosira
granulata.
The White River
The White as sampled at County Highway D, seven miles east
of Princeton is a clear water trout stream. The plankton was
composed primarily of benthic escapes and organic debris along
with a few filaments of Melosira granulata and Microcystis
aeruginasa .
The Puckyan River
The Puckyan River as sampled from County Highway A east
of Princeton is a shallow stream having warm water and many
submerged aquatic plants. Skeins of Spirogyra sp . streamed from
the vegetation and the plankton was primarily epiphytes and
benthic escapes, but a few euglenoids and Chlamydomonas sp.
were present. Epiphytes included Navicula spp ., Cocconeis pla-
centula, Rhoicosphenia curvata , Cymatopleura sp. and Cymbella sp.
The Wolf River
The Wolf River at New London has a composite plankton com¬
posed primarily of euplankters such as Melosira ambigua, and
M. granulata with lesser numbers of M. italica , Fragilaria leptos-
tauron , F. capucina , Synedra acus, Navicula scutelloides , Cymbella
tumida , and Surirella caproni Breb. No blue-greens were present,
but considerable numbers of benthic forms such as Navicula
cryptocephala Kiitz. var. crypto cephala, M. varians, Nitzschia
sigmoidea , N. cuspidata , and Achnanthes lanceolata were present.
At Fremont, the Wolf River had a plankton composed almost
entirely of euplankters. These were dominated by Melosira granu¬
lata (including the variety angustissima) and M. ambigua . On
24 July 1966 no blue-greens were present. Other forms present
included Pediastrum duplex var. gracillimum , Synedra sp M.
italica , Chroococcus elegans , Pediastrum boryanum) Stephanodiscus
niagarae , M. varians , Surirella sp. and Cocconeis pediculus.
The Waupaca River
The Waupaca River below the reservoir at Weyauwega is a
warm-water stream containing mostly Melosira ambigua and Oscil-
latoria tenuis , with some M. granulata. Other forms present in¬
cluded Actinastrum sp Nitzschi sp Synedra acus , Meridian
142 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
circular e, Diatoma vulgar e, Cyclotella glomerata , Fragilaria ley-
tostauron, Melosia varians, Gyrosigma sp ., Opephora martyi,
Navicula crypto cephala, Cymhella tumida, and Synedra delicatis-
sima, var. angustissima.
The Walla Walla Creek
The Walla Walla Creek at County Highway EE north of Lake
Poygan is a small stream containing much floating plant debris.
The plankton was composed almost entirely of epiphytes and
benthic escapes, but a few filaments of Melosira ambigua and M.
granulata were present. Also present were Amphora ovalis, Navi¬
cula scutilloides , N. crypto cephala, Fragilaria leptostauron,
Opephora martyi, Cymhella tumida, Meridion circular e var. con-
strictum, Navicula accommada Hust. var. accommada, Cymato-
pleura solea, Cyclotella meneghiniana, Diatoma vulgare, Nitzschia
spp. and Synedra ulna var. amphirhynchus (Ehr.) Grun.
The Pine River
The Pine River (1 mile east of Waushara County Highway W
on 26th Street) near Saxville is a clear trout stream. The seston
is composed primarily of organic debris and benthic escapes. These
include Gomphonema constrictum Ehr. var. capitata (Ehr.) Cleve,
Fragilaria pinnata Ehrenberg, Melosira varians, Diatoma vulgare,
Navicula spp., and Cocconeis placentula var. lineata .
Below the reservoir at Poysippi, the Pine contains primarily the
gigantic Oscillatoria princeps Vaucher. About 25% of the diatoms
were Melosira ambigua and M. granulata. Also present were Am¬
phora ovalis, Navicula gastrum var. gastrum, Cocconeis placentula
var. lineata, Achnanthes saxonica Krasske var. saxonica, A. lanceo-
lata, and Synedra ulna var. contracta.
The Willow River
The Willow River at County Highway D near Rorth is a sluggish
stream containing many macrophytes and a wide variety of Chloro-
phytes and diatoms. These included Scenedesmus bijuga (Turpin)
Lagerheim, S. quadricauda, Anabaena planktonica Brun., Co sa¬
marium botrytis (Bory) Meneghini, Gloeotrichia natans (Hedwig)
Rabenhorst, Oocystis elliptica W. West, Merismopedia convoluta
de Brebisson in Kiitz, Synedra ulna var. contracta S. socia Wal¬
lace var. socia, Pinnularia sp., Cyclotella Meneghiniana, Cymhella
cuspidata Kiitz., C. sp., Cocconeis placentula var. lineata , Hantz-
schia sp., and Nitzschi spectabilis (Ehr.) Ralfs.
At County Highway Q above the Auroraville millpond, the Wil¬
low is a fast flowing trout stream and contains mostly detritus and
benthic escapes such as Melosira varians, Nitzschia spp. and
Neidium productum (Wm. Smith) Pfitzer.
1972] Sloey and Blum — Algae of the Winnebago Pool
148
The Pumpkinseed Creek
The Pumpkinseed Creek at County Highway D near Borth is
also a sluggish, warm-water stream. The plankton is predominated
by Synedra pulchella Ralfs ex. Kiitz. var. pulchella and S. acus.
Also present were Eunotia curvata (Kiitz.) Lagerst. var. curvata,
Eudorina elegans Ehrenberg, Gomphonema constrictum Ehr. var.
capitata Ehr. Cleve., Scenedesmus quadricauda var. longispina,
Staurastrum longiradium, Rhoicosphenia curvata and Melosira
varians.
Summary
General Observations
During this study, 106 taxonomic entities of algae were encoun¬
tered in the Winnebago Pool and additional forms were found in
the upper Fox and Wolf Rivers and their tributaries. Total stand¬
ing crops of phytoplankton were very high (volumes ranged to
110 X 106 jxs ml-1; Sloey, 1970).
Lake Butte des Morts and the Winnebago Pool exhibit phyto¬
plankton typical of shallow, eutrophic lakes and large rivers (Raw-
son, 1961, Fritsch, 1931, Eddy, 1934, and Hustedt, 1930a, 1930b) .
Plankton similar to that in the Pool predominated in the upper
Fox as far upstream as Endeavor and in the Wolf as far upstream
as New London. The Pool, then, as concerns its phytoplankton
populations, represents little more than a widened expanse of these
rivers.
In general, the upper Fox contained higher populations of most
species than the Wolf or the lakes of the Pool. Lake Butte des
Morts contained higher populations than Winneconne-Poygan
(represented by station 3 at town of Winneconne) , and higher pop¬
ulations of all predominant forms, except Melosira granulata, than
Lake Winnebago.
Throughout the year, except for brief periods during blooms
of blue-greens, the centric diatoms Melosira ambigua and M . gran¬
ulata and Stephanodiscus niagarae and S. Hantzschii dominated
the plankton. M. ambigua was most abundant during early summer,
M. granulata during late summer, S. niagarae during autumn and
S. Hantzschii during winter. In late autumn and early spring, the
araphids, Synedra spp. and Asterionella formosa reached maximum
numbers. When water temperatures exceeded 15 C, the “nuisance”
blue-greens, Anabaena spp., Aphanizomenon flos-aqua and Micro¬
cystis aeruginosa were evident and at times dominated the plankton.
Generation Time as a Measure of Growth Conditions
Minimum generation times for a number of algae were deter¬
mined. These ranged from 1-3 days for bloom forming blue-greens
144 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Table 1. A Comparison of Generation Times of Phytoplankton in the
Winnebago Pool with Those Reported by Other Authors*
*References Cites: Holland, R., 1969; Fogg, G. E., 1965, and Verduin, J., 1952.
such as Anabaena and Microcystis to 16 days, for Stephano discus
niagarae. The generation times reported here are reasonably con¬
sistent with those from the literature (see Table 1) .
A shortened generation time of a population apparently reflects
favorable growth conditions. In this study no species continued
to increase without interruption until population maximum. In
each case numerous dips were observed which probably resulted
from sampling error, basin flushing or other (generally unknown)
external causes. However, the rate of each recovery after a dip
undoubtedly reflected the optimality of the environmental condi¬
tions for that organism. Nutrient levels were lowest when total
plankton counts were highest and basin flushing affected stand¬
ing crops (Sloey, 1970). Correlations of generation times to sea¬
sons and particularly to water temperature were obvious, but other
positive correlations between populations or generation times of
any species and water chemistry (Le., nitrate, phosphate, etc.) were
not observed. Water temperature is a result of seasonal weather
patterns and daylength; as such it might mask photoperiod re¬
sponses of the flora and it functions indirectly by altering meta¬
bolic rates, nutrient availability and flotation.
References Cited
Bozniak, E. G. and L. L. Kennedy. 1968. Periodicity and ecology of phyto¬
plankton in an oligotrophic and eutrophie lake. Canadian J. Bot., 46(10) :
1259-1271.
Eddy, S. 1934. A study of fresh-water plankton communities. Illinois Biol
Monogr., 12(4): 1-92.
Fogg, G. E. 1965. Algal Cultures and Phytoplankton Ecology. Univ. Wisconsin
Press. Madison, Wisconsin, 126 p.
Fritsch, F. E. 1931. Some aspects of the ecology of fresh-water algae. J. EcoL,
19(2): 16-272.
1972] Sloey and Blum — Algae of the Winnebago Pool
145
Holland, R. E. 1969. Seasonal fluctuations in Lake Michigan diatoms. Limnol.
Oceanogr., 14(3) : 423-436.
Howmiller, R. P. and W. E. Sloey. 1969. A horizontal sampler for investiga¬
tion of stratified waters. Limnol. Oceanogr., 14 (2) : 291-92.
Hustedt, F. 1930a. Bacillariophyta. In: Pascher, A. Die Siisswasser-Flora
Mitteleuropas. 10: 1-466. Jena.
- . 1930b. Die Kieselalgen Deutsehlands, Osterreichs und der Schweiz
mit Beriicksichtigung der iibrigen Lander Europas sowie der angrenzenden
Meeresgebiete. In: Dr. L. Rabenhorst’s Kryptogamen-Flora von Deutsch¬
land, Osterreich-reichs u. der Schweiz. Akademisch Verlags-Gesellschaft,
Leipzig.
Leuschow, L., J. Helm, D. Winter and G. Karl. Trophic nature of selected
Wisconsin lakes. Wis. Academy of Science, Arts and Letters, 58 : 237-264.
Marsh, C. D. 1903. The plankton of Lake Winnebago and Green Lake. Wis¬
consin Geol. Nat. Hist. Sur. Bull. 12, 94 p.
McMabb, C. D. 1960. Enumeration of freshwater phytoplankton concentrated
on the membrane filter. Limnol. Oceanogr., 5(1) : 57-61
Patrick, R. and C. W. Reimer. 1966. The Diatroms of the United States, Vol. 1.
Monograph of the Academy of Natural Sciences of Philadelphia, No. 13,
Philadelphia, Pa. 688 p.
Prescott, G. W. 1962. Algae of the Western Great Lakes Area. W. E. Brown,
PubL, Dubuque, Iowa 977 p.
Kawson, D. S. 1961. A critical analysis of the limnological varieties used in
assessing the productivity of northern Saskatchewan lakes. Verb, Int.
Verein. Limnol., 14: 160-166.
Sloey, W. E. 1970. The limnology of hyp ereu trophic Lake Butte des Morts,
Wisconsin. Proc. 13th Conf. Great Lakes Res. 1970: 951-968: Internat.
Assoc. Great Lakes Res.
Smith, G. M. 1920. Phytoplankton of the Inland Lakes of Wisconsin. Part I :
Wisconsin Geol. and Nat. Hist. Surv., Bull. 57, Sci. Series, No. 12. 243 p.
- . 1924. Phytoplankton of the Inland Lakes of Wisconsin. Part II:
Desmidiaceae. Wis. Geol. and Nat. Hist. Surv., Bull. 57, 227 p.
Tiffany, L. H. and M. E. Britton. 1952. The algae of Illinois. Univ. Chicago
Press, Chicago.
Verduin, J. 1952. Photosynthesis and growth rates of two diatom communities
in western Lake Erie. Ecology, 33(2): 163-168.
KINETICS OF ORTHOPHOSPHATE UPTAKE
BY PHYTOPLANKTON POPULATIONS IN LAKE WINNEBAGO
Steven Bartell and Sumner Rickman
Abstract
The growth of natural phytoplankton populations of Lake Winne¬
bago was investigated with regard to orthophosphate limitation.
An ascorbic acid-antimony modification of a standard molybdenum
blue colorimetric test for orthophosphate showed a concentration of
94.0 fig • liter-1 in late November, 1970. This concentration steadily
decreased through the winter months to a minimum of 29.0
fig • liter-1 in early February, 1971. With the removal of ice cover
and increased allochthanous addition of phosphate associated with
the spring thaw, the orthophosphate concentration increased to
109.0 fig • liter-1 in early April.
A standard acetone extraction procedure showed that chlorophyll
concentrations followed a similar trend in seasonal variation. The
2.2 mg • m-3 concentration found in the January 9, 1971 sample
decreased to a minimum of 0.4 mg • m-3 for the January 24
sample. The chlorophyll concentration gradually increased to 20.8
mg • m-3 by early April.
Primary productivity measured by a standard 14C technique,
increase in biovolume determined by the Model B Coulter Counter,
and chlorophyll production were used to study the kinetics of
orthophosphate uptake in enrichment experiments where varied
amounts of orthophosphate, ranging from 20 to 200 fig • liter-1,
were added to samples of Lake Winnebago water.
The resulting changes in the rates of these parameters suggest
that uptake followed the Michaelis-Menten equation, modified for
nutrient limitation theory, for enriched samples collected on Febru¬
ary 2 and February 20, 1971. This model may also apply to the
November and April populations; however, a limitation in ex¬
perimental design prohibited the calculation of Vmax and Kt.
Orthophosphate was determined to be a limiting factor for
productivity in Lake Winnebago over the winter months.
I. Introduction and Background
Eutrophication refers specifically to the natural or artificial
addition of nutrients to aquatic ecosystems. This term has been
more broadly interpreted to include the physical, chemical, and
147
148 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
biological implications of nutrient enrichment. When these effects
are undesirable the process is a form of pollution. One of the
effects of increasing importance in aquatic biological research is
the role of nutrients in relation to the productivity of the system.
Because of the accompanying degradation of water quality for
recreational and industrial use, much concern has been given to the
increase in biological productivity that results from the addition
of nutrients to the aquatic system. While the process is natural
and a function of the age of the particular system, man’s industrial,
agricultural, and domestic activities may effectively increase the
natural rate of eutrophication. In order to determine how to best
minimize the effects of man’s activities on the rate of eutrophica¬
tion, the role of nutrients in relation to productivity must be
determined. This not only necessitates the elaboration of effects of
individual nutrients upon the system, but also includes the deter¬
mination of the synergistic relationships among the various plant
nutrients. For example, iron has been found necessary in some
systems to enhance the availability of other major nutrients, such
as nitrate and phosphate (Schelske, 1962). The particular nutri¬
ent (s) that determines the trophic nature of a system varies
greatly from lake to lake and from fresh water to marine environ¬
ments.
Ryther found nitrate to be the limiting factor in primary produc¬
tivity in Great South Bay off Long Island, New York (1971).
Schelske determined that iron was the limiting nutrient in several
Michigan marl lakes (1962). Micronutrients such as sulfur, potas¬
sium, magnesium, calcium, boron, zinc, copper, cobalt, sodium,
and chloride are also essential nutrients for growth (Lee, 1970).
Molybdenum and manganese have been shown to limit productivity
in Castle Lake, California (Goldman, 1965). Provasoli (1969)
has shown that certain organic compounds, such as vitamin B12j
thiamine, and biotin are necessary requirements for several marine
phytoplankters. However, the nutrient most often implicated as the
limiting factor in biological production is phosphorous (Wentz and
Lee, 1969).
Considering its importance as a vital structural component of
DNA, RNA, and protein, as well as its functional significance in
intermediary metabolism; the common occurrence of phosphate as
a limiting factor of productivity is logical from a theoretical view¬
point. In addition phosphate usually occurs in minute concentrations
compared to other nutrients in lakes (Tucker, 1957) .
The purpose of this study was to investigate the influence of
phosphorous on the realization of productivity potential in Lake
Winnebago by comparing the uptake of phosphate enrichments
by natural phytoplankton assemblages as the natural available
1972] Bartell and Rickman ■ — Orthophosphate Uptake 149
phosphate concentrations in the lake varied over a six month period.
While previous research of this nature has employed the direct
measurement of nutrient uptake by means of radioactively labelled
nutrient sources, such as 32P04 and 15N03 (Dugdale, 1967 ; Mac-
Isaac and Dugdale, 1969), this project emphasized a more indirect
approach. The rate of phosphate uptake was measured in terms
of three biological parameters. Increases in chlorophyll concentra¬
tion, population size in terms of biovolume, and primary production
were monitored as a function of phosphate concentration by en¬
riching samples of Lake Winnebago water with orthophosphate.
The use of these parameters lends more biological meaning to the
determined uptake kinetics. One might follow not only how the
nutrient is taken up, but also how the nutrient is utilized by the
plankton.
IL General Lake Information
Lake Winnebago is a large, shallow, fresh water lake located
in northeastern Wisconsin, with over 137,000 acres of surface area
and a maximum depth of 21 feet (Lueschow, Helm, Winter, and
Karl, 1970). The western shore is extensively developed. The
four largest cities on the lake, Fond du Lac, Oshkosh, Neenah, and
Menasha, are located on this shore. The remaining shoreline is
under private ownership, agricultural cultivation, or recreational
use, in the form of county and state parks. Lake Winnebago,
typically characterized by frequent plankton “blooms”, high
nutrient content, and low Secchi disc readings (annual mean of
2.3 meters) has been classified as one of the more eutrophic lakes
in Wisconsin (Lueschow, Helm, Winter, and Karl, 1970).
This lake is also important as the source of the lower Fox River,
which is its major outlet. This river ultimately flows into Green
Ray.
HI. Methods and Procedures
A. Sampling
Twelve nineteen liter samples of Lake Winnebago water were
collected biweekly over a period beginning in early November,
1970 and ending in early April, 1971. The samples were collected
by means of a battery operated pump from a depth that ranged
from 3 to 4 meters through surgical tubing to a Nalgene poly¬
propylene carboy. A piece of #11 plankton net (0.145 mm) placed
over the mouth of the carboy prevented the introduction of zoo¬
plankton to the sample.
During the period of ice cover, samples were collected approxi¬
mately one quarter mile offshore from Sportsman's Park, which is
located on the northern shore several miles southeast of Appleton
150 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
at fire lane #8. All other samples were collected near High Cliff
State Park, nearly two miles away from Sportman’s Park.
B. Phosphate
Usually 24, but no later than 48 hours after collection, 250 mis.
of the sample were Millipore filtered (.45/*). One hundred mis.
were used for an orthophosphate determination.
Because of the amount of controversy concerning the preserva¬
tion (Lee, 1969), storage (Hassenteufel et al., 1963; Strickland
and Parsons, 1965; and Lee, 1969), and measurement of ortho¬
phosphate in lake water; the polypropylene carboy was chosen for
sample storage. Similarly, all phosphate enrichment experiments
were performed in Nalgene polypropylene bottles to minimize loss
of phosphate by container absorption.
Due to the nature of the experiments, no preservatives were
added to the samples. The samples were stored at 15 C until tested.
Workers generally agree that refrigeration is the most preferred
method of sample preservation (Schelske, personal communication) .
Glass distilled water blanks, standard orthophosphate solutions,
and samples were all treated with a colorimetric test developed
by Murphy and Riley (1962), which uses ascorbic acid-antimony
as the reducing agent. This method is reported to have a low
temperature coefficient, stable color, and no salt error according to
Strickland and Parsons, as cited by Lee (1969). A Bausch and
Lomb Spectronic 20 equipped with red phototube and filter meas¬
ured absorbance of blanks, standards, and samples at 830 /*.
A one thousand milligrams per liter solution was made by adding
2.007 grams of Na3P04 • 12H20 to 500 mis. of glass distilled water.
This stock solution was the phosphate source for the enrichment
experiments and the phosphate test standard solutions.
On the same day that the initial phosphate tests were performed,
the enrichment experiments were begun. Subsamples were trans¬
ferred from the carboy so as to completely fill 1.98 liter Nalgene
polybottles, which had been previously autoclaved at 120 p.s.i. for
twenty minutes. Care was taken to thoroughly mix the carboy
sample in order to ensure the homogeneity of the five subsamples
in the experiment. Predetermined amounts of phosphate were
added to four of the subsamples. The fifth bottle, which contained
the phosphate present in the lake at the sample date, served as the
control. The samples were incubated for five days on individual
magnetic stirrers under constant temperature (15 C) and incident
white light (1000 ft. candles) saturation, which appears to be the
only light requirement for enrichment experiments of this nature
(Maclssac, 1969).
1972] Bartell and Richman — Orthophosphate Uptake 151
C. Particle-size distributions
The Model B Coulter Counter proved to be a fast, efficient device
for obtaining the distribution and abundance of phytoplankton
in the initial and nutrient enriched Lake Winnebago samples. Size
distributions were taken both at the beginning and at the end of the
experiments with a 100 /i aperture. The initial size distribution
represented the current population of phytoplankton in the lake.
The final distributions represented population growth due to phos¬
phate enrichment. The difference in biovolume divided by the dura¬
tion of the experiment in hours expresses a rate of biovolume
increase (cubic microns per ml. per hr.) as a function of phosphate
concentration.
The amount of detritus collected in several of the samples was
measured by means of the Coulter Counter to determine its sig¬
nificance in the enrichment experiments (Sheldon and Parsons,
1967). The one per cent NaCl electrolyte solution produced a max¬
imum of only one to two per cent error in sample counts at the
most sensitive machine settings.
D. Chlorophyll
In this study changes in chlorophyll concentration were used to
determine increases in phytoplankton populations as a function of
phosphate enrichment. Four replicate samples were taken from
the original Lake Winnebago sample at the beginning of the experi¬
ment. Similarly, chlorophyll was extracted in quadruplicates from
the control and experimental bottles at the end of the experiments.
The chlorophyll concentration was determined by a standard
trichromatic acetone extraction procedure (Strickland and Par¬
sons, 1965). The extracted volume ranged from 100 to 250 mis.,
depending upon the phytoplankton population density present in
the initial Lake Winnebago sample; however, the volume remained
internally consistent for any experiment the author performed.
A Bausch and Lomb Spectronic 20 measured absorbance at 630,
645, and 665 jx. The optical densities for all samples were punched
onto computer cards and fed into an IBM 1620 data processing
system. The duration of the experiment (in hours) and a dilution
factor were also included in the input of the CHLOREG program,
which calculates the concentrations of chlorophyll a and b using
the revised coefficients of Parsons and Strickland (1963) . Because
concentrations for chlorophylle were often less than zero, which is
attributed to a fault of the trichromatic test (Strickland and Par¬
sons, 1965), chlorophylL was deleted from the experimental design.
Having calculated the concentrations of chlorophyll a and b in
all samples, the CHLOREG program calculated the rate of chloro¬
phyll production in the control and enriched bottles which is
152 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
expressed as mg Chlorophyll • m-3 • hr-1. Finally, all possible
comparisons were made between the five populations to test
whether the production rates were significantly different from one
another. A contrast test, instead of the standard t test, was used
to calculate significant differences between chlorophyll production
rates at the .05 level (Dixon and Massey, 1969).
E. Primary Productivity
On the final day of the experiment, the rate of carbon fixation
was measured for each of the populations in the enrichment experi¬
ment by the standard 14C method (Vollenweider, 1969). A standard
phenolphthalein-methyl orange test was used to measure total alka¬
linity as ppm CaC03 (Standard Methods, 1965). This value was
corrected to total available carbon-12 by a factor determined by
the pH of the sample (Bachman, 1959). A Corning pH meter was
used to measure endpoints in the sample titrations for alkalinity,
as well as to measure sample pH.
The source of available 14C was labelled NaHC03 purchased from
New England Nuclear in ampoules containing one microcurie in
one ml. of sterile water (2.22 x 106 dpm). In all experiments, the
available 14C was one microcurie.
Ten ml. aliquots were filtered (.45 jx) at two hour intervals over a
six hour incubation from the light and dark bottles. This volume
was used to minimize loss of activity during period of filtration
(Arthur and Rigler, 1967).
The filters were dried in a vacuum at 35 C overnight. After dry¬
ing, the filters and 10 mis. of a scintillation cocktail consisting of
5.0 grams of PPO (2,5 diphenyloxazole) dissolved in 1000 mis.
of reagent grade toluene were placed in glass scintillation vials
and loaded onto a Beckman LS-230 Liquid Scintillation System.
The efficiency of counting this primary fluor system ranged from
an initial 87% to 92% in later experiments. The efficiency was
determined by making a set of standards with NaHC03 (4.422 x
105 dpm per ml.) in toluene as the 14C source (Packard Instrument
Company). Also, known amounts of 14C (NaHC03) were added
to several vials which were recounted to calculate efficiency. Count¬
ing efficiency was determined for each experiment for calculation
of carbon fixation. An isotope correction factor of 1.06 was used.
The carbon fixation rates were calculated by means of a least
squares analysis program available on a RAX IBM 360/44 com¬
puter. The program calculated slope (photosynthetic rate), y-inter-
cept, and correlation coefficient (r). In all cases, the value of r was
.90 or greater for the rate calculation.
1972] Bartell and Richman— Orthophosphate Uptake 153
IV. Results
A. Seasonal Phosphate and Chlorophyll Concentrations
The variation of phosphate and chlorophyll concentrations in
Lake Winnebago over the period of study as illustrated in Figure 1
demonstrates a somewhat parallel trend. While the measured
decrease in phosphate and chlorophyll occurs almost simultaneously
through the month of January, the increase in chlorophyll appears
to follow that of phosphate by about a one week interval through
the month of April. An increase in orthophosphate during one
week is followed by a similar rise in chlorophyll the following week.
This seems to substantiate two essential points in the enrichment
study. First, the soluble inorganic phosphate measured by the test
appears to be a legitimate form of phosphorous that is utilized by
the phytoplankton. Second, this form of phosphorous is a limiting
factor for population growth in Lake Winnebago, at least from
January to late March or early April. Hilsenhoff (1967) found
phosphorous to fluctuate greatly over a four year study of Lake
Winnebago, 1961-1964, but reports that phosphorous concentra¬
tions were lowest in the winter and highest in the summer. He
reports an annual mean of 2.0 /xg • liter-1 for a station near the
sampling site for this study. Soluble phosphate levels determined
by Sloey (1970) for a station in Lake Winnebago two miles east
of Oshkosh were lowest in late October and November. The values
range from 20.0 to 30.0 /xg • liter-1. However, he reports no values
for January or February.
The fivefold increase in the phosphate concentration from Janu¬
ary to April probably reflects the large amount of phosphorous
that enters the lake by agricultural and urban runoff during the
spring thaw. Phosphate concentrations as great as 1000 /xg- liter-1
have been measured in samples of surface runoff (Biggar and
Corey, 1969).
B. Seasonal Variation in Phytoplankton
The size distributions of the Lake Winnebago samples indicate
an interesting succession over the course of the study. The sample
collected November 24, 1970 exhibits two predominant populations
at 9.4 and 29.9 micron diameters (Figure 2). Figure 3 depicts the
tremendous decrease in the abundance of phytoplankton by Janu¬
ary 9, 1971. The shift in peak location indicates an apparent change
in the species composition by January 24. The 3.0 micron diameter
particle clearly dominates the community, while a sizeable decrease
has occurred at all other particle diameters. It must be pointed out
that this peak may be due to detritus or background noise at the
upper limit of machine sensitivity for the 100 /x aperture. The
February 2 sample displays the lowest abundance of phytoplankton
154 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
ifov» Dec, Jan. Feb. March April
Figure 1. Seasonal variation of phosphate (ortho) and chlorophyll <ft+b> con¬
centrations in Lake Winnebago from November, 1970 to April, 1971. Sample
depth is approximately 3 meters. Solid line = i^g • liter-1 P04„ (Encircled
point is from Lueschow, Helm, Winter, and Karl, 1970). Dashed line = mg
Chlorophyll(a+b> m-3.
of all samples collected. (Figure 4). It is interesting to note that
the lowest phosphate concentration was determined for this sample
(Figure 1), though light must have also been limiting under the
1972] Bartell and Richman — Orthophosphate Uptake
155
PARTICLE DIAMETER
</*>
Figure 2. Size distribution determined by Coulter Counter (100 /* aperture)
for a L. Winnebago sample collected November 24, 1970.
thick cover of ice and snow present at this date. By February 12,
the size distribution indicates the emergence of a population that
can maintain and reproduce itself under these severe environmen¬
tal conditions (Figure 4-b) . This phytoplankter at a diameter of
9.4 microns has convincingly established itself as the dominant
156 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
PARTICLE DIMETER
in
Figure 3. Size distribution determined by Coulter Counter for L. Winnebago
samples collected January 9 and January 24, 1971 (100 ^ aperture).
species of the winter population by February 20 (Figure 4-c).
Qualitative observations with a Zeiss inverted plankton scope
revealed a species of Asterionella to be the most abundant form in
this sample. The distribution of the March 1 sample reflects a sub¬
stantial change in the community structure with regard to the
1972]
Bartell and Rickman — Orthophosphate Uptake
157
PARTICLE DIMETER
oo
Figure 4. Size distributions for L. Winnebago samples (a) collected on
February 2, (b) collected February 12, and (c) collected on February 20,
1971 as determined by the Coulter Counter (100 aperture).
previous sample (Figure 5-a). The appearance of two new peaks
may illustrate the growth of new populations as the physical and
chemical parameters of the lake change with the spring thaw.
Figure 1 shows that the phosphate concentration has substantially
increased by this time. The evident blending and reduction of these
158 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
PARTICLE DIAMETER
(/O
Figure 5. Size distribution for L. Winnebago samples collected (a) on March
1, and (b) on March 13, 1971 (100 n- aperture).
peaks (Figure 5-b) by March 13 might suggest that they were
transient species with a very narrow tolerance range for the
nutrient conditions in the lake during March. This is a common
occurrence as community structure may change significantly in
natural phytoplankton assemblages within a week’s time (Schelske,
Callender, and Stoermer, 1969).
1972] BarteU and Rickman— Orthophosphate Uptake 159
The final distribution (Figure 6) emphasizes the increase in bio¬
volume in the sample collected on April 2, 1971, By this time, phos¬
phorous had reached the highest concentration measured during
this study (Figure 1),
PARTICLE DIMETER
CM)
Figure 6, Size distribution for L. Winnebago sample collected on April 2,
1971 (100 fi aperture).
160 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
C. Phosphate Uptake Kinetics
The enrichment experiments performed with the samples col¬
lected on February 2 and February 20, 1971 yielded rates of bio¬
volume increase, primary production, and chlorophyll production
relative to orthophosphate concentration that approximated the
model proposed by Dugdale (1967) and Maclsaac and Dugdale
(1969) for the kinetics of nutrient uptake by phytoplankton. This
model is derived from the Michaelis-Menten equation for the effect
of substrate concentration on the rate of an enzyme catalyzed reac¬
tion as expressed by:
__ vmaxs
“ Km + S
(1)
where,
v = rate of catalyzed reaction
Vmax = maximum rate of reaction
S = substrate concentration
Km = substration concentration at which ^Vmax is calculated
(Lehninger, 1970).
This equation describes the curve in Figure 7-a. Two modifications
of the Michaelis-Menten equation have been derived in order to
more accurately calculate Vmax and Km. By taking the reciprocal of
both sides of ( 1 ) and simplifying, a linear function results :
l__Kin 1 _JL
v Vmax ’ S + Vmi
(2)
This is the Lineweaver-Burk equation, which is graphically repre¬
sented in Figure 7-b.
The Eadie-Hofstee modification is derived by multiplying both
sides of (2) by Vmax(v) and rearranging to yield:
V = • -g- + V max (3)
which is illustrated in Figure 7-c. These modifications will be of
importance in the application of the Michaelis-Menten model to the
measured uptake kinetics of phosphate in this study.
The units of variables in the Michaelis-Menten model are slightly
different in application to the study of nutrient uptake. In this
paper,
v = uptake velocity measured as rate of
increase in biovolume, cubic microns • ml-1 • hr-1
primary productivity, mg Carbon • m-3 • hr-1
chlorophyll production, mg Chlor. • m-3 • hr-1
Vlnax = maximum value of above rates
S — fig • liter-1 orthophosphate
Km = concentration of o-P04 where %Vmax is realized.
1972] Bartell and Richman— Orthophosphate Uptake
161
C
Eadie-Hofstee Plot
Figure 7-a. Michaelis-Menten curve. Ymax = maximum reaction rate. Km =
substrate concentration where reaction rate is one half of maximum velocity.
7- h. The Lineweaver-Burk linear modification of the Michaelis-Menten curve.
7— c. The Eadie-Hofstee linear modification of the Michaelis-Menten curve.
(Lehningerj 1970).
162 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Rather than use the designation Km, Kt or “transport constant”
will be used to emphasize that a mathematical and not a biochem¬
ical equivalence of the Michaelis-Menten model is being used. This
follows the notation developed by Wright and Hobbie, as cited by
Dugdale (1967). However, if one considers an enzyme mediated
mechanism for nutrient uptake, a biochemical equivalence may be
inferred from this particular application of the model. From this
brief description of the model, one can analyze the rates of bio¬
volume increase, primary production, and chlorophyll synthesis as
a function of increasing phosphate concentrations in the Lake Win¬
nebago samples collected during the month of February.
In order to compare the increase in biovolume of the enriched
samples to that of the control for any given experiment, size dis¬
tributions were taken at both the beginning and end of the experi¬
ment. This is necessary because the control sample contains the
concentration of orthophosphate innate to Lake Winnebago at the
time of sample collection. When incubated under conditions of light
saturation and 15 C, the population utilizes the available phosphate
supply. Figure 8 shows a typical growth pattern of a control sample
that has been incubated under the described conditions. The meas¬
ured rate of increase could then be plotted as a function of the
orthophosphate present in the Lake Winnebago sample and com¬
pared with the rates of biovolume increase of the enriched samples.
Clearly not all of the measured biovolume of any given size dis¬
tribution consists of living algal cells. Sheldon and Parsons (1967)
have determined a method for calculating the amount of detritus in
a sample that takes advantage of the assumption that the growth
rate of detritus is zero. The total volume of the initial size distribu¬
tion for a January 9, 1971 sample was 11.2 x 105 p3 • ml-1. The
amount of detritus measured in this sample was 5.72 x 105 p3 • ml-1,
which is 51% of the initial total volume. However, at the end of
the enrichment experiment, the measured biovolume was 320 x 105
p3 . ml-1. Assuming a growth rate of zero, the previously measured
amount of detritus is only 1.5% of the total biovolume. This seems
to indicate that in the enrichment experiments the amount of
detritus in the final analysis of samples does not contribute any
significant source of error in calculating rates of increase. The
same technique revealed a detritus level of 1.8 x 106 p3 • ml-1 in the
April 2, 1971 sample. However, this is still only 51% of the initial
biovolume of the sample, 3.5 x 106 p3 • ml-1. This might tend to
indicate that the amount of detritus present in Lake Winnebago is
directly proportional to the phytoplankton population size. How¬
ever, much more data is needed to validate this hypothesis.
The use of biovolume to study the kinetics of phosphate uptake
provides evidence that there is no appreciable change in species
1972] Bartell and Rickman— Orthophosphate Uptake
163
PARTICLE DIAMETER
c/o
Figure 8* Increase in biovolume of the control sample during* period of
incubation determined by Coulter Counter (100 m aperture). Sample collected
Nov. 24, 1970.
composition in the samples as a result of enrichment. Figure 9
illustrates this. The sample, collected February 2, 1971, was en¬
riched to phosphate concentrations ranging from 50 ug • liter-1 to
200 ixg • liter-1. The shape of the distribution remained very similar
during the period of incubation.
The use of biovolume increase as a function of phosphate enrich¬
ment provided meaningful data for application to Miehaelis . Men-
ten kinetics in only one experiment. This was an enrichment experi-
164 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
PARTICLE DIAMETER
CM )
Figure 9. Increase in biovolume at various particle diameters as a function
of orthophosphate enrichment. Lowest solid line i— control. Dashed line = 50
ppb 0-PO4 addition. Middle solid line = 100 ppb o-POi addition. Upper solid
line == 150 ppb 0-PO4 addition. Dash-dot line = 200 ppb o-PO* addition.
ment performed with a sample collected on February 20, 1971.
Figure 10 illustrates the increase in biovolume • ml 1 • hr-1)
for particles arbitrarily categorized into three size ranges accord¬
ing to particle diameter. The curve for the circled dots shows that
the greatest increase in growth rate as a function of phosphate
addition occurred for particles in the size range of 5.9 to 14.9
microns. This is interesting in view of the initial size distribution
of this sample (Figure 4-c). The growth rates of the smaller cells
(3.0-4.7 n), though considerably lower, also suggest an approxima¬
tion to Michaelis-Menten kinetics. The growth rates for the larger
1972]
Bartell and Richman — Orthophosphate Uptake
165
(>ug* liter-1)
Figure 10. Rate of biovolume increase as a function of increasing o-POi
concentration for different diameter particles. Circled dots =: 5.9 — 14.9 v
diameter. Solid dots = 3.0 — 4.7 diameter. Open circles = 18.8 — 29.9
/a diameter. From enrichment experiment with February 20, 1971 sample.
166 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
particles (18.8-29.9 y) appear to approximate the curve for the
smaller particle growth rates. If one assumes that at zero phos¬
phate there is no increase in biovolume, the data suggests a lag
period in population growth as a function of phosphate enrichment.
The validity and significance of this assumption will be discussed
later.
Figure 11 illustrates the rates of carbon fixation and production
of chlorophyll a and b as a function of increasing phosphate con¬
centrations in an enrichment experiment with a sample collected
February 2, 1971. The data seems to demonstrate that phosphate
is being taken up in a Michaelis-Menten relationship. To determine
the validity of fitting the data to this particular curve, the data was
subjected to the Lineweaver-Burk and Eadie-Hofstee modifica-
100 200
(^g. liter"1)
Figure 11. Rates of carbon fixation and production of chlorophyll a and b as
a function of increasing concentrations of orthophosphate. L. Winnebago
sample collected February 2, 1971. A — mg Carbon • m~3 • hr-1. Solid dots =
chlorophylls. Open circles := chlorophylli*
1972] Bartell and Richman — Orthophosphate Uptake 167
tions. The values of Vmax, Kt, and a linear regression correlation
coefficient could then be calculated. These values are presented in
Table 1. As might be expected, the values calculated by both mod¬
ifications were highly comparable for any given sample as seen in
Table 1. For convenience only the Lineweaver-Burk modifications
are illustrated in this paper. Figure 12 is the application of this
modification to the data presented in Figure 11. The values of r for
primary production, chlorophyll^ and chlorophyll production, are
respectively, .97, .99, and .97. These values suggest the fit of the
data to the Michaelis-Menten curve is valid. All lines were deter¬
mined by least squares analysis on the RAX IBM 360/44. The con¬
trast test mentioned earlier shows that rates of chlorophyll produc-
Table 1. Comparison of Vmax and Kt Calculated by Lineweaver-Burk
AND EADIE-HOFSTEE MODIFICATIONS OF MICHAELIS-MENTEN EQUATION
r— - - - - - _ _ _ _ _ a
b
c
a = primary production rates
b - rates of chlorophylla synthesis
c = rates of chlorophyll^ synthesis
168 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 12. Application of the Lineweaver-Burk modification to the data
presented in Fig. 11. For carbon fixation (triangles), Vmax == 129.9 mg
Carbon • nr3 • hr-1. Kt = 25.7 vg • liter-1 o-PCh, r t= .97. For chlorophylls,
Vmax i= .022 mg chlor. • m-3 • hr-1. Kt c= 19.7 rg • liter-1 o-P04, r s= .99
(solid dots). For chlorophyll (open circles), Vmax = .023 mg chlor. • m-3
• hr-1. Kt i= 22.5 vg • liter-1 o-P04, r ■= .97.
tion at the steep part of the curve are significantly different from
those points along the plateau of the curve; however, as expected,
the rates along the plateau are not significantly different from one
another (P = .05) .
The rates of carbon fixation and chlorophyll synthesis deter¬
mined from the enrichment of a sample collected February 20,
1971 again demonstrate Michaelis-Menten phosphate uptake as
illustrated in Figure 18. The values of Vmax and Kt for both primary
production and chlorophyll synthesis rates are presented in Table
1. At this time there is no obvious reason for the differences in
Vmax and Kt calculated at these two dates for these parameters.
1972] Bartell and Richman — Orthophosphate Uptake 169
However, the values of Vmax for production of chlorophyll are
quite comparable, .023 and .024 mg Chlor. • m-3 • hr-1, as calcu¬
lated by the Lineweaver-Burk modification. To determine the fit of
these rates to the Michaelis-Menten curve, the data was again sub¬
jected to least squares analysis. The value of r = .51 for carbon
fixation rate as a function of phosphate enrichment. In the deter-
(ju g. liter"1')
Figure 13. Rates of carbon fixation and chlorophyll a and b production as a
function of increasing concentrations of orthophosphate. L. Winnebago sample
collected February 20, 1971. A c= mg Carbon • nr3 • hr-1. Solid dots =
chlorophyll*. Open circles = chlorophylls
mg Carbon •
170 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Figure 14. Application of the Lineweaver-Burk modification to the data
presented in Fig. 13. For carbon fixation, (triangles), Vmax = 25.6 mg
Carbon • nr3 • hr1-. Kt = 16.2 vg • liter-1, r = .51. For chlorophyll
omitting point at — ,= .029 (solid dots), Vmax — .007 mg chlor. • m-3
• hr-1. Kt = 39.4 i^g • liter-1, r == .98. For chlorophyll (open circles),
Vmax = .024 mg chlor. • m-3 • hr-1. Kt = 195.0 vg ■ liter-1 o-POt, r t== .99.
mination of r for the rate of chlorophyll synthesis as a function of
phosphate addition, the value of— for the point at TJtr. was omitted
V Jr (J4
because of its extreme variation from the other points which other¬
wise suggest a strong linear correlation, r = .98. For the rate of
chlorophyll,, production, r = .99. Again the M ichaelis-Menten rela¬
tionship seems valid as a model for phosphate uptake.
Rates of primary productivity in enrichment experiments per¬
formed with samples collected in November, 1970 indicated that
1972] Bartell and Rickman — Orthophosphate Uptake 171
orthophosphate was not a limiting factor for growth. The rate of
carbon fixation for the control in an experiment with a November
18 sample was 17.8 mg Carbon • m-3 • hr-1. In a 50 jug • liter-1
phosphate enriched sample for this experiment, the rate was only
14.0 mg Carbon • m-3 • hr-1. A t-test for small sample sizes (Hoel,
1967) showed that these rates are not statistically different at the
.05 level of predictability. The addition of 1.0 and 2.0 mg • liter-1
equivalents of orthophosphate to a sample collected on November
9, 1970 yielded similar results. The rate of productivity in the con¬
trol sample was 46.0 mg Carbon • m-3 • hr-1. The rates for the
enriched samples were 30.7 and 28.8 mg Carbon • m-3 • hr-1,
respectively. Because these rates are not sample means, the sig¬
nificance of the decrease cannot be tested. However, the data shows
that there was no increase in productivity with the addition of
phosphate.
Similarly, productivity rates for a sample collected on April 2,
1971 indicated that phosphate was not limiting in the spring. The
rate of productivity for the control was 154 mg Carbon • m-3 • hr-1,
while the rate for the most highly enriched sample of the series
was only 158 mg Carbon • m-3 • hr-1. These rates are not signif¬
icantly different at the .05 level. The rates of productivity for a
sample collected April 18 and enriched with 20, 40, and 60 ^g •
liter-1 orthophosphate are 21.3 mg Carbon • m-3 • hr-1 for the
control and 21.5, 28.3, and 28.5 mg Carbon • m-3 • hr-1 for the
enriched samples. The significance of the increase is questionable,
however the values are not means so they cannot be tested. Phos¬
phate is probably no longer limiting; however, this statement is
made with some reservation due to the absence of statistical
verification.
No chlorophyll production rates were measured during late
autumn when orthophosphate was present in high concentrations.
However, the rate of chlorophyll synthesis was found to decrease
somewhat with increasing additions of phosphate to a Lake Win¬
nebago sample collected April 2, 1971, when orthophosphate was
measured as 105.0 jig • liter-1 (Figure 15). Such inhibition also
occurred in the enrichment of a sample collected on April 18, 1971.
This seasonal pattern of nutrient uptake as measured by increased
chlorophyll synthesis rate seems to parallel that of carbon fixation.
V. Discussion
The results of the enrichment experiments indicate that the use
of biological parameters to indirectly measure the kinetics of phos¬
phate uptake is a valid approach to nutrient limitation theory and
its relation to natural phytoplankton populations. Primary produc¬
tivity and chlorophyll production seem particularly suited for this
172 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
(/* g • liter"1 )
Figure 15. Mean chlorophyll^ + » production rates as a function of increasing
orthophosphate concentrations in an enrichment experiment with a sample
collected April 2, 1971.
type of study. However, care must be taken in the use of these
parameters.
Results obtained by means of the 14C technique for measuring
carbon fixation are subject to uncertainty in interpretation. The
addition of phosphate may influence the photosynthetic apparatus
specifically. In other words, a certain concentration of the nutrient
may specifically alter the physiological efficiency of photosynthesis
just as other metabolic processes are regulated by pH, temperature,
or other similar environmental factors. Once the nutrient is depleted
beyond the critical concentration, the productivity of the system
will return to its level measured prior to enrichment. The effect of
enrichment is short lived. However, this type of error is normally
associated with the effect of micronutrients, according to Fogg in
1965, cited by Dugdale (1967).
Because phosphate can increase productivity as these enrichment
experiments illustrate, any compound that displays a rapid syn¬
thesis in response to slight increases in the level of productivity
1972] Bartell and Rickman — Orthophosphate Uptake 173
would be a sensitive parameter for potentially measuring the uti¬
lization of phosphorous in the system. Chlorophyll is such a com¬
pound (Margalef, 1968). This sensitivity in relation to phosphate
enrichment as shown in Figures 11 and 13 support the use of this
parameter for measuring phosphate uptake quite well. It must be
remembered, however, that the interpretation of this induced
chlorophyll increase in relation to growth includes not only
increased cell numbers, but also the synthesis of maximum chloro¬
phyll content per cell prior to division.
