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Transactions
Carl N. Haywood, Editor
134 Schofield Hall
University of Wisconsin-Eau Claire
Eau Claire, Wisconsin 54701
Production Editor
Janis Haywood
Poetry Editor
Bruce Taylor
Transactions welcomes articles that explore features of the State of Wisconsin and its people. Articles
written by Wisconsin authors on topics other than Wisconsin sciences, arts, and letters are also occa¬
sionally published.
Manuscripts, queries, and other correspondence should be addressed to the new editor after 1 April
1992:
William Urbrock
Associate Dean
College of Letters and Sciences
University of Wisconsin-Oshkosh
Oshkosh, Wisconsin 54901
Transactions is a publication of the Wisconsin Academy of Sciences, Arts, and Letters, 1922 University
Avenue, Madison, Wisconsin 53705-4099.
© 1992
Wisconsin Academy of Sciences, Arts, and Letters
Manufactured in the United States of America
All rights reserved '
TRANSACTIONS
of the Wisconsin Academy
of Sciences, Arts, and Letters
Volume 80
1992
Contents
The Ordeal of Being a Test Case: 1
In Quest of the Right to Practice Medicine in Wisconsin
Hania W. Ris
In 1949 the State Board of Medical Examiners denied a young, foreign-bom medical
doctor a license to practice medicine in Wisconsin, or even an opportunity to sit
for the examination for the license. The fact that she was licensed to practice in
two other states and had been on the faculty of two noted medical schools made
no difference. But Hania Ris was different, and her account of her successful
challenge of the Board’s decision and procedure will intrigue scholars and general
readers alike.
1988 Drought Impacts Among Wisconsin Dairy Farmers 21
John A . Cross
When a severe drought occurs in an area unused to this hazard, how are the farmers
affected? John Cross divided the state into nine districts and examined specific
drought-related problems. He also examined the question of why some farmers
went out of business during this period. The conclusions contain surprises.
Distribution, Abundance, Larval Habitats, and Phenology 35
of Spring Aedes Mosquitoes in Wisconsin (Diptera: Culicidae)
Jeffrey W. Gilardi and William L. Hilsenhoff
The spring mosquitoes that are a nuisance in wooded areas of Wisconsin belong
to the twenty-three species of Aedes . This paper summarizes previous studies in
Wisconsin and examines new larval and adult Aedes collected in nine representative
areas of the state. Two species were collected for the first time in Wisconsin.
Symbolism in the Cave of Montesinos 5 1
James T. Abraham
In a famous episode in Don Quixote the knight enters the legendary cave of Mon¬
tesinos, where he has a strange dream about the enchanted inhabitants of the cave.
This paper uses psychoanalytical theory to discuss the symbolism of the cave and
the dream and what they reveal about the psyche of Quixote and perhaps Cervantes
himself.
57
The Distribution of Franklin’s Ground Squirrel
in Wisconsin and Illinois
Timothy L. Lewis and Orrin J. Rongstad
Eastern populations of Franklin’s ground squirrel have declined in the past two
decades. The authors studied the range of this squirrel to determine whether a
reduction in range accompanied the population decline.
That Eyes May Be Free: 63
Mary North Allen Talks with Transactions Editor
Carl Haywood
What started as conversation about photography between Transactions editor Carl
Haywood and photographer Mary North Allen ended up covering much more.
Readers will glimpse the complexity, intellect, and vision of one of Wisconsin’s
premier photographers.
The Photography of Mary North Allen 72
Community Response to Floodplain Relocation 87
in Soldiers Grove, Wisconsin
Graham A. Tobin
The citizens’ response to frequent flooding in Soldiers Grove was to relocate the
downtown and several residential properties. This project is often cited as a suc¬
cessful example of multipurpose, community-sponsored planning. Although the
relocation was effective in preventing most flooding, Graham Tobin raises questions
about the social impact.
Depth, Substrate, and Turbidity Relationships 97
of Some Wisconsin Lake Plants
Stanley A . Nichols
In his continuing studies of Wisconsin’s lake plants, the author examines their
tolerance for varying growing conditions. The information gathered will be useful
in managing the state’s lake plant resources.
Poetry 119
Poetry editor Bruce Taylor has selected poems from some of Wisconsin’s well-
known poets and some of our newer, most promising poets for inclusion in this
volume.
IV
139
First Report of Natural Bridges in Eastern Wisconsin
Richard A . Pauli
Although natural bridges are well-known features in the Driftless Area of Wisconsin
and adjacent states, none of these delicate land forms is documented in the recently
glaciated region of Wisconsin. This paper describes natural bridges in two localities
along the Silurian escarpment in eastern Wisconsin.
Live Capture Methods of Sympatric Species 149
of Flying Squirrel
Thomas C. Engel, Michael J. Lemke, and Neil F. Payne
Standard methods of capturing other tree squirrels are not as effective for flying
squirrels. The objective of this study was to determine the trapping success for
sympatric species of flying squirrels relative to tree species, trap type, and the height
of the trap in the tree.
Range Extension of Northern Flying Squirrels 153
Thomas C. Engel, Michael J. Lemke, and Neil F. Payne
The authors examine the range habitats of northern and southern flying squirrels.
Their evidence extends the known range of the northern squirrels farther south into
Portage County.
The Modem Spiritual Condition and the Ancient Wisdom 155
of the I Ching
Claire E. Matthews
The I Ching or Book of Changes is an ancient Chinese manual of divination and
wisdom. It preceded Confucius, who wrote commentaries on it. Later Carl Jung
asserted his belief in its predictions. Claire Matthews presents a case for using the
1 Ching as a means of access to our own society’s beliefs.
Distribution, Abundance, and Diversity of Mollusks 163
(Bivalvia: Unionidae) from the
Lower Chippewa River, Wisconsin
Terry Balding
For four summers the author collected mussels from the Chippewa River between
Eau Claire and the Mississippi. Because freshwater mussels are a good ecological
indicator of water quality, these raw data will be especially valuable as we monitor
our environment during the next twenty-five to fifty years.
v
From the Editor
It was five years ago that I became editor of Transactions. It hardly seems that long, but
my commitment of five years as editor is fulfilled, and this volume is my last. This has been
a period of change in the activities of the Wisconsin Academy and in all its publications. But
the great tradition of the Academy in providing a mechanism for collecting and preserving
the creative, intellectual life of our state, and presenting that information to our citizens, has
continued. It is a history in which the Academy justifiably takes pride, and it has been an
honor to be associated, even for this short period, in the continuing process.
The changes in Transactions during the past five years were built on the groundwork
established by previous editors, particularly Kathryn and Philip Whitford, whose dedication
and work kept the journal alive. During my years as editor there have been a number of
changes. Poetry and photography have been added as regular sections. In addition to regular
issues, three special issues have been published: a history of the limnology program at UW-
Madison {Breaking New Waters), a poetry anthology ( Wisconsin Poetry), and a book on
treaty rights {Chippewa Treaty Rights). There has also been started an occasional series that
will result in at least two other publications. I am happy with these accomplishments, but
they are not those of a single editor. Patricia Duyfhuizen and Bruce Taylor deserve much
credit, and it is my pleasure to recognize their contributions and to thank them on behalf of
all who have enjoyed the results of their work.
Bruce built the poetry section from an idea into one of the exciting places to sample the
poetry of Wisconsin authors each year. It was Bruce who solicited and chose the poetry and
then wrote the introduction to the special issue entitled Wisconsin Poetry (Vol. 79, No. 2),
which has been extraordinarily well received. Bruce’s poetry readings at the annual convention
and around the state have done much to establish the Wisconsin Academy as one of the
leaders in encouraging Wisconsin poets.
Our former Production Editor, Patricia Duyfhuizen, enabled us to expand our services to
authors by increasing the level of professional scrutiny of the journal. Her advice, professional
judgment, and dedication added immensely to the journal. And her work with student interns
brought a level of professionalism that had not been available to previous editors.
When Patricia was unable to continue as Production Editor for this last volume under my
editorship, Transactions was fortunate to find Jan Haywood to produce the current issue.
Only those involved in publishing realize the seemingly unlimited number of mistakes possible
in the process. The professional scrutiny and advice Jan has provided have continued the high
quality of production our readers have come to expect. I am grateful to her for picking up
the work on short notice and for doing so with good humor.
Virtually every piece of paper, telephone call, or message associated with Transactions for
the past five years has been handled by Jan Kroll, a member of the Arts and Sciences staff
at UW-Eau Claire. Perhaps only she and I realize her role in the success of this journal. She
would, of course, dismiss it with “it’s my job,’’ but I know better and acknowledge her
contributions with sincere gratitude.
The current volume of Transactions contains some absorbing articles. The lead essay is an
examination of why a physician, licensed to practice medicine in two states and on the staff
of two noted medical schools, would be denied a medical license by reciprocity or even an
opportunity to sit for the examination to practice medicine in Wisconsin. Hania Ris’ story is
captivating. There are also the poetry section, an interview with Dresen Award-winning
vii
photographer Mary North Allen, along with a selection of her photographs, and articles on
subjects as diverse as the impact of the 1988 drought on Wisconsin dairy farmers, lake plants,
natural land bridges, spring mosquitoes, and ancient Chinese wisdom. And there is even more
for the reader to discover.
Bill Urbrock becomes the new editor with the publication of this volume. All further
correspondence, articles, and proposals should be sent to him at the address on the inside
front cover.
Carl N. Haywood
Editor, 1987-1992
The Ordeal of Being a Test Case:
In Quest of the Right
to Practice Medicine in Wisconsin
Hania W. Ris
When my husband Hans Ris accepted
an appointment as Associate Profes¬
sor in the Department of Zoology at the Uni¬
versity of Wisconsin-Madison in 1948, I was
intrigued. That was the year Life magazine
(6 September) ran its famous cover story
identifying Madison, Wisconsin, as Ameri¬
ca’s best place to live. Although I had an
interesting and prestigious position as a pe¬
diatrician in the Cornell Medical School De¬
partment of Pediatrics, I looked forward to
the move with anticipation. After diligently
studying the Life article, I became even more
enthusiastic. I learned that Madison, with a
population of 80,000, had three lovely lakes,
that the streets were lined with elms and ma¬
ples, that its many parks were maintained
with a very ample city appropriation. Its
“intelligent and alert populace’’ had a lit¬
eracy rate of 98%, and 17% had attended
college. The schools had an excellent repu¬
tation, pertinent information for a couple ex¬
pecting their first child in March 1949. Baby¬
sitters were easily available because of a large
student population. The city had many cul¬
tural groups. The university supported sev¬
eral “artists in residence’’ including a painter
as well as musicians. Drama was provided
Hania W. Ris, M.D., has been a member of the De¬
partment of Pediatrics, UW -Madison Medical School,
for thirty -five years. She is a peace activist and champion
of women’s rights, reproductive rights, the prevention
of teenage pregnancy, quality day care, and national
health insurance. She writes newspaper and magazine
features, as well as scientific papers. An abstract painter,
she has had several one -woman exhibits.
by the Lunts, who lived nearby and usually
opened their new plays in Madison. The ar¬
ticle even referred to the importance of the
Madison League of Women Voters and its
influence on civic decisions.
At the time of the University of Wisconsin
offer we were living in New York City. My
husband, a biologist, had been working in
the field of cytology (structure, function, and
pathology of the cell) at the Rockefeller In¬
stitute for Medical Research. I was working
with Dr. May Wilson at the Children’s Car¬
diac Clinic of New York Hospital as a Fellow
in Pediatrics as well as teaching on the staff
of Cornell University Medical School.
Hania Ris today
1
Wisconsin Academy of Sciences, Arts and Letters
Before my departure I had to certify my
medical documents. To my amazement and
amusement, I was warned by the physician
in charge that I was moving to a “socialist
state”!
In order to learn something about Wis¬
consin state politics, I read about “Fighting
Bob” La Follette and his Progressive Party,
which lost to the Democrats after the Second
World War. Although La Follette had been
dead for twenty years, his ideas of social
reforms, including care of the unemployed
and the elderly, had left permanent marks on
Wisconsin. It was the first state to pass a
workmen’s compensation law, in 1932, and
to prohibit child labor. Its law became a
prototype for other states. La Follette also
promoted the idea that the state government
should use the university as its first resource
and the university in turn should exert its
influence on the entire state.
All this information added to our convic¬
tion that Madison would be an interesting and
stimulating place to live and to raise a family.
My husband was especially urged to accept
the position by a friend and colleague, Charles
Leonard Huskins, Professor of Botany at the
university. A Canadian, he had been Profes¬
sor of Botany at McGill University in Mon¬
treal until he moved to Madison in 1945. He
and his wife Margaret and their three children
befriended us and offered their home when
we arrived in Madison in June 1949 with our
three-month-old son, Christopher. The
Huskins lived in a spacious older house on
Vilas Avenue near the Vilas Zoo. We stayed
with them a fortnight, and we could not have
had a warmer and more gracious welcome.
The Huskins remained our friends and wise
advisors until their deaths in 1953.
In July 1949 we moved to the University
Houses, built by the Wisconsin Alumni As¬
sociation the previous year for faculty and
families, and later given to the state. A gen¬
erous gesture, but the architecture left some¬
thing to be desired. Imagine a kitchen with
one drawer and minimal counter space! As
an ardent admirer of Frank Lloyd Wright, a
native son of Wisconsin, I could never un¬
derstand why the “Williamsburg” style of
architecture had been chosen. But taking into
account the prevailing housing shortage, we
were grateful.
After getting settled I was ready to con¬
tinue my pediatric work. In preparation for
the move to Wisconsin, I had had the appli¬
cation form for medical licensing in Wis¬
consin completed and certified by the Board
of Medical Examiners of Maryland in June
1949 for the purpose of licensure by reci¬
procity. I had been granted a license to prac¬
tice medicine by the Maryland Board of Ex¬
aminers in 1941, after passing on the first
attempt what was considered a very rigorous
examination, and after I had worked and
studied in this country for only one and a
half years. I had obtained my New York li¬
cense to practice medicine in 1942 through
reciprocity — a standing agreement between
two state boards of medical examiners. Wis¬
consin and Maryland also had a reciprocity
agreement, and I expected to obtain a Wis¬
consin license in the same way. I was there¬
fore totally unprepared for what followed.
I submitted my duly certified application
and my curriculum vitae to the Wisconsin
State Board of Medical Examiners (hereafter
The new Dr. Ris, circa 1937
2
In Quest of the Right to Practice Medicine in Wisconsin
referred to as the Board). My vitae outlined
my experience:
1 937— Graduation with Doctor of Medicine
degree from the Medical School of the Uni¬
versity of Zurich, Switzerland.
1937-39 — Assistant in Pediatrics at the
Children’s Hospital of the University of Zurich.
1939- 40 — Year’s internship in Baltimore.
1940- 41 and 1943-49 — A total of seven
years at the Johns Hopkins Pediatrics Depart¬
ment, including one year of work with Dr. Helen
Taussig, cardiologist, the originator of the world-
famous operation which corrected the defects
in the hearts of “blue babies.’’
1942-43 — Resident in Pediatrics, Chil¬
dren’s Hospital, and Instructor of Pediatrics,
University of Cincinnati Medical School.
1948-49 — Assistant Pediatrician, New York
Hospital, and Fellow in Pediatrics, Cornell
University Medical School.
All of these positions required teaching of
medical students. My experience included
clinical work in syphilis and diabetes, and I
had conducted several Well Baby Clinics for
the Baltimore City Health Department. I also
included letters of recommendation from
prominent physicians with whom I had
worked.
Round I
To my amazement, on 15 July 1949 all of
my credentials were returned to me with the
following arbitrary denial: “The Board of
Medical Examiners of the State of Wisconsin
is not licensing foreign graduates at the pres¬
ent time. It is hoped that within the not too
distant future we will be able to get reports
on the foreign schools which might enable
us to license graduates of some of them.”
To my further dismay, I learned that the
State Board, at a meeting held in 1937, had
adopted a policy of refusing any qualifying
examinations for graduates of foreign uni¬
versities, with the exception of graduates from
approved Canadian schools. This ruling co¬
incided with the immigration to the United
States of a number of physicians threatened
by racial and political persecution in Ger¬
many and other parts of Europe. Critics sug¬
gested that the Board’s 1937 ruling was self-
serving, aimed at eliminating competition and
creating a monopoly under the guise of pro¬
tecting the health of Wisconsin citizens. Other
states had passed similar measures.
Some in government questioned the Board’s
action. I learned that in 1948, Wisconsin
Representative Ruth Doyle had proposed a
bill requiring the Board to provide any ap¬
plicant denied the right to take the exami¬
nation with written notification of the reasons
for denial. It also provided that the Board be
subjected to judicial review of its decisions
in the same manner as other state boards and
commissions. The purpose was to provide
the applicant with an orderly procedure through
the courts. It was blocked by the successful
lobbying of both the Board and the Wiscon¬
sin State Medical Society (hereafter referred
to as the Medical Society).
There is indisputable evidence that the Board
was making exceptions to the 1937 ruling,
however. In reviewing the minutes of the
Board in 1989, 1 discovered, to my surprise,
that it had allowed Dr. Harry Leeb, an
American-born graduate of the University of
Bern Medical School, Switzerland, to obtain
his license. Although denied that right on his
first appearance before the Board on 1 1 Jan¬
uary 1938, Dr. Leeb was granted licensure
at the Board’s subsequent meeting on 27 June-
1 July 1938, during which Dr. Leeb’s case
was discussed “at length” by Mr. Resh of
the attorney general’s office. A resolution
was adopted unanimously that since Dr. Leeb
had received his medical education at the
University of Bern prior to the adoption of
the 1937 ruling, and because of the “mis¬
taken assumption based upon correspon¬
dence with the Board that the Board would
recognize such a school,” Dr. Leeb was per¬
mitted to take the examination. This reso¬
lution also stipulated that the permission would
not extend to other graduates of foreign med¬
ical schools.
Additionally, the Milwaukee Journal, 5
December 1948, reported that an American-
born, Swiss-trained physician had been li¬
censed within the previous three years by the
3
Wisconsin Academy of Sciences, Arts and Letters
Board, after passing an examination given
especially for him.
Ironically, there was a shortage of physi¬
cians at the time in Wisconsin, as frequently
reported in the Milwaukee Journal and other
newspapers. Indeed, prior to our move to
Wisconsin, I had been contacted in January
1949 by Dr. Amy Hunter, Director of Ma¬
ternal and Child Health, Wisconsin Depart¬
ment of Health, who offered me a position
contingent on my obtaining a license. She
had written me after hearing from Dr. Leona
Baumgartner, Director of the New York City
Health Department, that I would be available
for employment.
I responded to the notification of my re¬
jection by the State Board with a letter dated
8 August 1949, pointing out that I had grad¬
uated from the University of Zurich in 1937,
before the war, that the medical school was
considered one of the best on the European
continent, that I had had ten years of expe¬
rience in the United States teaching in three
leading medical schools, including Johns
Hopkins, and that I was certified in 1944 by
the American Board of Pediatrics, a national
professional organization that certifies com¬
petence in the field.
The Board did not keep me long in sus¬
pense. In its 12 August 1949 reply, it stated:
“It is a definite policy of the Board ... at
this time to grant no licensure to graduates
of foreign schools other than Canadian.” Fol¬
lowing the second refusal, I asked for the
privilege of appearing before the Board at its
10 January 1950 meeting. It met regularly
only twice a year, in January and July. Per¬
mission was granted.
My presentation was factual and legalistic.
I emphasized that the University of Zurich,
from which I had graduated, was comparable
to the American schools with which I was
familiar: Johns Hopkins, Cincinnati, and
Cornell. All my records, European and
American, were available for review. I pointed
out that the Board had, in 1925, licensed Dr.
Karl F. Schlaepfer, a graduate of the Uni¬
versity of Zurich Medical School and an
American citizen. (I had become an Amer¬
ican citizen in 1944.) Dr. Schlaepfer was
practicing at that time in Milwaukee, and his
licensure showed that the Board had already
accepted Zurich as a reputable school.
In addition, I stated that I had been advised
that my credentials could be submitted by the
proper authorities for evaluation by Dr. Helen
Dwight Reid, Chief, European Section, Di¬
vision of International Educational Rela¬
tions, Federal Security Agency, Office of Ed¬
ucation, Washington, D.C. Dr. Reid had also
advised me as follows: “The fact that you
have been accepted for postgraduate training
in American institutions and licensed in two
other states should be helpful in obtaining
recognition, if the State can make any ex¬
ception to its general regulation.” I informed
the Board that I had been considered for a
position with the State of Wisconsin Board
of Health which called for a person with my
pediatric training. I added to my previously
submitted letters of recommendation one from
Dr. Helen Taussig, the world-renowned child
cardiologist with whom I had worked at Johns
Hopkins from 1940-41, testifying to my
character and professional competence and
the reputability of the University of Zurich
Medical School.
To my continuing dismay and growing sense
of unreality, I was once again denied the right
to be admitted for examination for licensure
to practice medicine in Wisconsin. The three
and a half nonchalant lines in the Board’s
minutes hardly reflect the impact that this
decision had on my life: “Dr. Hania Ris,
graduate of the University of Zurich, Switz¬
erland, in 1937, appeared. She answered
questions asked by the Board members rel¬
ative to her professional education and his¬
tory, following which she was informed that
her application was temporarily, at least, re¬
fused. Dr. Ris left the meeting.”
Inner politics of Board and
Medical Society
In the course of preparing for my 10 Jan¬
uary 1950 meeting with the Board, I had been
advised by a respected senior pediatrician,
Dr. Horace Tenney of Madison, to consult
4
In Quest of the Right to Practice Medicine in Wisconsin
with two officers of the Medical Society. I
found Mr. C. H. Crownhart, attorney and
secretary of the Society, and Mr. Tom Doran,
Society employee, both courteous and will¬
ing to advise. It was my impression that they
were sympathetic to my plight and interested
in my obtaining the license. As members of
a professional organization dealing with the
public, they appeared to be concerned about
the image the Medical Society was projecting.
By contrast, the State Board was a legal
body; its eight members (seven physicians
and one osteopath) were appointed by the
governor for a period of four years. I learned
that Dr. C. A. Dawson, the powerful sec¬
retary of the Board and a homeopath, had
run for the office of lieutenant governor. When
Dawson was defeated, Governor Goodland
appointed him to serve as secretary of the
Board of Medical Examiners. The remaining
seven members of the Board were appointed
at Dr. Dawson’s suggestion.
All the members of the Board, with the
exception of the osteopath, were members
of the Medical Society and active in its
affairs. Since the Board and the Medical
Society always presented a united front be¬
fore the legislature in matters such as the
Medical Practice Act, in the eyes of the
public they were not differentiated. It is
germane to note that the Medical Society
had always been a conservative body, cau¬
tious in endorsing new ideas.
A letter from the past sheds light
In my recent search— some thirty-four years
after my original request for licensure— for
documentation indicating covert deliberation
regarding my case, I found nothing other than
my original application in my file or under
various headings such as “Foreign Gradu¬
ates,” at the Historical Society Archives or
in the office of the State Board. My written
testimony presented to the Board, various
documents submitted, the Board’s corre¬
spondence with the Council of Medical Ed¬
ucation of the American Medical Association
(AMA) and with the University of Zurich
Medical School, and the correspondence be¬
tween my attorney and the Board cannot be
found even at the office of the Board. But
my file at the Medical Society (not available
to the public) contained, among other infor¬
mative documents, a letter by C. H.
Crownhart, secretary of the Medical Society.
This letter was sent to me in 1984 by courtesy
of Mr. Earl Thayer, at the time secretary of
the Medical Society. A consultation in 1989
with the attorney general’s office revealed
that it should have been part of the public
documents of the State Board. It was either
suppressed or overlooked, to my detriment.
Crownhart’ s letter of 23 November 1949
was addressed to Dr. H. H. Christofferson
of Colby, Wisconsin, a member of the Board,
a member of the Council of the Medical So¬
ciety, and its president-elect. It was written
six weeks before my 10 January 1950 hearing
before the Board. Three and one-half pages,
single-spaced, the letter was a legal attack
on the Board’s position against licensing for¬
eign graduates in general, and its refusal to
license me in particular.
Mr. Crownhart pointed out that he did not
recall any instance in which there was a di¬
vergence of opinion between the Medical So¬
ciety and the Board. When, in 1948,
“Assemblywoman Ruth Doyle, along with
other legislators interested in the problem of
the foreign graduate, brought in a proposal
to amend the law to make it easier for these
people to qualify, the State Board and the
State Society saw eye to eye on the effect of
that bill. As a matter of fact the Secretary of
the State Board and the Secretary of the State
Medical Society appeared at the hearing and
explained the problem.” Both secretaries must
have been very persuasive in defending the
status quo; the bill did not pass.
However, Mr. Crownhart also pointed out
in his letter that before the Board adopted the
1937 ruling against admitting graduates of
foreign medical schools to examinations, the
graduates of such schools could apply and
their credentials would be verified. As a re¬
sult, there were many foreign-educated phy¬
sicians practicing in Wisconsin. Then Mr.
Crownhart cited my case, emphasizing that
5
Wisconsin Academy of Sciences, Arts and Letters
I had graduated from the University of Zurich
in 1937 before the war, that I had had ten
years of postgraduate training in this country
in addition to teaching in three medical
schools, and that I was accredited as a spe¬
cialist by the American Board of Pediatrics
and was a member of national medical so¬
cieties. Furthermore, he noted that I was being
considered for appointment in one of the state
agencies on the basis of my credentials and
recommendations .
“It is my feeling,” Mr. Crownhart con¬
tinued in his letter,
that the odds are about even that this particular
situation may ultimately result in wide public
knowledge of her problem. ... It seems to me
that the public would feel that this woman was
entitled to the examination. As a matter of fact
they would feel that even if her school of grad¬
uation should have been inferior to American
schools in the type of training offered, that her
subsequent training as an intern and as a resi¬
dent, and her acceptance on the teaching faculty
of several schools would have overcome what¬
ever deficiencies she might have had from her
academic training.
Mr. Crownhart continued:
I have read the Medical Practice Act many times.
As I have told the Board I, as an attorney, fail
to find in it authority under which the Board
may adopt any blanket rule. The burden of
proof is, undoubtedly, upon the applicant. I
would not question that for one minute, but
unless the Board considers the application and
the applicant’s qualifications and gives that in¬
dividual an opportunity to fulfill the burden that
is upon her or him, it seems to me that the
Board has failed to follow the spirit or the letter
of the law.
Unaware of this letter as I was for forty years,
I could not have known at the time what an
advocate I had — a competent attorney in an
official position at the Medical Society who
was cognizant of the political scene.
My case gains notoriety , or
“Can’t Examiners Examine?”
Indeed my 10 January 1950 appearance
before the Board generated a great deal of
publicity, accurately reported, and all of it
critical of the Board. Europeans traditionally
eschew publicity; I was crushed! I was to
grow more accustomed to and more grateful
for the press as my case was championed over
the following year on the editorial pages of
the Capital Times, the Wisconsin State Jour¬
nal, the Milwaukee Journal, and the Mil¬
waukee Sentinel. Headlines were often bluntly
critical of the establishment: “The State Board
of Medical Examiners Continues to Operate
a Closed Shop,” “Medical Monopoly Still
Upheld,” “The State Board of Medical Ex¬
aminers Continues Its Stubborn Policy,” and
“Can’t Examiners Examine?” It is hard to
remember any other such instance when these
four newspapers, with their otherwise diver¬
gent opinions, acted in such unison.
During this period of publicity the Board
cited cases in defense of its policies. Another
physician, Dr. Ralph Smith, a graduate of
Edinburg Medical School and formerly a pro¬
fessor in Canada, was denied a license to
practice in Wisconsin. He was later found to
be a drug addict.
Another case was that of a Dr. Dubin. In
1930 he had presented himself as a graduate
cum laude of Maximilian University in
Wurzberg, Germany. He was permitted to
write the examination eight times, with fail¬
ure each time. It was later discovered that he
had never actually graduated from Maximi¬
lian University and that by a special dispen¬
sation he had been permitted to take an ex¬
amination and present a doctoral thesis.
I was confronted with this case of forgery
when I was referred by some prominent Wis¬
consin physicians to an administrator asso¬
ciated with the Wisconsin State Laboratory
of Hygiene in the hope that he might inter¬
vene in my behalf. It was devastating for me
to have this prominent administrator insin¬
uate that my own veracity might be
questionable.
In a similarly distressing encounter, a highly
placed medical educator told me that aca¬
demic medicine would be closed to me for¬
ever because of the publicity, that one does
6
In Quest of the Right to Practice Medicine in Wisconsin
From the Capital Times, 17 February 1950 . Courtesy of the State Fiistorica! Society of Wisconsin.
not go public with such complaints in the
United States and especially in Wisconsin. I
informed him that I myself had been per¬
turbed by the publicity and explained that I
had no control over it. He indicated doubt
about this, and ironically an editorial critical
of the Board appeared in the Capital Times
the next evening.
I was not surprised that support for my
plight was not coming from physicians in
private practice, but this prejudiced treatment
from physicians in academic medicine was
unsettling. I was told that Dr. Amy Hunter,
who had offered me the state position on the
basis of my credentials, tried to intervene and
for her efforts was rebuffed by her superior
in the State Health Department. I grew
desperate.
A confidential source within the Medical
Society staff who began advising me about
this time informed me that several physi¬
cians, members of the Medical Society, had
gone to the Governor and asked him to in¬
tervene. But Dr. Dawson, secretary of the
Board, had anticipated their move and pre¬
sented my case to the Governor in a fashion
that made him believe the Board would be
breaking the law by granting me a license,
an interpretation that distorted the law. I was
also told that several physicians hoped I would
7
Wisconsin Academy of Sciences, Arts and Letters
take my case to court. But none of them had
the courage to protest the restrictive policy
openly.
The biggest blow to my morale was a press
release issued by the Medical Society un¬
conditionally endorsing the Board’s policy
against licensing foreign graduates. In re¬
sponse to criticism by the media, the Medical
Society, on 18 February 1950, issued a news
release commending the State Board for act¬
ing in “good faith in the matter of reviewing
qualifications of those educated in foreign
countries.” It complimented the State Board
for being a moving factor in initiating a study
of foreign medical schools by the AM A. At
that time, thirty-eight schools had been ap¬
proved, but Swiss schools had not as yet been
evaluated. To counteract any suspicion of
prejudice, the Medical Society added that the
“Board consists of highly respected individ¬
uals, many of whose immediate forebears
come from foreign countries.”
If the State Board had done any “fence¬
building,” of which it was accused, the news
release continued, it had done so only “to
protect the health of the people of this state
. . . through the legislative and judicial pro¬
cesses.” Yet this was the very process that
had been described privately by Mr.
Crownhart, the Medical Society’s attorney,
as a failure “to follow either the spirit or the
letter of the law.”
But the communique opened one door,
namely, that the Board would “ continue
[emphasis added] in the future to receive ap¬
plications from graduates of schools not as
yet formally qualified.” (Yet I had been de¬
nied this right when I had appeared before
the Board in January 1950, only five weeks
earlier.) The applicant would have the burden
of proof to demonstrate that “he was trained
under the same general conditions as are re¬
quired of those attending the medical school
of the University of Wisconsin.”
This statement gave me some hope. Ironi¬
cally, during that time many European and
American medical academics were critical of
the quality of institutions such as the Uni¬
versity of Wisconsin Medical School. For
instance, at Wisconsin the Department of Pe¬
diatrics was part of Internal Medicine and
did not become independent until 1957. Sim¬
ilarly, the Department of Psychiatry did not
become independent from the Department of
Neurology until 1956. In contrast, the De¬
partment of Pediatrics of my medical school
at the University of Zurich was headed by
the world-renowned Professor Guido Fanconi.
I had also been privileged to study under
Professor W. R. Hess, Director of the Phys¬
iological Institute of the University of Zurich,
who received the 1949 Nobel Prize in Med¬
icine and whose work greatly enhanced phys¬
iologic and psychiatric thinking throughout
the world. American physicians used to come
to Zurich for postgraduate training and to
work with such other famous department
chairpersons as Professor Guido Miecher
(dermatologist and venerealogist), Professor
Hans Rudolf Schinz (roentgenologist), and
Professor Otto Naegeli (hematologist). The
departments of pediatrics in which I had trained
in this country for ten years prior to seeking
the Wisconsin medical license were also in¬
dependent; the Department of Pediatrics at
Johns Hopkins had been independent
since 1914.
One other hopeful note in the Medical So¬
ciety release was its endorsement of a pro¬
posal that the Board consider “such addi¬
tional training as an applicant may have
acquired since coming to this country.” (The
Board never acted on this endorsement in
my case.)
In response to this press release, the Mil¬
waukee Journal printed an editorial entitled
“Whitewash for Doctors’ Fence” on 20 Feb¬
ruary 1950. It attacked the policy of the Board
in protecting its selfish professional interests:
“Doctors of unquestioned ability and repute
have been arbitrarily barred from examina¬
tion in Wisconsin by a policy of the Board
which was never imposed by the legislature,
courts or public. Communities and institu¬
tions in need of those doctors have been de¬
nied them — by the Board and by nobody else.’ ’
Yet Mr. Crownhart, secretary of the Med¬
ical Society, defended the Board’s actions in
8
In Quest of the Right to Practice Medicine in Wisconsin
a letter responding to this editorial which was
published in the Milwaukee Journal and re¬
produced in the Wisconsin Medical Journal,
March 1950. It seems impossible to reconcile
this public statement with his letter sent to
Dr. Christofferson in November 1949.
A steady source of encouragement
While battling what seemed a no-win sit¬
uation, I contacted my former teacher and
mentor, Dr. Edwards A. Park, recently re¬
tired and former chairman of the Department
of Pediatrics of the Johns Hopkins Medical
School. Though others helped, I am con¬
vinced that his tireless one and one-half year
interventions with the AMA were crucial to
my obtaining my license to practice medicine
in Wisconsin.
Dr. Park, a nationally and internationally
renowned pediatrician, became my steady
source of encouragement. At age seventy-
one he took it upon himself to fight my battle
with youthful vigor. He wanted to know every
detail of my dealings with the Board. I wrote
lengthy letters to him to which he always
responded promptly, frequently after con¬
sultation with individuals who he thought
might help. This correspondence became a
useful reference for documenting my case
(and was recently accepted by the Johns
Hopkins Medical Archives to broaden the
profile of Dr. Park).
In his comforting letter to me after the
Board’s second refusal to recognize my ap¬
plication, Dr. Park wrote on 25 January 1950:
“May I say that I was incensed at your treat¬
ment by the examining Board in Wisconsin.
... I am sure you will receive your license,
the examining Board will not dare refuse it
after their exposure by the press. They will
probably wait long enough to save their face.’’
Dr. Park wrote me on 8 March 1950: “The
whole affair makes me ashamed of my coun¬
try and particularly ashamed of the medical
profession.” In a letter dated 22 March 1950
he informed me: “ Time [magazine] has writ¬
ten that they will accept a letter from me on
your case in Wisconsin.” At the urging of
Dr. Donald G. Anderson, secretary of the
Council to the House of Delegates of the
AMA, Dr. Park postponed sending this letter
in order to allow Dr. Anderson to intervene
in my behalf. On 11 April 1950 Dr. Park
wrote to me again noting his request to Dr.
Anderson and further explained, “I hesitate
to take too open a part for the reason that I
am anathema, having headed the protest
against organized medicine. By some I am
regarded as having communistic leanings.”
Round II
In response to the Board’s statement that
it would accept new evidence from the ap¬
plicants as to the reputability of their medical
schools, I resubmitted my credentials on 5
April 1950 to Dr. Dawson, secretary of the
Board. It included the enumeration of every
lecture, every course and laboratory exercise,
certified by the Zurich Medical School. I also
sent a money order for fifty dollars to cover
the reciprocity fee with Maryland, for which,
as I was told by Dr. Dawson, I was to be
eligible, once the reputability of the Zurich
Medical School was established. I also asked
him to consider my application at the forth¬
coming meeting on 19 April 1950.
I accompanied my application with a letter
from Dr. Marion Sulzberger, an American-
born U.S. citizen, a world-renowned der¬
matologist and allergist who was at the time
professor and chairman of the Department of
Dermatology and Syphilology at the Post-
Graduate Medical School, New York Uni¬
versity. He had graduated from the Univer¬
sity of Zurich Medical School in 1926, just
eleven years prior to my graduation. He had
written several textbooks and more than a
hundred articles and had contributed greatly
to his fields of expertise. I could not have
had a better testimony to the reputability of
the University of Zurich Medical School. In¬
deed, some of my teachers were the same as
those of Dr. Sulzberger. At the time I was a
student, Dr. Sulzberger had returned to Zu¬
rich for postgraduate training. How could the
Board ignore these facts?
Providing another written testimony was
Professor Karl Meyer, a Swiss native and
9
Wisconsin Academy of Sciences, Arts and Letters
Professor of Experimental Pathology at the
University of California Medical Center- San
Francisco.
My application, dated 5 April, was ac¬
knowledged by Dr. Dawson on 10 April 1950.
His letter stated that although the one-day
meeting of the Board on 19 April would have
a heavy agenda, “I shall present your ap¬
plication in its entirety to the Board at that
time.’ ’ When I did not hear from Dr. Dawson
within the following two weeks, I wrote him
on 6 May to inquire about the action the
Board had taken in my case. Dr. Dawson
replied on 10 May saying that the matter of
foreign graduates had not been considered.
He added, “The fact of the matter is that no
change in the policy regarding foreign grad¬
uates is possible at this time inasmuch as no
addition has been made to the list of the ap¬
proved schools issued by the AMA.” This
statement reversed the Board’s alleged public
change of policy that it would honor the right
of the applicant to prove the reputability of
the school of graduation. It became clear to
me that the Board’s intention was not only
to stall but to deny me the license permanently.
Dr. Dawson also mentioned in his letter
that the next meeting of the Board would be
held in Milwaukee on 11-13 July 1950; if I
wished to appear I should let the Board know,
and they would notify me as to place and
time of my appearance. On 17 May I wrote
Dr. Dawson to confirm my interest, adding:
“Undoubtedly, you have by now reviewed
the standing of the Medical School of the
University of Zurich in the prewar period [on
the basis of documents I submitted] .... If
there is any further evidence that you would
like to have presented to prove that ... the
University of Zurich Medical School . . .
provided training equivalent to the Medical
School of the University of Wisconsin, I will
make every effort to obtain such evidence.’’
I had been warned by my confidential source
at the Medical Society that since Dr. Dawson
withheld information and communication from
other members of the Board, I should dis¬
tribute a copy of each communication to every
Board member. This was still a world with¬
out photocopy machines. If I did not advance
my medical career during this interim, I cer¬
tainly did advance my secretarial and para¬
legal skills.
The warning was not idle. I did not hear
from Dr. Dawson until I wrote him again on
3 July, this time sending a copy to each mem¬
ber of the Board. The time constraint was
nerve-wracking and intimidating. Had I missed
the semiannual meeting, I would have had
to wait another six months. Dr. Dawson re¬
sponded with a letter dated 5 July 1950 giving
me an appointment for 12 July at 2 p.m. at
the Pfister Hotel in Milwaukee. There was
no response to my inquiry as to whether the
Board wished to have additional documents
to prove the reputability of my medical school.
Approval of Swiss medical schools
By 1949, the year my fight for licensure
began, the problem of licensing foreign grad¬
uates had assumed national and political di¬
mensions. A report by the Council on Med¬
ical Education and Hospitals to the delegates
of the AMA on 6 June 1949 recognized these
dimensions: “In the past fifteen years more
than 10,000 foreign trained physicians have
migrated to the United States and it may be
expected in the years ahead that at least 1 ,000
foreign medical graduates will be coming to
this country annually.’’ The Council recog¬
nized that the state licensure boards had no
way by which to evaluate foreign medical
schools, and consequently some excluded all
foreign medical graduates, while others ad¬
mitted all foreign graduates to their exami¬
nation for licensure. (In 1949 only twelve
state medical examining boards admitted for¬
eign graduates to examinations for licen¬
sure.) The Council proposed that the House
of Delegates empower it to evaluate foreign
medical schools, which it did.
In preparing to appear before the Board I
learned that on 24 June 1950 the Council on
Medical Education and Hospitals of the AMA
approved five Swiss medical schools, in¬
cluding that of the University of Zurich. Again
my confidential source told me that Dr.
Dawson would try to suppress this infor-
10
In Quest of the Right to Practice Medicine in Wisconsin
mation in order to prevent me from obtaining
the license and to justify his original refusal.
I contacted Dr. Donald G. Anderson, Sec¬
retary of the AM A Council, and asked him
to notify Dr. Dawson about the Council’s
decision. Dr. Anderson kindly obliged on 8
July by letter. On 7 July I received a wire
from the Council of the AM A informing me
of the Board’s notification. Dr. Park’s cor¬
respondence with Dr. Anderson on my behalf
had paved the way to this unprecedented
cooperation.