The use of biovolume increase as a growth parameter must also
be carefully interpreted. Increase in cell numbers or biovolume
results from phosphate induced productivity only after the cellular
content of chlorophyll and carbon have been maximized. This may
explain the lag in the increase in growth rate as illustrated in Fig¬
ure 10. When intermediary metabolism in the cell has achieved its
optimum level or maximum efficiency, any remaining potential
energy from the phosphate induced increase in productivity may
be channeled into increasing the population size, as measured by
increased biovolume. This makes the use of biovolume increase to
estimate uptake kinetics less sensitive than the measurement of
chlorophyll synthesis or the direct measurement of primary pro¬
ductivity by the 14 C method.
Regardless of the approach taken to elucidate the kinetics of
nutrient uptake, there is one limitation to the experimental design
when dealing with natural populations. This limitation is equally
applicable to this study or to the direct measurement of uptake by
the use of labelled nutrients as performed by Dugdale (1967) and
Maclsaac and Dugdale (1969). In both studies, a lower limit to
the amount of possible enrichment is determined by the concentra¬
tion of the nutrient in the sample when collected. As enrichment
implies addition, no calculation of uptake could be measured for
concentrations of the nutrient below that already present in the
sample. If the rate of nutrient uptake by the plankton was already
at the plateau of a Michaelis-Menten curve, further additions of
the nutrient would fail to yield the necessary information to calcu¬
late values of Vmax or Kt. Uptake rates of enriched samples in this
case would simply approximate the plateau or show inhibition.
Maclsaac and Dugdale (1969) found this to be true in their study
of nitrate uptake kinetics by marine phytoplankton. The plateau
approximations and inhibition measured at high phosphate con¬
centrations in the fall and spring by both chlorophyll production
and primary production indicate a similar situation may exist for
populations in Lake Winnebago in regard to phosphate uptake.
The kinetics in the fall and spring populations may be still
174 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Michaelis-Menten in nature; however, the values of Vmax and Kt
cannot be determined with the particular experimental design. In
this study, Michaelis-Menten kinetics were best described by all
parameters in February when the phosphate concentrations were
at or near their lowest recorded values (Figure 1).
An alternate design might involve culturing different algal
species in low nutrient mediums. These species might then be sub¬
jected to enrichments to determine their particular values of Vmax
and Kt for various nutrients. Unfortunately, it is difficult to extra¬
polate results obtained in the laboratory to the behavior of the
organisms in a dynamic ecosystem. Values of Vmax and Kt calcu¬
lated under these conditions may be entirely different from those
values for a species in its natural environment. Furthermore, Ham¬
ilton (1969), working with species of algae collected from Cayuga
Lake in New York in controlled culture mediums, found the effects
of phosphate enrichment (5.0 ^g • liter-1 KH2P04) unclear as
determined by productivity measurements with 14C. In some cases,
this small concentration was even found to limit productivity. Cer¬
tainly more work needs to be done in this area of research.
The ecological significance of the Michaelis-Menten model for
nutrient uptake provides insight into the dynamics of nutrient
limitation. While the application of the model to natural popula¬
tions assumes that a single species of phytoplankton or a group of
species with similar uptake kinetics dominates the population, this
is usually not the case in most systems. Dugdale (1967) has calcu¬
lated values of Vmax for several species of algae and found them
to range from .268 • hr-1 for Skeletonema costatum (Grev.) to
.034 • hr-1 for Rhizoselenia alata Bright. These values are for
nitrate uptake. Eppley (1969) has determined Kt values for seven¬
teen oceanic and neritic species for nitrate and ammonium uptake.
His results range from 8.6 ^ moles • liter-1 to 0.1 /x moles • liter-1.
Unfortunately, the use of biological parameters does not allow the
author to compare uptake rates or Kt values with those of Eppley
and Dugdale. Furthermore, this study only involved phosphate.
Nitrate kinetics were not investigated.
Returning to Dugdale’s work, if the species with the lowest value
of Vmax, R. alata , also exhibits a lower Kt value than S. costatum ,
then R. alata would be able to maintain itself in lower nitrate con¬
centrations than S. costatum. Skeletonema would utilize nitrate
faster and consequently approach concentrations where it could no
longer compete with R. alata. This effect would be greatly amplified
in time if the nitrate concentration was being depleted by other
species of plankton as well. In general, species with lower values
of Vmax and Kt would successively dominate the algal community
as the concentration of the limiting nutrient decreased through
1972] Bartell and Rickman— Orthophosphate Uptake 175
time. These species would be evolutionarily selected for where com¬
petition for low available nutrient concentrations existed (i.e. the
Sargasso Sea, Dugdale, 1967). The values of Vmax and Kt found in
this study represent only one predominant species of phytoplank¬
ton, most likely a species of Agterionella. Therefore, no compari¬
sons can be made at this time with other Lake Winnebago species
to test the significance of Vmax and Kt in relation to phytoplankton
succession. However, it is interesting to note that the data illus¬
trated in Figure 15 indicates a greater probable value of Vmax for
the April 2, 1971 population than those that were calculated for
the February experiments in terms of chorophyll synthesis rates.
The measured phosphate concentration more than doubled during
this time period (Figure 1). More data is needed to pursue this
aspect of .succession in the Lake Winnebago phytoplankton popu¬
lations. The important concept that emerges from the above data
in terms of Michaelis-Menten kinetics is that nutrient limitation
is a dynamic process. Not only the available nutrient concentration
for a given element determines limitation, but the rate in which
this nutrient is utilized by the different species in the community,
as well as the rate in which the nutrient is supplied to the system
are equally important in describing limiting conditions.
Furthermore, the Michaelis-Menten model as applied to nutrient
limitation theory may provide valuable insight into the predictabil¬
ity of dynamic interactions between phytoplankton and nutrients
in aquatic systems. If Vmas and Kt values were determined for
many species in relation to the different macro and micronutrients,
a systems analysis approach with the analog computer might be
of great importance in determining the effects of varying nutrient
compositions on the productivity, species diversity, and abundance
of phytoplankton in both oligotrophic and eutrophic systems. This
predictive power would be of invaluable assistance in proper plan¬
ning for the location of cities, industry, and waste treatment plants,
as well as in calculating the permissible amount of nutrient load¬
ing for any system over time in order to minimize the effect of
urban, industrial, and agricultural activities on the natural rate
of eutrophication. Parker (1968) has been successful in integrating
some field investigations with laboratory information to calculate
the effect of increasing organic phosphate pollution of Kootenay
Lake, British Columbia, on populations of phytoplankton. By means
of a digital computer and a program based upon biological con¬
stants that approached reality for such parameters as nutrient con¬
centration, photoperiod, and temperature, Parker was quite suc¬
cessful in predicting the occurrence of algae “blooms” as a result
of the inflow of phosphate from a fertilizer plant located on the
Kootenay River, which flows into the lake.
176 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Smith (1969) has used the Michaelis-Menten expression to fol¬
low the effect of increased nutrient supply on a phytoplankton, zoo¬
plankton, predator food chain. He also investigated the situation in
which a secondary predator was added to the simulation. In the
first situation, Smith’s model shows the effect of nutrient increase
to be reflected in an increase in the population of the phytoplank¬
ton. Interestingly, there is also an increase in the density of the
predator population. In the second situation, which includes the sec¬
ondary predator, the system responds quite differently. The in¬
crease in nutrient concentration remains mostly in free form, while
there is a small increase in the zooplankton and secondary predator
populations. While such models are very simple in their ecological
approach, they are a vital step in the direction of understanding
the structural and functional dynamics of the trophic nature of
the aquatic ecosystem. Smith’s model indicates the subtle effects
that one trophic level has upon another in relation to nutrient
enrichment. Clearly, much more work of this nature, especially a
further elucidation of nutrient limitation dynamics in the form of
Michaelis-Menten theory, is vital in man’s understanding of the
process of eutrophication.
Another interesting aspect of the nutrient enrichment study
described in this paper pertains specifically to Lake Winnebago.
While the measured value of soluble phosphorous has increased
manifold from the 2.0 ^g • liter-1 annual mean reported by Hil-
senhoff for the 1961-1964 period to the 29.0 to 105.0 /zg • liter-1
range of concentrations determined over the course of this study,
the enrichment experiments performed in February, 1971 demon¬
strated rather substantially that soluble phosphorous can still be a
productivity limiting factor in this aquatic system. Lake Winne¬
bago appears to have not yet reached a state of year round phos¬
phorous saturation. Any concerned effort to effectively limit the
allochthanous input of phosphorous into this system would be of
significance in reversing the culturally induced acceleration of the
eutrophication processes. Strict regulation of urban, industrial,
and agricultural phosphorous loading would be proportionally
repaid by an increase in water quality with its associated biological,
chemical, and cultural implications.
Acknowledgement
Appreciation must be expressed to Mr. Peter Becker for his
invaluable assistance in the design of the CHLOREG program.
The data for the calculation of chlorophyll synthesis rates for
the April 2, 1971 sample as illustrated in Figure 15 was provided
by the members of the Experimental Ecology class at Lawrence
University.
1972] Bar tell and Rickman — Orthophosphate Uptake
177
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Bachman, R. W. 1959. On the use of radioactive tracers in limnology,
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Biggar, J. W., and R. B. Corey. 1969. Agricultural drainage and eutrophica¬
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Dixon, W. J., and F. J. Massey. 1969. Introduction to statistical analysis,
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Dugdale, R. C. 1967. Nutrient limitation in the sea: dynamics, identification,
and significance. Limnol. Oceanog., 12: 685-695.
Eppley, R. W., J. N. Rogers, and J. J. McCarthy. 1969. Half-saturation
constants for uptake of nitrate and ammonium by marine phytoplankton.
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Goldman, C. R. 1965. Micronutrient limiting factors and their detection in
natural phytoplankton populations, p. 121-135. In C. R. Goldman ed.,
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18 Suppl., University of California Press, Berkeley.
Hamilton, D. H. 1969. Nutrient limitation of summer phytoplankton growth
in Cayuga Lake. Limnol. Oceanog., 14: 579-590.
Hassenteufel, W. R., R. Jagitsch, and F. F. Koczy. 1963. Impregnation of
glass surface against sorption of phosphate traces. Limnol. Oceanog.,
8: 152-156.
Hilsenhoff, W. L. 1967. Ecology and population dynamics of Chironomus
plumosus (Diptera: Chironomidae) in Lake Winnebago, Wisconsin. Ann.
Entomol. Soc. Amer. 60: 1183-1194.
Hoel, P. G. 1967. Elementary statistics. John Wiley & Sons, Inc., New York,
p. 348.
Lee, G. F. 1969. Analytic chemistry of plant nutrients, p. 646-658. In NAS
Symposium, Eutrophication: causes, consequences, correctives. Washing¬
ton, D. C.
Lehninger, A. L. 1970. Biochemistry. Worth Publishers, Inc. New York.
p. 811.
Lueschow, L. A., J. M. Helm, D. R. Winter, and G. W. Karl. 1970. Trophic
nature of selected Wisconsin lakes. Trans. Wis. Acad. Sci. Arts Lett.,
58: 237-264.
MacIsaac, J. J. and R. C. Dugdale. 1969. The kinetics of nitrate and ammonia
uptake by natural populations of marine phytoplankton. Deep-Sea Re¬
search., 16: 45-57.
Margalef, R. 1968. Perspectives in ecological theory. University of Chicago
Press, Chicago, p. 111.
Murphy, J., and J. P. Riley. 1962. A modified single solution for the deter¬
mination of phosphate in natural waters. Anal. Chim. Acta., 27 : 31-36.
Parker, R. A. 1968. Simulation of an aquatic ecosytem. Biometrics., 24:
803-821.
Parsons, T. R., and J. D. Strickland. 1963. Discussion of spectrophotometric
determination of marine-plant pigments, with revised equations for
ascertaining chlorophylls and carotenoids. J. Marine Res., 21 : 155-163.
Provasoli, L. 1969. Algal nutrition and eutrophication, p. 574-593. In NAS
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ton, D. C.
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Ryther, J. H., and W. M. Dunstan. 1970. Nitrogen, phosphorous, and
eutrophication in the coastal environment. Science., 171: 1008-1013,
Schelske, C. L. 1962. Iron, organic matter, and other factors limiting primary
productivity in a marl lake. Science, 136: 45-46.
Schelske, C. L., E. Callender, and E. F. Stoermer. 1969. Nutrient enrich¬
ment experiments on phytoplankton populations in Lake Michigan. (32nd
annual meeting of the American Soc. of Limnol. and Oeeanog., Scripps
Institution of Oceanography, University of Cal., San Diego).
Sheldon, R. W., and T. R. Parsons. 1967. A practical manual on the use
of the Coulter Counter in marine research. Coulter Electronic Sales
Company, Toronto, Canada. 66 p.
Sloey, W. E. 1970. The limnology of hypereutrophic Lake Butte des Morts,
Wisconsin. Proc. 13th Conf. Great Lakes Res. 1970: 951-968. Internat.
Assoc. Great Lakes Res.
Smith, F. E. 1969. Effects of enrichment in mathematical models, p. 631-645.
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Washington, D. C.
Strickland, J. D., and T. R. Parsons. 1965. A manual of sea water analysis.
Bull. Fisheries Res. Board Can. No. 125. 203 p.
Tucker, A. 1957. The relation of phytoplankton periodicity to the nature of
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Vollenweider, R. A., (Ed.). 1969. A manual on methods for measuring
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F. A. Davis, Co., Philadelphia 221 p.
Wentz, D. A., and G. F. Lee. 1969. Sedimentary phosphorous in lake cores—
analytical procedure. Environ. Sci. Tech., 8: 750-759.
A RECORD OF THE FRESHWATER NEMERTEAN,
PROSTOMA RUB RUM, IN WISCONSIN
Robert F. Browning
Prostoma rubrum is the only freshwater species of the acoelo-
mate phylum., Nemertea (Rhynchocoela), which is known to occur
in North America. Although Coe (1943) implies they are widely
distributed throughout the United States, the only actual sites of.
occurrence he mentions are some freshwater ponds in the Woods
Hole area. Poluhowich (1968) reviewed various aspects of the biol¬
ogy of P. rubrum in an attempt to stimulate new collections of
this worm and extend the distribution records. In his review he
provides references to their occurrence in the Chicago area and
in Pennsylvania. He also reported that he collected numerous spec¬
imens from a small brook in Stratford, Connecticut. In a recent
letter to me, he indicated that he had received word of a few ad¬
ditional reports in response to his publication but not enough to
substantiate Coe's claim of nation-wide distribution. The present
report appears to be the first record of P. rubrum in Wisconsin.
A population of P. rubrum was discovered in Silver Creek, in
the city of Ripon (Fond du Lac County) from samples brought
into the laboratory from a class field trip in early September 1970.
The first specimen was isolated by two students, Joan Strewler
and Kathleen Spence, and from its general morphology, I tenta¬
tively identified it as P. rubrum . The identification was confirmed
later by observing the involvement of its probocsis in feeding and
the deposition of eggs in peculiar mucous tubes secreted by sexu¬
ally mature worms.
Using the collection method devised by Polohouwich (1968), I
isolated as many as 35-40 specimens from a liter of substrate on
several occasions during the Fall of 1970. The most productive
substrate consisted of a mixture of mud detritus, and filamentous
algae. In mid-February only two specimens were obtained from
substrate samples obtained under snow and ice cover. In early
April I found no specimens but this was probably due to the ex¬
tended period of high water associated with the spring thaw.
Although the high water undoubtedly flushed a great deal of the
preferred substrate downstream, it is likely that many of the
worms migrated deeper in the stream bed. I have not investigated
this possibility fully at the present time.
179
180 Wisconsin Academy of Sciences, Arts and Letters [VoL 60
The first specimens were taken approximately 100 yards down¬
stream from the spillway of Gothic Mill Pond, an impoundment
in a city park. A student, Mydin Shariff, made a preliminary semi-
quantitative study of their distribution in Silver Creek which sug¬
gested that the population was confined to that portion of the
stream from the pond overflow, downstream to the vicinity of the
sewage treatment plant, a distance of a little over one mile. This
work will be repeated quantitatively this summer and fall to pin¬
point the linear distribution of this species in Silver Creek. Such
a study may answer some very interesting questions concerning
the physical and biotic factors involved in the dispersal of this
species. According to Coe (1943), aThe species presumably was
carried to the western states with cultivated water plants.”
Thus far, attempts to establish laboratory cultures have been
unsuccessful although I have maintained individual specimens for
up to two months by feeding them a small length of tubifex worm
each week.
References Cited
Coe, W. R. 1943. Biology of the nemerteans of the Atlantic Coast of North
America. Trans. Conn.Acad. Arts and Sci. 35: 147-327.
Poluhowich, J. J. 1968. Notes on the freshwater nemertean Prostoma ruhrum .
Turtox News. 46 (1) : 2-7.
A RECORD OF GRASPED ACUST A SOWERBYI
IN WISCONSIN1
Richard P. Howmiller and G. M. Ludwig
Abstract
The freshwater jellyfish, Craspedacusta sowerbyi, is reported for
the first time from Wisconsin. Large numbers of medusae occurred
in a small artificial pond in late summer of 1969 and 1970. This
appears to be the northernmost locality for the species in the
Mississippi drainage.
The known North American distribution of the freshwater jelly¬
fish, Craspedacusta sowerbyi Lankester, is broad but appears to be
centered in the northeastern quarter of the United States. There
seem to be no previous records of this species as far north in the
Mississippi drainage as Wisconsin (Pennak 1957, Lytle 1960, Bush-
nell and Porter 1967). As pointed out by Pennak (1957) the lack
of reported occurrences in this region are remarkable because of
the considerable limnological knowledge of Wisconsin and
Minnesota.
A medusa brought to the Milwaukee Public Museum for iden¬
tification directed our attention to a population of C. sowerbyi in
a small pond on Skillet Creek Farm (N y2 Sec 15, T 11 N, R 6 E)
near Baraboo, Sauk County, Wisconsin. The pond is approximately
50 m in diameter, dug in glacial till and fed by water from Skillet
Creek. Dug six years ago, the pond still appears “new” with a bot¬
tom of sand and gravel and no colonization by aquatic macrophytes.
Medusae of C. sowerbyi have been noticed by the owner in the fall
of 1969 and 1970. When we visited the pond on September 13, 1970,
large medusae (1.0-1. 5 cm) were abundant; we captured more than
50 with small hand nets in an hour. Some medusae swam to the
surface of the water and floated downward again. Others lay,
pulsing slowly, on the bottom in patches of filamentous algae
( Spiro gyra) and detritus. Crayfish, amphipods, ostracods, chiro-
nomid larvae, mayfly nymphs and the gastropods, Campeloma and
Ferrissia, were common in the algae and detritus. Polyps of C.
sowerbyi were not observed.
1 Contribution No. 43, Center for Great Lakes Studies, University of Wisconsin-
Milwaukee, Milwaukee, Wisconsin 53201
181
182 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Surface water temperature at mid-day on September 18, 1970,
was 18 C. Medusae were common in the pond during the previous
two or three weeks, according to the property owner.
Many other records of C. sowerbyi, perhaps a majority, are from
artificial ponds or impoundments (Pennak 1957 ; Lytle 1960, Bush-
nell and Porter 1967). It has been suggested that long range dis¬
persal may have occurred in conjunction with shipments of aquatic
plans (Bushnell and Porter 1967). The owner of Skillet Creek
Farm stated that no aquatic plants and no animals other than
locally obtained bass, bluegills and crappies had been introduced
to the pond. Craspedacusta must have been introduced by some
agent other than man. Perhaps wood ducks, which frequent the
pond, carried in Craspedacusta polyps among their wet feathers.
References
Bushnell, J. H. and T. W. Porter. 1967. The occurrence, habitat, and prey
of Craspedacusta sowerbyi (particularly polyp stage) in Michigan.
Trans. Amer. Microsc. Soc. 86: 22-27.
Lytle, C. F. 1960. A note on distribution patterns in Craspedacusta. Trans.
Amer. Microsc. Soc. 79: 461-469.
Pennak, R. W. 1956. The fresh-water jellyfish Craspedacusta in Colorado with
some remarks on its ecology and morphological degeneration. Trans.
Amer. Microsc. Soc. 75: 324-331.
PEDOLOGY OF THE TWO CREEKS SECTION,
MANITOWOC COUNTY, WISCONSIN
Gerhard B. Lee and M. E. Horn
Introduction
The Two Creeks Forest Bed of eastern Wisconsin is a well known
marker of the interval of deglaciation between the Cary and Vald-
ers glacial advances. Frye and Willman (1960) have classified this
interval as the Two Creekan, a substage of the Wisconsinian that
occurred from about 12,500 to 11,000 radiocarbon years ago. Ac¬
cording to their classification, the Two Creekan followed a much
longer, earlier substage, the Woodfordian, and preceded the Vald-
eran, or youngest substage, recently described by Black (1966).
The Two Creeks horizon, near the village of Two Creeks, Wis¬
consin, consists of a thin paleosol, in which long-dead trees and
other plants are rooted, and the overlying, broken over remains
of the Two Creeks forest. Both are buried beneath younger drift.
At the surface a modern soil has been formed. While the age of
the buried horizon, its paleobiological nature and geologic history
have been the subject of considerable study, little has been written
about the site from a pedological point of view. The present paper
is a report of such a study.
Location op Site
The original Two Creeks site, first reported by Goldthwaite
(1907), was located along the west shore of Lake Michigan in
Manitowoc County, Wisconsin (Sec. 11, T21N, R24E). Other ex¬
posures in the same general area were subsequently studied by
Wilson (1932), (1936) , Thwaites. (1943), Thwaites and Bertrand
(1957), Horn (1960), West (1961), and Hole (1967). Similar
horizons have also been reported in the Fox River Valley by Law-
son (1902) and in the Duck Creek Ridges by Piette (1963), and
Janke (1962).
The section described in this study was found in the near ver¬
tical bluffs overlooking Lake Michigan just south of the Manito¬
woc . Kewaunee County line (NE^, Sec. 2, T21N, R24E). This
exposure was 2™3 kilometers north of the one described by Gold¬
thwaite in 1907 and that studied by Wilson in 1936.
Method of Study
Field studies were made of a 4.8 meter vertical section extend¬
ing from the top of the bluff to a talud at its base. This section,
183
184 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
which included both the modern soil and the Two Creeks paleosol,
was subdivided into soil horizons or sedimentary strata on the
basis of morphologic and/or stratigraphic features (Fig. 1).
In the laboratory, gravel, mainly 15 to 20 mm in diameter, was
separated from air-dried bulk samples with a No. 10 U. S. Stand¬
ard sieve. Gravel content was expressed as percent by weight of
each bulk sample. The particle-size distribution of the <2 mm
fractions was determined by the method outlined by Day (1956) ;
a hydrometer was used to determine the percentage of silt and
clay. Carbonates were not removed prior to analysis. Organic
matter in the surface horizon (Ap) of the modern soil was de¬
stroyed by treatment with 30% H202.
Percent calcium carbonate equivalent was determined by treat¬
ing 1 gram samples with an excess of IN HC1 and then back titrat¬
ing with IN NaOH. Milliequivalents of acid neutralized were
expressed as a percentage of the me that would be neutralized by
1 g of pure CaC03. Reductant soluble iron was determined accord¬
ing to the method of Aquilera and Jackson (1953) .
Organic carbon content of the peaty surface horizon of the buried
paleosol was determined by the method described by Walkley and
Black (Jackson, 1958) ; total nitrogen by the Kjeldahl method
(Jackson, 1958). Solubility in sodium pyrophosphate (Na4P207),
as a measure of the degree of decomposition of organic matter,
was measured by placing a sample on a spot plate and thoroughly
wetting it in a saturated solution of Na4P207; a strip of filter paper
was then inserted and the resultant color of the filter paper deter¬
mined. pH readings were made on soil pastes, with a glass elec¬
trode, after the soil had been saturated with distilled water and
then allowed to equilibrate for a % hour. Samples were ashed at
600 °C for 4 hours in a muffle furnace to determine ash content.
Results and Discussion
The soil developed on the modern surface at the study site was
formed in reddish brown, clay loam glacial till of Valderan age,
and thin loamy coverings likely of glacio-fluvial or glacio-lacustrine
origin (Fig. 2). This soil had been cultivated at one time, before
erosion of the headland had caused the bluff to retreat inland to
a point where cultivation of the site was no longer possible.
The pedon described was tentatively identified as a dark surface
variant of Hortonville loam, a Glossic Hapludalf. Its description
was as follows :
Ap 0-25 cm Black (10YR 2/1 — moist) and grayish brown
(10YR 5/2 — dry) loam; moderate, medium
granular structure; friable moist; neutral to
alkaline; abrupt boundary.
1972] Lee and Horn — Pedology of Two Creeks Section
185
Geologic
Co lumn
Depth
(cm)
Recent
colluvium
Valders
Till
100 _
-ft r
0 * •' . ‘ Q 1
' O )* '
200 -
Pro-Valders
silts and sands
contain ina
wood
fragments
Two Creeks
Peat
Drift
(loamy sand
and gravel)
Lacustrine
silts and
clays
400 -
500 -
' • .(.
&
f. « :
; PSA
<5^r- —
300 - - • n
&
7Wj
f'r-sf
d<&
Si:1-
Soil Profile
horizons
0
Ap
A2
Bi
UB2t
nci
nc2
Ob
Bgb
Clb
C2b
C3b
C4b
Figure 1. Sketch of section studied showing stratigraphic and morphologic
features.
186 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 2. Modern soil in bluff overlooking
Lake Michigan at Two Creeks site. Note dark
surface horizon, structured Bt horizon, and
underlying Valders till.
Brown (10YR 5/3 — moist) and pinkish gray
(7.5YR 6/2 — dry) fine sandy loam; weak,
medium platy structure; plates break into
weak, medium subangular blocky aggregates ;
A2 25-36 cm
1972] Lee and Horn — Pedology of Two Creeks Section
187
B1 86-46 cm
IIB2t 46-71 cm
IIC1 71-102 cm
IIC2 102-224 cm
patchy, yellowish -brown (10YR 5/3 — dry)
stains on ped surfaces; friable moist; a few
pebbles; alkaline; clear boundary.
Brown (7. SYR 5/4 — moist) and light brown
(7.5YR 6/4 — dry) silt loam; moderate, fine to
medium, subangular blocky structure; pink¬
ish gray (7.5YR 7/2) coatings on ped faces;
slightly sticky wet, hard dry; alkaline; clear
boundary.
Reddish-brown (5YR 5/3-4/3 — moist) clay
loam; moderate, coarse prismatic structure;
prisms break into strong, medium angular
blocky peds; clay coatings on both verti¬
cal and horizontal faces; sticky wet, hard
dry ; a few pebbles ; alkaline ; clear boundary.
Reddish-brown (SYR 5/3— moist) loam to
clay loam; moderate, coarse prismatic struc¬
ture; prisms break into moderate, medium,
angular blocky peds; sticky wet, hard dry; a
few, mainly dolomitic, pebbles and cobbles;
calcareous matrix; gradual boundary.
Reddish-brown (SYR 5/3 — moist) clay loam;
coarse prismatic structure; prisms break into
coarse blocky peds ; light brownish-gray (2.5Y
6/2 — moist) coatings on vertical faces of
prisms. These coatings effervesce strongly in
dilute acid; ped interiors effervesce moder¬
ately; a few, mainly dolomitic, pebbles and
cobbles; abrupt boundary with underlying
pro-Valderan mud flows.
As can be seen by its description, this soil was characterized by
a thick, dark colored, surface (Ap) horizon. Surface horizons of
this nature are not common to well or moderately well drained soils
of the Two Creeks area, as typical A1 horizons are thin, and under¬
lain by light colored eluvial layers. A likely explanation is that adja¬
cent, cultivated slopes had eroded during the post settlement era,
causing the accumulation of 18-20 cm of colluvium at the study
site. Tillage operations, concurrent with this period of colluvial
deposition, would have mixed these sediments with the original A1
horizon, forming the thick, uniformly dark colored and loamy,
mollic-like epipedon described. Field observations support this hy¬
pothesis as the study site was slightly concave in surface configu¬
ration, with adjacent, cultivated, slopes.
188 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Below the Ap horizon, A2 and Bl horizons were formed in thin,
sandy loam and silt loam strata, respectively. The low clay con¬
tent and numerous uncoated soil grains in the gray A2 horizon
indicate that both illimerization (loss of clay) and podzolization
(loss of iron) had occurred during soil formation. These observa¬
tions are supported by laboratory data on clay and iron content
(see Tables 1 and 2).
Yellowish-brown stains on peds in the A2 horizon suggest that
some secondary translocation of iron had occurred and that a Bir
horizon was beginning to form. This observation agrees with those
Table 1. Mechanical Composition and Textural Class of Modern Soil and
Underlying Deposits at Two Creeks Site.*
*C1 and C2 horizons are Valders till; C3, C4 and C 5 horizons are pro-Valders
deposits.
fGravel content was determined on bulk sample. Sand, silt and clay on < 2 mm
subsample.
JStandard abbreviations are used as follows: 1, loam; fsl, fine sandy loam; sil,
silt loam; cl, clay loam; s, sand.
Table 2. Content of Free Iron Oxides and Carbonates in Modern Soil
at Two Creeks Site.
1972] Lee and Horn — Pedology of Ttvo Creeks Section 189
made by Beaver and Lee (1963) which indicated that the Two
Creeks site is in a transitional soil zone where Spodosols form in
acid, sandy deposits; biseqnal soils (Alfic Haplorthods) in loamy
sediments underlain by calcareous, loamy to fine textured till at
moderate depth; and degraded Alfisols (Glossic Hapludalfs) in mod¬
erately fine to fine textured glacial drift having thin loamy surface
coverings. In the soil described, the loamy coverings were likely
not deep enough for an upper, Spodosol sequum to develop; how¬
ever, the rusty stains were indicative of juvenile development of
a Bir (spodic) horizon. Morphological observations of associated
soils along the face of the bluff lent support to this theory. Some
of these soils were formed in loamy and sandy coverings, 50 to
75 cm thick over Valders till, and exhibited bisequal profiles char¬
acterized by an upper, A2-Bhir sequum over A'2 and Bt horizons.
Other soils having very thin surface coverings were monosequal
in nature. It should be noted, however, that the latter soils were
characterized by a degraded B, or an A and B horizon indicating
strong podzolization.
Degradation of the B1 horizon in the soil studied was indicated
by the pinkish gray color of ped faces in that layer. This color
was indicative of the stripping away of iron coatings from sand
and silt grains, as could be observed with an ordinary (10X) hand
lens. The lower boundary of the B1 horizon marked a lithologic
discontinuity, as the underlying IIB2t horizon contained consid¬
erably more clay, as well as a few erratic pebbles and cobbles
(Table 1), suggesting that it was formed in Valderan till. Argil-
lans on ped faces in this layer indicated that illuvial clays were
present. Structural elements in the IIB2t horizon were extremely
well defined. This was likely due in part to exposure and desicca¬
tion of the profile, and in part to the moderately high content of
expanding layer silicate clay minerals characteristic of red clays
in this region (Petersen, Lee, and Chesters, 1966).
Total depth of A and B horizons in the modern soil was 71 cm;
the C horizon (unleached Valderan till) extended beyond this to
a depth of 224 cm. Laboratory data (Tables 1 and 2) showed this
till to be of heavy loam or clay loam texture in which carbonate
content ranged from 30 to 40%. Till of this texture is common to
the region (Lee, Janke and Beaver, 1962). Vertical faces of pris¬
matic elements in the C2 horizon were covered with brownish-gray
coatings; this same phenomenon has been noted by the authors in
other red clay deposits in eastern Wisconsin. The physical appear¬
ance of these coatings indicated that they were illuvial in origin;
their strong effervescence with HC1 suggested that they were com¬
posed in part of pedogenic calcite.
Below the Valderan till a layer of coarse and medium sand was
190 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
encountered (Fig. 1 and Table 1). This, and the two silty layers
beneath, resembled the pro-Valders fluvial deposits described ear¬
lier by Thwaites and Bertrand (1957). Occasional seams of sand
were noted in the upper part of the silty layers ; several wood frag¬
ments were noted in both layers.
The Two Creeks Soil
The Two Creeks paleosol was beneath the pro-Valders deposits
at a depth of 406 cm (Fig. 1). It consisted of a thin, compact or¬
ganic layer and a gleyed subsoil horizon (Fig. 3). Rooted in this
soil was a brokenover tree approximately 7 inches in diameter,
Figure 3. Two Creeks paleosol beneath light col¬
ored, silty, Pro-Valders deposits. The thin, dark
colored layer is the Ohb horizon. Note: the tape
is in inches.
1972] Lee and Horn — Pedology of Two Creeks Section
191
breast high (Fig. 4). It was lying toward the west-southwest,
corresponding to the generally assumed direction of Valders ice
flow at this point. Broeker and Farrand (1963) have dated wood
from the Two Creeks horizon at 11,850 ± 100 radiocarbon years
before present.
The gross morphology of the buried paleosol at this exposure
gave it the appearance of a thin, young, hydromorphic soil that
would be classified as an Aquent in the new (U. S.) system of soil
classification (Soil Survey Staff, 1967). Its description was as
follows :
Ohb 8-0 cm Black (10YR-5YR 2/1) fibrous peat containing
recognizable fragments of twigs; coarse platy
structure; very compact; brittle dry; slightly
acid (pH 6.5) ; abrupt smooth boundary.
Bgb 0-20 cm Grayish-brown (2.5Y 5/2) to light brownish
gray (2.5Y 6/2) loam; weak platy structure;
apedal; friable moist; many pebbles; calcare¬
ous; clear wavy boundary.
Figure 4. Broken over tree, rooted in the Two
Creeks paleosol. This tree points west-south¬
west, indicating the direction of Valders ice
flow at this point.
192 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Clb 20-26 cm
C2b 26-41 cm
IIC3b 41-46 cm
IIC4b 46-66 cm
Brown (10YR 5/3) loamy sand; single grain;
stratified; calcareous; abrupt boundary.
Brown (10YR 5/3) gravelly loamy sand; strati¬
fied; loose; iron stains noted; calcareous;
abrupt wavy boundary.
Pale brown (10YR 6/3) silt loam (laminated
silts, clays, and fine sands) ; very friable; cal¬
careous; abrupt boundary.
Reddish gray (SYR 5/2) silty clay loam (lam¬
inated silts and clays) ; massive, breaking with
conchoidal fracture; very hard dry and very
firm moist; calcareous. This deposit extended
to the level of the modern beach.
As can be seen from its description, the uppermost horizon of
the Two Creeks soil consisted of dark colored, fibrous, organic ma¬
terial, many wood fragments, and considerable mineral material;
its compact nature suggested that it had been considerably thicker
before being compressed by the weight of overlying deposits.
Analyses of this horizon (Table 3) showed that the organic frac¬
tion was highly soluble in Na4PL>07, indicating a more advanced
stage of decomposition than its fibrous nature and relatively wide
C:N ratio (33) would suggest. Chesters (1959) studied a sample
of material from this horizon and reported that its thermogram
“showed many lignin characteristics” ; he believed it to be more
highly decomposed than a sample of Horicon muck, a well decom¬
posed, contemporary Histosol. On the basis of these analyses the
horizon was classified as Oh, despite its fibrous appearance. In the
new system of classification (Soil Survey Staff, 1968), an Oh
(hemic) horizon is partially decomposed but may be fibrous in
appearance.
Other characteristics of this horizon that relate to its genesis
were its slightly acid pH and relatively high ash content (Table 3).
Free carbonates were not present except as they occurred in the
shells of mollusks or other soil fauna. This suggests that carbonate
Table 3. Chemical and Physical Properties of the Surface Horizon of
the Two Creeks Paleosol.
* Acronym for sodium pyrophosphate extract color (Munsell notation).
1972] Lee and Horn — Pedology of Two Creeks Section 193
laden waters did not move through this horizon, either vertically or
laterally, while it was buried.
The high content of ash in the Ohb horizon indicates considerable
incorporation of mineral sediments. A partial explanation is that
during its terminal stage, the Two Creeks Forest was inundated by
waters in front of the advancing Valders ice, and then buried by
mud flows. During this period, considerable sand, silt and clay were
deposited on the Ohb horizon; some of this sediment likely infil¬
trated the layer.
Another factor contributing to the high mineral content of the
Ohb horizon relates to its environment of formation. Wilson’s
studies (1932, 1936) of the paleocology of the Two Creeks horizon
showed that 3 biologic zones were present, corresponding to 3
periods of development during its genesis. The first period was
characterized by aquatic or semi-aquatic mollusks, indicating a
wet, early stage at which time muddy runoff waters were likely
present. The middle period was characterized by mosses character¬
istic of moist to dry woodlands. A major part of the Ohb horizon
at this site probably developed as the 01 or 02 horizon of an up¬
land forest soil; its high mineral content is likely due in part to
the mixing of mineral and organic material frequently seen in such
horizons.
Below the Ohb was the Bgb horizon, formed in a loamy deposit
containing many pebbles (Table 4). According to Thwaites and
Bertrand (1957), the sediments comprising this and underlying
layers were deposited in glacial Lake Chicago in front of Cary ice
as it melted and retreated northward.
Table 4. Mechanical Composition and Textural Class of Two Creeks
Soil and Underlying Deposits.
*Gravel content was determined on bulk sample; sand, silt and clay on < 2 mm
subsample.
jStandard abbreviations are used as follows: 1, loam; Is, loamy sand; gr Is,
gravelly loamy sand; sil, silt loam; sicl, silty clay loam.
194 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
The Bgb horizon was about 20 cm thick ; morphological evidences
of pedogenesis included its gray, reduced color, indicative of gleiza-
tion, and structural development. In comparison to the underlying
Clb and C2b horizons, it had a higher content of clay and free iron
and a somewhat lower content of carbonates (Tables 4 and 5). The
latter likely represents leaching during the relatively brief period
that this soil underwent pedogenesis. Free iron content is in part a
reflection of original texture, but it may also reflect illuvial accumu¬
lation ; clay content is likely related to sedimentary history.
The present day morphology of the B horizon suggests that it
was formed under water-logged conditions and that it was the sub¬
soil of a hydromorphic soil. This may not have been the case, how¬
ever, as the process of gleization is not strictly a pedogenic process
but occurs quite readily wherever there are reducing conditions,
providing a source of energy such as organic matter is available.
This can be demonstrated in the laboratory and is frequently ob¬
served in young soils, for example in waterlogged layers of recent
colluvium. Near Coopers Mill, a few km inland from the Two Creeks
site, gravel pit operations exposed logs buried in Valderan out-
wash. Examination of the sand around many of these logs revealed
a peripheral gleyed zone, several inches thick, and obviously de¬
veloped in situ since burial.
Beneath the Bgb horizon were thin layers of loamy sand, gravel,
and silt loam, underlain by reddish colored, calcareous, laminated
silts and clays which extended to the level of the modern beach. The
latter were similar to the Early Lake Chicago deposits described by
Thwaites and Bertrand (1957). According to their observations,
which were made when higher lake levels caused rapid removal
of talus from the base of the bluff, these deposits ranged from 7
to 20 feet in thickness and were underlain by gray, loamy, Cary-
age glacial till.
Table 5. Content of Free Iron Oxides and Carbonates in the
Two Creeks Paleosol.
1972] Lee and Horn — Pedology of Two Creeks Section 195
History of the Two Creeks Site
The chronology of events at the Two Creeks site has been de¬
scribed by Thwaites and Bertrand (1957), Wilson (1932), Hough
(1958), Murray (1953) and others as follows:
1. When Cary (Woodfordian) glacial ice retreated from eastern
Wisconsin, the Lake Michigan basin was flooded by waters
of glacial Lake Chicago, This was the Glen wood State (Wil¬
son, 1932, quoting Thwaites, unpublished) during which
lake waters were about 60 feet higher than at present or
about 640 feet elevation. Sediments, including varved silts
and clays, derived in part from reddish brown fine textured
elastics siphoned down from the Lake Superior basin (Mur¬
ray, 1953), (Peterson, Lee and Chesters, 1966), were de¬
posited in glacial Lake Chicago,, Green Bay, and the Lake
Winnebago lowlands.
2. When Cary ice retreated beyond the Straits of Mackinac,
freeing the northern outlet of the lake, water levels in Lake
Chicago fell to present levels (580 feet) or lower. Freshly
exposed soil material was invaded by aquatic and semi aquatic
mollusks, mosses, and other water loving plants, later by
trees, primarily black spruce (Picea mariana) in moist areas
(Wilson, 1932). This, Two Creekan substage estimated by
Frye and Willman (1960) to be not over 1500 years in
duration, was the period during which the Two Creeks soil
formed.
3. The Two Creekan period ended with the advance of Valders
ice. Water in front of the glacier rose to the Calumet outlet
(about 40 feet above present levels) ; the Two Creeks Forest
was inundated, and pro- Valders lake deposits were deposited
on the forest floor. Valders ice then passed over the area bury¬
ing the Two Creeks soil and overlying deposits beneath red
till.
4. When Valders ice retreated, about 7000 years ago (Frye,
Willman and Black, 1965), the area in front of the melting
ice was again flooded. Sand, silt and clay was deposited
locally on uplands. When the Straits of Mackinac were again
freed of ice, water levels receded to present day levels. For¬
mation of the modern soil at the Two Creeks site presumably
began between 7000 and 5000 years B.P., and has continued
until the present day.
Soil Formation
The Two Creeks soil formed from the vegetative remains of
plants growing on it, and in sediments presumably deposited in
glacial Lake Chicago during retreat of Cary ice. At the site studied
196 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
the varved silts and clays, described by Thwaites and Bertrand
(1957), were overlain by thin layers of sandy and loamy sedi¬
ments. The modern soil at the Two Creeks site was formed partially
in Valders till and partially in loamy post-Valders deposits, the
latter likely laid down in front of the melting Valders ice and later
reworked in part by wind. The till at this site is of clay loam tex¬
ture (about 35% clay) and is calcareous (40% calcium carbonate
equivalent). The clay fraction of glacial till in this region is com¬
prised mainly of montmorillonite, mica, vermiculite, chlorite and
interstratified minerals (Janke, 1962). Loamy cover sediments
were likely calcareous when deposited but included only 16-18%
clay (Table 1).
Forest vegetation at the Two Creeks site was comprised mainly
of black spruce (Wilson, 1932). A log identified as white spruce
(Picea canadensis) was also found by Wilson, however he believed
it to have been derived from a drier site. At his second site, Wil¬
son (1936) also found balsam. Other flora included mosses and
lichens. A number of the former were identified by Cheny (1930;
1931) who found them to be existing species that are generally
found north of the Two Creeks site under present-day climatic
conditions. The organic-rich surface layer of the soil also included
pollen spores indicating the presence of grasses (Gramineae), heath
plants (Ericaceae) , birch (Betula) and jackpine (Finns banksiana)
in nearby areas during Two Creeks time. Wilson (1932) concluded
from the boreal character of the fossil trees and the associated
biota that the climate during the Two Creeks interval was prob¬
ably like that of northern Minnesota today, or somewhat colder.
Later studies by Culberson (1955) of fossil mosses supported this
conclusion. Zumberge and Potzger (1955) , after studies of a nearby
area in western Michigan, came to a similar conclusion on the basis
of pollen studies and radio-carbon dating, finding evidence that a
spruce-fir forest and a cool to cold, moist climate existed in that
area about 11,000 years before present. West (1961), on the basis
of pollen studies in a Cary end moraine bog (outside the Valderan
drift), found that white spruce (Picea glauca) reached a maxi¬
mum during Two Creeks time suggesting that the vegetation at
that time had true boreal character. Later, when Valders ice ad¬
vanced, black spruce became prominent, indicating wetness.
If the climate during Two Creeks time was somewhat cooler
than at present, and the Two Creeks Forest consisted of moist to
dry woodlands for at least 300 years (Wilson, 1932, 1936; Broeker
and Farrand, 1963), the dominant soil forming process during the
period the site was covered by forest growth was likely podzoli-
zation rather than gleization, with the latter process being dom¬
inant prior to growth of the forest, and during and after inunda-
1972] Lee and Horn — Pedology of Two Creeks Section 197
tion and burial of the Forest Bed. Following this line of reasoning,
the horizon we now call Bgb may very well have been Bir or Bhir,
i.e., a spodic-like horizon characterized by an accumulation of illu¬
vial iron, or illuvial iron and humus, at the time of burial. Later,
iron in the subsoil horizon could have been reduced to produce the
gley-like horizon present today.
Climatic and biotic factors affecting the modern soil at Two
Creeks can probably best be inferred from West’s (1961) pollen
studies. West found that pioneer vegetation, following retreat of
glacial ice, consisted of scattered white spruce, weeds and shrubs.
These were followed by oak (Quercus) and birch (Betula) and as
a warming trend continued, jack pine, elm (Ulmus), oak and iron-
wood (Ostrya). Later, white and red pine (Pinus strobus) replaced
jack pine, and hardwoods such as elm, hickory (Cary a), iron wood,
walnut (Juglans) and basswood (Tilia) became prominent. As the
climate became warmer and drier, and lake levels dropped to the
Lake Chippewa stage during the xerothermic period that ended
about 3500 years before present, oak became the dominant tree
species. At the same time grasses and forbs (e.g., Gramineae,
Cyperaceae and Compositae) became common, indicating prairie¬
like vegetation. During a succeeding cooler and moister period,
oak was followed by pine and then beech, hemlock (Tsuga) and
birch. Hemlock then increased in importance up until settlement,
when forests were cleared and most of the land cultivated.
The humid cool and humid to subhumid temperate climatic re¬
gimes described, in combination with coniferous and deciduous
vegetation, would be conducive to soil forming processes such as
eluviation and illuviation of clay (illimerization) , and of iron and
humus (podzolization) . Soils in the region having distinct argillic
(clay enriched) subsoil horizons are generally believed to have
formed under hardwoods such as oak, hickory, maple and bass¬
wood, and in some cases prairie, alternating with hardwoods, under
humid to subhumid temperate climatic conditions. Forest species
such as red and white pine, white spruce, beech, and hemlock, espe¬
cially the latter, produce litter that is conducive to the formation
of podzol soils (Lee, 1955, Wilde, 1958) ; cool humid climatic con¬
ditions are associated with all major podzol (Spodosol) regions.
It appears therefore, that during the early stages of formation,
as the forest canopy closed above it, the modern soil accumulated
organic matter in and on its surface layer. This was followed by
the removal of carbonates from the upper part of the soil by pre¬
cipitation percolating through the litter layer and into the mineral
soil below, slowly leaching out the free carbonates present. Clay
was then dispersed in surface layers and translocated to subsoil
horizons where it was deposited on ped surfaces, and in pores, as
198 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
water was withdrawn into the soil matrix by capillary action. This
last process, occurring over a long period of time, produced the
argillic (Bt) horizon present in this soil today.
Following the xerothermic period about 1500 B. C., and the in¬
crease in pine, beech, and hemlock, podzolization likely became the
dominant soil forming process in the upper, leached and relatively
permeable part of the soil. While podzolization did not proceed
very far in the pedon studied, where loamy coverings were rela¬
tively thin, it was apparent that in adjacent pedons, where loamy
or sandy deposits extended to greater depths, podzolization was
much more pronounced.
The thick dark colored surface horizon of the modern soil can
best be explained as an anthropic feature brought about by man’s
cultivation of the land, resulting in erosion of surrounding slopes
and deposition of topsoil at the study site.
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clays. Soil Sci. Soc. Amer. Proc. 17: 359-364.
Beaver, A. J. and G. B. Lee. 1963. Distribution and characteristics of some
bisequal soils in eastern Wisconsin. Amer. Soc. Agron. Abstracts, p. 53.
Black, Robert F. 1966. Valders glaciation in Wisconsin and Upper Michigan —
A progress report. Great Lakes Research Division, Pub. No. 15, 169-175.
Broecker, W. S. and W. R. Farrand. 1963. Radiocarbon age of the Two
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Cheny, L. S. 1930. Wisconsin fossil mosses. 1930. Bryologist 33: 66-68.
Cheny, L. S. 1931. More fossil mosses from Wisconsin. Bryologist 34: 93-94.
Chesters, G. 1959. Soil aggregation and organic matter decomposition. Ph.D.
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LOWER WISCONSIN RIVER VALLEY SOIL
RESOURCES AND USE POTENTIALS
G. E. Musolf and F, D . Hole
The Wisconsin River flows for 90 miles (145 km) from the gla¬
cial end moraine at Prairie du Sac to the confluence with the Mis¬
sissippi River at Prairie du Chien. This stretch is called the Lower
Wisconsin River Valley of which the floor includes 145, 107 acres
in area (2272 mi; 587 km2). The valley floor consists of floodplain
and a sequence of stepped natural terraces, only 10 percent of
which, by area, are rock benches, the rest being of glacial outwash.
The geomorphic history of these features has been discussed else¬
where (Hole et aL, 1952; Musolf, 1970).
The westward to southwestward trend of this valley runs coun¬
ter to the major flow of people and freight between Chicago and
Twin Cities (Figure 1). “Nature-made highway’’ was the title
assigned to the Fox-Wisconsin waterway by Whitbeck in 1915.
The idea of improving the Lower Wisconsin River by local dredg¬
ing and excavation of supplemental side canals was promoted by
W. J. Nicodemus in an article in these Transactions in 1874, when
a passage for steamboats to carry grain eastward was seen as a
real need. Other more rapid means of transportation developed on
land before this “improvement” could be accomplished, as was
recognized by F. E. Williams (1921) who attributed “the passing
of an historic waterway” to the directional trend mentioned above,
to the shallowness that even impeded canoe passage, to frozen con¬
ditions in winter and to several other factors. The valley is rela¬
tively little trafficked even today and retains much of its natural
beauty. It is a major “environmental corridor” (Lewis, 1964)
which is attracting an increasing number of tourists from the
vicinities of the three nearest urban centers (Figure 1). The floor
of the Lower Wisconsin River Valley has five features that qualify
it as an environmental corridor : water, wetlands, floodplain, sandy
soils and escarpments at the edge of the terraces. The valley floor
is bracketed by scenic wooded bluffs and ridges, themselves “corri¬
dors”, although peripheral to the specific emphasis of this study.
It is the purpose of this paper to report on the soil resources
of the valley floor as a unit, and to suggest how a zoning ordinance
might function on the basis of the detailed soil map in such a way
as to avoid unwise land use in this unique area. We may look at
the valley from the point of view of a hypothetical Lower Wis-
201
202 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 1. Regional location of the Lower Wis¬
consin River Valley.
consin River Valley planning commission as it might begin with
a soil inventory to assess the potentialities of this long, narrow
strip of land, water and scenery with respect to agriculture, indus¬
try, recreation, wildlife, residences and esthetic enjoyment.
The data for the soil inventory have been collected over many
years in the portions of six counties which the valley includes
(Figure 2). The soil mapping has been done by soil scientists of
the Soil Conservation Service and cooperating soil scientists of
the University of Wisconsin. Musolf (1970) assembled the maps
for the entire valley and measured on them the acreages of the 356
different soil phases that had been listed previously on a county
basis (Robinson and Klingelhoets, 1959; Robinson and Klingel-
hoets, 1961 ; Slota and Garvey, 1961 ; Klingelhoets, 1962) . He also
arranged the soil map information in a way suitable for incorpora¬
tion into a zoning ordinance similar to that adopted in Buffalo
County, Wisconsin (Buffalo County Zoning Committee, 1965).
Grateful acknowledgement is made to Soil Conservation Service
1972] Musolf and Hole— Wisconsin River Valley Soil
203
Figure 2. Fluvial terraces and floodplain of the Lower Wisconsin River valley.
soil scientists R. W. Slota and C. Glocker for their assistance, as
well as to E. H. Hammond who, as Professor of Geography at the
University of Wisconsin at Madison, helped to initiate this study.
Acknowledgement is also made to the University of Wisconsin
Center System Faculty and Curricula Development Committee for
financial support of the senior author during this study.
Methods and Procedures
Field investigations were carried out by the senior author dur¬
ing the summers of 1962, 1965, and 1968. Natural river terraces
and rock benches and land use patterns were mapped. The junior
author was a participant in the soil mapping program in Richland
and Grant Counties (Hole, el ah, 1950; Hole, 1956) and in coopera¬
tive soil correlations in Iowa and Dane Counties.
Acreages of soil phases in the valley were determined by the
“cut and weigh” method by which an analytical balance was used
to determine actual weights of portions of the map. Care was taken
to establish controlled conditions for these measurements. Calcula¬
tion of proportionate areas of the map units on the basis of weight
was done with the aid of a desk calculator. Acreages were sum¬
marized by towns and counties of the valley, and for the entire
valley floor as a whole.
Versatility of the Lower Wisconsin River Valley and its
Relation to Multiple Land Use Potential
The versatility of the valley is indicated by the variety of its
soils which range from riverwash and peat to active sand dunes
and level productive loams, all enclosed by steep bluffs, 300 to 500
feet (100 to 170 m) high. Sixty percent of the soils of the natural
terraces, or about 56,000 acres (22,670 hectares) developed under
204 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
stands of tall prairie, 23 percent under forest and 17 percent under
bog and marsh vegetation, as interpreted from soil profile charac¬
teristics. The valley walls stand about four and one-half miles
(7.2 km) apart near the Cary end moraine at Prairie du Sac and
near Lone Rock, and only one-half mile (0.8 km) apart near the
confluence of the Wisconsin and Mississippi Rivers (Figure 2).
The abundance of groundwater would make possible some industry
(within the strict limits of modern environmental quality stand¬
ards) and irrigated truck cropping, such as is already practiced on
sandy and loamy soils. It is possible that sludge from Madison
sewage treatment plants might be used to fertilize truck crop fields
in the valley in northern Iowa County. The ribbon-like shape of the
area, however, precludes development of large-scale commercial
canning crop operations, which require roughly equidimensional
clusters of numerous 180-acre blocks of level soils irrigable by
self-propelled rotating sprinkler systems. The cultivable part of the
valley floor is irregularly partitioned and interrupted by sinuous
floodplain, railroads and highways, and old dune ridges. The valley
offers scenic beauty and opportunities for outdoor recreation,
coupled with production of vegetable crops and dairy products on
a limited scale. Farmers' roadside stands are familiar sights during
the growing season.The village of Spring Green in Sauk County is
a cultural center with a legitimate stage theatre that attracts
visitors in the summer. State, county and private facilities for
boating, camping, and a variety of forms of recreation are impor¬
tant to the economy of the area. The Lower Wisconsin River Valley
contains much of scientific interest with respect to geology, geog¬
raphy, botany, zoology, ecology, archaeology and soil science.
The Soil Resources
On the valley floor are nearly one hundred different types of
soil and miscellaneous land units. These are subdivided into the
356 soil phases on the basis of slope, degree of erosion and land¬
scape position. The intricate pattern of these soils lies on major
geomorphic units (Figure 2) as follows. Floodplain soils account
for 35 percent of the area. These include alluvial land and river-
wash (81.5%), peat and muck (11.5%) and marsh (7%) . Twenty-
six acres of cherty alluvial land are the result of recent flood
deposits by tributary streams. Terraces, mostly made of glacial
outwash, occupy the remaining 65 percent of the area, including
six small rock terraces in the lower half of the valley. Escarpments,
usually less than 25 feet in height, commonly mark boundaries
between flood plain soils and terrace soils, and between terraces
of different levels.
Soils information plotted in Figure 3 shows that the valley
floor is predominantly level to gently sloping, only slightly to
1972]
Musolf and Hole — Wisconsin River Valley Soil
205
Figure 3. Areal extent of soil physical characteristics and land use capability
in the Lower Wisconsin River valley.
moderately eroded, naturally excessively drained (drouthy) to well
drained, sandy in texture and fourth class in land use capability.
Bimodal features of the soils are 1) presence of both excessively
and poorly drained soils, and 2) presence of both sandy and silty
soils. Figures 4 through 6 illustrate soil ratings on major terrace
levels.
The silty soils are derived from loess (and their derivatives)
largely deposited during the period 29,000 years before present
(Hogan and Beatty, 1963) to the time of post-Cary loess deposi¬
tion. This later deposition was probably between 14,, 000 and
6,000 years before present judging by pedogenic analyses by Allan
and Hole (1968). The sand of the valley floor was deposited by
meltwater from Valderan glacial ice about 9,000 years ago, and
wind has redistributed it as dunes and valley Alls both southward
(as Chelsea sand in Grant County) and northeastward (Plainfield
in Richland County) (Hole, 1956; and Hole, et al., 1950). Some
of the loams of the terraces appear to be admixtures of the sand
and overwash of silt from tributary valleys. The mixing was
probably by biotic agents, particularly ants (Baxter and Hole,
1967).
The soils of the natural terraces differ from the valley floor as
a whole in being a little more sloping and eroded,, sandier,
drouthier, and very slightly higher in land use capability, accord¬
ing to the Soil Conservation Service system of rating. An experi¬
mental numerical productivity rating of soil used by Musolf (1970)
in the Lower Wisconsin River Valley gave the Plainfield loamy
sand a rating of 39, the Sparta loamy fine sand 57, the Dakota
206 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 4. Low terrace, Sauk City, Sauk County: Dakota
sandy loam; S.C.S. land capability — Class III-s; Soil pro¬
ductivity rating — 75.
sandy loam 75, the Richwood silt loam 97 (near the maximum
possible rating), and all the soils of the valley an average of 61,
as compared with a rating of 90 for the upland prairie (Argiudoll)
soil landscape of southern Grant County near Cuba City. It is true
that with irrigation and fertilization the sands of the valley ter¬
races could be brought up to the equivalent of the Dakota loam
in productivity. But the dissection of the soil bodies into narrow
strips, already referred to, precludes development of significant
crop units. The relatively low natural agricultural productivity of
the soils of the valley still dictates an emphasis on multiple land
use with special attention to recreational activities and forestry.
Suggested Use of the Detailed Soil Survey for Zoning Purposes
in the Lower Wisconsin River Valley
A well designed zoning ordinance makes, possible the avoidance
of objectionable land uses, such as misplacement of non-farm rural
homes on soils incapable of accepting septic system effluent and
and construction of hunting and fishing shacks on floodplains
(Yanggen et al, 1966). Primary rural land use districts may be
set up, as was done in Buffalo County (Buffalo County Zoning
Committee, 1965) under six headings: agricultural, residential,
1972] Musolf and Hole — Wisconsin River Valley Soil
207
>v'v- ' ' ^ '
Figure 5. Intermediate terrace, 1% miles west of Mazo-
manie, Dane County: Sparta loamy fine sand; S.C.S. land
capability — Class XV-s; Soil productivity rating — 57.
ipllllll
Figure 6. High terrace, 1 mile northwest of Wauzeka, Crawford County: Fay¬
ette silt loam , uplands; S.C.S. land capability — Classes Ill-e and IV-e; Soil
productivity rating — 75 and 48.
208 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
recreational, commercial, industrial and floodplain. These districts
are related rationally to specific parcels of land by using the de¬
tailed soil map as the zoning map, with four interpretive overlay
maps on it showing four soil districts : steep soils, wet soils, flood-
plain soils and suitable soils. The suitable soils are further sub¬
divided into sandy soils, medium-textured soils and clayey soils
so that lots may be made large enough for adequate on-lot sew¬
age disposal. Musolf (1970) has grouped the soils of the Lower
Wisconsin River Valley in this manner and has shown that the
area can be zoned under a uniform system. Musolf’s overlay group¬
ing (too voluminous for reproduction here) is suitable for use
with the Buffalo County Ordinance and detailed soil maps which
are published for Crawford, Grant, Iowa and Richland Counties.
Incorporation of a detailed official soil map into a zoning scheme
makes decisions about use of most parcels of land clear-cut and
unclouded by conflict of opinion. Where serious questions are raised,
additional field checking by soil scientists can quickly lead to a
satisfactory solution. Advantages of the use of the soil survey in
zoning outweigh the disadvantages (Yanggen et al., 1966) .
Summary
Since the exploratory canoe trip along the Lower Wisconsin
River Valley in 1673 by Pere Jacques Marquette and Louis Joliet,
European and American settlers have replaced the Indian occu¬
pants, exploited the forests and prairies and, in succession, prac¬
ticed wheat, hop, corn-hog, dairy and truck crop farming. Devel¬
opment of the Lower Wisconsin River as a main transportation
route never materialized. Erosion control practices, planting of
trees in shelterbelts and plantations, and protection of woodlands
from grazing have resulted from a growing awareness by the
inhabitants of the need for soil and water conservation and from
technical assistance provided to land operators by the Soil Con¬
servation Service and the College of Agricultural and Life Sci¬
ences working through cooperative Extension. Recreational activ¬
ities and residential developments have been increasing in the area.
Recent elevation of standards for the protection of quality of water
and other components of the environment, and an increasing ap¬
preciation of the scientific, esthetic and recreational values of this
principal environmental corridor of Wisconsin point to the need
for a practical land use zoning system in the valley. It is advan¬
tageous to base the zoning on the detailed soil maps that are now
available, along with interpretive overlay maps and zoning direc¬
tives that regulate land use according to site characteristics.
1972] Musolf and Hole — Wisconsin River Valley Soil
209
Literature Cited
Allan, R. J. and F. D. Hole. 1968. Clay accumulation in some Hapludalfs as
related to calcareous till and incorporated loess on drumlins in Wisconsin.
Soil Sci. Soc. Amer. Proc. 32: 403-408.
Baxter, F. P. and F. D. Hole. 1967. Ant (Formica cinerea) pedoturbation in
a prairie soil. Soil Sci. Soc. Amer. Proc. 31: 425-428.
Buffalo County Zoning Committee. 1965. Buffalo County Zoning Ordinance.
p. 28.
Hogan, J. D. and M. T. Beatty. 1963. Age and properties of Peorian loess
and buried paleosols in southwestern Wisconsin. Soil Sci. Soc. Amer. Proc.
27: 345-350.
Hole, F. D. 1956. Soil Survey of Grant County, Wisconsin. Bui. 80. Wis. Geo¬
logical and Nat. Hist. Survey, University of Wisconsin, p. 54.
Hole, F. D., F. F. Peterson and G. H. Robinson. 1952. The distribution of
soils and slopes on the major terraces of southern Richland County, Wis¬
consin. Trans., Wis. Academy of Sci., Arts and Letters. 41: 73-81.
Hole, F. D., G. H. Robinson and A. V. Miller. 1950. Soils of Richland County,
Wisconsin. County and Town Folder maps. Wis. Geological and Nat. Hist.
Survey, University of Wisconsin.
Klingelhoets, A. J. 1962. Soil survey of Iowa County, Wisconsin. Soil Con¬
servation Service, USD A, Series 1958, No. 22, U.S. Government Printing
Office, Washington, D.C. 101 pp., 44 map sheets.
Lewis, P. H., Jr. 1964. Landscape resources of Wisconsin. Wisconsin Blue
Book: 130-142.
Musolf, G. E. 1970. The geography of the Lower Wisconsin River Valley with
emphasis on soil resources of the fluvial terraces. Ph.D. Thesis, The Uni¬
versity of Wisconsin, Madison, p. 248.
Nicodemus, W. J., 1874. On the Wisconsin River improvement. Trans., Wis.
Academy of Sci., Arts and Letters. 2: 142-152.
Robinson, G. H. and A. J. Klingelhoets. 1959. Soil survey of Richland County,
Wisconsin, Soil Conservation Service, USD A, Series 1949, No. 9, U.S.
Government Printing Office, 42 pp., 51 map sheets.
Robinson, G. H. and A. J. Klingelhoets. 1961. Soil Survey of Grant County,
Wisconsin, Soil Conservation Service, USD A, Series 1951, No. 10, 98 pp.,
72 map sheets.
Slota, R. W. and G. D. Garvey. 1961. Soil survey of Crawford County, Wis¬
consin, Soil Conservation Service, USD A, Series 1958, No. 18, U.S. Gov¬
ernment Printing Office, 85 pp., 39 map sheets.
Whitbeck, R. H. 1915. The geography of the Fox-Wisconsin valley. Bui.
XLII, Wisconsin Geological and Natural History Survey, Madison, p. 27.
Williams, F. E. 1921. The passing of an historical waterway. Trans., Wis.
Academy of Sci., Arts and Letters. 20: 131-140.
Yanggen, D. A., M. T. Beatty and A. J. Brovold. 1966. Use of detailed soil
surveys for zoning. J. Soil and Water Conservation. 21, no. 4: 123-126.
-M
THE PITUITARY GLAND OF THE ALEWIFE IN LAKE
MICHIGAN: CYCLICAL CHANGES IN THREE
ADENOHYPOPHYSEAL CELL TYPES
Alexander H. H. Li and Eldon D. Warner
The population explosion of the alewife ( Alosa pseudoharengus ,
Wilson) in the Great Lakes is frequently accompanied by massive
spring and early summer mortalities. The possibility of endocrine
involvement in this phenomenon was first suggested by Hoar
(1952). He found histological evidence of thyroid hyperplasia and
exhaustion in alewives in Lake Ontario and postulated thyroid-
related osmoregulatory failure as a factor in the die-offs. The lack
of more recent endocrine information emphasizes the need for fur¬
ther study of this important regulatory system in relation to ale¬
wife physiology and mortality.
Since the pituitary gland plays a key role in a variety of hor¬
monal activities, it is a prime target for investigation. In elucidat¬
ing pituitary function a logical first step is to identify the specific
types of hormone-secreting cells and to study their annual patterns
of change. Investigations of this nature have been carried out in
many other species of teleost fishes. The earlier literature has
been reviewed by Pickford and Atz (1957). Olivereau (1963)
described six types of chromophilic cells in the teleost pars distalis
on the basis of her own and other work. The tentative functions
which she assigned to these cells have largely been supported by
subsequent histophysiological studies. The pars intermedia of
certain teleosts appears to contain two additional cell types.
(Olivereau, 1969) Thus, a total of eight kinds of cells have been
identified in the teleost adenohypophysis. Among the more recent
studies of cyclical pituitary changes are those of Sokol (1961),
Robertson and Wexler (1962 a and b), Sathyanesan (1963), Lagios
(1965), Olivereau (1967) and Sage and Bromage (1970).
The present report includes a description of the histology and
cytology of the alewife pituitary gland and an account of cyclical
variations in three adenohypophyseal cell types, the gonadotrophs,
thyrotrophs and corticotrophs.
Supported in part by Sea Grant Project #68-4203 from the National Science Foun¬
dation.
211
212 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Materials and Methods
Alewives were collected by trawl, seine or dip net from several
sites in southern Lake Michigan. The collections covered a period
from May 21, 1968 to April 27, 1969 with all months represented
except September and October. The samples of May through Au¬
gust 1968 were obtained off Saugatuck, Michigan while those of
November 1968 through April 1969 were taken along the western
shore of the Lake between Waukegan, Illinois and Port Washing¬
ton, Wisconsin. Adult specimens of both sexes were present in all
collections.
Immediately after capture, the fish were placed in Bouin-
Hollande fixative for at least three days. After fixation they were
air dried and at this time body weight, standard length and sex
were recorded and scale samples were taken for age determinations.
Prior to further processing they were stored in 70% ethanol. Fifty
adult specimens, all apparently healthy when captured, were se¬
lected for the pituitary study.
The dorsal portion of the head of each specimen was removed
and decalcified in 5% formic acid for five days. After decalcifica¬
tion, extraneous tissue was trimmed away, leaving a cube about
0.5 cm. square per side containing the pituitary gland and asso¬
ciated brain structures. The preparations were dehydrated in
ethanol, cleared in xylene and infiltrated and imbedded in para-
plast. Serial sagittal sections were cut at six micra. The staining
procedures employed were as follows: (1) periodic acid-Schiff
reagent (PAS), lead hematoxylin and orange G (modified from
MacConaill, 1956, McMannus and Mowry, 1958 and Elftmann,
1959 a and b) ; (2) aldehyde fuchsin, lead hematoxylin and orange
G (modified from Gomori, 1950, MacConaill, 1956 and Elftmann,
1959 a and b; (3) erythrosine, Mallory II and acid alizarine blue
(modified from Herlant, 1960).
Pituitary cell types were identified by tinctorial reaction, mor¬
phology and location and by reference to other histological and
histophysiological studies.
Several criteria were used in evaluating the cyclical secretory
activity of gonadotrophs, thyrotrophs and corticotrophs. Nuclear
diameters were measured with an ocular micrometer on midsagittal
sections of glands from four fish (two males and two females) for
each collection date. The mean nuclear diameter of each cell type
on a given collection date was based on a total of 40 measurements.
The Student-N ewman-Keu! multiple range test was employed to
compare differences among the means for statistical significance.
Results are shown graphically in Figure 1. Other criteria of secre¬
tory activity were cell size, degree of cytoplasmic granulation and
vacuolation, staining intensity and nucleolar prominence.
1972]
Li and Warner — Alewife in Lake Michigan
21
Figure 1. Annual Size Variations of Gonadotroph, Thyrotroph
and Corticotroph Nuclei.
Observations and Discussion
Histology and cytology.
The pituitary gland of the alewife is very similar to that of its
relative the herring ( Clupea harengus, L) described by Buchmann
(1940). The alewife gland is ovoid in shape with a tapering an¬
terior region that ends in a narrow hypophyseal duct. According to
Buchmann the duct is open to the pharynx in young herring but
closed in adults. No pharyngeal connection was observed in adult
alewives, although the duct extends for some distance in an antero-
ventral direction toward the pharynx. The major pituitary regions,
neurohypophysis and adenohypophysis, are readily distinguishable
in histological sections (Fig. 2).
The neurohypophysis consists largely of fiber tracts that origi¬
nate in the hypothalamus and extend through the infundibular
stalk to the posterior dorsal part of the gland. Here, the greatest
concentration of neurohypophyseal tissue is located. From this
area, fiber bundles of varying size ramify into the other pituitary
regions and form interdigitations with groups of adenohypophyseal
cells. Numerous glial cells with ovoid nuclei are scattered among
the fibers. Aldehyde fuchsin-positive globules, presumed to be
products of neurosecretion, are frequently present, especially in
214 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Figure 2. Mad-sagittal section of alewife pitui¬
tary gland (X 47). D, hypophyseal duct; H,
hypothalamus; I, pars intermedia; N, neuro¬
hypophysis; P, proximal pars distalis; R, rostral
pars distalis; S, infundibular stalk.
the posterior dorsal region. The neurohypophysis is well vascular¬
ized and small vessels are abundant in close proximity to adeno-
hypophyseal cells.
The adenohypophysis is subdivided into the three regions char¬
acteristic of most bony fishes. From anterior to posterior they are :
rostral pars distalis (pro-adenohypophysis), proximal pars distalis
(meso-adenohypophysis) and pars intermedia (meta-adenohypo¬
physis).
The rostral pars distalis constitutes about 40% of the adeno¬
hypophysis. The cells are arranged for the most part in follicles
of varying size and shape. (Fig. 3). With the techniques used,
little stainable material is seen in the follicular lumina. Neuro¬
hypophyseal fibers are interspersed among the follicles and the
latter are bounded by basement membranes.
Examination of serial sections reveals that the follicles are not
isolated units, but instead, have interconnecting lumina. Further¬
more ; every lumen is in contact directly or indirectly with the
lumen of the hypophyseal duct. Thus, the follicle cells form a con¬
tinuous, folded epithelium surrounding passages that are essen¬
tially ramifications of the hypophyseal duct. The functional sig¬
nificance of this morphological pattern is not clear.
The follicle wall consists principally of a layer of large columnar
cells whose outer surfaces are adjacent to the basement membrane.
Their nuclei are either basal in position or centrally located. The
cytoplasm usually contains granules that stain with erythrosine
and orange G although agranular, poorly stained cells are not
1972]
Li and Warner — Alewife in Lake Michigan
215
Figure 3. Portion of rostral pars distalis show¬
ing- follicular arrangement (X216). L, follicular
lumen; N, neurohypophyseal branch; P, prolactin
cells.
uncommon. Substantial evidence from several species of euryhaline
teleosts indicates that these cells produce a prolactin-like hormone
that promotes sodium retention in low saline environments (re¬
viewed by Ball and Baker, 1969). An extremely thin layer of
non-secretory squamous cells lines the follicular lumina and
covers the apical borders of the prolactin cells. This layer is con¬
tinuous with the lining of the hypophyseal duct.
A second secretory cell type occurs in inconspicuous clusters
between the prolactin cell follicles and the neurohypophyseal fiber
tracts in the dorsal part of the rostral pars distalis. The cells are
small and round or polyhedral in shape with central nuclei. Their
cytoplasm is scanty and contains granules that stain weakly with
lead hematoxylin. (Figs. 8 and 9). Similar cells in Poeceilia lati-
pinna and Anguilla anguilla show hyperactivity under the influence
of the adrenocortical inhibitor, metopirone, suggesting that they
are ACTS producing corticotrophs. (Ball and Olivereau, 1966).
The proximal pars distalis is situated ventral and posterior to
the rostral pars distalis. (Fig. 2). It comprises from 20% to 30%
of the adenohypophysis, attaining its greatest size before and
during the spawning period. The cells are arranged in cords or
masses around neurohypophyseal terminations. Three kinds of
cells can be identified.
One of the cell types is distinguished by the presence of fine
orange G-positive granules in the cytoplasm. These cells are rela¬
tively small, round or polyhedral in shape with large spherical
nuclei. They are most concentrated dorsally along the neuro¬
hypophyseal branches but are also scattered throughout the re-
216 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
mainder of the proximal pars distalis. In certain teleosts, changes
in these cells during the normal growth cycle (Olivereau, 1963) or
as a result of experimental alterations in growth (Sage, 1967)
suggest that they are somatotrophs which produce growth hor¬
mone.
A second cell type shows variations closely associated with the
reproductive cycle. These are large round or irregularly shaped
cells with large nuclei (Figs. 4 and 5). They are most abundant
in the centers of cell cords in the posterior and central areas of the
proximal pars distalis. Characteristically, the cytoplasm stains
strongly with the PAS technique and with aldehyde fuchsin, but
under certain circumstances cytoplasmic degranulation and vacuo-
lation are widespread. It is highly probable that these cells are
gonadotrophs since their cyclical activity, and tinctorial reactions
are essentially like those reported for this type of cell in other
teleost studies (Sokol, 1961, Robertson and Wexler, 1962a and b,
Sathyanesan, 1963, Olivereau and Herlant, 1964, Lagios, 1965, and
Olivereau, 1967 and 1969).
Cells of the third type also react positively to PAS and aldehyde
fuchsin, but they differ from gonadotrophs in several other ways.
They are usually smaller and cone-shaped or spindle-shaped with
eccentric nuclei. (Figs. 6 and 7). They are less numerous than
gonadotrophs, and are confined mainly to the anterior ventral
zone of the proximal pars distalis. Cytoplasmic granulas are finer
in texture and cyclical changes in granulation and vacuolation are
less pronounced. Histophysiological studies showing changes in
similar cells under the influence of hypothyroidism and hyperthy-
Figure 4. Pre-spawning gonadotrophs, March
(X2164), g, granulated cell, d, degranulated cell;
v, blood vessel.
1972]
Li and Warner — Alewife in Lake Michigan
217
Figure 5. Post-spawning gonadotrophs, Novem¬
ber (X2164) .
Figure 6. Pre-spawning thyrotrophs, March,
showing partial degranulation (X2164).
roidism suggest that they are thyrotrophs (Atz, 1953, Olivereau,
1954 and 1963,, Barrington and Matty, 1955, Sokol, 1955 and 1961,
Sage, 1967, Brornage and Sage, 1968 and Sage and Bromage, 1970).
The pars intermedia makes up 40% or less of the alewife
adenohypophysis. It is closely associated with the main trunk of
the neurohypophysis in the posterior part of the gland (Fig. 2).
The cells are aggregated in masses around short, broad neuro¬
hypophyseal terminations. Two types of small, faintly acidophilic
cells are recognizable, one type, spherical, with a central nucleus,,
and the other, angular, with an eccentric nucleus. Olivereau (1969)
described two types of pars intermedia cells in Leuciscus rutilis.
218
Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 7. Post-spawning thyrotrophs, November
(X2164).
The functions of these cells are unknown although in fishes as
in other vertebrates the pars intermedia is assumed to produce
the hormone, intermedin.
Cyclical pituitary changes.
Variations in pituitary cytology are evident when glands from
different collection dates are compared. The gonadotrophs, thyro-
trophs and corticotrophs were selected for detailed studies of these
changes. To facilitate description the annual pituitary cycle is
arbitrarily subdivided into three periods related to reproductive
activity. They are the pre-spaivning phase from early January to
mid April, the spawning phase from late April to early August
and the post-spawning phase from mid August to late December.
Annual variations in mean nuclear diameters of gonadotrophs,
thyrotrophs and corticotrophs are shown in Figure 1. In the three
cell types studied, nuclear size is at a maximum during the pre¬
spawning phase in March and at a minmum near the end of the
post-spawning phase in December. The differences between max¬
ima and minima for all cell types are statistically significant.
Gonadotroph nuclear diameters increase to the pre-spawning
peak and then gradually decrease during the remainder of the
year. Nuclear size in thyrotrophs is somewhat more variable. A
decline occurs in April followed by a rise in May, but neither is
statistically significant. Thereafter, mean nuclear diameters de¬
crease with minor fluctuations to the December minimum. Mean
diameters of corticotroph nuclei show the greatest variability.
The annual maximum in March as well as two secondary size
peaks in May and November all are significantly greater than
mean nuclear diameters for preceding and succeeding months.
1972] Li and Warner— Alewife in Lake Michigan 219
The more subjective cytological variations are considered sepa¬
rately for each cell type. The gonadotroph cycle parallels the re¬
productive cycle, but precedes it by several weeks. Maximum cell
size is attained during the pre-spawning phase in March (Fig. 4).
Coarse, intensely stained granules are abundant in the cytoplasm.
Relatively few cells show cytoplasmic degranulation and vacuola-
tion. At the peak of spawning activity during June and July,
gonadotrophs are much more variable. Large heavily granulated
cells, partially degranulated cells and agranular vacuolated cells
may be found closely associated. This variability may indicate
functional differences in individual cells as regards hormone pro¬
duction, storage and depletion. Decreasing size and increasing
degranulation and vacuolation are characteristic of late spawning
and early post-spawning gonadotrophs. Some nuclear pycnosis
is present, but widespread cellular degeneration as noted by Rob¬
ertson and Wexler (19626) in Pacific salmon does not occur. The
post-spawning picture is incomplete because of the lack of Sep¬
tember and October specimens, but during November and Decem¬
ber cell size reaches a minimum. The cytoplasm shows typical
PAS and aldehyde fuchsin staining reactions although granules
are either very small or absent (Fig. 5).
Thyrotroph size is also at a maximum during the pre-spawning
period in March. Most cells are elongated and cone-shaped (Fig. 6) .
Partial cytoplasmic degranulation is widespread. This condition
has often been related in previous pituitary studies to a high level
of thyrotroph activity. During the spawning phase thyrotrophs
vary widely in size, although the characteristic shape is main-
Figure 8. Pre-spawning’ corticotrophs, March,
showing large size, reticular cytoplasm and large
vacuole (X2164).
220 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Figure 9. Post-spawning Corticotrophs, Novem¬
ber (X2164) .
tained. The more elongated cells are about twice the length of the
shorter ones. Cytoplasmic degranulation is rare, indicating a
lower level of activity than during the pre-spawning phase. Unlike
gonadotrophs during the spawning phase, vacuolation does not take
place. Except for a general size decrease, little change is seen in
post-spawning thyrotrophs. (Fig. 7).
As in the other two cell types, the peak of corticotroph activity
occurs during the pre-spawning phase in March. At this time cell
size is at a maximum. (Fig. 8). Nucleoli attain their greatest de¬
gree of prominence. The relatively abundant cytoplasm presents
a reticular appearance suggestive of degranulation. Occasional
very large cytoplasmic vacuoles are present. Spawning and post¬
spawning corticotrophs show decreases in size, nucleolar promi¬
nence and cytoplasmic reticulation and vacuolation (Fig. 9). Sub¬
jective evaluation of corticotroph activity is difficult because of
their small size and sparse granulation. Variation in mean nuclear
diameters over the annual cycle appears to be a more reliable
indicator of the corticotroph functional state.
From the foregoing cytological observations it may be inferred
that high levels of gonadotrophic, thyrotrophic and adrenocorti-
cotrophic hormones are secreted by the alewife pituitary gland just
prior to the spring spawning migration. The expected result is
stimulation of the appropriate target organs, the gonads, thyroids
and interrenals. This is obviously true of the gonads which undergo
marked growth and increased functional activity. Histological evi¬
dence of thyroid stimulation in alewives during the spring was
noted by Hoar (1952) and Boyles (unpublished communication,,
1970. 1 In certain other teleosts, increased thyroid function parallels
1 Dr. Marcia Boyles, Biology Department, Grand Valley State College, Michigan.
1972] Li and Warner— Alewife in Lake Michigan 221
reproductive activity (Berg, et al, 1959, Bromage and Sage, 1968).
There is no published information concerning the alewife inter-
renal but in Pacific salmon, extreme interrenal hyperplasia and
elevated plasma levels of 170H corticosteriods were found during
the spawning migration (Robertson and Wexler, 1959, Hane and
Robertson, 1959). In the alewife and other migratory fishes, the
increased endocrine activity associated with spawning may be
partly an adaptive response to environmental changes encoun¬
tered during their shoreward migration.
The possible effects of hormonal variations on alewife mortality
may now be considered. It seems unlikely that pituitary gonado¬
trophins and gonadal steroids are primarily involved since sexually
immature fish and adults in spawning condition are both abund¬
antly represented in the spring dieoffs (Brown, 1968).
Changing levels of thyroid hormones may be more significant.
Thyroid function in fishes is not well understood, but effects on
osmoregulation, growth and migratory and motor behavior have
been noted (Gorbman, 1969). The original suggestion of Hoar
(1952) that thyroid induced osmoregulatory failure may be a fac¬
tor in alewife mortality should receive further attention.
Changes in interrenal function may be even more pertinent.
Corticosteroids cause electrolyte and fluid shifts in fishes but their
exact roles in normal osmoregulation are not clear. These hor¬
mones also mediate responses to stress. (Chester Jones, et al,
1969). Stanley (1969) found significant shifts of sodium from
plasma to muscle in apparently healthy alewives taken from Lake
Michigan in June and July. In a laboratory study, Stanley and
Colby (1971, in press) demonstrated that cold shock lowered
plasma sodium levels in alewives maintained in fresh water. Holzer
(1971) obtained similar results, and in addition found plasma
sodium depletion and tissue hydration in dying alewives. These
changes may denote partial or complete osmoregulatory failure
possibly related to interrenal insufficiency.
In the present study, there are cytological indications of de¬
creases in thyrotroph and corticotroph activity after pre-spawning
peaks in March. These may signify reduced TSH and ACTH se¬
cretion during the spawning migration. A resulting decline in
thyroid and interrenal function could, therefore, contribute to the
mass alewife mortalities in June and July.
Since firm conclusions cannot be drawn from cytological evidence
alone, it is obvious that endocrine-related physiological data must
be obtained before the role of hormones in alewife mortality can
be fully evaluated.
222 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Summary
Alewives were collected from Lake Michigan during all seasons
to study annual variations in pituitary cytology. The alewife pitui¬
tary gland like those of other isospondylons teleosts has a hypo¬
physeal duct and a follicular type rostral pars distalis. A total of
seven adenohypophyseal cell types were recognized, two in the
rostral pars distalis, three in the proximal pars distalis and two in
the pars intermedia. Detailed studies of gonadotrophs, thyrotrophs
and corticotrophs reveal basically similar annual patterns of
change in secretory activity. Maximal stimulation is indicated in
March just prior to the shoreward spawning migration and mini¬
mal activity occurs in December after the fish have returned to
deep water. Thyrotrophs and corticotrophs are somewhat more
variable in their cyclical patterns than gonadotrophs. Decreasing
thyrotroph and corticotroph activity during the spawning run may
lead to thyroid and interrenal deficiencies that are related to the
annual spring and early summer mortalities.
Acknowledgements
Our appreciation is expressed to the following individuals for
furnishing many of the specimens used in the present study: Dr.
Peter Colby, formerly of the U.S. Bureau of Commercial Fisheries,,
Ann Arbor, Michigan; Dr. Marcia Boyles, Grand Valley State
College, Michigan; Mr. James Miller, Miller Fisheries, Milwaukee
and Mr. Lelond LaFond, LaFond Fisheries, Milwaukee.
Thanks are also extended to Dr. Douglas Dunlop for advice in
photography and to Mr. Alex Tucker for slide preparation.
Authors
Alexander H. H. Li — Ph.D. candidate in Interdisciplinary Endo¬
crinology of Reproduction Program, University of Wisconsin,
Madison. Present Address — Zoology Research Building, University
of Wisconsin, Madison. (Present article based on Mr. Li’s thesis
for the M.S. Degree in Zoology at UWM, completed 1969 under
Dr. Warner’s supervision.)
Eldon D. Warner - — Professor of Zoology, University of Wiscon¬
sin — Milwaukee.
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Atz, E. H. 1953. Experimental differentiation of basophil cell types in the
transitional lobe of the pituitary of a teleost fish, Astyanax mexicanus.
Bull. Bingham Oceanogr. Coll. (Yale), 14: 94-116.
Ball, J. N. and B. I. Baker. 1969. The pituitary gland: anatomy and histo-
physiology. In “Fish Physiology” (W. S. Hoar and D. J. Randall, eds.).
Vol. II, pp. 1-110 (Academic Press, New York).
1972]
Li and Warner — Aleivife in Lake Michigan
223
Ball, J. N. and M. Olivereau. 1966. Experimental identification of ACTH
cells in the pituitary of two teleosts, Poecilia latipinna and Anguilla an -
guilla. Correlated changes in the interrenal and in the pars distalis re¬
sulting from administration of metopirone (SU4885), Gen, and Comp.
Endo., 6, 15-18.
Barrington, E. J. W. and A. J. Matty. 1955. The identification of thyro-
trophin-secreting cells in the pituitary gland of the minnow (Phoxinus
phoxinus). Quart. J. Micr. Sci., 96: 193-201.
Berg, 0., A. Gorbman and H. Kobayashi. 1959. The thyroid hormones in
invertebrates and lower vertebrates. In “Comparative Endocrinology”
(A. Gorbman, ed.) pp. 307-413 (John Wiley and Sons, Inc., New York).
Bromage, N. R. and M. Sage. 1968. The activity of the thyroid gland of Poe¬
cilia during the gestation cycle. J. Endocrinol. 41: 303-311.
Brown, E. H. 1968. Population characteristics and physical condition of ale-
wives, Alosa pseudoharengus, in a massive die-off in Lake Michigan, 1967.
Great Lakes Fishery Commission. Technical Report #13.
Buchmann, H. 1940. Hypophyse and Thyroidea im Indiviualzyklus des Her-
ings. Zool. Jb., Abt. 2 Anat. Ontog., 66: 191-262.
Chester Jones, I., D. K. O. Chan, I. W. Henderson, and J. N. Ball. 1969.
The Adrenocortical steroids, adrenocorticotropin and the corpuscles of
Stannius. In “Fish Physiology” (W. S. Hoar and D. J. Randall, eds.) .
Vol. II (Academic Press, New York).
Elftman, H. 1959a. Combined aldehyde fuchsin and PAS staining of the pitu¬
itary. Stain Technology, 34: 77-80.
Elftman, H. 1959b. Aldehyde-fuchsin for pituitary cytochemistry. J. of His-
tochem., 7 : 98-100.
Go mori, G. 1950. Aldehyde-fuchsin: a new stain for elastic tissues. Am. J. of
Clinical Pathology, 20: 665-666.
Gorbman, A. 1969. Thyroid function and its control in fishes. In “Fish Physi¬
ology” (W. S. Hoar and D. J. Randall, eds.). Vol. II, pp. 241-274. (Aca¬
demic Press, New York).
Hane, S. and O. H. Robertson. 1959. Changes in plasma 17-hydroxycorticoids
accompanying sexual maturation and spawning of the pacific salmon
( Oncorhynchus tschawytscha) and rainbow trout (Salma gairdnerii) . Proc.
Nat. Acad. Sci., 45 (6) : 886-893.
Herlant, M. 1960. Etude critique de deux techniques nouvelles destinees a
mettre en evidence les differentes categories cellulaires presente dans la
glande pituitaire. Bull. Micr. Appl., 10 (3) : 37-43.
Hoar, W. S. 1952. Thyroid function in some anadromous and landlocked tele¬
osts. Trans. Roy. Soc. Can., 46 : 39-53.
Holzer, J. 1971. The effects of thermal shock, disease and morbidity on elec¬
trolyte metabolism in the alewife. Unpublished M.S. thesis, University
of Wisconsin — Milwaukee.
Lagios, M. D. 1965. Seasonal changes in the cytology of the adenohypophysis,
testes and ovaries of the black surf perch ( Embiotica jacksoni), a vivip¬
arous percomorph fish. Gen. and Comp. Endo., 5: 207-221.
MacConaill. 1956. In “A Practical Manual of Medical and Biological Stain¬
ing Techniques,” by E. Gurr, ( Interscientific Publishers, Inc., New York).
McMannus, J. F. A. and R. W. Mowry. 1958. Effects of fixation on carbohy¬
drate histochemistry. J. of Histochem. and Cytochem., 6: 309-316.
Olivereau, M. 1954. Hypophyse et glande thyroide chez les poissons. Etude
histophysiologique de quelques correlations endocriennes, en particular
chez Salmo salar. L’Ann. Inst. Oceanogr., 29: 95-296.
224 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Olivereau, M. 1963. Cytophysiologie du lobe distal de l’hypophyse des Agna-
thes et des poissons, a l’exclusive de cells concernat la fonction gonado-
trope. Colloques Internationaux du Centre National de la Recherche Sci-
entifique No. 128 Cytologie de FAdenohypophyse. (J. Benoit and C.
de Lage).
Olivereau, M. 1967. Observations sur l’hypophyse de l’anguille femelle, en
particular lors de la maturation sexuelle. Z. Zellforsch. mikr. Anat., 80:
286-306.
Olivereau, M. 1969. Donnees cytologiques sur Fadenohyphphyse du gardon,
poisson teleosteen. Gen. and Comp. Endo., 12: 378-384.
Pickford, G. E. and J. W. Atz. 1957. The Physiology of the Pituitary Gland
of Fishes. New York Zool. Soc., New York.
Robertson, O. H. and B. C. Wexler. 1959. Hyperplasia of adrenal cortical
tissue in pacific salmon (Genus Oncorhynclius ) and rainbow trout (Salmo
gairdnerii) accompanying sexual maturation and spawning. Endocrinol.
65: 225-238.
Robertson, O. H. and B. C. Wexler. 1962a. Histological changes in the pitui¬
tary gland of the rainbow trout (Salmo gairdnerii) accompanying sexual
maturation and spawning. J. Morphol., 110: 157-169.
Robertson, 0. H. and B. C. Wexler. 1962b. Histological changes in the pitui¬
tary gland of the pacific salmon (Genus Oncorhynclius) . J. Morphol., 110:
171-185.
Sage, M. 1967. Responses of pituitary cells of Poecilia to changes in growth
induced by thyroxine and thiourea. Gen. and Comp. Endo., 8: 314-319.
Sage, M. and N. R. Bromage. 1970. The activity of the pituitary cells of the
teleost, Poecilia, during the gestation cycle and the control of the gonado¬
tropic cells. Gen. and Comp. Endo., 14: 127-136.
Sathyanesan, A. G. 1963. Histologic changes in the pituitary and their cor¬
relation with the gonadal cycle in some teleosts. La Cellule, Tome 63:
283-290.
Sokol, H. W. 1955. Experimental demonstration of thyrotropic and gonado¬
tropic activity in the adenohypophysis of the guppy, Lebistes reticularis.
Anat. Rec., 122: 451.
Sokol, H. W. 1961. Cytologic changes in the teleost pituitary gland associated
with the reproductive cycle. J. Morphol., 109: 219-235.
Stanley, J. G. 1969. Seasonal changes in the electrolyte metabolism in the
alewife, Alosa pseudoharengus, in Lake Michigan. Proc. 12th Conf. Great
Lakes Res. 91-97.