But there was a problem lurking in the
Council decision. In its evaluation of the Swiss
medical schools, the Council of the AM A
had reviewed their status after 1940, at which
time a new degree was introduced for non-
Swiss citizens: Akademische Zeugnis or the
Certificat d’ Etudes Medic ales (Certificate of
Medical Studies). The only degree available
to non-Swiss citizens like myself at the time
of my graduation in 1937 had been the M.D.
degree, which the Council of the AM A did
not approve. The Council also approved a
second degree, the Swiss Federal Diploma,
for which only Swiss citizens were eligible.
At the time of my study in Switzerland I was
a Polish citizen, although I later became Swiss
through marriage.
The AMA recommendation specified that
the requirement for both approved degrees
was at least eight semesters of study. I was
in my tenth semester when I passed the ex¬
amination for my M.D. degree and thereafter
completed three additional semesters of post¬
graduate study. I took the same courses and
lectures required of the Swiss students eli¬
gible for the Federal Diploma.
Immediately following my degree in 1937
I was granted a position as Assistant in Pe¬
diatrics at the Children’s Hospital of the Uni¬
versity of Zurich, working under the re¬
nowned pediatrician, Professor Guido
Fanconi. I performed the same duties as my
Swiss colleagues holding the Federal Di¬
ploma. I held this position for two years be¬
fore coming to the United States in March
1939, and I had a statement from Professor
Fanconi attesting to these facts. In spite of
this evidence of my training, I surmised that
the Board would use the AMA evaluation as
a weapon against me. I did not err.
On the advice of my confidential source,
I engaged a lawyer. My fortunate choice was
James Doyle, who later became a federal
judge and was married to Assemblywoman
Ruth Doyle.
The date of my appearance before the Board
on 12 July to defend my case was fast ap¬
proaching. On 8 July an important letter was
written by Dr. Kenneth McDonough, As¬
sociate Professor of Pediatrics, University of
Wisconsin, to Dr. J. W. Smith, president of
the Board:
Dr. Ris has made rounds and attended staff
meetings at the Wisconsin General Hospital in
Madison during the past year. I have been im¬
pressed with her intelligence, her knowledge
of medicine in general, and her understanding
of pediatrics, the field in which she is partic¬
ularly interested and for which she has excellent
training. We also have the opportunity to know
her professionally and believe that she will make
a fine practitioner. She will render a valuable
service to the community and state.
On the crucial day, I drove with my at¬
torney to Milwaukee. I did not plan to have
him appear with me at the meeting unless
there was nothing further to lose. Dr. Park
feared that having me represented by a law¬
yer might antagonize the Board. Members of
the press representing major Wisconsin
newspapers were also present.
I spoke and supplemented my oral presen¬
tation with two concise written statements.
One addressed itself to the reputability of the
University of Zurich Medical School and the
other to the approval of the Swiss medical
schools in general. I specifically explained
why I could not possess the Certificate of
Medical Studies. It seemed so simple. This
particular degree was introduced in 1940, and
I had graduated in 1937.
The Board at first denied knowledge of the
AMA’s approval of the University of Zurich
Medical School. When I showed them the
wire from the AMA stating that the Board
11
Wisconsin Academy of Sciences, Arts and Letters
had been notified, its response was that the
AMA’s letter was not official, because the
approval had not yet appeared in print in the
Journal of the American Medical Association.
Though some members of the Board asked
questions during my forty-five-minute ap¬
pearance, the attorney for the Board, John
W. Davison, questioned me most often. After
the futile battle, I asked to be represented by
my counsel. Davison agreed. Though the
presence of James Doyle appeared not to an¬
tagonize the Board, it simply continued to
stall. My attorney suggested that the Board
should send my credentials for evaluation to
the Council on Medical Education and Hos¬
pitals of the AMA and that the Board afford
me opportunity to present my case personally
to the Council. The Board appeared to accept
this suggestion. When my attorney asked for
a written confirmation of this agreement at
the conclusion of the meeting, the Board’s
attorney stated that this request might anger
the Board. There is no evidence that the Board
ever sent my credentials to the AMA. This
is no surprise since my attorney told me after
the meeting that he had never seen such a
disorganized state body as the Board of Med¬
ical Examiners. Most of the time, he ob¬
served, they did not seem to know what they
were discussing.
The following are the “postmortem” min¬
utes of the 11-13 July 1950 Board meeting
as they relate to my case:
The Dr. Hania Ris case was reviewed briefly
by Mr. Davison, having been announced by
the President as the first order of business. Dr.
Ris had previously requested permission to at¬
tend the meeting, and she was admitted to it at
this time. Mr. Spaulding of the Milwaukee
Journal, and a reporter from the Milwaukee
Sentinel, were also present. Dr. Ris made a
short statement to the Board and answered sev¬
eral questions put to her by the members. Dr.
Ris then requested that her attorney, Mr. James
Doyle, Madison, be admitted to the meeting,
and her request was granted. Mr. Doyle at¬
tempted to clarify Dr. Ris’ position on the mat¬
ter of recognition of Swiss schools, particularly
the University of Zurich, and the matter of her
diploma.
The Board did not take any formal action
on the matter of Dr. Ris’ application, and indi¬
cated to her that nothing further could be done
until a decision had been reached by the Coun¬
cil on Medical Education of the American Med¬
ical Association. Dr. Ris and Mr. Doyle left
the meeting.
Following the meeting, Dr. J. W. Smith,
Board president, spoke to me privately. He
said he had tried to convince the members
of the Board to grant me a license but they
would not listen. He apparently wanted me
to reassure him that he had been fair to me.
The Council on Medical Education and
Hospitals of the AMA and the Executive
Council of the Association of American
Medical Colleges officially reported their ap¬
proval of the University of Zurich Medical
School on 14 July 1950, three days after my
appearance at the Board meeting. My attor¬
ney, James Doyle, sent a written reminder
on 25 July to Davison, the Board’s attorney
(as well as precautionary copies to all mem¬
bers of the Board), to send my credentials
for evaluation to the AMA Council as per
agreement. In the letter Mr. Doyle pointed
out that the language of the AMA report of
14 July was almost identical to the language
of the letter from Dr. Anderson of the AMA
Council that had been in the possession of
the Board at the 11 July meeting.
It was not certain that the Board would
honor its agreement even after an article ap¬
peared in the Milwaukee Journal on 30 July
1950 stating that the Board intended to send
my records to the Director of the Swiss Health
Bureau for evaluation. The article added that
the Board might consider it necessary to send
my records to the AMA Council for inter¬
pretation after the Swiss director’s reply had
been received. In order to avoid further un¬
necessary delays, my attorney requested, in
a 31 July 1950 letter, that the Board send my
credentials simultaneously to Switzerland and
the AMA Council. Mr. Doyle also asked to
receive a copy of the letter the Board was
to send to the Swiss Health Bureau and re¬
quested to be informed which of my records
had been sent. There was no response to Mr.
12
In Quest of the Right to Practice Medicine in Wisconsin
Doyle’s letters of 25 July and 31 July until
he sent a written reminder on 7 August.
The Board’s attorney responded on 9 Au¬
gust: “The case of Dr. Ris has been brought
to the attention of the Director of the Swiss
Health Bureau. To date none of the records
has been sent to Switzerland. It may be nec¬
essary to do so in the future. At the present
time I am not able to furnish you with a copy
of the Swiss correspondence.” We were at a
loss to understand what objection there could
be to our request to see the Board’s corre¬
spondence with the Director of the Swiss
Health Bureau. The disconcerting explana¬
tion was that the Board had not included my
records, even though the basic controversy
centered around the duration and character
of my study.
The delays continued. On 17 November
the Board requested a copy of the regulation
sent me by the Dean of the University of
Zurich Medical School, defining the eligi¬
bility for obtaining the Certificate of Medical
Studies introduced in December 1940. “If I
find it necessary to obtain this information
from Switzerland,” warned the Board’s at¬
torney, “it may take several weeks.” I had,
however, already submitted this six-page,
single-spaced document in the original and
with a translation at my 12 July 1950 ap¬
pearance before the Board!
I learned not to underestimate the Board’s
creative stalling tactics. This same letter from
the Board requested certified copies of my
marriage license. Records from 20 Novem¬
ber 1950 show that the Board’s attorney re¬
quested the marriage certificate “in order to
definitely establish the identity of the appli¬
cant.” This request was made sixteen months
after my original application for the license
and after my two appearances before the
Board! I found it insulting. My attorney was
appalled.
The 1950 document spelling out why the
Board needed my marriage license epito¬
mized the Board’s tortuous rationale against
licensing me. It referred to a new course in¬
troduced by the Swiss Medical Schools in
1940 leading to a Certificate of Medical Stud¬
ies, equivalent to the course leading to the
Federal Diploma. Noted the Board’s attor¬
ney: “Dr. Ris would have been eligible for
this course had she been attending school at
the time. The fact that she was not able to
enroll in this course for the reason that it was
not offered at the time can in no way be held
against her, but neither can the Board be
criticized because of her inability to do so.”
However ridiculous the reasons, it became
clear that the Board hoped above all that its
rationalizations would preserve its credibility
with the public over this issue.
The reply of the Swiss Director of Health
to questions submitted by the Board was nec¬
essarily general in nature, because the Board
had not included my individual records de¬
spite our urging. Later, after I had sent the
records myself and the Swiss director had
reviewed them, he concluded that I had taken
more courses, lectures, and clinics than re¬
quired for admission to the examination for
the degree of Doctor of Medicine; that, in
fact, I had received the same training as Swiss
citizens then received; and that I would have
been admitted to examination for the Certif¬
icate of Medical Studies, if such examination
had existed at the time.
Round III
News of the Board’s refusal on 1 1 July to
grant me the right to take an examination for
licensure reached Dr. Park at his vacation
cottage in Canada. In a letter dated 13 August
this dignified and gentle human being ex¬
pressed his profound outrage: “lam incensed
over the action of the WI. Licensing B’d.
... I have again written to Dr. Anderson
[secretary of the AMA Council] . If this does
no good I shall consider getting Dr. Weech
and Levine to unite with me in some public¬
ity. [Dr. Weech was chairperson of Pediat¬
rics, University of Cincinnati Medical School,
where I worked in 1942-43. Dr. Levine was
chairperson of the Department of Pediatrics,
Cornell University Medical School, where I
worked in 1948-49.] I shall not give up. . . .
I am filled with shame that you should be
treated so.”
13
Wisconsin Academy of Sciences , Arts and Letters
My other champions, the Wisconsin press,
meanwhile continued to advocate my licen¬
sure. In fact, the source of our information
about the Board’s contemplated action was
often newspaper articles. An article of 6 Oc¬
tober 1950 indicated that the Board said I
would be notified of the meeting and per¬
mitted to present my case to the Board on 10
January 1951. I was glad to read this, since
I had not been personally notified.
A Capital Times editorial on 9 October
again defended my case: “Dr. Ris is a dis¬
tinguished member of the profession . . . but
here in Wisconsin the political fuddy-duddies
who dominate the Board of Medical Exam¬
iners and whose competence is far inferior
to that of Dr. Ris are allowed to sit in judg¬
ment of her case.” Referring to the critical
shortage of physicians, the editorial urged the
passage of a bill introduced by U.S. Rep¬
resentative Andrew Biemiller of Wisconsin
to provide federal aid to medical schools, a
bill which it accused the AM A lobby of ‘ ‘knif¬
ing in Congress.” This federal aid was deemed
imperative by deans of the major medical
schools “to insure even the barest minimum
of doctors for future civilian and military
needs,” noted the Capital Times. President
Harry Truman had termed the bill “the most
vital health legislation before Congress”
(“Washington Merry-Go-Round,” syndi¬
cated column by Jack Anderson and Fred
Blumenthal, Capital Times, 18 August 1950).
At the end of November I received a long
letter from Dr. Park outlining a strategy to
enlist the aid of Dr. Anderson of the AM A
Council. Dr. Park intended to visit Dr.
Anderson in Chicago and to “be guided of
course by his advice, provided his advice
appears to me in your interest and wise.” He
planned to seek Dr. Anderson’s consent to
send my credentials to the AMA Council for
adjudication, and mailed him records of my
educational qualifications and reports of the
Board’s action. In late December Dr. Park
counseled me to write Dr. Anderson directly
asking for the adjudication before the AMA
Council, if the licensing Board would be
willing to refer my records. Warning me not
to mention his name, Dr. Park suggested that
I try to secure a wise physician-advisor in
Madison to guide me step by step so as to
avoid political mistakes. No one was willing
to take an open stand in what had become a
controversial issue.
In another letter from Dr. Park on 27 De¬
cember, just two weeks prior to the Board’s
meeting on 10 January 1951, he indicated
that he had written Dr. Anderson “that if the
Board did not grant your request at their ap¬
proaching meeting, ... I could no longer
restrain myself. ... I should not be surprised
if Dr. Anderson exerted some pressure on
the Board, for he said to me over the tele¬
phone, ‘Let’s wait and see what they do on
January 10.’ . . .He expressed a belief that
they would pass ‘Hania’ on that date ... if
they fail to pass Hania on that date, he [Dr.
Anderson] would recommend some action.”
Dr. Park also wrote of the possibility of
seeking publicity to expose the Board’s re¬
fusal to license me, perhaps consulting the
New York Times or the Washington Post. In
his strategy letter of 29 November, he had
written: “It might be possible to create enough
sentiment in Wisconsin so that the Board
would be forced out.’ ’ I do not think Dr. Park
realized the political power of the Board.
Meanwhile, the communications between
my attorney, James Doyle, and the Board’s
attorney, John W. Davison, accelerated. Be¬
tween 12 July and 31 October there were
seven such exchanges. In November they ex¬
changed ten letters; in December thirteen, in
addition to a number of telephone calls. There
were always delays in Davison’s answers to
my attorney’s letters, in spite of the fact that
we were critically short of time. For instance,
the letters from the Swiss authorities evalu¬
ating my credentials dated 15 August and
addressed to the Board were not forwarded
to us until 15 November, in spite of several
earlier requests. And at this late juncture the
Board asked me to translate the documents!
Another example of delay and harassment:
On 29 November my attorney requested that
the record of my two semesters (1942-43)
in the Graduate School of the University of
14
In Quest of the Right to Practice Medicine in Wisconsin
Cincinnati, which was in the possession of
the Board, should become part of my official
record. Although the “record book” with
entries constituted an “official transcript” as
the term is commonly used, the Board’s at¬
torney now insisted that we obtain a certified
copy of the official transcript from the uni¬
versity. We complied.
On 12 December my attorney reminded
Davison that the Board now possessed two
documents verifying that my studies at the
University of Zurich had included more se¬
mesters and more courses than required for
the Certificate of Medical Studies, which was
now approved by the AM A Council. (These
were from Dean F. Schwarz, of the Univer¬
sity of Zurich Medical School, and from Dr.
P. Vollenweider, Director of the Federal
Health Department.) My courses of study
would have entitled me to examination for
the Certificate of Medical Studies had such
a certificate been offered at the time I com¬
pleted my studies. Furthermore, Mr. Doyle
reminded the Board that I had taken the same
medical courses as those taken by Swiss cit¬
izens then entitled to examination for the
Federal license.
Mr. Doyle wrote: “I assume that any pre¬
vious uncertainty has now been dispelled by
the AM A Council’s formal approval of the
Medical School of the University of Zurich,
coupled with the unequivocally favorable
evaluation of Dr. Ris’ credentials. ... Dr.
Ris will very much appreciate your early ad¬
vice as to the time and place in January at
which she will be expected to appear before
the Board on her application for licensure by
reciprocity.”
But the Board was not yet willing to accept
defeat. Davison stated in a 14 December re¬
ply that the two documents mentioned by my
attorney were in the process of being trans¬
lated. Translation was hardly the obstacle this
implied. The records were, after all, in Ger¬
man, not Sanskrit, and were only two
pages long.
Attorney Davison’s letter continued: “It
appears to me that the facts which you an¬
ticipate being included in the letters from [the
University of Zurich] . . . could be very eas¬
ily established by procuring from the Uni¬
versity of Zurich the course of study required
for a Certificate of Medical Studies and an
official transcript of Dr. Ris’s credits. If a
comparison of these two documents reveals
that she has taken all of the courses required
for a certificate of medical studies, it would
seem that that particular question would be
definitely answered.”
They were asking for documents they al¬
ready had! They had possessed, since 1949,
the official transcript of my credits and had
had the official documents from Switzerland
concerning courses required for the Certifi¬
cate of Medical Studies since 11 July 1950.
Previously the Board had agreed it was
willing to rely on the direct evaluation of the
Swiss authorities. Apparently it had intended
to honor this only if the result was detrimental
to my record. Now the Board was proposing
a different procedure and adopting new cri¬
teria less than a month before the meeting
where my professional future was to be de¬
cided; I could not interpret this in any way
other than that the Board had been acting in
bad faith.
On 26 December the Board’s attorney called
my attorney to solicit his help in making a
comparison of my courses with those re¬
quired for the Certificate of Medical Studies.
On 27 December Mr. Doyle made the com¬
parison using two parallel columns. I came
off with flying colors. Yet at this late date I
still had not been granted permission to ap¬
pear before the 10 January 1951 meeting of
the Board.
On 27 December my attorney wrote a two-
page letter to Dr. J. W. Smith, president of
the Board (with copies to members of the
Board and its Council), summarizing my one
and a half year struggle for licensure. Mr.
Doyle pointed out that I was entitled to be
informed without delay whether the Board
would grant me permission to appear at its
upcoming meeting.
This was the last document in my own and
attorney James Doyle’s files of my case. What
followed must have been transacted over the
15
Wisconsin Academy of Sciences, Arts and Letters
telephone because of time constraints.
I was told I would be permitted to appear
before the Board on 10 January 1951 to take
the oral examination for licensure by reci¬
procity. Dr. Edwards Park, my advocate,
awaited the outcome anxiously. Dr. Anderson
of the AM A Council wired Dr. Park on 5
January:
Your letter of January 3 just received. Have
telephoned Dr. Christofferson, chairman of the
Wisconsin Board, who assured me without re¬
servation that Dr. Ris will receive exactly same
type of oral examination as that given to all
physicians seeking licensure in Wisconsin by
reciprocity. Written examination was waived
for her as it is for other candidates for reci¬
procity to spare unnecessary ordeal. I feel con¬
fident that Dr. Christofferson will insure Dr.
Ris a fair examination, [signed:] Donald G.
Anderson MD.
Round IV: I Am Finally Licensed
My appearance before the Board was sum¬
marized in the rather anticlimactic language
of the 10 January 1951 minutes of the Board:
“Dr. Hania Ris, applicant number 25, was
ushered into the room. Dr. Ris’ application
has been reviewed again in the light of letters
from the school from which Dr. Ris gradu¬
ated, giving information that she had re¬
ceived the same education and had taken the
same examination as those students who had
received the accepted degree following which
she left the room.”
My name appeared later in the minutes
among the list of candidates receiving the
Wisconsin license by reciprocity. After one
and a half years of painful negotiation with
the State Board of Medical Examiners, I fi¬
nally experienced one humane act. In mid-
January 1951 I received a letter from Dr.
C. A. Dawson (erroneously dated 13 January
1950 instead of 1951), stating: “Knowing
you are naturally anxious as to the outcome
of your examination, I am telling you con¬
fidentially that you were successful. . . . The
list of all newly licensed physicians will be
furnished shortly.” The first congratulatory
call came from Mrs. Edwin B. Fred (Rosa),
the wife of the president of the university,
who had kept in touch with me throughout
the struggle. Her support was typical of the
non-medical community.
It is ironic, however, that the State Board
of Medical Examiners likely was not follow¬
ing the 1937 law when they denied me a
license to practice medicine in Wisconsin. In
a February 1991 Legislative Reference Bu¬
reau legal opinion, Mr. Barry J. Stem, leg¬
islative attorney, indicates the following:
In my opinion, while the board appears to have
had the authority under the 1937 law to adopt
a policy of accepting an application for ex¬
amination for licensure to practice medicine
from any graduate of a foreign medical school
that was classified in the American Medical
Association (A.M.A.) rating, the board did not
appear to have the authority under that law to
accept an application from a foreign graduate
only if the applicant was a graduate of one of
the A.M.A. classified schools. On its face, the
1937 law, which required an applicant to have
a diploma from a “reputable professional col¬
lege approved and recognized by the board,”
would appear to have required the board to
provide a foreign applicant who was a graduate
of a school that was not classified by the A.M.A.
with an opportunity to show that his or her
school was reputable. [Personal correspon¬
dence, 4 February 1991]
Of course, the saga of my quest for licensure
in Wisconsin does not end on the date of 10
January 1951 . I paid a considerable price for
being a test case, in addition to the price of
being a woman challenging the medical es¬
tablishment. I had many experiences as a
persona non grata; one incident stands out.
While awaiting the decision of the Board,
I attended clinical conferences held regularly
at the University Hospital. At one confer¬
ence, a prominent professor of gastro¬
enterology approached me during a lecture
and said: “You have to leave, you did not
register.” There were approximately forty
participants in the room, which had a large
capacity. I knew I was not displacing any¬
body by my presence, but I received a public
rebuke because of my controversial status.
16
In Quest of the Right to Practice Medicine in Wisconsin
Four decades later, the professor’s command
still rings in my ears. However, at an Alumni
Conference reception in 1981, some thirty
years after the episode, the same professor,
then approaching ninety years, came up to
me, shook my hand and then kissed it (which
was quite unusual for someone without a Eu¬
ropean background), and said, “I had to do
what I have done.” “I forgive you,” I replied.
It is true that the struggle to be recognized
for my professional credentials and expertise,
to have the right to practice medicine in the
state of Wisconsin, left some personal scars.
But there were rewards in winning the battle,
not just for me but for the many foreign phy¬
sicians who followed.
Aftermath: The Status of Foreign
Graduates
Since the conclusion of my personal battle,
Wisconsin laws pertaining to foreign-educated
applicants have been liberalized. The law of
1957 provided that if an applicant had grad¬
uated from a foreign medical school that was
not approved or recognized by the Board,
but had postgraduate training in this country
substantially equivalent to training at the
University of Wisconsin, the Board might
admit the applicant to examination. How¬
ever, this law allowed no more than twenty-
five licenses a year to be granted under such
conditions, and the ruling was to expire in
1961. After that date the fixed quota of for¬
eign medical graduates who could be licensed
each year was increased to fifty. In 1969 the
Board started to rely selectively on exami¬
nations conducted by the Educational Coun¬
cil for Foreign Medical Graduates. Since 1970
Wisconsin law has governed the licensure of
graduates of foreign medical schools under
provisions similar to those of 1957 but with¬
out the limitation to fifty licenses annually.
In the opinion of Mr. Earl Thayer, who
was employed by the Medical Society from
1947 to 1957 as public relations person, later
serving as assistant secretary (1957-70) and
as secretary of the Society (1970-87), my
test case forced the Board to rethink and re¬
vise its policy and to accept the AMA Coun¬
cil’s approval of some foreign medical schools.
In the years 1930 to 1949, among active phy¬
sicians in Wisconsin who were counted in a
five-year period (Wisconsin Division of
Health, Center for Health Statistics), the
number of foreign graduates ranged between
6 (0.7%) and 44 (5.1%). Between 1950 and
1954 the number of foreign graduates in¬
creased to 140 (16.2%) (I was the first con¬
tributor to this increase), and in the years
1955-60 it increased to 218 (25.3%).
The Woman Question
What role, if any, did the fact that I was
a woman play in the Board’s attitude? I have
never been sure. My perception, no doubt,
was colored by almost a decade of earlier
positive working experiences in friendly,
congenial atmospheres where colleagues,
professors, and administrators had gone out
of their way to be helpful. The first time I
experienced discrimination was when I came
to Madison.
One authority is persuaded that being a
woman and being aggressive were pivotal
factors. Being aggressive was a positive trait
in the world of men, but it was negative when
applied to women. Mr. Earl Thayer, the
Medical Society’s respected secretary, re¬
cently told me that he had been appalled at
the way the Board operated not only in my
case but in general. He said I had been viewed
as “aggressive” and the Board had hoped its
tactics would discourage me.
The general lack of recognition and respect
given to women in medicine was certainly a
factor in my struggle to gain the right to
practice medicine in Wisconsin, but perhaps
it is illustrated even more clearly by a job
offered to me in 1951 in Milwaukee. I was
the mother of a fourteen-month-old infant at
the time and had no private transportation,
which prohibited my commuting. Neverthe¬
less I was offered a position by the Bureau
of Maternal and Child Health for the City of
Milwaukee, which is about eighty miles from
Madison. At first it was suggested that I hitch
a ride daily with a truck driver at truck stops !
Although I have never been conventional, I
17
Wisconsin Academy of Sciences, Arts and Letters
rejected that idea. It was then suggested that
I take a bus which left daily from Watertown,
Wisconsin, at 6:30 a.m. and arrived in Mil¬
waukee at 7:45 A. M. “In other words, it is
only necessary for you to find transportation
from Madison to Watertown [a distance of
about thirty miles] to make your daily jour¬
ney here possible.’ ’ This kind of sacrifice was
expected of a woman physician in the 1950s:
a willingness to sacrifice her motherhood, her
child, her personal life, for the privilege of
having a position in the field of public health.
These suggestions would have been less
shocking and more amusing had they not come
from a woman physician who was herself a
promoter of maternal and child health.
Women have had to persevere in an arena
and during times when medicine was con¬
sidered male territory. Statistics bear this out.
At the time I received my licensure, in 1951,
there were 204 women physicians (5.5%) in
Wisconsin, compared to 3,492 male physi¬
cians. In 1960 the percentage went down to
4.5% (3,833 males, 183 females). It has risen
steadily since that time: 5.9% in 1978, 7.4%
in 1980, and 9.9% in 1984, the last year for
which statistics are available (Department of
Health and Social Services, Center for Health
Statistics).
But even today there are relatively few
women in medical academia. In 1981 women
constituted only 17% of all medical school
faculty. Few women chair medical school
departments, and few are in leadership po¬
sitions in professional organizations. A case
in point is the American Academy of Pedi¬
atrics, which was established in 1930, and
now has a membership of 37,000 (25% of
whom are women). Only in 1986 did a woman,
Dr. Betty Lowe, become a member of the
nine-person Executive Board. The Acade¬
my’s first woman president, Dr. Antoinette
Eaton, became vice-president and president¬
elect in 1989, by a majority vote of the Acad¬
emy membership.
“Living the Good Life”
While public health medicine appealed to
me, I turned to a much more reasonable and
agreeable alternative. My family and I looked
for a house in Madison that could lend itself
to combining living quarters with a physi¬
cian’s office, where I could practice without
outside pressures and spend as much time
with each patient as necessary. We found
such a house, surrounded by large trees, at
2306 Van Hise Avenue, across the street from
West High School before its expansion. This
work arrangement was rather unusual in
Madison but quite common in the East. I
would have preferred an academic position
or an association with an obstetrician in an
office, but this was unrealistic; I was still
perceived as too controversial and too much
a risk for close professional associations such
as these.
Many of our friends, university teaching
staff, people whom I came to respect and
admire, entrusted me with their children. As
a pediatrician, I later became a specialist in
adolescent medicine, and then part-time
medical director of a school for delinquent
girls. I developed a comprehensive, multidis¬
ciplinary health program for the underprivi¬
leged young women, which led to clinical
research in the field of sexually transmitted
diseases and to the position of medical di¬
rector of all Wisconsin state correctional in¬
stitutions under the jurisdiction of the De¬
partment of Health and Social Services,
Division of Health. Prevention of teenage
pregnancy through sex education and the
elimination of legal barriers to control of re¬
production for teenagers and adults has been
an important part of my activities.
Despite the dire predictions that I would
be barred from academic medicine forever,
since 1956 I have been a member of the Uni¬
versity of Wisconsin Medical School faculty
and am currently a Clinical Professor of Pe¬
diatrics. I have published a number of profes¬
sional articles, mainly in the field of sexually
transmitted diseases in young adults.
Of course, since Life magazine’s 1948 ar¬
ticle about ‘ The Good Life in Madison, Wis¬
consin,” many things have changed. Never¬
theless, I have enjoyed my forty years as a
Madison resident immensely and would still
18
In Quest of the Right to Practice Medicine in Wisconsin
contend that it is one of the best places in
America to live. My fondness for Madison
is probably even stronger because I had to
fight for the right to make it my home, to be
able to practice my profession without dis¬
crimination; I have always believed that is
the right of every American citizen. My per¬
severance has been amply rewarded.
Acknowledgments
It should be obvious to the reader that I
did not persevere without a great amount of
encouragement and support. The press of
Wisconsin championed my case with nu¬
merous articles, and without its airing of the
issues my efforts would have been far more
difficult. The late James Doyle, my attorney,
exhibited incredible skill and patience in
dealing with the machinations of the Board
of Medical Examiners. He contributed greatly
to bringing my case to a successful resolution.
I wish to reserve a special place to cele¬
brate the late Dr. Edwards A. Park, physi¬
cian, scientist, champion of medical care for
the poor, early supporter of Medicare and
Medicaid, and devoted friend. Dr. Park
championed my cause out of his intense com¬
mitment to human decency, fairness, and jus¬
tice. He was among the first to oppose the
AMA’s conservative policy regarding social
health issues. In his teaching and by the ex¬
ample he set, he instilled in people the im¬
portance of the search for knowledge, the
pursuit of truth, honesty, and high standards
in all aspects of life. He published over a
hundred articles, and until his death in 1969
at age ninety-one, his expertise was sought
by authors of scientific publications. Much
of the material needed to reconstruct the events
described in this article came from the vo¬
luminous correspondence I had with Dr. Park,
who obtained confidential information and
advice from many individuals.
19
1988 Drought Impacts
Among Wisconsin Dairy Farmers
John A. Cross
Abstract . Drought such as occurred throughout the American Midwest during 1988 was an
unusual experience for Wisconsin' s farmers , who lost half their hay and corn crops . Dairy
operators, who represent nearly half of the state' s farmers , faced added hardships in main¬
taining their herds in face of feed shortages and rising feed prices. This paper reports the
findings from a survey of Wisconsin dairy farmers concerning the drought impacts and the
adoption of various drought mitigation measures. The consequences of the drought were most
severely felt by farmers already experiencing a variety of economic stresses. Although three-
quarters of the dairy farmers reported receiving federal drought assistance payments, 73%
of these farmers would have survived without such relief. Farmers are pessimistic about future
drought occurrences.
Drought is a frequent and ever-present
hazard for farmers tilling subhumid and
semi and lands and has been most studied
within such environments (Hurt 1981; Ro¬
senberg 1978; Saarinen 1966; and Warrick
1975). Although the rare drought events within
normally humid environments have received
less attention, the impacts of unusual drought
occurrences can be highly significant and are
worthy of study. This paper reviews the im¬
pacts of the 1988 summer drought upon dairy
John A . Cross is an associate professor and chair of
the Department of Geography at the University of Wis-
consin-0 shkosh. His other recent publications have dealt
with geographic literacy, natural hazards, and the Dairy
Termination Program in Wisconsin.
Funding for the research survey was provided by a grant
from the Natural Hazards Research and Applications
Information Center of the University of Colorado, Boul¬
der, through their quick-response research program .
Any opinions, conclusions, or recommendations ex¬
pressed in this paper are those of the author and do not
necessarily reflect the view of the Hazards Center or its
funding agencies.
farmers in Wisconsin. At the time of the
drought, 45% of Wisconsin’s 81,000 farmers
were engaged in dairying, leading the nation
in milk production.
Drought conditions were felt throughout
Wisconsin during the summer of 1988, when
“43% of the area of the contiguous United
States was in the severe or extreme drought
category” (Trenberth, Branstator, and Arkin
1988). In Wisconsin the drought resulted in
the loss of approximately half of the state’s
hay and com (maize) crops. During the win¬
ter and spring of 1989 dairy farmers faced
not only the consequences of these feed losses,
but also the possibility of continuing drought
conditions. This paper summarizes findings
from a survey concerning the impacts of the
1988 drought at a time many dairy farmers
would be expected to be experiencing hay
and feed grain shortages resulting from the
substantially diminished 1988 harvest. The
1988 drought provided an excellent oppor¬
tunity to study drought impacts, mitigation,
and perception among the population of a
normally humid environment that has rarely
had to deal with such a hazard.
21
Wisconsin Academy of Sciences, Arts and Letters
1988 Drought Conditions
Extreme drought conditions, as defined by
the Palmer Index (Fig. 1), occurred in six of
Wisconsin’s nine agricultural reporting dis¬
tricts during the summer of 1988 {Weekly
Weather and Crop Bulletin 1988). Precipi¬
tation from April through August 1988, as
shown in Figure 2, was the lowest recorded
in ninety-three years of records in both south¬
western and southeastern Wisconsin, with
three additional districts recording their sec¬
ond or third driest growing seasons. Rainfall
was particularly deficient in May and June,
with a Milwaukee weather station reporting
a two-month total of 0.99 inch and Green
Bay recording 0.73 inch — 11.6% of normal
(U.S. Department of Commerce 1988). Ab¬
normally hot temperatures accompanied the
drought, with six of Wisconsin’s nine agri¬
cultural reporting districts reporting their
highest June through August mean temper¬
atures in records dating back to 1895 (U.S.
Department of Commerce 1989b). Although
a portion of eastern Wisconsin received sub¬
stantial rainfall in August and September,
over half of the state experienced an annual
precipitation shortfall of at least six inches,
with the southwestern comer of Wisconsin
receiving fifteen inches below normal pre¬
cipitation (Clark 1989a).
The 1988 com harvest was 60% below the
1987 harvest, alfalfa hay was down 45%,
other varieties of hay were off 44%, and oats
were down 54% (Wisconsin Department of
Administration 1989). Furthermore, because
1987 harvests had fallen from even greater
1986 harvests as a result of less severe drought
conditions in 1987, the 1988 harvests of al¬
falfa hay and com were 46% and 36%, re¬
spectively, of their 1986 harvests. Although
the tonnage of com silage harvested in 1988
was down only 2%, this was accomplished
by a doubling of the harvested acreage, largely
an effort to salvage wilted cornfields that had
been planted for grain (Wisconsin Agricul¬
tural Statistics Service 1988 and 1989). Un¬
fortunately, because of its lower protein con¬
tent, the substitution of such com silage for
alfalfa without additional protein supple¬
ments reduced milk production (Howard and
Shaver 1988).
The dollar value of the 1988 hay harvest
exceeded that of the 1987 harvest by 18.2%
because of the shortage and rapidly escalating
prices. Indeed, hay prices doubled or tripled,
with the price of top-grade alfalfa hay reach¬
ing $250 per ton. Price increases failed to
keep pace with lost production for other crops.
For example, the cash value of the com har¬
vest was down by 48.9%, and the oat harvest
22
1988 Drought Impacts Among Wisconsin Dairy Farmers
generated 25.5% less revenue than in 1987
(Wisconsin Agricultural Statistics Service 1988
and 1989). However, higher commodity prices
were not advantageous to most Wisconsin
dairy farmers. They normally consume their
crop on their farms, and the higher prices
simply translated into higher costs of feeding
their herds to stay in business (Rodefeld
1988a).
Drought-induced crop losses were not uni¬
formly distributed across Wisconsin, with the
western and southern portions of the state
reporting the greatest declines in production
between 1987 and 1988 (Fig. 3). For ex¬
ample, in Polk County the 1988 com crop
was 17.8% of the 1987 harvest, and in Mar¬
athon County — the state’s foremost milk
producing county — the com crop was only
24.5% of the previous year’s harvest. Al¬
though the decline in alfalfa production
(Fig. 4) was not as dramatic as the drop in
the com harvest, similar spatial patterns of
crop losses were noted, with the greatest
drought losses occurring in the north central,
northwestern, and southernmost portions of
Wisconsin.
Press reports during the winter and early
spring of 1989 painted a bleak picture of
conditions facing Wisconsin dairy operators.
Large proportions of farmers had either ex¬
hausted their feed or were expected to do so
before their next harvest. A Wisconsin Ag¬
ricultural Statistics Service survey in early
November 1988 determined that 14% of the
state’s livestock farmers (dairy, cattle, and
hog) expected to have exhausted their hay by
January, 42% by March, and 74% would be
out of hay by May. Grain or grain concentrate
supplies were expected to be similarly ex¬
pended {Wisconsin Farm Reporter 9 Novem¬
ber 1988). Wisconsin’s hay stocks in De¬
cember 1988 were down 3.9 million tons
from December 1987 (Rodefeld 1988b). Re¬
placement of this lost hay and haylage state¬
wide was estimated to cost from $600 to $700
million, and replacement of com stocks was
estimated between $200 and $400 million
(Wisconsin Department of Administration
1989).
Weather conditions during the winter and
spring of 1989 caused further concern at the
same time dairy farmers were facing feed
shortages. Freezing rains (rather than snow)
during January 1989 had seriously damaged
the alfalfa fields (Clark 1989b). Precipitation
during the spring of 1989 was well below
normal, with many areas by early May hav¬
ing received less precipitation since the be¬
ginning of the year than in 1988 (U.S. De¬
partment of Commerce 1989a). The 6 May
23
Wisconsin Academy of Sciences, Arts and Letters
1989 Palmer Index ( Weekly Weather and Crop
Bulletin 1989) indicated that severe drought
was occurring in southwestern Wisconsin,
moderate drought was present in central Wis¬
consin, and mild drought was present in five
of the state’s other seven agricultural re¬
porting districts.
Methodology
Concerns about how Wisconsin dairy
farmers had managed to deal with the drought-
induced hay and feed grain shortages during
the winter and spring of 1988-89 and about
these farmers’ vulnerability to future stress —
drought and otherwise — prompted the sur¬
vey of Wisconsin dairy farmers that provided
most of the data reported in this paper. An
eight-page questionnaire was mailed on 4 May
1989 to 506 dairy operators throughout the
state. The initial mailing of the questionnaire,
accompanied by a cover letter and a business
reply envelope, was followed six days later
by a reminder post card. This card thanked
participants and encouraged recipients to
complete and return the survey. Farmers not
responding to the initial survey were mailed
a second copy of the questionnaire, a new
cover letter, and a second business reply en¬
velope on 24 May 1989. Completed surveys
were received from 283 farmers, represent¬
ing 57% of the eligible members of the sam¬
ple who received the survey.
Farmers receiving this survey were se¬
lected by a stratified systematic sampling
procedure from an early April 1989 Wiscon¬
sin Department of Agriculture listing of the
35,611 dairy operators whose herds had had
the Brucellosis Ring Test, which is required
quarterly for all commercial milk producers.
The sampling was designed to select 1.2%
of the dairy operators in the six largest ag¬
ricultural reporting districts. In the remaining
three districts, which would have fewer rep¬
resentatives, an additional 1-2% of the farm¬
ers were selected so that district-to-district
comparisons could be made. In addition, a
short survey was sent to agricultural exten¬
sion agents throughout Wisconsin requesting
data on their observations of farmers in their
various counties.
Impacts of 1988 Drought
The 1988 drought had numerous impacts
upon Wisconsin’s dairy farmers, including
substantial crop losses, losses of income, and
shortages of hay and feed grains, together
with a wide assortment of economic stresses
(Table 1). Some even lost their farms.
Crop losses and feed purchases
Crop losses led to a cascade of drought
impacts for Wisconsin’s dairy farmers. The
Wisconsin State Agricultural Stabilization and
Conservation Service in late August 1988 es-
Table 1. Impacts of the 1988 drought
“Indicate to what degree (if any) you have experienced the following as a result of last
summer’s drought?’’
24
1988 Drought Impacts Among Wisconsin Dairy Farmers
ti mated statewide crop losses of 50% for both
hay and com (U.S. Department of Agricul¬
ture 1988). Crop yields for 1988, as reported
by dairy farmers responding to my survey,
were close to these estimates, with their hay /
alfalfa harvest averaging 45% of normal and
their com crop averaging 50% of normal.
When surveyed in May-June 1989, only 37%
of the farmers indicated they had “sufficient
feed grain supplies to last until the next
harvest.”
Purchases of hay or alfalfa hay had been
made by 62% of the dairy operators between
September 1988 and May 1989, and 72% had
purchased feed grains. Statewide, 75.5% of
Wisconsin’s dairy operators reported making
purchases of either hay or feed grains, with
the proportion ranging from 68 to 83% of the
farmers in the various agricultural reporting
districts. In a normal year, over 40% of the
surveyed farmers buy neither hay nor feed
grains. Purchases of hay and feed grains dur¬
ing 1988-89 averaged $14,616, with a me¬
dian purchase cost of $9,000. The mean cost
of these purchases in a normal year averaged
$6,733 with a median cost of $2,000. The
greatest increase in hay and feed purchases
was reported by dairy farmers in southwest¬
ern Wisconsin.
Changing feed, reducing herd size
Farmers also took a variety of other actions
(Table 2) to mitigate the feed shortages. For
example, 35% changed the type of feed given
to their animals, 22% reduced the amount of
feed, and 38% of the respondents reduced
the size of their herds. Conversely, 27.8%
of the survey respondents were able to in¬
crease their herd size.