Stanley, J. G. and P. J. Colby. 1971. Effects of temperature on electrolyte
balance and osmoregulation in the alewife (Alosa pseudoharengus) in
fresh and sea water. Trans. Am Fish. Soc. (in press).
OVIPOSITIONAL SITE PREFERENCES OF THE OAK
DEFOLIATING GRASSHOPPER, DENDROTETTIX
QUERCUS,1 IN WISCONSIN2'3
Douglas A. Valek and Harry C. Coppel
Nymphs and adults of Dendrotettix quercus Packard feed largely
on oak foliage (Valek and Coppel, 1971). The species has a 2 year
life cycle in Wisconsin with most individuals in a particular popu¬
lation appearing during the same summer. Complete defoliation
occurs occasionally, but tree mortality is infrequent. Previous
observations indicated that most of the grasshopper damage oc¬
curred near the area of eclosion. Bruner (1887) reported, indi¬
rectly, that D. quercus egg pods, in Texas, were “deposited in the
ground about the bases of the trees or indifferently scattered about
the surface among the decaying leaves.” The purpose of this study
was to characterize the sites preferred for oviposition and thereby
aid in the determination of sites predisposed to oak defoliation.
Emphasis was placed upon the relationship of light and amount
of non-woody organic matter to oviposition.
Methods and Materials
The study was conducted in Jackson County, Wisconsin, in 1969,
on a site with a large D. quercus population. The study area was
sandy and covered mainly by the sedge Carex pensylvanica Lam.
and 15-20 ft. black and northern pin oak ( Quercus velutina Lam.
and Q. ellipsoidalis E. J. Hill) with occasional quaking aspen
(Populus tremuloides Michx.), white oak ( Q . alba L.), and red pine
( Pinus resinosa Ait.). A sandy surfaced road passed through the
study area. The road had 20-30 ft. margins which contained 3-6 ft.
coppice black and northern pin oaks. Four parallel lines of sample
units were established on the ground, crossing the road perpendicu¬
larly. Each line contained 33 sample units, each 1 ft. square, 4 ft.
apart, center to center. The center of each sample unit was marked
by a stake in the ground. The lines were randomly spaced 45-60
1 Orthoptera : Acrididae.
2 Research supported by the College of Agricultural and Life Sciences, University
of Wisconsin, Madison, and in part by the Wisconsin Department of Natural Resources
and the University of Wisconsin Research Committee of the Graduate School with
funds supplied by the Wisconsin Alumni Research Foundation. Received for publica¬
tion January 13, 1972.
3 Part of a thesis submitted by the senior author in partial fulfillment of the re¬
quirements for the Ph.D. degree in Entomology at the University of Wisconsin.
225
226 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
ft. apart. The beginning of each line was arbitrarily set in regard
to its distance from the road, but each line was crossed by the road
near its center.
Light intensity measurements were taken over each sample unit
during September 18-21, when most oviposition had already oc¬
curred. A Weston Illumination Meter, Model 756 (Weston Electric
Instrument Co., Newark, N.J.) was used. The mean of 3 light
intensity measurements, randomly taken over each sample unit,
was recorded. Light intensity under bracken fern, raspberry, or
other plants taller than the average herbaceous cover was meas¬
ured, but the light meter was held high enough to minimize the
influence of herbaceous plants directly over the meter’s sensor.
One mean light intensity measurement was obtained over each
sample unit in early morning, late morning, early afternoon, and
late afternoon. The time necessary to take readings at each point
plus the traveling time through the entire study area caused a
variation in the measurements due to changes in ambient light
with passage of time. This was especially apparent in the early
morning and later afternoon readings. Therefore, a measurement
of ambient light intensity near the ground was made in the center
of the road, near the middle of each line, when that point was
reached during the operation. These 4 measurements served as
standards to which the mean measurements in each line were cor¬
rected by the factor that its standard varied from the mean of the
4 standards for each general day period. The combined means of
the light intensity measurements taken at the 4 general day pe¬
riods, in their corrected forms, produced light intensity values for
each sample unit which were compared with the others. The term,
‘flight intensity index”, described the final light intensity value
derived from each sample unit.
When oviposition ceased, but before appreciable numbers of
leaves had fallen, the sod units with all vegetation and other non-
woody organic matter, in situ, were cut from the ground. Each
unit was removed by cutting around a square foot wood template
with a flat shovel and placed into a large plastic bag for transpor¬
tation and storage. In the laboratory, the vegetation on each
sample unit was removed by first scraping off loose material such
as decaying leaves or grass. Rooted plants were clipped at the soil
line. The remaining material, which consisted largely of plant rem¬
nants, fungal hyphae, and humus, was swept from the surface of
the sample unit with a stiff, long-bristled brush until only mineral
soil and small roots remained. The soil was examined for egg pods
by cutting the square into 6-8 strips with a heavy knife and tear¬
ing the strips into pieces small enough to allow detection of the
pods when squeezed with the fingers. Plant material removed from
1972] Valek and Coppel—Oak Defoliating Grasshopper 227
the samples was dried at room temperature for 3 or more days.
Most of the sand was removed from the dry organic matter with
cold water and detergent. The sand settled to the bottom of the
wash container as the material was squeezed and stirred by hand.
Floating organic matter was strained from the water and drained.
All woody plant parts such as twigs over 0. 2 inch in diameter
and current year’s acorns were arbitrarily removed. The wet ma¬
terial was dried at 60 °C for a minmum of 4 days before it was
weighed.
Results
No egg pods were found in the 20 sample units taken from the
nearly barren sand road. It was felt that the information gained
from the analysis of the data on this relatively unimportant situa¬
tion was not particularly useful and the extreme effect of high
values in light intensity and low values in ground cover on the total
analysis warranted the deletion of these units. It may be argued
that sample units taken from the road had no pods because they
were located the maximum distance from the grasshopper’s food
sources. However, adult grasshoppers commonly traversed the road
and there was sufficient opportunity for oviposition there.
In the remaining 112 sample units, 37 contained 1 or more egg
pods each. There was a significant difference between the light
intensity indices of those sample units containing egg pods and
those not containing pods (99% level, chi square in a 2 X r con¬
tingency table, Steel and Torrie, 1960). The mean light intensity
index of the sample units with at least 1 egg pod was 2149 ± 1630
(SD) foot-candles and it was 775 ± 1056 ft.-c. for the units with¬
out pods.
Egg pod density was highest in the highest light intensity index
classes (Fig. 1). The number of pods per sample unit remained
below 2.0 pods/sq. ft. in the 3 lowest light intensity index classes,
but increased to 6.78 and 4.62 pods/sq. ft .in the 2 highest classes.
The relationship of non-woody organic matter to oviposition ac¬
tivity revealed a significant difference in the amount of organic
matter between those sample units with at least 1 egg pod and
those with none (99% level, chi square in a 2 X r contingency ta¬
ble, Steel and Torrie, 1960). The mean dry weight of organic
matter on the 37 sample units with at least 1 egg pod was 60.7 ±
21.8 g., whereas on those without pods it was 70.0 ± 17.1 g.
The highest mean density of egg pods was found generally in the
median weight classes (Fig. 2). Since the results from the 20 sam¬
ple units from the road were not considered, the length of the bar
in the 0-9 g. weight class in the figure reflects the influence of the
228
Wisconsin Academy of Sciences, Arts and Letters [VoL 60
LIGHT INTENSITY INDEX CLASS ( FT.- CANDLES )
* NO. SAMPLE UNITS IN CLASS
Figure 1. The relationship between the light intensity index class and the mean
number of D. quercus egg pods per sq. ft.
DRY WT. ORGANIC MATTER CLASS (GRAMS)
*NO. SAMPLE UNITS IN CLASS
Figure 2. The relationship between the dry weight of organic matter class and
the mean number of D. quercus egg pods per sq. ft.
1972]
Valek and Coppel — Oak Defoliating Grasshopper 229
Figure 3. A profile of the spatial distribution of organic matter, light inten¬
sity, and D. quercus egg pods.
unusual case of 19 pods in 1 of the 2 units falling into the class.
No explanation can be given for the large amount of oviposition
in this area. Pods were sometimes concentrated in small areas of
soft soil caused by burrowing animals, but it is not known if such
a disturbance occurred in this instance.
The spatial relationship of the egg pod density to the distribution
of sample units of varying light intensity indices and organic
matter weight is presented as a profile (Fig. 3) where the values
of the sample lines have been combined. The 5 sample units from
each line falling in the road have been placed together to align
the sample lines and the running means of the columns of sample
units have been used for the plotted values. The plotted values of
the light intensity indices in the figure show that the light intensity
was the lowest at the left and right margins of the profile, corre¬
sponding to the light reaching near soil level under the large trees,
and highest toward the center of the profile, corresponding to the
light reaching the soil in the open area. The dry weight of non-
woody organic matter appeared uniform at both margins of the
profile, corresponding to the areas near or under the tall trees, but
fell uniformly to 0 in the center of the road. There was no rela¬
tionship between the light intensity index and dry weight of
organic matter (regression analysis, 95% level). Density of egg
pods was greatest in the areas between the tall trees and the road.
This distribution of egg pods is in accord with previous observa-
230 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
tions by the authors in which pods and emerging nymphs were
most commonly encountered near the margins of woods.
The highly attractive nature of certain small areas to the ovi¬
positing females indicated that the measurement of light intensity,
and especially the amount of organic matter covering the soil, could
probably be conducted more profitably with a sample unit of 0.67
or 0.50 sq. ft.
References Cited
Bruner, L. 1887. Report on locusts in Texas during the spring of 1886. U. S.
Dep. Agr. Div. Entomol. Bull 13 : 9-19.
Valek, D. A. and H. C. Coppel. 1971. Bionomics of an oak defoliating grass¬
hopper, Dendotettix quercus, in Wisconsin. Manuscript submitted to the
Annals, Entomol. Soc. Amer. May 1971.
Steel, R. G. D. and J. H. Torrie. 1960. Principles and procedures of statis¬
tics. McGraw-Hill. New York. 481 p.
WILD RIVERS OF NORTHEASTERN WISCONSIN
(WILD RIVERS COOPERATIVE RESEARCH PROJECT)
George Becker
The birth of the wild river concept in Wisconsin must surely
go back many years. Perhaps the best-known elegy came from
Aldo Leopold who in 1943 wrote the essay “Flambeau — the Story
of a Wild River. ” Modern-day Wisconsin voyageurs like Joe Mills
and J. J. Werner of the John Muir Chapter (Sierra Club) probed
the white waters of the state with their canoes. They made mental
notes of unusually primitive waters and began talking up wild
rivers.
During the early 1960’s the Wisconsin Conservation Depart¬
ment fought against proposed construction of dams on the Wolf
and Popple rivers in northeastern Wisconsin. Perhaps the biggest
thrust was made by Walter E. Scott, then administrative assistant
to the director. In 1964 Walter (as his many friends call him)
delivered an address in Madison entitled “Preserving Wisconsin’s
Wild Rivers.” He summarized clearly the outlook for wild rivers
as being both “bitter and sweet,” as having “great possibilities
as well as serious setbacks and failures.”
When he became President of the Wisconsin Academy of Sci¬
ences, Arts and Letters (1964-65), Walter proposed that the
Academy initiate a research program on Wisconsin’s wild rivers.
There were two reasons for this. First he recalled that in the
early 1940’s the Conservation Department made a series of basic
studies on the famous Brule River in northwestern Wisconsin.
These studies appeared in installments over several issues of the
Transactions. They have since become definitive and useful refer¬
ences. Second, a wild river needs basic research which can be shown
to the State Legislature. This is one of the paradoxes of human
nature — secrets must be unlocked before the organism, in this
case the river, is allowed to keep its secrets. The bill for setting
aside the Pine, Popple and Pike rivers of northeastern Wisconsin
was already in the legislative hopper. A going program of research
on these streams would hopefully influence the Legislature to pass
the bill.
1 Paper No. 1 in a series, “Studies on the Pine-Popple Wild Rivers Area of North¬
eastern Wisconsin”, which will appear in this and succeeding- issues of the Trans¬
actions of the Wisconsin Academy. As an account of the history, objectives and
development of the project it is written, quite properly, by Professor George Becker,
who as its coordinator was largely responsible for its organization, progress and super¬
vision. — Lowell E. Noland, editor.
233
234 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Early in 1965 Walter asked me to assume organization of a
wild river study for the Pine, Popple, Pike and Wolf rivers. I
accepted the task with some misgivings. There were no funds at
hand for getting such a program under way, nor did there appear
to be much opportunity for getting financial aid. Also the people
we thought most likely to perform the research were already
actively engaged in other research and publishing. Could they
be persuaded to turn time, effort and money in this direction?
I spent several days talking to these people. They were en¬
thusiastic about the wild rivers of northeastern Wisconsin and
about the mysteries to be unravelled. On October 16, 1965, the
organizational meeting was held at the Hill State Office Building
in Madison. Present were James Anthony, Robert Dicke, William
Dickinson, Lewis Posekany, Edward Schneberger, Walter Scott,
Howard Young, Stan Welsh and James Zimmerman. The com¬
mittee decided to complete its report over a five-year period,
culminating with the Academy Centennial in 1970. Plans were to
collect all information into a bound book which would be dis¬
tributed to members of the Academy and to other interested
agencies. During later deliberations the committee voted to direct
its research primarily toward the Pine and Popple basins, leaving
the Pike and Wolf basins until after the first phase was completed.
The Wild Rivers bill became law in November, 1965. It set up a
program for the preservation of the Pine, Pike and Popple rivers
in Florence, Forest and Marinette counties. It designated the Con¬
servation Commission to provide leadership in the development
of a practical management policy. Late in 1965 the Wisconsin
Society for Ornithology, Inc., donated to the Wisconsin Academy
a sum of $2500 for the study of birds in the wild rivers area.
Perhaps the most memorable meeting of the Wild Rivers Coop¬
erative Research Project took place at the Trees for Tomorrow
Camp at Eagle River, October 22-23, 1967. Arranged by Art
Oehmcke, it was designed to show members of the committee and
their families the beauties and the scars of the Pine and Popple
rivers. Speaker for the occasion was Philip Archibald, then Forest
Supervisor, Nicolet National Forest, who discussed “The U. S. For¬
est Service and Its Management of Wild Rivers.” The text of this
paper appeared in the 1966 Fall-Winter issue of the Wisconsin
Academy Review, pp. 77-80.
At the very start of our research it became apparent that ad¬
verse activities were going on in the basins of the Pine and Popple.
Some of this activity was initiated by individuals and towns who
feared that unless “improvements” were made immediately, wild
river policy restrictions, which were in the process of being de-
1972] Becker — Wild Rivers of Northeastern Wisconsin 235
veloped by state and federal agencies, would forbid the desired
“improvements.”
These conflicting activities were called to my attention by Art
Oehmcke, then area supervisor with the Conservation Department
at Woodruff. Late in 1966, with permission from the Academy
Council, I named the Wild Rivers Advisory Policy Subcommittee,
with Oehmcke as chairman. Members appointed were Phil Archi¬
bald, Joe Mills and Calvin Erickson. This committee would be
advisory to the Conservation Department’s wild rivers policy com¬
mittee, but its main role would be that of watchdog. It would
attempt to forestall any possible activities which appeared to be
detrimental to the wild river program.
During the subsequent months, conflicting encroachments within
the wild rivers basins were observed and appropriate action was
taken. We are grateful to this sub-committee for its vigilance,
which preserved a number of wild features that would otherwise
have been lost.
It was a forester, Aldo Leopold, who said “The best way to
manage a wild river is to let it be.” Because sectors of the newly-
named wild rivers are used for many purposes, the Wisconsin
Department of Natural Resources in its policy statement has
established a zoning system, allowing considerable man-use in
some sectors. The U. S. Forest Service has developed yet another
plan which, even within its “wild river” zones, feels man-directed
embellishments are permissible. For instance, mature timber to a
forester demands cutting, and, keeping things “wild” means proper
landscaping.
I wonder, for instance, whether a lightning fire will be allowed
to run its course or whether an insect infestation will avoid treat¬
ment? I wonder if the down-tree in the water must really be re¬
moved to make easy passage for the canoer, or whether the stabili¬
zation of naturally-eroding stream banks should be “top priority
work within the zone?”
I personally take a dim view of a wild river program which
prohibits damming of the main stem but makes no similar provi¬
sion for its life-giving tributaries. These are but a few of the many
objections which may be raised against “management” of our wild
rivers.
Men are of many persuasions; men tend to relate the concept
of wilderness to their own training and interests. Unfortunately
the present state and federal criteria allow for considerable en¬
croachment on the wildness of the area. It is my hope that our
experts will soon come to the concept that “the best way to man¬
age a wild river is to let it be,” and that wilderness is its own
master.
236 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
During the course of committee activity several reports were
published and, in a sense, belong to the series which follows.
Among these are: Olson, Gerald W. and Francis D. Hole “The
Fragipan in Soils of Northeastern Wisconsin,” Trans. Wis. Acad.
Sci., Arts and Letters, 56 (1967-68), pp. 173-184; and M'ason,
John W. and Gerald D. Wegner “Wild Rivers Fish Populations
(Pine, Popple and Pike Rivers)” Dept, of Nat. Resources, Madi¬
son, Wis., Research Report 35 (1970), 42 pp.
The following topics and prospective authors constitute the Wild
Rivers series. This list is not arranged according to order of publi¬
cation ; nor is there assurance that all of these topics will appear.
At this date a number of manuscripts have been received and are
indicated by asterisks before the names of the authors.
Soils — *Prof. Francis Hole, Soils Dept., U.W., and Director of
Soils Survey, Wis. Geol. & Nat. Hist. Survey, Madison (Co¬
author of another paper published in 1968 and mentioned
above) .
Water, resource planning and management — C. L. R. Holt, District
Chief, U. S. Geol. Survey, Madison; Ed Oakes, Hydrologist,
U. S. Geol. Survey, Madison; * Gerald L. Paul, Chief Hydrolo¬
gist, Northeastern Wis. Regional Planning Comm., Appleton.
Inventory of surface waters — *C. W. Threinen, Administrative
Assistant, Wis. Dept, of Nat. Resources, Madison.
Vascular plants — Prof. S. Galen Smith, Dept, of Biology, Wis.
State Univ., Whitewater; Prof. Hugh litis, Dept, of Botany,
Univ. of Wis., Madison; Dr. James Zimmerman, Naturalist,
Univ. of Wis. Arboretum, Madison.
Non- vascular plants — * James A. Jesberger, Dept, of Biology, Univ.
of Saskatchewan, Saskatoon.
Forest resources — Robert Train, Supervisor, Timber Management
Staff Officer, Nicolet National Forest, Rhinelander.
Aquatic insects — *Prof. William Hilsenhoff, Dept, of Entomology,
Univ. of Wis., Madison.
Fish parasites — Prof. James D. Anthony, Dept, of Zoology, Univ.
of Wis., Milwaukee.
Fish populations studies — Jack Mason, Biologist, Wis. Dept, of
Nat. Resources, Madison. (Paper published in 1970 and men¬
tioned above).
Distributional list of fishes — Prof. George Becker, Dept, of Biology,
Wis. State Univ., Stevens Point.
Amphibians and reptiles — *Dr. William E. Dickinson, Curator of
Lower Zoology, Milwaukee Public Museum.
1972] Becker — Wild Rivers of Northeastern Wisconsin 237
Mammals — *Prof. Robert McCabe, Chmn. Dept, of Wildlife Ecol¬
ogy, Univ. of Wis., Madison.
Birds-— * Prof. Howard Young, Dept, of Biology, Wis. State Univ.,
La Crosse.
Wild rivers — *Joe Mills, Wild Rivers Chmn., Izaak Walton League
of America, and John Muir Chapter of Sierra Club, Ripon.
History — -John Winn, Field Representative, Office of Field Serv¬
ices, State Historical Society of Wisconsin, Madison.
Maps and mapping (historical) — Walter E. Scott, Asst, to the
Deputy Secretary, Wis. Dept, of Nat. Resources, Madison.
Aboriginal occupants — *Prof. Robert Salzer, Logan Museum of
Anthropology, Beloit College.
Literature and arts — Prof. Robert E. Gard, Director, Wisconsin
Idea Theater, Univ. of Wis., Madison.
Climatology— -Hans Rosendal, Weather Bureau Wisconsin State
Climatologist, Madison.
Economic development — Pres. Walter Peterson, Univ. of Dubuque.
Development of a wild rivers policy — * Arthur A. Oehmcke, Asst.
Dir. of the Bur. of Fish Management, Dept, of Nat. Resources,
Madison.
Limitations imposed by zoning ordinances — Calvin Erickson, Edi¬
tor, Florence County Mining News, Florence.
Case for public ownership of wild river stream banks — *John Chaf¬
fin, Forest Supervisor, Nicolet National Forest, Rhinelander.
In addition to the above I wish to recognize the following for
their many contributions : Perry Olcott, Lewis Posekany, Ed
Schneberger, Lyle Christenson, Lloyd Andrews, Eunice Bonow,
Steve Field, Larry Seeger, and George F. Hanson; and, if special
thanks are allowed, I wish to acknowledge the following for their
steadfast encouragement and assistance: Walter E. Scott, Arthur
A. Oehmcke, David J. Behling (Acad. Pres. 1966-67), John W.
Thomson (Acad. Pres. 1967-68), Norman Olson (Acad. Pres.
1970-71), Jack Arndt (Editor, Wis. Acad. Review 1964-67).
I speak in behalf of the entire committee in expressing gratitude
to Professor Lowell E. Noland (Academy President 1946-47) who
has consented to edit this wild rivers series.
Finally, as an indication of the psychological effect of the Pine-
Popple wild rivers region on those who have spent some time there,
I submit the following poem written about 1930 by James M. Wood¬
man, then a sports writer for the Chicago Tribune, and made avail¬
able to me through the kind offices of Prof. L. G. Sorden, of the
University of Wisconsin.
238 Wisconsin Academy of Sciences , Arts and Letters [VoL SO
Where the Popple Joins the Pine
Far away from all. the glitter of the busy city's life,
Where calm contentment drives away all worldly grief and strife,
Where the melody of songbirds lulls a fellow's soul to rest
When the slanting shadows greet him as the sun sinks in the west — ■
Tis a spot that Nature moulded in a manner most divine,
Just a place of matchless beauty — Where the Popple joins the Pine.
There is music when the water ripples o'er the polished stones;
There is sadness when the balsam bows before the gale and moans;
And my heart leaps wild with rapture when I roam along the streams
Living o’er once more my boyhood in a mass of daylight dreams.
So I snuggle close to Nature claiming all her charms for mine
In that place of tranquil splendor — Where the Popple joins the Pine.
There I gaze upon the glory of the river’s mirrored sky
And the magic of the boulders where the speckled beauties lie.
I can hear the partridge drumming to his faithful feathered mate —
Oh, it fills my heart with gladness and it drives away all hate
As I loiter in the shadows with my rod and reel and line,
Courting Nature in her homeland — Where the Popple joins the Pine.
When my brow by Time is furrowed and my hair grows silver white,
When my eyes are dimmed though eager for a never failing light,
When the Lord who in His wisdom sends a summons unto me,
When I leave this earthly turmoil for a Land-that-is-to-be,
I would lie forever sleeping where the sun and stars may shine
Through the branches of the hemlocks — Where the Popple joins the Pine.
(Written for and dedicated to my friend Oscar Franknecht, whose beautiful
home occupies a most inspiring position where the Popple joins the Pine. —
James M. Woodman.)
CANOEING THE WILD RIVERS PINE AND POPPLE
Joe Mills1 2 3
Captain Jefferson Cram, of the U.S. Topographic Engineers, was
among the first explorers to describe the Pine River. In his detailed
report of 1840s on the Michigan-Wisconsin Boundary Survey, he
wrote :
The tributary of the Menominee called the Mus-kos Se-pe, is so low in
summer as to be unnavigable for any but the smallest of canoes, and in
some seasons it is almost dry. . . . The valley of this river is long, and
contains deer in great abundance; and consequently, much resorted to by
Indians .... for the winter hunt. This river is called by some Pine River.
Captain Cram’s impression of the country was far from favor¬
able:
The country .... has an exceedingly desolate appearance; all the timber
which was once pine has been consumed by fire, as far as the eye can
reach all around on every side. The prospect is one of a broken land¬
scape of barren hills, studded here and there with scarred pine stubs,
with scarcely a living tree, except the second growth of white birch and
poplar.
Making his observations during late summer and viewing the
Pine River from the low profile of his canoe midstream on the
Menominee, Captain Cram’s conclusions were quite accurate, but
far from complete. Hidden from his vision behind the first bend
was a beautifully wild river, serene in its quiet stretches, boisterous
as it dashed over falls and rapids, coursing along through a verdant
forest.
Chippewa Indians inhabited the area. Those occupying the burned
and barren district were referred to as the Badwater Indians.
The stretch of the Menominee River here was known as “bad-
water”. The Indians grew only potatoes, as it was too far north for
the growing of corn. To the west at the headwaters of the Pine
and its main tributary, the Popple, lay a vast region of forest and
1 This is Paper No. 2 in the series “Studies on the Pine-Popple Wild Rivers Area of
Northeastern Wisconsin”. It is included at the beginning- of the sequence because it
gives a vivid picture of the two rivers and adjacent land, as seen by a canoeist
traversing their lengths.
2 Mr. Joe Mills (688 Gary St., Ripon, Wis. 54971) is an enthusiastic member of the
Sierra Club, and an honorary trustee of the Wisconsin Natural Resources Foundation.
— Editor.
3 Cram, Captain T. J., 1840. Report on the survey of the boundary between the
State of Michigan and the territory of Wiskonsin. U. S. Senate Document 151, 26th
Congress, 2nd session.
239
240 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
swamp. Much of the land was a flat pinery dotted with stands of
mixed hardwoods. The swamps consisted of open bogs edged with
cedar, tamarack, spruce and balsam. Deer were abundant, yarding
in the thick cedar swamps to escape the heavy snows of winter.
It was here that Chief Ca-sha-o-sha came with his band from their
summer planting ground on Lake Vieux Desert for the hunt that
was to supply them with meat for survival through the cold
Wisconsin winters. From the Brule River they pushed their canoes
south up Elvoy, Brule and Alvin Creeks. A short quarter-mile
portage from the headwater springs of Alvin Creek put them on
the North Branch of the Pine River. The western terminus of the
portage trail was at the present location of the Forest Service
canoe landing northwest of Windsor Dam Campground.
I began my exploration of the Pine River there (Fig. 1,1) on a
bright summer day in 1963, with Nancy, my teen-age daughter in
the bow of our light-weight aluminum canoe. I shoved hard, step¬
ping lightly as I felt the keel leave the landing. We dug deeply
into the sluggish current to gain momentum. Just around the point
of land below the landing we found ourselves in the midst of a
collection of loose-fitting logs. However, we had no problem extri¬
cating ourselves, as the logs moved easily from the pressure of
our paddles. Out on the open river, zigzagging through an open
swamp surrounded with white birch, popple, balsam and spruce,
we settled back to enjoy the scenery. As we rounded a bend, sud¬
denly a large boulder reared its dark form dangerously close to
the bow. A rapid swish of the paddle and we averted a collision
with only an inch to spare. Alert now, we cautiously avoided the
rocks that appeared regularly ahead of us as we proceeded. A
creek came in on the right, then the marsh broadened : we were in
the flowage area of Windsor Dam (Fig. 1, 2).
Built in 1891 for the purpose of facilitating the driving of logs,
Windsor suffered the fate of all logging dams — abandonment and
gradual deterioration. Later the fill was utilized in the approaches
to the bridge for a public road, now designated as Forest Road
2174. Some of the original dam timbers can still be found imbedded
in the river under the bridge.
Two other logging dams were constructed on the North Branch
of the Pine. Gillett Dam, built a half mile below the entrance of the
Lake Howell outlet (Fig. 1,3), occupied the open swamp in Sections
24 and 25. A mile downriver was Stones Dam.
The Pine River, as it flows from Butternut Lake, is too shallow
and badly clogged with windfalls to be floated in a canoe. Further¬
more, access is difficult. However in August 1967 Dr. Galen Smith,
professor of biology at Whitewater State University, managed to
get his canoe to the river over an abandoned logging-railroad grade.
1972] Mills — Canoeing the Wild Rivers Pine and Popple
241
242 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
He embarked from a crude bridge in Section 18, and found the
river meandering a great deal, but with water of sufficient depth
for good paddling.4
Gary Werner, a University of Wisconsin student, with two
companions, paddled a canoe upriver from the Forest Service land¬
ing, in April 1966, as far as the Section 18-19 line. A minor log
jam, several beaver dams and shallows in the vicinity of the Jeep
trail across Section 17 hampered their travel. While the experiences
of Werner and Smith imply that at least two miles of the upper
North Branch are canoeable, the best choice for the beginning of a
canoe trip is the Forest Service canoe landing, with the first portage
at Windsor Dam.
All that is necessary at Windsor Dam is to slide the canoe under
the bridge into the pool below. A small island obstructs the outlet.
Immediately beyond, the river bottom becomes very rocky. Unless
the river is extremely high, it is utterly impossible to paddle a
loaded canoe. Wading is not difficult as the current is slow, posing
no threat to canoe or contents. This condition persists for a half
mile. Finally the banks flatten into an alder swamp and the river
deepens. From this point on we enjoyed good canoeing for about a
mile and a half. Once we caught a glimpse of an otter. Twice later
we sighted the animal and attempted to catch up with it as it
swam downriver. As we entered big timber, rocks began ripping
the surface of the river, and from ahead came the sounds of a
rapids. We got as far as the logging-road bridge A clearing fifty
yards north of this is the location of a logging camp. The road on
the other side of the river continues into the timber inviting further
exploration. The residue of a campfire tells us that this is an ideal
campsite.
Wading the canoe through the rocks, we find the river turning
southward and improving. Once more we clamber back into our
seats, and proceeding onward pass dense thickets of balsam and
spruce towered over by an occasional white pine. The remains of
another logging bridge slip by. Three huge boulders, probably
rolled there by loggers clearing the channel many years ago, doze
in the sun. Then one of us points to the spinning propeller of a
windcharger over the tops of balsams ahead of the canoe. What
a shocking intrusion into a wild river area! Presently we come to
a road, a bridge and a log cabin, the retreat of some city dweller.
We tarry long enough to pull the canoe over the bridge and to
glance disapprovingly in the direction of the cabin.
Paddling again, we discover the North Branch taking on the
characteristics of a beaver meadow (Fig. 1, J). The river divides
into several channels. Sloughs lead off on both sides. Abandoned
4 Smith, Galen S., and Robert K. Rose, 1967. Canoeing the Pine and Popple Rivers.
1972] Mills — Canoeing the Wild Rivers Pine and Popple 243
beaver lodges dot the clumps of willow and alder. Due ahead a
break in the conifers marks the location of the South Branch,
emerging to join the North Branch (Fig. 1, 8).
The Forest Service rates the South Branch an excellent canoe
trail, and has provided a fine landing at the Pine River Camp¬
ground bridge. The two mile stretch to Jones Dam is ideal for a
leisurely family outing. The current is slow in the sinuous river
as it courses, fringed with willows and alders, through an open
swamp. A number of beautiful wild swamp-river-forest vistas
open up. Almost all of the scenic views of the South Branch include
tall sentinel white pines which miraculously escaped the axes of
loggers.
Built in the same year and for the same purpose as Windsor
Dam on the North Branch, Jones Dam (Fig. 1, 5) was once the
site of a farm. Ramsay and Jones, an outfit operating camps in
the area, cleared the land for the raising of potatoes and the pas¬
turing of horses. When the company moved out, the buildings were
abandoned, but not for long. A woods character known as White-
water Mike moved in.5 In those days there were always a number
of lazy, smelly bums inhabiting the woods eking out a living by
trapping, poaching game and stealing provisions from the logging
camps. One night a barn burned at one of the camps destroying
eight fine teams of horses. Whitewater Mike, who had had trouble
with James Holmes, the tough camp boss, was suspected of setting
the blaze. Sometime later Whitewater Mike was found shot dead.
The forest fire which destroyed Whitewater Mike’s last abode
also consumed the timbers of Jones Dam. Civilian Conservation
Corps boys in the 1930s planted the potato fields with Norway
pines.
Downriver from Jones Dam is a much longer and more varied
canoe trip with a rapids to be lined or portaged. A full day ought
to be scheduled for passing through it. The current, for the most
part, is slow, and the river continues to meander from side to
side touching the fringes of spruce and balsam. Beaver are numer¬
ous, and one often encounters their dams. A mile below McDonald
Creek (Fig. 1, 6) the river bottom becomes rocky, a fitting prelude
to Wildcat Rapids (Fig. 1, 7). Nicolet National Forest game biolo¬
gist Ed Wilder rates Wildcat Rapids high as a forest beauty spot.
“The area is a green garden of beauty”, he wrote in a report to
forest supervisor Phil Archibald, “with giant moss-covered boul¬
ders, a nearly solid canopy of conifers and the greatest fall of
water per distance involved of any rapids on the river.”6
5 Information obtained from James Huff in 1969 ; personal communication.
6 As cited by Edwin Wilder in a 1967 report to Philip Archbald, of the U. S. Forest
Service, Rhinelander, Wis.
244 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
A deer trail, conveniently located on the left, can be used for the
100-yard portage, or the canoe can be lined through the rapids.
A log jam just above the confluence with Wildcat Creek requires
another pullout. Farther on, a low footbridge spans the river, and
the plywood camp of a beaver trapper follows on the right. The
river then bends north. The remnants of a bridge cribbing mark
the river where it leaves Argonne Township to enter Alvin Town¬
ship. An access road from Highway 55 terminates here at the
location of a farmstead long abandoned. A log cabin occupies a
grove of pines on the east bank, while on the opposite side a barn
overgrown by popples decays into oblivion. The canoe float may
be terminated at this location or continued to the confluence with
the North Branch and a landing at the highway bridge.
A second road from the highway provides access to three cabins
standing in a clearing a mile downriver. An improvised log bridge
spans the river here, barely high enough to provide clearance for
a canoe. The South Branch joins its counterpart, the North Branch,
in a wide open flat (Fig. 1, 8). Merged into a broader stream, the
river swings southeast between high banks covered with popple,
white birch and balsam, occasionally dominated by tall white pines.
As we pass a large log cabin on the left, the bridge looms into
sight. The landing is on the left. Brush almost hides an unsightly
dilapidated cabin. A summer home and two additional cabins stand
on the roadside under the shade of many pines.
From Highway 55 to Highway 139 the Pine is wild, fast-flowing,
and an adventure to canoe. It is no longer a quiet river for family
outings. Only the daring, white-water canoeist should venture onto
these waters; and any attempt to canoe them should be made
during above-normal water conditions of early spring, or following
periods of heavy rains, preferably with a minimum of gear. The
low water levels of late summer expose long stretches of rocks,
and to canoe at this time would necessitate dragging for miles.
Leaving the Highway 55 bridge behind, the river moves moder¬
ately fast between pleasantly timbered banks which soon widen
into an open swamp, the flowage site of Forks Dam. The Pine cuts
through a narrow opening, pouring into a large, deep, circular pool.
The exit from the pool is at the extreme right under the canopy
of a huge balsam. The canoe is immediately caught by the fast cur¬
rent of a sharp pitch. A hundred yards of Grade 1 rapids lie ahead.
For two miles the river is a series of fairly easy rapids, separated
by brief stretches of quiet water. A large clearing on the left is the
location of a logging camp once operated by the Holt Lumber
Company (Fig. 1, 9). Low rectangular mounds outline the shape
of the buildings. A nearby spring was the camp’s source of water.
1972] Mills — Canoeing the Wild Rivers Pine and Popple 245
On the opposite side of the river is the overgrown site of an older
camp.
A half mile downriver the sagging timbers of a railroad bridge
arch overhead. Known as Lindels Spur (Fig. 1, 10) it penetrated
the timber of the upper Pine River. Over its tracks went the hard¬
wood logs passed up during the river drives. The right-of-way,
virtually all of it over lands controlled by the Forest Service,
crosses the river again north of Long Lake.
Near Zepp Farm the river traverses private land. Cottage de¬
velopments threaten the wild character of the banks. One new
A-frame home has been constructed in a manner overhanging the
river. The Zepp buildings stand in a field empty and abandoned.
Below Upper Zepp Bridge the current slows perceptibly, and the
river deepens as it swings to and fro through swampy bottomland.
Two cottages with outbuildings and a collection of tin cans, bottles
and assorted junk flank the river. Lower Zepp Bridge is an arched
wooden structure with a locked gate on its south approach. The
excellent gravel road is public, but the bridge and both banks are
privately owned.
Ten minutes of paddling beyond this point will put the canoeist
at the head of another long series of rapids. In the middle of the
first rapids an island is approached. The canoe should be directed
into the left channel to be followed with a course directly in the
center of the river. The last rapids in this series is in Section 16.
After leaving the island at the base of the rapids, the canoeist
can relax a bit.
After we had descended the rapids safely, the weather took a
turn for the worse. We were enveloped in one of those early spring,
wet, sticky snowstorms sweeping the country. Despite the thickly
falling flakes, we discerned a large bird perched in the top of a dead
pine. We surmised it was a bald eagle, and not wishing to alarm
the bird we headed the two canoes toward a suitable landing. The
moment we rose from our seats the suspicious eagle spread its
broad wings and soared out of sight into the storm. Feeling frus¬
trated we climbed the high river bank, built a fire in the shelter
of several protecting pines, and ate our lunches while we listened
to the moan of the wind above our heads. After this we were glad
to pick up our paddles, for the activity would warm our shivering
bodies.
Shortly thereafter we pass a huge boulder on the left where a
small stream draining a swamp enters the Pine. The terrain is flat
and the current sluggish. The northeastwardly flowing river sud¬
denly turns sharply southwest. We shoot a short rapids before
Kingstone Creek (Fig. 1, 1,1) enters the river. Then comes Pine
Dam No. 3 (Fig. 1, 1,2), a huge affair of rock, gravel and protruding
246 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
timbers. The water flowing’ through the sluiceway is deep enough
for the passage of a canoe, but the menacing protruding spikes
pose a serious menace to the canoes. So we drag them over the
jumble of rocks and timbers to the river a safe distance below.
Having escaped one hazard we confidently resumed our journey
downriver, but misfortune strikes one of the canoes in a boulder-
strewn rapids. In a moment of carelessness the bow of the light¬
weight Grumann strikes a midstream rock, and the rear paddler
fights desperately against the current to prevent the stern from
swinging dangerously downstream. Jammed against another rock,
the canoe is doomed. The two occupants in hip-deep icy water
watch the canoe as it buckles from the pressure of tons of water
against its frail frame. For an hour we struggled with rope and
poles to free the wrecked craft from the clutches of the river. On
a gravel bar we straightened the canoe as best we could, sealed
the rivet holes with adhesive tape from a first aid kit, and with
a lone paddler in the damaged canoe resumed the slow trip to the
next bridge three miles downriver. Some of the romance and
glamor of canoeing a wilderness stream had gone out in the humili¬
ating upset. Running the remaining rapids to Forest Road 2169
seemed devoid of challenge and excitement. Our spirits depressed,
we concentrated on getting through to our cars and a change into
dry clothes.
On another date and in a better frame of mind, we pushed the
canoe into the current of the Pine River, leaving behind Forest
Road 2169 bridge. We pass several cottages that flank the river.
One of the few active farms on the Pine is located on the right
bank. Cattle graze the fields, and a small sawmill supplements the
income from the land. On the left a substantial home has been
erected on the river’s floodplain. A short rapids precedes Stevens
Creek (Fig. 1, 13). Then follows a mile-long straight shot of fast
water that in the right stage affords a safe and exciting run. The
canoe bounces from wave to wave, sweeps around a bend, cuts
through Lindel Spur right-of-way, then settles down in the gentle
current, as Highway 139 bridge (Fig. 1, 14) is reached. The stretch
to the next bridge is devoid of interest. The sounds of highway
traffic, the highway trestle, and the odor of septic tank discharges
from nearby homes detract significantly from the river’s attrac¬
tiveness. Past the bridge and beyond the next bend, charm returns
to the Pine. The only intrusion on the landscape is the power line
to Lost Lake (Fig. 1, IS).
Powers Dam is soon reached (Fig. 1, 16). It was one of the four
dams constructed on the Pine River by the Menominee River Boom
Company, a conglomerate of the logging companies operating on
the Menominee and its tributaries. In all, the company operated
1972] Mills — Canoeing the Wild Rivers Pine and Popple 247
41 dams. The purpose of the dams was to store water so that a
sufficient river level was available to float the logs over falls and
rapids. During the winter months the various logging camps
assembled huge piles of logs at landings on the river banks. Just
prior to the breakup in the spring, the Company sent expert scalers
to estimate the quantity of logs awaiting the drive so that equitable
tolls could be assessed. The peak of logging on the Pine and Popple
rivers occurred in the winter of 1895-98, when 31% million feet
were banked on the Pine, 9 million on the South Branch, 3 million
on the North Branch and 22% million on the Popple. Thereafter
the Boom Company handled a steadily decreasing volume. In the
winter of 1916-17, the mill companies banked their last crop of
logs on the Menominee watershed. In 1918 the Marinette & Menomi¬
nee Paper Company purchased all deadhead logs piled in rollways
along the river. The curtain rang down on the drama of the river
drives in 1919 when the Roper Lumber and Cedar Company drove
its winter cut of cedar on the Pine River down to Marinette.7
Below Powers Dam is Chipmunk Rapids (Fig. 1, 17). Several
white cottages occupy a farm field. A road skirts the left bank,
and if the decision is not to run the rapids the canoe can be landed
where the river bends sharply east to go into the first pitch. A
good canoeist will find Chipmunk Rapids not difficult although a
reading of the river prior to the run will help assure safe passage.
Beyond the lower pitch, rocks continue to rip the fast current, but
these diminish as the bridge and campground are neared.
Don Quinn and I departed from Chipmunk Rapids Campground
early in the day with 20 miles of river ahead of us, much of it
rapids. The day before, we had attempted to gain access at a num¬
ber of points. A washout in the Goodman Grade forced us to turn
around. The road at Seven Mile Creek was too soft for car travel.
The Highway 101 bridge would have to be our exit. We were pre¬
pared for a long, hard day filled with adventure.
A moderate current carried the canoe along at a rapid pace.
On the left charred stumps indicated that the area had once been
ravaged by a forest fire; but the new forest of white birch, popple
and balsam had made remarkable recovery. Tall elms again stood
on the wide flats. Wild ducks were constantly flushing ahead of
the canoe. A kingfisher rattled his disdain at our invasion of his
domain. Attention could not be divorced fully from the river, for
occasional rocks had to be spotted and avoided. There was the addi¬
tional hazard of cedar sweeps close to the banks. We passed a
weatherbeaten hunting camp in a clearing on the left. The hunters
had stretched a cable overhead to give them access to the woods
on the south bank. Entering Section 4, we passed a log jam in
7 Burke, Fred C., 1946. Logs on the Menominee.
248 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 2. View of lower Pine River, in Florence County. Photo taken in
October 1938 by Dorothy Ferguson; print furnished by Walter E. Scott,
courtesy of Wisconsin Department of Natural Resources.
midstream, then noted a quickening current. We were nearing
Snaketail Eapids (Fig. 1, 18) and, upon catching the first sound of
the white water, began hugging the left bank. Backpaddling, we
eased along and eventually located the beginning of the portage
trail.
The river boils into a furious frenzy among the rocks, filling the
surrounding forest with the noise of rushing water. Very few
canoeists who dared accept the challenge of Snaketail Rapids have
made it. The wild beauty of the spot has attracted many. The
portage trail is marked by the coals of numerous campfires. Com¬
plete privacy is unattainable because of a recently constructed
log cabin. Unfortunately Snaketail Rapids is just outside the east
1972] Mills— Canoeing the Wild Rivers Pine and Popple 249
boundary of the Nieolet National Forest. The cabin owner has
access by road from the north. The road is private and the public
is not encouraged to use it.
The portage trail is unimproved; it winds between the boulders
and tends to peter out at its eastern terminus. The canoe can be
launched in the wide eddy below the rapids for the short 100-yard
float to Lower Snake tail Rapids, which can be safely run by an
expert canoeist. An island is followed by a quick left and right
turn, then a short, easy rapids. The current continues fast. Rips
and riffles are common.
The Pine River in the Calumet Hecla lands is truly delightful.
The practice of selective logging, pioneered in Wisconsin by the
Goodman Lumber Company, the former owners of the tract, is
being continued. One is struck by the majesty of the forest here
in contrast with the second growth of the upper river.
Ten minutes below Lauterman Creek (Fig. 1, 19) is a short
rapids easily run on the left. Beyond Kieper Creek, Meyer Falls
is located in Section 36. Rocks All the river in the approach to the
falls. Wading becomes necessary. The river plunges 6 or 8 feet
through a narrow cut in the outcropping of rock. The portage on
the right bank is about 20 yards long. A cottage on a leased loca¬
tion intrudes on the scene.
The river to Wakefield Creek (Fig. 1, 20) has a number of small
islands, all of them with lodged logs and debris. A clearing on the
right was the location of one of the Goodman Lumber Company
camps. Logs were concentrated here in a huge landing for trans¬
portation by rail to the company’s mills at Goodman, Wisconsin.
A root cellar with its sagging roof is all that remains of the camp
buildings. The camp was large, and a stroll through the site will
reveal the location of many buildings. The railroad crossed the
Pine River to penetrate the company timber to the north. For a
number of years after steel was pulled, the bridge was maintained
for the big trucks hauling logs. But with the advent of hard¬
surfaced public roads the usefulness of the bridge ended; and,
with maintenance discontinued, it is no longer safe. In its sad state
of neglect the bridge has a picturesque quality about it, marred
only by the distracting presence of a hunting camp.
A half mile below the bridge, in Section 31 where the river bends
east, a rather long series of brawling rapids begins. Some maps
indicate Bull Falls here, but this is an exaggeration. Good canoeists
successfully run these rapids. Those not wishing to risk an upset
can resort to wading or lining their canoes. A mile and a half will
leave the worst rapids behind. Small rapids continue to characterize
the river. The banks, where Seven Mile Creek enters, are high and
sunny, a good place to look for early spring flowers. The river
250 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
bottom is shallow and gravelly. The river has exposed a slaty schist
type of rock. In the vicinity of Bessie Babbet Lake, the river is
very shallow. In low water, canoeing would be no joy in this area.
The increasing number of cottages and cabins hint of heavy pri¬
vate ownership. Some of the higher banks are eroded. One worried
landowner has constructed a bulkhead to shore up his slumping
real estate. The river is being encroached upon from all sides. A
“No Park — Police’’ sign reminds us, by contrast, of the “Bide-A-
Wee” signboard observed by Aldo Leopold on the Flambeau River.
As Don and I approached the Highway 101 bridge (Fig. 1, 22),
our thoughts were of the famous conservationist. Our experience
had been the same as his. We had been seesawed between the
serenity and beauty of the Nicolet National Forest, the Goodman
timber, and the ugly degradation of civilization.
Beyond Highway 101, the Pine River flows languidly, shallowing
where the rock crop lies close to the surface. On a shelf above the
river on the left an unusual formation thrusts its granite form
upward. In the distance to the north are two cottages. A cluster
of white cottages and green lawns overlooks the confluence of the
Popple River with the Pine (Fig. 1, 23). The Popple sparkles in
the sun as it emerges from the forest and flows purposefully to join
its parent. At this location a trading post grew to a sizable settle¬
ment named LaSalle by 1868. The discovery of iron ore at Florence
and Commonwealth drained the population. The bridge washed out
in a flood, erasing the last vestige of a busy community of a bygone
era; but the river murmurs on, a sense of immortality in its
movement.
The Pine, a big river now, moves along slowly, its youthful vigor
seemingly expended. During the log drives thousands of feet of
logs became lodged in the silt of the slow current. None of the
boom companies bothered to collect the deadheads. But in 1947
a couple of enterprising sawmill operators, Mr. Walter Buza and
Mr. Emmanuel Konell, initiated a salvage project. From the sand
and debris they pulled out an estimated 250,000 feet of hemlock
logs in an excellent stage of preservation. Still discernible in the
butt ends were brands of at least three different boom companies.
Over the line in Section 25 on the right (Fig. 1, 2U) is a new
log cabin owned by Cal Erickson, editor and publisher of the
Florence County Mining News. If Cal is about, he will more than
likely wave the canoeist in for a cup of coffee. When Cal isn’t trout
fishing or deer hunting he is at his typewriter making a living.
In many an acid editorial he has attacked the despoilers of nature
and wild areas. Cal has particularly criticized the building of roads
into the county’s last bits of wilderness. He is fighting a proposed
road to LaSalle Falls (Fig. 1, 25), an outstanding scenic attrac-
1972] Mills — Canoeing the Wild Rivers Pine and Popple 251
tion in the next section. The falls is at present accessible only
by river or trail.
Pine Island is passed enroute. The faint, muffled roar of the Falls
will be picked up around the next bend. The portage is on the
left just past a small island and log jam. Extreme caution must
be exercised in the approach to the portage to avoid being caught
in the fast current. The crash of the water over the 20-foot falls
is almost deafening. Below, the Pine River boils through a half
mile gorge. The portage is easily followed through a pleasant birch
woods. The lower end of the carry is down a steep and rocky
bank. On the opposite side of the river a trail skirts the gorge,
climbs the crest of rocks overlooking the falls, from which excel¬
lent photographs can be made. The trail continues to the head
of the falls where the timbers of a log sluiceway used during
the drives rot into dust. All about are signs of heavy public
use. Fishermen come up the river from Pine River Dam (Fig.
1, 26). Some walk in on the trails. A few come down by canoe,
make the carry, photograph the falls, and continue downriver
perhaps to camp on the shores of the dam. Wisconsin-Michigan
Power Company has provided excellent access facilities on the
north and south shores of its power dam. The shoreline is wooded,
and not a single man-made structure intrudes on the scene. The
rocky points and quiet bays make ideal campsites; indeed all the
sites are occupied on weekends and holidays. No wonder, for in
looking over the scene of rocks, water and forest, one is reminded
of the Canadian bush country.
Construction of the dam commenced in 1920 immediately on the
heels of the last log drive in 1919. The 42-foot high structure
produced a 170-acre reservoir. Drowned were two falls, each
about 8 feet high, a half mile rapids, and a third falls of 12 feet.
LaSalle Falls was the only survivor. One of the obliterated falls
can be felt where the river suddenly drops a foot a short distance
above its confluence with Halls Creek.
The most popular embarkation point for down-river canoe trips
is the County Highway N Bridge (Fig. 1, 27). The float can be
lengthened by beginning at the power plant. To reach the river
it is necessary to clamber down a high embankment with canoe
and gear. The river below the plant is broad and shallow in mid¬
summer. The surrounding hills are the highest anywhere in the
watershed. The river forms three oxbows in the next eight miles.
The Indians eliminated five miles of paddling with a half-mile
portage beginning a short distance upriver from the Highway N
bridge. The modern day canoeists might indulge in a bit of in¬
triguing diversion by trying his hand at tracing the probable
route of the portage. The trail went up the small stream west
252 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
of the gravel pit, turned northeast to cross the present blacktop
roughly four tenths of a mile north of the river. If highway trucks
haven't hauled away the bones, an Indian grave remains near the
gravel pit.
The oxbows are not without interest. The banks are either very
low or quite high. The forest is a delightful mixture of red and
white pine (some of them leaning giants), birch, balsam, and (on
the flats) alder and elm. While the first appearance of jack pine
is between LaSalle Falls and Pine River Dam, the species becomes
common here. The gravelly richer soils of the upper river are giv¬
ing way to poor sand in the lower stretches. A large boulder mid¬
stream has formed a log jam, but there is no difficulty in finding a
passage.
The end of the second oxbow finds the river almost touching
Highway N north of the bridge. A roadside clearing is traversed
by a faint road and an eroded path down the bank to a muddy land¬
ing on the river. Canoeists use this access, the likely eastern termi¬
nus of the Indian portage trail mentioned earlier (Fig. 1, 28).
Past the elm and birch flat below the landing the left bank rises
steeply. Like other similar banks downriver it is marked perpen¬
dicularly by otter slides. Beyond Johnson Creek, where the river
bends south, a panoramic view of river and forest can be had by
climbing to the top of the left bank. The overlook on county forest
lands is an advertised tourist stop on a recently improved gravel
road. Majestic pines stand on the river bank as the river swings
south and east around the next bend. In the next section a local
group of sportsmen, who call themselves Sons of the Pioneers,
have built a new lodge with four picture windows glaring on the
river. There simply had to be an unobstructed view of the river,
so when the bulldozers went to work not a bush or blade of grass
remained. As a final act of desecration the low shoreline was pushed
into the river. One can understand why men who love the outdoors
are attracted by a river such as the Pine, but what impels men
to such thoughtless destruction of nature? This sort of ravishment
repeated over and over would ruin a river.
The open slopes east of Lepage Creek (Fig. 1, 29) represent
samples of the barrens Captain Cram observed at the mouth of the
Pine while on his mission of surveying the Wisconsin-Michigan
boundary. On the south side of the Pine River, up over the fringe
of timber, is another segment of the barrens. The scene is typical,
— scattered clumps of gnarled popples, the outcrop of rock, the
scent of sweet fern. To complete the picture there ought to be a
flock of sharptail grouse. The birds were quite numerous every¬
where in northern Wisconsin 25 to 50 years ago. In the succession
of logging, fires and farming, sharptails found the combination of
1972] Mills— Canoeing the Wild Rivers Pine and Popple 253
openings and forest ideal habitat. The suppression of fires and the
gradual closing in of the openings set in a decline, so that today
it is doubtful whether a single remnant flock remains on the Pine.
Ellwood Lake Outlet (Fig. 1, 30) enters noisily into the river.
Tethered to a cedar nearby is a green flat-bottomed boat named
“Agnes.” A path leads into the woods, and we wonder if there
isn’t a beaver trapper’s cabin among the trees. If so, he has more
compassion for the natural environment than do the Sons of the
Pioneers. He has hardly left a trace of his whereabouts.
Ellwood Lake landing is a small space in the rocks and sweet
fern to turn around. A rutted road climbs up the slope away from
the river. I fervently pray that it will never be improved and
blacktopped, that the barren here will always remain undisturbed,
for this is the very same barren that Captain Cram saw and wrote
about 125 years ago. The rapids are gone, buried by the dead-
waters of Henry Ford Dam on the Menominee River, but the land
is little changed. Let us hope that the landowners, Florence County,
W isconsin-Michigan Power Company, and several individuals
aware of the historical significance of their ownerships, will leave
a heritage to the future, — a barren.
The Popple
There are few historical references to the Popple. Since it was
away from the principal canoe routes of the Indians and explorers,
fame was to come later with the penetration of the watershed by
loggers and settlers, and later still it acquired a reputation as an
outstanding trout stream. Free flowing throughout its length, the
Popple was threatened early in the 1960s by a dam. With statewide
attention centered on the controversy, the Public Service Commis¬
sion denied a construction permit, explaining it had done so to
keep the river free flowing. The Pete Blankenheim and Paul Bab-
bington canoe trip in September 1960 proved that 50 miles of the
Popple was floatable.8 The exploration of the river from Highway
55 to the confluence with the Pine in 1967 by Sierra Club canoeists
confirmed the wild, rugged character of the Popple River Canoe
Trail.
The Popple River is born in the swamps west of Highway 55
(Fig. 1, 31). Flowing serenely northeast, the stream’s nature is
deceptive. Within a mile it narrows to a noisy, stony brook flowing
under countless windfalls through tangles of willows and alders.
Bending southward the stream meanders through a beaver meadow.
Although the water is deeper, canoeing is difficult because of the
8 From an article, “One dcwn cn Aspen Lake’’ in the Wisconsin Academy Review,
1961, spring issue.
254 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
dense alder growth. Though it is virtually a jungle of water and
alders we finally stumbled upon a passageway cut by a beaver
trapper for his boat. An interesting feature of the beaver meadow
is the dozen or so dead pine rampikes rearing their dark huge
hulks above the alders. Upon examining one of the fallen monsters,
we found the trunk so solid as to defy the sharpest axe.
The bridge in Section 12 on the access road from the highway is
safe, but is barred by a locked gate. Clearings occupv both banks.
The river is rocky, and progress by wading or paddling is neces¬
sarily slow. Signs of recent logging litter the banks. A plank bridge
abandoned by loggers has been washed out on the boulders. A
green tar-paper hunting camp stands on a glacial moraine to the
north. A shaky footbridge spans the river where an old tote road
once crossed. On the south side is a watercress-filled soring hole,
source of fresh water for the camps. Away from the river, under
the spruces at the edge of the clearing, is another camp, a log
cabin. East of the cabin the clearing is littered with the rusty
furnishings of a camp destroyed by fire.
The river meanders in tight bends through a narrow swamp
edged heavily with alders. In a deep, dark stream the South Fork
merges with the Popple. The black spruce swamp from which the
South Fork emerges is so inviting that we paddle into it to dis¬
embark from the canoes where a logging road crosses it. Our
topographic map indicates the location of a camp a short walk
up the road. We searched in vain concluding it was moved out or
had completely disintegrated in the wet swampy environment. In
the search we found several huge yellow birch trees growing on
the higher ground. We wondered why they had not been harvested
long ago.
Back in the canoes on the Popple River, paddling northeast, we
feel the quiet remoteness of the river and the vast black spruce
swamp. Black ducks flush ahead of us, and a blue hereon flaps over
the timber. An occasional white pine rears its ragged top above
the spruce. Dead snags point to the sky. The old tote road skirting
the river on the north side crosses the river in Section 8, and here
at the bridge a log jam has formed. A half mile further on, the Rat
Lake outlet (Fig. 1, 32) enters on the left and another unnamed
stream from the right. The remaining half mile to Forest Road
2167 is broad, shallow in places, and a number of boulders dot the
channel. A good landing has been provided by the U. S. Forest
Service beside the bridge, a new steel and concrete structure.
This is a nice location except for a summer cottage on the north
bank. No downriver canoe trip should be commenced here, however.
Below the bridge and around the first bend a huge log jam
has formed. Below the jam the river channel is shallow, extremely
1972] Mills— -Canoeing the Wild Rivers Pine and Popple 255
rocky, and utterly impassable for a canoe. The rapids terminate
in an oblong pool a half mile southeast of the bridge and just a
short carry from Forest Road 2167.
In April, when Tom Sbonik and I carried our canoe to the pool,
we found it still ice-covered. Snow lingered in the timber, but the
river was open. To overcome the chill bite of the early morning
air, we paddled briskly, passing an occasional floating chunk of
river ice. In a half hour we were at the head of a challenging
rapids. Large boulders were numerous, but the first pitch didn’t
appear very difficult. We attempted a run and made it, experienc¬
ing one heart-stopping moment when the canoe hung up momen¬
tarily on a rock. The second pitch we approached with a degree of
trepidation. The drop here was considerable ; and, with the river in
flood stage, an upset in the icy water would spell disaster. Tom and
I prudently decided to portage. The third pitch, though not severe,
was a long one. We continued to drag the canoe through the open
timber on the right bank, the canoe sliding easily over the foot-
deen snow. In a final frenzied dash through a narrow chute the
river calms in a pool shadowed by surrounding hemlocks. We
returned the canoe to the river, and over placid water paddled
between banks of popple, white birch, elm, maple and scattered
thickets of balsam. A large swamp opened into a beaver meadow.
The current began picking up in Section 11; boulders were again
appearing, and the banks were higher. A log jam had formed at a
large midstream boulder. A minute or two of paddling below the
jam brought us to the head of a rapids 100 yards long. We chose
the more open timber on the right, avoiding the dense pines and
balsams on the opposite bank. After the portage the river meanders
between flat alder banks. Fence posts and barbed wire suggest that
we are approaching abandoned farmlands along the town road.
Tom and I landed the canoe at the edge of a field on the left, and
carried it northeast along the base of a low hill and over the road
to the river below the rapids. The portage we had made was over
private lands. Tom and I felt guilty of trespassing; and, had we
been accosted by the owners, were prepared humblv to confess
our guilt. Local people are generally quite forgiving. It is usually
non-residents highly protective of their property rights who put
up “No Trespassing” signs.
The river sweeps along at a good pace as it moves into the forest.
Boulders dot the channel. Railroad Rapids (Fig. 1, 33) will soon
be heard. Tom and I found the alder flat at the head of the rapids
flooded. We headed the canoe toward higher ground on the right,
and found an old road along which we carried the canoe up and
over the railroad embankment to the river on the other side. A
plantation of red pine in neat rows flanks the railroad. Soon we
256 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
were away from the sounds of white water and in the silence of
a broad elm flat, interrupted only by the twitter of birds. Tom, who
had always lived in a big city, was overwhelmed by the quiet soli¬
tude. The serenity we experienced was short-lived, however, for
all too quickly Highway 139 bridge (Fig. 1, 34) hove into sight.
Later by canoe and white water boat, I accompanied Marlene
and Gil Bortleson in the exploration of the Popple from Highway
139 to the confluence with the Pine. The water level gauge at the
bridge read 4.60 when in mid- April we pushed the canoe into the
current. Swinging gradually to the right the river half circles a
roadside picnic grounds built by the Civilian Conservation Corps
boys in the 1930s. The site fell into disuse, and is now overgrown
with popples; but hidden in the grass and briars are fireplaces
constructed of mortar and stone, still sound, evidence that the
boys had built solidly. Their camp stood in a clearing a mile north
on Highway 139.
Ahead the river's meandering was to take us through extensive
swamps. Far from being a dull monotonous float, this stretch of¬
fered some of the finest skylines in the wholewatershed. Off to the
south were the ragged spires of pines piercing the sky. Closer at
hand the blue river was rimmed with the waving dead marsh grass
framed with dark stands of spruce and balsam. Birch, maple and
elm occupied the higher sites.
Nailed to nearby elms were nesting boxes put up by a conserva¬
tionist interested in the welfare of the colorful wood duck. On the
other hand beaver trappers, encouraged by the Wisconsin Con¬
servation Department, were relentlessly pursuing the beaver. We
found their wicked steel traps everywhere. We wondered how the
animals could survive the onslaught of trappers and a state agency,
but somehow enough remain to keep the Popple populated with
beavers. For years the Department has been coping with the prob¬
lem of too many beavers. Trout fishermen contend that the de¬
cline of trout fishing in the Popple is due to the activities of
beavers, and the Conservation Department, in an attempt to quell
the cry, has liberalized seasons and bag limits, not to completely
eradicate the beaver, but to control the number of dams to a
point at which water temperatures tolerable to trout could be
maintained.
Beyond Martin Creek (Fig. 1, 35) the Popple turns east, and a
huge log jam fills the river from bank to bank. We discovered the
high water of the river spilling out among the trees. We poked
the bow of the canoe into the alder brush, and by weaving this
way and that we managed to float around the jam. This feat could
not be accomplished during normal water levels. A short paddle
and we were at the head of McDougal Rapids (Fig. 1, 36). While
1972] Mills— Canoeing the Wild Rivers Pine and Popple 257
Gil and Marlene went to scout the rapids I began the tough portage
on the left. The carry was fairly flat but very difficult because of
stumps, brush, rocks and windfalls. We finally met at the foot of
the rapids, and Gil reported that the descent consisted of three
pitches, every one of them swift. Though Gil rated the rapids “as
No. 1 or No. 2 in difficulty, he felt that canoeists would be well
advised to scout these rapids for log obstructions before attempting
a run.
Resuming our journey between pleasant banks, we pass through
a cut-over land with an occasional white pine missed by loggers.
Beavers keep the riverside popples pretty well harvested. An un¬
painted hunting camp is our clue that we are nearing a road' ‘We
stop to chat with a beaver trapper, and learn that he is from Sha¬
wano and was attracted to the Popple because of the abundance
of beaver reputed to be here. His car is parked by the bridge.
Ten minutes of paddling below Forest Road 2398 is over a broad,
deep and slow current. The original forest here was almost a pure
stand of pine, one of the largest in the entire watershed. Logging
and the fires that followed changed the ecology so drastically, that
only a few scattered pines remain today.
Burnt Dam Rapids (Fig. 1, 37) begins around the bend below
the hunting camp on the right bank. Garnet Tinsman, of Newald,
who owns the camp, states that a logging dam was located at the
head of the rapids to supply sufficient water to take logs through
it, but a forest fire consumed the timbers of the structure so
thoroughly that it is difficult today to believe that a dam ever
existed there.
Gil and Marlene decide to run the rapids. Before entering the
timber on the left to scout out a possible portage route, I watch
the couple cautiously pass around a huge floating log, then, caught
in the fast current of the first pitch, whisk downriver, bouncing
easily over the white crests. Half way through the rapids "a log
jam divides the river into two channels; Gil and Marlene find them¬
selves among the trees trying to find a route back to the main chan¬
nel. The divided waters ultimately regroup into a straight chute
of white water for the remaining distance to the base of the rapids.
Meanwhile I learn that a portage would not be difficult. Deer
trails and logging roads provide a clear path, with only a short
struggle through the brush at both ends of the portage.
Leaving the rapids, the left bank is high with a light scattering
of balsams, Norway pines, white birches and popple. Sweet ferns
blanket the open ground. Of the original pinery only the charred
stumps remain. A Forest Service plantation of pine is growing
well. The river passes a hunting camp, located high on the left
bank under a clamp of pines, bends, then enters the broad willow
258 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
flats, once the flowage behind Podunk Dam. The south horizon is a
ridge of ragged pine tops.
Where the river cuts through a glacial esker, uncanny loggers
constructed Podunk Dam (Fig. 1, 38). The esker runs north-south,
then bends southwest, its location in the forest marked by tall
pines. Don Quinn and I had discovered Podunk Dam a year earlier
at the end of a faint road. Parking the Jeep in the brush, we
climbed to the top of the pine-needled esker; and, as we gazed out
over the sweep of swamp and forest to the west, resolved to come
back another time with tent and pack to really savor the solitude
we felt.
The opportunity came the following spring. We hiked in over
a trail from the south, set up our tents, and built a fireplace with
rocks salvaged from the dam fill. After supper, as the setting sun
was painting the western horizon vivid red and yellow, I wandered
off to investigate the bogs we had skirted coming in. Suddenly
the evening stillness was broken by the yipping of a coyote to the
north, followed by a chorus of yips and yaps from a pack to the
east. Another coyote responded from the west. The blended voices
of all the coyotes filled the forest with wild, vibrating sounds. A
shiver ran up my spine. Were they communicating my presence?
Or were they just giving expression to their free-roaming way of
life? Timber wolves have disappeared from Wisconsin along with
the wilderness, but the coyotes remain to add character to the
North Woods.
Old timers tell an interesting tale about Podunk Dam. The story
illustrates graphically the shrewd bargaining methods of the lum¬
ber barons. It is said that a Scandinavian farmer by the name of
Annunsen from Winnebago County had exchanged his rich farm¬
lands near Lake Poygan for pine lands on the Popple River. To get
the logs out, the river had to be improved, dams built, the chan¬
nel cleared of boulders. Having exhausted his funds on the land
exchange, Annunsen went to his friend Philetius Sawyer of Osh¬
kosh, a man once a farmer who had become wealthy from dealings
in timber. Sawyer, sensing an opportunity for a prime grab,
quickly put up the money, to the tune of $30,000. Before Annunsen
could establish his camps and begin cutting, Sawyer foreclosed on
the loan. Sawyer’s ruthlessness left Annunsen destitute. Sawyer
later becme U. S. Senator, and continued to negotiate for govern¬
ment timber. As if to atone for his misdeeds, Sawyer donated a
library to the City of Oshkosh.9
Riley Creek (Fig. 1, 39) enters the Popple below Podunk Dam
in a broad expanse of open marsh. To the north is a vista of for-
9 The information in this paragraph was related to me in 1969 in a personal com¬
munication from Clarence Harrison.
1972] Mills— Canoeing the Wild Rivers Pine and Popple 259
ested hills. Snags protrude above the willows. Sentinel pines grace
the swamp edges; jack pine forest, a high sandy moraine on the
south. From the top of the moraine there is a view of a large open
bog, part of it flooded. A corrugated metal hunting camp on the
left is followed by a white summer cottage on the right. The river
current picks up and protruding boulders appear. The first pitch
encountered is very short, and can be run with a good water level.
A pool of quiet water about a hundred yards distant leads into the
main rapids. In high water the lower rapids are “tops” for an
exciting run. Gil Bortleson rated the difficulty a solid Grade 2.
Those who will shoot the rapids will find heavy “curlers” and “boil¬
ers” in the sharp drops. The current is very swift, producing waves
two to three feet high. The river bends sharply in two places,
requiring good control of the canoe to avoid smashing into large
boulders. Below the second bend, the course is straight ahead to
the bridge. Under the bridge the rapids terminate in a broad pool.
A parking area has been provided, and trout fishermen have
tramped an easily followed trail along the river. The canoe can be
portaged over the trail, or a logging road farther north can be used.
One-fourth mile below Forest Road 2159 Bridge, the South
Branch of the Popple River (Fig. 1, 40) joins the main Popple.
Though an important tributary, the South Branch is unsuited for
canoeing. Beaver dams, windfalls, old bridges, and log jams fre¬
quently obstruct passage. Heddin Dam, a logging dam between
Forest Road 2159 and 2383 is in an excellent state of preservation.
The canoeist will find the shallows below the dam difficult to navi¬
gate, even in high water. Canoeing the South Branch would entail
unfavorable conditions at any time.
The next two miles of the main Popple is punctuated with nu¬
merous rocks, a number of riffles, and several short rapids. In
places the river is shallow and rocky. A dense forest blankets both
sides. Masons Rapids (Fig. 1, 41) is in two pitches. The first pitch
is sharp but short, followed by a hundred yards of fast but safe
canoeing. The canoe can be landed on the right below the lone pine
tree leaning over the river from the north bank. The river is white
water as it circles to the north in a half moon course. Nearing the
foot of Masons Rapids, the river becomes a virtual rock garden.
Any lead chosen for the canoe will result in a hang-up. Any canoe¬
ist running Masons Rapids can expect to wade to extricate his craft
from the rock pile.
An old corduroy road cuts across the neck of land half circled by
the river beginning at the landing below the white pine. It can be
used for the portage. An excellent popular campsite is located at
the base of the rapids. Fishermen coming up the river in motor
boats to fish the rapids often camp here.
280 Wisconsin Academy of Sciences , Arts and Letters [Vol. '60
The Popple below Masons Rapids is known locally as “Dead-
water”. The setting is an open grassy marsh. Tall hemlocks stand
where Rock Creek enters. Masons Creek is next on the right, with
beautiful vistas of pine on the north. Since leaving the bridge we
have been traversing the timber lands of Calumet Heela. Where
the river crosses the line between Sections 27 and 22 is the site of
Camp No. 1 used when Goodman Lumber Company operated a rail¬
road to haul logs to their mills. Deer hunters salvaged some of the
materials from the original buildings to build a crude hunting
shack. Of the remaining buildings only their outlines can be
observed. Fishermen launch their boats here for the cruise up to
Masons Rapids. The railroad bridge over the Popple River is being
maintained and is safe for travel. Beyond, cottages appear on the
right, some distance back from the river. The steel town bridge
has replaced an older wooden structure. A gauging station has been
erected here by the U. S. Geological Survey. The embankment next
to the gauging station is the embankment of Anderson Dam, named
for Rudy Anderson who drowned here in a logging accident. The
current bridge rests on a rock ledge over which the Popple River
drops.
None but the most daring canoeists should attempt the stretch
of the Popple between the Iron Bridge and Highway 101. “This is
no ordinary stretch of the river,” testified Ralph Hovind at the
Public Service Commission hearing on the proposed Aspen Dam.
Voicing the Conservation Department’s opposition, Hovind con¬
tinued, “It is a tumbling series of falls, rapids and quiet areas
thrown together in a jumbled mixture in a remote, hilly country.”10
Little Bull Falls (Fig. 1, Jp2) is a half mile below the Iron Bridge.
A weatherbeaten, beautifully designed log cabin with steep roof
overlooks the falls from the left, while a new cabin stands on the
right. Spruce, balsam and cedars shade the outcrop of rock which
constitutes the falls. Grade 2 rapids continue half a mile below
Little Bull Falls. Fisherman trails border the river.
Murphy Rapids (Fig. 1, US) is reached after passing a high rock
outthrust on the left. The approach is over a wide, sluggish stretch
of quiet water. The first brief pitch of Grade 1 difficulty ends in a
short pool followed by a quarter-mile of Grade 1 and 2 whitewater
including a sharp bend. An island marks the end of Murphy Rap¬
ids. Deer use Murphy Rapids to cross the river; their trails con¬
verge on both sides.
Enroute to Nine Day Rapids (Fig. 1, 44) a log jam requires a
short portage. The name Nine Day Rapids originated during the
river drives. One spring a huge log jam occurred. Loggers
10 From same reference as Footnote 8.
1972] Mills— Canoeing the Wild Rivers Pine and Popple 261
struggled for nine days to free the logs, and ever since the rapids
have been referred to as the Nine Day Rapids.11 They begin as
Grade 2, tapering to Grade 1 whitewater, with plenty of rock-
dodging thrown in. The current throughout is very swift. Low wa¬
ter summer conditions would make running difficult.
Hendricks Creek (Fig. 1, 1+5) is the beginning of fast water lead¬
ing into Big Bull Falls. Inexperienced canoeists should not proceed
into the fast current, but are advised to begin the portage on the
right. The falls are approximately 10 feet high over a formidable
rock formation. A trail approaches the falls from downriver on the
left bank. The canoe, launched into the pool at the base of the falls,
can be paddled a hundred yards to rapids which continue almost
to Highway 101 bridge. During high water a thrilling ride is in
store. On the downriver side of the bridge is a wayside and landing.
This beautiful stretch of the Popple River would have been lost
had Aspen Dam been built just above Hendricks Creek. Nine Day
Rapids, Murphy Rapids and the river all the way to Little Bull
Falls would have been obliterated. The 35 foot high structure would
have blotted out over half of the vertical drop from bridge to
bridge. Local sporting interests and the Florence County Board
supported the Elco Corporation's application to build the dam ; but
conservationists, assisted by the Wisconsin Conservation Depart¬
ment, presented convincing testimony. Paul E. Klopsteg, a riparian
owner, testified that he “would must prefer a first class river,
which the Popple is, to a third class lake, which the Aspen might
occur. . . . The river is most attractive as it now is ; I cannot believe
that an artificial lake would make it so”. On April 3, 1961, the Pub¬
lic Service Commission denied a permit. In its decision, the Com¬
mission ruled that the Popple River in its natural state offers
greater scenic value for the public than the proposed flowage. The
dam would violate enjoyment of natural scenic beauty by the pub¬
lic, the Commission stated. The Elco Corporation sold its river
holdings to the State following the PSC denial. “A large stretch
of this segment of the Popple River is now public property,” wrote
Walter Scott in the spring 1961 issue of the Wisconsin Academy
Review . “This preservation is vitually important for the enjoyment
of those who want to fish the rapids of a stream so rough that it
can hardly be waded, so precipitous and rocky that canoeing is a
serious challenge, and so scenic that it embodies rugged northwoods
beauty at its best.”
The rapids continue for a mile below the wayside. The gradient
is not severe and the canoeist needs only to avoid the white-tails
in the fast current. A road skirts the right bank, while a farm is
11 Personal communication from Horace McClain, 1969.
262 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Figure 3. View of rapids, looking upstream, in lower Popple River, in
Florence County. These rapids would have been submerged and lost, had the
“Aspen Lake” dam been built. Photo taken May 1, 1960, by Lewis Posekany;
print furnished by Walter E. Scott, courtesy of Wisconsin Department of
Natural Resources.
located on the left. Considerable destruction of the bank and cover
has occurred. A trash dump is an eyesore. The river is broad and
sluggish to Burnt Bridge, where the Girard Lumber Company
logging railroad crossed years ago. Two fishing camps are located
on the right near Montagne Creek. Fishermen use this area heavily.
Below Burnt Bridge the river grows increasingly faster. Two large
boulders, one on the left, the other on the right, are a warning to
1972] Mills— -Canoeing the Wild Rivers Pine and Popple 263
begin a portage over a brush flat on the left to the pool below Wash¬
burn Falls (Fig. 1, 4,6) .
Jennings Falls (Fig. 1, 47) is two miles farther down. The cur¬
rent is swift with numerous riffles. Deadhead logs, occasional rocks,
and small jams are obstructions to look for and avoid. Sweepers
reach out over the river where cedar swamps are traversed. The
river is wide and, in the vicinity of Woods Creek, shallow with a
gravel bottom. The river then bends south, and ahead is the sound
of Jennings Falls. By hugging the right bank the canoeist may con¬
tinue cautiously for another 100 yards to a small cove, where the
portage can begin northeast on the right. Immediately ahead, the
river narrows into a deep gorge to roar and twist among the
boulders. Carrying the canoe one should stay in the depression,
avoiding the higher rock on either side. At best the portage is a
difficult one. Jennings Falls is very impressive in its wild, rocky
setting overgrown with pine and balsam.
The remaining mile and a half to the Pine River is prime trout
water. The terrain is rough and the river’s current fast, as it moves
over a shallow gravelly bottom. Rock outcrops are numerous on
both banks. A few rise in high bluffs. Rounding a final bend the
Popple flows eagerly toward its union with the Pine River.
Thus, with Sierra Club members as my companions, I had canoed
the Pine and Popple, two of northeastern Wisconsin’s wild rivers.
In a series of weekend trips during the springs of 1966 and 1967
we had paddled the sparkling waters of the rivers, felt the excite¬
ment of running rapids, sensed the solitude of wild stretches, gazed
at the vistas of river, swamp and forest. Of the pages of history
written on land and river we could only imagine the scenes : Indians
with flashing paddles in birch bark canoes, traversing a thin blue
thread in a vast unbroken forest. In a stump-studded clearing a
lonely logging camp. Lumberjacks cursing the elements as they
struggle to break a log. A settler scanning his domain of fire-
charred stumps and barn. We observed nature struggling to heal
the scars inflicted by man. We thought of coming generations.
Would their heritage include a wild river?
AN ARCHAEOLOGICAL SURVEY
OF THE PINE, PIKE AND POPPLE RIVERS
Robert J. Salzer
The area of northeastern Wisconsin which is located in the
drainages of the Wild Rivers-— the Pine, Popple, and Pike — is, like
so many other regions in the Upper Great Lakes area, almost com¬
pletely unknown to the culture historian. It is certainly true that
the major prehistoric developments in eastern North America will
never be properly understood until these uninvestigated regions
have received the close scrutiny of the trained investigator. This
report contains the results of an attempt to make a preliminary
assessment of the archaeological and anthropological resources of
one such area.
During the month of July, 1968, a survey crew composed of
Beloit College students under the direction of the author and under
the field supervision of Mr. J. Edson Way, was sent into the Wild
Rivers area of northeastern Wisconsin with the purpose of locat¬
ing and describing sites of prehistoric human activity. Surface col¬
lections of the debris resulting from these activities were made
and are now housed in the collections of the Logan Museum of
Anthropology, Beloit College. The survey program was run at the
same time that excavations were being conducted in nearby Oneida
and Vilas counties by the Beloit College Archaeological Field
School. These latter activities limited the author’s opportunities to
accompany the Wild Rivers survey crew, with the result that the
bulk of the actual work in the field was under Mr. Way’s super¬
vision.
One of the results of the experience derived from four years’
archaeological operations in northern Wisconsin is the knowledge
of techniques which are best suited to survey procedures in a
heavily-forested environment. However, the particular character¬
istics of vegetation cover and drainage in the Wild Rivers area
made surface observation of prehistoric aboriginal remains par¬
ticularly unproductive and frustrating. In addition, the wide¬
spread absence of access roads to the rivers and the small number
of farms adjacent to the rivers made the work of the survey crew
unusually difficult. As a result of these factors and as a result of
1 This is Paper No. 3 in the series “Studies on the Pine-Popple Wild Rivers Area
of Northeastern Wisconsin.” Submitted November 1, 1969.
265
266 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
limited time and funds, our 1968 survey was limited to restricted
areas along the banks of the streams and lakes within the drainages
of these three wild rivers. The various loci of prehistoric habitation
which were located in the course of the survey do not, therefore,
represent more than a fraction of the total number of such sites
to be found in the area.
Not only were such archaeological sites difficult to find, but the
heavy vegetation cover encountered precluded the recovery of
extensive surface collections of cultural debris. Some of the sites
are represented in our collections by a mere handful of the wastage
which results from aboriginal stone-working techniques. Such small
assemblages seldom include artifact forms which can be considered
to be diagnostic of even very broad temporal or cultural divisions
of Upper Great Lakes prehistory. This is a highly significant limita¬
tion of our data, particularly in view of the generally scanty and
imperfect understanding of Upper Great Lakes prehistory which
is now available.
Fortunately, our work in Oneida and Vilas counties and the work
of Dr. Ronald J. Mason in Door County to the southeast, provide
some measure of control and furnish some sort of framework
within which our data may be assessed.
The following list of archaeological sites constitutes the results
of our survey. Each site described has been given a site name, a
Wisconsin Archaeological Survey codification number immediately
following the site name, and a catalog number in the collections of
the Logan Museum of Anthropology, Beloit College (e.g. LMA #)
at the end of the paragraph dealing with that site. For convenience,
the sites are grouped according to the particular drainage system
in which they are located.
The Pine River Drainage
The Franknecht Site (F12) is situated on the north bank of the
Pine River at the confluence with the Popple River. It is located
in the SW % of the SE % of the SW % of Section 23, Township
39 North, Range 17 East, in the Town of Fern, Florence County.
Surface collections from the eroded bank include two quartz ham-
merstones, two chert flakes, 51 quartz flakes, nineteen quartz cores,
six bipolar cores (Figure 1, I), and 12 irregular quartz chunks.
(LMA #21460)
The Two Banks Site (F13) is located in the SW% of the NW^
of the SE 1/4 of Sec. 7, T 39 N, R 18 E, Town of Commonwealth,
Florence County. It is situated on a ridge on the southeast side of
the stream which crosses County Road D at Emily Lake. Surface
collections from the road cut consist of one welded tuff hammer-
1972] Salzer— Survey of the Pine, Pike and Popple Rivers 267
stone, a utilized quartz flake, 38 unmodified quartz flakes, and 3
quartz cores. (LMA #21461)
The Troika Site (F14) appears to be confined to the north side
of County Road D on the northwest side of the stream leaving
Emily Lake, in the SW % of the NW 1/4 of the SE 3 4 of Sec. 7,
T 39 N, R 18 E, Town of Commonwealth, Florence County. It is
possible that this site is a continuation of the Two Banks Site. One
broken chert side-notched projectile point (Fig. 1, C), one broken
quartz small triangular projectile point with serrated edges (Fig.
1, A), two utilized quartz flakes, one utilized quartzite flake, 11
unmodified quartz flakes, one chert flake, and three quartz cores
were found on the surface. (LMA #21462)
The End of Road Site (F15) is on a point on the southeast side
of the junction of the Pine River and the drainage of Bessie Bab¬
bit Lake. It lies in the SE % of the NE % of Sec. 3, T 39 N, R 17 E,
in the Town of Fern, Florence County. The surface collection con¬
sists of 304 unmodified quartz flakes, 48 quartz chunks, 36 quartz
cores, 2 utilized quartz flakes, and two quartz wedges (Fig. 1,
E and F). Wedges are tools which are thought to have been used
in the manufacture of bone tools. (LMA #21463)
The North End Site (F16) lies on a rise to the west of
the stream which flows out of the north end of Long Lake, in the
NE % of the NW % of Sec. 19, T 39 N, R 15 E, Town of Long-
lake, Florence County. The surface of the site yielded 17 plain
shell-tempered bodysherds (fragments of pottery vessels), 2 plain¬
surfaced, grit-tempered bodysherds, and 3 unidentifiable fragments
of pottery. Also collected were three burned chert flakes, one rhyo¬
lite flake, 14 quartz flakes, and two bipolar cores — one of quartz
(Fig. 1, H) and one of quartzite (Fig. 1, J). The entire site ap¬
pears to have been plowed at some time in the past. (LMA #21464)
The Merganzer Point Site (Fr3) is situated on the large penin¬
sula on the northwest shore of Franklin Lake in the SW % of the
NW % of Sec. 21, T 40 N, R 12 E, Town of Hiles, Forest County,
in the Nicolet National Forest. Surface collection yielded 7 quartz
flakes and one quartz bipolar core (Fig. 1, G). The site appears
to have been badly disturbed by erratic digging activities. (LMA
#21465)
The Pike River Drainage
The Eichenger Ring Site (Mt33) is located on the west bank
of the Menominee River about one-quarter mile upstream from
the mouth of the Pike River. It is situated on a high ridge in the
NW % of the SW 1/4 of Sec. 2, T 34 N, R 21 E, Town of Amberg,
Marinette County. The site is intriguing because of the presence
of two low earthen “dance rings”, some small earthen mounds which
268 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Figure 1. Artifacts found in the archeological survey of the Pike, Pine
and Popple Wild Rivers Area. A, small, triangular projectile point, with
serrated edges, from Troika site; B, small triangular projectile point frag¬
ment, from Eichenger Ring site; C, side-notched projectile point, from Troika
site; D, side-notched projectile point base, from Eichenger Ring site; E and
F, quartz wedges, from End of Road site; G, quartz bipolar core, from
Merganser Point site; H, quartz bipolar core, from North End site; I, quartz
bipolar core, from Franknecht site; J, quartzite bipolar core, from North
End site.
1972] Salzer— Survey of the Pine , Pike and Popple Rivers 269
may be burial mounds, and several regular depressions which may
be graves. The site is narrow and about one-quarter mile long.
Surface collection yielded artifacts which suggest that the site
was occupied several times by aboriginal groups. Included are:
a broadly side-notched chert projectile point base (Fig. 1, D),
a small triangular quartz projectile point fragment (Fig. 1, B),
one utilized chert flake, 3 quartz flakes, one chert chunk (burned?),
and a quartz bipolar core. Historic artifacts found on the sur¬
face include bowl and stem fragments of a white kaolin pipe, a
rivet-type cast brass sleigh (dance?) bell, and fragments of glass
and glazed ceramic vessels. Also collected are several small frag¬
ments of both burned and unburned animal bone. Although this
site technically lies outside the Pike drainage, it is included here
because of the interesting information it contributes to the gen¬
eral area. (LMA #21466)
The Mathis Farm Site (Mt34) is located on a hill to the south
and east of the old Mathis farm house in the SE *4 of the NW *4
of Sec. 36, T 35 N, R 19 E, Town of Athelstane, Marinette County.
Due to extensive and prolonged corn agriculture between 1900
and 1935, sand from the back of the hill has been blown over the
site, in places to depths of five feet. The owners report that they
have found many “arrowheads” over the years on the site. Our
collections were extremely meagre and include one quartz flake
and one chert flake. (LMA #21467)
Twin Oaks Site (Mt35). This site is located in the NE of the
SE of Sec. 3, T 34 N, R 21 E, in the Town of Wausaukee, Mari¬
nette County, on a ridge on the north bank of the Pike River about
150 yards upstream from its confluence with the Menominee River.
The surface collection includes one felsite flake, 4 quartzite flakes,
28 quartz flakes, two quartz cores, two chert cores, and a chert
wedge. (LMA #21468)
The Dolan Lake Site (Mt36) is located on a ridge which sepa¬
rates Dolan and Coleman lakes, to the southwest of a small con¬
necting stream, in the SE % of the SW 14 of Sec. 10, T 35 N,,
R 19 E, Town of Athelstane, Marinette County. The site is largely
undisturbed and does not appear to be very large or to have a
deep deposit. The surface collection consists of 2 felsite flakes, and
5 quartz flakes. (LMA #21469)
The Popple River Drainage
No sites were found in the Popple basin. The survey crew felt
that this apparent lack of aboriginal activity may very well be
explained by survey limitations. Access roads are few and widely
separated and much of the river is in low, swampy areas with
heavy growths of alder along the banks.
270 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Discussion
The results of our field survey are frustratingly meagre, which
is not surprising in view of the limitations under which it was
conducted. In spite of our small collections, however, some rather
important observations can be made. The most obvious of these
is the conclusion that the drainages of the Pike, Pine and prob¬
ably also the Popple were clearly exploited by aborigines in both
recent and prehistoric times. Also, the discovery of village and
campsite debris gives an important clue to the sorts of exploitative
patterns which were present. The relative lack of diagnostic tool
forms which were recovered from these deposits by our survey
crew makes the task of determining when and by whom these
artifactual remains were deposited a difficult one.
The impression that the area contains a small number of small
sites with shallow deposits should be considered a function of the
limitations of surface collection techniques in a heavily forested
area. Experience derived from the application of similar techniques
in Oneida and Vilas counties suggests that such surface indications
are seldom representative of the quantity of debris or the depth
of the deposit which lies beneath the humus.
If our primary data are meagre, and do not permit a direct
evaluation of local culture history, recent discoveries by Dr. Ron¬
ald J. Mason in the Door County-Green Bay area to the southeast,
and in the lacustrine district of Oneida and Vilas counties to the
west provide bases for at least a preliminary assessment of the
broad outlines of human history in the Pine, Pike, and Popple
drainages. The general chronology used here is a modified version
of that used by George I. Quimby in his Indian Life in the Upper
Great Lakes (1960).
The Paleo-Indian period (10,000 to 7,000 B.C.)
No direct evidence for human exploitation of the Wild Rivers
area during this period is known. Indeed, intensive investigations
in adjacent areas clearly indicate that only the southern margins
of the Great Lakes region were settled by the mastodon hunters
which moved north during the Twocreekan. In Wisconsin, occupa¬
tions by these big-game hunters are found only in the southern
half of the state.
The Late Paleo-Indian period (7,000 to 5,000 B.C.)
With the retreat of the Valders glacial advance, and the climatic
amelioration which followed, northern Wisconsin and other areas
of the Great Lakes apparently become more attractive to human
settlers. While we have no evidence of occupations during this
1972] Salzer—Survey of the Pine , Pike and Popple Rivers 271
period in the Pine, Pike and Popple basins, we do have information
from adjacent areas such as Green Bay (Mason and Mason 1960)
and from Oneida and Vilas counties (Salzer 1969a, 1969b) to indi¬
cate the presence of at least two phases of the cultural and temporal
continuum which is identified as “Late Paleo-Indian”. In addition,
a 1968 survey crew from Beloit College discovered remains which
date from this period in the Upper Wolf River valley and it seems
likely that future research in the Pine, Pike, and Popple area will
disclose similar evidences of such occupations.
The Archaic period (5000 to 500 B.C.)
Data from our survey are inconclusive but do not rule out the
possibility of occupation during this period. The Archaic period
is a lengthy and complex affair in the Great Lakes and includes
innovations in technology such as ground stone tools (axes, gouges)
and annealed native copper tools. In the Green Bay area, the Oconto
cemetery site (Ritzenthaler and Wittry 1957) has provided radio¬
carbon assays which range from about 5600 B.C. to 3600 B.C. On
the other hand, dates from the Riverside cemetery site (Hrushka
1967) located in the present city of Menominee, Michigan, range
from 500 B.C. to about A.D. 1. In Oneida County, the Squirrel Dam
Site and the Burnt-Roll ways Site provide additional information
on domestic, rather than burial, activities during this long period
(Salzer 1969a, 1969b) . It is likely that the copper tools which were
found some years ago near Long Lake in Florence County repre¬
sent the occupation of the Wild Rivers area by aborigines at this
time (Ritzenthaler 1957).
The Early Woodland (500 B.C. to 100 B.C.) and the Middle Wood¬
land periods (100 B.C. to A.D. 500)
During the first of these periods, pottery technology appears in
the Great Lakes area, and again, although no diagnostic artifacts
were found in our survey of the Wild Rivers area, data from adja¬
cent areas strongly suggest that occupations dating from these two
periods will ultimately be located. These two periods are somewhat
difficult to distinguish in northern Wisconsin. This is largely due
to the presence of at least two distinctive local developments
which have been discovered in the area and to our still meagre
understanding of the details of their origins and elaboration. The
recently defined North Bay Culture (Mason 1966) is found in the
Green Bay-Door County area and dates from around A.D. 100-200.