Farmers most likely to reduce the size of
their dairy herds were located in central Wis¬
consin (where over half took this action), as
well as the southwestern, northeastern, and
south central Wisconsin regions. With the
exception of southwestern Wisconsin, which
experienced the greatest precipitation deficits
in 1988, districts where dairymen were less
likely to purchase feed supplies were those
most likely to have reductions in herd size.
Farmers in east central, west central, and
southeastern Wisconsin (42 to 50%) were most
likely to change the type of feed given to
their herds, while those in northeastern and
south central Wisconsin were least likely (18
to 23%). Reduction of the amount of hay or
feed grains fed to the dairy cows was reported
by over 30% of the dairymen within central,
southwestern, and southeastern Wisconsin,
while fewer than 14% of the dairymen within
Table 2. Actions taken because of hay or feed grain shortages
% % % %
‘Because Wisconsin’s dairy farmers are not evenly distributed among the nine agricultural reporting districts,
this total was calculated by weighting the responses from each district by the proportion of Wisconsin’s
dairy farms that operate in that district.
25
Wisconsin Academy of Sciences, Arts and Letters
the northern third of the state put their herds
on short rations.
Drop in farm income
Net farm income was down for the ma¬
jority of the dairy farmers, yet for better than
one in ten, income was above average. When
asked an open-ended question, “Your net
income (from all sources) for 1988 was about
what percent of normal?” one-quarter of the
farmers indicated 100% or more (Table 3).
Dairy farmers reporting above normal in¬
come — with some reporting their best year
ever — typically had substantial hay supplies
carried over from 1987. Conversely, 11%
reported net incomes of 50% or less of nor¬
mal, with the mean net farm income being
84.5% of normal (median was 90%). These
estimates are similar to those of the agricul¬
tural extension agents, who estimated that
average income was 89% of normal. Al¬
though one out of seven of the agents esti¬
mated that the average farmer within his or
her county earned a greater than normal in¬
come, in none of the agricultural reporting
districts was the average net income (either
mean or median), as reported by the farmers,
greater than 90% of normal. The greatest
departures from normal were reported by dairy
farmers in central and southwestern Wisconsin.
Sixty-seven percent of the dairy farmers
surveyed indicated that decreases in their net
farm income were strongly or moderately felt,
an even larger proportion than those farmers
reporting decreases in gross farm income
(56%, Table 1). Half of the farmers indicated
Table 3. Change in net farm income from
normal during 1988
“Your net farm income (from all sources) for
1988 was about what percent of normal?”
that shortages in hay or alfalfa were “strongly
felt,” with an additional 24% claiming these
shortages were “moderately felt.” Never¬
theless, increases in farm indebtedness as a
result of the drought were strongly or mod¬
erately felt by only 38% of the farmers.
Drought-induced bank foreclosure threat was
strongly or moderately felt by 8% of the dairy
farmers, and the sale of lands or farm equip¬
ment was similarly felt on 10% of the farms.
Thirty-five percent of the farmers indicated
that a “greater need of off-farm income” was
strongly or moderately felt.
Spatial differences in “strongly felt” im¬
pacts of the drought were noted (Table 4).
Farmers in southwestern, central, and east
central Wisconsin were most likely to report
strongly felt shortages of hay or feed grains.
Decreases in both gross and net farm income
were most frequently reported in northeast¬
ern, central, and southeastern Wisconsin, all
areas where dairy operators had faced above
average economic stresses and declines in the
previous decade (Cross 1989). Dairymen in
the central Wisconsin region were signifi¬
cantly more likely to have strongly felt in¬
creases in farm indebtedness, while farmers
in the north central, west central, and central
agricultural reporting districts were most likely
to have a greater need for off-farm income.
Although these spatial patterns are not en¬
tirely consistent, farmers in central Wiscon¬
sin consistently reported above average lev¬
els of concern about all the potential drought
impacts.
Drought-induced declines in both gross and
net incomes, shortages of hay and feed grains,
and increased indebtedness were experienced
by a broad spectrum of Wisconsin dairy
farmers. No differences in drought impacts
were noted among the farmers based upon
the age of the farmer, the number of years
as farm operator, farm acreage, size of dairy
herd, the farmer’s land tenure status, or
whether the farm was a grade A or grade B
operation. On the other hand, the responses
of the farmers to the shortages of hay and
feed supplies were related to a number of
these characteristics (Table 5). For example,
26
1988 Drought Impacts Among Wisconsin Dairy Farmers
Table 4. Impacts of the 1988 drought in Wisconsin’s agricultural reporting districts
Percent of dairy farmers indicating that the condition was
“strongly felt’’ as a result of the summer 1988 drought
‘Because Wisconsin’s dairy farmers are not evenly distributed among the nine agricultural reporting districts,
this total was calculated by weighting the responses from each district by the proportion of Wisconsin’s
dairy farms that operate in that district.
Table 5. Significant relationships between dairy farmer characteristics and responses to
drought-induced hay/feed shortages
Farmer responses
*NS indicates chi-square not significant at .1000 significance level.
farmers most likely to have already pur¬
chased hay or feed grains as a result of the
drought were the youngest, those with lower
net incomes, those with the largest number
of cows, and those with off-farm incomes.
Drought Relief
Wisconsin dairy farmers were asked to
evaluate the importance of various factors in
helping their farms financially survive the
1988 drought and its aftereffects. The two
most important factors were government
drought relief payments and increased milk
support prices, cited as “very important” by
44 and 41% of the farmers, respectively. Per¬
sonal savings were cited as “very important”
by 26% of the dairy operators, bank credit
by 22%, and off-farm income by 17%. Crop
insurance payments were “very important”
to only 8% of the surveyed farmers.
Several federal and state government pro¬
grams provided assistance to Wisconsin
farmers. Lands in the Conservation Reserve
Program and Conservation Use (set-aside)
27
Wisconsin Academy of Sciences, Arts and Letters
program were opened to both grazing and
haying. State-owned lands and highway right-
of-ways were opened to haying. Additional
federal funds were authorized to purchase
ground beef, assuring that dairy farmers who
liquidated their herds could do so with a rea¬
sonable market for their cows. A scheduled
drop in the federal milk support price was
postponed, and support prices were in¬
creased. Property tax credits and a guaran¬
teed loan program for farmers were approved
by the state (Richards 1988; Wisconsin De¬
partment of Administration 1989). The larg¬
est relief program was provided by the U.S.
Disaster Assistance Act of 1988, which au¬
thorized compensation to farmers with crop
losses exceeding 35% of normal production.
In general, farmers received no compensa¬
tion for their first 35% of lost production,
were compensated at 65% of the target price
(approximately the pre-drought average mar¬
ket price) for the loss of 36 to 75% of their
harvest, and 90% of the target price for the
loss of 76 to 100% of production. Thus, the
program did not “prevent farmers from ex¬
periencing substantial declines in their in¬
comes” (Jones 1988).
Financial assistance through the Disaster
Assistance (Drought Relief) Act was re¬
ported by 75% of the Wisconsin dairy farm¬
ers surveyed, a greater proportion than those
who reported that this assistance was either
“important” or “very important” in helping
their farm financially survive the drought.
Thirty-seven percent indicated that these
drought relief payments were the primary
source of funds for their hay and feed grain
purchases. Thirty-five percent relied primar¬
ily upon their farm income (milk check) or
withdrawal of funds from their savings to
purchase hay or feed. Thirteen percent bor¬
rowed funds from banks or other financial
institutions.
Seventy-three percent of the surveyed
dairymen who received drought relief pay¬
ments indicated that they would have been
able to remain in the dairy business even
without the aid. In contrast, Wisconsin ag¬
ricultural extension agents estimated that
without the drought relief payments only 9%
of the dairy farmers would succumb. Dairy¬
men in southeastern, south central, and west
central Wisconsin expressed the greatest con¬
fidence that they would have survived with¬
out any drought relief payments. Conversely,
dairy farmers in the central, east central, and
north central agricultural reporting districts
were least confident about their ability to have
survived the drought without drought relief
payments. In central Wisconsin only 53% felt
they would have survived without the
payments.
If we consider only those farmers who re¬
ceived drought relief payments, these pay¬
ments were most significant in the survival
of Grade B dairy farms, the farmers with off-
farm income or employment, those farmers
who reported that decreases in net farm in¬
come were moderately or strongly felt, and
the farmers reporting the greatest hay and
com crop losses in 1988. Indeed, 41% of the
dairy farmers receiving drought assistance who
had lost over two-thirds of their hay/alfalfa
crops doubted their ability to survive without
those payments (Table 6).
Drought relief payments and other federal
and state benefits, estimated at $565 million
for Wisconsin farmers, covered approxi¬
mately 37% of feed expenses and lost cash
crop revenues of Wisconsin farmers. Thus,
Wisconsin farmers (both dairy and crop) had
estimated uncompensated losses averaging
over $11,000 each (Wisconsin Department
of Administration 1989). Since even a year
before the drought 16% of all Wisconsin farms
(a total of 12,800 farms) were considered by
the U.S. Department of Agriculture to be
experiencing “extreme financial stress,” for
many farmers such losses were unbearable.
Furthermore, Rodefeld (1988) indicated,
“Many farmers who survive the coming win¬
ter will have high levels of stress in future
years because of their higher debt loads and
tighter cash flows from this year’s drought.”
Loss of Dairy Farms
Statewide, an estimated 960 farmers had
already terminated their dairy operations by
28
1988 Drought Impacts Among Wisconsin Dairy Farmers
Table 6. 1988 hay/alfalfa crop loss and ability of drought relief recipients to survive
without payments*
*Chi-square = 16.010, 2 degrees of freedom, significance = .00033.
early May 1989 as a direct result of the 1988
drought, based upon estimates of county-level
agricultural extension agents. Between March
1988 and March 1989 the total number of
commercial herds in Wisconsin dropped by
1,351, a decline of 3.7% (Wisconsin Agri¬
cultural Statistics Service 1988 and 1989).
Thus, the drought would appear to be the
leading cause of herd losses. Between March
1989 and March 1990 Wisconsin lost another
1,768 dairy herds, a decline of 5%. How¬
ever, to keep these losses in perspective, we
should note that the number of dairy opera¬
tions in Wisconsin fell by 9.8% (a total of
4,026 herds) between 1986 and 1988 (40%
were participants in the Dairy Termination
Program) and fell by 19.2% between 1982
and 1988 (Cross 1989). Although the annual
loss between March 1988 and March 1989
was smaller than within the previous few years,
many agricultural extension agents felt in May
1989 that it was still too early to determine
the total number of casualties from the 1988
drought. Hence, many dairy farmers ceasing
operations between 1989 and 1990 should
also be considered victims of the drought.
The distribution of the losses of dairy
farmers between 1988 and 1989 (Fig. 5) was
similar to that over the previous half decade
(Cross 1989), with a few notable exceptions.
For example, above average declines were
noted in several counties of northwestern
Wisconsin and in central Wisconsin, simply
continuing trends that existed before the
drought. On the other hand, the above av¬
erage losses during 1988-89 in several
southwestern Wisconsin counties (notably
Grant and Iowa) are in sharp contrast to their
considerably below average declines be¬
tween 1981 and 1988. Losses in dairy herds
between 1989 and 1990 more closely parallel
long-term pre-drought trends (Fig. 6), which
saw the greatest declines in northern Wis¬
consin, parts of central Wisconsin, and near
the Milwaukee metropolitan area in south¬
eastern Wisconsin.
Vulnerability to continued
drought stresses
A third of the dairy farmers surveyed in¬
dicated that, if there was a drought during
29
Wisconsin Academy of Sciences, Arts and Letters
Figure 6
the summer of 1989, they would no longer
be in business by the spring of 1990. (Al¬
though five of Wisconsin’s nine agricultural
reporting districts received less average
district- wide precipitation in 1989 than in
1988, strategically spaced rainfall spared most
crops [Naber 1990].) However, 7% of the
surveyed farmers did not expect to still be in
business the next year, even if rainfall amounts
were normal during the summer of 1989.
Furthermore, 29% of the dairy farmers af¬
firmatively answered the question, “Would
you like to sell your farm?” This was par¬
ticularly prevalent throughout the northern
third of Wisconsin, where 40% of the dairy
farmers wished to sell. It should be noted the
entire northern portion of the state saw the
highest rates of farm abandonment over the
past decade and the greatest participation rates
in the Dairy Termination Program (Cross
1989).
Farmers who expressed the greatest con¬
cern about their vulnerability to continued
drought were typically those with smaller than
average farm acreages, with below average
herd sizes, and with Grade B operations
(Table 7). Thus, farmers most threatened by
the drought were the same group the Wis¬
consin Dairy Task Force (1987) had identi¬
fied even before the drought as having the
bleakest chance of economic survival be¬
cause they both lacked economies of scale
and could not “afford relatively expensive
new technologies.” Strong statistical rela¬
tionships were noted between the farmers’
expectations that they could survive a 1989
drought and their crop losses (especially com)
in 1988 and the adequacy of their hay and
feed grain supplies (Table 8). Those farmers
who indicated they did not expect to still be
in business— drought or no drought — by 1990
were generally the oldest, with the greatest
number of years as farm operators, and with
smaller than average herd sizes. However,
those expecting to quit had suffered drought-
Table 7. Significant chi-square relationships between drought vulnerability of farmers and
characteristics of farmers
*AII respondents were asked this question, both those who had and those who had not received any drought
relief payments.
fNS indicates chi-square not significant at .1000 significance level.
30
1988 Drought Impacts Among Wisconsin Dairy Farmers
Table 8. Significant chi-square relationships between 1 988 drought impacts and future
vulnerability of dairy operations
*NS indicates chi-square not significant at .1000 significance level.
induced crop dosses and feed and hay short¬
ages that were no different than the remaining
farmers.
Drought Mitigation for 1989
Many dairy farmers made no efforts to
mitigate possible drought losses in 1989, al¬
though virtually all Wisconsin dairy farmers
had suffered crop losses in 1988 and 70%
expected a drier than normal 1989 growing
season. Nevertheless, crop insurance cov¬
erage expanded, and nearly half the farmers
took some action to reduce future drought
losses.
Crop insurance
Crop insurance to cover drought losses had
been obtained by only 8% of the surveyed
dairy farmers in 1988, although 36% had
obtained insurance to cover hail losses. For
their 1989 crop season, 51% of the dairy
farmers reported that they either had or would
obtain crop insurance to cover drought losses.
Such a low figure is surprising because multi¬
peril crop insurance was required of drought
relief recipients who lost at least 65% of their
crops as a condition for receiving their pay¬
ments (U.S. Public Law 100-387, Section
207). Nevertheless, 16% of the surveyed
farmers whose hay and com crops were both
under 35% of normal — but who still had re¬
ceived drought assistance-had not pur¬
chased drought (or multi-peril) insurance.
Furthermore, several other respondents ob¬
tained only minimal crop insurance coverage
because of its cost. Although the legal re¬
quirements mandating multi-peril insurance
in exchange for drought assistance did not
receive universal compliance, dairy opera¬
tors with the largest crop losses (whether or
not they received financial assistance) were
significantly more likely to obtain drought
insurance for the next year.
Crop planting
The 1988 drought prompted 42% of the
dairy farmers to make changes in their crop
planting plans for 1989. Farmers in north¬
western, north central, and east central Wis¬
consin were significantly more likely to re¬
port these changes. Numerous changes in
cultivation techniques and crops were un¬
dertaken, although only a few farmers men¬
tioned changes in plowing/planting dates, re¬
duced tillage, or fertilizer and herbicide usage.
Farmers with the greatest acreages were
most likely to report making changes in their
crop planting plans for 1989. Likewise,
younger and middle-aged farmers were sig¬
nificantly more likely to report making changes
than the older farmers (those over sixty years
of age). On the other hand, the land tenure
status of the farmers, the size of the farmer’s
dairy herd, and whether the herd was a Grade
A or Grade B operation were not statistically
related to crop planting changes. The deci¬
sion to make changes in crop planting plans
was significantly related to both the farmers’
perceptions of the likelihood of drought dur¬
ing the summer of 1989 and their perception
31
Wisconsin Academy of Sciences, Arts and Letters
that drought possibilities are a problem in
Wisconsin.
The prominence of hay/alfalfa, com, and
oat production on Wisconsin dairy farms re¬
mains unchanged following the drought.
Ninety-five percent of the surveyed farmers
reported planting com in 1988, and 94% in¬
tended to grow com in 1989. Similar pro¬
portions produced hay and/or alfalfa. Oats
were cultivated on 73% of the dairy farms in
1988, the same proportion that planned to
grow oats in 1989. However, a slightly greater
amount of crop diversification was planned
for 1989, and the proportion of farms pro¬
ducing many of the lesser grown crops in¬
creased. For example, dairy farmers planting
sudan grass increased from 9.6% in 1988 to
13.3% in 1989 and sorghum from 5.2% to
7.3%.
Irrigation is a rarity on Wisconsin dairy
farms. Only 3.2% of the surveyed farmers
had irrigation systems in place before 1988,
with an additional 1.4% installing systems
during 1988. Only two (of the 283 farmers
responding to the survey) planned to install
an irrigation system during 1989. Statewide,
only 250 of Wisconsin’s 81,000 farms in¬
stalled emergency surface water irrigation
systems during the summer of 1988 (Wis¬
consin Department of Administration 1989).
Conclusions
The drought of 1988 has provided us with
a unique opportunity to study drought per¬
ceptions and drought mitigation among farm¬
ers who have rarely dealt with this hazard.
Drought is but one of many conditions that
threaten the livelihood of Wisconsin dairy
farmers. When asked to evaluate drought and
a variety of other problems, farmers more
frequently mentioned everyday economic
concerns as being major problems than any
natural hazard, including drought, hail, and
flood. Indeed, milk support prices were con¬
sidered a major problem by 53% of the farm¬
ers, property taxes by 51%, and drought by
36%. Only 9% of the farmers ranked drought
possibilities as the single most important
problem facing dairy farmers in their Wis¬
consin county, compared with 45% who cited
either milk support prices or wholesale milk
prices.
The final toll of the 1988 drought upon
Wisconsin’s dairy farmers will take years to
tally fully. However, between March 1988
and March 1990 Wisconsin lost 3,119 dairy
operations, an 8.4% decline. The economic
stresses caused by the drought-induced di¬
minished feed stocks and high hay and feed
grain prices were somewhat mitigated by in¬
creased milk production per cow, rapidly es¬
calating milk prices, and drought relief pay¬
ments. The greatest stresses of drought did
not necessarily occur in those areas experi¬
encing the greatest meteorological drought or
crop losses, but in areas where farmers were
already under economic stress, and thus lack¬
ing in the resilience to respond successfully
to another threat. In this respect, Wisconsin
dairy farmers are no different from farmers
in Mexico, where Liverman (1990) made
similar observations. For many Wisconsin
dairy farmers, high debt loads remain, only
increased by the stresses of the drought.
Wisconsin dairy farmers are generally pes¬
simistic about the possibility of future
droughts. Another drought, as severe as the
1988 drought, is expected within ten years
by nearly half of the farmers surveyed (Cross
1990). If predictions of climatologists about
climatic warming because of the Greenhouse
Effect are accurate (Schneider 1989), Wis¬
consin farmers must learn to deal with an
increasingly capricious environment. Al¬
though the Greenhouse Effect cannot be
blamed for an individual drought such as that
during 1988, “the greenhouse effect may tilt
the balance such that conditions for droughts
and heat waves are more likely” (Trenberth,
Branstator, and Arkin, 1988). We should re¬
member what we have learned from this
drought experience, which nationally was
overshadowed only by the droughts of the
1930s and 1950s. Indeed, we should not for¬
get the advice of Miewald, who wrote after
another drought, “If we learn nothing from
the current drought, then it may be said that
32
1988 Drought Impacts Among Wisconsin Dairy Farmers
the worst impact is no real impact at all”
(1978).
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Wisconsin Academy of Sciences, Arts and Letters
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34
Distribution, Abundance, Larval Habitats,
and Phenology of Spring Aedes Mosquitoes
in Wisconsin (Diptera: Culicidae)
Jeffrey W. Gilardi and William L. Hilsenhoff
Abstract. In 1988 10,651 larval and 1 ,612 adult Aedes were collected in nine representative
areas of Wisconsin during the spring and early summer; an additional 3,146 larvae were
collected during the spring in 1989 and 1990. From these collections and previous studies
we determined the distribution, relative abundance, larval habitats, and probable phenology
of the twenty-three species of Aedes that breed in water from melting snow and early spring
rainfall. Aedes decticus and A. euedes were collected for the first time in Wisconsin. Adult
longevity and flight ranges were also studied.
In May and June Aedes stimulans was the predominant mosquito biting humans in central
and southern Wisconsin, while A. communis and A. punctor were the most important pests
in the north. Adults of A. vexans were late spring pests in most areas in 1988 but were absent
in 1989 and restricted to west central Wisconsin in 1990. Adults of A. canadensis, A.
excrucians, A. fitchii, and A. provocans were minor pests in most areas of the state but were
abundant locally. Adults of A. cinereus were troublesome biters in woodland areas throughout
Wisconsin.
Almost every year in May and June mos¬
quitoes become a nuisance in wooded
areas throughout Wisconsin. This problem is
created by females of twenty-three species
of Aedes that breed in temporarily flooded
areas, which have resulted from snowmelt
and early spring rains. The mosquito nui¬
sance is especially severe after heavy snow¬
melt and/or heavy rains. Adults of these spring
Aedes mosquitoes have a limited flight range
and tend to remain in areas near larval de¬
velopment sites. Most are relatively short-
William Hilsenhoff is Professor of Entomology at the
University of Wisconsin-Madison. As a graduate stu¬
dent, he worked with mosquitoes. Since 1957 he has
studied all types of aquatic insects that inhabit Wiscon¬
sin’s streams, lakes, ponds, and other habitats. Know¬
ing that Wisconsin’s mosquito fauna is poorly known,
he began to study mosquitoes again six years ago.
Jeffrey W. Gilardi teaches at the University of Illinois,
Urbana, in the Department of Entomology.
Research supported by the College of Agricultural and
Life Sciences, UW-Madison, and by Hatch Research
Project 3050.
lived, and after June, mosquitoes that have
emerged in spring are rarely a nuisance.
Fifty-three species of mosquitoes are known
from Wisconsin; twenty-eight of them are
Aedes. All adult mosquitoes probably feed
on floral nectar and other plant liquids (Grim-
stad 1973), but only females take blood meals.
Females of some species feed on reptiles and
amphibians, others prefer birds, but those of
at least forty species, including all Aedes,
readily attack humans and other mammals.
This study was undertaken to determine the
distribution, relative abundance, larval hab¬
itats, and nuisance potential of each species
of Aedes that emerges during the spring and
to summarize previous studies of these spe¬
cies in Wisconsin.
Larvae of all Aedes in Wisconsin develop
in areas that are temporarily inundated with
water, ranging from small cavities, holes,
and depressions to marshes, ponds, bogs, and
swamps. They also develop in fluctuating
margins and intermittent shallow areas of more
permanent habitats. Areas included in this
35
Wisconsin Academy of Sciences, Arts and Letters
study fall into six categories: (1) marshes,
primarily open areas vegetated with cattails
(Typha) and/or sedges and rushes ( Carex ,
Eleocharis, Scirpus) and associated plants;
(2) Sphagnum bogs and swamps, usually with
leatherleaf ( Chamaedaphne calyculata ),
tamarack ( Larix laricina ), and/or black spruce
( Picea mariana), and with water often lim¬
ited to isolated pockets; (3) woodland pools
surrounded by coniferous and/or deciduous
trees, with a layer of leaf litter, and without
aquatic vegetation; (4) drainage ditches with¬
out associated aquatic vegetation; (5) grassland
pools surrounded by non-woody vegetation
and without aquatic vegetation; and
(6) temporary to semipermanent ponds with
aquatic vegetation.
Species of Aedes in Wisconsin can be di¬
vided into two groups, both of which over¬
winter as eggs. The first group has a single
generation each year, with eggs entering an
obligatory diapause that is terminated by ex¬
posure to several weeks of cold winter tem¬
peratures. In Wisconsin this group includes
A. abserratus, A. aurifer, A. communis, A.
decticus, A. diantaeus, A. euedes, A. excru¬
cians, A.fitchii, A.flavescens, A. grossbecki,
A. implicatus, A. intrudens, A. provocans,
A. punctor, A riparius, and A. stimulans.
Hatching of eggs is related to flooding from
snowmelt and rain and also to warming in
March and early April; it may be delayed in
heavily shaded areas and in habitats that re¬
main relatively cold.
The second group consists of species whose
eggs also hatch in the spring, but these spe¬
cies are capable of having additional broods
throughout the warm months of the year. Ad¬
ditional eggs hatch each time the habitat is
reflooded after it has become dry. This group
includes A. atropalpus, A. campestris, A.
canadensis, A. cinereus, A. dorsalis, A. hea¬
der soni, A. nigromaculis, A. spencerii, A.
sticticus, A. triseriatus, A. trivittatus, and A.
vexans. Some of these species may have only
one brood in some northern locations. Aedes
nigromaculis (Ludlow, 1907) and A. trivit¬
tatus (Coquillett, 1902) were not studied be¬
cause their larvae develop in late spring or
summer. Also not studied were A. header -
soni Cockerall, 1918 and A. triseriatus (Say,
1823), which breed in tree holes and artificial
containers, and A. atropalpus (Coquillett,
1902), which develops in rock pools.
While several studies of mosquitoes have
been carried out in Wisconsin, the relative
abundance, statewide distribution, and larval
habitats of species of spring Aedes have re¬
mained poorly known. Dickenson’s mono¬
graph (1944) provided the first account of
Wisconsin mosquitoes, listing thirty-eight
species. It was followed by Allen’s summary
(1950) of Wisconsin mosquito studies and
his preliminary survey of species in the Uni¬
versity of Wisconsin-Madison Arboretum.
He documented the only statewide survey of
adult mosquitoes, which was conducted in
twenty-three counties by the Wisconsin State
Board of Health in 1941; univoltine species
of Aedes were poorly represented in this sur¬
vey because many counties were sampled only
in mid or late summer. More recently, Siverly
and DeFoliart studied larvae (1968a) and adults
(1968b) in northeastern Wisconsin, signifi¬
cantly contributing to our knowledge of spring
Aedes. A study by Porter and Gojmerac (1970)
identified A. stimulans as the most important
pest in Point Beach State Forest, Manitowoc
County, and another study (Gojmerac and
Porter 1969) compared pest species of Point
Beach State Forest with those of Wyalusing
State Park, Grant County, where A. com¬
munis group species and A . vexans predom¬
inated. Amin and Hageman (1974) identified
A. stimulans and A. vexans as important
springtime pests in southeastern Wisconsin.
Other studies that included county records or
other information pertinent to this study were
carried out by Ryckman (1952), Patel (1959),
Thompson (1964), Thompson and Dicke
(1965), Thompson and DeFoliart (1966), Loor
and DeFoliart (1970), Wright and DeFoliart
(1970), Wright et al. (1970), Grimstad (1973),
and Kardatzke (1979).
The bionomics of mosquitoes, including
almost all species of Aedes, was summarized
by Carpenter and LaCasse (1955) for North
America and by Wood, Dang, and Ellis (1979)
36
Spring Aedes Mosquitoes in Wisconsin
for Canada. Several previous studies in Wis¬
consin also provided ecological information,
especially those by Siverly and DeFoliart
(1968a, 1968b). Additional information on
larval ecology in nearby states and provinces
appeared in Owen (1937), Barr (1958), and
Price (1963) for Minnesota; Knight and Wonio
(1969) for Iowa; Ross (1947) for Illinois;
Matheson (1924), Irwin (1942), Obrecht
(1949, 1967), Beadle (1963), and Wilmot,
Henderson, and Allen (1987) for Michigan;
Christensen and Harmston (1944) and Siverly
(1959) for Indiana; Venard and Mead (1953)
for Ohio; and Beckel and Atwood (1959) and
Steward and Me Wade (1960) for Ontario. In
our “Account of Wisconsin Species,” which
follows, these references are not cited unless
the information differs from our findings.
Methods and Materials
Study areas
Larval and adult Aedes populations were
surveyed in nine approximately 24-mile-square
areas defined by Billmyer (1971) in con¬
junction with a survey of Wisconsin stone-
flies (Fig. 1). These areas were selected as
representative of Wisconsin based on topog¬
raphy, geology, soil types, vegetation, and
climate. Study areas were located as follows:
Northern: North of T47N, R4-7W in Bayfield
and Ashland counties.
Northwestern: T37-40N, R15-18W in Burnett
and Polk counties.
Northeastern: T36 -39N, R15-18E in Flor¬
ence, Forest, and Marinette counties.
North central: T33-36N, R2-5E in Lincoln,
Oneida, Price, and Taylor counties.
West central: T23-26N, R1 1-14W in Buffalo,
Dunn, and Pepin counties.
East central: T15-18N, R19-22E in Calumet,
Fond du Lac, Manitowoc, and Sheboygan
counties.
Central: T16-19N, R7-10E in Adams, Mar¬
quette, and Waushara counties.
Southwestern: T9-12N, R1E-3W in Craw¬
ford, Richland, and Vernon counties.
Southeastern: T2-5N, R14-17E in Jefferson,
Rock, Walworth, and Waukesha counties.
Larval collections
Using a long-handled 350-ml dipper, Gilardi
collected mosquito larvae from twenty to thirty
sites in each study area on two dates between
4 April and 19 May 1988 (Gilardi 1990). The
first set of collections was made when most
larvae were early instars. The second set was
made just prior to, or coinciding with, the
first appearance of adults. Larvae were not
present in all localities when the first collec¬
tions were made, and many sites had dried
up before they were sampled again. Five dips
were taken from each site on each collection
date. Each dip was taken from a different
area within the site because numbers and spe¬
cies composition may vary with location
(Service 1976). Because larvae submerge
when disturbed, they were allowed one min¬
ute to return to the surface following a dis¬
turbance of the habitat (Hocking 1953) before
each sample was collected. Larvae were reared
to the fourth instar and pupae were reared to
adults to facilitate species identifications.
Ten sites in each area were selected for
additional larvae collections by Hilsenhoff in
1989 and 1990. Collections were made just
prior to first emergence in 1989 (18 April-
10 May) and somewhat after first emergence
in 1990 (25 April- 8 May), with ten dippers
of larvae or a maximum of fifty larvae being
collected from each site. Identifications of
larvae and adults were based on keys and
descriptions by Barr (1958) and Wood, Dang,
and Ellis (1979). Voucher specimens are in
the University of Wisconsin Insect Collection.
Adult collections
Gilardi (1990) also collected adult mos¬
quitoes with an aspirator during the daytime
from 6 June to 21 July 1988 as they at¬
tempted to feed. Although the propensity to
feed in daylight varies among species, ef¬
fective biting responses were obtained by
collecting in heavily vegetated areas where
mosquitoes rest during the day, and by dis¬
turbing vegetation before obtaining samples.
Two sets of collections were taken. The first
37
Wisconsin Academy of Sciences, Arts and Letters
Figure 1. Nine approximately 24-mile-square study areas selected to be representative of
Wisconsin (Billmyer 1971), with a number for each county.
set approximated peak adult populations. The
second was obtained five to six weeks later.
Collections were made for a ten-minute pe¬
riod at ten previously identified breeding sites
within each study area and also at locations
one to five miles from all known breeding
areas. Mosquitoes were collected from all
areas of the body that could be reached with
an aspirator; the head and back were pro¬
tected with a repellent.
Because female adults of several species
of Aedes cannot be readily identified after
more than a few days of flight, some were
often identified only to species group. Two
groups are typically recognized for species
in this region. The A. stimulans group in¬
cludes the following band-legged species: A.
euedes, A. excrucians, A. fitchii, A. flaves-
cens, A. riparius, and A. stimulans. The A.
communis group includes the following black¬
legged species: A. abserratus, A. cinereus,
A. communis, A. decticus, A. diantaeus, A.
implicatus, A. intrudens, A. provocans, A.
punctor, and A. sticticus.
38
Spring Aedes Mosquitoes in Wisconsin
Results and Discussion
Weather
Prior to the study period in 1988, precip¬
itation ranged from near normal to more than
40% below normal. An early thaw occurred
and was followed by cold temperatures, which
may have affected the larvae of some species.
Kardatzke (1979) determined that larvae of
A. ahserratus, A. communis, A. diantaeus,
A. provocans, and A. punctor may appear
during early thaws in northern Wisconsin and
Michigan but ^subsequently become vulner¬
able to refreezing. James (1962) documented
mortality of A. provocans larvae trapped in
ice following a thaw in Ontario.
The 1988 study period was characterized
by unusually warm temperatures and the on¬
set of a record drought. Small or shallow
breeding areas remained dry, and an unu¬
sually large proportion of habitats dried be¬
fore larval development was completed. The
spring of 1989 was very dry, and some sites
that had been sampled in 1988 contained no
water. Very warm temperatures and melting
snow in mid-March of 1990 caused an early
hatch of Aedes larvae, but unseasonably cold
temperatures followed, retarding larval de¬
velopment. The onset of very warm weather
during the last ten days of April accelerated
development and caused rapid pupation and
emergence of mosquitoes in most study areas.
As a result, larvae of early emerging species
were missed or underrepresented at some sites.
Except for the northeastern study area, which
remained very dry, water levels in 1990 were
similar to those in late April of 1988.
Collections
A total of 10,651 larvae and 1,612 adults
were collected in 1988, representing twenty
of the twenty-three species of spring Aedes
known to occur in snowmelt habitats in Wis¬
consin. Included were the first collections of
A. euedes within the state. The 1988 collec¬
tions are summarized in Tables 1 and 2. Lar¬
val collections from ten selected sites in each
area numbered 1,546 in 1989 and 1,600 in
1990, which compares to 1,658 (total ad¬
justed for differences in collecting proce¬
dures) from the same sites in 1988 (Table 3).
The 1989 samples included the first records
of A. decticus in Wisconsin.
Species distributions
Crossing the state diagonally is a region
of climatic and ecological transition that is
reflected in a tension zone of varying width
between two major floral regions, the North¬
ern Hardwood-conifer province and the Oak-
prairie Province (Curtis 1959). Aedes com¬
munis, A. decticus, A. diantaeus, A. euedes,
A. implicatus, and A. intrudens are boreal in
Wisconsin, and the southern limit of their
range apparently parallels this floral tension
zone. Ranges of A. aurifer, A. canadensis,
A. cinereus, A. dorsalis, A.fitchii, A.flaves-
cens, A. sticticus, A. stimulans, and A. vex-
ans encompass the entire state. Aedes ab-
serratus, A. campestris, A. excrucians, A.
provocans, A. punctor, A. riparius, and A.
spencerii are probably also present through¬
out Wisconsin, but may reach their southern
limit in southern Wisconsin or northern Il¬
linois. Aedes campestris, A. provocans, and
A. riparius have not been reported from Il¬
linois. Aedes grossbecki is represented in
Wisconsin by a single specimen from Dane
County, which probably represents the north¬
western limit of its range.
Geology and soil type also influence the
distribution of mosquitoes. Two sections in
the Central Lowlands Geomorphic Province
of the United States are represented in Wis¬
consin (Hole 1976), the Wisconsin Driftless
Section in the southwestern part of the state,
and the Great Lakes Section elsewhere. Mos¬
quito breeding was confined primarily to
floodplain marshes and pools in the Wiscon¬
sin Driftless Section (southwestern and west
central study areas), which limited somewhat
the diversity of species collected. Larval hab¬
itats were more varied and numerous in the
Great Lakes Section (all other study areas).
Paleozoic bedrock is present in nearly all areas
except a southern extension of the Cana¬
dian Shield into the northern third of the
state. Certain northern species of Aedes were
39
Table 1. Larvae and collection sites (in parentheses) for species of Aedes collected from nine 24-mile-square areas of Wisconsin in the
spring of 1988
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40
Table 2. Adults and collection sites (in parentheses) for species of Aedes collected at known breeding sites in nine 24-mile-square
areas of Wisconsin in 1988
Spring Aedes Mosquitoes in Wisconsin
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associated with the latter area (Wood, Dang,
and Ellis 1979).
Larval phenology
Based on larval collections and rearing,
the order of appearance of fourth instar larvae
approximated the following sequence: (1) A.
spencerii; (2) A. implicatus, A. intrudens, A.
provocans; (3) A. communis , A. diantaeus,
A. punctor, A. sticticus, A. stimulans; (4) A.
abserratus, A. aurifer, A. excrucians, A.
euedes, A. flavescens; (5) A. canadensis, A.
fitchii, A. riparius; (6) A. cinereus, A. vex-
ans. Larval development of most species was
synchronized, with nearly all members of a
given species entering the fourth instar within
a week or less in the same region. Exceptions
were A . cinereus and A . vexans, which often
had staggered emergence periods.
The central sandy uplands and plains, and
northern loamy uplands and plains, respond
quickly to seasonal warming (Hole 1976).
Larval development in these regions was
considerably advanced relative to develop¬
ment on other soils.
Adult dispersal
Aedes canadensis, A. cinereus, A. excru¬
cians, A. fitchii, A. provocans, A. stimulans,
and A. vexans were frequently collected within
two miles of known breeding sites. Collec¬
tions at greater distances did not occur for A .
excrucians and were infrequent for the other
six species. These observations are similar
to reports of Aedes flight ranges by Jenkins
and Hassett (1951), Neilsen (1957), Burst
(1980), Washino (1984), and Joslyn and Fish
(1986), although many workers have cited
much greater flight ranges for A. vexans dur¬
ing the summer. Horsfall et al. (1973) in¬
dicated that dispersal varies with environ¬
mental conditions and may be thwarted in A .
vexans populations that emerge in the spring
before evening temperatures are conducive
to flight.
41
Wisconsin Academy of Sciences, Arts and Letters
Table 3. Larvae of each species of Aedes collected from ten selected sites in each of nine 24-
mile-square study areas in 1988, 1989, and 1990
Continued on next page
42
Spring Aedes Mosquitoes in Wisconsin
Table 3— Continued
Adult longevity
Because drought prevented additional
broods of multivoltine spring Aedes in 1988,
this study provided an unusual opportunity
to identify persistent components of spring
populations. The second set of adult collec¬
tions yielded A. aurifer, A. campestris , A.
cinereus , A. stimulans group, and A. vexans .
Aedes stimulans group adults are frequently
reported to persist into August and occasion¬
ally September. Carpenter and Nielsen (1965)
reported a seventy two -day biting period for
A. campestris , and Horsfall et ah (1973) con¬
cluded that the typical lifespan of spring gen¬
eration A. vexans females is about seventy -
two days. At the opposite extreme, the nui¬
sance potential of A. provocans was offset
by its brief adult lifespan. Adult collections
of this species were primarily a measure of
how recently it had emerged in a given area.
Wood, Dang, and Ellis (1979) noted that adults
of this early species were seldom seen in
Canada after other spring Aedes had emerged.
Account of Wisconsin species
Aedes abserratus (Felt and Young, 1904)
= Aedes implacabilis of authors before 1954,
except Walker, 1848. County records (Fig. 1):
2, 6, 9, 11-13, 18, 32, 45, 48,* 49, 51, 52,
55, 63,* 64, 65,* 69. (Asterisks indicate
published records only.)
Aedes abserratus was fairly common state¬
wide. Larvae were almost always associated
with Sphagnum in swamps, bogs, shrubby
marshes, and woodland pools. Larvae de¬
veloped in essentially the same habitats oc¬
cupied by A. punctor, but A. abserratus lar¬
vae were more prevalent in sandy regions,
and A. punctor larvae were more prevalent
in and around Sphagnum bogs. Other re¬
searchers also noted the association of larvae
with Sphagnum among shrubs or trees.
Aedes aurifer (Coquillett, 1903). County
records (Fig. 1): 2, 13,* 17,* 18,* 35,* 43,*
46,* 51,* 52,* 57,* 61,* 63,* 70.
This statewide species is apparently rare
in early spring breeding sites in Wisconsin.
In 1988 a single larva was collected from a
large, open cattail pond in the southeastern
study area, and five adults were collected
near a small woodland lake in the northern
study area. Other studies in Wisconsin re¬
sulted in the collection of a limited number
of specimens. Breeding sites reported from
43
Wisconsin Academy of Sciences, Arts and Letters
nearby states and provinces include perma¬
nent and semipermanent bodies of water,
cranberry bogs, river-overflow areas, wood¬
land pools, and roadside habitats, with larvae
frequently being collected away from shore¬
line areas.
Aedes campestris Dyar and Knab, 1907.
County records (Fig. 1): 2, 52,* 63.*
Aedes campestris is apparently rare in
Wisconsin in the spring; it may occur more
commonly in the summer. A single adult was
collected in the northern study area from a
row of trees surrounded by open farmland.
This species was found during three surveys
in Dane County and was represented by only
three specimens in the State Board of Health
General Survey (Allen 1950). Elsewhere in
the United States and Canada, it was usually
found in open areas; larvae were reported to
develop in alkaline prairie pools, especially
those with a high organic content.
Aedes canadensis (Theobald, 1901).