Its development seems to have been strongly influenced by con¬
temporary Middle Woodland groups in Illinois, Ontario, and New
York. Adjacent to our area on the west is another Middle Wood¬
land manifestation, termed the Nokomis Phase (Salzer 1968,
272 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
1969b), which is largely affected by still unclear connections to
southeastern Wisconsin, although trade with North Bay is evident
also. The Pine, Pike, and Popple, together with the Menominee
river, certainly served as transportation devices in contact between
these two northern Wisconsin areas and we can expect that
further research in the Wild Rivers area will help to elucidate
the mechanisms by which this trade and contact was accomplished.
It is possible that the Troika Site (F14) and the Eichenger Ring
Site (Mt33) may eventually supply some of these details.
The Late Woodland period (A.D. 500 to 1600)
This long time period is one of extremely complex developments
and population movements in the Upper Great Lakes area, and
it is very important to our understanding of the cultural processes
which led to the formulation of the various Indian groups which
were in the area at the time of contact with the Europeans. In
Oneida and Vilas counties, this period is one of population growth.
Not only are villages more numerous and of larger size, but the
quantity and depth of debris and domestic garbage at these sites
clearly indicates intensive and extensive sedentary life (Salzer
1969b). In the Wild Rivers area, the Troika Site (F14), the Eichen¬
ger Ring Site (Mt33), and the North End Site (F16) date from
this period. Based on our experience from Oneida and Vilas coun¬
ties, it is likely that most, if not all, of the sites described in this
report were occupied during this period. Small triangular projectile
points are diagnostic of the period and, in northern Wisconsin,
crushed shell temper for pottery vessels was used only during this
period.
The large quantity and universal occurrence of quartz chipping
debris at the sites in the Wild Rivers area is instructive also,
since in the area immediately to the west, this raw material was
used almost exclusively for the manufacture of stone tools during
this period. However, the low conical and linear burial mounds
found to the west are conspicuously absent in the Wild Rivers
area, with the possible exception of the Eichenger Ring Site.
Mounds are similarly absent at Late Woodland sites in the Door
County area (Mason 1966). However, burial mounds are present
along the Menominee River in Menominee County, Michigan (Brose
1968). A similar situation prevails to the south along the Wolf
River (Barrett and Skinner 1932) „ and also in the Peshtigo River
drainage (Sperka 1962). The significance of these distributions
can only be food for speculation until more research is accom¬
plished in northeastern Wisconsin.
The major ceramic styles in all these areas are similar and in¬
volve round or somewhat conical jars of various size which are
1972] Salzer — Survey of the Pine, Pike and Popple Rivers 273
covered with impressions of a cord-wrapped paddle. Decoration
is common and is usually found near the rim and consists of im¬
pressions of single twisted cords or cord-wrapped sticks or strings.
Such pottery will undoubtedly be found in the Wild Rivers area
as more work is done.
The Historic period (A.D. 1600 to the present)
At least one of the sites located by our survey, the Eichenger
Ring Site (Mt33) , appears to have been occupied by Indians during
the Historic period. However, the artifactual remains would seem
to suggest that this occupation was quite recent and probably
does not date much before the middle of the 19th century. Two
sites which were located in Oneida County in 1966 are similar in
that they also provide evidence of modern artifacts, low earthen
“dance rings”, and shallow depressions suggestive of graves. In
these instances, local traditions attribute the sites to occupations
by the Potawatomi tribe in the late 19th century, and it is pos¬
sible that the Eichenger Ring Site will eventually be similarly
identified. The Forest band of the Potawatomi tribe have settle¬
ments in Forest County to the southwest of the Popple River.
However, representatives of the Chippewa tribe have also been
known for some time in the same general vicinity and they may
have ranged into the Wild Rivers area in the recent past.
To the south and east of the area, the Menominee tribe has
apparently maintained a relatively long-term residence, and, in
fact, claims to have come into being as a recognizable social entity
near the mouth of the Menominee River (Skinner 1913, p. 8).
They were encountered at this locality in 1634 and they appear
to have had additional villages in the general area at this time
(Quimby 1960). It is probable, but by no means certain, that the
Late Woodland occupations in the Wild Rivers area and in adja¬
cent areas represent the material culture remains of the cultural
ancestors of this tribe. Certainly, the Potawatomi, Chippewa, and
other tribes known to have been in the area in recent times are
late prehistoric and early historic immigrants and cannot account
for the Late Woodland debris in the area.
Conclusions
A short-term archaeological survey of the proposed Wild Rivers
area of extreme northeastern Wisconsin during the summer of
1968 succeeded in locating ten loci of prehistoric and historic cul¬
tural activities. Since the literature contains no examples of re¬
sponsible excavation in this area, and, since the exigencies of sur¬
face survey in a largely undisturbed heavily-forested region se¬
verely limit the size and quality of artifactual recoveries, it is
274 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
necessary to rely upon the data and interpretations resulting from
research in nearby areas to evaluate the present data. A reason¬
able, although tentative, sequence of major cultural events for
the Wild Rivers area can be offered on these bases. This culture
history begins shortly after the retreat of the Valders ice sheet
and terminates in the recent past.
It is hoped that full scale excavations will be conducted in this
area in the future since research of this sort can be expected to
produce information on several problems of local and regional sig¬
nificance. For example, we can anticipate the recovery of data
relating to the specific mechanisms and modes of colonization by
human groups in post-Valders times. The area should also pro¬
vide details of the relationships between Middle Woodland groups
in adjacent areas. Hopefully, some insight into the prehistoric basis
of the Menominee tribe might result from such research also.
Certainly, archaeological investigations beyond the scope of our
preliminary survey will be required to fully assess the scientific
and educational resources of the area.
Bibliography
Barrett, S. A., and Alanson Skinner, 1932. Certain Mounds and Village
Sites of Shawano and Oconto Counties, Wisconsin. Bull. Public Museum
of the City of Milwaukee 10 (5).
Brose, David S., 1968. The Backlund Mound Group. Wisconsin Archeologist
49 (1) : 43-51.
Hrushka, Robert, 1967. The Riverside Cemetery Site: A Late Archaic Mani¬
festation in Michigan. Wisconsin Archeologist 48 (3) : 145-260.
Mason, Ronald J., 1966. Two Stratified Sites on the Door Peninsula of
Wisconsin. Anthropol. Papers 26. Museum of Anthropol., Univ. of Michi¬
gan. Ann Arbor.
Mason, Ronald J., and Carol Irwin, 1960. An Eden-Scottsbluff Burial in
Northeastern Wisconsin. American Antiquity 26: 43-57.
Quimby, George I., 1960. Indian Life in the Upper Great Lakes: 11,000 B.C.
to 1800 A.D. Univ. of Chicago Press. Chicago.
Ritzenthaler, Robert E., 1957. Six old copper implements from Long Lake,
Florence County. Wisconsin Archeologist 38 (1), 35 pp.
Ritzenthaler, Robert E., and Warren Wittry, 1957. The Oconto Site: An
Old Copper Manifestation. Wisconsin Archeologist 38 (4) : 222-243.
Salzer, Robert J., 1968. “Middle Woodland Occupations in Northern Wis¬
consin.” Paper read at a symposium on Laurel and its Neighbors. Central
States Anthropological Society Meeting. Detroit, May 3, 1968.
- , 1969a. “Preceramic Occupations in North-Central Wisconsin.”
Paper read at the Canadian Archaeological Association Meetings. Toronto,
Mar. 16, 1969.
■ - , 1969b. An Introduction to the Archaeology of Northern Wisconsin.
Unpublished Ph.D. dissertation. Southern Illinois University. Carbondale,
Illinois.
Skinner, Alanson, 1913. Social Life and Ceremonial Bundles of the Menomini
Indians. Anthropol. Papers Amer. Mus. Nat. Hist. Vo. 13, Part I.
Sperka, Roger, 1962. The Senator Lake Site. Wisconsin Archeologist 43 (4) :
94-106.
THE MAMMALS OF THE PINE AND POPPLE RIVER AREA1
Robert A . McCabe
The Pine and Popple rivers are located in Florence and Forest
counties. These counties have perhaps fewer published natural his¬
tory accounts than any others in the state. The human population
there has always been low and the area is remote from institutions
likely to investigate natural history. Historically, the region is also
relatively sterile since there were no trading posts on either river.
Records from such posts usually present data on occurrence and
abundance for larger mammals and occasionally scattered records
on noncommercial smaller mammals.
The primary source of information on Wisconsin mammals
comes from H. H. T. Jackson’s book, Mammals of Wisconsin
(1961). This publication was the major reference used in forming
this status report for the Pine and Popple rivers. Although the
reference material is now over 15 years old, to my knowledge there
are no additional data from this region. Jackson must be considered
a “splitter” as a mammalian taxonomist. Treatment here will tend
to “lump” rather than split since the approach will be ecological
rather than taxonomic. The only other major work on Wisconsin
mammals is that of Cory (1912). Although the taxonomy and the
range maps are outdated, the records and life history data are
very good, and in its day, the Cory bulletin was an outstanding
publication.
There has been no systematic mammal survey of the Pine and
Popple river watersheds, nor can this present report in any way
be considered definitive. Dr. Howard F. Young, Professor of Biol¬
ogy at Wisconsin State University, La Crosse, and I spent four
days trapping small mammals in the various habitat types along
the two rivers. From June 13-16, 1966, our total of about 520
trap nights produced only the common small mammals: short-
tailed shrew, field mouse, deer mouse, masked shrew, and least
chipmunk.
Purdue University has a forestry summer camp on the Pine
River from which two colleagues, R. E. Mumford and C. M. Kirk¬
patrick of that institution, have contributed data on the small
mammals and observations on the larger forms.
1 This is paper No. 4 in the series, “Studies on the Pine-Popple Wild Rivers Area of
Northeastern Wisconsin.”
275
276 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Mr. and Mrs. Carey Anderson of Sea Lion Lake in Fern Town¬
ship, Florence County, have collected specimens and sight records
from that area and allowed me to use them in this report. Charles
A. Long, Director of the Museum of Natural History at Wisconsin
State University, Stevens Point, has also made available his records
from Chipmunk Rapids and Lost Lake in Long Lake Township of
Florence County.
All game and fur harvest figures are those of the Wisconsin
Department of Natural Resources (formerly the Wisconsin Conser¬
vation Department) . To eliminate repeated reference to the same
work, information from the major sources will be designated by
letters in brackets as follows:
[J]
[A]
[C]
[P]
[L]
[G]
[S]
[DNR]
[WCD]
— H. H. T. Jackson, 1961
— data contributed (in personal communication) by
Mr. and Mrs. Carey Anderson
— Cory, 1912
— Purdue University Forestry Camp staff
— Aldo Leopold, personal notes
— Records of Charles A. Long
— A. W. Schorger papers, 1942-65
— Wisconsin Department of Natural Resources re¬
ports
— Wisconsin Conservation Department, now the
Department of Natural Resources
Isabel Brackbill of Madison compiled data from Department of
Natural Resources records, and aided in numerous ways in pre¬
paring the manuscript.
The assessment below will follow the taxonomic sequence (but
not necessarily the subspeciation or common names) of Jackson
(1961). Walker (1964) was consulted and followed in some cases.
All specimen records, observational data and kill records are
assigned whenever possible to either Florence or Forest counties.
Whenever the data are specific, the locality is identified, particu¬
larly within the watersheds of the two rivers involved.
CLASS MAMMALIA
Order Marsupialia
Family Didelphidae.
Didelphis virginiana (opossum). The opossum is the only mem¬
ber of this order found in Wisconsin, but there are no records in
Jackson for either Florence or Forest counties. However, the kill
reports [DNR] show that four were taken in Florence County in
1972] McCabe — Mammals of Pine and Popple River Area 277
1946, and 51 in 1954. These are the only two years for which I
could find harvest information.
Order Insectivora
Family Soricidae (shrews).
Sorex cinereus (cinereous shrew, masked shrew, common
shrew). Six specimens from Florence County, three from the
vicinity of Florence, and three from Spread Eagle Lake [ J] . “I
have examined specimens from various localities in the interior
and several of the most northern counties including Douglas, Iron,
Florence and Vilas” [C, p. 411]. Cory also lists three specimens
from Spread Eagle Lake. These may be the same specimens re¬
corded later by Jackson. There is a specimen in the Purdue Uni¬
versity collection from Lost Lake area of Long Lake Township,
Florence County [P] and the Andersons have a preserved specimen
of this shrew taken in Fern Township of Florence County [A].
Long collected a masked shrew on August 19, 1968 at Chipmunk
Rapids [A]. Although I found no authentic record of this shrew
in Forest County, it doubtless exists there in the same kinds of
habitats (e.g., moist woods, marsh edges, along streams) in which
it is found in Florence County.
Blarina brevicauda (short-tailed shrew, mole shrew). Seven
specimens are recorded from Florence County, one from Florence
and six from Spread Eagle; one specimen from Forest County
(T 84 N, R 14 E). These are recorded as B. b. kirtlandi [J]. This
species has also been taken at Lost Lake [P] , and at Sea Lion Lake
in Florence County [A]. I also trapped this shrew at Tipler in
Florence County. The animal is apparently a common small mam¬
mal in both the Pine and the Popple watersheds.
Condylura cristata (star-nosed mole). One specimen is recorded
from Newald in Ross Township [J]. Cory records a specimen
from “Newbold” in Forest County. This may be a misspelling of
the Newald location also recorded by Jackson. Another specimen
was taken at Sea Lion Lake in Florence County [A].
Order Chiroptera
Family Vespertilionidae (common bats).
Myotis lucifugus (little brown bat). Although extremely common
throughout the state, there is only one authentic record of this
bat, taken at Sea Lion Lake, Fern Township, in Florence County,
on July 8, 1968 [A]. This bat is recorded as observed at Lost Lake
in Florence County [P] . There are no other records of other bat
species in Florence or Forest counties, largely because there have
been few, if any, collections made there.
278 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Order Lagomorpha
Family Leporidae (hares and rabbits).
Lepus americanus (varying hare, snowshoe hare). This some¬
times superabundant hare has only one authentic record each for
Florence and Forest counties (no specimens examined) [J] . This
species was also observed at Lost Lake, Florence County [P] . A
specimen was taken by Long near the entrance to the Purdue
Science Camp above Lost Lake [G]. I also saw several hares in
Florence County June 1966, in the Pine River watershed near
Chipmunk Rapids. Leopold (1945) shows both Florence and Forest
counties to be well within the range of this species in Wisconsin.
The average annual kill of this hare for an 18-year period ending
in 1956 was 2,605 for Florence County (peak in 1949 with a bag
of 5,481) and 3,177 for Forest County (peak also in 1949, when
8,163 hares were taken) [DNR].
Sylvilagus floridanus (cottontail rabbit). There is one authen¬
tic record from both Florence and Forest counties (no specimens
examined) [J]. The Forest County record is not in the Popple
River watershed. Leopold (1945) records the first cottontail seen
in Forest County in 1914.
The progress to abundance has been slow, and it is doubtful that
either of these two watersheds will ever support high populations
of cottontail rabbits. In the 18-year period prior to 1956 the aver¬
age yearly kill was 342 for Florence County and 613 for Forest
County. By way of comparison, a county with a high cottontail
population, such as Dodge County, in 1956 produced a harvest
of 63,384 cottontails [DNR].
Order Rodentia
Family Sciuridae (squirrels and allies).
Marmota monax (woodchuck, groundhog, marmot). One authen¬
tic record exists for each of Florence and Forest counties. The
Forest County record is not in the Popple watershed (no specimens
examined) [J]. Young saw woodchucks in the Pine-Popple water¬
sheds in May 1969.
Tamias striatus (chipmunk, gray chipmunk). There is one au¬
thentic record from Forest County, and three specimens from
Florence County (two from Spread Eagle Lake; one at T 40 N,
R 16 E). The record from Forest County is not in the Popple
River watershed. Jackson also lists the specimens recorded by
Cory [C] . The Andersons have a specimen from Sea Lion Lake in
Florence County [A] ; as does the Purdue forestry camp for Lost
Lake in Florence County [P]. I also saw numerous chipmunks in
both Florence and Forest counties.
1972] McCabe — Mammals of Pine and Popple River Area 279
Eutamias minimus (least chipmunk, little chipmunk). Forest
County has one authentic record (but not in the Popple River
watershed) , and two specimen records for Florence, Florence
County [J]. There are also specimens from Chipmunk Rapids
where this species was observed abundant [G] . I collected a speci¬
men in June 1966.
Sciurus carolinensis (gray squirrel). This squirrel is now com¬
mon in all parts of Florence and Forest counties. There is one
authentic record in Florence County prior to 1900 [J]. Schorger
(1949, p. 204) records: “The shooting of a gray squirrel at Flor¬
ence in 1886, and again in 1895, in both instances induced the
remark that this species was very rare in Florence County.” As
a hunted animal, the average annual kill (8-year record) has been
322 for Florence County, and 719 for Forest County [DNR].
Sciurus niger (fox squirrel). Although there are no authentic
records of this species prior to 1900 [J] or any specimens ex¬
amined, there has been an open season on fox squirrels in both
Florence and Forest counties since 1948. A six-year average end¬
ing in 1956 shows that the annual harvest in Florence County
was 122, and for eight years in Forest County the average harvest
was 226 [DNR].
Tamiasciurus hudsonicus (red squirrel, chickaree). There is one
authentic record since 1900 in Forest County. Two specimens were
examined for Florence County (one from Richardson Lake; one
from Section 26, T 40 N, R 16 E) [J]. The Richardson Lake
specimen should have been recorded for Forest County, since
Richardson Lake is in Sections 10 and 11, T 35 N, R 14 E of that
county. There is at least one specimen from Lost Lake in Florence
County [P] . This species is common in both the Pine and Popple
watersheds. In the two years for which there are hunting statis¬
tics (1947 and 1948), 96 red squirrels were taken in Florence
County and 1,002 in Forest County [DNR]. This small squirrel
cannot be considered very highly as either a game or fur animal
in Wisconsin.
Glaucomys sabrinus (flying squirrel). There is one authentic
record for southern Forest County [J], one specimen record from
Sea Lion Lake [A], and one from Lost Lake in Florence County
[P], This species is undoubtedly much more abundant than the
records indicate since the species is nocturnal and is not sought
after as a game animal.
Family Castoridae (beavers).
Castor canadensis (beaver). There were no specimens examined
from either county. There is, however, a substantial population of
these animals in both the Pine and Popple watersheds. In 1950-
280 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
1952 the population was relatively stable in Florence County and
increasing in Forest County (Knudsen, 1953). The beaver has
been observed in the Lost Lake area of Florence County [P] .
Knudsen (1963), in his history of the beaver in early Wisconsin,
lists the first record of beaver in Florence County from Tipler
Township in 1920. In 1920 also the first record is given for Forest
County in the town of Alvin. He also states that there were more
beavers in the 1930’s than in the 1960’s. Schorger (1965, p. 167),
however, points out that “A black beaver was caught by Paul
Miller on Pine River, town of Commonwealth, Florence [County],
1886,” and further that “Insofar as known, the beaver was never
exterminated [from Forest County].”
The beaver, once on the verge of extinction, has been an impor¬
tant furbearer in both counties. In the 18 years for which we have
harvest records, the average annual take of beavers for Florence
County was 164, and 240 for Forest County [DNR]. Beavers
have also been pests; between 1938 and 1948, for example, there
were 29 complaints in Florence County and 38 in Forest County
(Hovind, 1948). Such complaints have doubtless increased in
recent years.
Family Cricetidae (mice, voles, muskrats).
Peromyscus maniculatus (woodland deer mouse) . Five specimens
were examined from Florence County (four from Spread Eagle
Lake; one from T 40 N, R 16 E) [J]. There are also specimens
from Lost Lake, Florence County [P], and five specimens from
Chipmunk Rapids [G]. This is doubtless a common species in
both watersheds since Young and I trapped it in both watersheds
in June 1966.
Synaptomys cooperi (lemming mouse). A specimen was collected
at the Purdue Science Camp above Lost Lake, October 1959 [P].
There are no records from Jackson in either Forest or Florence
county. This species perhaps occurs more regularly than the mea¬
ger record shows. There appear to be large areas of suitable
habitat.
Clethrionomys gapperi (red-backed vole). Six specimens were
examined from Florence County (four from Spread Eagle Lake;
two from Florence). One specimen was examined from Forest
County (from Crandon) [J]. At least one specimen was taken at
Lost Lake in Florence County [P] , and one from Chipmunk Rapids
[G] . I caught no specimens in either county while trapping in
likely habitat (June 1966).
Microtus pennsylvanicus (meadow mouse). Four specimens were
recorded from Florence County (all from Spread Eagle Lake), and
1972] McCabe — Mammals of Pine and Popple River Area 281
one authentic record was from Forest County [ J] . Specimens exist
in the Purdue collection from Lost Lake, Florence County [P] .
At Chipmunk Rapids one specimen was preserved and several
were discarded [G] . I caught this species in both watersheds in
June 1966.
Ondatra zibethicus (muskrat). There are two authentic records,
one from each county [J]. The Forest County record was not in
the Popple River watershed. Although there are no specimens from
either county from 1927 to 1957, the trapping records for these
counties show a take of 79,838 pelts: 22,806 from Florence
County and 57,032 from Forest County [DNR] .
Family Muridae (Old World rats and mice).
Rattus norvegicus (Norway rat). There are two authentic
records, one in northeastern Florence County and the other in
southwest Forest County [J]. Neither was from the Pine-Popple
watershed, although it is doubtful that farms in this watershed
are completely free of Norway rats.
Mus musculus (house mouse). There is one authentic record of
the house mouse in southwest Forest County, and one in Florence
County by the Pine River [J] . The Andersons obtained a specimen
of this mouse on September 8, 1968, about one mile from the Pine
River at Sea Lion Lake [A]. It is very likely a common species.
Family Zapodidae (jumping mice) .
Zapus hudsonius (jumping mouse). There were two specimens
examined in Forest County, one at Crandon and the other at
Richardson Lake (T 34 N, R 14 E) ; and one specimen was ex¬
amined in northern Florence County (T 40 N, R 16 E) [J].
Five jumping mice were seen in the Chipmunk Rapids area [G],
and one was taken by the Andersons on September 13, 1968, be¬
side Sea Lion Lake, Fern Township, Florence County [A].
Family Erethizontidae (American porcupines).
Erethizon dorsatum (porcupine). There are two authentic
records of the Canada porcupine, one in southwest Forest County,
the other by the Pine River in Florence County [J]. Porcupines
were seen in the Lost Lake area (1966-68) [P]. I also saw porcu¬
pines in the Pine-Popple watershed, June 1966.
Order Carnivora
Family Canidae (wolf, coyote and foxes).
Canis latrans (coyote, brush wolf). Three authentic records are
shown for southern Forest County, and one in northeast Florence
County [J]. Coyotes were heard in the Lost Lake area, Florence
County, 1966-68 [P], and Young saw a crippled coyote on June 9,
282 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
1969, two miles east of the town of Florence. The spread of the
coyote in Wisconsin needs documentation and appraisal. The re¬
placement of the wolf by the coyote was perhaps relentless until
it was complete. The “brush wolf” was reported as common in
Forest County in 1937 (Anon., 1937). While deer hunting in 1962
I found much sign and heard coyotes frequently in the Armstrong
Creek area.
Canis lupus (timber wolf). There is one authentic record in
Florence County along the Pine River and two in Forest County
[J], all made after 1900. One specimen was taken in Forest
County, Hiles Township [J]. There has been no reliable census of
Wisconsin timber wolves in recent years. In the winter of 1941-42
Daniel Q. Thompson reported (to Leopold) wolves in the town of
Tipler, northern one-third of the town of Long Lake, northwestern
part of the town of Florence in Florence County, and the eastern
one-half of the town of Alvin in Forest County [L] . Thompson’s
later paper (1952) does not repeat this detail of wolf range in
Wisconsin. These were not, however, the only wolves in Wisconsin
at that time. Oliver Flannery is reported (Anon., 1939) to have
collected $185 in wolf and wildcat bounties in one week at Crandon
in Forest County. Young talked to a fisherman in the Florence
(town) area, who claimed he had seen a pair of timber wolves
during the winter of 1968-69. There is little doubt that the timber
wolf is a rare and endangered species in Wisconsin. The timber
wolf, like a wild river, now requires the understanding, apprecia¬
tion and protection deserving of a natural resource so intimately
associated with Wisconsin history and heritage.
Vulpes fulva (red fox). There are two authentic records, one
each in Forest and Florence counties. One specimen was examined
in Forest County in Crandon [J]. Red foxes were seen in the Lost
Lake area, Nicolet National Forest, Florence County (1966-68)
[P]. The red fox apparently burst onto the Florence and Forest
county scene in 1938 when 33 were recorded in the WCD [DNR] kill
statistics. Prior to that time only gray foxes had been taken.
Following 1938 an average annual kill of 204 animals occurred
through 1955. At present it is the most abundant of the two resi¬
dent foxes common to both the Pine and Popple watersheds.
The DNR harvest records show an average take in Florence and
Forest counties to be 204 red fox (18 years) and 25 gray fox
(25 years). These averages do not include years when there were
no records. The first records which began in 1927 show a limited
kill of gray foxes and no red fox. Eleven years later red fox har¬
vest numbers increased markedly as the gray fox kill declined. This
trend was maintained until 1955 when the last report of game
1972] McCabe — Mammals of Pine and Popple River Area 283
harvest by species by counties was available. The trend is shown
in Fig. 1.
Urocyon cinereoargenteus (gray fox) . There are two authentic
records, one in central Forest County and the other in the Popple
watershed in Florence County [J].
Family Ursidae (bears).
Euarctos americanus (black bear) . Four authentic records are
listed, three from Forest and one from Florence counties between
1915 and 1935 [J]. Three authentic records are listed for Forest
County and one for Florence County in Pine River watershed since
1935. Three specimens were taken in northeastern Florence County,
west of Spread Eagle Lake [J] . Adult and cub tracks were seen on
22 July 1967, one mile east of Purdue Forestry Camp [G] . Black
bears were also seen in this same area by the Purdue Camp staff
[P] (1966-1968). In May 1969 Young sighted a black bear in
the Pine River area near Chipmunk Rapids.
Black bears have undoubtedly been common in the Pine and
Popple watersheds for many years. In 1937, when counties were
given the option of closing or keeping open hunting season on
Figure 1. Red and gray fox harvest for Florence and Forest coun¬
ties [DNR].
284 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
bears, Florence and Forest counties chose to keep the season
open and concurrent with the deer season (Grimmer, 1937). Ten
years later (Scott, 1947) the estimated black bear population
was 105 for Florence County and 200 for Forest County. The
annual harvest of black bear in these two counties is shown in
Fig. 2. In 34 years between 1934 and 1968 when data were re¬
corded, 923 black bears were harvested in Florence and Forest
counties [DNR].
Family Procyonidae (raccoons).
Procyon lotor (raccoon). One authentic record exists for Forest
County and one specimen was examined in Forest County at
Laona [J] . Long treed a raccoon on 22 July 1967 in the Chipmunk
Rapids area [G]. The Department of Natural Resources harvest
record for the raccoon in Florence and Forest counties is erratic. In
30 years ending in 1956, raccoons were taken in only 10 years in
Florence County, and in 11 years in Forest County. The average
annual harvest was 12 and 23 raccoons for the respective counties.
Family Mustelidae (weasels and allies).
Martes americana (American marten). One authentic record is
shown for the Pine River watershed, Florence County [ J] . Charles
Cory wrote in 1912 that he had been informed that “martens are
still to be found in the counties of northern Wisconsin .... based
Figure 2. Black bear harvest for Florence and Forest
counties [DNR].
1972] McCabe — Mammals of Pine and Popple River Area 285
upon personal knowledge or the testimony of reliable hunters and
trappers and he listed Florence County among others
[C., p. 383]. This furbearer was apparently present in the Pine
and Popple watersheds but obviously not in great densities. In the
personal records of Aldo Leopold is a notation that Mr. Jack Zatic
saw a marten near Wabikan Lake, town of Laona in 1933 [L] .
I found no published records of recent date. The marten is not on
the list of furbearers that can legally be trapped in Wisconsin.
Maries pennanti (fisher). There are two authentic records for
Forest County, one in the north near the Michigan line, the other
on the Popple River [J]. Both records date from 1923. Tony
Chester, of Antigo, in 1923 claimed he saw fishers in northern
Forest County just east of the Argonne game refuge and believes
they are still there (as of 1939). Eugene Mayo agrees with Ches¬
ter, since he saw fishers in the same area in 1937 (Scott, 1939).
Fourteen fishers (6 males; 8 females) were stocked under Conser¬
vation Department [WCD] auspices in a 40,000-acre “Fisher Wild¬
life Management Area” in a wilderness area within the Pine River
watershed in northern Forest County, 1955-1957 (Bradle, 1957).
Stocking continued so that 60 fishers in total were released in the
Nicolet Forest from 1956 to 1960. They were shortly thereafter
surviving and extending their range (Olson, 1966). Sightings
have been made up to the present.
Mustela erminea (short-tailed weasel, ermine). One authentic
record is listed for southwestern Forest County and one at the
Popple River in Florence County [ J] . The Andersons took a speci¬
men on 18 October 1968 near Sea Lion Lake, Fern Township,
Florence County [A]. This little mustelid is, however, a common
species in both the Pine and Popple watersheds. A 30-year har¬
vest record shows an average annual take from Florence County
of 436, and 732 from Forest County. In 1927, 3,424 weasels were
reported as taken from Forest County alone [DNR] .
Mustela vis on (mink) . There is one authentic record in north¬
western Forest County and one in Florence County along the Pine
River [J] . Mink are doubtless present along both rivers and their
tributaries. A 28-year average lists a harvest of 290 mink an¬
nually for Florence County and 359 for Forest County [DNR].
In an article on the Popple River, Know a River (Erickson, 1962)
is the line: “You must watch for the mink that plays along the
bank. . . ”
Taxidea taxus (badger). One authentic record is listed for
southwestern Forest County and one in Florence County near the
Popple River [J]. Badgers are not abundant anywhere in Wis¬
consin and harvest records for both counties show that in the
286 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
years when at least one badger was reported the average annual
harvest was 9 for Florence County and 6 for Forest County [DNR] .
Since 1955 badgers have been a protected species.
Mephitis mephitis (skunk). One authentic record is located in
south Forest County and one in Florence County south of the
Pine River near Lake Michigan [J] . Three juveniles were ob¬
served in the Chipmunk Rapids area on 8 August 1968 [G], and
a skunk was also seen in the Lost Lake area [P] . Skunks are com¬
mon throughout Wisconsin, with the average annual kill of 109
for Florence County (28 years), and 89 for Forest County (29
years) [DNR].
Lutra canadensis (otter). There is one authentic record for
Florence County in the Pine River watershed, and three authentic
records for Forest County, one of these on the Popple River
near its delta [J]. There is one specimen examined from Cran-
don, Forest County [J] . On a map showing the relative abundance
of the otter in 1951 to 1953, Knudsen (1956) shows the otter to
be fairly common in both Forest and Florence counties. In Leo¬
pold’s personal notes he lists a Mr. F. Bell as seeing an otter on
the Pine River in the town of Florence in 1924 [L].
The otter is rarely abundant anywhere in its Wisconsin range.
The harvest data from Florence County show an annual harvest
of 15 otters over a 22-year period, and 21 for Forest County over
a 27-year period.
Family Felidae (cats and allies) .
Lynx canadensis (lynx). There is one authentic record of the
lynx for Florence County [J]. There could have been and per¬
haps there still is an occasional lynx on either the Pine or Popple
rivers. Published records, however, are meager. Fur-return records
do not distinguish between lynx and the more abundant bobcat.
The average annual harvest of “wildcats” for Florence and Forest
counties is 62 (both counties over a 29-year period) [DNR].
Lynx rufus (bobcat) . Two authentic records are listed, one in
south-central Forest County, and one on the Pine River where it
divides and turns north in Florence County [J]. One specimen
was examined in Florence County (no locality) and three in
Forest County, one at Laona and two at North Crandon [J].
Order Artiodactyla
Family Cervidae (deer and allies).
O do coileus virginianus (white-tailed deer). Seventeen specimens
have been examined from Florence County, 16 in the Spread
Eagle area, and one in Florence. There are no records from Forest
County [J]. Dahlberg and Guettinger (1956), in a range map
1972] McCabe— Mammals of Pine and Popple River Area 287
of deer, show Forest and Florence counties as principal forest
range, but that probable deer densities in both of these counties
before 1800 were only about 10-15 deer per square mile. White¬
tailed deer were seen in the Lost Lake area (1966-1968) [P]
and by Young in the Pine and Popple watersheds in May 1969.
Schorger (1953, p. 224) records the first mention of deer for
Florence County was in 1882, and also that “The following year,
a party of four Ohio hunters . . . killed 18 deer in 25 days on the
Popple River in 1888.” Of Forest County Schorger says (p. 225),
“There is no early information. Much game, including deer, was
obtained by hunters in 1888. Indians bringing venison to Cran-
don in 1889 reported that there was not much game. Deer was
scarce the following season. Indians had only fair success with
deer in 1892 ; however, M. S. Barker bought 1000 pounds of veni¬
son from them at Armstrong. On October 26, 1893, John Bowers
brought to Eagle River ten deer that were killed in the northern
part of Forest County. The complaint was made that deer were
being exterminated by market hunters so that few were left for
the local people. Very few were killed at Three Lakes.”
The deer harvest record for Florence and Forest counties is
shown in Fig. 3 [DNR] . About twice as many deer are harvested
in Forest County as in Florence County.
Figure 3. Deer license sales and deer harvest for Florence and Forest counties
[DNR].
288 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Alces alces (moose). There is one record of moose in south¬
western Forest County (Schorger, 1956). He states (p. 7), “There
is no authentic record of a native moose [for Florence County] .”
However he continues, “In October, 1885, a party of Indiana
and Ohio hunters was reported to have killed 8 deer and a large
moose. The locality was not stated. At this time most of the
‘foreigners’ hunted in Florence County (p. 4.) And further: “A
bull moose that was supposed to have wandered down from Lake
Superior, was killed at Rice Lake by Indians on March 18, 1873. . . .
Rice Lake, town of Crandon, Forest County.” It thus appears
that there are no authentic (i.e., verified) records of native moose
from either Florence or Forest Counties.
Discussion
In spite of the remoteness of Florence and Forest counties from
centers of population, the first state game refuge, it appears, was
in Forest County. In the personal records of Leopold there is a
letter from W. F. Grimmer, then Superintendent of Game Manage¬
ment for the Conservation Department, State of Wisconsin, dated
August 1, 1936. It reads: “On checking our records it appears
that the first state refuge in Wisconsin was established by legis¬
lative act (Chapter 310, Laws of 1915, approved by the governor
on June 23, 1915) and was known as the Forest County Game
Refuge. The description of the refuge was Township 38 North
Range 12 East and Township 38 North Range 13 East.” Town¬
ship 38 at that range includes the Pine and Popple rivers. Today
that area is part of the Nicolet National Forest. The rationale
for establishing a refuge at that time and place is vague at best.
As of 1970, there are 41 mammals recorded for Florence and
Forest counties, any one of which either has been or is likely to
be in the watersheds of the Pine and Popple rivers. The list is
not complete for all possible resident species, nor has the sub¬
specific grouping been thoroughly explored.
The fact that so little is known about the mammal fauna of the
Pine and Popple watersheds should surprise no one. There has
never been a county-by-county mammal survey of Wisconsin.
Jackson’s work (op. cit.) is an excellent starting point, as is Cory’s
(op. cit.) earlier report, but even in the former volume, published
in 1961, the field work was completed ten years earlier, and much
of it as many as 42 years earlier.
The much-needed survey would be expensive, but not exorbitant.
The results could be used in problems of education, recreation,
wildlife management, forestry, and in regional planning. It re¬
mains only for a public agency or teams of agencies to assume
the initiative.
1972] McCabe — Mammals of Pine and Popple River Area 289
Some mammals, like wild rivers, may need to be protected and
cared for so that they will remain on the Wisconsin scene as part
of our historical heritage.
Literature Cited
Anon., 1937. Timber and brush wolves. Wis. Cons. Bull. 2 (10) : 30.
Anon., 1939. Outdoor briefs. Ibid. 4 (12) : 61.
Bradle, Bernard J., 1957. The fisher returns to Wisconsin. Ibid. 32 (11) :
9-11.
Cory, Charles B., 1912. The mammals of Illinois and Wisconsin. Field Mus.
Nat. Hist., Publ. 153. Zool. Series 11. Chicago. 505 pp.
Dahlberg, Burton L., and Ralph C. Guettinger, 1956. The white-tailed deer
in Wisconsin. Wis. Cons. Dept., Techn. Wildlife Bull. 14. Madison. 282 pp.
Erickson, Calvin, 1962. Know a river. Wis. Cons. Bull. 27 (2) : 3.
Grimmer, W. F., 1937. The 1937 bear season. Ibid. 2 (10) : 4-6.
Hovid, James, 1948. Beaver trouble. Ibid. 13 (7) : 15-18.
Jackson, Hartley H. T., 1961. Mammals of Wisconsin. Univ. of Wis. Press.
Madison. 504 pp.
Knudsen, George J., 1953. Beaver population trends, 1950-51. Wis. Wildlife
Research ( Pittman-Robertson Quarterly Progress Reports) 11 (4) : 53-59.
Mimeo.
- , 1956. Preliminary otter investigations. Ibid. 15 (2) : 131-147. Mimeo.
- , 1963. History of beaver in Wisconsin. Miscell. Research Report 7
(Game), Wis. Cons. Dept. Madison. 15 pp.
Olson, Herman F., 1966. Return of a native. Wis. Cons. Bull. 31 (3) : 22-23.
Schorger, A. W., 1949. Squirrels in early Wisconsin. Trans. Wis. Acad. Sci.,
Arts and Lett. 39 : 195-247.
- , 1953. The white-tailed deer in early Wisconsin. Ibid. 42: 197-247.
- , 1956. The moose in early Wisconsin. Ibid. 45: 1-10.
- - — , 1965. The beaver in early Wisconsin. Ibid. 54: 147-179.
Scott, W. E., 1939. Rare and extinct mammals of Wisconsin. Wis. Cons.
Bull. 4 (10) : 21-28.
- - — , 1947. The black bear in Wisconsin. Ibid. 12 (11) : 3-10.
Thompson, D. Q., 1952. Travel, range and food habits of timber wolves in
Wisconsin. Jour, of Mammal. 33 (4) : 429-442.
Walker, Ernest P., et al., 1964. Mammals of the world. The Johns Hopkins
Press. Baltimore. 3 vols.
THE AVIFAUNA OF THE PINE-POPPLE WATERSHED
Howard Young
Introduction
The following account is based on observations made during
eleven trips to the Pine-Popple area between May 1966 and June
1969. Acknowledgement for assistance in field work is made to
Dr. Richard Bernard, Mr. Robert Fiehweg, Dr. William Hilsen-
hoff, Prof. Frederick Lesher, and Dr. Robert McCabe. This study
was supported through funds made available by the Wisconsin
Society for Ornithology.
Additional information was gleaned from back issues of the
Passenger Pigeon (1939 to date), field records of the Milwaukee
Museum, hunting kill estimates of the Wisconsin Department of
Natural Resources, and personal correspondence with Dr. Alvin
Throne, Waukesha, Wis., Mr. Samuel Robbins, Cadott, Wis., and
Mr. William Hummel, Merrill, Wis.
The nature of the regular work assignments of people cooperat¬
ing in this study made it possible for extensive observations to be
made only during early summer. Most trips to the area were made
in early June. Many of the early migrants were therefore missed.
This is particularly noticeable in the case of waterfowl and shore-
birds. In the former case some records of occurrence were obtained
from hunting reports.
Another shortcoming involved the lack of time for intensive nest
hunting, therefore the observers elected to spend their time census-
ing rather than searching for nests. Most of the definite breeding
records were obtained from the field work of Pelzer and Stevens
for the Milwaukee Museum in 1940. Indirect evidence was obtained
for other species, but the records of breeding are very incomplete.
Careful quantitative measurements usually were not feasible;
and, in the annotated list of 169 species which follows, statements
of relative abundance are subjective. Those species simply listed
without comment may be considered as generally common.
Species Observed
Gaviiformes. Common loon3 (Gavia immer) : observed at Lost
Lake (Throne), Butternut Lake (Lesher), Stevens and Porcupine
1 This is paper No. 5 in the series, “Studies on the Pine-Popple Wild Rivers area
of Northeastern Wisconsin."
s Assumed breeder in the area of this study.
* See Table I, Duck Kill Reports.
291
292 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Lakes (Young), and Emily Lake, 1968. While there are no pub¬
lished records of nests, this is well within the breeding range.
Podicipediformes. No records. Grebes undoubtedly occur in the
area, at least as transients.
Ciconiiformes. Great blue heron5 {Ardea herodias) : seen in
small numbers throughout the area ; not common along Pine
and Popple Rivers; rookery found in Goodman Timber, 1968.
Green heron3 (Butorides virescens) : one individual at Howell Lake
(Fiehweg) ; not common along Pine and Popple Rivers. American
bittern3 ( Botaurus lentiginosus) : observed along Pine River, 1940
(Pelzer and Stevens).
Anseriformes. Canada goose ( Branta canadensis) : recorded
October, 1957 (Passenger Pigeon 20 (1) : 33). Snow goose ( Chen
hyperhorea) : Hilsenhoff observed a large mixed flock of this and
the following species over Keyes Lake in November, 1966. Blue
goose ( Chen caerulescens) : see preceding. Mallard3-4 ( Anas platy-
rhynchos) : common; most frequently reported species in water-
fowl kill estimates for Florence County. Black duck3-4 ( Anas
rubripes) : probably fairly common migrant. Gad wall4 ( Anas
strep erus) : waterfowl kill estimates suggest that a few migrate
through the study area. Pintail4 ( Anas acuta). Green-winged
teal4 ( Anas carolinensis) . Blue-winged teal3-4 ( Anas discors) :
seen in Franklin-Butternut area June 12, 1963, by Bernard;
fairly common along Pine and Popple Rivers (Hilsenhoff). Ameri¬
can widgeon4 ( Mareca americana) . Shoveler4 ( Spatula clypeata) .
Wood duck4 (Aix sponsa) : pair on Halsey Lake, June 1967
(Lesher). Redhead4 {Ay thy a americana). Ring-necked duck3-4
{Aythya collaris). Canvasback4 ( A thya valisneria). Scaup3-4 ( Athya
spp.) . Common goldeneye4 {Bucephala clangula) : a record for
August 2, 1968 (Hummel) probably represents a sick or injured
bird, though there are several breeding records for northern Wis¬
consin (Kumlien and Hollister 1951). Bufflehead4 {Bucephala
albeala) . Ruddy duck4 {Oxyura jamaicensis) . Hooded merganser5
{Lophodytes cucullatus) : female with young seen on Riley Lake;
single female seen on Halsey Lake. Common merganser3 {M erg us
merganser) : reported from the Pine River, July 1940 (Pelzer and
Stevens). Red-breasted merganser3 {Merganser serrator) : re¬
ported present in good numbers, August 1941. (Pass. Pigeon 3
(9): 82).
Falconiiformes. Turkey vulture {Cathartes aura) : seen in
April 1946 (Pass. Pigeon 18 (3): 129). Goshawk3 {Accipiter
atricapillus) : seen in November 1943 (Pass. Pigeon 4 (4) : 98) :
5 Breeding records exist for this species in the area of this study.
1972] Young— Avifauna of the Pine-Popple Watershed 293
not common, but has also been recorded on Christmas censuses
from adjacent Forest County. Sharp-shinned hawk3 (Accipiter
striatus) : Pine River July 1940 (Pelzer and Stevens) ; Franklin
Lake 1966 (Lesher) : not common. Cooper’s hawk (Accipiter
cooperii ) : one seen at Lost Lake July 1939 (Throne) ; also on
Pine River July 1940 (Pelzer and Stevens). Red-tailed hawk3
(. Buteo jamaicensis) : fairly common. Broad-winged hawk3 ( Buteo
platypterus) : probably the most abundant hawk of the study area.
Rough-legged hawk ( Buteo lagopus) : November 1957 (Pass.
Pigeon 20 (1) : 34) ; also thirteen were seen March 27-28, 1968,
and three on April 12, 1968 (Hilsenhoff) . Bald eagle5 (Haliaeetus
leucocephalus) : immatures seen by Throne in June 1940, and by
Lesher in 1966; adult at Stevens Lake 1966, and active nest near
there 1968 and 1969; formerly nested at Lost Lake; apparently
scarce in the area at present. Marsh hawk3 (Circus cyaneus) :
two individuals seen (Young, Lesher 1968). Osprey5 (Pandian
haliaetus) : several individuals seen (Throne 1940, Young and
Fiehweg 1966) ; active nest near Long Lake in 1967, 1968 and
1969; not common. Sparrow hawk5 (Falco sparverius) : fairly
common; seen in agricultural areas; nest with young found in
Goodman Timber July 1968 by Bernard.
Galliformes. Spruce grouse3 (Canachites canadensis) : Scott
(1943) lists the last record for Florence County as a single bird
seen in 1932, and estimates that as of 1951 the population for this
area might lie between 10 and 50; no evidence of their presence
since then has been discovered. Ruffed grouse5 (Bonasa umbellus) :
common; seen with young chicks along Pine River 1940 (Pelzer
and Stevens) ; estimated harvest for Florence County in 1967
was 8,832. Sharp-tailed grouse5 (Pediocetes phasianellus) : seen
with young chicks along Pine River, 1940 (Pelzer and Stevens) ;
reported at intervals since then, with last published record for
April 1964 (Pass. Pigeon 27 (1) : 32) ; estimated hunter kill in
Florence County for 1960 was 200.
Gruxformes. Virginia rail3 (Rallus limicola) : reported in Sep¬
tember 1945 (Pass. Pigeon 7 (4) : 124). Coot3 (Fulica americana) :
reports of hunters indicate the following harvests for Florence
County: 23 in 1938, 515 in 1939, 34 in 1940, 30 in 1941, 181 in
1942, 183 in 1943.
Charadriiformes. Killdeer3 (Charadrius vociferus) : fairly com¬
mon, lake edges and streams. Woodcock3 (Philohela minor) : two
seen near Pine River, 1968; estimated hunter kill in Florence
County for 1966 was 536. Upland plover3 (Bartramia longicauda) :
one seen in June 1967 (Lesher), a pair in June 1969 in Florence
cemetery (Young) . Spotted sandpiper5 (Actitis macularia) : seen
294 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
with young- along Pine River in 1940 (Pelzer and Stevens) and
with young on the Pine in 1968 (Hilsenhoff) ; not common in the
watershed. Solitary sandpiper ( Tringa solitaria) : one individual
seen along Pine River (Young, June 1967), two near Fence (Hum¬
mel, July 1968). Herring gull ( Larus argentatus) : seen in sum¬
mer of 1966 (Fiehweg). Ring-billed gull ( Larus delawarensis) :
flock of about 20 (Lesher, 1966). Common tern3 ( Sterna hirundo) :
sighted by Bernard at Franklin Lake in 1963, and by Young at
Halsey Lake in 1966. Black tern3 ( Chlidonias niger) : seen on
several lakes in the watershed area.
Columbiformes. Mourning dove3 ( Zenaidura macroura) : com¬
mon in residential and agricultural areas.
Cuculiformes. Yellow-billed cuckoo3 ( Coccyzus americanus) :
uncommon; Franklin-Butternut area (Bernard, 1963; Lesher,
1966). Near Fence (Young, 1968). Black-billed cuckoo3 ( Coccyzus
erythrophthalmus ) .
Strigiformes. Great horned owl3 (Bubo virginianus) : heard by
Young, 1966; Bernard, 1963. Barred owl3 ( Strix varia) : heard on
several occasions.
Caprimulgiformes. Whip-poor-will3 ( Caprimulgus vociferus) :
recorded Sept. 1956 (Pass. Pigeon 19 (1) : 39) ; also commonly
Table I. Duck Kill Reports.
Compiled from mandatory hunter reports made to the Wisconsin Depart¬
ment of Natural Resources for the years 1938 to 1943 inclusive. These figures
should be viewed with caution, since many hunters are not accurate in their
identifications. However, all species shown here are common migrants in
Wisconsin.
1972] Young— -Avifauna of the Pine-Popple Watershed 295
heard during this study. Nighthawk3 (Chordeiles minor) : fairly
common on lower Pine River.
Apodiformes. Chimney swift3 (Chaetura pelagica) . Ruby-
throated hummingbird5 (Archilochus colubris) : nest with young,
July 1940 (Pelzer and Stevens).
Coraciiformes. Belted kingfisher3 (. Megaceryl alcyon) : common
on streams and in lake areas.
Piciformes. Flicker3 ( Colaptes auratus) : very common. Pileated
woodpecker3 ( Dryocopus pileatus) : not rare; also has been re¬
corded in Christmas censuses from Forest County. Red-headed
woodpecker3 (Melanerpes erythrocephalus) : found in less densely
forested areas. Yellow-bellied sapsucker5 ( Sphyrapicus varius) :
nest found July 1940 (Pelzer and Stevens). Hairy woodpecker3
( Dendrocopus villosus) : while there is evidence that D. villosus is
more abundant than D. pubescens in some parts of northern Wis¬
consin (Young, 1961), limited quantitative data suggest that they
occur in approximately equal numbers in the study area. Downy
woodpecker3 (Dendrocopus pubescens) : see preceding. Black-
backed woodpecker3 (Picoides arcticus) : rare; seen along Pine
River by Pelzer and Stevens (1940) ; another was reported Oct.
1943 (Pass. Pigeon 4 (4) : 98). Northern three-toed woodpecker3
(Picoides tridactylus) : one seen in 1966 (Pass. Pigeon 29 (4) :
120).
Passeriformes : Tyrannidae. Eastern kingbird3 (Tyrannus
tyr annus) . Great crested flycatcher3 (Myiarchus crinitus) . Phoebe5
(Sayornis nigricans ): nest found by Pelzer and Stevens, 1940;
several nests also found in 1967 and 1968 (Hilsenhoff) . Yellow-
bellied flycatcher3 (Empidonax flaviventris) : May 31, 1969; one
bird at Franklin Lake (Bernard). Traill’s flycatcher3 (Empidonax
traillii) . Least flycatcher3 (Empidonax minimus) : abundant. Wood
peewee3 (Contopus virens) : very common. Olive-sided flycatcher3
(Nuttallornis borealis) : recorded by Lesher at Franklin Lake,
1966; apparently not common in the Pine-Popple area.
Alaudidae. Horned lark3 (Eremophila alpestris) : recorded Feb.
1964 (Pass. Pigeon 26 (3) : 151), and March 1967 (Bernard).
Hirundinidae. Tree swallow5 (Iridoprocne bicolor) : common;
seen nesting at Florence and at Sea Lion Lake; many immatures
seen at other places. Bank swallow5 (Rip aria riparia) : one colony
near Fence, also near Lake Emily. Rough-winged swallow5 (Stel-
gidopteryx ruficollis) : two seen at Pine River bridge, 1968; nest¬
ing near Lake Emily, 1969. Barn swallow5 (Hirundo rustica) :
common; seen flying to nests. Cliff swallow5 (Petrochelidon pyr-
rhonota) : common; seen flying to nests. Purple martin5 (Progne
296 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
subis) : common; seen flying to nests; arrival date of April 12,
1964 (Pass. Pigeon 27 (1) : 37).
Corvidae . Gray jay5 ( Perisoreus canadensis) : fairly common;
seen in Goodman Timber and along Pine and Popple Rivers ; young
bird seen in May 1964 (Pass. Pigeon 27 (1) : 38). Blue jay5 (Cyano-
citta cristata) : seen feeding young, July 1940 (Pelzer and Stevens).
Raven3 ( Corvus corax) : common, timbered areas. Crow5 ( Corvus
brachyrhynchos ) : seen feeding young, July 1940 (Pelzer and
Stevens) .
Paridae. Black-capped chickadee3 ( Parus atricapilius) . Boreal
chickadee3 ( Parus hudsonicus) : three seen in November 1949
(Pass. Pigeon 11 (2) : 87) ; also reported Sept. 1956 (Pass. Pigeon
19 (1) : 40) and Aug. 1959 (Pass. Pigeon 21 (2) : 86).
Sittidae. White-breasted nuthatch3 ( Sitta carolinensis) : seen
more commonly than the following during the period of this study.
However, Christmas census records indicate that S. canadensis is
the more abundant of the two in this region (Young, 1965). Red¬
breasted nuthatch3 ( Sitta canadensis) : see preceding.
Troglodytidae . House wren5 ( Troglodytes aedon) : common ; ob¬
served nesting in houses (Florence), but also often heard in woods
away from dwellings. Winter wren3 ( Troglodytes troglodytes) :
uncommon. Short-billed marsh wren3 ( Cistothorus platensis) :
found in several low lying meadows.
Mimidae . Catbird5 ( Dumetella carolinensis) : nest with 4 eggs
near Frog Lake, June 9, 1969. Brown thrasher5 ( Toxostoma
rufum) : nest with 3 young ready to fledge, June 9, 1969, near
Frog Lake.
Turdidae . Robin5 ( Turdus migratorius) : common nester, many
local young observed. Wood thrush3 (Hylocichla mustelina) . Her¬
mit thrush5 ( Hylocichla guttata) : nest record for Lost Lake, 1939
(Pass. Pigeon 3 (2): 13). Swainson’s thrush ( Hylocichla ustu -
lata): seen June 12-14, 1963, at Butternut Lake (Bernard).
Veery3 (Hylocichla fuscescens) : abundant. Bluebird3 ( Sialia
sialis) .
Regulidae . Golden-crowned kinglet3 ( Regulus satrapa) : recorded
June 1970 (Robbins). Ruby-crowned kinglet3 ( Regulus calen¬
dula) : uncommon; bird seen July 9, 1968, was probably a breed¬
ing bird (Hilsenhoff) ; pair seen June 1970 (Robbins).
Bombycillidae . Cedar waxwing5 (Bomby cilia cedrorum) : abun¬
dant; nest with young, July 1940 (Pelzer and Stevens).
Laniidae. Loggerhead shrike3 ( Lanius ludovicianus) : observed
in Oct. 1957 (Pass. Pigeon 20 (1) : 40).
1972] Young — Avifauna of the Pine-Popple Watershed 297
Sturnidae. Starling5 (Sturnus vulgaris) : common in residential
areas, nesting in martin houses ; numerous young observed.
Vireonidae. Yellow-throated vireo ( Vireo flavifrons) : uncom¬
mon. Solitary vireo3 ( Video solitarius ) : four records (Lesher,
1966; Hilsenhoff, 1967 ; Bernard and North, 1968; Robbins, 1970).
Red-eyed vireo5 (Vireo olivaceus) : abundant; seen feeding young
1940 (Pelzer and Stevens). Warbling vireo3 ( Vireo gilvus) .
Parulidae. Black and white warbler3 ( Mniotilta varia) . Golden¬
winged warbler3 (V ermivora chrysoptera) : fairly common along
Pine and Popple Rivers (Hilsenhoff). Tennessee warbler (V ermi¬
vora peregrina) : one record (Young, June 1967). Nashville
warbler5 (V ermivora ruficapilla) : abundant; seen feeding young,
1940 (Pelzer and Stevens). Parula warbler3 (Parula americana) :
uncommon. Yellow warbler3 ( Dendroica petechia). Magnolia
warbler5 ( Dendroica magnolia) : seen feeding young 1940 (Pelzer
and Stevens) ; uncommon. Cape May warbler ( Dendroica tigrina) :
one record, May 27, 1968 (Hilsenhoff). Myrtle warbler3 (Dendroica
coronata) : observed by Fiehweg, 1966. Black- throated green
warbler3 (Dendroica virens) : fairly common in maple areas.
Blackburnian warbler3 (Dendroica fusca) . Chestnut-sided warbler5
(Dendroica pennsylvanica) : nesting record July 1940 (Pelzer and
Stevens) ; very common. Bay-breasted warbler (Dendroica cas-
tanea) : single bird seen by Lesher, June 1966. Blackpoll warbler
(Dendroica striata) : several seen by Lesher, June 1966. Pine
warbler3 (Dendroica pinus) . Palm warbler (Dendroica palmarum) :
Hilsenhoff reported a single bird for May 10, 1968. Ovenbird3
(Seiurus aurocapillus) : abundant. Northern water thrush3 (Seiu-
rus novehoracensis) : fairly common along Pine and Popple rivers.
Mourning warbler5 (Oporornis Philadelphia) : nesting record at
Lost Lake 1939 (Throne) ; also 1940 (Pelzer and Stevens). Yellow-
throat5 (Geothlypis trichas) : abundant; nesting record at Lost
Lake 1940 (Throne). Canada warbler3 (Wilsonia canadensis) : un¬
common. Redstart5 (Setophaga ruticilla) : abundant; nesting rec¬
ord at Lost Lake 1940 (Throne).
Ploceidae. House sparrow5 (Passer domesticus) : abundant in
towns and near farms; numerous nests observed.
Icteridae. Bobolink3 (Dolichonyx oryzivorous) : common in grass¬
land areas, particularly in eastern half of watershed. Eastern
meadowlark3 (Sturnella magna) : common in grassland areas, often
in close proximity to S. neglecta. Western meadowlark3 (Sturnella
neglecta) : see preceding. Redwing5 (Agelaius phoeniceus) : abun¬
dant near lakes; nest with 3 downy young near Commonwealth,
June 9, 1969. Baltimore oriole5 (Icterus galbula) : nest found July
1940 (Pelzer and Stevens). Brewer’s blackbird3 (Euphagus cyano-
298 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
cephalus) : common in agricultural areas. Bronzed grackle5 ( Quis -
coins versicolor) : abundant; seen carrying food, Florence, 1969.
Cowbird3 ( Molothrus ater) : abundant.
Thraupidae. Scarlet tanager5 (Piranga olivacea) : nesting record
July 1940 (Pelzer and Stevens).
Fringillidae . Cardinal3 ( Richmondena cardinalis) : uncommon.
Rose-breasted grosbeak3 ( Pheuticus ludovicianus) . Indigo bunting3
( Passerina cyanea) . Evening grosbeak3 ( Hesperiphona vesper -
tina) : Fiehweg saw 5 late in June 1966; Hilsenhoff saw 25 on the
Pine River in June 1968; also several winter records. Purple finch5
(Carpodacus purpureus) : nesting record July 1940 (Pelzer and
Stevens). Pine grosbeak ( Pinicola enucleator) : one record Jan.
1967 (Bernard) ; reported quite regularly on Forest County Christ¬
mas censuses. Pine siskin3 ( Spinus pinus) : Hilsenhoff had March,
April and May records in 1968. Goldfinch5 ( Spinus tristis) : nest¬
ing record July 1940 (Pelzer and Stevens). Red crossbill3 (Loxia
curvirostra) : “numerous” in Florence County during deer season,
1960 (Pass. Pigeon 23 (2): 74). Twelve seen in March 1965
(Pass. Pigeon 28 (1) : 41). White winged Crossbill ( Loxia leu-
coptera) : a flock on the lower Pine in November 1966 (Hilsen¬
hoff). Rufous-sided towhee5 (Pipilo erythrophthalmus ) : nest with
3 eggs, June 1968 (Young). Savannah sparrow3 (Passer cuius sand-
wichensis) . Vesper sparrow3 ( Pooecetes gramineus) . Slate-colored
junco3 (Junco hyemalis) : uncommon. Tree sparrow (Spizella ar-
borea) : recorded Sept. 1959 (Pass. Pigeon 21 (2) : 89). Chipping
sparrow3 (Spizella passerina) . Clay-colored sparrow3 (Spizella pal¬
lida) : found in several areas in vicinity of Florence; previous 1943
record (Pass. Pigeon 5 (3) : 74). Field sparrow3 (Spizella pusilla) :
uncommon. White-throated sparrow3 (Zonotrichia albicollis) . Fox
sparrow (Passerella iliaca) : four individuals seen by Hilsenhoff,
April 11, 1968. Swamp sparrow3 (Melospiza georgiana) . Song
sparrow5 (Melospiza melodia) : nesting record July 1940 (Pelzer
and Stevens) ; abundant. Lapland longspur (Calcarius lapponicus) :
recorded Oct. 1957 (Pass. Pigeon 20 (1) : 45). Snow bunting (Plec-
trophenax nivalis) : Nov. record 1946 (Pass. Pigeon 9 (1) : 34) ;
Bernard found them in agricultural areas, Jan. and March 1967.
Discussion
Early in the study, two 25-mile transects were established. One
(Transect A) ran essentially west to east mainly through the
Goodman timber, north of Highway 70, but also cut across some
agricultural land for about 3 miles. The second (Transect B)
started at Florence and ran primarily north to south along Highway
101, then curved west. This route traversed much agricultural
area, but also intersected some timber.
1972] Young— Avifauna of the Pine-Popple Watershed 299
These transects were run by automobile, with stops at half-mile
intervals where observations were made for 3 minutes. Transect A
was run 5 times, and a total of 82 species was observed. Transect
B was run 6 times, with a total of 77 species observed. The
transects give some opportunity for comparing the avifauna of
the two areas traversed. The data are summarized in Table II.
If the two areas had identical avifauna, all species would be
common to both transects; if they were entirely different, they
would have no species in common. In this case we have 64/95 = .67
homogeneity. However, of the 28 species restricted to one transect
or the other, 22 (79%) were seen only on a single run. It appears,
therefore, that the observed differences between the two transects
are probably fortuitous.
The considerably greater agricultural development in the eastern
part of the watershed has not as yet resulted in any significant
change in the composition of the avifauna. This reflects the per¬
sistence of large wooded areas in the agricultural area, some pene¬
tration of farming into the western portion, and the mobility of
the birds.
The data are not adequate for a detailed discussion of species
abundance. In the more heavily forested area (Transect A) there
were more records of ruffed grouse, redstart and scarlet tanager.
Along the B transect the following were distinctly more common
than along A : starling, English sparrow, eastern meadowlark, west¬
ern meadowlark, bronzed grackle. All of these species, however,
were recorded on both transects.
Another view of the avifauna may be obtained by examining
the results of both transects which were run simultaneously on
July 14, 1968. Transect A was run by Drs. Richard Bernard, of
Wisconsin State University, Superior, and Charles North, Wiscon¬
sin State University, Whitewater. Transect B was run by myself
and Dr. Steven Goddard, Wisconsin State University, River Falls.
The results are shown in Table III.
Typical patterns of abundance appear. On Transect A a total of
61 species and 324 individuals was recorded. Of these, 21 (34%)
Table II. Summary Data, Transects A and B. Column 1, Number of
Individuals; Column 2, Percentage of the Total.
300 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Table III. Frequency of Bird Species Observed on July 14, 1968.
Ft. = frequency, or percentage occurrence, i.e., number of stops at which
the species was seen divided by the total number of stops. No. — total indi¬
viduals of the species seen that day on the whole transect. Trans. A =
Transect A. Trans.B = Transect B.
were represented by a single individual. Twelve species (20%)
were represented by 10 or more individuals. Together they ac¬
counted for 191 (59%) of all individuals seen. The crow was the
species recorded at the greatest number of stops; the red-eyed
vireo had the largest number of individuals recorded. On Transect
A the number of species per stop ranged from 0 to 15, with an
average of 5.2; individuals recorded per stop ranged from 0 to 22,
with an average of 7.5.
1972] Young — Avifauna of the Pine-Popple Watershed 301
On Transect B a total of 54 species and 383 individuals was
recorded. Of these, 14 (26%) were represented by a single indi¬
vidual. Twelve species (22%) were represented by 10 or more
individuals. Together they accounted for 243 (63%) of all indi¬
viduals seen. The robin was the species seen at the greatest number
of stops; the starling had the greatest number of individuals
recorded. On Transect B the number of species observed per stop
ranged from 0 to 12, with an average of 5.3; individuals recorded
per stop ranged from 0 to 46, with an average of 8.1.
In general it may be said that the Pine-Popple watershed has the
typical bird life of northern Wisconsin.
Hypothetical List
The following group of 47 species would be expected to occur
in the study region according to distributional maps (Gromme
1963) and correspondence with Robbins (1971), but have not been
recorded in the literature for this specific area, and were not ob¬
served during the project. Forty (85%) of these are transient
visitants; only 7 supposed breeders have not been recorded. Also,
almost forty per cent of the species on the hypothetical list are
shore birds, most of which had moved farther north by the time
the observers reached the study area.
If all species on this hypothetical list do in fact occur in the
Pine-Popple watershed, the total species list is raised to 216,
suggesting that our efforts resulted in the recording of about 78%
of the species probably found in the area. Those not yet recorded
are listed below.
Horned grebe ( Colymbus auritus) , pied-billed grebe3 ( Podylim -
bus podiceps) , double-crested cormorant {Phalacro corax auritus ),
whistling swan (Olor cy gnus) , sandhill crane ( Grus canadensis ),
Sora3 ( Porzana Carolina ), semi-palmated plover ( Charadrius semi-
palmatus) , golden plover ( Pluvialis dominica) , black-bellied plover
( Squatarola squatarola) , ruddy turnstone ( Arenaria interpres) ,
common snipe3 ( Capella gallinago) , greater yellowlegs ( Totanus
melanoleucus) , lesser yellowlegs ( Totanus flavipes), pectoral sand¬
piper (Erolia melanotos) , white-rumped sandpiper ( Erolia fus-
cicollis) , Baird’s sandpiper {Erolia bairdii) , least sandpiper ( Ero¬
lia minutilla) , dunlin {Erolia alpina) , stilt sandpiper ( Micro -
palama himantopus) , semipalmated sandpiper {Ereunetes pusil-
lus) , sanderling {Crocethis alba), short-billed dowitcher {Lim-
nodromus griseus) , long-billed dowitcher {Limnodromus scolo-
paceus) , Wilson’s phalarope {Steganopus tricolor), Bonaparte’s
gull {Larus Philadelphia), Forster’s tern {Sterna forsteri) , Cas¬
pian tern {Hydroprogne caspia) .
302 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Snowy owl ( Nyctea scandica) , long-eared owl (Asio otus) , short¬
eared owl ( Asio flammeus) , brown creeper3 ( Certhia familiaris) ,
long-billed marsh wren3 ( Telmatodytes palustris) , gray-cheeked
thrush ( Hylochichla minima), water pipit ( Anthus spinoletta) ,
Bohemian waxwing ( Bombycilla garrulus) , northern shrike
( Lanins excubitor).
Orange-crowned warbler (Vermivora celata) , Cerulean warbler
( Dendroica cerulea) , Connecticut warbler ( Oporornis agilis) ,
Wilson’s warbler ( Wilsonia pusilla) , yellow-headed blackbird3
( Xanthocephalus xanthocephalus), rusty blackbird ( Euphagus
carolinas) , common redpoll ( Acanthis flammea) , grasshopper spar¬
row3 (Ammodramus savannarum) , sharp-tailed sparrow ( Ammo -
spiza caudacuta) , white-crowned sparrow ( Zonotrichia leuco-
phrys) , Lincoln’s sparrow ( Melospiza lincolnii) .
References Cited
Gromme, 0. J., 1963. Birds of Wisconsin. Univ. Wis. Press. 220 pp.
Hummel, W., 1969. Personal communication.
Kumlien, L., and N. Hollister, 1951. The Birds of Wisconsin (with revisions
by A. W. Schorger). Wis. Soc. Ornith., 122 pp.
Passenger Pigeon, 1941-1966. Field Notes Section, specifically the following
citations: 3(2) :13; 3(9) :82; 4(4) :98; 5(3) :74; 7(4) :124; 9(1) :34;
11(2) :87; 18(3) :129; 19 (1) :39— 40; 20 (1) : 33,34,40,41,45; 21(2) :86,89:
23(2) :74; 26(3) :151; 27 (1) :32,37,38; 28(1) :41; 29(4) :120.
Pelzer, W., and L. Stevens, 1940. Unpublished field notes. (On deposit with
the Public Museum, Milwaukee, Wisconsin).
Robbins, S., 1971. Personal communication.
Scott, W. E., 1943. The Canada Spruce Grouse in Wisconsin. Passenger
Pigeon 5 (3) : 67-72.
Throne, A., 1966. Personal communication.
Young, H., 1961. The Downy and Hairy Woodpeckers in Wisconsin. Passenger
Pigeon 23 (1) : 3-6.
- , 1965. An Analysis of Christmas Bird Counts: White-breasted and
Red-breasted Nuthatches. Passenger Pigeon 27 (1) : 16-19.
THE AMPHIBIANS AND REPTILES OF FOREST,
FLORENCE AND MARINETTE COUNTIES WITH SPECIAL
REFERENCE TO THE PINE, POPPLE AND PIKE WATERSHEDS
William E. Dickinson
Introduction
Some of the factors that probably influence the amphibian
and reptile population of the watersheds under consideration are
the temperature (mean: 39° to 40° Fahrenheit), rainfall (36-40
inches annually), soils (some peat, sand and Kenon Loam) and
the soil chemistry (acid to slightly acid). The area in general has
a gently rolling aspect without drastic dispersal barriers. In gen¬
eral it is not being cleared for agriculture to the extent that the
reptiles and amphibians are much disturbed. Food is not a primary
concern : there seems to be an abundance.
The 44 collecting sites in the three counties involved in the
survey vary in types of soil, vegetation, habitats and moisture.
Previous collecting in the areas considered has been generally
sketchy. The first records were apparently those of vacationers.
They came mainly from certain limited lake areas, and this re¬
sulted in wide gaps in the data. Intensive collecting has been done
by Howard Suzuki, 1945-47 (908 specimens) , Dr. C. A. Long (400
specimens), and Dr. George Becker and his students, of Wis¬
consin State University at Stevens Point, Wisconsin, 1966-67 (338
specimens) . Edgren and Levi also worked in the region.
When the present survey committee was formed, the available
records of species and subspecies (12) for the area appeared quite
small, but subsequent collections and research have added several
more. In all, there are now some 35 species and subspecies recorded
or observed for this area.
Many records, though outside the actual watersheds involved,
are nevertheless included (see Table I). These extraneous species
and subspecies have been listed because of similarity of habitats,
and because their range may well be found, in the future, to in¬
clude the watersheds of the Pine, Popple and Pike rivers.
A great deal of collecting has been done in Vilas and Oneida
counties by vacationers. Some of the species thus obtained might
be expected to get into the watersheds of the three rivers here con-
1 This is paper No. 6 in the series, “Studies on the Pine-Popple Wild Rivers Area
of Northeastern Wisconsin.”
303
304 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
sidered. However, most streams in the two counties mentioned
above are in the Mississippi drainage basin, rather than in the
Great Lakes watershed, which is the area under discussion here.
Despite intermittent rumors of poisonous species seen by some
visitors or even residents of the area, no poisonous species have
up to the present been deposited in collections or reported by ex¬
pert observers.
Species Collected or Observed
In the following list certain items of information are given for
each animal, and in the order stated: (1) the name of the species;
(2) the collector; (3) date of collection or observation; (4) the
station or locality at which the specimen was taken or observed;
(5) the catalog number of the specimen in the museum where it is
deposited; (6) the name of the museum where it is deposited.
To save space the following abbreviations will be used: Sta, sta¬
tion or locality of collection or observation ; Cat. No., catalog num¬
ber under which the specimen is stored in the museum ; MPM , Mil¬
waukee Public Museum; WSUSP, museum of the Wisconsin State
University, at Stevens Point, Wisconsin ; UW, museum of the Uni¬
versity of Wisconsin, at Madison, Wisconsin; CMNH, Chicago Mu¬
seum of Natural History, at Chicago, Illinois. For the location of
the collecting stations, listed by number, consult Table I.
Class Amphibia
Order Caudata (tailed amphibians).
Mudpuppies. Nec turns maculosus maculosus (mudpuppy) : range
shown by Bishop (1943, Map 1) and by Conant (1958, Map 154)
as including the Pine-Popple-Pike area. A subspecies, N. m. stictus
is also reported by Conant (1958, Map 154) as ranging into this
area.
Salamanders. Amby stoma laterale and A. j effersonianum (Jef¬
ferson salamander) : Kuony, 8/10/36, Sta. 29, Cat. No. 2568, MPM;
Burant, 7/8/69, Sta. 44, Cat. No. 3274, MPM. Plethodon cinereus
cinereus (red-backed salamander) : Kuony, 8/10/36. Cat. No.
2569, MPM; Strelitzer, 7/5/69, Sta. 44, Cat. No. 3273, MPM.
H emidactylium scutatum (four -toed salamander) : Linds trom,
5/29/49, Sta. 15, Cat. No. 2667, MPM. Diemictylus viridescens
(newt) : Suzuki, 9/7/49, Sta. 42, Cat. No. 2793, MPM.
Order Salientia (jumping amphibians).
Toads. Bufo terrestris americanus (common American toad) :
Long, 7/21/67, Sta. 9, Cat. No. 203, WSUSP; Becker, 6/15/66,
Sta. 19, Cat. No. 3198, MPM ; Suzuki, 10/30/66, Sta. 37, Cat. No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
i — Amphibians and Reptiles in Watersheds 305
I. Collection and Observation Stations: Forest,
Florence and Marinette Counties.
L and 13 are in the Pine River watershed; Station 14 in the
:e basin; Stations 12, 16, 19, 21, 24-27, 30, 32 and 37 are in
watershed; the remainder are in adjacent areas within the
involved.
306 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
2841, MPM; Groh, 7/7/69, Sta. 44, Cat. No. 3277, MPM ; also
observed by Dickinson, 7/7/28, at Sta. 10.
Frogs. Pseudacris triseriata triseriata (3-striped chorus frog) :
listed by Conant (1958, Map 235) as including the Pine-Popple-
Pike area in its range, as does also the subspecies P. t. maculata
(Conant, 1958, Map 235). Hyla crucifer crucifer (spring peeper) :
Surges and Schmidt, 7/9/69, Sta. 44, Cat. No. 3272, MPM. Hyla
versicolor versicolor (tree frog) : Clowes, 9/12/07, Sta. 13, Cat.
No. 789, MPM; Kuony, 8/6/35, Sta. 29, Cat. No. 2562, MPM;
Suzuki, 8/24/47, Sta. 40, Cat. No. 2873, MPM. Rana catesbiana
(bullfrog) : Suzuki, 8/24/47, Sta. 40, Cat. No. 2892, MPM. Rana
clamitans melanota (green frog) : Long, 7/21/67, Sta. 9, Cat. No.
205, WSUSP ; Becker, 7/23/66, Sta. 11, Cat. No. 3250, MPM;
Becker, 6/23/66, Sta. 7, Cat. No. 3263, MPM; Becker, 6/17/66,
Sta. 24, Cat. No. 3257, MPM; Becker, 6/16/66, Sta. 33, Cat. No.
3267, MPM; Kuony, 8/1/35, Sta. 29, Cat. No. 2567, MPM; Suzuki,
8/12/48, Sta. 38, Cat. No. 2996, 2997, MPM. Rana palustris
(pickerel frog): Becker, 6/23/66, Sta. 6, Cat. No. 3255, MPM;
Suzuki, 8/21/48, Sta. 43, Cat. No. 2894, MPM. Rana pipiens (leop¬
ard frog) : Long, 7/21/67, Sta. 9, Cat. No. 206, WSUSP; Kuony,
8/6/35, Sta. 29, Cat. No. 2565, MPM; Suzuki, 9/2/49, Sta. 39,
Cat. No. 2947, MPM; Suzuki, 8/12/48, Sta. 38, Cat. No. 2946,
MPM; Becker, 6/30/65, Sta. 17, Cat. No. 3194, MPM; Becker,
7/12/66, Sta. 28, Cat. No. 3195, MPM. Rana septentrionalis
(mink frog): Becker, 6/5/66, Sta. 19, Cat. No. 3221, MPM;
Becker, (date?), Sta. 36, Cat. No. 3222, MPM; Becker, 6/15/66,
Sta. 16, Cat. No. 3223, MPM; Becker, 6/24/66, Sta. 21, Cat.
No. 3224, MPM; Becker, 8/24/66, Sta. 4, Cat. No. 3226, MPM,
Becker, (date?), Sta. 1, Cat. No. 3278, MPM; Becker, 7/11/66,
Sta. 35, Cat. No. 3232, MPM; Long, 7/21/67, Sta. 9, Cat. No.
207, WSUSP; Becker, 6/23/66, Sta. 12, Cat. No. 3266, MPM;
Becker, 6/23/66, Sta. 6, Cat. No. 3254, MPM ; Becker, 6/16/66,
Sta. 26, Cat. No. 3259, MPM; Becker, 6/14/66, Sta. 32, Cat. No.
3260, MPM; Becker, 6/24/66, Sta. 34, Cat. No. 3268, MPM;
Becker, 6/23/66, Sta. 8, Cat. No. 3261, MPM ; Becker, 6/29/66,
Sta. 5, Cat. No. 3262, MPM ; Becker, 6/23/66, Sta. 7, Cat. No. 3264,
MPM; Becker, 6/30/66, Sta. 18, Cat. No. 3197, MPM; Becker,
6/13/66, Sta. 25, Cat. No. 3265, MPM; Becker, 6/21/66, Sta. 27,
Cat. No. 3258, MPM. Strelitzer, 7/9/69, Sta. 44, Cat. No. 3276,
MPM. Rana sylvatica (wood frog) : Long, 7/21/67, Sta. 9, Cat. No.
204, WSUSP.
Class Reptilia
Order Chelonia (Turtles). Chelydra serpentina (snapping
turtle): Becker, 7/3/66, Sta. 30, Cat. No. 3216, MPM; Suzuki
1972] Dickinson — Amphibians and Reptiles in Watersheds 307
observed this species on 8/26/48 at Sta. 42, and on 8/26/48 at Sta.
39. Chrysemys picta belli (Western painted turtle) : Long, 7/21/67,
found one dead on Hwy. 8, Forest Co. Chrysemys picta marginata
(Midland painted turtle) : Becker, 6/10/66, Sta. 31, Cat. No. 3218,
MPM ; Witman, 7/11/69, Sta. 44, Cat. No. 3279, MPM. Clemmys
insculpta (wood turtle) : Johnson collected one in the Pine-Popple-
Pike area, date not recorded, Cat. No. 2689, MPM. Emys blandingii
(Blanding’s turtle) : shown by Conant (1958, Map 29) as occurring
in the area of this study.
Order Sauria (Lizards). Eumeces fasciatus (5-lined skink).
Smith (1946, Map 26) indicates that the range of this lizard
includes the area of this study ; Pelzer collected one in this area in
1939, Cat. No. 2653, MPM.
Order Serpentes (Snakes). Diadophis punctatus edwardsii
(ringneck snake) : Ziebel, Sept./1954, Sta. 41, Cat. No. 3059, MPM.
Elaphe vulpina (fox snake) : Long, 7/21/67, Sta. 14, Cat. No. 208,
WSUSP (snake was dead when found) ; Riker, 6/21/66, Sta. 20,
Cat. No. 3219, MPM; Grow, 10/17/32, Sta. 41, Cat. Nos. 2432-
2435, MPM; Kuony, Aug. 1936, Sta. 29, Cat. No. 2558, MPM.
Heterodon contortrix (hognose snake) : Pelzer, 1940, Popple R.,
Cat. No. 2606, MPM; Grow, 10/17/32, Sta. 41, Cat. Nos. 2430-
2431, MPM; Fullerton, 9/10/56, Marinette Co., Cat. No. 3075,
MPM. Lampropeltis doliata triangulum (milk snake) : (Schmidt,
1941, p. 190). Opheodrys vemalis (green snake) : Long, 7/21/67,
Sta. 9, Cat. No. 210, WSUSP; Levi, July 1949, Sta. 3, UW; Suzuki,
8/30/49, Sta. 39, Cat. No. 2775, MPM; Cooper, 1927, Sta. 23, Cat.
No. 2200, MPM. Pituophis catenifer sayi (bull snake) : Dickinson
observed this snake 7/5/28 at Sta. 30; Conant (1958, Map 123)
indicates that its range includes this area. Storeria dekayi dekayi
(brown snake) : Suzuki, 9/2/49, Sta. 42, Cat. No. 2773, MPM.
Storeria occipitomacnlata (red-bellied snake) : collected by Rein-
hard, 8/31/07, in this area of study, Cat. Nos. 763, 788, 812, MPM;
Long, 7/21/67, Sta. 9, Cat. No. 209, WSUSP ; collected by Arch¬
bald, undated, in area of this study, Cat. No. 3312, CMNH ; Suzuki,
9/3/49, saw this snake at Sta. 42. Thamnophis sauritus proximus
(ribbon snake) : Suzuki, 9/2/49, Sta. 39, Cat. No. 2777, MPM.
Thamnophis sirtalis sirtalis (common garter snake) : Becker,
10/28/66, Sta. 2, Cat. No. 3214, MPM; Levi, July 1949, Sta. 3, UW;
Becker (date?), Sta. 19, Cat. No. 3215, MPM; seen by Suzuki,
9/4/49 at Sta. 39, and by Dickinson, 6/6/28, Sta. 3; Witman,
7/11/69, Sta. 44, Cat. No. 3278, MPM; Natrix sipedon sipedon
(water snake) : this snake is indicated by Conant (1958, Map 850),
Schmidt (1941, Map 219), and Wright and Wright (1942, Map 42)
to have a range including the area of this study.
308 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Remarks on Distribution
Perusal of the above records reveals that only two species have
been reported officially from the Popple River watershed : Elaphe
vulpina (the fox snake) and Heterodon contortrix (the hognose
snake). The Pine River watershed is represented by ten species:
American toad ( Bufo terrestris) , tree frog ( Hyla versicolor) , green
frog (Rana clamitans) , pickerel frog ( Rana palustris) , leopard
frog {Rana pipiens) , wood frog {Rana sylvatica) , mink frog {Rana
septentrionalis) , green snake {Opheodrys vernalis) and garter
snake {Thamnophis sirtalis) . The Pike watershed has only five
representatives : the toad, the green frog, the mink frog, the snap¬
ping turtle {Chelydra serpentina) and the garter snake.
The fact that so few common species have so far been found
in these drainage basins is not because they are not there, but
because so little collecting has been done in these wild areas. In
fact many of them have been found in the adjacent county areas.
A great deal more collecting must be done in the whole wild rivers
area of northeastern Wisconsin before any adequate idea can be
formed regarding the distribution, abundance and range of am¬
phibians and reptiles in this remote area.
Bibliography
Bishop, S. C., 1943. Handbook of Salamanders. Comstock Publ. Co., Ithaca,
N.Y. 555 pp.
Committee on Herpetological Common Names, 1956. Common Names for
North American Amphibians and Reptiles: Copeia 3: 172-185.
Conant, Roger A., 1958. A Field Guide to the Reptiles and Amphibians.
Riverside Press, Boston. 366 pp.
Dickinson, W. E., 1950. Recent Additions to the Reptiles of Wisconsin.
Trans. Wis. Acad. Sci., Arts and Lett. 40 (1) : 71.
Edgren, R. A., 1944. Notes on Amphibians and Reptiles from Wisconsin.
Amer. Midi. Nat. 32: 495-506.
Pope, T. E. B., and W. E. Dickinson, 1928. The Amphibians and Reptiles of
Wisconsin. Milwaukee Public Mus. Bull. 6 (1). 139 pp.
Schmidt, K. P., 1941. Field Book of Snakes. Putnam, New York. 365 pp.
Smith, Hobart M., 1946. Handbook of Lizards. Comstock, Ithaca, N.Y. 557 pp.
Suzuki, Howard K., 1951. Recent Additions to the Amphibians of Wisconsin.
Trans. Wis. Acad. Sci., Arts and Lett. 40 (2) : 215.
Wright, A. A., and A. H. Wright, 1942. Handbook of Frogs and Toads.
Comstock, Ithaca, N.Y. 284 pp.
ANNOTATED LIST OF THE FISHES OF THE
PINE-POPPLE BASIN1
George C. Becker 2
Introduction
Perhaps the first systematic survey of fishes of the Pine-Popple
basin occurred in the late 1920?s when E. Willard Greene (1935)
made his survey of the fishes of Wisconsin. From some 20 collec¬
tions he recorded the following1 19 species: rainbow trout, brook
trout, white sucker, blacknose dace, longnose dace, creek chub,
pearl dace, northern redbelly dace, finescale dace, common shiner,
bluntnose minnow, central mudminnow, yellow perch, johnny
darter, Iowa darter, smallmouth bass, largemouth bass, mottled
sculpin and brook stickleback.
For many years the Wisconsin Department of Natural Resources
has surveyed the waters of the Pine-Popple basin in both their
research and management programs. Aside from the game and
panfish species there appeared to be little interest in fish systematics
until Mason (1966) listed all of the species he encountered at 92
different locations. Several additional species were listed in an
expanded paper (Mason & Wegner 1970).
In the present survey 34 species of fish were collected from the
Pine River drainage and 29 species from the Popple River drainage.
Examples of these have been placed in the collections of the
Museum of Natural History at the University of Wisconsin—
Stevens Point.
Materials and Methods
The fish were captured by seines 10 to 25 feet long with *4"
bar mesh, or with electrofishing equipment. Two complete electro¬
fishing units were used (one for each collecting crew). One unit
was a C & H alternating current generator rated for 1000 watts
and 120 volts at 8.5 amperes. The other was a Sears Alternator
Portable light plant rated for 900 watts and 115 volts at 7.8 am¬
peres. These were used from a boat or from the bank. One hundred-
foot electrode cords enabled shocking in particularly brushy or
1 This is Paper No. 7 in the series, “Studies on the Pine-Popple Wild Rivers
Area of Northeastern Wisconsin”. Received Nov. 28, 1971.
2 Curator of Pishes, Museum of Natural History, University of Wisconsin — •
Stevens Point, Stevens Point, Wisconsin. 54481
309
310 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
inaccessible areas. The electrodes themselves were fashioned from
heavy gauge copper wire forming long oblong hoops upon which
V^-inch hardware cloth had been sweated.
The method of capture was determined upon arrival at the
collection site. Frequently both seines and shocker were used.
Collection time at a site averaged two hours. All possible niches
were sampled and the captured fish placed in jars containing
10% formalin. Most game fish and large non-game forms were
returned to the water, but a record was kept of all fish captured.
Station data were placed on cards and then were inserted into
the jars long with the fish.
All the preserved fish were brought back to the laboratory at
Stevens Point where they were rinsed, sorted and identified to
species. Examples of all species were placed into the collections
of the Museum of Natural History at Stevens Point.
Physical and chemical data of each stream site were taken during
or immediately following the fish collections. These are recorded
in Tables III and IV.
Distribution of Fishes
Collection sites are listed in Table I (Pine River basin) and in
Table II (Popple River basin) . All collection sites are numbered on
Map 1. When used in the annotated list they are coded as follows:
Pi — Pine River basin
Po — Popple River basin
Each basin has a separate numbering system. After the name
of each species I have indicated by code and numbers all of the
stations where it was captured. If one species appeared at several
consecutive collecting sites, the first and the last are separated
by a hyphen; e.g., Pi 23-26 represents capture at sites 23, 24, 25
and 26 within the Pine River basin.
Frequency represents percentage of stations within the basin
at which that species of fish appeared. E.g., the bluntnose minnow
appeared at 8 out of a total of 38 stations in the Pine basin with
a frequency of 21%.
Forty-two species of fish are known from the Pine-Popple
basin. These are listed below. Another 6 species have been captured
near the Pine-Popple basin and may — with more intensive study—
be shown to occur there. These are listed under PROBLEMATICAL
SPECIES.
The common and scientific names used follow Bailey (1970).
1972] Becker — List of the Fishes of the Pine-Popple Basin
311
basins. The. collecting stations that were selected art in two series , one for each basin , numbered
consecutively from the mouth of each river toward its headwaters. A special line on the map marks
the divide between the watersheds of the two rivers. The location of the Pine-Popple Wild
Rivers Jirea in the State of Wisconsin is shown in the small inset map at the right.
312 Wisconsin Academy of Sciences , Arts and Letters ,[VoL 60
Table I. Pine River Basin Collections.
Information regarding each collecting station listed below is given in the
following sequence: (1) site number, (2) river, creek or lake from which col¬
lection was taken, (3) location of station by section, town and range, (4)
county in which station is located, (5) date of collection, (6) collecting tech¬
nique used, (7) names of collectors. Map 1 shows location of stations.
Pi 1 Pine R., Sec 24 T39N R18E, Florence Co., June 24, 1965, 25 ft. seine,
K. and D. Becker, J. Copp.
Pi 2 Pine Cr., Sec. 25 T39N R18E, Florence Co., June 25, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Pi 3 Pine R., Sec. 22 T39N R18E, Florence Co., June 22, 1965, 25 ft. seine
K. and D. Becker, J. Copp.
Pi 4 Halls Cr., Sec. 31 T39N R18E, Florence Co., July 2, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Pi 5 Halls Cr., Sec. 11 T38N R17E, Florence Co., July 2, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Pi 6 Small tributary of Pine R., Sec. 8 T39N R18E, Florence Co., June 23,
1965, electrofishing, K. and D. Becker, J. Copp.
Pi 7 Creek connecting Sealion and Grass L., Sec 2 T39N R17E, Florence
Co., June 22, 1965, electrofishing, K. and D. Becker, J. Copp.
Pi 8 Pine R., Sec. 15 T39N R17E, Florence Co., June 25, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Pi 9 Seven Mile Cr., Secs 27 & 33 T40N R17E, Florence Co., June 23, 1965,
electrofishing, K. and D. Becker, J. Copp.
Pi 10 Wakefield Cr., Sec. 25 T40N R16E, Florence Co., June 22, 1965, elec¬
trofishing, K. and D. Becker, J. Copp.
Pi 11 Pine R., Sec. 1 T39N R15E, Florence Co., June 13, 1965, 25 ft. seine,
K. and D. Becker, J. Copp; — July 22-23, 1967, seines, C. Long.
Pi 12 Johnson Cr., Sec. 21 T40N R15E, Florence Co., June 14, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Pi 13 Halsey L., Sec. 21 T39N R15E, Florence Co., June 15, 1965, 10 ft. and
25 ft. seines, K. Becker and J. Copp.
Pi 14 Long L. outlet, Sec. 19 T39N R15E, Florence Co., June 14, 1965, 25 ft
seine, K. Becker and J. Copp.
Pi 15 Coldwater Cr., Secs. 25 & 26 T39N R14E, Forest Co., June 27, 1966,
G. Becker team.
Pi 16 Pine R., Secs. 5 & 6 T39N R15E, Florence Co., June 14, 1965. K. Becker,
J. Copp.
Pi 17 Lilypad Cr., SE1! Sec. 14 T40N R14E, Forest Co., June 27, 1966,
Pi 18 Lilypad Cr., NW% Sec. 20 T40N R14E, Forest Co., June 27, 1966,
E. Peters team.
Pi 19 Stevens L., Secs. 23 & 26 T40N R14E, Forest Co., June 28, 1966,
G. Becker team.
Pi 20 Meadowbrook Cr., SE^ Sec. 35 T40N R14E, Forest Co., June 28,
1966, electrofishing and seining, G. Becker team.
Pi 21 Pine R., NE1^ Sec. 12 T39N R14E, Forest Co., June 28, 1966,
G. Becker team.
Pi 22 Bastile L., S% Sec. 28 T39N R14E, Forest Co., June 29, 1966, E. Pe-
Pi 23 Kingstone Cr., NE1^ Sec. 21 T39N R14E, Forest Co., June 27, 1966,
G. Becker team.
Pi 24 Pine Cr., Secs. 35 & 36 T39N R13E, Forest Co., June 27, 1966,
G. Becker team.
1972] Becker — List of the Fishes of the Pine-Popple Basin 313
Table I. Pine River Basin Collections. (Continued)
Pi 25 Jones Cr., Secs. 13 & 24 T39N RISE, Forest Co., June 24, 1966,
G. Becker team.
Pi 26 Sawyer Cr., SW % Sec. 14 T39N R13E, Forest Co., June 24, 1966,
G. Becker team.
Pi 27 Pine R., Secs. 25, 35, & 36 T40N R13E, Forest Co., June 24, 1966,
E. Peters team.
Pi 28 McDonald Cr., NE1^ Sec. 17 T39N R13E, Forest Co., June 27, 1966,
E. Peters team.
Pi 29 South Branch Pine R., NW1^ Sec. 30 T39N R13E, Forest Co., June
24, 1966, E. Peters team.