County records (Fig. 1): 2, 5,* 6, 9, 11-13,
17,* 18, 22, 30, 34,* 35,* 37,* 45,* 46,*
48,* 49, 52, 55, 56 (unpublished), 57,* 58,*
63,* 64, 69, 70, 71,* 72.*
This species was fairly common statewide.
Larvae were most abundant in woodland
seepage pools in the south and boggy areas
in the north but also occurred in sedge-cattail
marshes. Many researchers in nearby states
to the south of Wisconsin noted that wood¬
land pools, especially those associated with
streams, are a preferred habitat. Others in
northern states and provinces noted an as¬
sociation with Sphagnum.
The limited number of adults that were
collected may be attributed to the wide range
of hosts that are attractive to this species.
Limited biting activity was also noted in Wis¬
consin by DeFoliart (1967), and Carpenter
and LaCasse (1955) observed that this spe¬
cies is seldom a pest in the eastern half of
its range, even when present in considerable
numbers. Several authors have noted that
adults often feed on turtles (Crans 1964; Hayes
1965; Nolan, Moussa, and Hayes 1965;
DeFoliart 1967; Crans and Rockel 1968).
Nevertheless, adults are known to readily at¬
tack humans, and they may be an important
pest in some areas of Wisconsin, most no¬
tably in woodland seepage areas.
Aedes cinereus Meigen, 1818. County
records (Fig. 1): 1,* 2, 4,* 5,* 6, 7,* 9,
11-14, 17,* 18, 19,* 22, 23,* 30, 32, 34,*
35-37,* 39,* 41,* 43,* 44, 45,* 48,* 49,
51, 52, 55, 57,* 58,* 61-63,* 64, 65, 66,*
69, 70, 71,* 72.*
Larvae were fairly common in a wide va¬
riety of habitats but were mostly found in
sedge and cattail marshes or in bogs. They
were especially common in the northern,
northwestern, and north central study areas.
Often several instars were present at the same
time, indicating a staggered emergence. The
wide variety of larval habitats was noted pre¬
viously in Wisconsin and by many workers
in nearby states and provinces.
Adults attacked readily throughout the day
in woodlands, where they were often en¬
countered in considerable numbers. This spe¬
cies was identified as a pest throughout Wis¬
consin in the State Board of Health General
Survey (Allen 1950); it was the second most
numerous species biting humans in Iowa
County (Loor and DeFoliart 1970).
Aedes communis (De Geer, 1776).
County records (Fig. 1): 2, 5,* 9, 11-13, 18,
35,* 48,* 63,* 72.*
This boreal species is an important pest of
the Canadian Shield. Larvae were collected
only in the northern, north central, and north¬
eastern study areas, where they were the pre¬
dominant or only species in certain sites. They
occurred only within soils of loamy uplands
and plains. Here they were found in vernal
ponds, mostly in woodlands and partially
shaded areas, and along margins of swamps
and leatherleaf bogs. Larvae were especially
common in 1989; they were much less com¬
mon in 1988 and 1990. Rapid drying of hab¬
itats in 1988 and possible emergence before
completion of sampling in 1990 may have
contributed to lower numbers of larvae in
these years. Siverly and DeFoliart reported
this to be the most numerous species in larval
collections from Forest County (1968a), and
the second most abundant spring mosquito
44
Spring Aedes Mosquitoes in Wisconsin
in adult collections from five northeastern
counties (1968b). Workers in nearby states
and provinces reported A. communis larvae
from habitats similar to those described above
and also noted that they often occur exclu¬
sively or nearly so in large numbers. Irwin
(1942) found that larvae were particularly
abundant in rapidly drying pools and shallow
habitats in central Michigan; Gjullin et al.
(1961) indicated that adults frequently emerged
just before larval habitats in Alaska had dried.
Aedes decticus Howard, Dyar, and
Knab, 1917. County records (Fig, 1): 12, 18.
In 1989 six larvae were collected in north¬
eastern Wisconsin from two sites that con¬
tained Sphagnum. They represent the first
records of this species for the state. Four
larvae were found in an open leatherleaf bog;
the other two were collected from a spruce-
tamarack swamp. In the western Great Lakes
region other studies associated larvae of this
relatively rare species with Sphagnum.
Aedes diantaeus Howard, Dyar, and
Knab, 1917. County records (Fig. 1): 2, 8,*
9, 11,* 12, 13, 18.
This boreal species occurred uncommonly
in northern Wisconsin. Larvae were collected
from margins of swamps or from pools in
alder ( Alnus ) thickets and were usually as¬
sociated with Sphagnum and alders and al¬
most always with larvae of A. punctor. Siverly
and DeFoliart (1968a) identified a productive
breeding site in Forest County where larvae
occurred in stump holes “formed by cedar
and hemlock windthrow.” They also ob¬
tained modest numbers of adults from five
northeastern counties (1968b). Prior to their
study this species was represented in Wis¬
consin by a single adult from Sawyer County
(Smith 1952). Other studies in our region
associated larvae with Sphagnum pools and
alders or woodland pools.
Aedes dorsalis (Meigen, 1830). County
records (Fig. 1): 19,* 63,* 64,* 72.*
Aedes dorsalis adults may emerge later than
those of most species of spring Aedes in Wis¬
consin, where they have usually been col¬
lected in small numbers, mostly during the
summer. No specimens were collected in this
study. In other states and provinces larvae of
this species were reported to occur in alkaline
and saline pools and also in ponds that are
rich in organic matter. Outside of the prairie
region larvae were usually associated with
industrial wastes.
Aedes euedes Howard, Dyar, and
Knab, 1917 = Aedes barri Rueger, 1958.
County records (Fig. 1): 11, 13, 49, 51, 52.
Aedes euedes larvae were found only in
the northeastern, central, and east central study
areas, where they were uncommon. Seven
were collected in 1988 and seven more in
1990. These are the only records of this spe¬
cies from Wisconsin. Larvae were collected
in association with cattail marshes containing
scattered ash trees ( Fraxinus ) or from rather
open pools adjacent to woodlands. Wood,
Dang, and Ellis (1979) associated larvae with
large open marshes having dense accumu¬
lations of decomposing sedges and cattails.
Wilmot, Henderson, and Allen (1987) re¬
ported larvae from several woodland pools
in Michigan, while Price (1963) found larvae
in nearly all habitats he sampled in northern
Minnesota.
Aedes excrucians (Walker, 1856).
County records (Fig. 1): 2, 4,* 6, 7,* 9,
11-14, 18, 28,* 30, 39,* 41,* 42,* 44, 45,
48,* 49, 51, 52, 55, 57,* 58,* 63,* 64, 65,
66,* 68,* 69, 70, 71,* 72.*
Aedes excrucians larvae were common in
all study areas and occurred in more different
sites and habitats than larvae of any other
species. They were almost always found in
association with larvae of other species and
usually were not the dominant species. The
relatively ubiquitous larvae were most com¬
mon in marshes and margins of swamps. It
was the most abundant larva that Siverly and
DeFoliart (1968a) collected in Lincoln County,
where it occurred in unshaded grassy pools
and ditches. In nearby states and provinces
all researchers reported larvae from a variety
of habitats.
Aedes fitchii (Felt and Young, 1904).
County records (Fig. 1): 2, 4,* 5,* 6, 7,* 9,
11-14, 18, 19,* 22,* 28,* 30, 35,* 37-39,*
43,* 44, 45, 48,* 49, 51, 52, 55, 57,* 58,*
45
Wisconsin Academy of Sciences, Arts and Letters
63,* 64, 65, 66,* 69, 70, 71.*
Larvae of A. fitchii were fairly common
statewide. They were almost always asso¬
ciated with larvae of A . excrucians and/or A .
stimulans and were never the dominant spe¬
cies in collections. Larvae were most com¬
mon in marshes but also occurred in drainage
ditches, woodland pools, and margins of
swamps and bogs. Siverly and DeFoliart
(1968a) reported that A. fitchii larvae oc¬
curred in a wider range of habitats than A.
excrucians in northeastern Wisconsin. Other
researchers in our region found that larvae
inhabited mostly marshes and rarely were as¬
sociated with Sphagnum.
Although A. excrucians outnumbered A.
fitchii in larval collections, the opposite was
true in adult collections. This may be par¬
tially attributed to the greater propensity of
A. fitchii to invade woodlands, where biting
counts were made. In addition, since adults
of A. fitchii emerge later than those of A.
excrucians, they would have experienced less
mortality prior to the collection of adults in
June and July.
Aedes flavescens (Muller, 1764). County
records (Fig. 1): 2,* 5,* 32, 43,* 49, 60,*
63,* 65,* 69, 72.*
This species was rare in our study. It was
reported to be sporadic over most of its range
and common only in prairies (Wood, Dang,
and Ellis 1979). Three larvae were collected
in the southern half of the state, one from a
woodland pool, another from a cattail pond,
and the third from a glacial kettle. Siverly
and DeFoliart (1968b) collected a single adult
from northeastern Wisconsin. In other areas
of the western Great Lakes region larvae were
associated with grassland pools and marshes,
cattail ponds, and by Irwin (1942) with
woodland pools.
Aedes grossbecki Dyar and Knab, 1906.
County record (Fig. 1): 63.*
Aedes grossbecki is represented in Wis¬
consin by a single adult from the University
of Wisconsin-Madison Arboretum (Thomp¬
son and DeFoliart 1966). This southern
woodland species was reported to be com¬
mon in southern Illinois and rare northward
(Ross 1947; Ross and Horsfall 1965).
Aedes implicatus Vockeroth, 1954 =
Aedes impiger of authors before 1954, except
Walker, 1848. County records (Fig. 1): 2,
9 (unpublished), 11,* 48.*
Aedes implicatus is apparently a rare bo¬
real species in Wisconsin. In the northern
study area a single adult was reared from a
pupa that was collected from a marsh that
contained willows and was next to a stream.
Siverly and DeFoliart, who first reported this
species in Wisconsin, obtained a single larva
from a coniferous woodland stump hole in
Forest County and one adult nearby (1968a);
they also collected one adult from an un¬
specified location in northeastern Wisconsin
(1968b). Porter and Gojmerac (1970) re¬
ported small numbers of adults emerging from
woodland pools in Point Beach State Forest,
Manitowoc County. In Colorado, Smith (1965)
found larvae in “small shallow pools left by
receding streams and shaded by willow thick¬
ets”; Wood, Dang, and Ellis (1979) obtained
large numbers of larvae from a similar habitat
in Ontario. Other workers found larvae mostly
in temporary woodland habitats and
Sphagnum bogs.
Aedes intrudens Dyar, 1919. County
records (Fig. 1): 2, 5,* 6, 9 (unpublished),
11-13, 18, 28,* 30, 44, 48,* 49, 63,* 72.*
Larvae were rare, but those that were col¬
lected were found in all types of habitats
except ditches. The varied larval habitat was
also noted by other researchers in our region,
with Sphagnum bogs and woodland pools the
most frequently mentioned habitats.
Aedes provocans (Walker, 1848) =
Aedes trichurus (Dyar, 1904). County rec¬
ords (Fig. 1): 2, 6, 9, 11-14, 18, 30, 32, 44,
45, 49, 51, 52, 55, 64, 70.
Aedes provocans larvae were very com¬
mon in all study areas. Although they were
collected most frequently from marshes and
woodland pools, larvae were rather common
in all types of habitats that were sampled,
especially open, temporary sites such as grassy
ditches and grassland pools. In northeastern
46
Spring Aedes Mosquitoes in Wisconsin
Wisconsin, Siverly and DeFoliart (1968a) as¬
sociated larvae with grassy pools. The varied
nature of the larval habitat was also reported
in studies in nearby states and provinces.
Aedes punctor (Kirby , 1837). County
records (Fig. 1): 2, 5,* 6, 9, 11-13, 18, 30,
35, 45,* 48,* 49, 52, 55, 57,* 65,* 69.
Larvae of A. punctor , an important pest
of the boreal forest, were common in the
north central and northeastern study areas and
uncommon elsewhere. They were the most
numerous larvae in the northeastern study
area, and also in the study by Siverly and
DeFoliart (1968a) in northeastern Wisconsin.
Among species we commonly collected, A.
abserratus and A. punctor were most often
coincident in larval habitats; A. punctor lar¬
vae appeared in more than three-fourths of
the sites known to be inhabited by larvae of
A. abserratus . During a seven-year study in
Minnesota, Price (1963) observed that A.
punctor larvae occurred in all habitats that
had yielded A. abserratus larvae. While lar¬
vae were collected from a variety of habitats,
they most frequently occurred in Sphagnum
bogs and, to a lesser extent, in woodland
pools with Sphagnum, Most researchers in
nearby states and provinces mentioned that
larvae were associated with Sphagnum and
shrubs or trees. The exception was Steward
and Me Wade (1960), who found larvae in
Ontario in virtually all types of standing water,
most commonly in woodland pools.
Aedes riparius Dyar and Knab, 1907.
County records (Fig. 1): 6, 9, 12, 28,* 43,*
45, 49, 51,* 63,* 64, 65.*
Aedes riparius is apparently rare in Wis¬
consin. Only four specimens were found in
the State Board of Health survey (Allen 1950).
Larvae were collected in small numbers from
several habitats but most frequently were found
in marshes. Researchers in nearby states and
provinces also collected larvae mostly from
marshes that frequently contained some scat¬
tered trees or shrubs, or had trees along their
margins.
Aedes spencerii (Theobald , 1901).
County records (Fig. 1): 2, 5,* 6, 9, 12, 13,*
32,* 51,* 52,* 58,* 63,* 69.*
Aedes spencerii is apparently uncommon
in Wisconsin. A few pupae were collected
in northern areas during the first set of larval
collections in 1988; in 1989 and 1990 adults
had probably emerged before larval collec¬
tions were made. All pupae were collected
from open habitats, including grassland pools,
a sedge marsh, a Sphagnum bog, and a small
pond. In adjacent states larvae were reported
from grassland pools, marshes, and bogs.
Wood, Dang, and Ellis (1979) indicated that
this species has been underrepresented or
overlooked in several studies because of its
early appearance, noting that pupae were
present when larvae of other spring Aedes
were about half grown.
Aedes sticticus (Meigen, 1838). County
records (Fig. 1): 2, 6, 11, 18, 30, 32, 35,*
42,* 48,* 49, 52, 55, 67.*
Larvae develop throughout the state pri¬
marily in flood water pools along streams. Al¬
though rare in this study because of the drought
in 1988, A. sticticus larvae may become ex¬
ceptionally numerous near streams after un¬
usually heavy snowmelt and/or rain. It was
identified as a pest in Manitowoc County
(Porter and Gojmerac 1970). Adults were
collected in considerable numbers in Wood
County (Wright et al. 1970) and in five north¬
eastern counties (Siverly and DeFoliart 1968b).
Cook, Bodine, and Wermerskirchen (1974)
studied the biology of this species in the Twin
Cities area just west of Wisconsin. Eggs were
laid along the periphery of flooded areas and
accumulated in river floodplains and bottom¬
lands during intervals between extensive
floods. They remained viable for several years
under drought conditions and hatched after
flooding. In Canada, Wood, Dang, and Ellis
(1979) observed that A. sticticus populations
are almost always associated with A . vexans,
but that the converse is seldom true “because
A . vexans develops in summer rainpools after
local flooding, whereas A. sticticus requires
extensive flooding, which only follows wide¬
spread excessive precipitation.”
Aedes stimukms (Walker , 1848). County
47
Wisconsin Academy of Sciences, Arts and Letters
records (Fig. 1): 2, 4,* 5,* 6, 11-14, 18,
19,* 22, 28,* 30, 32, 37,* 39,* 44, 45, 47,
48,* 49, 51-53, 55, 57-59,* 61,* 63,* 64,
65, 66,* 69, 70, 72.*
Larvae of A. stimulans, the most numerous
species in this study, were found in a variety
of habitats but were collected most frequently
and in largest numbers from marshes, tem¬
porary ponds, and woodland seepage pools,
including river valley sites. Some ditches and
grassy pools near marshes also harbored large
populations. In other studies larvae were most
often associated with woodland pools. In
Minnesota, however, Owen (1937) men¬
tioned grassland pools as a preferred larval
habitat, while Price (1963) observed that they
prefer marshes but often occur elsewhere.
Adults were the most important nuisance
in central and southern Wisconsin in May
and early June, but they were uncommon in
much of the north. Siverly and DeFoliart
(1968a) noted a virtual absence of this spe¬
cies in northeastern counties. A similar pat¬
tern is evident in other studies in this region.
Wood, Dang, and Ellis (1979) associated large
populations south of the Ottawa area with
paleozoic sediments, and the general scarcity
northward with the Precambrian Shield, im¬
plicating acidity as a possible limiting factor.
The largest collection of larvae from the four
northern study areas in 1988 (outnumbering
all other northern collections combined) was
taken from a site in the northwestern study
area that is not on the Precambrian Shield.
Aedes vexans (Meigen, 1830). County
records (Fig. 1): 2, 5,* 6, 9, 11, 13, 14, 17,*
18, 22, 29,* 30, 32,* 34,* 35,* 37-39,*
41,* 42,* 44, 46,* 47, 48,* 49, 50,* 51, 52,
53,* 54,* 55, 56-58,* 61-63,* 64, 65-68,*
69, 70, 71,* 72.*
Aedes vexans, the most important pest
mosquito during the summer and fall in most
areas of Wisconsin, was a variable compo¬
nent of spring populations. It was the second
most abundant species in larval collections
in 1988, larvae were absent in 1989, and
larvae were numerous only in the west central
study area in 1990. This was probably the
result of more extensive flooding of habitats
by early spring rains in 1988. Adults, how¬
ever, were found in small numbers in 1988
because many habitats dried before larval de¬
velopment was completed. Larvae were col¬
lected from all types of habitats, but more
than 85% were found in marshes and grass¬
land pools, and only about 1% were from
woodland pools and Sphagnum bogs. Shal¬
low, open, grassy depressions were identified
as primary breeding areas by workers in nearby
states and provinces. However, Horsfall et
al. (1973) and Wood, Dang, and Ellis (1979)
observed that woodland habitats can also har¬
bor large populations.
Larvae were collected most frequently and
in largest numbers within areas containing
sandy or loamy soils, which corroborates
findings in other states (Horsfall et al. 1973).
Siverly and DeFoliart collected substantial
numbers of larvae (1968a) and adults (1968b)
in parts of a five-county area in northeastern
Wisconsin that were dominated by such soils.
Horsfall et al. (1973) indicated that this spe¬
cies is very local or absent in northern Mich¬
igan wherever black-legged species such as
A. communis and A. punctor are abundant.
A predominance of A. communis group spe¬
cies and scarcity of A. vexans were evident
in most larval collections by Siverly and
DeFoliart (1968a).
Possible additional species
Aedes pionips Dyar, 1919. This uni-
voltine northern species was reported from
Itasca State Park, Minnesota (Barr 1958; Price
1963), Isle Royale, Michigan (Cassani and
Newson 1980); and Ontario (Wood, Dang,
and Ellis 1979). It was reported to be com¬
mon in the boreal forest region and rare or
local southward.
Aedes pullatus (Coquillett, 1904). Ear¬
lier reports of A . pullatus from Michigan by
Irwin (1942) were questioned by Barr (1958)
and Wood, Dang, and Ellis (1979), and be¬
cause of their distance from established rec¬
ords and an apparent lack of more recent
material, they were discounted by Darsie and
Ward (1981). However, based on unpub¬
lished data of Wagner and Newson from 1971,
48
Spring Aedes Mosquitoes in Wisconsin
Cassani and Newson (1980) reported A. pul-
latus from six counties in the northern half
of Michigan. Other records show an unusual
disjunct distribution of this univoltine species
in northwestern and northeastern North
America, which may be a result of glacial
history (Wood, Dang, and Ellis 1979).
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G. R. DeFoliart, G. Seawright, and R. P.
Hanson, 1970. Isolations of LaCrosse Virus of
the California group from Tabanidae in Wis¬
consin. Mosq. News. 30:600-603.
50
Symbolism in the Cave of Montesinos
James T. Abraham
The episode of the Cave of Montesinos
in Don Quixote has become one of the
most analyzed and interpreted incidents in
modem literary history. Most critics agree
that it is the crucial moment in the work
because, as Gethin Hughes states, “We wit¬
ness the confrontation of two worlds in Don
Quixote’s mind — the chivalric and the real”
(112). The novel takes a different direction
after Don Quixote resurfaces and tells the
story of his “grande ad ventura de la cueva
de Montesinos.” This is the only adventure
that Don Quixote faces alone, so it gives us
the best opportunity to study his psycholog¬
ical state. By analyzing the dream, we can
better understand Don Quixote’s madness.
This paper discusses the symbols and their
meanings in the dream and the significance
of the dream itself.
Sigmund Freud wrote in his book, The
Interpretation of Dreams, that all dreams have
meaning. The meaning of a dream can be
interpreted through the symbols that appear
in the dream itself. The incident in the Cave
of Montesinos is rich in symbolization. In
the dreamwork Don Quixote descends into a
legendary cave and encounters a beautiful
landscape and crystal palace. He meets sev¬
eral famous characters, all demonstrating bi¬
zarre behavior. Upon seeing his enchanted
mistress, the knight is elated but confused by
her actions. Finally Don Quixote ascends back
James T. Abraham is a doctoral candidate at the Uni¬
versity of Arizona in Tucson, specializing in Golden Age
theater and literary theory. He has recently received his
M.A. degree in Foreign Language and Literature at
UW -Milwaukee . He has lived and studied in Spain,
Mexico, and Brazil.
into the “real” world only to find his friends
criticizing the cave, the adventure, and Don
Quixote himself. Analyzing dreams and their
significance is part of the psychoanalytic pro¬
cess developed by Freud around the turn of
the century. Psychologists today still use the
guidelines set down by Freud to analyze
dreams. The ideas presented in this paper
suggest possible explanations for the dream
and even Quixote’s madness by using psy¬
choanalytical theories.
The first symbol we encounter is the cave
itself. Don Quixote is familiar with the leg¬
end surrounding the cave and insists on stop¬
ping at it on his way to Barcelona. He wants
to descend into the cavern and see “si eran
verdaderas las maravillas que de ella se de-
cfan por todos aquellos contomos (if they
were true, the wonders that were spoken of
the cave in those parts)” (Cervantes, Don
Quixote, 435). Before entering the cave, Don
Quixote must chop his way through thick
brush to find the entrance and fend off bats,
owls, and other nocturnal birds. Quixote has
intense drive and needs to experience what
the cave holds. He is not afraid of his destiny
and is, in this case, actively pursuing his
future.
The cave itself is “a maternal symbol that
excites curiosity” (Becker, 149). It is a posi¬
tive symbol because caves were often used
as oracles. In the pastoral novel of Spain, the
cave was the entrance to the underworld. Ac¬
cording to Frederik de Armas, caverns were
“the source of power of magicians, wisdom
of prophets and inspiration of poets” (Ar¬
mas, 337). They were used to communicate
with the dead. Carl Jung believed the cave
represents the unconscious. Cervantes did not
use caves in his pastoral novels because of
51
Wisconsin Academy of Sciences, Arts and Letters
their demonic connotations but reversed his
fear in Don Quixote and La cueva de Sala¬
manca.1 The use of a cave by Cervantes sig¬
nals something very important. In this case,
the cave symbolizes a mystical realm in the
unconscious mind of Don Quixote where he
can come in contact with the souls of his
fallen brethren and chivalric heroes from an
age regrettably now past.
This magical place where knights live is
known to Alonso Quijano only through the
books of chivalry that have driven him mad.
The old man’s love and belief in the Age of
Chivalry, yet total lack of experience in it,
mirror in reverse his adventures. In the nov¬
els of chivalry, the aspiring knight learned
of rules and codes of conduct to be followed
by all in order to establish order amidst chaos.
But El Caballero de la Triste Figura, upon
putting on armor and accepting the charge of
knighthood, has come to learn that being a
knight is not an easy endeavor. He is battered
and trampled, disgraced and dishonored.
Needing rest and a return to the code of chiv¬
alry, Quixote sets out on a pilgrimage to the
cave of Montesinos. But the cave is much
more than just another place to explore. He
seeks in the maternal symbol a return to his
novels, to security, to the womb. This is
evident when Don Quixote resurfaces and
begins to tell his story. Sancho, solidly fixed
in the “real” world of the novel, calls the
cave a pit. Don Quixote becomes enraged
and demands that Sancho and the scholar not
call it a pit. With great emotion, he chastises
them saying, “Dios os lo perdone, amigos;
que me habeis quitado de la mas sabrosa y
agradable vida y vista que ningun humano
ha visto ni pasado (God forgive you friends
because you have taken me from the most
pleasurable life that any human has seen or
experienced)” (Cervantes, Don Quixote, 438).
On the surface Quixote may be defending the
honor of his mother since the cave is a ma¬
ternal symbol. However, I believe he is really
defending his entire chivalric world and the
fact that it exists. No part of his “world”
may be put into question because it is founded
on such unsteady ground that it could easily
be toppled.
Deeper into the symbolism, we can look
at the bell that Don Quixote forgets to put
on. The bell is a symbol of reality and his
link to the world outside the cave. It is proof
that he is moving in the flesh and blood world
of reality. If he had been wearing the bell,
Sancho and the scholar would surely have
yanked him back into reality when it stopped
making a sound, thus ending any possible
adventure. Without it, however, the knight
is free to roam in the unknown and experi¬
ence all the “maravillas” of the cave. He is
free to step out of reality and explore the
world of fantasy.
Another symbol that is part of the old
knight’s entrance into the cavern is the rope
used to lower him. Becker states that a rope
represents the sexual act (84). The smooth
walls of the cave represent erect bodies ac¬
cording to Freud (109). Therefore, Don
Quixote’s descent by means of a rope, past
the smooth walls of the cave, symbolizes
what Don Quixote knows little about—sex.
The effect is to emphasize the fact that Don
Quixote is breaking new ground with respect
to the world of his heroes and his own
sexuality.
Once inside the cave, Don Quixote sees a
beautiful place that only nature could create.
He is not sure whether he is asleep or awake
but soon convinces himself that he has all his
faculties. Could the knight be in paradise? I
believe so, at least the paradise that the knight
Don Quixote de la Mancha has envisioned.
The beautiful landscape emphasizes the idyl¬
lic nature of the dream. In this ideal land,
Quixote spies a crystal castle. Crystal rep¬
resents the self (Becker, 154), and here Don
Quixote is looking into a mirror. He sees
himself as a majestic, strong, and royal per¬
sonage equal to his ideals. He has created in
his mind the perfect reincarnation of himself
and his beliefs.
Soon after creating his heaven, Quixote
meets its first inhabitant, the old Montesinos.
The knight describes Montesinos as “vestido
52
Symbolism in the Cave of Monte sinos
con un capuz de bayeta morada, que por el
suelo le arrastraba, cernale la cabeza una gorra
milanesa negra, y la barb a, camsima, le pas-
aba de la cintura (He was clad in a long
mourning cloak of purple baize, which trailed
upon the ground; over his shoulders and breast
he wore a kind of collegiate tippet of green
satin, his hoary beard reaching below his gir¬
dle)” (Cervantes, Don Quixote, 439). Is this
the image of God in Don Quixote’s mind?
Not exactly. Becker says kings or queens
represent parents (85); Montesinos represents
Don Quixote’s father. Quixote further proves
this when he says, “El continente, el paso,
la gravedad y la anchfsima presencia, cada
cosa de por si y todas juntas, me suspendi-
eron y admiraron (His mien, his gait, his
gravity, and his goodly presence each singly
and conjointly filled me with surprise and
admiration)” (Cervantes, Don Quixote, 439).
Just as a child respects his parents, Don
Quixote looks up to Montesinos and respects
him.
Montesinos has been analyzed by many
critics, and all have found different things
about him strange. He is a figure from medi¬
eval Spanish ballads in the region of La Man¬
cha. Carrol B. Johnson, in his book, Mad¬
ness and Lust, remarks that although
Montesinos is dressed in a scholarly way, he
does not know all the answers. He does not
know how to disenchant all the people in the
cave or whether he used a “daga” (dagger)
or a “punal buido” (dirk) to take out his
friend Durandarte’s heart. Even more star¬
tling to Quixote is the fact that Montesinos
is not familiar with the beautiful Dulcinea
(163). Johnson interprets these uncertainties
as Don Quixote’s own or, much more likely,
as those of his father (163). Just as a rebel¬
lious teenager believes that his parents do not
know what is best, Don Quixote questions
Montesinos.
A theme discussed by E. C. Riley is the
absurdity of Montesinos. The picture given
us of Montesinos does not fit the image of a
great knight. He is holding a rosary with
beads the size of chestnuts and ostrich eggs.
He does not evoke awe or fear as a great
knight of his time would but rather appears
ridiculous. Further highlighting the absurdity
of Montesinos is the fact that his best friend,
Durandarte, was to have had a heart that
weighed two pounds.2 Riley writes, “These
ridiculous details puncture the fabric of his
(Don Quixote’s) chivalric vision” (142). He
believes these elements are meant to mock
Don Quixote and his principles (142). They
emphasize the ridiculous nature of the dream
and the old knight himself.
Soon after meeting Montesinos, Don
Quixote asks the old man if the legend sur¬
rounding the removal of his best friend’s heart
is true. Montesinos answers “yes,” and the
fact that there is a question about the dagger
leaves us to wonder about its significance.
The knife is a masculine symbol. The ap¬
pearance of a masculine element within the
womb causes much fear in Don Quixote.
Johnson believes the dagger symbolizes
Quixote’s fear of castration and the castrating
female (167). This symbol is a manifestation
of his inability to interact with women and
probably stems from an unresolved problem
in the Oedipal stage of his childhood (John¬
son, 167).
Next, Don Quixote meets the zombie Du¬
randarte. Named for Roland’s sword, he is
Don Quixote’s image of the ideal knight.
Johnson believes Durandarte is identifiable
to Don Quixote because they are both knights
and both have hairy, bony hands that show
great strength. Don Quixote identifies with
him but is afraid when he realizes that Duran¬
darte is no longer a powerful knight. Because
Durandarte and a sword are so closely re¬
lated, Johnson associates Durandarte to the
phallus through symbolization (164). The fact
that Durandarte is a “sword-phallus rendered
useless by bloody mutilation” (Johnson, 164)
points to impotency. Because Durandarte and
Quixote are essentially the same, Duran¬
darte’s impotency points to fears of impo¬
tency in Don Quixote. Johnson goes as far
as to say that this element of impotency
“bring[s] together some of the most pervasive
53
Wisconsin Academy of Sciences, Arts and Letters
themes of Don Quixote’s psychic life, with
some of the most deep-seated fears about
himself and his manhood” (164).
The first female character that the knight
encounters is the noble woman, Belerma.
Quixote describes Belerma as clad in black,
with a slightly up-turned nose and a large
mouth with colored lips. Johnson sees three
different themes in the character of Belerma.
First, he believes that she represents all the
older women in Quixote’s life — his mother,
his grandmother, and others (165). The al¬
lusions to the age and sensuality of Belerma
are signs of an Oedipal attraction in the knight’s
past. Next, there is a relationship between
Belerma and Dulcinea since Belerma is to
Durandarte what Dulcinea is to Don Quixote,
mainly the object of courtly love (164). Be¬
lerma, according to Johnson, represents the
reason for his dysfunction, something from
his childhood that has forced him to create
Dulcinea (167). Finally, because they both
have bad teeth and are sexually inoperative,3
Don Quixote and Belerma are identifiable as
one (Johnson, 168). Belerma was a legendary
beauty, but when Don Quixote sees her he
is disappointed and disillusioned. Hughes sees
the symbolism and applies it to Dulcinea. She
believes that it means if Belerma can be made
ugly, through enchantment, so too can his
beautiful Dulcinea (110). It is important to
remember that Don Quixote’s picture of the
enchanted Dulcinea is the ugly maid Sancho
pointed out to him. The image of ugliness
through enchantment bolsters Quixote’s be¬
lief in the existence of an enchanted world
and its need for his help.
Separately, each of these people has major
significance. Is there any significance to the
three being together? Johnson believes there
is. He states, “All three of the chivalric char¬
acters are projections of different aspects of
our hero himself” (167). This idea fits with
Freud’s theory of condensation; that is, many
unrelated elements may come together in a
dream. All share nearly the same age and the
fact that their lives are at a standstill (John¬
son, 167). The three inhabitants of the cave
are sentenced to live forever in legend, while
Don Quixote, although still part of the “real”
world, takes time out from the continuing
action above ground to join them in fantasy
below ground. By joining the three characters
into one, we complete the psychic picture of
the bent knight. Montesinos, according to
Johnson, projects a number of intellectual
insecurities. Durandarte projects Quixote’s
fear of castration and impotence, and Be¬
lerma reflects his fear of aging (167).
Throughout the dream, Quixote is analyzing
himself and struggling with questions that run
deep into his psyche.
Finally, Don Quixote comes face to face
with the “incarnation of his chivalric world”
(Hughes, 109), Dulcinea. She is with two
other damsels and runs away at the sight of
the great knight. One of the damsels soon
comes back and asks if Don Quixote might
lend Dulcinea six reales. He has only four,
but gives them to her anyway. Johnson be¬
lieves the money represents Dulcinea’ s sex¬
ual needs and Quixote’s prowess (158). The
fact that he is unable to give her the total of
six reales once again symbolizes his fears of
impotency and capability of loving his mis¬
tress. Hughes believes the monetary aspect
destroys Don Quixote and his chivalric world
(112). The money is not part of the chivalric
code and thus proves that this world cannot
and does not exist. It is this event that later
(on his deathbed) permits Don Quixote to
accept Sanson Carasco and the priest. Dul¬
cinea’ s simple request for money shows him
that his ideals are fantasy, and he cannot
survive in the current age of realism.
By looking at the entire episode of the
Cave of Montesinos, we can get a good look
at Don Quixote’s psychological state. Per¬
haps the best picture comes not from a
psychoanalyst, but from one of Spain’s great
authors, Miguel de Unamuno. Donald Pal¬
mer, in his article entitled, “Unamuno, Freud
and the Case of Alonso Quijano,” points to
Unamuno’s book Vida de Don Quijote y
Sancho4 as the first psychoanalytical account
of Don Quixote (243). In fact, Unamuno and
Freud were contemporaries, and Unamuno’s
book was published while Freud was doing
54
Symbolism in the Cave of Montesinos
his major work on psychoanalysis. In the
book, Unamuno discusses sublimation. Freud
defines sublimation as “the sexual trend
abandoning its aim of obtaining a component
of reproductive pleasure and taking on an¬
other which is related genetically to the aban¬
doned one but is itself no longer sexual”
(Brill, 179). Palmer reports that Unamuno
believed Quixote repressed his amorous feel¬
ings for his teenage maid and, after letting
the repression boil for twelve years, finally
went mad. Claiming that aspects of higher
culture come from sublimations of repressed
instinctual drives (Brill, 215), Freud does not
condemn this sublimation as evil but calls it
“a triumph of spirituality over the senses”
(Brill, 217). Quixote’s quest for the pure,
the noble, and the chivalric is the outcome
of his sublimation of amorous feelings. His
need to contact past heroes is just another
manifestation of his repressed desires. Al¬
though hiding a dark secret, he retains his
honor and dignity (at least in his own eyes).
Johnson agrees with this idea, stating that
Don Quixote has repressed his feelings for
his niece from ‘ ‘just below conscious to deep
unconscious level” (156).
Another plausible explanation for the
dreamwork in Don Quixote’s dream is the
theory of wish fulfillment. Freud writes: “A
dream is a (disguised) fulfillment of a (sup¬
pressed or repressed) wish” (Brill, 57).
Quixote has been struggling with the en¬
chantment of his damsel since Book I, Chap¬
ter 10, when Sancho invented her. Hughes
believes the dream allows him to solve the
problem of Dulcinea’s enchantment (108)
through wish fulfillment. Don Quixote’s wish
for a land where the laws of chivalry are
upheld and adventure involving his damsel
is obvious in the dream. He identifies with
all the people in the dream world, and his
supreme chivalric act would be to disenchant
all its inhabitants. This theory rationalizes the
dream as merely an escape into fantasy land
for the gallant knight, thus having no psy¬
chological value other than to manifest his
aspirations.
The final explanation for this episode is
that it is a look into the psyche of Cervantes
himself. Becker states, “The work of art and,
even more, dreams in works of art have been
considered as confessions of the artist’s un¬
conscious personality, his affective conflicts
and especially his sexual complexes” (103).
He outlines how dreams may be used in lit¬
erature. First, the author may use the dream
explicitly to further the main theme in the
work. Second, he may use the dream im¬
plicitly as an invisible support system for the
structure of the work. Applying this theory
to Don Quixote, we might say Cervantes uses
Don Quixote’s lunacy explicitly to satirize
the Chivalric Age. The dream, once again,
reinforces the madness of the old knight and
the absurdity of the Chivalric novel. Implic¬
itly, however, the dream creates a picture of
the reasons for Don Quixote’s madness. It
subtly shows us that Don Quixote is not just
mad but that there are concrete reasons for
his condition. He has suppressed his amorous
desires for all the women he has ever known
and now must deal with all the repercussions.
He struggles relentlessly against tremendous
obstacles. It is sad that Don Quixote will
never know love, but it is noble that he will
fight until death to keep the hope for it alive.
Endnotes
‘Miguel de Cervantes, “La cueva de Sala¬
manca,” in Entremeses de Miguel de Cervantes
Saavedra, ed. Adolfo Bonilla y San Martin (Mad¬
rid: Asociacion de la Librerfa de Espana, 1963).
2The legend of the day stated that the size of a
man’s heart is directly proportional to his bravery.
Belerma was postmenopausal and Don Quixote
impotent.
4Miguel de Unamuno, Vida de Don Quixote y
Sancho Segun Miguel de Cervantes Saavedra, Ex-
plicada y Comentada por Miguel de Unamuno
(Madrid: Renacimiento, 1928).
Works Cited
Armas, Frederik de. 1985. Caves of fame and
wisdom in the Spanish pastoral novel. Stud.
Philol. 82:332-58.
Becker, Raymond de. 1968. The understanding
of dreams. New York: Hawthorn.
Brill, A. A. , ed. and trans. 1938. The basic writings
55
Wisconsin Academy of Sciences, Arts and Letters
of Sigmund Freud. New York: Modem Library.
Cervantes, Miguel de. 1986. El ingenioso hidalgo
don Quijote de la Mancha. Madrid: Coleccion
Austral.
Gutheil, Emil A. 1951. The handbook of dream
analysis. New York: Liveright.
Hughes, Gethin. 1977. The cave of Montesinos:
Don Quixote’s interpretation and Dulcinea’s
disenchantment. Bull. Hisp. Stud. 54: 107-13.
Johnson, Carrol B. 1983. Madness and lust. Los
Angeles: University of California Press.
Palmer, Donald. 1971. Unamuno, Freud and the
case of Alonso Quijano. Hispania 54: 243-48.
Riley, E. C. 1986. Don Quixote. London: Allen
and Unwin.
56
The Distribution of Franklin’s Ground Squirrel
in Wisconsin and Illinois
Timothy L. Lewis and Orrin J. Rongstad
Abstract. Eastern populations of Franklin s ground squirrel (Spermophilus franklinii) have
declined in the past two decades. We studied the current range of this squirrel in Wisconsin
and Illinois to determine whether a reduction in range accompanied the population decline.
We contacted 236 biologists in Wisconsin and Illinois by mail and telephone to determine the
extent of recent sightings. We found a range extension in northwestern Wisconsin but a range
reduction in southwestern Wisconsin and northwestern Illinois. Several possible explanations
for the range reduction are discussed.
Agrowing body of evidence indicates
that in recent years the number of
Franklin’s ground squirrels has declined at
the eastern extent of its range (Van Petten
and Schramm 1972; Panzer 1986; Johnson
1988). We studied the distribution of this
squirrel in Wisconsin and Illinois to deter¬
mine its current range.
In Indiana Franklin’s ground squirrel was
listed as a “species of special concern” (Pan¬
zer 1986). Recent trapping work in Indiana
indicated a substantial reduction in the ground
squirrel’s distribution (Johnson 1988). At least
two reintroductions of Franklin’s ground
squirrel in Illinois have succeeded in counter¬
ing this decline (Van Petten and Schramm
1972; Panzer 1986). In Wisconsin the squir¬
rel is currently managed by the Bureau of
Endangered Resources.
Franklin’s ground squirrel is reclusive, hi¬
bernating from late September until April each
Timothy L. Lewis is Assistant Professor of Biology at
Wittenberg University, Springfield, Ohio. He was for¬
merly with the Department of Wildlife Ecology at the
University of Wisconsin-Madison .
Orrin J. Rongstad is with the Department of Wildlife
Ecology at UW -Madison.
year (Sowls 1948; Panzer 1986), and is strictly
diurnal. Thus it may spend 90% of its life
below ground (Sowls 1948). Its habitat is
native prairie, brushy borderlands, fence rows
bordering cropland and railroad tracks, or
marshland edges (Cory 1912; Sowls 1948;
Jackson 1961).