Pi 30 Kimball Cr., Secs. 27 & 28 T39N R12E, Forest Co., June 23, 1966,
Pi 31 McDonald Cr., NW% Sec. 12 T39N R12E, Forest Co., June 23, 1966,
G. Becker team.
Pi 32 Pine R., NW1! Sec. 21 T40N R13E, Forest Co., June 25, 1966, E. Pe¬
ters team.
Pi 33 Howell L. and outlet, Sec. 13 T40N R12E, Forest Co., June 29, 1966,
G. Becker team.
Pi 34 Pine R. (outlet of Butternut L.) , N% Sec. 26 T40N R12E, Forest Co.,
June 23, 1966, G. Becker team.
Pi 35 Butternut L., SW1^ Sec. 34 T40N R12E, Forest Co., June 29, 1966, gill
nets (100 ft. 1 in. bar, 100 ft. IV2 in. bar) set at depth of 25-30 ft.,
60-80 yds. off southwest shore, G. Becker, P. Holden.
Pi 36 Butternut L., SW^ Sec. 28 T40N R12E, Forest Co., June 23, 1966,
seines, G. Becker team.
Pi 37 Franklin L., SW% Sec. 21 T40N R12E, Forest Co., June 23, 1966,
seines, E. Peters team.
Pi 38 Franklin L., NE% Sec. 16 T40N R12E, Forest Co., June 23, 1966,
seines, E. Peters team.
Table II. Popple River Basin Collections.
The sequence of items in each of the site entries listed below are in the
same order as in Table I: site number; stream or lake being surveyed; loca¬
tion by section, town and range; county in which station is located; date of
collection; collecting technique used; names of collectors. See Map 1 for loca¬
tion of each site within the Popple River drainage basin.
Po 1 Lamon-Tangue Cr., Sec. 4 T38N R17E, Florence County, July 1, 1965,
10 ft. seine, K. and D. Becker, J. Copp.
Po 2 Lamon-Tangue Cr., Sec. 28 T38N R17E, Florence Co., June 20, 1965,
K. and D. Becker, J. Copp.
Po 3 Hilbert L., NW% Sec. 8 T37N R17N, Marinette Co., June 21, 1966.
Po 4 Lunds Cr., Sec. 21 T38N R17E, Florence Co., July 1, 1965, electro¬
fishing, K. and D. Becker.
Po 5 Woods Cr., Secs. 28 & 29 T39N R17E, Florence Co., June 30, 1965,
electrofishing, K. and D. Becker.
Po 6 Price L., Sec. 17 T39N R17E, Florence Co., June 28, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Po 7 Woods Cr., Sec. 23 T39N R16E, Florence Co., June 28, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
Po 8 Woods Cr., Sec. 25 T39N R15E, Florence Co., June 21, 1965, electro¬
fishing, K. and D. Becker, J. Copp.
314 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Table II. Popple River Basin Collections. (Continued)
Po 9 Popple R., Sec. 5 T38N R17E, Florence Co., June 30, 1965, electro*
fishing*, K. and D. Becker.
Po 10 Popple R., Sec. 23 T38N R16E, Florence Co., June 21, 1965, electro¬
fishing and 25 ft. seine, K. and D. Becker, J. Copp.
Po 11 Artificial L., Sec. 26 T38N R16E, Florence Co., June 18, 1965, 25 ft.
seine, K. Becker, J. Copp.
Po 12 Rock Cr., Sec. 29 T38N R16E, Florence Co., June 18, 1965, electro¬
fishing, K. Becker, J. Copp.
Po 13 Laura L., NW% Sec. 9 T37N R16E, Forest Co., June 30, 1966,
G. Becker team.
Po 14 South Branch Popple R., Sec. 19 T38N R16E, Florence Co., June 18,
1965, electrofishing, K. Becker, J. Copp.
Po 15 Simpson Cr., NW*4 Sec. 1 T37N R15E, Forest Co., June 30, 1966,
25 ft. seine, G. Becker team.
Po 16 Ross L. and outlet, SW*4 Sec. 16 T38N R15E, Forest Co., June 30,
1966, G. Becker team.
Po 17 South Branch Popple R., Sec. 26 T38N R15E, Florence Co., June 16,
1965, 10 and 25 ft. seines, K. Becker, J. Copp.
Po 18 Popple R., Sec. 19 T38N R16 E, Florence Co., June 17, 1965, electro¬
fishing and 10 ft. seine, K. Becker, J. Copp.
Po 19 Morgan Cr., Sec. 19 T38N R16E, Florence Co., June 17, 1965, electro-
fishing, K. Becker, J. Copp.
Po 20 North Branch Popple R., Sec. 22 T38N R15E, Florence Co., June 16,
1965, 25 ft. seine, K. Becker, J. Copp.
Po 21 Popple R., Sec. 6 T38N R15E, Florence Co., June 16, 1965, seine,
K. Becker, J. Copp.
Po 22 Little Popple R., SE1^ Sec. 27 T38N R14E, Forest Co., June 29, 1966,
E. Peters team.
Po 23 Popple R., Secs. 4 & 8 T38N R14E, Forest Co., June 29, 1966, E. Pe¬
ters team.
Po 24 Popple R., NW% Sec. 12 T38N R13E, Forest Co., June 24, 1966,
G. Becker team.
Salmonidae — trouts, whitefishes
Lake Whitefish — Coregonus clupeaformis (Mitchill).
Reported from Keyes L. (DNR files — Woodruff).
Cisco — Leucichthys artedii Lesueur.
Reported from Butternut, Franklin, and Keyes lakes (Wis.
Cons. Dept. 1964).
Rainbow trout — Salmo gairdneri Richardson.
Pi 11 ; Po 9, 10, 14. These rainbows were taken in 1965. Accord¬
ing to Mason (1966) 1,900 fingerlings were stocked in Pine R.
during the summer of 1966. Between September and October 1967
he captured 32 by electrofishing in the mainstem between the Pine
R. flowage and the juncture of the North and South Branches
(Mason and Wegner 1970). Reported from Keyes and Anna lakes
(DNR files— Woodruff).
1972] Becker — List of the Fishes of the Pine-Popple Basin 315
Brown trout — Salmo trutta Linnaeus.
Pi 11 ; Po 9, 10, 22, 24. Brown trout distribution and incidence
of marked and unmarked trout is documented by Mason (1966)
and Mason and Wegner (1970).
Brook trout — Salvelinus fontinalis (Mitchill).
Pi 4, 11, 21, 23-26, 28, 30; Po 1, 2, 4, 5, 7-9, 14, 19, 22, 24. Our
only native trout, the brook trout, occurred in 21% of the stations
sampled in the Pine R. basin and 49% in the Popple R. basin. See
Mason (1966) and Mason and Wegner (1970) for additional in¬
formation. Reported from Sand and Patten lakes (DNR files — •
Woodruff) .
Umbridae — mudminnows
Central mudminnow — Umbra limi (Kirtland).
Pi 2 4-6, 8, 9, 15, 18, 19, 23-25, 27-30, 34; Po 1, 4, 9, 12, 14, 24.
Frequency in the Pine basin was 45% ; in the Popple basin 25%.
Reported from McKinney and Two Sisters lakes (DNR files —
Woodruff) .
Esocidae — pikes
Northern pike — Esox Indus Linnaeus.
Pi 1, 7, 8, 15, 21. Frequency in the Pine basin was 13%. Eight
individuals were captured at Pi 1. Reported from Pine, Butternut,
Bastile, Stevens, Two Sisters, Harriet, Lilypad, Long, Fay, Lost,
Lauterman, Sealion, Keyes, Emily, Pine R. flowage, Halsey, Reis-
ner, Lake of Dreams, Bessie Babbett, Loon lakes (DNR files—
Woodruff). Also reported from Forest, Halsey, Seidel, Cosgrove,
Hilbert lakes (Wis. Cons. Dept. 1964).
Muskelunge — Esox masquinongy Mitchill.
Reported in Sealion and Emily lakes and from the Pine R. flow-
age (Wis. Cons. Dept. 1964). Report from Quartz L. (DNR file —
Woodruff) .
Cyprinidae — minnows
Brassy minnow — Hybognathus hankinsoni Hubbs.
Pi 3-6, 14, 18, 20, 25, 28, 30, 31, 37; Po 1, 4, 11, 12, 15, 17,
19-21, 23, 24. Frequency in the Pine basin was 32% in the Popple
basin 46%. Largest numbers were captured at Pi 18 (80) and Po
21 (153).
Hornyhead chub — Nocomis biguttatus (Kirtland).
Pi 1, 3-5, 7, 8, 11, 14, 16, 17, 21, 27, 32, 34, 38; Po 1, 9, 10, 14,
18-21, 23. Frequency in the Pine basin was 39%; in the Popple
basin, 38%. Largest numbers captured were 180 and 131 at Po
316 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Table III. Pine River Basin — Station Data.
1972] Becker — List of the Fishes of the Pine-Popple Basin B17
20 and 23 respectively. Condition of both sexes at Pi 1, 7, and 34
indicates spawning- toward the end of June 1965 and 1966.
Two hornyhead nests were observed just above the outlet of
Butternut L. (Pi 34) on June 23, 1966. One nest was constructed
in 1 ft. of water, the other in 3 feet. Both nests were of pebbles
!/2 to 1 inch in diameter, diameter of nests 18 inches and mounds
up to 3 inches in height. Two females give the following data:
total weight 25.22 and 23.87 gms., ovarian weight 3.00 (11.9%)
and 2.10 gms. (8.8%) respectively.
Common shiner — N o tropis cornutus (Mitchill).
Pi 1, 3-8, 11-14, 16-18, 20, 21, 27-29, 31-38; Po 1, 5, 9-12, 14, 15,
17-24. Frequency in the Pine basin was 71% ; in the Popple basin,
67%. A lake as well as a stream inhabitant in this sector of the
state. Largest numbers taken at Pi 3 (368), Po (603), Po 21
(637). This is the most abundant species in the Pine-Popple basin
and over 5300 were collected in this survey.
Blackchin shiner — Notropis heterodon (Cope).
Pi 5, 13, 14. This species is generally taken in lakes or lake
outlets. Twenty-four specimens were captured at Pi 5 ; 17 at Pi 13 ;
1 at Pi 14.
Blacknose shiner — Notropis heterolepis Eigenmann & Eigenmann.
Pi 4, 5, 8, 9, 12-14, 18, 20, 31, 33, 37; Po 1, 3, 10, 11, 15, 16,
19, 20, 23. Frequency in the Pine basin was 32%; in the Popple
basin, 38%. This species is associated with lakes or slow-moving
streams. Largest numbers were taken at Pi 13 (186) and Po 11
(605). At the latter site a gravid female with loose eggs appeared
ready for spawning.
Golden shiner — Notemigonus crysoleucas (Mitchill).
Pi 4, 11, 13, 14, 29, 33, 37, 38; Po 6, 10, 11. Frequency in the
Pine basin was 21% ; in the Popple basin, 13%. Largest numbers
captured were 77 (Pi 14) and 319 (Po 11). Terry McKnight, DNR
biologist, reports individuals 6 to 8 inches long from Patten L.
Bluntnose minnow — Pimephales notatus (Rafinesque) .
Pi 5, 6, 13, 14, 32, 34, 37, 38; Po 3, 5, 10, 15, 20, 21, 23. Fre¬
quency in the Pine Basin was 21%; in the Popple basin, 29%.
Largest numbers were captured at Pi 14 (138 plus), Pi 37 (276),
Pi 38 (566), Po 3 (126). Reported from Patten L. (DNR files — -
Woodruff) .
Fathead minnow — Pimephales promelas Rafinesque.
Pi 6, 8, 11, 13, 18, 20, 25, 29, 31 ; Po 1, 8, 10, 11, 14-17, 19, 21,
23. Frequency in the Pine basin was 24%; in the Popple basin,
46%. Largest numbers were captured at Pi 31 (36) and Po 16
(48). The fathead and bluntnose occurred together only inf re-
318 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
quently : twice in the Pine basin samples and four times in the
Popple basin samples. Reported from Frog. L. (DNR files—
Woodruff) .
Northern redbelly dace — Phoxinus eos Cope.
Pi 4-6, 8-10, 13, 14, 18, 20, 28, 30, 32-34, 37; Po 1, 2, 4,
10-12, 14, 15, 17-24. Frequency in the Pine basin was 42% ; in the
Popple basin, 67%. Largest numbers were taken at Pi 10 (137),
Po 15 (411), Po 19 (358). A female, 2.52 gm, taken at Pi 33
(June 29, 1966) had ovaries weighing 0.13 (5.1% of body weight).
Deep red flanks were noted on 3 males taken June 28, 1966 (Pi 20)
from Meadowbrook Cr.
1972] Becker — List of the Fishes of the Pine-Popple Basin 319
Finescale dace — Phoxinus neogaeus Cope.
Pi 2, 6, 9, 10, 12, 18, 20, 26, 31, 34; Po 1, 4, 11, 12, 14, 15, 17,
19-22, 24. Frequency in the Pine basin was 26%; in the Popple
basin, 50%. Stations at which 25 or more were captured are Po 1
(26), 11 (25), 19 (35), 21 (38). Prefers slow-moving water over
a wide variety of soft bottoms. The 10 collection sites in the Pine
basin can be summarized as follows :
3 sites stream width 2' to T , water depth 2" to 13"
5 sites stream width V to 13.7', water depth 3" to 36
2 sites— bog ponds
This species appears to be best represented in the state of Wiscon¬
sin in the counties of Florence, Forest and Marinette.
Blacknose dac e—Rhinichthys atratulus (Hermann).
Pi 4, 5, 8-12, 15-18, 20, 21, 23, 25, 27-30; Po 1, 5, 8-10, 12-15,
17-24. Frequency in the Pine basin was 50% ; in the Popple basin,
71%. Largest numbers captured were 106-plus (Pi 10) and 81 (Po
22). On June 18, 1965, males from the So. Branch Popple R. (Po
14) were still in breeding color and all the females had not spawned
as yet. On June 28, 1966, two females taken from Meadowbrook
Cr. (Pi 20) were gravid and eggs loose, showing spawning
readiness.
Longnose dac e—Rhinichthys cataractae (Valenciennes).
Pi 8, 11, 16, 20, 21, 25, 28; Po 5, 8-10, 14, 18. Frequency in the
Pine basin was 18% ; in the Popple basin 25%. Largest numbers
captured were 41 (Po 15) and 35-plus (Po 18)
Creek chub — Semotilus atromaculatus (Mitchill).
Pi 2, 4-6, 8-10, 12, 14-18, 20, 21, 23-25, 27-32, 34, 37; Po 1, 5,
9, 10, 12, 14, 15, 17, 18, 20-24. Frequency in the Pine basin was
68% ; in the Popple basin, 58%. Largest number captured was 67
(Pi 20). Breeding tubercles present on males from Pi 6 (June 23,
1965) although the females were no longer gravid.
The range and average number of lateral line scales in creek
chubs from northeastern Wisconsin is high. A Wakefield Cr. sample
of 35 ranged from 52 to 64 with an average count of 58.43. Of
these, eight individuals (23%) had lateral line scale counts of more
than 60. A Popple R. sample of 61 ranged from 50 to 65 with an
average of 58.48. Of these, 18 individuals (30%) had lateral line
counts of more than 60. For comparison, a central Wisconsin
(Portage Co.) sample had a range of 55 to 63 with an average of
58.03. A southern Wisconsin (LaFayette Co.) sample had a range
of 49 to 62 with an average of 55.1.
Pearl dace* — Semotilus margarita (Cope).
Pi 6, 9-12, 17, 20, 25, 26, 29-32; Po 1, 4, 6, 10-12, 14, 15, 17, 19-24.
Frequency in the Pine basin was 34 % ; in the Popple basin 63 % .
320 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Largest numbers captured were 186 (Po 22) and (Pi 20) . A Febru¬
ary, 1968, DNR collection indicated this species from Patten L.
Catostomidae - — suckers
White sucker — Catostomus commersoni (Lacepede) .
Pi 3-7, 9, 13, 14, 16-21, 25, 27, 29, 32, 34-37; Po 1, 5, 7, 9-12, 14-24.
Frequency in the Pine basin was 58% ; in the Popple basin, 75%.
The young-of-the-year taken June 22, 1965, at Pi 3 had total length
of 17-23 mm. Largest numbers of suckers captured were 157 (Po
23) and 92 (Po 24). Reported from Butternut, Franklin, Three
Johns, Quartz, Fay, Lost, Perch, Emily, Frog, Elwood, Patten, and
Hilbert lakes (DNR files — Woodruff).
Ictaluridae — freshwater cat fishes
Black bullhead — Ictalurus melas (Rafinesque) .
Pi 5, 7, 9, 37 ; Po 9-12, 16, 20, 23. Frequency in the Pine basin
was 11%; in the Popple basin, 29%. No more than 3 individuals
were taken at any station.
Yellow bullhead — Ictalurus natalis (Lesueur).
Pi 7, 9. One individual was captured at Pi 7 and 4 at Pi 9.
Brown bullhead — Ictalurus nebulosus (Lesueur).
Pi 7, 14. Three individuals were captured at Pi 7 and 1 at Pi 14.
This species is frequently found in inlet or outlet streams near
lakes.
Tadpole madtom — Noturus gyrinus (Mitchill).
Pi 8, 13, 14. Ten individuals were captured at Pi 8 and singles
at the other two sites.
Gadidae — codfishes
Burbot — Lota lota (Linnaeus).
John Mason (letter,, June 18, 1969) reports that he had recently
captured 4 or 5 individuals from the lower Pine river SW 1/4 Sec. 23
and SE % Sec. 23 T39N R18E.
Gasterosteidae — sticklebacks
Brook stickleback — Culaea inconstans (Kirtland).
Pi 2, 6, 9, 10, 12, 18, 20, 23-25, 28-31 ; Po 1, 4, 8, 10, 12, 14-17,
19-21, 23, 24. Frequency in the Pine basin was 37 % ; in the Popple
basin, 58%. Largest numbers were captured at Pi 25 (24), Pi 31
(24) , Po 17 (64) , Po 20 (25) , Po 21 (32) , Po 24 (87) .
C entrarchidae — sun fishes
Rock bass — Ambloylites ruyestris (Rafinesque)
Po 5. Reported from Sealion, Keyes, Pine R. flowage, Cosgrove,
Anna, Elwood, Patten lakes (DNR files — Woodruff).
1972] Becker — List of the Fishes of the Pine-Popple Basin 321
Green sunfish — Lepomis cyanellus Rafinesque.
Reported from Lost, Emily, Cosgrove, Anna, Bessie Babbet,
Grass, Bass (Sec. 15 T40N R17E), Oneata lakes (DNR files —
Woodruff).
Pumpkinseed — Lepomis gibbosus (Linnaeus).
Pi 4, 6-8, 11, 14; Po 3, 6, 11. Reported from Franklin, Lost,
Perch, Emily, Halsey, Reisner, Lake of Dreams, Patten, Hilbert
lakes (DNR files— — Woodruff) .
Bluegill — Lepomis machrochirus Rafinesque.
Pi 7=9, 14, 16, 33, 37, 38; Po 3, 6, 12. Reported from Franklin,
Three Johns, Quartz, McKinney, Fay, Porcupine, Lauterman,
Keyes, Emily, Cosgrove, Anna, Halsey, Reisner, Lake of Dreams,
Loon, White Bass, Elwood, Savage, Price, Oneata, Patten, Hilbert
lakes (DNR files-==Woodruff).
Smallmouth bass — Micropterus dolomieui Lacepede.
Pi 35, 36, 37. The smallmouth is an important game species in
Butternut and Franklin lakes. Also reported from Quartz, Keyes,
Cosgrove and Elwood lakes (DNR files — Woodruff). The Wisconsin
Department of Natural Resources (1968) reports it in the Pine R.
from La Salle Falls downstream to its juncture with the Menomi¬
nee R.
Largemouth bass — Micropterus salmoides (Lacepede).
Pi 5, 16, 19, 37 ; Po 3, 5, 12. Reported from Butternut, Franklin,
Three Johns, Stevens, Harriet, Harmony, Indian Camp, Howell,
Ritter, Rogers, Wapoose, Long, Fay, Lost, Porcupine, Lauterman,
Keyes, Cosgrove, Anna, Morgan, Halsey, Reisner, Lake of Dreams,
Bessie Babbett, Grass, Loon, White Bass, Bass (Sec. 15 T40N
R17E), Elwood, Nona, Savage, Price, Oneata, Patten, Hilbert lakes
(DNR files — Woodruff).
Black crappie — Pomoxis nigromaculatus (Lesueur).
Pi 14, 33. Reported from Stevens, Fay, Porcupine, Perch, Lauter¬
man, Keyes, Emily, Halsey lakes (DNR files — Woodruff).
Percidae — perches
Iowa darter — Etheostoma exile (Girard)
Pi 6=9, 13, 14; Po 2, 11, 16, 23. Twenty were captured at Pi 13.
Recorded from Patten L. (DNR files — Woodruff) . Shallows of lakes
and slow-moving streams.
Johnny darter — Etheostoma nigrum Rafinesque.
Pi 1, 8, 11, 13, 17, 18, 20, 21, 25, 28, 29, 32, 33, 36, 37; Po 5;
9, 10, 14, 15, 17-24. Frequency in the Pine basin was 39% ; in the
Popple basin, 54%. Largest numbers were captured at Pi 37 (24)
and Po 22 (35). Recorded from Patten L. (DNR files — Woodruff).
322 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
Yellow perch — Perea flavescens (Mitchill).
Pi 1, 3, 11, 13-15, 17, 19, 22, 27, 33, 34, 36-38; Po 6, 10, 11, 13, 14.
Frequency in the Pine basin was 39% ; in the Popple basin, 21%.
Common at most sites where found. Reported from Butternut,
Franklin, Bastile, Three Johns, Stevens, Quartz, McKinney, Two
Sisters, Harriet, Lilypad, Indian Camp, Howell, Wapoose, Fay,
Lost, Porcupine, Perch, Lauterman, Keyes, Emily, Pine R. flowage,
Cosgrove, Halsey, Reisner, Lake of Dreams, White Bass, Bass
(Sec. 15 T40N R17E), Elwood, La Fave, Oneata, Patten, Hilbert
lakes (DNR files- — Woodruff) .
Logperch — Percina caprodes (Rafinesque) .
John Mason (letter, June 18, 1969) reported recently taking
“quite a few” from below the Pine River power dam in Secs. 22
and 23 T39N R18E.
Walleye — Stizostedion vitreum vitreum (Mitchell) .
Pi 35. Reported from Butternut, Stevens, Laura, Long, Forest,
Halsey, Fay, Sealion, Keyes, Emily, Cosgrove, Hilbert lakes (Wis.
Cons. Dept. 1964). Reported from Harmony, Howell, Wapoose,
Pine R. flowage, Frog, Patten lakes (DNR files— Woodruff) .
Cottidae — sculping
Mottled sculpin — Coitus bairdi Girard.
Pi 2, 4, 5, 8, 11-13, 15-21, 22-30, 32, 34; Po 2, 4, 5, 7-10, 14, 15,
17, 22, 24. Frequency in the Pine basin was 66%; in the Popple
basin, 67%. Largest numbers were captured at Pi 20 (88) , Pi 28
(75), and Po 5 (43) . An individual 4%// total length from Pi 28
had a 2-inch fish (mudminnow?) in its stomach.
Problematical Fishes
Following are species not recorded from the Pine-Popple basin
but which may be expected on the basis of proximity.
Northern brook lamprey— Ichthyomyzon fossor Reighard &
Cummins.
An adult was taken June 21, 1966, near the mouth of the Pike R.,
Sec. 4 T34N R21E, Marinette Co.
Silver lamprey — Ichthyomyzon unicuspis Hubbs & Trautman.
On July 19, 1970, several adults were taken from fish in the
Menominee R., T34N R21E, Marinette Co., by Terry McKnight,
DNR biologist.
Lake sturgeon — Acipenser fulvescens Rafinesque,
This species is a common inhabitant of the Marinette Co. sector
of the Menominee R. into which the Pike R. empties. The range
map by Priegel and Wirt h (1971) shows this species in the Menomi¬
nee R. through its entire contact with Florence Co, If the sturgeon
1972] Becker — List of the Fishes of the Pine-Popple Basin 323
still occurs in the Menominee R. at the point where the Pine R.
enters, it may be expected in the lower Pine R. between the Pine R.
flowage and its mouth.
Longnose sucker — Catostomus catostomus (Forster).
This species is present in Kentuck L., Forest Co., less than a
mile from the Pine-Popple drainage. A specimen was given to the
Univ. of Wisconsin— Stevens Point Museum by Terry McKnight,
DNR biologist. It was netted in May, 1970.
Rosyface shiner — Notropis rubellus (Agassiz).
Greene (1935) recorded this species at the juncture of the Pike
and Menominee rivers in Marinette Co.
Blackside darter — Percina maculata (Girard).
In 1966 I captured this species from near the mouth of the
Pike R., Marinette Co.
The Fishery, — Past, Present, Future
Although much of the area is wild and both the Pine and Popple
rivers have been designated “wilderness rivers”, there is much
evidence that many profound changes have taken place in the
basin which detract from their wildness. These changes have
altered the fish composition of the basin.
In discussing the trend in trout fishing with a number of resi¬
dents who have fished these waters over the years, it has become
evident that trout fishing has fallen off. Within recent years trout
fishing for good-sized trout was excellent at LaSalle Falls. This
is no longer the case.
After the first few weeks of trout fishing in spring the fishing
drops off rapidly. Water temperatures of the mainstem, especially
the Pine River, respond readily to the rising air temperatures
(Table 3 & 4). With a few days of warm spring weather fly
fishing for trout ends. Water temperatures become critical for the
brown trout and especially the brook trout.
Both the Pine and Popple rivers begin as slow-moving non-trout
streams. With the increase in gradient and the admixture of numer¬
ous springs and spring-fed tributaries the mainstems soon become
trout water. Originally the only trout were native brook trout.
Later the temperature-tolerant brown was introduced.
At the turn of the century the Pine-Popple basin became cut¬
over. There are many openings visible from the Pine River which
are still reminiscent of this activity. Old abandoned farms provide
additional open areas. Cottages, summer homes, campsites, road
bridges, and canoe landings pushed back the vegetation and let in
the sunlight. Man-made dams on mainstem and tributaries do the
same while allowing the water behind the dams to warm up
sharply.
324 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
Opening the forests provided an important wedge for the beaver
which became abundant in many Wisconsin trout streams in the
1930’s :
“. . . Their ponds caused warming of waters, chemical changes, siltation,
blocking of spawning runs, and decreased spring flow, according to those
people making investigations.” (Christenson et al. 1961)
The same source showed that in 1936 203 beaver dams were re¬
moved from Jones Creek of the upper Pine River basin alone. One
hundred and ten dams were removed in Florence county with 58
from La Montagne Creek. Although La Montagne Creek lies just
north and outside of the Pine River basin the numbers are indica¬
tive of the pressure which the beaver were placing on the trout
streams in this region of the state.
From 1945 to 1952 beaver reached their all-time peak of abun¬
dance. Special trapping seasons reduced their numbers but “by 1958
and 1959, the beaver problem on trout streams had again reached
such proportions that NEA fisheries personnel became concerned
and requested a policy directive to cope with the situation.” Beaver
activity was high on the South Branch Popple River, Popple River
(main), McDonald Creek, Meadowbrook Creek, South Branch Pine
River, North Branch Pine River, Pine River (main), and Kimball
Creek.
The upshot of man’s activities combined with that of the beaver
was to put the trout fishery in jeopardy. The result is summarized
by Mason and Wegner (1970) :
“Warmwater fish predominated in the lower Pine. Trout were found in
all three rivers (Pine, Popple, Pike), but productivity is restricted by low
water fertility, high summer temperatures and cold winter temperatures.”
We may be dealing with wild rivers ; however, fish species com¬
position has changed with the use of fish poisons on a number of
waters within the basin. According to Department of Natural Re¬
sources files at Woodruff, the following lakes in the Pine-Popple
basin have been poisoned: Three Johns (1965), McKinley (1968),
Two Sisters (partially treated, 1964), Harriet (treated for perch,
1966), Sand (1962), and Morgan (1962). Treated lakes frequently
show little diversity in fish speciation. After treatment the Depart¬
ment stocks only a few game fishes, panfish, and one or two species
of forage fishes. The natural complement of fishes has been removed
through the poisoning program. Disadvantages in such a program
are summarized by Becker (1971).
That fish poisoning programs should continue in a wild river
system is inexcusable unless there is built into the system a well-
designed preliminary research program followed by a well thought-
out management program. Fish poisons are useful tools, but they
1972] Becker — List of the Fishes of the Pine-Popple Basin 325
are a last-ditch measure. Effects upon the aquatic environment are
devastating and could lead to serious ecological imbalance. I should
like to suggest that the use of fish poisons be entirely banned in
wild river basins. Poisons produce problems, and, unfortunately
there are too many possible side effects to the solution of which
we have not even begun to direct ourselves.
The Pine and Popple are not free from airborne chemical con¬
taminants. Pesticide contamination in the form of DDT and dieldrin
residues have been detected in trout taken from these rivers (Klein-
ert et al. 1967). Recent reports indicate that DDT tests may be
confused with PCB’s (polycholorinated biphenyls) which also are
highly persistent industrial chemicals of widespread distribution.
The point is that although the contamination of fish with these
toxic chemicals is low, civilization’s pollutants are reaching into
our most isolated wilderness areas. Our concern must extend be¬
yond mere provision for wilderness vegetation strips along the
banks. If at all possible, the technological crunch should rightfully
be stemmed well behind the wilderness Maginot line.
Continued pressure is being placed by landowners and organiza¬
tions upon the Department of Natural Resources to allow dam¬
ming up of tributary streams in the Pine and Popple basin. Justi¬
fication for such action is cited as “recreational area, waterfowl
resting and nesting area” and the like. Several permits for such
impoundments have been granted with the Department agent stat¬
ing that such action will constitute no damage to the fishery on
the mainstem. Such reasoning is questionable when we consider
the damaging effects of ponds on increasing the temperature of
outlet waters. Beaver ponds are an example. A number of these
warmed-up tributaries are bound to raise the water temperature
of the mainstem Pine and Popple. It is inadvisable to allow further
damming even in the lower Pine River below the La Salle Falls
area which is considered non-trout water until adequate study has
been made of water conditions necessary to support the fishery
desired.
When the Pine, the Popple and the Pike were declared wild rivers
under Wis. Statute 30.26, Chapt. 363, Laws of 1965, the Wisconsin
Conservation Commission was empowered to draw up a plan for
its administration. On November 3, 1967, the Wis. Conservation
Commission (1967) adopted its policy on wild and scenic rivers
“In order to afford the people ... an opportunity to enjoy natural
streams” and because “it is in the interest of this state to preserve
some rivers in a free-flowing condition and to protect them from
development. . . .”
326 Wisconsin Academy of Sciences, Arts and Letters [Vol, 60
The preamble goes on to state that a wild river area
“is a stream or section of a stream, tributary, or river — and the related
adjacent lands — located in a sparsely populated, natural and rugged en-
vironment where the river is free-flowing and unpolluted or where the
river should he restored to such condition , in order to promote sound wa¬
ter conservation, and promote the public use and enjoyment of the scenic,
fish, wildlife, and outdoor recreation values.”
I have italicized the items in the above statement with which I
would take issue. If this means that we are going to allow the river
to revert to its wild or primitive state, if it means that we are going
to pull man and his devices back from the banks, that we are going
to remove as much as possible things such as buildings, roads,
campsites, power and telephone lines, then I can go along with the
statement. If pulling man back is synonymous with “restoring,”
then I can see eye-to-eye with the statement. But, “to promote
sound water conservation, and promote the public use and enjoy¬
ment . . imply man stepping into the act and “improving” on
what nature can do herself. Stabilizing eroding banks and restoring
bank cover are measures which promote sound water conservation.
A trained technician knows this and he has been hired for exercis¬
ing his expertise.
Do we want this kind of manipulation on a wild river? And, by
promoting the public use and enjoyment, by encouraging more use
by the public, are we protecting the wild river, the “natural and
rugged environment.” The more people using such a river, the
more evidence there will be of human presence, and, in fact, the
less “wild” it will be.
The State Department of Natural Resources, in order to provide
solace for purists has designated a zone known as the Primitive
Zone :
“A true natural wilderness zone devoid of ALL man-made efforts, devel¬
opments or improvements of any type, in the water, or within 400 feet
of the stream bank or to the visual horizon, whichever is greater, except
for statutory requirements of an emergency nature.”
Unfortunately only small fragments of the Pine and Popple are so
zoned.
The other zones are: Scenic and Aesthetic Management Zone,
Roadside Public Use Zone, Timber Management Zone, Agricultural
Zone, Developed Recreation Zone, Developed Zone. All of these un¬
fortunately allow wide latitude for manipulation by man.
Even more disappointing is the wild river plan adopted by the
Forest Service (U.S. Dept, of Agric. 1969). That portion of the
Pine River from its origin downstream to a point seven miles east
of State Hwy. 139 lies within the Nicolet National Forest. Within
the Forest, 65% of the land adjacent to the Pine River is in Na-
1972] Becker — List of the Fishes of the Pine-Popple Basin 327
tional Forest ownership. Hence, what happens on Forest lands will
pretty much determine what happens to the river.
In the Federal plan for this wild river we read (italics are mine) :
“Every effort will be made to maintain enjoyable visual conditions within
this Zone by perpetuating, restoring , or improving vegetative cover to
reflect a pleasing Forest environment. Within this Zone, the goal will be
to present a vegetative condition that appears undisturbed by man.”
Cannot nature “restore” or “improve” on her own? Is a wild river
one that simply “appears undisturbed by man?”
uCanoeable stretches of the Pine will be maintained to allow relatively
free passage. Maintenance will include the removal of fallen trees which
block the movement of canoes, and selective moving of key boulders .”
Does the canoeist, seeking out a wild river, want a manicured river
bed or would he prefer a true wilderness stream with all of its
hazards? How will such landscaping help the fishing for trout?
Don’t down logs and key boulders provide trout cover already?
The Pine River will be managed “to increase the likelihood of
observing wild animals. . . . Ruffed grouse, and especially deer ob¬
servations, will be greatly increased by enlarging and improving
existing wildlife openings along the River.” Do we need a Disney¬
land of nature just because a river has been declared wild? Won’t
opening up the landscape also accelerate warming of the waters,
thereby cancelling out hope of returning to primitive trout and
aquatic conditions?
The plan calls for blasting waterfowl potholes “in suitable marsh
areas in and adjacent to the Water Influence Zone.” Has any study
been made as to what effect this will have on the groundwater
temperatures next to the potholes and the ultimate effects on the
temperature of the water in the river?
The plan calls for possible use of “natural-appearing structures
to improve the fish habitat. . . .” There is no question that water
conditions for trout production can be improved through the use
of management devices. This has been well documented (White &
Brynildson 1967). However, won’t this foresake completely the
concept of a wild river?
In the plan there is no such thing as a no-cut zone along each
bank, except through the approval of the Forest Supervisor “if
needed to protect the Wild River character of the Pine.” Indeed,
logging will be permitted right up to the bank of the stream and
“All timber cutting and cultural work will be designed for the
primary purpose of improving the present and future aesthetic
value of this zone.” Just recently this regulation was violated by a
logging contractor. The matter was reported to the Forest Service
who admitted that the contractor was in error, that he had violated
328 Wisconsin Academy of Sciences, Arts and Letters [Vol. 60
the cutting principles and that accordingly he was to be fined up
to $100.
From the plan we learn that “Stabilization of eroding stream
banks and existing road ditches and back slopes will be top priority
work within the zone.” Also “Conifers and shrubs can be planted
along the Pine to stabilize the banks of the River.” Are we manag¬
ing a wild river or a trout stream in rural southern Wisconsin?
In all fairness, there are many excellent features of the Nicolet
National Forest Wild River Plan for the Pine River. I would ad¬
vocate adoption of many of these in future wild river planning.
But I am concerned with those features mentioned above which
disrupt and interfere with the mechanisms which nature uses for
establishing the primitive. I am disturbed by a federal logging pol¬
icy which will forever keep a managed forest lining most of the
banks of the Pine River. Primeval wilderness with downed trees
and large decaying logs play no role in the Forest Service’s wilder¬
ness program, at least none as it alfects the Pine River.
It almost appears that in setting policy for managing a wild
river, both the state and federal plans call for more management.
The federal plan is expanding its multiple-use program for forest
lands — meaning more activities for more people. The mentality
operating is one which equates wildness, the wilderness and the
primitive with “restoration.” We appear to be headed for another
Quetico-Superior wilderness slum. The Pine-Popple wilderness
streams will through over-management and overuse lose much of
their wilderness quality.
Sixty-nine percent of the trout captured by Mason and Wegner
(1970) were stocked brook, brown and rainbow trout, many of
them recently stocked fingerlings. At least some natural reproduc¬
tion in brook and brown trout occurs each year. Water tempera¬
tures which readily fluctuate in accordance with the air tempera¬
tures are undoubtedly the limiting factor in trout reproduction in
the mainstem.
The Forest Service’s wild river policies, many of which will
open rather than close the vegetative canopy, will operate against
the trout fishery in the mainstem. This is certain to happen unless
the mainstem is heavily managed for improving trout habitat.
There is little likelihood that management for improving trout
habitat will be extensive enough to improve the character of the
river for trout production and to offset the affects of the anticipated
multiple uses.
In summation I look for no improvement of fishing — rather a
gradual deterioration. This will continue until a radical change in
state and federal policy is instituted. At this time there appears
little hope that such a change will occur.
1972] Becker — List of the Fishes of the Pine-Popple Basin 329
Acknowledgments
Field work was done under my direction during the summers of
1965 and 1966. I am grateful to Kenneth Becker, who was in charge
of the 1965 collections, assisted by James Copp and Dale Becker.
My able assistants in 1966 were Dale Becker, Robert Fiehweg,
Paul Holden, Dan Keppie, John Palmisano, Ed Peters and David
Becker. Dr. Charles Long, director of the Museum of Natural
History — Stevens Point, gathered valuable data with his field class
in 1967. I am grateful to Jack Mason, Terry McKnight and Arthur
Oehmcke, all of the Department of Natural Resources — Woodruff
Headquarters, who supplied me with specimens and made available
to me their data and files. Map 1 was redrawn and lettered by Dr.
Lowell E. Noland.
References
Bailey, Reeve et al., 1970. A list of common and scientific names of fishes
from the United States and Canada. 3rd. ed. Amer. Fish. Soc. Spec. Pub¬
lication No. 6. 150 p.
Becker, George C., 1971. Preliminary survey of the fishes of the Rock River
basin, Wisconsin, with suggestions for management. Fauna & Flora Re¬
port #5, Museum of Natural History, Univ. of Wis. -Stevens Point. 16 p.
Christenson, L. M. et al., 1961. Beaver-trout-forest relationships. Wis.
Cons. Dept., Madison. 34 p. plus 2 appendices.
Greene, C. Willard, 1935. The distribution of Wisconsin fishes. Wis. Con¬
servation Comm., Madison. 235 p.
Kleinert, Stanton J., Paul E. Degurse, Thomas L. Wirth, and Linda C
Hall, 1967. DDT and dieldrin residues found in Wisconsin fishes from
the survey of 1966 (Preliminary report). Research Report No. 23 (Fish¬
eries), Research and Planning Div., Wis. Conservation Dept., Madison.
29 p. (mimeo.)
Mason, Jack, 1966. Annual report, wild rivers fish population studies for the
period July 1-December 21, 1966. Water Resources Section, Wis. Dept, of
Natural Resources. 56 p.
Mason, John W. and Gerald D. Wegner, 1970. Wild rivers fish populations
(Pine, Popple and Pike rivers). Research Report 58, Dept, of Natural
Resources, Madison. 42 p.
Priegel, Gordon R. and Thomas L. Wirth, 1971. The lake sturgeon — its life
history, ecology and management. Publ. 240-70. Wis. Dept, of Nat. Res.,
Madison. 20 p.
U. S. Dept, of Agriculture, 1969. Wild river plan — Pine river. Nicolet Na¬
tional Forest Headquarters, Rhinelander, Wis. 10 p. plus map.
Wis. Conservation Commission, 1967. Conservation commission policy on wild
and scenic rivers. Dept, of Natural Resources, Div. of Conservation, Madi¬
son. 6 p.
Wisconsin Conservation Dept., 1964. Wisconsin lakes. Publication 218-64,
Wis. Conservation Dept., Madison. 67 p.
BIOGRAPHIES
STEVEN BARTELL is a graduate of Lawrence University and is cur¬
rently a graduate student in the Department of Botany at the University of
Wisconsin, Madison. DR. SUMNER RICHMAN is Chairman, Department of
Biology, Lawrence University, Appleton, Wisconsin.
DR. GEORGE C. BECKER is Professor of Biology at Wisconsin State
University — Stevens Point. A former Vice President of Sciences of the Acad¬
emy, he has been the driving force behind the Wild Rivers Cooperative Re¬
search Project.
DR. ROBERT F. BROWNING is Assistant Professor of Biology at Ripon
College. His interest is the biology of invertebrates.
WILLIAM E. DICKINSON lives at 730 Euclid Avenue, Milwaukee.
DR. DONALD EMERSON is Professor of American Literature at the
University of Wisconsin — Milwaukee.
DR. KENNETH J. GRIEB is currently Associate Professor of History
and Coordinator of Latin American Studies at Wisconsin State University —
Oshkosh. This article was presented as a paper at the Fifth Annual Northern
Great Plains History Conference at the University of North Dakota, Octo¬
ber 23, 1970.
DR. RICHARD P. HOWMILLER is at the Center for Great Lakes Studies
at the University of Wisconsin — Milwaukee. G. M. LUDWIG is Curator of
Ecology at the Milwaukee Public Museum.
DR. GERHARD B. LEE is Associate Professor of Soil Science at the
University of Wisconsin, Madison. M. E. HORN is Senior Soil Scientist, Dames
and Moore, Consulting Engineers, Park Ridge, Illinois. The research was sup¬
ported by the College of Agricultural and Life Sciences, University of Wis¬
consin, Madison and by the Geological and Natural History Survey, Univer¬
sity Extension.
331
332 Wisconsin Academy of Sciences , Arts and Letters [Vol. 60
DR. ROBERT A. McCABE is Professor and Chairman of Wildlife Ecology
at the University of Wisconsin, Madison. He is a member of the research
advisory council for the Wisconsin Department of Natural Resources.
DR. TED J. McLAUGHLIN is Professor of Communication at the Uni¬
versity of Wisconsin — Milwaukee. As of October, 1971, he was appointed As¬
sociate Dean of the UWM Graduate School.
JOE MILLS lives at 688 Gary St., Ripon, Wisconsin. A canoeing and
hiking enthusiast, he is dedicated to preserving wild areas and quality en¬
vironments in Wisconsin.
L. G. (LARRY) MONTHEY is travel-recreation specialist with University
Extension, The University of Wisconsin. During the period 1970-71, he served
as executive officer of the Wisconsin Academy. ROBERT A. RICKETTS is a
business analyst with Arthur Anderson and Associates and is located in
Philadelphia.
GENE E. MUSOLF teaches at the University of Wisconsin — Marathon
Campus, Wausau. DR. FRANCIS D. HOLE is at the University of Wisconsin,
Madison.
MARGARET A. HARNEY and DR. CARROLL R. NORDEN are in the
Zoology Department, University of Wisconsin — Milwaukee.
NORMAN OLSON, an officer of Northwestern Mutual Insurance Com¬
pany of Milwaukee, was President of the Academy 1970-71.
CHARLES REDENIUS is in the Department of Political Science at Car-
roll College, Waukesha, Wisconsin.
DR. ROBERT J. SALZER is Associate Professor of Anthropology, Beloit
College, Beloit, Wisconsin.
DR. WILLIAM E. SLOEY is Assistant Professor of Biology at the Wis¬
consin State University, Oshkosh. DR. JOHN L. BLUM is Professor of Biology
at the University of Wisconsin — Milwaukee.
1972]
Biographies
333
DR. KENTON M. STEWART is in the Department of Biology, State Uni¬
versity of New York, Buffalo, New York.
DOUGLAS A. VALEK is a Specialist in Entomology at the University
of Wisconsin, Madison. DR. HARRY C. COPPEL is Professor of Entomology
at the University of Wisconsin, Madison.
DR. HOWARD F. YOUNG received all of his higher academic training
at the University of Wisconsin — Madison with studies concentrated in the
field aspects of Biology. He is currently Professor of Biology at Wisconsin
State University — La Crosse.
DONALD ZOCHERT is a reporter for The Chicago Daily News . He has
published in American Heritage, Natural History and other periodicals, and
has studies in press at the Journal of the Illinois State Historical Society and
Western American Literature.
■
'
>
WISCONSIN ACADEMY OF SCIENCES, ARTS & LETTERS
Madison, Wisconsin
OFFICERS 1971-72
President
F. Chandler Young
University of Wisconsin —
Madison
Vice-President (Sciences)
Elizabeth McCoy
University of Wisconsin —
Madison
Vice-President (Arts)
Edgar L. Obma
Dodgeville
Vice-President (Letters)
Flazel S. Alberson
Madison
President-Elect
Louis W. Busse
University of Wisconsin —
Madison
Secretary
Martha G. Hanson
Madison
Treasurer
George E. Sprecher
Madison
Librarian
Jack A. Clarke
University of Wisconsin —
Madison
APPOINTED OFFICIALS
Editor — Transactions
Walter F. Peterson
Dubuque University, Dubuque, Iowa
Editor — Wisconsin Academy Review
James R. Batt
W.A.S.A.L. Office, Madison
Chairman — Junior Academy of Science
LeRoy Lee
W.A.S.A.L. Office, Madison
ACADEMY COUNCIL
The Academy Council includes the above named officers and officials and
the following past presidents:
A. W. Schorger
Henry A. Schuette
Otto L. Kowalke
Katherine G. Nelson
Ralph N. Buckstaif
Joseph G. Baier
Stephen F. Darling
Robert J. Dicke
Henry A. Meyer
Carl Welty
J. Martin Klotsche
Aaron J. Ihde
Walter E. Scott
John W. Thomson
A. A. Suppan
W. B. Sarles
Norman C. Olson
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