This squirrel’s natural range was almost
exclusively in the tall- and mid-grass prairie
region (Hall 1981; Hall and Kelson 1956).
The general range of the squirrel has not
changed much in recent times, although De
Vos (1964) reported a slight range extension
along the Indiana-Michigan border, and An¬
derson (1947) reported a range extension in
Manitoba. De Vos (1964) attributed the ex¬
tension along the Indiana-Michigan border to
human-influenced disturbances. Smith (1957)
attributed the Manitoba extension to climatic
changes.
The Franklin’s ground squirrel has prob¬
ably always been an uncommon species in
Wisconsin and Illinois. The squirrel is more
abundant farther west in Minnesota, the Da¬
kotas, and north into the plains of Canada.
Wildlife biologists in the eastern range of the
squirrel feel that the abundance of the ground
squirrel has declined during the past twenty
years. This survey was conducted to deter-
57
Wisconsin Academy of Sciences, Arts and Letters
mine whether there have been any changes
in the ranges found in Wisconsin and Illinois.
Methods
We identified wildlife biologists and state
park naturalists as the people most likely to
be familiar with the Franklin’s ground squir¬
rel. In October 1986 we surveyed each bi¬
ologist by mail and by follow-up telephone
interview about recent and past ground squir¬
rel sightings. In the past, sightings were often
recorded because of the squirrel’s destructive
role as a nest predator (Sowls 1948; Sargeant
et al. 1987). Each biologist was asked to
report any Franklin ground squirrel sightings
made in the past ten years.
In Wisconsin we contacted all 68 Depart¬
ment of Natural Resources wildlife biologists
and managers, as well as 4 U.S. Fish and
Wildlife biologists at Horicon National Wild¬
life Refuge. In Illinois we contacted 22 of
25 wildlife managers and all 6 natural heri¬
tage biologists. In addition, we wrote to each
state park supervisor or naturalist at Wiscon¬
sin’s 61 state parks and recreation areas, the
Illinois Department of Conservation’s 71 state
parks and recreation areas, and 4 forest pre¬
serve districts. A sample of nonrespondents
was made to determine nonresponse bias.
Information on Franklin’s ground squirrel
was also solicited from the general public
through wildlife managers, radio programs,
and personal contacts in areas where Frank¬
lin’s ground squirrels were previously found.
Most such sightings reported by the public
were other small mammals; however, several
sightings were later confirmed by personal
observation. Each potential sighting location
in Wisconsin was visited and livetrapping
attempted at five locations.
Results
We received 70 responses from 126 bi¬
ologists (many responded jointly) of the 236
biologists originally surveyed. Follow-up
telephone calls to 15 nonrespondents indi¬
cated nonresponse was due to lack of sight¬
ings to report.
Wisconsin
The Franklin’s ground squirrel was re¬
ported in 14 of 72 counties in Wisconsin.
There were 35 sightings reported for 28 lo¬
cations (Fig. 1). Concentrations of squirrels
Fig. 1. The reported locations of Franklin’s
ground squirrel sightings by Illinois and Wis¬
consin biologists for 1985 and 1986. Note the
lack of sightings in the unglaciated southwest
portion of Wisconsin and northwest Illinois.
58
Distribution of Franklin s Ground Squirrel
were found in Douglas, Burnett, and Rusk
counties in northwest Wisconsin, and in
Waukesha, Racine, and Kenosha counties in
southeastern Wisconsin, as well as thinner
groupings in between in an area ranging from
Marathon County to Dodge County. In ad¬
dition, one Franklin’s ground squirrel was
observed during trapping near Horicon Na¬
tional Wildlife Refuge, and two were col¬
lected in northwestern Douglas County.
Illinois
Franklin’s ground squirrels were reported
in 22 locations in 16 of 102 counties in Il¬
linois (Fig. 1). Squirrels were reported in the
northeast in Cook, DuPage, and Will coun¬
ties. All other sightings were in a band of
central counties from Henderson and Green
counties to Ford, Vermilion, Coles, and
Champaign counties. Squirrel range in Illi¬
nois showed no new extensions; no sightings
were reported in northwestern Illinois, con¬
tiguous to an area of southwestern Wisconsin
where there were no squirrels.
Discussion
Wisconsin
Cory (1912) listed the range of Franklin’s
ground squirrel in Wisconsin as southern and
western Wisconsin. His map (Fig. 2 A) de¬
picted the range from Burnett County south¬
east to Walworth County, west of Lake
Michigan on the Illinois border. No specific
sightings were listed.
Hall and Kelson (1956) drew the range line
closer to Lake Michigan in the southeast,
including Racine and Kenosha counties, south
of Milwaukee (Fig. 2B). They listed only
one specific sighting in Wisconsin, at Lake
Delavan, and relied on sightings in Minne¬
sota and Illinois to place the range line in
Wisconsin.
Jackson (1961), dealing specifically with
Wisconsin mammals, as had Cory (1912),
listed 32 sightings and museum specimens
dating from pre-1900 to 1960. Jackson’s
range for the Franklin’s ground squirrel
(Fig. 2C) is the most accurate to that date,
Fig. 2. The historical distribution of the Franklin’s ground squirrel in Illinois and Wisconsin.
Figure A shows the early distribution in both states according to Cory (1912). Figure B shows
the distribution according to Hall and Kelson (1956). Figure C gives the Illinois distribution of
Hoffmeister and Mohr (1957) and the Wisconsin distribution after Jackson (1961). Figure D
shows the most recent range estimates for both states as given by Hall (1981).
59
Wisconsin Academy of Sciences, Arts and Letters
based upon museum specimens and sightings
from authorities. He confirmed the range de¬
scribed by Hall and Kelson (1956) in Racine
and Kenosha counties with actual records
and moved the northwest range more south¬
erly to Polk County, just south of Burnett
County.
Hall (1981) revised the 1956 range map in
Wisconsin (Hall and Kelson 1956) in light
of Jackson’s 1961 work and one additional
sighting in Hibbing, Minnesota, and one in
Duluth. Using no records from Wisconsin,
Hall estimated the range to extend north in
northwest Wisconsin into Douglas County
(Fig. 2D). We found five Franklin’s ground
squirrels, including two livetrapped, for
Douglas County, verifying Hall’s 1981 range
estimate. All of the Wisconsin ranges listed
are close to the tension zone described by
Curtis (1959).
Illinois
Cory (1912) placed the range of the Frank¬
lin’s ground squirrel in Illinois as the entire
northern two-thirds of the state except Lake
County in the far northeast (Fig. 2A). His
southern line ran from Madison to Clark
counties.
Hall and Kelson (1956) established the range
80 km farther south based on one sighting in
St. Clair County (Fig. 2B). They also in¬
cluded one sighting in Lake County in the
northeast. Hall (1981) did not modify his
earlier range map (Hall and Kelson 1956)
after twenty-five years (Fig. 2D).
Hoffmeister and Mohr (1957) were more
conservative with their range line (Fig. 2C).
Their line of known locations was farther
north from Adams County to Vermilion
County, with a disjunct population in St. Clair
County.
Our results tended to follow a line from
Madison County to Clark County in the south,
though sightings were reported from St. Clair
County. There were no sightings in northwest
Illinois contiguous with the area in south¬
western Wisconsin that had no recent Frank¬
lin’s ground squirrel sightings.
Changes in distribution
It appears from our distributional data that
Franklin’s ground squirrels have a relatively
stable range in Wisconsin and Illinois. How¬
ever, we found no sightings in southwestern
Wisconsin or northwestern Illinois, where a
few squirrels had previously been reported
(Jackson 1961).
Illinois naturalists familiar with Franklin’s
ground squirrel think it has declined over the
past thirty years, although precise data are
lacking. Jim Grude of the McHenry County
Conservation Department attempted to trap
ground squirrels in the county but found none
during the summers of 1986 or 1987 (pers.
com.). Van Petten and Schramm (1972) wrote
twenty years ago of the “increasing rarity”
of Franklin’s ground squirrel in Illinois. Many
of the responses to our survey also included
comments suggesting the loss of squirrels,
or at least the perception of loss from de¬
creased frequencies of sightings. In order to
counter the decline in Illinois, Van Petten
and Schramm (1972) in Knox County and
Panzer (1986) at the Markham Prairie have
with some success attempted reintroduction
into the former range.
There are several reasons that the Frank¬
lin’s ground squirrel may no longer be found
in southwestern Wisconsin. The squirrels may
never have been common in the unglaciated
portions of Wisconsin and Illinois. This area
is covered by a thin layer of unconsolidated
material less than fifty feet thick, and often
only inches thick. Erosion can be severe and
could create a problem with burrow
construction.
Land-use changes seem to be a primary
candidate for causing a decline, as suspected
in places in Minnesota as early as 1 892 (Her¬
rick). Sowls (1948) related a comment from
C.C. Fumiss that the squirrel “appears to be
retreating before the advance of agriculture.”
Van Petten and Schramm (1972) blamed cul¬
tivation, mowing, and grazing for the decline
of Franklin’s ground squirrel. However, Cory
(1912) felt the squirrel was not greatly af¬
fected by the cultivation of land.
60
Distribution of Franklin’ s Ground Squirrel
The general loss of prairie habitat alone
may not be entirely responsible for the de¬
cline. The Franklin’s ground squirrel is often
locally abundant while nearby areas have none
(Jackson 1961). Recent trappings in the plains
of Canada by A. Sargeant revealed small
concentrations of the squirrel isolated by large
areas without them, despite apparently ho¬
mogeneous habitats (pers. com.).
Isolation of these “islands” could easily
lead to long-term numerical declines. New-
mark (1987) found that over 40% of all spe¬
cies of lagomorph, carnivore, and artiodactyl
(12 species) found in western national parks
have become extinct. The loss of park species
was attributed to the loss of mammals on
adjacent lands, isolating the park popula¬
tions. The populations within the park were
smaller, less stable, and isolated from po¬
tential recolonizers. Habitat fragmentation in
agricultural areas could similarly isolate ground
squirrel populations.
Another factor contributing to a decline
may be that the populations are cyclic. Erlien
and Tester (1984) noted a ten-year cyclic
population pattern in Franklin’s ground squirrel
that they linked to predator shifts during cyclic
lows in the snowshoe hare population. Sowls
(1948) noted a six-year cycle at Delta, Man¬
itoba, that he attributed to climate, infertility,
and disease. Normal cyclic declines in frag¬
mented populations could eliminate some
populations even though habitat is suitable,
and the isolation would prevent reoccupa¬
tion, leading to a general decline. However,
the apparent decline in Franklin’s ground
squirrels at the eastern extent of the range
has been noted for more than twenty years,
and farther south than other cyclic popula¬
tions. There seems to be no macroclimatic
change that could exclude the squirrels from
the area, as they are found farther south in
Illinois, farther north into Canada, farther
east into Indiana, and farther west into Illi¬
nois, Iowa, and the Dakotas.
Further work on site- specific changes in
habitat should be done to examine changes
over time in areas that may have lost or gained
Franklin’s ground squirrels. Also necessary
are studies of reproductive success and
survivorship.
Acknowledgments
We would like to thank the many biolo¬
gists in Illinois and Wisconsin for their co¬
operation with this research, and S. Craven,
R. Guries, and an anonymous reviewer for
comments. Support for this study was pro¬
vided by the Max McGraw Wildlife Foun¬
dation, the University of Wisconsin-Madi-
son College of Agricultural and Life Sciences,
and the University of Wisconsin-Madison
Graduate School.
Works Cited
Anderson, R. M. 1947. Catalogue of Canadian
recent mammals. Canada Dept, of Mines and
Resources Bulletin 102, Biol. Ser. no. 31.
Curtis, J. T. 1959. The vegetation of Wisconsin.
Madison: University of Wisconsin Press.
Cory, C. B. 1912. The mammals of Illinois and
Wisconsin. Publication 153. Chicago: Field
Museum of Natural History.
De Vos, A. 1964. Range changes of mammals in
the Great Lakes region. Am. Midi. Natur. 71:210—
31.
Erlien, D. A., and J. R. Tester. 1984. Population
ecology of sciurids in northwestern Minnesota.
Can. Field-Nat. 98:1-6.
Grude, J. Telephone conversations with author, 6
October 1986 and 11 September 1987.
Hall, E. R. 1981. The mammals of North Amer¬
ica. New York: Wiley.
Hall, E. R., and K. R. Kelson. 1956. The mam¬
mals of North America. New York: Ronald Press.
Herrick, C. L. 1892. Mammals of Minnesota.
Geologic and Natural History Survey of Min¬
nesota, Bulletin 7. Minneapolis: Harrison and
Smith.
Hoffmeister, D. F., and C. O. Mohr. 1957.
Fieldbook of Illinois mammals. Urbana: Natural
History Survey Division.
Jackson, H. H. T. 1961. Mammals of Wisconsin.
Madison: University of Wisconsin Press.
Johnson, S. 1988. Status and distribution of the
Franklin’s ground squirrel. Indiana Wildl. Mgmt.
and Research Notes, no. 407 (March).
Newmark, W. D. 1987. A land-bridge island per¬
spective on mammalian extinctions in western
North American parks. Nature 325:430-32.
Panzer, R. 1986. Franklin’s ground squirrel
61
Wisconsin Academy of Sciences, Arts and Letters
translocated to an Illinois prairie preserve. Res¬
toration and Management 4(1): 27.
Sargeant, A. B. Conversation with author, 10
September 1987.
Sargeant, A. B., M. A. Sovada, andR. J. Green¬
wood. 1987. Responses of three prairie ground
squirrel species, Spermophilus franklinii, S. ri-
chardsonii, and S. tridecemlineatus, to duck
eggs. Can. Field-Nat. 101:95-97.
Smith, P. W. 1957. An analysis of post- Wisconsin
biogeography of the prairie peninsula region
based on distributional phenomena among ter¬
restrial vertebrate populations. Ecol. 38:205-19.
Sowls, L. K. 1948. The Franklin’s ground squir¬
rel Citellus franklinii (Sabine) and its relation¬
ship to nesting ducks. J. Mammal. 29:113-37.
Van Patten, A., and P. Schramm. 1972. Intro¬
duction, dispersal, and population increase of
the Franklin’s ground squirrel, Spermophilus
franklinii, in a restored prairie. In Proceedings
of the Second Midwest Prairie Conference, ed.
J. H. Zimmerman. Madison: University of
Wisconsin.
62
That Eyes May Be Free: Mary North Allen
Talks with Transactions Editor Carl Haywood
In 1990 the Wisconsin Academy of Sci¬
ences, Arts, and Letters established the
Dresen Award in memory of David Dresen,
longtime photographer with the University of
Wisconsin Photo Media Center.
Mary North Allen was the first recipient,
and the award was presented at the Acade¬
my’s annual meeting held at UW- Platte ville.
Looking at the photographs, I was intrigued
with the question, “What are Mary North
Allen and her work about?’ ’ After the awards
luncheon I sought her out to talk about the
possibility of publishing some of her pho¬
tographs in Transactions, perhaps with an
interview regarding her work and her view
of photography. A year later we met for the
interview at the Academy meeting at UW-
Superior. I thought it would be a relatively
easy assignment, but that was before I knew
Mary North Allen. After talking to her for
hours, listening to the tapes, and reading
transcriptions, I found the results complex
and subtle. Months of correspondence have
followed, with telephone discussions for
clarifications, and Mary has put her answers
in writing, but the answer to my question
remains elusive. As with so many things, the
interview published here does not do justice
to the complexity and richness of one of Wis¬
consin’s remarkable people. The purpose will
have been served if the reader comes to see
the beauty in both the person and her work.
What started out as an assignment has be¬
come a joy, and it is my hope to share that
with our readers.
Transactions: When we talk about “ photog¬
raphy ? ’ what idea or definition comes to mind?
MNA: On a simple level photography is a
way of making pictures by bringing light to¬
gether with a light-sensitive surface. A cam¬
era is a light-tight box containing a device
for holding film. Opposite the film is a hole
through which the light can pass. For opening
and closing the hole a cork would do or a
piece of electrician’s tape. All the stuff you
pay so much money for is there to let you
control exactly how the light will enter the
camera and impinge upon the film. But most
people buy cameras with automatic controls
to coordinate the various actions. Some of
us prefer to do our own coordinating.
The light entering your camera is, of course,
not pure light. What enters the camera is light
that is reflected from the surfaces in front of
the camera. The configuration of light and
shadow which impinges on the film is thus
a reflection of the surface appearance of what’s
in front of the camera. But surface appear¬
ance changes every hour of the day, every
time a cloud passes, every season of the year.
Although the camera is often figuratively
described as an “eye,” it cannot see the way
a person sees. The purpose of photography,
in most cases, is that you want to share with
somebody your interest in what you are seeing.
The camera is just a box. It has no way of
knowing about your interests.
Transactions: In our discussions you have
often used the words ‘ ‘ seeing ’ ’ and ‘ ‘ vision ! ’
Do these have special meaning in your ap¬
proach to photography?
MNA: Vision is more than reception of raw
data, more than identification, more than
seeing enough to avoid bumping into things.
Vision is a thought process, interrelated with
all of mind’s activity, with all our formal and
experiential learning, and with all the bag¬
gage, both essential and superfluous, that we
all carry. A configuration of various inten¬
sities and wave lengths of light impinging
upon the retina of our eyes generates nerve
impulses that travel the optic nerve to enter
an intricate network of thousands of brain
cells. Herein lies the intuitive part of the visual
63
Wisconsin Academy of Sciences, Arts and Letters
process — what the photographer depends
upon.
I believe that each person’s vision is as
unique to that particular person as his or her
voice. We recognize the voice of a friend
whom we haven’t seen for years. But how
can any of us know our friend’s vision — or
our own vision — unless we communicate it
in some way? And my way is to make some¬
thing — to create a photograph.
Transactions: Have you always had this sense
of seeing or did something in your back¬
ground develop it?
MNA: I am one of those who can’t draw, or
so I always thought. When I was a child in
rural New York State, there was no art taught
in the local schools. Nor was there music,
except for a singing class, for which the teacher
screened us in first grade. He determined that
roughly half of the pupils were listeners, not
singers. Through twelve grades we sat while
others sang.
We lived on a farm and I played in the
woods. I waded in a clear, clean, sparkling
brook. In my trusty oatmeal carton I collected
flowers and pretty stones, twigs of interesting
shape, colored leaves, hemlock cones, wild
strawberries and blackberries and raspberries
(a patch of white raspberries grew by the
remains of an old stone wall where trees met
the upper field) and beechnuts, if I could get
them ahead of the squirrels. When I lugged
my treasures home, my mother didn’t scold
about the holes in the knees of my stockings
or my hair ribbon tattered with briars. In¬
stead, she admired my precious findings and
listened to my tales of adventure. It was
imaginative, inventive play.
After public schools I majored in biology
at Mills College, then washed laboratory
glassware at Hopkins Marine Station, where
I marvelled at intertidal fauna and studied
microbiology. Biology appealed to me as a
new way of seeing the living earth I felt so
close to.
In 1946 I came to Wisconsin with my hus¬
band, who was for thirty years professor of
botony at the University in Madison. We raised
three children on a farm near Mt. Horeb. I
had long been interested in photography and
in the 1960s decided to study it seriously. I
entered the UW- Department of Art, where
I was fortunate to work with George Gamb-
sky, who taught photography and who him¬
self had been a student of Minor White.
In biology I had been asked, ‘ ‘On the basis
of what observations do you draw your con¬
clusions?” There was some important dif¬
ference in the seeing I was now learning to
practice. I was engaged in yet another kind
of learning, another mode of thinking new
to me. Gambsky was reminding us that mak¬
ing a photograph is sending a message. ‘‘Don’t
you think you ought to know what sort of
messages you are sending?” We would sit
in class, in long, concentrated silence, look¬
ing at our own and each other’s prints, before
anybody spoke up.
The photographer has to grasp what Cartier-
Bresson called the “precise moment,” when
all elements of your picture come together,
and this can only be done intuitively. Only
later, when studying prints, can you assess
the message content.
In 1971 I hung my first one-person show.
I was already past fifty, but I had finally
found the right work for me. I was elated to
discover that there is such a thing as grasping
a concept aesthetically, that image-making
opens a whole new approach to understand¬
ing. Above all, I had the joyful feeling that
some part of me that had been totally ne¬
glected was coming to life. I’m still learning,
along with my students, many of whom have
experienced a similar liberation themselves.
Only after several more decades did it dawn
on me that the driving question of my life
has been to make sense of the dilemmas of
being human, the paradoxes that have been
puzzling me as long as I can remember, and
that this is the stuff of art — all the arts. The
arts are of and for all of us, and in simpler
eras they played essential roles in daily living.
It is photography as an art form that par¬
ticularly interests me. There are countless
other applications of photography, from X ray
64
Interview with Mary North Allen
to remote sensing to advertising, about which
I know very little.
For ten years I taught photography with
the UW-Extension, which for much of that
time had one of the best photography pro¬
grams in the country, with more than a dozen
instructors under the direction of Tom
Mclnvaille. In 1985, after Tom had left and
the peak of that excellence had passed, I re¬
signed to found CAMERA WORKS, making
my home into an independent center for pho¬
tographic study.
In the CAMERA WORKS program be¬
ginners do an intensive eight-week sequence
of structured assignments that enable them
to see with their own eyes how the technical
adjustments they have made influence their
images. The basic building block of photog¬
raphy is the one-stop difference in exposure.
Making a series of photographs of the same
subject with a one-stop difference, they see
how change in the intensity of light looks and
how it alters relationships between the ob¬
jects in the picture. At one end of the series,
detail in the shadows is more visible; at the
other end, detail in the highlights. Often there
are several acceptable possibilities, and the
photographer must make a choice: What will
be revealed, what will remain hidden?
From experiments with light, students go
on to learn about depth of field as they work
on problems of making a two-dimensional
picture out of the three-dimensional world.
Finally they work on problems of making a
still photograph out of the hustle and bustle
of a world endlessly in motion.
Encouraged to be aware of the design ele¬
ments all around them, and to see with their
own eyes, students apply the discipline of
the photographic craft to whatever they see
in their daily lives. They begin to discover
new worlds on their own doorsteps. Some¬
times in class we all spark each other. We
stretch ourselves. We outdo ourselves, and
we’re all amazed at the photographs that
emerge.
CAMERA WORKS has been enormously
successful as a pedagogical experiment. I
found that I love teaching, and I’ve come to
see teaching as inseparable from learning.
Take the problems of teaching depth of field.
It is hard to find words for speaking of re¬
lationships in space, as distinct from the mea¬
surement of distance. There was the hockey
coach still mystified after two class sessions.
Well, I venture, sometimes he must be
watching just one player, and other times he
must watch the movements of several. How
does he get just one in focus for one picture,
and then for another picture get several play¬
ers in focus all at once? Right away he sees
the problem, and I have a new way of pre¬
senting it.
I tell a family story to illustrate the com¬
plexities of thinking spacially. Many years
ago we took our children to Niagara Falls.
Here we are standing at the top of the cliff.
I am getting nervous at the height, wanting
to get away before somebody falls. Suddenly
the four-year-old, all excited, is pointing to
the boatload of people at the bottom of the
falls. “Look at all the little people!” he
exclaims.
A small child, in the habit of looking up
at adults towering above him, has found some
adult-type people who are the size of his little
fingernail — what a discovery! Somewhere
along the line we learn to use that illusion of
diminished size as a measure of distance. But
we are not aware we ever learned it until a
child shows us.
Vision is not a simple mirroring of the
world. It’s a capability we have to learn to
use. As I study the responses of students to
my assignments. I realize that I am not clear
enough in what I ask of them. As I search
for ways to clarify my class presentations,
writing and rewriting assignments, I gradu¬
ally see the problems more clearly and in
larger perspective. How the camera functions
in relation to light, in relation to space, in
relation to time — those camera functions be¬
come more understandable when I realize that
vision has got to do with seeing things in
relation to one another — brighter/darker,
nearer/farther, stationary/in motion. I think
about people functioning in relation to light,
to space, to time. Thinking “in relation to”
65
Wisconsin Academy of Sciences, Arts and Letters
is different from thinking absolutes. Visual
thinking, with its own logic, known as de¬
sign, supplements, complements, enriches
all of mind, all of living.
Half way through the foundation course at
CAMERA WORKS I can begin to detect that
some students have a strong sense of form,
some a subtle sense of timing, some a feel
for color — capabilities they probably did not
know they had. I have watched countless
people free their vision from the blinders of
stereotype, but there is absolutely no pre¬
dicting who is going to do well in photography.
Transactions: Is photography what we see,
a representation of something, or does it
mainly connect us with something we have
experienced?
MNA: A simplistic answer is “all of the
above.” If you want to consider the question
in depth, you have to ask whether “what we
see” is physical world or mental construct.
I have ideas about this, but I’ll leave the
discussion to others more qualified.
We can use photographic materials and
processes in whatever way suits our pur¬
poses, and the results will bear some resem¬
blance to whatever the camera is pointed at.
The critical question then is the kind of re¬
semblance and how it relates to your purpose:
What is your interest in your subject?
You can use words to write a government
document, a business letter, a sonnet, or a
shopping list. So with photography. There
are so many applications of photographic ma¬
terials and methods that I cannot possibly
speak for all. To me photography is a means
of representing a person’s vision/perception
of a subject.
Transactions: If you see photography con¬
nected with vision/perception, do you also
see it as a symbol, like language?
MNA: Yes, the photograph is a symbol in
that it stands for or represents. The photog¬
rapher learns technical basics as tools for
building images with some sort of message
content. When I speak of symbol, I do not
mean just the flag or the cross. I refer to
symbol systems and all the various carriers
of message that human beings have devised:
spoken and written words, numbers, music,
dance, theater, as well as all the visual arts.
A photograph is, after all, a two-dimensional
object, an emulsion on paper. But it can bear
a powerful illusion of the three-dimensional
world we walk around in. In a sense all rep¬
resentation is illusion. I sometimes think that
a photograph is much like theater. It con¬
denses. It intensifies. It demands the willing
suspension of disbelief.
In an anecdote told about Picasso, a vil¬
lager, finding the master in his garden, ap¬
proaches him with a question, “Why don’t
you paint a woman the way a woman really
looks?”
“How does a woman really look?” asks
Picasso.
The man takes from his pocket a snapshot.
“Here, this is how a woman looks. This is
my wife.”
Picasso looks intently, points to the pho¬
tograph and asks, ‘ ‘This really is your wife?’ ’
“Oh, yes, that’s my wife.”
“She’s rather small, isn’t she? And flat?”
Other creatures communicate (we are just
beginning to learn how extensively), but we
are the ones who have devised the elaborate,
complex symbol systems we think with — and
thereby structure the worlds we inhabit. It is
a peculiar predicament we human creatures
have made for ourselves, trying to live in
physical world and symbolic world simul¬
taneously. Here we sit at the junction, Janus-
like, looking both ways: we are the joining,
the transforming link our genius and our ex¬
quisite vulnerability.
I like to think of the first woman drawing
the first picture on the wall of a cave. I can
see her bursting with need . . . need to share
more than food and shelter . . . need ... to
share vision. Suddenly she picks up a charred
stick, rubs it against the rock, and out flows
form. Need and capability and tools and vi¬
sion, enhancing each other, evolving to¬
gether . . . ; today we can scarcely separate
the strands.
66
Interview with Mary North Allen
Transactions: Is what the woman produced
“truth”? It is often said that cameras
never lie.
MNA: That is like saying that words never
lie, or statistics never lie. It is not the symbol
system or technological device that does or
does not speak truth. It is the human being.
People can lie in any language, with or with¬
out technological device, if lying is their intent.
Often, I think, we confuse logical types.
It is important to distinguish between au¬
thenticity, raw data, factual information, lit¬
eral content, subliminal content, verbal inter¬
pretation. A photograph does not necessarily
have any verbal equivalent. You can talk about
it in words, but that is interpretation. Is the
picture on your driver’s license a “true” pic¬
ture of you? A teenager may want to be pho¬
tographed to look like the most recent celeb¬
rity. The teenager’s parents and grandparents
won’t think photographs of themselves are
“good pictures” if their warts show. Con¬
vention dictates that a studio portrait will make
you appear attractive in accord with current
fashion. Whether it reveals anything of your
personality or character is another question.
When a studio photographer uses lighting,
makeup, retouching, etc., to make you look
the way you would like to look, when this
is carried a step further to create fantasy for
marketing shirts, shoes, beer, or candidates
for public office, she/he is following con¬
ventional procedures. Is this lying? Wherein
does truth abide?
It is not the mechanical equipment, but the
skill and hard work of the photographer that
make portraits and ceremonial photographs
look the way we expect them to. The pho¬
tographer has to work fast, ever alert to in¬
clude all elements necessary for a coherent
image, and to exclude all distractions. The
last time I photographed a wedding is a fine
example of the hazards awaiting any pho¬
tographer who suffers a momentary lapse of
concentrated attention. Just before the cere¬
mony I came upon a great scene— -the bride
and her mother conferring with the judge in
a hallway. I failed to notice that behind them
was a door slightly ajar. I failed to observe
evidence that somebody back there was using
the phone. My flash picked up the white tele¬
phone cord and suspended it, shiny-bright
and droopy, from the nose of the mother of
the bride.
Transactions: You appear to be saying that
a photographer creates something from
her/his vision. How would the vision of the
observer relate to the photograph?
MNA: The viewing of a photograph can be
as complex as human vision. It is possible
to return again and again to a truly memo¬
rable image and experience anew the thrill
of discovery. Just as words, used to write
legal or scientific documents or stories or
poetry or lists, will be read with differing
expectations, so with photography. Most im¬
ages invite interpretation. Some require inter¬
pretation by specially trained analysts. Oth¬
ers are adventures in seeing.
In 1979, as part of an Extension photog¬
raphy instructors exhibit, I tried an experi¬
ment which I called “Participatory Photog¬
raphy.” Along with two of my photographs
I provided a little book in which viewers were
invited to write comments. The photographs
I used are the first two in the collection that
follows. I hope readers will take time to look
at these two photographs and then make their
own observations before reading further. Here
are samples of what exhibition visitors wrote:
“Not too different — both couples appear fi¬
nancially secure. One couple waits and the good
things come to them. The other goes out to get
the good things and has to work for them.”
“Two well-to-do couples. Right: Self-made
hard-working couple proudly posing before their
home.”
“I think they are my grandparents.”
“The beauty of old age — especially the beauty
of relationships that have lasted a long time.
My grandmother used to say, This is the last
for which the first was made.’ ”
“Old age is not worth the price of admission.”
“At age 52— AMEN!”
“The hell you say — I know. I’m 67 and I
love it.”
67
Wisconsin Academy of Sciences, Arts and Letters
“The secret of old age is dying young at the
last possible moment.’’
“They look so unhappy.’’
“In the final analysis we are all alone. That
is the bottom line.’’
“Pictures are grey just like the people.’’
“The photos are ten years old. Do you want
to talk them to death? It seems time to make
some new ones, Mary.’’
“Good work should be shown and reshown.”
“It is a relief to see some pictures of the
elderly as opposed to glossy-color photos of
young models.”
“The first one is great. If I knew why, it
wouldn’t be art, would it?”
“Left: Not paying attention to each other —
space between them has an uncomfortable feel
to it. Right — together.”
“It’s interesting, the man walking behind the
woman.”
“Left: Unhappy situation because of female
dominance.”
“Unhappy for whom?”
“What dominance?”
“I hadn’t even realized that the two people
in the left photo were supposed to be to¬
gether! — chilling comment on marriages of too-
long standing.”
“Do you know, for sure, that they’re
together?”
“The two people are not actually a couple.”
“Left: The action and viewing angle are
interesting.”
“Left: Crop the picture more at top and left
side to cut out extra people.”
“Extra people are vital to composition. They
balance the photograph.”
“No, they unbalance.”
“It’s easy to read too much into these pic¬
tures, but again it’s fun to think about them.
It’s good for the imagination.”
“In the end, it seems appropriate to note that
not only are the photographs themselves inter¬
esting and enlightening, but that the comments
and reflections of others add greatly to the over¬
all growing experience of the work. It should
be added that this book, as a growing statement
on art, is actually a living and growing example
of art. For our perceptions and reflections are
as significant a part of our experience as any
tangible physical items. — Thomas R. G — ’’(last
name illegible)
“It’s interesting how much people tell about
themselves when they comment on the photos.
I found the comments much more interesting
than the photos.”
“Comment on the comments: Most people
seem to have lost the gift of ‘just looking’ at
an image without (at least subconsciously)
naming it, interpreting verbally, editorializing,
or otherwise bogging it down with words. —
JT Beers”
I had selected these two photographs not
because I thought they are the finest, but
because of their potential as a pair. The pho¬
tograph on the left I took as I was sitting on
the steps of the National Gallery in London,
watching people, wondering where they came
from, where they were going, who they might
be. It is an example of image seen intuitively,
caught on the instant.
The photograph on the right: I had asked
the Grabandts, retired grocer and his wife of
Verona, Wisconsin, whether I might pho¬
tograph them. They chose to present them¬
selves exactly as they appear.
Transactions: So the photograph of the
strangers is just an image without any in¬
tended meaning or message; the viewer has
to bring meaning to it?
MNA: Two people can be seen in each pho¬
tograph, but the message content is not the
same. One photograph clearly questions, while
the other presents a statement. Format and
structure contribute to the sense of doubt of
the one, the certainty of the other. The jux¬
taposition of the two pictures reinforces this
difference. Each photograph has become part
of the context in which the other is seen.
Viewers did not find much to say about the
one on the right; the picture has said most of
what there is to say. Did you notice how
many of the comments were attempts to an¬
swer implied questions that viewers read into
the picture on the left?
Transactions: Does this experiment rule out
the possibility that a picture can contain a
universal idea? Can it have anything in com¬
mon with all people?
MNA: Are you asking whether by means of
photography a universal idea might be given
68
Interview with Mary North Allen
some form accessible to all people? I think
that is asking a bit much for any medium.
Between the message sent and the message
received “falls the shadow.”
Everyone is bom and dies and in between
experiences troubles and satisfactions. It is
not difficult to take a picture that refers to
the commonality of human experience, on
some level. More often than not, the effect
is to trivialize. A photography student taking
a picture of a beautiful sunset discovers that
the result is not automatically a beautiful im¬
age. A sensitive photographer who has wit¬
nessed and photographed a terrible event, such
as an act of war, will be painfully aware that
the photograph does not come anywhere near
conveying the full sense of tragedy.
I believe that the true archetype lies very
deep. The more profound, the more difficult
to transform into profound image. I think that
all great art arises from experience felt so
deeply that the artist feels compelled to labor
to give it form.
Viewer response will vary from individual
to individual and culture to culture. Occa¬
sionally a work breaks barriers, and the im¬
age becomes part of cultural heritage. This
can happen in photography. Almost everyone
knows Dorothea Lange’s “Migrant Mother.”
I well remember the first time I saw an
original Edward Weston print, a Minor White,
a Cartier-Bresson. For full appreciation of
fine photographs, people need to have access
to original prints. The subtleties are lost in
reproduction for publication unless the very
finest book papers are used and the very finest
printing methods, all of which are expensive.
Fortunately, it is now possible to see original
prints in many galleries, and I urge everyone
to go and to take time looking. If you have
never seen exhibition quality prints, you may
be surprised to discover what a photograph
can be.
Transactions: Michael Brenson, in a review
of ‘ An Uncertain Grace, the Photographs of
Sebastaio Salgado,” said that the photog¬
rapher had, in fact, and Tm quoting, “ turned
African tribesmen and women into Biblical
kings and queens ” And he went on to talk
about the emotion and energy in the photo¬
graphs of the people. Does his conclusion
incorporate your earlier point about what the
viewer brings to the photograph? How can
Brenson conclude that the photograph has
changed, even metaphorically, a tribesman
into a king? He implies that the photograph
creates nobility.
MNA: I’m glad that you mentioned Salgado.
He is one of the greats, one of those who tie
me to humanity on a very deep level. Salgado
came from Latin America where he had seen
at first hand terrible poverty. He became an
economist, but decided that nothing would
be done about poverty until the world moved
beyond thinking of poverty as an abstraction
and the poor as statistics. Instead of suppos¬
ing that anybody living under degrading con¬
ditions must be despicable, Salgado felt that
human beings who manage to survive ap¬
palling circumstances without losing self-
respect must have great inner strength. That
is the quality Salgado sensed and made visible.
It is the rare artist who can give form to
human suffering endured with spiritual
strength. Only a person of great conviction
would even try. Salgado has not given up
hope for humankind. And, he has succeeded
brilliantly in bringing his vision to the rest
of us. His photographs are powerfully beau¬
tiful. The reviewer was reminded of Biblical
kings. I am reminded of the writings of Elie
Wiesel and Viktor Frankl. Same fundamental
archetype expressed in different forms.
Transactions: If I showed you my photo¬
graph of my grandparents, you could eval¬
uate it technically. When I look at it, I bring
to it a set of memories and experiences pro¬
foundly different from yours. Can we ever
bridge this space? Can there be any common
understanding?
MNA: The way you phrased the question
brings to mind a student I had in class a dozen
years ago. Every photograph was, for him,
evidence of this or that lens or gadget, and
nothing more. I labored the entire semester
to find ways to encourage him to see the
image, and eventually he did. Of course,
69
Wisconsin Academy of Sciences, Arts and Letters
technical decisions influence the way a pic¬
ture looks, but concern for technique should
never obscure vision.
Sometimes the photographic technology of
the time has an influence we need to take
into account when we look at old pictures.
Were all those stem ancestors of a century
ago really humorless people? Probably they
were not greatly different from us today. You
are observing the consequences of photo¬
graphic materials and equipment now obso¬
lete. Films were slow, lenses were slow. Por¬
trait studios were equipped with head rests
and other props so that people could hold still
for up to several minutes. You might like to
try photographing your next family reunion
that way and see how you look and how it
feels.
I had grandparents too, and I am a grand¬
parent. I might look at your picture and say,
“Hmm, Carl looks like his grandfather. Or,
grandmother has keen eyes. Interesting dress
she is wearing . . . ” I don’t have to share
your family memories to enjoy looking at
your picture and to pick up information about
those two people.
I have included in this volume several pho¬
tographs that might be considered of the fam¬
ily album type. Readers of Transactions are
likely to find the pictures accessible, even
though they have no acquaintance with the
people photographed.
Outside the family, such memorabilia may
be of great interest to cultural historians and
social anthropologists who are wanting evi¬
dence of living habits and customs of ordi¬
nary folk. There is a scholarly journal called
Visual Anthropology, and John Collier’s
classic book of that title is now back in print.
Recently I received Robert Levine’s book,
Images of History, in which he discusses cri¬
teria for historians to use in determining au¬
thenticity of a photograph as document and
for evaluating content.
Transactions: What about the other pictures
you have chosen to include in these pages?
I can see that many are about nature, but
they don’t look quite like most of the nature
photographs I have seen. How do people re¬
spond to your photographs?
MNA: A forester making pictures for a field
guide for trees would be certain that criteria
for identifying species were clearly visible in
his photographs. A plant pathologist would
make sure that disease symptoms in trees
were clearly visible. Their photographs would
not look like mine, nor will the romantic
nature pictures of vacationers and departments
of tourism. All of these photographs are related
to nature, but there the similarity ends.
I’ve been inquisitive for as long as I have
memory. The life of being human presents
dilemmas. I keep trying to reconcile it with
the life of living earth. The older I grow, the
less confident I am of easy answers or ulti¬
mate answers. Not surprisingly, my photo¬
graphs raise questions, tweak the imagina¬
tion, and, I hope, set viewers to wondering.
When my intuitive vision is working, the
resulting photographs might speak some po¬
etic truth. I believe that is possible, but I do
not know.
The woodland pictures came out of a deeply
troubled time in my life. I didn’t know it
then, but the making of those photographs,
in helping me to see my life in a larger con¬
text, was an act of affirmation.
I walk in the woods until some sight com¬
pels my attention. Attention ... I surrender,
becoming completely absorbed in seeing . . .
spaces shaped by light and shadows and by
trees . . .
In the last dozen years I have moved from
exploring light and spaces to exploring mo¬
tion and color: intensity and wave length,
passage of time. I have been tinkering with
color ever since a UW-Art Department course
in color theory. Color photography simply
requires use of a film that is sensitive to the
full spectrum of visible light, but I wanted
to use color to structure the image as a painter
would, for making visible what I could see
in mind’s eye — an interplay of life processes
and human experience. I wanted the added
factor of interacting color. Thinking color is
different from thinking monochrome. It took
quite a few years of experimenting before I
70
Interview with Mary North Allen
began to get the sort of imagery I wanted.
I am envisioning all this in multidimen¬
sional forms for which I know no words. I
have been reading systems theory and chaos
theory. I’m imagining the joys and terrors of
ordinary lives of ordinary folk flowing and
ebbing like river currents joining and sepa¬
rating, mixing and rejoining, slowly wearing
channels through soil, through rock, through
improbable time.
I’m wading in a little stream, so shallow
that rocky bottom colors show, a bit of sun¬
light catches a ripple, an autumn leaf floats
by ... I come in with a close-up lens on a
little red rock barely breaking surface ... I
drop a pebble, the current shifts ... I slow
down the shutter, reflections pull out like
taffy ... I am eliminating any key to scale.
The craft remains disciplined while result be¬
comes deliciously unpredictable.
I’m a child playing; the old oatmeal carton
has turned into a camera. I’ve got treasures
to share with kindred spirits.
My work is an attempt to see eye to eye
with other human beings. All I ask is an
honest response. If one person sees a lament
for vanishing forests and another itches for
a chainsaw, so be it.
A visiting professor from Japan looked long,
without words, and took home several tree
prints.
A farmer from southwest Wisconsin, un¬
known to me, saw one of my color photo¬
graphs on the wall of the Johnston Gallery
in Mineral Point. Later he told me that the
picture jumped right out of the frame.
To see eye to eye with another human being
is rare.
71
Wisconsin Academy of Sciences, Arts and Letters
The Photography of Mary North Allen
72
Photography of Mary North Allen
73
Wisconsin Academy of Sciences, Arts and Letters
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Photography of Mary North Allen
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Wisconsin Academy of Sciences, Arts and Letters
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Photography of Mary North Allen
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Wisconsin Academy of Sciences, Arts and Letters
Photography of Mary North Allen
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Wisconsin Academy of Sciences, Arts and Letters
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Photography of Mary North Allen
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Wisconsin Academy of Sciences, Arts and Letters
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Photography of Mary North Allen
83
Wisconsin Academy of Sciences, Arts and Letters
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Photography of Mary North Allen
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Wisconsin Academy of Sciences, Arts and Letters
86
Community Response to Floodplain Relocation
in Soldiers Grove, Wisconsin
Graham A. Tobin
Abstract. The Soldiers Grove downtown relocation project is often cited in the hazards
literature as a successful example of multipurpose, community-sponsored planning. Frequent
flooding has been alleviated and the local economy stimulated. Nevertheless, spatial changes
in the physical structure of the community suggest that social impacts should be examined.
It was hypothesized that unforeseen problems arising from the relocation project could lead
to a degree of disillusionment. A questionnaire survey of residents revealed that views were
mixed. Conflicts have arisen because of perceived inequalities in the distribution of benefits
accruing from the relocation. Additionally, internal dissatisfaction has disrupted community
spirit. However, Soldiers Grove remains a model for community involvement and has been
successful in many respects, although further lessons regarding spatial planning can be learned
from this project.
The relocation of Soldiers Grove, Wis¬
consin, is frequently cited as a success¬
ful example of multipurpose, community-
sponsored planning. The stimulus for the
project was a recurring flood problem ex¬
acerbated by a declining local economy. The
plans included the complete removal and re¬
location of the downtown business district,
the razing of several residential properties,
flood proofing of other structures, and the
incorporation of building ordinances zoning
the new town for solar energy. In terms of
flood alleviation, the project has been highly
successful. The old downtown area now boasts
a park and recreational facilities. However,
the social impacts of the project have not
been addressed. This research, therefore, looks
at residents’ opinions of relocation now that
Graham A. Tobin is an associate professor and head of
the Department of Geography, University of Minnesota
in Duluth. His research and publications have been
concerned with issues of water resources, particularly
flood hazard, wetland management, and groundwater
policies.
all major components of the project have been
completed.
The flood hazard literature includes a num¬
ber of examples of small-scale projects in
which real estate property has been relocated
out of the floodplain. Shawneetown, Illinois,
for instance, is often presented as a “plan¬
ning” failure since ultimately a portion of
the town opted to remain at the old site (Mur¬
phy 1958). It is interesting to note that the
new Shawneetown is now more prosperous
than the decaying remnants of the old com¬
munity. More recently, as the emphasis on
flood alleviation has broadened to incorpo¬
rate a greater range of nonstructural measures
(Dzurik 1979), other attempts at partial re¬
location have been made. These have usually
involved local residential areas subject to fre¬
quent or catastrophic flooding, rather than
downtown business districts. A few exam¬
ples include the following: Arnold, Missouri
(U.S. Water Resources Council 1981); Rob-
indale and Nelson, Pennsylvania; Clinch-
port, Virginia; Tulsa, Oklahoma; Rapid City,
South Dakota (U.S. Department of Housing
and Urban Development 1978); and Prairie
87
Wisconsin Academy of Sciences , Arts and Letters
du Chien, Wisconsin (Miller et al. 1983).
However, none of these involved the com¬
prehensive planning and community initia¬
tives that were pursued in Soldiers Grove.
The Soldiers Grove Relocation
Project
Soldiers Grove, along with several other
communities situated on the Kickapoo River,
had experienced severe flooding on eight oc¬
casions in the twentieth century. The U.S.
Army Corps of Engineers’ response was to
begin construction on a storage dam at La
Farge, thirty-six miles upstream from Sol¬
diers Grove, and to plan levee systems for
several communities. Building of the dam
started in 1969, and levee plans were pre¬
sented to the community in 1974 (U.S. Army
Corps of Engineers 1975). Since the dam
would protect only 9% of the hundred-year
floodplain area in Soldiers Grove, the levee
system was considered an essential compo¬
nent of the project. However, the high costs
of the levees and the large annual mainte¬
nance charges forced Soldiers Grove to look
for alternative solutions. Furthermore, in 1977,
President Carter imposed a moratorium on
water projects, which stopped dam construc¬
tion and left the Kickapoo communities with¬
out any flood protection. By this time, sub¬
stantial capital investment had already been
made in the project (Tobin and Peacock 1982).
The declining economic base of the town
accentuated the need for a major planning
initiative if the community were to survive.
Consequently, the flood alleviation project
grew to include major socioeconomic changes
incorporating several different goals: (1) to
eliminate the flood problem, (2) to enhance
local employment opportunities, and (3) to
stimulate the local economy. The main thrust
of the project entailed moving the central
business district to a safer location out of the
floodplain. In addition, plans were made to
create further recreational facilities by con¬
verting the old downtown area into a park
and erecting a community center in the new
town. The project called for a general revi¬
talization of Soldiers Grove. For detailed ac¬
counts of development, timing, and imple¬
mentation of the relocation project, see Becker
(1983); David and Mayer (1984); National
Science Foundation (1980); and Tobin and
Peacock (1982).
The flood hazard literature has often touted
the Soldiers Grove relocation plan as a sig¬
nificant advance in floodplain management.
For example, the successful relocation of
property away from flood-prone areas has
been commented on by Becker (1983) and
by the U.S. Department of Housing and Ur¬
ban Development (1978). Others have de¬
scribed the economic advantages accruing to
the community (David and Mayer 1984) and
the energy savings from the solar zoning or¬
dinance (Jenson and Fantle 1979). The com¬
prehensive planning initiated and undertaken
by the local community has been praised by
various authors (Becker 1983; Pierce and
Hagstrom 1978; Time 1981; Tobin and Pea¬
cock 1982). Some criticisms have been raised,
but generally these have focused on failings
of federal government programs to make any
long-term commitment to the locally spon¬
sored project rather than on specific
community-related problems (National Sci¬
ence Foundation 1980). Similar thoughts on
appropriate mixes of local, state, and federal
involvement are echoed in the Interagency
Task Force on Floodplain Management re¬
port (FEMA 1986).
In many ways relocation has been worth¬
while. The community clearly benefits from
projected savings from flood losses and cur¬
rent economic enhancement. The population
of Soldiers Grove grew from 530 in 1978 to
622 in 1980 (U.S. Department of Commerce
1982), although there has been little change
in recent years, according to local officials.
The community tax base, however, has in¬
creased by two million dollars (David and
Mayer 1984). The economic success of the
project, therefore, seems well established.
However, what has not been seriously ad¬
dressed is the impact the relocation plan may
have had on the local population. What do
residents of Soldiers Grove think of the pro¬
ject? In what ways has it affected daily liv-
88
Floodplain Relocation in Soldiers Grove
ing? The literature suggests that residents were
overwhelmingly supportive of the original re¬
location plan, and this aspect has often been
cited as a reason for the apparent success
(Pierce and Hagstrom 1978). To what extent
is this true now that relocation has been com¬
pleted? This paper examines some of the so¬
cial impacts of the relocation project on the
population of Soldiers Grove through a con¬
sideration of residents’ perceptions.
Research Questions
It was hypothesized that residents of Sol¬
diers Grove would perceive the relocation
plan as socially beneficial to the community.
The project was locally initiated and hence
demonstrated all the characteristics of self-
determination necessary to “guarantee” some
degree of success (Hoggart and Buller 1987).
However, it was further hypothesized that
some internal conflicts would materialize be¬
tween groups as the distribution of costs and
benefits was gradually realized. Hoggart and
Buller, for instance, warn against assuming
that rural communities constitute homoge¬
neous populations with uniform ideologies.
In particular, the physical separation of the
business district from the larger residential
areas might be expected to present problems,
especially to the elderly and those without
adequate transportation.
Apart from age and location of residence,
other factors thought to influence perceived
satisfaction with the relocation project in¬
cluded household income, gender, level of
education, and years of residence in the com¬
munity. In the literature, there is ample evi¬
dence that such socioeconomic traits play im¬
portant roles. Youmans (1977) suggested that
the elderly rarely benefit from development
projects and invariably carry any increased
costs associated with transportation difficul¬
ties. Since the elderly are also often women
in the lower income groups, they can be se¬
verely disadvantaged by community changes.
In addition, attitudinal contrasts often exist
between the elderly, who frequently look to¬
ward the past, and younger residents, who
are usually more forward-looking. In other
words, if benefits have accrued exclusively
to local businesses, that is, the economic elite
of Soldiers Grove, then others may now ex¬
press some dissatisfaction with the relocation
project.
Methodology
A personal, door-to-door questionnaire
survey was conducted of Soldiers Grove res¬
idents in August 1988. A stratified random
sample of households was surveyed such that
a representative proportion was drawn from
each residential area within the community.
At the time of the 1980 census, the town had
a population of 622, which included 249
households (U.S. Department of Commerce
1982). Of this population 54% were female,
77% were adults, and 37% of the adults were
over sixty-five years of age. The survey strat¬
egy called for interviewing one adult from
each of the randomly selected residences.
Eighty-five interviews were successfully
conducted with three rejections. An esti¬
mated accuracy rate for the overall survey,
based on the response/no response rate, was
plus or minus 1.5%, using the formula pro¬
posed by Moser and Kalton (1971).
The questionnaire was designed to elicit
opinions on three aspects of the relocation
plan and accumulate socioeconomic infor¬
mation on the residential population. The first
set of questions required respondents to as¬
sess Soldiers Grove as a place to live in an
attempt to determine community “spirit.” The
second set was used to examine the perceived
impacts of the relocation plan on various as¬
pects of the community, and the third set
focused specifically on the flood hazard.
Data were analyzed using standard statis¬
tical techniques including chi-squared and
simple frequency counts. Some recoding of
raw data was undertaken to organize re¬
sponses into larger groups. For instance, in¬
come was aggregated into three categories:
less than $10,000, $10,000 to $20,000, and
over $20,000. These were used to determine
whether any significant differences existed
among the responses of different groups with¬
in the community. Independent variables
89
Wisconsin Academy of Sciences, Arts and Letters
included residential location, age, sex, num¬
ber of years lived in Soldiers Grove, level of
education, and household income. Except
where otherwise stated, a probability level
of .05 was used to determine significantly
different responses.
Characteristics of Respondents
The survey sample consisted predomi¬
nantly of elderly people, with nearly 50%
over the age of fifty-five years and only 7%
under twenty-five years. Most respondents
had lived in Soldiers Grove for some time;
the modal category was over twenty-five years
(46%). Thirty-two percent of the respondents
had moved to the town since the last flood
in 1978, but only 15% since final decisions
were made regarding the relocation plan
(Table 1). It was expected that this combi¬
nation of age and length of residence would
contribute to a good understanding or aware¬
ness of the problems faced by the community
and how the relocation strategy had sought
to accommodate various interests.
Several other socioeconomic characteris¬
tics were collected. Sixty percent had grad¬
uated from high school, and nearly 15% had
a college-level education. Reported house¬
hold income for 1987 confirmed the generally
low-income nature of the community that had
been reported in the earlier studies (U.S . Sen¬
ate Oversight Hearings 1975, 7149). Forty-
nine percent of reporting households had in¬
comes lower than $10,000 and 24% more
than $20,000. As might be expected, level
of education was positively related to income
and inversely correlated with age of respon¬
dent. Consistency with the census data was
maintained across the sexes with 58% of re¬
spondents being female.
Location of residence within Soldiers Grove
was considered important, since the plan had
resulted in distinct spatial changes to the
physical structure of the community. In par¬
ticular, the relocation of the downtown busi¬
ness district was expected to present trans¬
portation difficulties for the majority of
residents. Initially, five areas were deter¬
mined, but these were later regrouped to re¬
flect proximity to the new downtown
(Table 1). Most of the respondents resided
in two areas, the Flats, which was flooded
in 1978, and the Hill. These residential areas
are immediately adjacent to the area that for¬
merly housed the old central business district
but are approximately 1.2 and 1 .5 miles from
the new town. A substantial number of re¬
spondents also lived in areas around Church
Street and Pine Street. These two were some¬
what closer to the new business district, less
than 1 and 0.75 mile, respectively. Finally,
very few respondents actually lived in the
new town. The flooding that inundated the
town in 1978 not only created problems for
Table 1. Characteristics of respondents
90
Floodplain Relocation in Soldiers Grove
the businesses but also caused extensive
damage throughout the Flats and parts of the
Pine Street area.
Results
(1) Community spirit
As a general introductory question, re¬
spondents were asked to rate Soldiers Grove
as a place to live on a scale from poor to
excellent (Table 2). Of those responding, 75%
rated the town satisfactory or better and only
5% as poor. There were no significant dif¬
ferences between responses based on the in¬
dependent variables.
Two further questions required respon¬
dents to be more specific regarding perceived
“community spirit.’’ Participants were asked
to assess the level of community spirit in
Soldiers Grove both currently and in the pre¬
move period (Table 3). Opinions on the sta¬
tus of community spirit indicated a distinct
decline. Whereas 31% rated pre-move com¬
munity spirit in the highest category, this
figure fell to 7% for the current situation.
Similarly, the proportion rating community
spirit as little or none rose from approxi¬
mately 13% to 35%. These responses were
significantly different. In 1988, therefore,
residents perceived a downturn in community
spirit in Soldiers Grove following imple¬
mentation of the relocation plan. While this
is not conclusive proof that social conditions
had deteriorated, since selective memory is
probably playing a role here, it is an indi¬
cation at least that not everyone is entirely
happy with current community affairs.
Perception of community spirit produced
significantly different results based on in¬
dependent variables of household income and
age of respondent. There was a tendency for
those with the highest incomes and to some
extent those in the youngest age group (this
latter variable was significantly different at
the .1 level) to perceive a lower level of
community spirit than others (Tables 4 and
5). Not surprisingly, it was found that those
respondents who rated Soldiers Grove less
favorably as a place to live also perceived
little current community spirit.
(2) Impacts of the relocation project
Nearly 70% of the respondents thought that
problems associated with the relocation plan
outweighed any advantages. Contrary to pre¬
vious findings described in the literature re¬
view, those in the oldest category (over fifty-
five years) were less likely to report problems
than those in the two younger age groups
(significantly different at the . 1 level, Table 6).
Since more specific information on the
perceived impacts of the relocation project
would help define the problems, participants
were asked to respond to a series of questions
related to service utilities such as water, elec¬
tricity supply, and garbage collection; quality
of the neighborhood environment; access to
public facilities and businesses, including
libraries, banks, and grocery stores; and
Table 2. Rating of Soldiers Grove as a
place to live
Table 3. Perceived community spirit*
*X2 = 19.08, 2 degrees of freedom, p = .001.
91
Wisconsin Academy of Sciences, Arts and Letters
Table 4. Perceived community spirit (1988) by household income*
personal finances (Table 7). The perceived
impact on public services and utilities was
predominantly in the no change category
(67%), although there was a large minority
(23%) who perceived that service facilities
had declined with implementation of the pro¬
ject. Those respondents who had perceived
little or no spirit in the community were more
likely to perceive a negative impact on ser¬
vices from the relocation plan (significance
level .1). The distribution of responses re¬
garding impact on neighborhood environ¬
ment was more varied, but still nearly 50%
perceived no change. However, a large pro¬
portion of respondents (36%) felt that the
relocation project had enhanced neighbor¬
hood quality. Similar results were apparent
for impact on personal finances. Over 65%
perceived no change due to the relocation
plan. It was noticeable, however, that no one
from the new town area perceived any det¬
rimental effect on their personal finances.
The greatest negative effect of the relo¬
cation plan appears to have been the impact
on access to public and private facilities. Forty-
seven percent of respondents believed that
access had got worse since the relocation plan
was implemented. However, 22% thought
the opposite. Responses from different areas
within Soldiers Grove were significantly dif¬
ferent (Table 8). Residents living in the new
town were more likely to perceive either no
change or some improvement in access,
whereas those living in the area farthest from
the new business district (the Flats and Hill
area) were most likely to see increasing prob¬
lems with access. This question generated the
most open-ended comments from partici¬
pants, many of whom complained about the
distance to stores and the general lack of
focus of the community.
The relocation project has had a recogniz¬
able impact on the businesses of Soldiers
Grove. Both David and Mayer (1984) and
92
Floodplain Relocation in Soldiers Grove
Table 7. Perceived impact of the relocation project
Table 8. Perceived impact on access by residential area*
*X2 = 7.619, 2 degrees of freedom, p = .022.
Becker (1983) have described the changes
that actually occurred during implementation
of the project. Eight businesses were lost,
including a restaurant, grocery store, meat
locker plant, laundromat, three bars, and the
local newspaper, while there were seven gains,
a restaurant/hotel, dental clinic, real estate
agency, craft store, pharmacy, an insurance
office, and an expansion to a nursing home.
On the other hand, there was a net gain in
permanent jobs of 46.5 (Becker 1983). The
number of persons employed in the business
district increased from 66 full-time equiva¬
lent jobs to 123. Residents’ perception, how¬
ever, was one of decline; 76% believed the
number of businesses had fallen. There was
a significantly different response based on
length of residence in Soldiers Grove. Those
residents new to the community were more
likely to perceive no change or an increase
in the number of businesses operating in Sol¬
diers Grove (Table 9). This response may
reflect the gradual growth and change in em¬
phasis of the new businesses. Many basic
commercial enterprises were replaced by those
related to secondary services. It was also no¬
ticeable that residents in the new town were
more likely to perceive an improvement than
residents from other parts of town.
Respondents were also asked to comment
on the relative success of remaining busi¬
nesses (Table 10). Twenty-nine percent be¬
lieved that businesses were poorer than be¬
fore the move compared to 41% who perceived
that they were more successful. Once again,
respondents living in the new town perceived
greater success than other respondents (sig¬
nificantly different at the .1 level).
(3) Flood hazard
Sixty-six percent of respondents indicated
that flooding was no longer a problem for
Soldiers Grove. This is a high negative re¬
sponse rate and certainly does not reflect the
serious nature of the flood hazard. Parts of
the residential community are still prone to
inundation, and public facilities, roads, sew¬
ers, etc. will suffer periodically from flood¬
ing. A failure to maintain old levees in the
area could also increase the incidence of
flooding in lower parts of the new parkland.
Nevertheless, this is a typical reaction, and
the hazard literature is full of discussions fo¬
cusing on the “false sense of security” gen¬
erated by implementing alleviation projects
(Burton, Kates, and White 1978).
The Soldiers Grove relocation plan had
been developed and financially supported to
93
Wisconsin Academy of Sciences, Arts and Letters
Table 9. Perceived number of businesses by length of residence*
*X2 = 8.093, 2 degrees of freedom, p = .02.
Table 10. Perceived success of local
businesses
a large extent by the local community. In
light of this direct experience, two questions
were asked regarding responsibility toward
flooding (Table 11). Financial responsibility
for correcting flood problems was placed pri¬
marily on the federal government (72%) and
secondly on state government (47%). Local
sponsorship of adjustments to flooding was
suggested by only 25% of the respondents.
In this question respondents were permitted
to nominate more than one option. These
results conflict with what actually occurred
in Soldiers Grove, where local financial com¬
mitment amounted to a substantial share of
total costs. However, some explanation is
forthcoming from the next question: Who
should take responsibility to oversee flood
control work? Here the response pattern was
different. While 51% still believed the fed¬
eral government should be held responsible,
significant groups supported state (38%) and
local (45%) control. These results indicate
that respondents would like a greater role by
the federal government financially but would
also like to retain some control over what is
actually undertaken in the community.
As a final analysis respondents were asked
their opinions on completion of the La Farge
dam several miles upstream from Soldiers
Grove. This dam had been shelved by the
Presidential moratorium on water projects in
1977 even though it was almost complete and
approximately fifty million dollars had al¬
ready been spent on it (Tobin and Peacock
1982). During discussions on the relocation
project the lack of protection offered by the
dam had been used as a strong argument for
an alternative project for Soldiers Grove. In
spite of this, 64% of respondents believed
that the dam should be completed, 14% said
no, and the rest (22%) did not know. Sig¬
nificantly different responses were found be¬
tween the sexes. Males were overwhelmingly
in favor of completing the dam, whereas fe¬
males were less likely to express an opinion.
Discussion and Conclusions
The response of residents in Soldiers Grove
to the relocation plan can be explained by
current thinking on rural communities. Not
surprisingly, given the degree of change,
Table 11. Responsibility for flooding
Financial Work control
94
Floodplain Relocation in Soldiers Grove
problems have arisen following implemen¬
tation of the project. In particular, there is a
consensus that community spirit is not good.
Over 25% rated the town as only fair or lower
as a place to live. This attitude was partic¬
ularly evident amongst those younger resi¬
dents of the town. Furthermore, respondents
perceive an inequality in the distribution of
costs and benefits accruing from the project.
Residents are now paying the costs of busi¬
ness revitalization. The spatial disassociation
prevalent within the community is especially
troubling for many individuals. Conse¬
quently, more attention needs to be addressed
to these questions before Soldiers Grove
can be cited as the “planning ideal” (David
and Mayer 1984) or model for small com¬
munity floodplain planning (Tobin and
Peacock 1982).
It is clear that residents came to accept the
idea of major change in the community and
gradually overcame any fears and uncertainty
about the future. Undoubtedly, residents per¬
ceived many gains in comparison with few
losses from relocation, which pushed the pro¬
ject toward the certainty end of the scale
(Becker 1983, 38). The severe nature of the
flood hazard (combined with a timely re¬
minder of flooding in 1978) and the declining
economic base of the community must have
been powerful incentives to accept this rad¬
ical change. These stimuli may not be found
in other communities contemplating such
drastic action.
Residential opinions on authority involve¬
ment could be explained within the context
of the theoretical structure described earlier.
The Soldiers Grove relocation project in¬
volved a high level of local commitment and
support and hence was perceived as a suc¬
cessful planning venture. Criticisms were
generally leveled at the federal government
for its vacillating policies and intermittent
funding (Hirsch 1980; National Science
Foundation 1980). Given the high local con¬
tribution to funding and the difficulties in
obtaining money from other sources, it is not
surprising to see respondents requesting greater
federal financial commitment. At the same
time, many residents wished to maintain strong
local control over any projects, thus retaining
an element of self-determination in com¬
munity affairs.
The socioeconomic elite in Soldiers Grove
had been responsible for developing the re¬
location project in the first place. The local
newspaper and several businesses had taken
very active roles in promoting the acceptance
of the project (Becker 1983, 20). This also
conforms to the theoretical structure de¬
scribed at the beginning of the paper, with
local leaders taking the initiative. It was very
much a locally inspired project that retained
local control, but this has also generated con¬
flict within the society. The heterogeneity of
groups within the community has led to dif¬
ferences of opinion about the perceived suc¬
cess of the project.
While the community tended to present a
united face to the problems confronting Sol¬
diers Grove, it is clear that some internal
conflicts now exist. Many residents are dis¬
satisfied with the relocation project and see
it as destroying the spirit, or sense, of com¬
munity. Results of the questionnaire survey
showed that residents are not entirely happy
with how benefits have been distributed. Many
residents perceive costs to have been borne
by all the community, especially regarding
the changing physical structure of the town.
Most residents now must drive to the new
downtown for groceries, for instance. Con¬
sequently, while the basic cost-benefit anal¬
ysis for the relocation plan was favorable
(David and Mayer 1984), attention should
also be devoted to particular gainers and los¬
ers of the project.
In conclusion, the relocation project in
Soldiers Grove has not been the solution to
all the community’s problems. Certainly, there
have been economic gains, and flood losses
should no longer devastate the town. How¬
ever, the social costs have also been high,
and it remains to be seen whether Soldiers
Grove can recapture or develop a sense of
community that at present appears to be lack¬
ing. The structural changes in use of space
throughout the town have clearly had
95
Wisconsin Academy of Sciences, Arts and Letters
repercussions on the community. The plan¬
ning process in Soldiers Grove, therefore,
can continue to serve as a model for other
communities, but it may also serve as an
example of the need to monitor carefully so¬
cial implications of such changes.
Acknowledgments
This research was supported in part by an
Undergraduate Research Opportunities Grant
awarded to Heidi Kroening by the University
of Minnesota. Heidi Kroening was instru¬
mental in administering the questionnaire
survey.
Works Cited
Becker, W. S. 1983. Come rain, come shine: A
case study of a floodplain relocation project at
Soldiers Grove, Wisconsin. Madison: Wiscon¬
sin Department of Natural Resources.
Burton, I., R. W. Kates, and G. F. White. 1978.
The environment as hazard. New York: Oxford
University Press.
David, E., and J. Mayer. 1984. Comparing costs
of alternative flood hazard mitigation plans: The
case of Soldiers Grove, Wisconsin. Am. Plan.
Ass. J. 50 (1): 22-35.
Dzurik, A. A. 1979. Floodplain management:
Some observations on the Corps of Engineers’
attitudes and approaches. Water Re sour. Bull.
15:420-25.
Federal Emergency Management Agency. 1986.
A unified national program for floodplain man¬
agement. Interagency Task Force on Floodplain
Management. Washington: Federal Emergency
Management Agency.
Hirsch, T. 1980. Personal communication, March.
Hoggart, K., and H. Buller. 1987. Rural devel¬
opment: A geographical perspective. London:
Croom Helm.
In Wisconsin: Kicking the Kickapoo habit. Time
8 (26 January 1981).
Jenson, C., and W. Fantle. 1979. Soldiers Grove:
Moving into the solar age. Altern. Energy
Sources 43:7-12.
Miller, D. J., C. E. Simpkins, J. L. Rooney, and
V. L. Golenzer. 1983. The St. Paul District-
IWR Prairie du Chien interim evaluation study.
In Seminar proceedings: Implementation of
nonstructural measures, ed. U.S. Army Corps
of Engineers. Policy Study 83-G520, 299-325.
Washington: GPO.
Moser, C. A., andG. Kalton. 1971. Survey meth¬
ods in social investigation. London: Heinemann.
Murphy, F. C. 1958. Regulating flood-plain de¬
velopment. Department of Geography Research
Paper no. 56. Chicago: University of Chicago
Press.
National Science Foundation. 1980. A report on
flood hazard mitigation. Washington: National
Science Foundation.
Pierce, N. R., and J. Hagstrom. 1978. One com¬
munity’s answer to flood relief. Nat. J. (Oc¬
tober): 1648-51.
Tobin, G. A., and T. Peacock. 1982. Problems
and issues in comprehensive planning for a small
community: The case of Soldiers Grove, Wis¬
consin. Environ. Prof. 4 (1): 43-50.
U.S. Army Corps of Engineers. 1975. Review of
alternatives for flood damage reduction on the
Kickapoo River. Washington: GPO.
U.S. Congress. Senate Oversight Hearings. 1975.
Committee on Public Works. Hearings before
the subcommittee on water resources. 94th
Cong., 1st sess., 6734-49, 7141-60.
U.S. Department of Commerce. 1982. Census of
population 1980. Vol. 1, Characteristics of
population; Part 51, Wisconsin. Washington:
GPO.
U.S. Department of Housing and Urban Devel¬
opment. 1978. Disaster information. Washing¬
ton: Office of Federal Disaster Assistance
Administration.
U.S. Water Resources Council. 1981. State and
local acquisition of floodplains and wetlands:
A handbook on the use of acquisition in flood-
plain management. Prepared by Ralph M. Field
Associates.
Youmans, E. G. 1977. The rural aged. Ann. Am.
Acad. Poli. Soc. Sci. 429 (January): 81-90.
96
Depth, Substrate, and Turbidity Relationships
of Some Wisconsin Lake Plants
Stanley A. Nichols
Abstract. The depth distribution and substrate and turbidity preferences are described for 78
aquatic plant taxa found in 68 Wisconsin lakes. In general it was found that 1) taxa numbers
decrease with increasing depth; 2) taxa richness is not different between substrate types; 3)
many species are restricted to shallow water, while others are broadly tolerant of water depth
variation; 4) the depth distribution of many species is skewed towards shallow water; 5) the
maximum growth depth for many species is highly variable; 6) there is a significant linear
relationship between water clarity and maximum depth of plant growth; and 7) water clarity,
water depth, turbidity tolerance, and substrate preference influence species association.
Water depth, substrate, and turbidity are
important factors affecting the growth
and distribution of aquatic plants in lakes
(Spence 1967; Swindale and Curtis 1957;
Pearsall 1920; Barko, Adams, and Clesceri
1986; Lind 1976; Dale 1981). Shallow-water
plants may be limited by mechanical damage
from ice, waves, or fluctuating water levels;
deep-water plants may be restricted by light
penetration, temperature, or nutrients. Tur¬
bidity decreases light penetration and acts
selectively, favoring species more adapted to
turbid conditions. Nutrient concentrations,
texture, amount of organic matter, and sil-
tation rate are some substrate parameters that
influence plant growth and distribution. Water
depth, turbidity, and substrate are inter¬
related. Increasing water depth decreases soil
particle size, turbulence, and light (Spence
1967).
Stanley A. Nichols is Professor of Environmental Sci¬
ences, University of Wisconsin-Extension. He is a bi¬
ologist with the Wisconsin Geological and Natural His¬
tory Survey, and he is associated with the Environmental
Resources Center and the Department of Liberal Studies
at UW -Madison. Past articles in Transactions have dealt
with aquatic plant resources in Wisconsin waters.
Is is difficult to describe habitat prefer¬
ences without detailed ecophysiological stud¬
ies of individual species. Variations in life
cycle, morphology, physiology, and repro¬
duction determine how a species relates to
the aquatic environment. Detailed life history
and ecophysiological studies are available for
relatively few, usually nuisance, species
(Nichols and Shaw 1986).
The depth, substrate, and turbidity pref¬
erences of 78 species of submergent, emer¬
gent, and floating-leaved plants in 68 Wis¬
consin lakes are described in this paper.
Specifically, the depth range where species
were found; the species preference for hab¬
itats described by substrate, water depth, and
turbidity; and the species response to various
substrate-depth habitats are described. In ad¬
dition, growth and distribution of species in
Wisconsin lakes are compared to responses
at other geographic locations. This infor¬
mation is useful for conducting further eco¬
physiological studies of individual species and
for management purposes.
Methods and Analysis
Between 1975 and 1983 detailed macro¬
phyte surveys were completed for 68 Wis-
97
Wisconsin Academy of Sciences, Arts and Letters
consin lakes. The lakes were sampled by
Wisconsin Department of Natural Resources
(WDNR) field staff or by private consultants
for the WDNR Office of Inland Lake Re¬
newal (OILR). The primary purposes of the
surveys were to design lake management
strategies or to collect benchmark limno¬
logical data.
The lakes represent a broad range of Wis¬
consin lake types with regard to geographic
distribution (Fig. 1), water chemistry
(Table 1), and human impact. Physical and
chemical data for each lake were collected
during macrophyte sampling or were col¬
lected earlier as part of surface-water re¬
source inventories for each county.
Field methods
To assure geographical coverage of a lake,
the surveyors selected sampling points using
a grid system. Grid size and the number of
sampling points per lake varied with lake
size, i.e., larger lakes contained more sam¬
pling points on a larger grid.
At every sampling point water depth was
measured to the nearest 0.1m, and substrate
was categorized as being hard (type 1: sand
or gravel) or soft (type 2: silt, muck, or floc-
culent). All plants within a circle 2 m in di¬
ameter around the sampling point were re¬
Figure 1. Location of sampled lakes
corded and were assigned a 1 to 5 density
rank based on the criteria established by lessen
and Lound (1962). Unknown species were
collected and sent to the Wisconsin Geolog¬
ical and Natural History Survey for identi¬
fication. Plant identification followed Fassett
(1969). Specimens were then sent to the Uni¬
versity of Wisconsin-Madison herbarium as
voucher specimens.
Analysis
Because the study is meant to determine
where plants grow, only quadrats with plants
were analyzed. Due to differing water clar¬
ities, plant depth for some analyses is ex¬
pressed as a percentage of the maximum depth
at which plants grew in each lake. Depth
classes of 0-25%, 26-50%, 51-75%, and
76-100% of maximum growth are reported
as depths 1, 2, 3, and 4.
Data were analyzed using standard de¬
scriptive statistics, boxplots, chi-square,
analysis of variance, correlations, and linear
regression (SAS Institute 1985; Lotus De¬
velopment Corporation 1985). Because more
information is available about common spe¬
cies than about rare ones, different levels of
analysis were necessary.
Results
Species occurrence and habitat richness
A total of 123 plant taxa were found in the
68 lakes. The numbers of taxa are nearly the
same for the two substrate types but declined
with relative depth (Table 2). The decreasing
taxa number with increasing depth was ex¬
pected. The similar number of taxa for both
substrates was not expected.
The similarity of species occurrence was
also compared for each depth- substrate class.
This was done by calculating the relative fre¬
quency of species occurrence for each depth-
substrate class from information provided in
Table 3. The classes were compared using
the similarity index 2W/A+B (Bray and Curtis
1957). The vegetation in the shallow-water,
hard-substrate habitat (i.e. , depth 1 , substrate
1) was least similar (i.e., most dissimilar) to
98
Specific Depth of
Total conductance H20 Quadrats maximum Biotic
Lake name County alkalinity pH at 25° C Secchi Area >3m <1m sampled plant growth influence
Depth, Substrate, and Turbidity Relationships of Lake Plants
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Lake name County alkalinity pH at 25° C Seech i Area >3m <1m sampled plant growth influence
Depth, Substrate, and Turbidity Relationships of Lake Plants
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other stands (Table 4). The shallowest water
was the least similar of all the depth classes.
There was little difference in similarity be¬
tween hard and soft substrates.
Taxa with fewer than ten total occurrences,
taxa identified only to the generic level ex¬
cept for Char a spp. and Nitella spp., and
free floating species such as Lemna spp. and
Wolfia spp., which have little relationship to
depth and substrate, were eliminated from
further consideration. Ceratophyllum demer-
sum was the most common species. It oc¬
curred in 34% of the quadrats and in two-
thirds of the lakes. All species with more than
two hundred occurrences are common aquatic
plants in Wisconsin. All occurred in at least
10% of the lakes. Many of the less common
species are either emergent, or they are found
in bogs or extremely hard- or extremely soft-
water lakes. They have the most restricted
habitat requirements.
Depth distribution and maximum depth of
growth
The maximum depth of any macrophyte
was 7.8 m in Clear Lake, Oneida County.
Clear Lake also had the greatest secchi disk
reading (Table 1). A least-squares regression
of 2.72 + 0.62 X describes the linear rela¬
tionship between maximum growth depth and
secchi depth in these lakes (Fig. 2). There is
a significant positive correlation between the
two factors (r = .58, N = 68, p < .001).
A maximum growth depth versus secchi disk
regression was calculated for lakes where a
charophyte ( Char a sp. or Nitella sp.) oc¬
curred in the deepest quadrat. This regression
was not significantly different from the
regression in which only non-charophytes were
found in the deepest quadrat.
The linear relationship between the max¬
imum depth of growth and secchi depth was
tested for the 43 species that occurred in five
or more lakes (Table 3). A significant posi¬
tive correlation (p < .05) was found for 13
species (Table 5). Eriocaulon septangulare
showed the strongest correlation, and all but
one of the species showing correlation were
submergent.
101
Wisconsin Academy of Sciences, Arts and Letters
Table 2. Distribution of taxa by relative depth-substrate class
Grand total taxa = 123
Boxplots (Fig. 3) show the depth distri¬
bution of individual species with ten or more
occurrences. The species are arranged in de¬
scending order of median depth. More than
75% of all plants were found in less than 3 m
of water. Chara spp. was found at 7.8 m,
the maximum depth for any species. It also
occurred over the broadest depth range. Na-
jas flexilis followed closely behind Chara with
a maximum depth of 7.5 m. Both species had
a broad outlier range. Nitella spp. had the
greatest median depth and the greatest depth
for the 75% quartile. Myriophyllum hetero-
phyllum, M. farwellii, and Isoetes macro-
spora had the broadest depth range when out¬
liers were not considered.
Species common in deep water were also
found in shallow water, but species common
in shallow were often not found in deep water.
Generally speaking, species with shallow
median depths are emergent species. How¬
ever, Potamogeton foliosus, P. oakesianus,
and P. vaginatus are submerged species with
a shallow median depth.
The maximum depths of plant growth for
46 species were compared to literature values
(Sheldon and Boylen 1977; Wilson 1941;
Schmid 1965; Denniston 1921; Lillie 1986;
Lind 1976). Twenty-three species were found
at a greater maximum depth in other lakes
(Table 6), including six species that were
found at greater maximum depth in a later
study of Devil’s Lake (Lillie 1986). Some
differences in maximum depth probably re¬
late to limnological conditions and others to
the sampling technique used to establish
maximum depth (see Spence, 1967, for a
discussion of problems related to determining
maximum depth of growth).
Various depth statistics were tested, using
correlation analysis, to determine how useful
they might be for predicting the sequential
order of maximum growth depth in a lake
(i.e., in a lake with a given flora, which
species will have the deepest maximum depth
of growth, the second deepest, and so forth).
The statistics tested were median depth, the
trimmed maximum depth (i.e. , the maximum
depth or the maximum depth not considering
outliers, whichever is the most shallow), the
maximum depth, the median of the maximum
depth for species that occurred in five or more
lakes, and the median of the maximum depth/
secchi disk for species that occurred in five
or more lakes (Table 6). Based on median
correlation values, the best predictor of the
sequential order of maximum growth depth
is median maximum depth (Table 7). Median
depth and median of maximum depth/secchi
ratio predicted maximum depth order nearly
as well. Maximum depth was the poorest
predictor of maximum depth order. On the
average, all methods were better at predicting
maximum depth order for Wisconsin lakes
than for non- Wisconsin lakes.
Substrate and depth preference
Substrate and depth preferences were tested
using a chi-square analysis on species oc¬
currence. The hypothesis tested was that the
distribution of a species is not significantly
different from the distribution of all vege¬
tated quadrats (Table 8).
Because each species acts as an individual,
significant variation from the all-species dis¬
tribution is expected. More interesting and
informative are how and to what degree each
species varies. A Z score of (observed/
102
Table 3. Number of species occurrence
Depth, Substrate, and Turbidity Relationships of Lake Plants
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Depth Depth Depth Depth Depth Depth Depth Depth Occurrence of No. lakes Association of
Species 1 2 3 4 1 2 3 4 turbid water occurring turbid water *
Wisconsin Academy of Sciences, Arts and Letters
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104
Depth Depth Depth Depth Depth Depth Depth Depth Occurrence of No. lakes Association of
Species 1 2 3 4 1 2 3 4 turbid water occurring turbid water *
Ranunculus trichophyllus 50204400 3 Nocalc
Sagittaria graminea 20208300 3 Nocalc
Sagittaria iatifolia 9000 21 230 10 9 Nonsig
Depth , Substrate, and Turbidity Relationships of Lake Plants
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105
Wisconsin Academy of Sciences, Arts and Letters
Table 4. Similarity of stands based on relative frequency
Figure 2. Secchi disk reading versus maximum depth of plant growth
expected)/(square root expected) was calcu¬
lated for each cell of the chi-square table. If
the score was less than ± 1 Z, the association
is weak (designated + or — in Table 9). If
it ranged from 1 to 2 Z, the association is
moderate (designated + + or - in
Table 9). If Z was greater than 2, it is a strong
association (designated + + + or - - in
Table 9).
A valid chi-square test requires a minimum
number of occurrences in each cell. To fit
the criteria of a valid test, species with dif-
106
Depth , Substrate, and Turbidity Relationships of Lake Plants
Table 5. Positive correlation between
maximum plant growth depth and
secchi depth (p < .05)
ferent numbers of occurrences had to be tested
in different ways. All species with 50 or more
occurrences were tested with the full eight¬
cell chi-square pattern of four depth classes
and two substrate types. Species with 25 to
50 occurrences were analyzed separately in
a four-cell depth preference test and a two¬
cell substrate preference test. Species with
15 to 25 occurrences were analyzed with a
two-cell substrate preference pattern. Species
with fewer than 15 occurrences were not
analyzed.
The patterns of positive and negative as¬
sociations were sorted until species with like
patterns occurred close to each other in a list.
The list was subjectively split, and species
groups were labelled with habitat preference
based on the pattern of positive and negative
associations (Table 9).
Twenty-six species showed a preference
for soft sediment; 14 species preferred hard
bottoms. A depth preference with no sub¬
strate preference was evident for 27 species.
The majority of these species showed a pref¬
erence for shallow water. No species showed
a unique preference for hard bottom and deep
water. With minor exceptions, species showed
a smooth transition between adjacent habi¬
tats. No species showed a strongly bimodal
distribution.
Myriophyllum heterophyllum, M. verticil-
latum, Potamogeton epihydrus, P. pectina-
tus, P. illinoensis, Heteranthera dubia, and
Char a spp. are classified as shallow species
by this technique. They may have a deeper
distribution than mid- to deep-water species
when boxplots are compared. Depth in this
test is relative to the maximum depth of plant
growth in a lake, whereas boxplots compare
absolute depth. Therefore, the two tests need
not give the same results.
Species association with turbid water
Twenty-one percent of the quadrats oc¬
curred in turbid lakes (lakes with secchi disk
readings of 1.5 m or less, Table 1). A chi-
square test was done on the 43 species that
occurred in five or more lakes to determine
whether they were found more or less fre¬
quently than expected in turbid water
(Table 3). A Z score was calculated as noted
previously to describe the strength of the
association.
No significant association was found for
14 of the species, 15 species showed a pos¬
itive association for turbid water, and 14 spe¬
cies showed a negative association with tur¬
bid water (Table 3).
Species density and habitat type
Differences in species density ranking for
the four depth classes and two substrate types
were tested using a two-way analysis of var¬
iance. Because of differences in occurrence,
the test had to be modified for some species.
Originally all species with 50 or more oc¬
currences were tested for depth, substrate,
and depth-substrate interaction. Because of
data limitations, two-way analysis of vari¬
ance could not provide a valid interaction
model for some species. In these cases the
interaction test was dropped and the analysis
was recalculated for only depth and substrate.
This test asks whether there is a significant
difference in mean density rank for a single
species between habitats where it is found.
A probability of F < .05 and at least a .5
difference between the largest and smallest
mean density were the criteria established to
107
Wisconsin Academy of Sciences, Arts and Letters
Species
0 1m 2m 3m 4m 5m 6m 7m 8m
i - 1 — - n - 1 - — i - r— r— — — i — 1
Nitella spp.
Myriophyllum heterophyllum
Ranunculus reptans
Myriophyllum farwellii
Isoetes macrospora
Potamogeton robbinsii
Myriophyllum tenellum
Myriophyllum exalbescens
Myriophyllum spicatum
Ceratophyllum demersum
Potamogeton praelongus
Najas marina
Potamogeton berchtoldii
Potamogeton pectinatus
Potamogeton richardsonii
Najas gracillima
Isoetes echinospora
Myriophyllum verticillatum
Potamogeton zosteriformis
Lobelia dortmanna
Elodea canadensis
Potamogeton filiformis
Potamogeton amplifolius
Utricularia gibba
Cyperus engelmannii
Potamogeton illinoensis
Vallisneria americana
Najas flexilis
Chara spp.
Gratiola aurea
Utricularia intermedia
Heteranthera dubia
Potamogeton diversifolius
Eleocharis robbinsii
Potamogeton obtusifolius
Potamogeton pusillus
Utricularia vulgaris
Utricularia geminiscapa
Megalodanta beckii
Eriocaulon septangulare
Potamogeton vaseyi
Potamogeton epihydrus
Potamogeton strictifolius
Potamogeton gramineus
Nymphaea odorata
Brasenia schreberi
Ceratophyllum echinatum
Nuphar variegatum
I- -
I -
— H
-I
. —I
■—I
H
H
I - •=£>
Figure 3. Boxplots of species depth distributions. Definitions follow Ryan, Joiner, and Ryan
(1981)
108
Depth, Substrate, and Turbidity Relationships of Lake Plants
Species
Eleocharis palustris
Polygonum amphibium
Zanichellia palustris
Potamogeton crispus
Ranunculus trichophyllus
Potamogeton natans
Ranunculus longirostris
Potamogeton nodosus
Nuphar advena
Eleocharis acicularis
Nymphaea tuberosa
Zizania aquatica
Leersia oryzoides
Sparganium chlorocarpum
Pontederia cordata
Sagittaria rigida
Scirpus americanus
Potamogeton foliosus
Sparganium angustifolium
Potamogeton oakesianus
Scirpus validus
Duiichium arundinaceum
Sagittaria graminea
Carex aquatilis
Elatine minima
Sparganium eurycarpum
Typha latifolia
Typha angustifolia
Sagittaria latifolia
Potamogeton vaginatus
1m
2m 3m 4m 5m 6m 7m
8m
Figure 3— Continued
test for significant differences. For any depth-
substrate cell in a single test to be signifi¬
cantly different from another cell, the p <
.05 criterion was used, but the mean density
difference of .5 was not used.
Twenty-six species showed no significant
difference across the range of depths and sub¬
strates. Five species showed significantly
higher densities on soft substrates (Fig. 4).
Seven species showed significantly different
densities according to depth (Fig. 5). Density
of Elodea canadensis and Eriocaulon sep¬
tangular e differed depending on depth and
substrate, but the depth- substrate interaction
was not significant (Fig. 4 and 5).
The density of the remaining ten species
109
Table 6. Comparison of maximum plant growth depths
Wisconsin Academy of Sciences, Arts and Letters
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110
Trimmed Median Median 1* 2* 3* 4* 5* 6*
Median maximum Maximum maximum maximum/ Lake Devil’s Trout SE Minn. Long Lake
Species depth depth depth depth secchi Mendota Lake Lake lakes Lake George
Depth, Substrate, and Turbidity Relationships of Lake Plants
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111
Wisconsin Academy of Sciences, Arts and Letters
Table 7. Correlation coefficients of depth order predictors
Table 8. Distribution of vegetated quadrats by depth-substrate type
Depth
varied significantly in the interaction between
depth and substrate (Fig. 6). Plants com¬
monly displayed significant density differ¬
ences across depth on one substrate but not
the other. This is the pattern displayed by
Ceratophyllum demersum, Heteranthera du-
bia, Myriophyllum verticillatum, Nymphaea
odorata, and Vallisneria americana. The first
two species showed no significant density
difference with depth on hard bottom; the
other three species showed no significant
density difference with depth on soft bottom.
Another common pattern is increasing
density with depth on soft substrates and de¬
creasing density with depth on hard sub¬
strates. Potamogeton berchtoldii, P. grami-
neus, and P. richardsonii displayed some
variation of this response. P. robbinsii was
the only species that showed an increased
density with depth on both substrates.
Discussion
The decreasing number of taxa found with
increasing depth confirmed results found by
Lind (1976) and Sheldon and Boylen (1977).
Most deep-water species also occur in shal¬
low water, and species distributions are skewed
toward shallow water. Lind (1976) nicely
summarizes this relationship by stating,
“Many species are restricted to shallow water
while others are broadly tolerant of water
depth variation.”
Since hard substrates appear less suitable
for plant growth (Barko, Adams, and Clesceri
1986), it is surprising that taxa richness is
not influenced more by substrate type. Where
species density is influenced by substrate,
higher densities are found on soft substrates.
This is especially true in deep water. The
shallow-water, hard-bottom communities are
probably most dissimilar to other areas be¬
cause they contain higher frequencies of
emergent and rosette species. Emergent spe¬
cies are not found in deep water, and rosette
species are found less frequently in deep water
and in shallow, soft- sediment areas.
The 1 .2-to-7.8-m range of maximum plant
growth depths for lakes in this study is similar
to that reported by Hutchinson (1975) and
112
Depth, Substrate, and Turbidity Relationships of Lake Plants
Table 9. Substrate-depth relationships
Continued on next page
113
Wisconsin Academy of Sciences, Arts and Letters
Table 9 — Continued
*+-,+ + — ,+ + + - represents weak, moderate, or strong association. Nonsig = nonsignificant
association using chi-square test (p < .05).
broader than the 1 .0-to-4.5-m range reported
by Lind (1976) for eutrophic lakes in south¬
eastern Minnesota. They are more shallow
than the 12-m maximum depth for Lake
George, New York (Sheldon and Boylen
1977), the 1 1-m depth for Long Lake, Min¬
nesota (Schmid 1965), or the 9 -in depth of
Devil’s Lake, Wisconsin (Lillie 1986). They
are considerably more shallow than the 18-m
maximum depth for Utricularia geminiscapa
(Singer, Roberts, and Boylen 1983) in Silver
Lake, New York, the 20- m maximum depth
for bryophytes in Crystal Lake, Wisconsin
(Fassett 1930), or the approximately 150-m
maximum depfh for charophytes and bry¬
ophytes in Lake Tahoe, California (Frantz
and Cordone 1967).
This study supports the findings of Hutch-
114
Depth, Substrate, and Turbidity Relationships of Lake Plants
Figure 4. Mean density of species by substrate
inson (1975), Dunst (1982), Chambers and
Kalf (1985), and Canfield et al. (1985) that
there is a significant regression between sec-
chi disk and maximum depth of plant growth.
The regression line is not significantly dif¬
ferent from one based on Hutchinson’s data
and one reported by Chambers and Kalf
(1985). Not enough statistical information is
provided by Canfield et al. (1985) to compare
regressions. The regression calculated in this
study is significantly different from Dunst ’s
regression (1982). His equation predicts deeper
plant growth. This is surprising because the
Dunst regression is based on data from 51
lakes in southeastern Wisconsin (Modlin 1970;
Belonger 1969).
A possible explanation for the difference
is the strong presence of charophytes or Na-
jas flexilis in the lakes used by Dunst. This
study shows that Char a spp. and Najas flex¬
ilis had the deepest maximum growth depths
and Nitella spp. had the greatest median and
75% quartile depths. However, a regression
equation for lakes where charophytes oc¬
curred in the deepest quadrat was found to
be significantly different from and would not
Eleocharis acicularis
Potamogeton diversifolius
Potamogeton illinoensis
Depth
Nuphar advena
Figure 5. Mean density of species by depth class
— ! - - I'"
Scirpus validus \
Elodea canadensis
Myriophyllum spicatum
Potamogeton foliosus
Eriocaulon septangulare
115
Wisconsin Academy of Sciences, Arts and Letters
g 3-
-a
I2'
Ceratophyllum demersum
Heteranthera dubia
D D
Potamogeton berchtoldii
C A
Nymphaea odorata
A
II
Potamogeton gramineus
□ Substrate 1
□ Substrate 2
g2
2
1. Same letter indicates depth classes are not signifi-
candy different on the same substrate.
* Indicates substrates are significantly different at the
same depth class.
1* 2 3* 4
Depth
Figure 6. Mean density of species by substrate type and depth class
predict plant growth as deep as Dunst’s
regression.
Decreases in maximum depth of plant
growth with increased turbidity have been
previously reported (Vander Zouwen 1982;
Hutchinson 1975; Spence 1982). The lakes
in this study with the shallowest secchi read¬
ing and shallow plant growth were strongly
influenced by carp and algae (Fig. 2). Their
influence appears to be turbidity related. All
lakes with carp problems had maximum plant
growth depths below the regression line. All
lakes but one with heavy algae blooms had
maximum plant growth depths above the
regression line. This may provide insights
into the overall impact these two biotic fac¬
tors have on plant growth.
Similar to the findings of Davis and Brinson
(1980), a linear relationship was found be¬
tween secchi disk reading and maximum plant
growth for some submerged species. Theo¬
ries that ascribe the maximum depth of plant
growth to a single factor are deficient. Actual
limitation may be brought about by a com¬
bination of factors (Singer, Roberts, and
Boylen 1983). Therefore, it is not surprising
116
Depth, Substrate, and Turbidity Relationships of Lake Plants
that maximum depth growth for many sub¬
merged species was not correlated with sec-
chi reading. Zizania aquatica, the only emer¬
gent species significantly correlated to secchi
reading, is an annual species that grows from
seed each year. Light penetration could in¬
fluence its growth. It is also a species that is
highly susceptible to water turbulence at a
critical period in its life cycle. Clear water
could be an indication of quiet water.
This study found no correlation between
maximum depth/secchi ratio, which Davis
and Brinson (1980) call a turbidity tolerance
index, and the association with turbid water
(Table 3). The results of the chi-square hab¬
itat preference test and the analysis of the
variance density test need not be comple¬
mentary. If they are complementary, habitat
preference is reflected in species density. This
could mean that depth and/or substrate has a
strong influence on species density. Species
that show a similarity between tests are Ni-
tella spp., Potamogeton praelongus, P. pus-
illus, Utricularia intermedia, Myriophyllum
spicatum, P. diversifolius, P. foliosus, Elo-
dea canadensis, Ceratophyllum demersum,
Myriophyllum verticillatum, P. richardsonii,
P. robbinsii, and Vallisneria americana.
Lack of similarity is more difficult to ex¬
plain. One possibility is that a species was
found only in preferred habitats, so densities
were similar wherever it was found. A sec¬
ond, and probably more likely, possibility is
that something that was not measured in this
study limits species distribution or density.
One easily overlooked possibility is inter¬
specific competition, which could limit a spe¬
cies in a lake even though the habitat is suit¬
able for its growth.
Summary
This study reinforces information from other
studies that taxa numbers decrease with in¬
creasing depth; that many species are re¬
stricted to shallow water, while others are
broadly tolerant of water depth variation; that
the depth distribution of many species is
skewed towards shallow water; that the max¬
imum growth depth for many species is highly
variable; that there is a significant linear re¬
lationship between water clarity and maxi¬
mum depth of plant growth; and that water
clarity, water depth, turbidity tolerance, and
substrate preference influence species asso¬
ciation. It was an unexpected finding that
taxa richness is not different between sub¬
strate type. This study differs from other
studies because it provides descriptions of
depth distribution and substrate and turbidity
preferences for a variety of Wisconsin lake
plants. The information should be very useful
for managing Wisconsin’s lake plant re¬
sources and for doing further ecophysiolog-
ical studies on individual species.
Acknowledgments
Murray Clayton, Peter Crump, and
Emmanuel Maurice of the University of Wis-
consin-Madison, College of Agriculture and
Life Sciences, provided statistical and com¬
puting assistance. Sandy Engel and Richard
Lillie of the Wisconsin Department of Nat¬
ural Resources and Michael Adams of the
University of Wisconsin-Madison, Depart¬
ment of Botany, are gratefully acknowledged
for critically reviewing the manuscript. The
Wisconsin Department of Natural Resources
provided lake vegetation and water chemistry
data. The Institute for Environmental Stud¬
ies, University of Wisconsin-Madison, pro¬
vided partial funding for data entry and
analysis.
Works Cited
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1986. Environmental factors and their consid¬
eration in the management of submersed aquatic
vegetation: A review. J. Aquat. Plant. Manage.
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Belonger, B.J. 1969. Aquatic plant survey of ma¬
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Bray, J.R., and J.T. Curtis. 1957. An ordination
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Wisconsin Academy of Sciences , Arts and Letters
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Chambers, P.A., and J. Kalf. 1985. Depth dis¬
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Dale, H.M. 1981. Hydrostatic pressure as a con¬
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Davis, G.J., and M.M. Brinson. 1980. Response
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plants of Lake Mendota. Trans. Wis. Acad. Sci.
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Dunst, R.C. 1982. Sediment problems and lake
restoration in Wisconsin. Environ. Int. 7:87-92.
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- . 1969. A manual of aquatic plants. Mad¬
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vations on deepwater plants in Lake Tahoe,
California and Nevada. Ecology 48:709-14.
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Wiley.
lessen, R., and R. Lound. 1962. An evaluation
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Lillie, R.A. 1986. The spread of Eurasian water-
milfoil Myriophyllum spicatum in Devil’s Lake,
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Lind, C.T. 1976. The phytosociology of sub¬
merged aquatic macrophytes in eutrophic lakes
of southeastern Minnesota. Ph.D. diss., Uni¬
versity of Wisconsin-Madison .
Lotus Development Corporation. 1985. 1-2-3 ref¬
erence manual. Cambridge, Mass.
Modlin, R.F. 1970. Aquatic plant survey of major
lakes in the Milwaukee River watershed. Wis.
Dept. Nat. Resour. Res. Rep. no. 52. Madison.
Nichols, S.A., and B.H. Shaw. 1986. Ecological
life histories of three aquatic nuisance plants,
Myriophyllum spicatum, Potamogeton crispus,
and Elodea canadensis . Hydrobiologia
131:3-21.
Pearsall, W.H. 1920. The aquatic vegetation of
English lakes. J.Ecol. 8:163-201.
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tistics. Version 5 ed. Cary, N.C., SAS Insti¬
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etation as measured by line intercept with
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Sheldon, R.B., and C.W. Boylen. 1977. Maxi¬
mum depth inhabited by aquatic vascular plants.
Am. Midi. Nat. 97:248-54.
Singer, R.D., D.A. Roberts, and C.W. Boylen.
1983. The macrophyte community of an acidic
lake in Adirondack (New York, U.S. A.): A
new depth record for aquatic angiosperms. Aquat.
Bot. 16:49-57.
Spence, D.H.N. 1967. Factors controlling the dis¬
tribution of freshwater macrophytes with par¬
ticular reference to the lochs of Scotland. J.
Ecol. 55:147-70.
- . 1982. The zonation of plants in fresh¬
water lakes. Adv. Ecol. Res. 12:37-125.
Swindale, D.N., and J.T. Curtis. 1957. Phyto-
sociology of the larger submerged plants in
Wisconsin lakes. Ecology 38:397-407.
Vander Zouwen, W.J. 1982. Vegetational change
in University Bay from 1966 to 1980. Trans.
Wis. Acad. Sci. Arts Lett. 70:42-51.
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Acad. Sci. Arts Lett. 33:135-46.
118
From Wisconsin Poets
Three years ago when Carl Haywood asked me to collect and edit a selection of poetry for
each issue of Transactions, I leapt at the chance to share the joy and insight that I find in
poetry with members of the Academy and other readers of the journal. I have been nothing
but astounded and immensely pleased at the level of support I have had for this project from
the Academy Board, the staff of Transactions, and the poets themselves, who have given
unstintingly of their work and their time. This ground swell of interest and support found
perhaps its epitome with the recent publication of the special issue of Transactions entitled
Wisconsin Poetry, an anthology of three hundred poems by sixty-five Wisconsin poets, which
represents the largest and most comprehensive collection of Wisconsin poetry published in
nearly fifty years.
I am especially thankful for the interest and support expressed to me by members of the
Academy I have met at readings throughout the state held in conjunction with publication of
that special issue. It is a direct result of that interest, and the efforts of Carl Haywood, that
there will be an ongoing session devoted to Wisconsin poetry at the yearly meeting of the
Academy, as well as sponsorship of the publication of individual collections of poetry by
Wisconsin poets.
The editorship of Transactions is changing, along with the site of its publication. I am sure
that poetry will continue to be a part of Transactions as well as a part of the mission and
service of the Wisconsin Academy of Sciences, Arts, and Letters.
Bruce Taylor
119
Wisconsin Academy of Sciences, Arts and Letters
About the Poets
Art Lyons manages tutoring programs at UW-Eau Claire. He has published a text-workbook
titled Writing for Workplace Success (Paradigm 1991). His poems have appeared in Wisconsin
Poetry, Slant, Confluence, Upriver 4, Wisconsin Dialogue, and Black Buzzard Review.
Jeri McCormick, of Madison, teaches creative writing at senior centers and elderhostels
and works as an editor in the State Department of Administration. She has published The
Sun Rides in Your Ribcage, a chapbook of poems, and co-authored Writers Have No Age,
a text for older adults. Her poems have appeared in the Wisconsin Academy Review, the
Wisconsin Poets’ Calendar, Poet Lore, Isthmus, and Wisconsin Poetry.
Kyoko Mori received her doctorate from UW -Milwaukee and teaches English at St. Norbert
College. She has published a book of short fiction, The Ritual in Roses and Silk. Her poems
have appeared in the South Florida Poetry Review, The Forbidden Stitch, the Denver Quar¬
terly, and the Madison Review.
Thomas R. Smith is a poet and essayist living in Minneapolis. He has one book of poems,
Keeping the Star, from New Rivers Press and a second, Horse of Earth, nearly completed.
He is an Associate Editor for Ally Press, where he is editing a festschrift for Robert Bly.
Jean Tobin lives in Black River, along the shore of Lake Michigan. She is Professor of
English at the University of Wisconsin Center -Sheboygan.
Marilyn Taylor recently earned her doctorate at UW -Milwaukee, where she received the
1991 Academy of American Poets Prize. Her work has appeared in Poetry, Wisconsin Review,
and Poetry Northwest. Her first collection, The Accident of Light, was published by Thorntree.
Laurence Giles is a Madison physician who says, ‘ ‘ Medicine is my life, but poetry is one of
my more positive obsessions." He has published a slender volume entitled Goat Cottage
Dream Poems, and his poems have appeared in Abraxas and The Literary Preview. He is
also a licensed private pilot and a scuba diver.
Joan Rohr Myers, Director of Human Relations and Affirmative Action at UW-Eau Claire,
has received the Catholic Press Association award for poetry as well as the Wisconsin Regional
Writers' Association Jade Ring and Bard's Chair awards for her poems. Her poetry has
appeared in over a hundred journals and anthologies.
Bruce Taylor, the Poetry Editor of Transactions, is Professor of English at UW-Eau Claire.
He has published two chapbooks, Idle Trade: Early Poems (Wolf song Press) and The Darling
Poems (Red Weather Press). His poems have appeared in The Nation, New York Quarterly,
and the New Orleans Review.
Ralph Schneider is Professor of English at UW-Eau Claire. A country dweller, he finds
much of his poetic material in the Wisconsin natural environment. He spent the spring semester
of 1991 on a poetry-writing sabbatical, and “ Breakfast " was one of the poems that developed
during that regenerative period.
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Wisconsin Poetry
Child to Father (later)
That all truths have their time, you never knew.
In your white ranch house, your Buick four-door,
you never knew the lies time chose for you.
You loved the timely truth of screw or be screwed;
in your time, a fuller garage bought a higher score.
That all truths have their time, you never knew,
so you hated every nigger, every spic and jew,
but loved white men whose cars out-buicked yours.
You never knew the lies time chose for you,
and wily time chose lies for me too -
truths against your lies— but let me ignore
that all truths have their time. You never knew,
I’m sure, that my spiteful words with every new
disgust meant I doubted you, doubted myself more;
you never knew the lies time chose for you.
Out of time now, your body tells the truth:
We live in time and time decides what for.
That all truths have their time, you never knew.
You never knew the lies time chose for you.
— Art Lyons
121
Wisconsin Academy of Sciences, Arts and Letters
Looking at Skulls
The first skull I saw
gleamed from a stereoscope
in Grandma’s front room.
The scene was a Mideast catacomb,
Jerusalem, perhaps, or Babylon;
Grandma favored the Bible lands.
In three scoped dimensions
two shadowy orbs leered at me
from a knob of chalky bone.
I scrambled for the next card —
benign camels crossing the desert
or the somber stillness of Golgotha,
anything to squelch that deep vacancy.
Now, all these years later,
I’ve met Hamlet and others
who do not turn from skulls;
I’ve lost Grandma
whose hidden remains still comfort,
whose skull is surely beautiful.
And I’ve made peace
with my own scaffolding —
femur, tibia, clavicle —
gliding me through this life
like a fine ghost ship,
at sea with lofty captain
intent on solid grace,
yet content with the usual gear:
a nose, an ear or two, eyes,
the accoutrements of face.
— Jeri McCormick
122
Wisconsin Poetry
Vehicles of Change
Our last week together, I borrow his
car to move my boxes. Daily, I cross
the bridge with books, dishes. The brake shoes are
slipping. By Wednesday, every stop grinds
to the roar of airplanes descending,
urgent upon the car roof. I am crash-landing
through lost time. I maneuver my craft
to lose it. I want to walk away from its
burning in some farmer’s field as the camera
crew rushes to record the miracle
of my get-away. In the ditches, cow
parsnips rotate purple shafts; rank white
crowns rise above the odor of burning
metal. Black smoke trails my path and becomes
a pack of cats skulking in my shadow.
ii.
The First of June, my friend’s
driving the panel van, the radio’s
stuck on a country station, and
I’m in the passenger’s seat
with furniture rattling in
back. The world shrinks up and
jumps into the side mirrors:
the lanes are parallel and skewed
both like corridors in
perspective drawings. You could
wrap them around the earth’s
core without their crossing. Never
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Wisconsin Academy of Sciences, Arts and Letters
the twain shall meet. I tower
over traffic; below, sunlight
glints off car tops like
pebbles I could flick over
water. Across the bridge, the wind
takes the words from my mouth and
erases.
iii.
Unpacking, I re-enact the Apartment
Within I’ve carried from place to place like
an absurd parody of the Soul, the God
imprinted in my heart. Its universals
include cups eye-level on a kitchen
shelf, scissors in the left-hand utility
drawer, the vacuum cleaner plopped among
coats conspicuous as a widower
in a grocery line. In this enactment,
white walls scrape easily to reveal
a fuzzy grey almost of cardboard. I
welcome this lightness— no more solid
doors, dark cabinets. I perform my life
inside a pop-up book, every moving part
collapsible, seamed to the center.
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Wisconsin Poetry
iv.
The next day in his absence, I
clean the house, wipe away the traces —
the dust of shed skin, an ear-ring
long lost, thrums from scarves woven
for gifts, and the inevitable
hair— thrums of daily life
unwoven. The house unravels
into a place where I’ve paid
rent. Already, its hallways
darken and merge into
others, each room floats up
disjointed in my mind. My
steps no longer connect them.
v.
In my dream that night, a giraffe
wades leisurely in the wake
of a barge across shallow
waters. The big cats, one of each,
feed from my hands while I wait
in a windowed cubicle for
a ride in some vehicle I
cannot begin to imagine. Far
off, planes dive into the sea
to rise as dolphins, whales
breach and send up a horizon¬
ful of sheep clouds, and the world
spins back into flickers of light
struggling to become the animals.
—Kyoko Mori
125
Wisconsin Academy of Sciences, Arts and Letters
End of Summer
1.
September rain slants into the heavy alfalfa. Water pours from the
square and brutish mouth of the spout and runs away into the boyish grass.
Circles of rust at the bottom of the bucket are thoughts, growing inward, of a
mind simplified by solitude, monotony of rain. . .
2.
Things done or not done for a long time lodge in odors, in old clothes
furrowed with brooding, in soot of bygone cooking black on the widower’s
stove, in the small varnished cross nailed to a bedroom door, and in the dirty
rose curtain on tarnished brass rings, all left as they were when the old man
died.
3.
Did he paint near the end so as not to have to watch night shambling
toward the bams? We find on the obverse of a cornfield scene with pheasants
a far older landscape of lacquered trees and rocks. Implacably symmetrical,
halved precisely by the glass knife of a falls, the left side is green, the right side
ochre, this picture turned toward the wall. . .
— Thomas R. Smith
126
Wisconsin Poetry
A Readiness to Weep
Five years later, we have returned
to the lake where we spent our
honeymoon, two people who knew
even less about marriage than we do.
I had refused for years, then
gave in because I feared some damage
to your happiness with me.
“I did it,” I told you,
“but I don’t know what it means,”
and wept. Today, a day clear
as that one we drove down the dirt
road to the cabin after the wedding,
I squat in these marshy woods
that have always been, in my mind, October
and give again my gift from
five years ago, a readiness to weep.
I sit quietly in the still sunlight
with you and feel the years, and
do not feel them. I let the peach-
flushed leaf I saved fall from my hand —
Yes, I want time to stop for us,
doesn’t everyone? — No, I let it go.
The fallen branch we sat on breaks
suddenly but lets us down unharmed —
the ground isn’t in the least sentimental.
We laugh together, then the tears again.
I soak my knees leaning to cup
in hands chill water from the lake
to splash over my burning eyes.
When I look up, every man
and woman ever married look with me
out across the autumn lake
toward the gold pavilions of uncertainty.
— Thomas R. Smith
127
Wisconsin Academy of Sciences , Arts and Letters
Poems from Paintings
Dream of MOM A
(Rousseau’s Yadwiga’s Dream, 1910)
A confounded round-eyed lion looks straight out
the canvas. She, poised calmly, lying nude
upon a jungle divan hears intrude
a black man garbed in rainbow loincloth, doubt¬
lessly hears melodies to ravish. His
seductive clarinet beguiles the moon,
now rising round and ghostly, begs full blooms
of heavy-headed lotus, blue and reddish
pink, entreats gold- winged, long birds to listen
to his song on silent canvas. Toward
her lifting high his trunk, an elephant
is hidden in the background— leaves that glisten
brightly, edged in yellow, sharp as swords,
precisely painted from Jardin des Plantes.
—Jean Tobin
128
Wisconsin Poetry
Miss Martin at Four O’Clock
I am LaShanda’s teacher
and to LaShanda that is all
I am. Every day she waits
for this, the breakaway hour
when the windows of my eyes
start to blacken behind
the neat rows of paper cutouts
facing the street
and the wide broom of darkness
comes, pushing blood-red
dust along my corridors.
LaShanda thinks I sleep
in a wooden drawer, folded
on a bed of thumbtacks-— my
left hand gripping a bone
of chalk that screams
by day, while my right
brandishes a scarlet
Eversharp, scattering
the swarm of butterflies
that will drift forever
in LaShanda’s head,
-—Marilyn Taylor
129
Wisconsin Academy of Sciences, Arts and Letters
Tercets from the Train
Human dramas implode without trace.
— Marge Piercy
Gorgeous, they are gorgeous, these two women getting
on the train, one in lime green silk, black hair
a mile wide, the other slim as a whip, coiled
in red linen. Their two small boys, grinning,
have squirmed into facing seats, bubbling with spare
energy, the cuffs of their designer jeanlets rolled
at the ankles, their studded shirts glinting.
I overhear the women talking over what to wear
to some convention (should it be the gold
Armani or the St. Laurent?) while the boys are gazing
through the rain- spattered window, practicing their
locomotive lingo in shrill, five-year-old
voices, demanding information: are we going
faster than a plane, where is the engineer,
does this train have electricity or coal?
But the women’s eyes are fierce, they are grumbling
over Lord & Taylor, which was once a store
to be reckoned with, although the one with wild
hair points out that even Bloomingdale’s is growing
unmistakably more K-Martish than it was before.
Don't you ever interrupt me, child,
she hisses to the boy who wonders why the train is grinding
so slowly through the towns, and where
the bathroom is and what the ticket-man is called
until she bends over him, glaring
from beneath her shadowed eyes, a crimson flare
on either cheek. You're interrupting me, she growls.
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Wisconsin Poetry
Now you'll be sorry. His mouth is gaping
as the flat of her pale hand splits the air,
annihilating two long rows of smiles.
I warned you, didn't I, darling?
Now don't you dare cry. Don't you dare.
Up and down the aisle, the silence howls.
— Marilyn Taylor
131
Wisconsin Academy of Sciences, Arts and Letters
Are There Sounds You Shouldn’t Hear?
(A Poem for William Stafford)
Are there sounds you shouldn't hear?
No, not trumpets of god
the walls of cities falling
the lamentations of Jeremiah
the doors closing at Auschwitz
Nor whispers behind my back
nor drill sergeants’ words
the slavers’ whips
newborn whimpers for food in a starved world
The end of history, the voice of Zeus in shrieking wind
the convulsive roar of the Minoan volcano 35 centuries into time
the Aegean tidal wave that pummels ships to death
the screaming horses that trample my children
The conductor dropping his baton
the crack of the firing squad
the drums of Autumn
birds and no singing, like Beethoven and the Song of Joy
The trap slam of King Henry’s scaffold
the avalanche in the High Himalayas
the teeth of the shark that grind my bones
the fall of the dagger in the Aztec Temple
The crash of trees into fire and prairie
the engine that sputters out over oceans
the ice freezing in great cracks round Arctic ships
the man who cries with his tongue cut out
132
Wisconsin Poetry
The gnawing of rats in dark dungeons
the forever farewell from voices of love
the sound boom of the comet that streaks to my feet
the whistling train over the dynamited trestle
The singer breaking glass and ear drums with notes gone cracked
the frenzied still of the oscilloscope beat in the Intensive Care Unit
the suicide striking the pavement
Russian Roulette and no click from the pistol
The rafters smashing in the mine shaft, the end of Welsh and Appalachian songs
the snort of the Cape Buffalo in my frozen face
the flutter of Robert Scott’s tent in the South Polar wind
the screams of the castrati under the knife
The prayers of every prisoner and captured woman
the cries of “Bring out your dead,” in plague-filled Warsaw, and Angkor-wat
the birth of babies, and no breath
the planes, the bombs ripping houses into falling night and screaming rain
No, none, not one
we shall not listen, heed, or obey
but
there is, yes, finally, one sound
I should not hear:
the last sound, the last
vaporized into a shadow on the wall
we shall see that sound as light before we hear it
the last sound, last.
— Laurence T. Giles
133
Wisconsin Academy of Sciences, Arts and Letters
December Lights
Beyond the mall’s warrens of promise
we drive to snow-stilled streets
where trees dance in small lights.
Before every bright fresco, I wonder
how things create lives —
how houses hand out habits
like how much to drink
and when to wake up,
how kitchens coax hearts
to always want more.
If the owner’s away
from the fading grey stucco
and we pass out of our bodies
and through the glass
to a past accrued
on porcelain plates,
would we learn
what a body softens to
when love is traced
on monogrammed sheets
and hands are shaped
by the garden gate?
Or are we bound
in the warm current
of this car
to watch tints of the known
expand
into the opening arms
of the galaxy?
— Joan Rohr Myers
134
Wisconsin Poetry
Flight
(from the notebooks of Leonardo da Vinci)
See tomorrow to all these matters and the copies. Leave them in Florence so that
if you lose those you take with you » the invention will not be lost.
[birds]
The science of birds
is the science of the wind
which is the science of water.
If you would know how things fly,
you must first study
what floats and what falls.
[of man]
The life of birds conforms
better to the needs of flight
than the will of man,
especially in the almost
imperceptible movements which
preserve an equilibrium.
[with drawings ]
spring of horn
of steel fastened upon wood,
of willow encased in reed—
Let A be the first movement.
Undo one and remove.
Double canes- soaped .
of rag, of skin, of flying fish.
spring with lock,
wire that holds the spring,
spring of wing—
Tomorrow morning
the second of January
I’ll make the thong and the attempt.
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Wisconsin Academy of Sciences, Arts and Letters
[ in which the figure of the man is seen
exerting force with arms and legs]
If you stand up on the roof at the side of the tower
the men at work on the cupola will not see you.
The machine should be tried over a lake
and you should carry a large inflated wineskin so
if you fall you will not drown.
Let the machine be 12 braccia high
and let the span of the wings
be 40 braccia
and the body from stem to prow
20 braccia
and the outside all covered
with cane and with cloth —
Ladder for ascending and descending.
[the atmosphere]
The air moves like a river
and carries the clouds with it
just as moving water carries
all things that float upon it.
Surface is the name
of that division which the body
makes with the bodies it encloses.
It does not partake
of the body which surrounds it,
or of the body which it surrounds.
Surface has a name
but not a substance
for that which has
substance has place.
136
Wisconsin Poetry
[words crossed out in manuscript]
Just as a stone thrown
into water becomes the center
and the cause of various circles,
so a motion made in the air
spreads itself out in circles.
So every body in the luminous air
spreads itself out in circles
and fills the sky with
infinite images of itself.
— Bruce Taylor
137
Wisconsin Academy of Sciences, Arts and Letters
Breakfast
After feet find the dark floor
after the pulling-on of socks and shirt and pants
after building up the fire with oak and breath
and putting the kettle on
there’s the sizzle of cold potatoes browning
as tomato’s upstart bite jangles the mouth
and chunks of yesterday’s ham color the potatoes’ hiss
and a brighteyed egg tops it off
with a sprinkle of salt for savor.
Then coffee in a chair
while the red comes up behind the horizon
past the silhouettes of hill and branch
that turn to gray and then
to glowing brown detail:
bark and leaf and blade of grass.
The empty cup is the origin of philosophy
as is the end of this dawn
fallen to the dingy day incapable
of this gray illumination.
— Ralph Schneider
138
First Report of Natural Bridges
in Eastern Wisconsin
Richard A. Pauli
Abstract. Although natural bridges are well-known features in the Driftless Area of Wisconsin
and adjacent states , none of these delicate landforms is documented in the recently glaciated
region of Wisconsin. This report describes natural bridges at two localities along the Silurian
escarpment in eastern Wisconsin.
A bridge 40 feet (12 m) high with a span of 14.5 feet (4.4 m) has developed in the Lower
Silurian Mayville Dolomite at Fonferek Glen in Brown County, 4.3 miles (7 km) south of
Green Bay. This feature was created in two stages. The first was differential undercutting of
bedrock on the outside of a meander at a higher elevation than present-day Bower Creek
during the waning stage of glaciation in late Wisconsinan time. Collapse of the inner part of
the overhang along prominent joints in the Holocene completed the bridge.
Six bridges have developed at the contact between the Lower Silurian Mayville Dolomite
and the overlying Middle Silurian Byron Dolomite at Oakfield Ledges in Fond du Lac County,
10.5 miles (17 km) east ofWaupun. The bridges vary in dimensions from 15 feet (4.5 m) to
20 feet (6 m) high, 3.3 feet (1 m) to 15 feet (4.5 m) wide, and 4 feet (1 .2 m) thick, with spans
of 4 feet (1.2 m). These features formed by the opening of three solution-enlarged joints in
the Mayville Dolomite by downslope movement at the escarpment after retreat of the Green
Bay glacial lobe. The Byron Dolomite remained in place as the underlying Mayville moved
outward. This created a roof of Byron Dolomite for these bridges. It is possible that the
present-day bridges along each joint were once part of cave-like features.
Natural bridges are common in the Drift¬
less Area of Wisconsin and parts of
immediately adjacent states (Fig. 1). Ac¬
cording to Martin (1932, 353), “All of these
features ... are of the sort that can exist only
in the Driftless Area. They are relatively fragile
and would certainly have been eroded away
or buried in glacial deposits.” The one ex¬
ception known prior to this paper is a small
natural bridge in Middle Ordovician dolomite
at Krape Park, Freeport, Illinois (Pauli and
Pauli, 1980, 85) (Fig. 1). The genesis of this
Richard A. Pauli is Professor of Geosciences at the
University of Wisconsin-Milwaukee, where he has taught
since 1962. Prior to that time, he worked for Jersey
Production Research Company, now Exxon.
bridge is similar to the developmental history
for the one at Fonferek Glen described below.
This report describes seven natural bridges
developed in dolomite along the Silurian es¬
carpment in the recently glaciated area of
eastern Wisconsin. One of these bridges is
in Fonferek Glen south of Green Bay, Brown
County, Wisconsin (Fig. 1). I first observed
this feature while studying Ordovician and
Silurian rocks in this area in 1967. Donn P.
Quigley of the Neville Public Museum, Green
Bay (verbal communication 1988), had dis¬
covered this bridge earlier in the 1960s.
At least six natural bridges are present in,
or adjacent to, the Wisconsin Department of
Natural Resources Oakfield Ledges Scientific
Area in southern Fond du Lac County (Fig. 1).
A descriptive article about this scenic locality
139
Wisconsin Academy of Sciences, Arts and Letters
bridges in Wisconsin and adjacent parts of
Iowa and Illinois. Most of these delicate land-
forms are within the Driftless Area (HN and
LH, Hornets Nest and Luncheon Hall, Martin
1932; MC, Maquoketa Caves, Pauli and Pauli
1980; MV A, Mt. Vernon Arch, Martin 1932;
NBSP, Natural Bridge State Park, Martin 1932;
Pauli and Pauli 1977; RA and ER, Rockbridge
Arch and Elephant Rock, Martin 1932; Pauli
and Pauli, 1980; and RMA, Readstown Mul¬
tiple Arch, Martin 1932), but several occur
within the recently glaciated region (FG, Fon-
ferek Glen Bridge; KP, Krape Park Bridge,
Pauli and Pauli 1980; OL, Oakfield Ledges
Bridges). The bridges in eastern Wisconsin
(FG and OL) are described in this paper.
by Peter Toepfer (1979) suggested that nat¬
ural bridges might be present here, and this
proved to be correct.
When I started this project, I thought I
knew what a natural bridge is. The deeper I
have delved into the problem, the more un¬
certain I have become. The Glossary of Ge¬
ology (Bates and Jackson 1980, 442) defines
a natural bridge as an arch-like rock forma¬
tion created by erosion that spans a drainage,
or the remnant of the partial collapse of the
roof of a cave. The water becomes muddier
when the editors referenced above define an
arch as a natural bridge resulting from ero¬
sion, or a landform similar to a natural bridge
not formed by erosive agencies. Leading
geomorphology textbooks fail to clarify the
issue. One thing that all features called bridges
and arches have in common is a relatively
resistant uppermost lithologic unit that forms
the span. It is also apparent that these delicate
features are short-lived geologic landforms.
In this paper I define a natural bridge as a
free-standing rock formation that allows hu¬
man passage across a relatively narrow span.
Such features could result from either erosion
or selective gravity-induced movements or a
combination of the two processes. Specifi¬
cally excluded, however, are down-dropped
blocks of rocks that result in “bridging” across
openings (usually joints). Examples of this
type of feature are Devil’s Doorway at Dev¬
il’s Lake State Park (Pauli and Pauli 1977,
127) and several rock fall “bridges” at Oak-
field Ledges.
Fonferek Glen Natural Bridge
A single natural bridge occurs in the Lower
Silurian Mayville Dolomite along Bower
Creek in Fonferek Glen, about 800 feet
(240 m) downstream from Fonferek Falls
(Polish Falls in older literature) (Fig. 2 and
3). The glen is a narrow, deeply cut reentrant
in the east-west trending Silurian escarp¬
ment. This locality is 1.15 miles (1.85 km)
east of the crossroad community of Kolb along
County Highway MM, and about 4.3 miles
(7 km) south of Green Bay in Brown County
(SW!/4, SElA, NElA, NWj/4, Sec. 34, T.23N.-
R.21E., Bellevue 7.5' quadrangle, 1954)
(Fig. 2). The Fonferek natural bridge is 40
feet (12 m) high, 5 feet (1.5 m) wide, and 5
feet (1.5 m) thick. It spans 14.5 feet (4.4 m)
at the top and opens to more than 40 feet
(12 m) in the erosional alcove below (Fig. 4
and 5).
The genesis of the natural bridge at Fon¬
ferek Glen has a long geologic history. The
140
Natural Bridges in Eastern Wisconsin
Figure 2. Location map of Fonferek Glen Natural Bridge in Brown County, Wisconsin. Fonferek
Falls, at the head of the glen, is a southerly reentrant cut by Bower Creek into the Silurian
escarpment, which trends east-west across the northern part of the map area.
Lower Silurian Mayville Dolomite at this lo¬
cality consists of three distinctive lithologic
units that are described in Figure 5. The up¬
per and lower units are resistant dolomite,
and the uppermost forms Fonferek Falls and
the span of the natural bridge. The middle
unit is a relatively nonresistant, chert-rich
dolomite that weathers more readily than the
overlying and underlying rock.
The Lower Silurian bedrock surface at
Fonferek Glen was polished and striated by
southerly moving glacial ice that deposited
reddish till of the Glenmore Member of the
Kewaunee Formation in late Wisconsinan
(Greatlakean) time (Need 1985, 1). The gently
rolling till plain in this area is locally overlain
by deposits that accumulated in Glacial Lake
Oshkosh, an impoundment that formed in the
Green Bay lowland behind the northeasterly
retreating ice dam.
Fonferek Glen developed when the late
Wisconsinan ice retreated far enough to allow
Glacial Lake Oshkosh to drain easterly into
ancestral Lake Michigan. Rapidly falling lake
levels allowed an earlier version of Bower
Creek to downcut across the Silurian escarp¬
ment. At this time, precipitation rates were
apparently high, and Bower Creek carried
significant amounts of runoff. As erosion
proceeded, the waterfall that initially was at
the edge of the Silurian escarpment retreated
upstream, leaving a narrow gorge behind.
When Bower Creek eroded downward
into the chert-rich middle unit of the Lower
141
Wisconsin Academy of Sciences, Arts and Letters
Figure 3. Fonferek Falls plunges over Lower
Silurian Mayville Dolomite at the head of Fon¬
ferek Glen.
Silurian Mayville Dolomite, it carved two
caves in this relatively erodible rock at the
outside of meanders in the canyon below the
waterfall (Fig. 6 and 7). At the upstream
meander an eddy current developed, and a
third cave formed (Fig. 6). As downcutting
continued into the basal resistant unit of the
Mayville, the bedrock valley narrowed.
Two sets of joints in the upper unit of the
Mayville Dolomite facilitated frost wedging,
and rock falls enlarged the eddy-formed cave
to create a large alcove. In the middle 1950s
two horses strayed onto the prominent over¬
hang and broke through the roof to create the
natural bridge (Norbert Fonferek, verbal
communication 1988) (Fig. 8). Both horses
were pulled free, and the opening was fenced
off for safety reasons. Joint blocks continue
to fall, and ultimately Fonferek Glen Natural
Bridge will collapse.
Figure 4. Fonferek Natural Bridge spans an alcove carved by fluvial erosion in the relatively
nonresistant, chert-rich, middle unit of the Lower Silurian Mayville Dolomite. The span of this
feature is formed from the resistant, upper unit of the Mayville Dolomite. The well-jointed nature
of the roof rock is apparent in the photo.
142
Natural Bridges in Eastern Wisconsin
PLEIST.
7? O
_i <
29’
13’
0’
DIAGRAMMATIC SKETCH OF FONFEREK GLEN NATURAL BRIDGE
Figure 5.
locality.
uumssssr*
A diagrammatic sketch of Fonferek Natural Bridge detailing the geology at this
Figure 6. A schematic sketch detailing the development of three caves in Fonferek Glen during
high stream flow at an earlier erosional level of Bower Creek.
143
Wisconsin Academy of Sciences, Arts and Letters
Figure 7. This cave was developed in the relatively nonresistant, chert-rich, middle unit of the
Lower Silurian Mayville Dolomite at the outside of the second meander downstream from
Fonferek Falls (Fig. 6). This feature is a precursor of the type of opening that developed into
the Fonferek Glen Natural Bridge upstream from this locality. A well-developed, open joint is
also visible to the left of the cave. Jointing facilitated the formation of the natural bridge in this
area.
Figure 8. A downward view of the Fonferek Natural Bridge from the top of the Silurian escarpment.
144
Natural Bridges in Eastern Wisconsin
Oakfield Ledges Natural Bridges
Six natural bridges occur within three joints
along the southwest-northeast trending Si¬
lurian escarpment in, and adjacent to, the
Wisconsin Department of Natural Resources
Oakfield Ledges Scientific Area. This local¬
ity is about 10.5 miles (17 km) east of
Waupun in the WV2, NWV4, SW!4, Sec. 27,
T. 14N.-R. 16E. , Fond du Lac County (Wau¬
pun 15' quadrangle, 1955) (Fig. 9). The
bridges range from 15 feet (4.5 m) to 20 feet
(6 m) high, 3 feet (1 m) to 15 feet (4.5 m)
wide, and 4 feet (1.2 m) thick, with spans
of 4 feet (1.2m). Other bridges may also be
present along joints in the talus-mantled,
westward-facing Silurian cliff face in this
general area.
The Silurian escarpment at Oakfield Ledges
consists of medium- to massive-bedded Lower
Silurian Mayville Dolomite and the overlying
thin- to medium-bedded, relatively resistant,
Middle Silurian Byron Dolomite (Shrock 1939;
Figure 9. Location map of Oakfield Ledges,
part of the Silurian escarpment in Fond du
Lac County, Wisconsin.
Mikulic and Klussendorf 1983) (Fig. 10).
These rocks dip gently eastward into the
Michigan Basin.
Silurian bedrock in this region was pol¬
ished and striated by the southerly moving
Green Bay glacial lobe in late Wisconsinan
time. The thin, reddish, bouldery till of the
Horicon Formation was also deposited during
this glaciation (Mickelson et al. 1984). This
lobe, and previous ice advances, scoured the
relatively nonresistant shale of the Upper Or¬
dovician Brainard Formation of the Ma-
quoketa Group to create the lowland region
west of the Silurian escarpment now occu¬
pied by Green Bay, Lake Winnebago, and
Horicon Marsh.
Upon retreat of glacial ice from the Green
Bay lowland, the Silurian dolomite overlying
the relatively soft shale of the Brainard For¬
mation was unsupported. Rock falls from the
cliff face and rotational downslope move¬
ment of Silurian rock along the Brainard sur¬
face were facilitated by solution-enlarged joints
that generally parallel the cliff face (Fig. 11).
This rotational type of slope failure along the
Silurian escarpment in Brown County was
previously detailed by Stieglitz (1980, 83).
The unique aspect of the downslope move¬
ment of jointed masses of Silurian rock at
Oakfield Ledges is the differential separation
along individual joints in the Mayville and
Byron formations (Fig. 12). This allowed the
Mayville to move along the unstable Brainard
surface, while part of the Byron remained in
place to bridge the gaps over the enlarging
joints (Fig. 13). This created caves along three
solution-enlarged joints. Selective collapse
of the roof rock created a series of three bridges
in one joint and two in another. A single
bridge is present along the third joint.
The natural bridges at Oakfield Ledges are
unusual features, but they meet the criteria
previously defined for natural bridges. As
downslope movement continues along these
major joints within the Mayville Dolomite,
the bridging roof rock of the Byron Dolomite
will become unsupported. Collapse will fol¬
low, and the natural bridges at Oakfield
Ledges, as well as at Fonferek Glen, will no
longer enrich the Wisconsin landscape.
145
Wisconsin Academy of Sciences, Arts and Letters
Figure 10. A schematic sketch defining the geology of Oakfield Ledges and illustrating the
formation of the natural bridges at this locality.
Figure 11. View along a major joint essentially parallel to the Silurian escarpment at Oakfield
Ledges shows the rotational down slope movement of a major mass of dolomite. The over¬
hanging rock at the top of the rotated block is the Middle Silurian Byron Dolomite.
146
Natural Bridges in Eastern Wisconsin
Figure 12. Overhang of the thin- to medium-bedded Middle Silurian Byron Dolomite across
an enlarged joint in the Lower Silurian May vi lie Dolomite at Oakfield Ledges.
Figure 13. View along a major joint at Oakfield Ledges illustrates bridging by the Middle
Silurian Byron Dolomite.
147
Wisconsin Academy of Sciences, Arts and Letters
Acknowledgments
The following individuals generously pro¬
vided information and advice during this study:
Genevieve Bancroft, Norbert Fonferek, Joanne
Klussendorf, Donald G. Mikulic, Meredith
Ostrom, Donn P. Quigley, and Ronald D.
Stieglitz. Field investigations and prepara¬
tion of the manuscript were aided by Rachel
K. Pauli.
Works Cited
Bates, R.L., and J.A. Jackson, eds. 1980. Glos¬
sary of geology. 2nd ed. Falls Church: Amer¬
ican Geological Institute.
Martin, Lawrence. 1932. The physical geography
of Wisconsin. Wis. Geol. Nat. Hist. Surv. Bull.
no. 36.
Mickelson, D.M., L. Clayton, R.W. Baker, W.N.
Mode, and A.F. Schneider. 1984. Pleistocene
stratigraphic units of Wisconsin. Wis. Geol. Nat.
Hist. Surv. Misc. Paper 84-1.
Mikulic, D.G., and J. Klussendorf. 1983. The
oolitic Neda iron ore (Upper Ordovician) of
eastern Wisconsin. Field Trip Guide Book for
the 17th Annual Meeting of the North-Central
Section, Geological Society of America. Wis.
Geol. Nat. Hist. Surv.
Need, E.A. 1985. Pleistocene geology of Brown
County, Wisconsin. Wis. Geol. Nat. Hist. Surv.
Informational Circular 48.
Pauli, R.K., and R.A. Pauli. 1977. Geology of
Wisconsin and Upper Michigan; including parts
of adjacent states. Dubuque: Kendall/Hunt
Publishing Co.
Pauli, R.K., and R.A. Pauli. 1980. Wisconsin and
Upper Michigan: Geology field guide. Du¬
buque: Kendall/Hunt Publishing Co.
Shrock, R.R. 1939. Wisconsin Silurian bioherms
(organic reefs). Bull. Geol. Soc. Amer.
50:529-62.
Stieglitz, R.D., ed. 1980. Guidebook for the 44th
Annual Tri-State Geological Field Conference:
Geology of northeastern Wisconsin.
Toepfer, P. 1979. The Oakfield Ledges: A scenic
inheritance of the ice age. Wis. Nat. Re sour.
Bull. 3:12-13.
148
Live Capture Methods of
Sympatric Species of Flying Squirrel
Thomas C. Engel, Michael J. Lemke, and Neil F. Payne
Standard methods of capturing other tree
squirrels are not as effective for flying
squirrels, which spend proportionately less
time foraging on the ground (Sollberger 1940;
Sonenshine et al. 1979). They can be cap¬
tured in natural or artificial dens (Sonenshine
et al. 1973) or in Sherman live traps attached
to trees (Sonenshine et al. 1979). Sumner
(1927) captured flying squirrels with rat (kill)
traps nailed in trees. Burt (1927, 1940), Jack-
son (1961), Sonenshine et al. (1979), and
Mowrey and Zasada (1984) reported that traps
set in trees are effective but did not present
trapping details or trap sympatric species of
flying squirrels. The objective of this study
was to determine the trapping success for
sympatric species of flying squirrel relative
to tree species, trap type, and height of trap
in tree.
The study area was the 83-ha Schmeeckle
Reserve, University of Wisconsin-Stevens
Thomas Engel graduated from the University of Wis¬
consin-Stevens Point in 1980 with an M.S. in wildlife.
Now based in Grand Rapids, he is the Forest Wildlife
Coordinator for northeastern Minnesota.
Michael Lemke graduated with a B.S. degree in natural
resources from the University of Wisconsin-Stevens Point
in 1980 and an M.S. in zoology from the University of
British Columbia-Vancouver in 1985. He currently is
a doctoral candidate in the Department of Biological
Sciences at Michigan Technological University studying
aquatic food chains.
Neil F. Payne teaches wildlife courses at the University
of Wisconsin-Stevens Point. He received a B.A. in bi¬
ology from the University ofWisconsin-Madison in 1961,
an M.S. in wildlife from Virginia Polytechnic Institute
and State University in 1964, and a Ph.D. in wildlife
from Utah State University in 1975.
Point, an area within the vegetational tension
zone (Curtis and McIntosh 1951; Curtis 1959)
that includes plants and animals typical of
both the prairie and boreal forest ecotone ex¬
tending northwest-southeast in Wisconsin.
Forest composition was 5.7 ha of mixed
hardwoods including oak ( Quercus spp.),
maple ( Acer spp.), elm ( Ulmus spp.), white
birch (Betula papyrifera), and quaking aspen
(Populus tremuloides); 14.3 ha of pine (Pi-
nus strobus, P. banksiana, P. resinosa);
15.6 ha of mixed woods containing mature
hardwoods and scattered mature white pine;
and 8.9 ha of oak savanna (Engel 1980).
Methods
To test trap type, we used wooden box
traps (Mosby 1955 in Day, Schemnitz, and
Taber 1980), Sherman sheet metal box traps
7.5 X 7.5 x 26 cm (Sonenshine et al. 1979)
and 13 X 13 X 45 cm, and Havahart wire
cage traps 13 X 13 x 45 cm; all were baited
with peanut butter. In 1978 we added tree
traps to three traplines consisting of wooden
box traps set on the ground 30 m apart, which
had produced 0.2 flying squirrels per 100
trapnights in 1977. Because the size and weight
of the wooden box traps made them awkward
to secure in trees, we set Sherman and
Havahart traps 60 m apart in trees next to the
ground traps. Trees for trap placement were
selected for convenience to the trapline; height
of trap placement was determined by the ease
of climbing without spikes. Trap heights were
grouped as 0 (ground), 1-3. 1 m, and >3.1 m.
Tree species of trap placement were pooled
as red maple (Acer rubrum), other hard¬
woods, jack pine (Pinus banksiana), red pine
149
Wisconsin Academy of Sciences, Arts and Letters
(P. resinosa), and white pine (P. strobus).
Traps were secured in trees with a rubber
band cut from an auto tire inner tube and
looped around one end of the trap and passed
beneath a branch, or around the tree trunk of
smaller trees and looped over the opposite
trap end. When large branch size precluded
this method, the rubber band was cut and
extended with a short length of light rope.
Traplines were operated 24 September-3
October, 10-20 October, and 22 October-2
November 1978 in three locations within the
study area and checked at dawn. Flying
squirrels were sexed, tagged in each ear with
a numbered aluminum fingerling tag No. 1
(National Band and Tag Co. , Newport, KY),
and released. After two flying squirrels died
in traps early in the study, we added shredded
wood packing materials and/or cotton to the
tree traps to provide insulation and help ab¬
sorb moisture from respiration. We used log-
linear models (Fienberg 1980) to analyze
capture data.
Results
In 1978 we captured 13 G. volans and 14
G. sabrinus in live traps 39 and 31 times,
respectively. Tree sets were nearly sixteen
times more effective than ground sets for
capturing flying squirrels ( X2 - 300.3, p <
.001, df = 2) (Table 1). No difference ex¬
isted (p > .05) in capture rates of traps set
1-3. 1 m from the ground and >3.1 m. Small
sample sizes precluded statistical compari¬
sons of the ease of trapping the two species,
although G. sabrinus seemed slightly more
predisposed to ground traps (Table 1).
The number of flying squirrels trapped per
100 trapnights (Nelson and Clark 1973) in
trees was 15.4 in wire traps, 10.8 in large
Sherman traps, and 10.2 in small Sherman
traps. These catch rates are not different, al¬
though wire traps might have been superior
had sample sizes been larger. No squirrels
died in the wire cage traps; squirrel mortality
was 10% in the other traps. No difference
existed in survival due to squirrel species or
trap type (G2 = 6.89, p = .44, df = 7),
but sample size was small.
We caught flying squirrels in all species
of trees (Table 2). The best loglinear model
(G2 = 6.33, p = .90, df = 12) indicates
that the tree species in which traps were placed
was more important than trap type or trap
height in determining capture rates. Indica¬
tions are that the most successful combina¬
tion of trap and tree used was a wire trap set
in a white pine (Table 2).
Discussion
The lack of differences among trap types
and the importance of ground versus tree
placement suggest that wooden traps set in
trees would be effective and that other types
of traps set on the ground would not, al¬
though study design precluded testing these
combinations. The relatively high capture rates
of both species in white pines in our study
area (Table 2) might not reflect habitat pref¬
erence or tree species as much as size of tree,
especially for G. volans, because pines were
available in all habitat types and were the
biggest trees. Most white pines in the study
area were taller than the forest canopy. Post-
Table 1. Trapping success for flying squirrels*
‘University of Wisconsin-Stevens Point, September-November 1978.
Adjusted for sprung traps (Nelson and Clark 1973).
150
Live Capture of Flying Squirrels
Table 2. Trapping efficiency for flying squirrels for traps set in various tree species*
‘Three types of trap were used, but there was no difference in catch rates. University of Wisconsin-Stevens
Point, September-November 1978.
Adjusted for sprung traps (Nelson and Clark 1973).
release observations of G. volans and G. sa-
brinus indicated that squirrels choose the most
direct route to a large (> 40- cm diameter at
breast height [dbh]) mature tree. Squirrels
climbed only 2-3 m up smaller (12-30 cm
dbh) trees before gliding to the base of an¬
other small tree in a direct route to a large
white pine or oak. Squirrels climbed to can¬
opy height and glided longer distances only
from large trees. Trees > 40 cm dbh typi¬
cally were selected as targets for glides ini¬
tiated at canopy heights. Sonenshine and Levy
(1981) and Ando and Imaizami (1982) also
found strong positive associations with ex¬
treme height and gliding. Bendel and Gates
(1987) suggested that trees > 40 cm dbh and
open upper-understory (> 10-15 m) aid
locomotion and escape, and that clearcuts
> 75 m wide are barriers. Mowrey and
Zasada (1984) suggested clearcuts not be
> 40 m wide, with < 20 m preferable.
Acknowledgment
D. Heisey provided statistical analysis and
help with interpretation of data.
Works Cited
Ando, M., and Y. Imaizami. 1982. Habitat uti¬
lization of the white-cheeked giant flying squir¬
rel (Petaurista leucogenys) in a small shrine
grove. J. Mammal. Soc. Japan 9:70-81.
Bendel, P. R., and J. E. Gates. 1987. Home range
and microhabitat partitioning of the southern
flying squirrel Glaucomys volans. J. Mammal.
68:243-55.
Burt, W. H. 1927. A simple live trap for small
animals. J. Mammal. 8:302-9.
- . 1940. Territorial behavior and popula¬
tions of some small mammals in southern Mich¬
igan. Misc. Publ. Mus. Zool. Univ. Mich.
45:1-58.
Curtis, J. T. 1959. The vegetation of Wisconsin.
Madison: University of Wisconsin Press.
Curtis, J. T., and R. P. McIntosh. 1951. An up¬
land forest continuum in the prairie-forest bor¬
der of Wisconsin. Ecology 32:476-96.
Day, G. I., S. D. Schemnitz, and R. D. Taber.
1980. Capturing and marking wild animals. In
Wildlife management techniques manual. 4th
ed., ed. S. D. Schemnitz, 61-88. Washington:
Wildlife Society.
Engel, T. C. 1980. Effects of urban development
on vertebrate wildlife populations in Schmeec-
kle Reserve, University of Wisconsin-Stevens
Point. M.S. thesis, University of Wisconsin-
Stevens Point.
Fienberg, S. E. 1980. The analysis of cross-
classified categorical data. Cambridge: MIT
Press.
Jackson, H. H. T. 1961. The mammals of Wis¬
consin. Madison: University of Wisconsin Press.
Mowrey, R. A., andJ. C. Zasada. 1984. Den tree
use and movements of northern flying squirrels
in interior Alaska and implications for forest
management. In Fish and wildlife relationships
in old growth forests, ed. W. R. Meehan, T. R.
Merrell, Jr. , and T. A. Hanley. Morehead City,
N.C.: Am. Inst. Fish Res. Biol.
Nelson, L., Jr., and F. W. Clark. 1973. Correc¬
tion for sprung traps in catch/effort calculations
of trapping results. J. Mammal. 54:295-98.
Sollberger, D. E. 1940. Notes on the life history
of the small eastern flying squirrel. J. Mammal.
21:282-93.
151
Wisconsin Academy of Sciences, Arts and Letters
Sonenshine, D. E., D. G. Cerrentani, G. Enlow,
and B. L. Elisberg. 1973. Improved methods
for capturing wild flying squirrels. J. Wildl.
Manage. 37:588-90.
Sonenshine, D. E., D. M. Lauer, T. C. Walker,
andB. L. Elisberg. 1979. The ecology of Glau-
comys volans (Linnaeus, 1758) in Virginia. Acta
Theriot. 24:363-77 .
Sonenshine, D. E., and G. F. Levy. 1981. Veg¬
etative associations affecting Glaucomys volans
in central Virginia. Acta Theriot. 26:359-71.
Sumner, E. L. 1927. Notes on the San Bemadino
flying squirrel. J. Mammal. 8:315-16.
152
Range Extension of Northern Flying Squirrels
Thomas C. Engel, Michael J. Lemke, and Neil F. Payne
While trapping small mammals in Ste¬
vens Point, Portage County, Wiscon¬
sin, we examined a northern flying squirrel
( Glaucomys sabrinus) collected on the Uni¬
versity of Wisconsin-Stevens Point campus
in 1976. In 1977 in the same area we captured
an adult female northern flying squirrel in a
Museum Special snap trap set on the ground
for small mammals. Dr. C. Long, museum
curator at the university, identified and re¬
tained the specimen (no. 4927). This evi¬
dence extends the known range of the north¬
ern flying squirrel south of the previously
known range, into Portage County, Wisconsin.
Our study area was the 83-ha Schmeeckle
Reserve, University of Wisconsin-Stevens
Point, an area within the vegetational tension
zone (Curtis and McIntosh 1951; Curtis 1959)
that includes plants and animals typical of
both the prairie and boreal forest ecotone ex¬
tending northwest-southeast in Wisconsin.
Forest composition was 5.7 ha of mixed
hardwoods including oak ( Quercus spp.),
maple (Acer spp.), elm ( Ulmus spp.), white
birch (Betula papyrifera), and quaking aspen
(Populus tremuloides); 14.3 ha of pine (Pi¬
rns strobus, P. banksiana, P. resinosa);
15.6 ha of mixed woods containing mature
hardwoods and scattered mature white pine;
and 8.9 ha of savanna (Engel 1980).
Our population estimates from live trap¬
ping (Overton 1965; Davis and Winstead 1980)
were 17 ± 2.5 southern flying squirrels (0.4
per ha) and 14 ± 2.8 northern flying squirrels
(0.3 per ha). Density of southern flying squir¬
rels in Virginia was 31-38 per ha (Sawyer
and Rose 1985); for northern flying squirrels
in Alaska density was 0.3 per ha (Mowrey
and Zasada 1984). The lower population es¬
timates of southern flying squirrels in our
study area, where sympatry occurred, might
be due to limited availability of large trees
for dens, suitable understory (Sonenshine and
Levy 1981; Bendel and Gates 1987), and
food in this type of presumably marginal hab¬
itat normally associated with range limitation.
A broad zone of potential sympatry of
northern flying squirrels and southern flying
squirrels exists in North America, coinciding
with northern hardwood or mixed vegetation
(Hall and Kelson 1959). But little actual
overlap in the ranges of the two species of
flying squirrel exists, with little evidence of
sympatry due to highly variable and often
exclusive niches (Weigl 1978). In Wiscon¬
sin, records of sympatry exist in Jackson
(Rausch and Tiner 1948), Clark (Jackson
1961), and now Portage counties, and in the
Upper Peninsula of Michigan (Stormer and
Sloan 1976). The potential zone of sympatry
in Wisconsin comprises the tension zone
(Curtis and McIntosh 1951; Curtis 1959)
within which Jackson, Clark, and Portage
counties occur. Sympatry of northern flying
squirrels and southern flying squirrels is likely
in other counties within the tension zone.
We found northern flying squirrels almost
exclusively in pine habitat and southern flying
squirrels mostly in mixed woods but also in
deciduous habitat. Weigl (1978) also found
northern flying squirrels associated with con¬
ifers and southern flying squirrels with de¬
ciduous or mixed woods in North Carolina,
where altitude influences habitat. Much (67%)
of our study area is pine or a mixture of oak
and pine, which Sonenshine and Levy (1981)
found southern flying squirrels to use less
than lowland deciduous areas. Wells-Gosling
153
Wisconsin Academy of Sciences, Arts and Letters
(1985) compiled a list of habitat types in
North America occupied by both species of
flying squirrel.
The northern range of mast trees limits
distribution of southern flying squirrels (Weigl
1978). Both species are omnivores, but
southern flying squirrels eat mainly mast in
winter, while northern flying squirrels eat fungi
and the abundant lichens which most animals
do not eat, resulting in an exclusive energy
source for northern flying squirrels and little
competition for food (Weigl 1978). Also,
northern flying squirrels feed on cached fungi
in red squirrel (Tamiasciurus hudsonicus)
middens (Mowrey and Zasada 1984); both
species generally are found together in con¬
iferous forests. Population densities were 0.4
per ha for red squirrels and 1.4 per ha for
gray squirrels (Sciurus carolinensis) in the
study area.
Southern flying squirrels use only tree cav¬
ities for dens (Weigl 1978). They are not
hibemators, and den in aggregations for
warmth in winter (Muul 1968). Northern flying
squirrels are larger, more thickly furred, and
thus more tolerant of cold temperatures. They
use tree cavities and outside nests. More tree
cavities are available in deciduous than con¬
iferous forests. Although smaller, southern
flying squirrels are more aggressive in de¬
fending a home range. When both species
occupy a deciduous forest, southern flying
squirrels can displace northern flying squir¬
rels into less suitable habitat, thus possibly
reducing their reproductive success (Weigl
1978).
Works Cited
Bendel, P. R., and J. E. Gates. 1987. Home range
and microhabitat partitioning of the southern
flying squirrel (Glaucomys volans). J. Mam¬
mal. 68:243-55.
Curtis, J. T. 1959. The vegetation of Wisconsin.
Madison: University of Wisconsin Press.
Curtis, J. T., and R. P. McIntosh. 1951. An up¬
land forest continuum in the prairie-forest bor¬
der of Wisconsin. Ecology 32:476-96.
Davis, D. E., and R. L. Winstead. 1980. Esti¬
mating the numbers of wildlife populations. In
Wildlife management techniques manual, ed.
S. D. Schemnitz, 221-45. Washington: Wild¬
life Society.
Engel, T. C. 1980. Effects of urban development
on vertebrate wildlife populations in Schmeec-
kle Reserve, University of Wisconsin-Stevens
Point. M.S. thesis, University of Wisconsin-
Stevens Point.
Hall, E. R., and K. R. Kelson. 1959. The mam¬
mals of North America. Vol. 1. New York:
Ronald Press.
Jackson, H. H. T. 1961. The mammals of Wis¬
consin. Madison: University of Wisconsin Press.
Mowrey, R. A., and J. C. Zasada. 1984. Den tree
use and movements of northern flying squirrels
in interior Alaska and implications of forest
management. In Fish and wildlife relationships
in old growth forests, ed. W. R. Meehan, T. R.
Merrell, Jr. , and T. A. Hanley. Morehead City,
N.C.: Am. Inst. Fish. Res. Biol.
Muul, I. 1968. Behavioral and physiological in¬
fluences on the distribution of the flying squir¬
rel, Glaucomys volans. Misc. Publ. Mus. Zool.
Univ. Mich. 134:1-66.
Overton, W. S. 1965. A modification of the
Schnabel estimator to account for removal of
animals from the population. J. Wildl. Manage.
29: 392-95.
Rausch, R., and J. D. Tiner. 1948. Studies of the
parasitic Helminths of the north central states.
I. Helminths of Sciuridae. Am. Midi. Nat. 39:
729-47.
Sawyer, S. L., and R. K. Rose. 1985. Homing
in and ecology of the southern flying squirrel
Glaucomys volans in southeastern Virginia USA.
Am. Midi. Nat. 113:238-44.
Sonenshine, D. E., and G. F. Levy. 1981. Veg¬
etative associations affecting Glaucomys volans
in central Virginia. Acta Theriol. 26:359-71.
Stormer, F. A., and N. Sloan. 1976. Evidence of
the range extension of the southern flying squir¬
rel in the upper peninsula of Michigan. Jack-
Pine Warbler 54:176.
Weigl, P. D. 1978. Resource overlap, interspe¬
cific interactions and the distribution of the flying
squirrels (Glaucomys volans and G. sabrinus).
Am. Midi. Nat. 100:83-96.
Wells-Gosling, N. 1985. Flying squirrels. Wash¬
ington: Smithsonian Institution Press.
154
The Modern Spiritual Condition and
the Ancient Wisdom of the I Ching
Claire E. Matthews
We are in a period of religious crisis,
Mircea Eliade tells us. Many ele¬
ments of modem life are attempts to recover
the sacredness of life and nature and to re¬
cover the religious dimension of authentic
and meaningful ‘ ‘human existence in the cos¬
mos.” As evidence, Eliade cites the contem¬
porary rediscovery of nature, uninhibited
sexual mores, and emphasis on “living in
the present.” He points to these as creative
and therefore unrecognizable answers to the
crisis and expressions of potentially new ex¬
periences of the sacred (Eliade 1969, preface).
Joseph Campbell states that “one of our
problems today is that we are not well ac¬
quainted with the literature of the spirit”
(Campbell and Moyers 1988, 3). Myths are
stories that give us a perspective on what is
happening to us. We have lost the function
of myth in contemporary society, and there
is nothing comparable to take its place. Themes
that have supported human life and informed
religions over the millennia are gone. Gone
also are the pieces of information that gave
us guidance concerning deep inner prob¬
lems, mysteries, and rites of passage. With¬
out them we are left to “work it out” our¬
selves (Moyers and Campbell 1989, 3f).
According to Eliade, when myth is living
and functioning in a society, it supplies models
of human behavior and gives meaning and
Claire E. Matthews maintains an interest in religion,
myth, and depth psychology. She is a graduate of Al-
verno College and is currently pursuing graduate studies
in depth psychology at Pacifica Graduate Institute in
Santa Barbara, California.
value to life (Eliade 1969, 2). Myth also nar¬
rates a sacred history and explains, through
the deeds of supernatural beings, how reality,
the cosmos, or a portion of it, came into being
(Eliade 1969, 5-6). Through his work in
comparative mythology, Joseph Campbell
found that there are certain timeless, univer¬
sal themes from every culture but with vary¬
ing cultural inflections. He also believes that
mythology is what lies behind literature and
the arts. Mythology likewise informs our per¬
sonal lives, particularly in relation to certain
life stages. Mythology imparts structure and
meaning to the initiation ceremonies that move
the individual from childhood to adult re¬
sponsibilities, from the unmarried to the mar¬
ried state, for example, or into a responsible
new role. Campbell maintains that the con¬
servative call for “old-time religion” is a
terrible mistake, that it would be trying to
return to something vestigial that no longer
serves us (Campbell and Moyers
1988, 10-12).
The / Ching or Book of Changes as it is
often called in English, is an ancient Chinese
manual of divination and wisdom that func¬
tions as a means of access to these transcul-
tural, mythic patterns. It did this historically
for the ancient Chinese and can do the same
contemporarily for Western humanity. It also
offers a paradigm of Eastern thought that has
implications for the Western mind. The kind
of assumptions that the / Ching makes — that
physical and psychological realities have a
connection at some deep level — have signif¬
icance for our Western society.
Marie Louise von Franz points out that the
unconscious aspect of the psyche is con-
155
Wisconsin Academy of Sciences, Arts and Letters
nected to matter; we know only that this is
so, and our scientific knowledge has come
to an end in this respect for the time being
(1980a, 79). In a sense, this perspective or
world view of the ancient Chinese points to
a religious attitude that instructs us never to
act only in accordance with conscious rea¬
soning, but with constant attention to what
could be termed irrational or unknown par¬
ticipating factors. This might mean consult¬
ing a valid oracle, such as the I Ching, or
concentrating on an attempt to get a sign from
within ourselves as to the right thing to do,
the proper path to take. In Chinese philos¬
ophy this would mean paying constant atten¬
tion to the Tao to see if the personal action
is in Tao. Applying Chinese philosophy to
Western thought would yield a new definition
of living in a religious way: being constantly
on the alert for those unknown powers that
guide one’s life. This may be a feeling as to
whether something is the right thing to do or
an instinctive feeling against it. As von Franz
puts it, “A bell does not always ring warning
us, but if it comes and one ignores it, then
something goes wrong” (1980a, 96).
One of the most important books in the
history of Chinese culture, the I Ching is one
of the Five Classics that, along with four
other works, made up the basic Confucian
canon (Gentzler 1988, 339). John Blofeld,
who made a modem translation of the I Ching
(1965) and who is obviously taken with its
value as an oracle, remarks that it was very
common for a Chinese individual to be Taoist,
Confucian, or Buddhist and something more
as well. He states, “There are whole seg¬
ments of traditional Chinese religion which
don’t fit into those categories and have ex¬
isted for several thousand years without ac¬
quiring a name” (1965, 38). The Book of
Changes reflects all of these Chinese attitudes
toward religion, cosmology, and meta¬
physics because much of it took shape before
distinctions between religions had arisen and
Taoism and Confucianism became separate
entities. According to Blofeld, the text con¬
tains the seeds or prototypes of both religions
and is not contradictory.
The archaic mode of expression used in
the / Ching adds to the difficulty in under¬
standing the oracles. Blofeld states that fre¬
quently the meaning is so esoteric that the
mind is baffled until intuition, careful thought,
or some unforeseen experience provides sud¬
den illumination. The obtuse language, as
well as the 2500 to 3000 years, creates a vast
period of time separating us from King Wen
and his contemporaries, who edited the I
Ching, and the disparity between Eastern and
Western culture further hinders clear under¬
standing (Blofeld, 32).
There are many varying explanations of
the origins of the I Ching, although the most
probable is that, like many other ancient works,
it assumed its present form through a long
process of evolution. According to R. L.
Wing, the Book of Changes was probably a
cooperative effort spanning many centuries.
The oldest stratum of ideas may have been
handed down from the elders of the nomadic
Siberian tribes. The early authors of the I
Ching observed all the cycles of life, natural
and human — the tides and the stars; plant and
animal life; the seasons; patterns of relation¬
ships in families and in societies, in business
and in warfare; and the eternal human dramas
of life, ambition, and conflict — and made a
guide to the way things change. This system
is not a fixed chart of the cosmos, but fluid
and interconnected. These writers created a
guide that offers a perspective on the eternal,
universal human drama (Wing, 8).
There are also discrepancies as to the exact
date of its conception. James Legge (1899)
states in his translation that the basic text was
prepared before 1000 B.c. in the last days of
the Shang Dynasty and the early part of the
Chou Dynasty. Confucius edited and wrote
commentaries on it that still exist as part of
some editions today. The Confucian com¬
mentaries often refer to the “superior man,”
the “chiin-tzu’’ ; there is also frequent ref¬
erence to the inferior man who is not ‘ ‘chiin-
tzu.’’ The commentary usually relates what
the “superior man” would do in a situation
and frequently uses politics or government
as an example of the arena (Blofeld 1965, 24).
156
Ancient Wisdom of the I Ching
The term “superior man” in the original text
was used to indicate a person striving to live
his life in the best possible way (Wing
1979, 30). It is reported that Confucius wished
he had fifty more years of life so that he could
study the I Ching. King Wen wrote com¬
mentaries on the social and political impli¬
cations of the hexagrams, making a monu¬
mental addition to the ancient hexagrams.
His son, the Duke of Chou, completed his
father’s work by writing commentaries on
each of the six lines within each of the sixty-
four hexagrams. Interestingly, both Taoist
and Confucian schools have claimed the /
Ching as their own classic. Later, even cer¬
tain Buddhists consulted, studied, and com¬
mented on it.
The core of the I Ching is a divination
manual overlaid with the explanations and
commentaries already mentioned. To consult
the oracle, we follow a simple divination rit¬
ual of tossing coins or sorting yarrow stalks.
We would approach the oracle as we would
a wise spiritual mentor, bringing the concerns
that we would like to see in a larger per¬
spective. In this way the more appropriate
action can be chosen. This is thought to work
because a mutual resonance echoes between
the currently active pattern informing the sit¬
uation under question, the objects used in the
divination (the coins or the yarrow stalks),
and the symbolic form described by the book.
Each coin toss (or sorting of the yarrow stalks)
is translated into a line which is either yin
(represented as broken: — ) or yang (repre¬
sented as unbroken: — ). The casting or med¬
itation is done six times, giving six lines that
are grouped in two sets of three lines each
called trigrams.
The six lines constituting the hexagram
represent the interrelationship of two fun¬
damental forces, “yin” and “yang,” two
concepts that are an integral part of ancient
Chinese philosophy and the Chinese spiritual
perspective. The Chinese frequently looked
to nature as a representation of the macro¬
cosm and microcosm. Yin and yang refer to
basic principles that are purely symbolic rep¬
resentations of energies of what we com¬
monly call “maleness” and “femaleness”
(Whitmont 1969, 171). Originally these words
referred to the sunny and shady sides of a
stream but were more generally symbolically
representative of the female (passive) and male
(active) principles in human beings, nature,
and the macrocosm. No moral verdict was
intended; neither principle is “better” or
“stronger” than the other. The Chinese saw
them as two equally potent, grounding prin¬
ciples on which all the world rests, and in
their interaction they inform, constitute, and
decompose all things. Their belief in this uni¬
versal diad also informs the I Ching (Camp¬
bell 1972, 119).
As previously mentioned, casting the coins
six times gives six lines that are either yin
or yang and form two groups of three lines
each, called trigrams. There is a traditional
belief that the legendary sage-emperor Fu-hsi
came by the idea for the trigrams from a map
found on the back of a horse or dragon-horse
(or, according to another source, a turtle) that
emerged from the Yellow River. The map
was supposedly preserved for some time but
has long since perished. It was composed of
a concentric configuration of lines made of
dark and light dots. In Legge’s opinion, the
purpose of perpetuating the legend was “to
impart a supernatural character to the tri¬
grams and produce a religious veneration for
them.” Legge (1899, 15-17) believed that
King Wen first used lines instead of circles
and was supposedly the first to combine two
trigrams to form a six-line figure called a
hexagram, of which I have more to say later.
The trigrams had an interesting evolution
from the supreme absolute as understood by
the ancient Chinese. They regarded the su¬
preme absolute as the yin and yang of the
cosmos out of which all that exists is pro¬
duced. They saw yang as always turning into
yin and yin in the process of becoming yang,
a process called enantiodromia, in which one
energy or thing turns into its opposite when
it has reached a zenith or nadir of develop¬
ment. This dynamic interplay in the cosmos
creates life, and this creative energy of life
manifests the cosmos. The ancient Chinese
157
Wisconsin Academy of Sciences, Arts and Letters
have this to say in reference to the creative
force: “From the Creative (yang) and the
Receptive (yin) emerge the ten thousand
things” (Wing 1979, 13). The yin and yang
lines combined in four different ways to rep¬
resent the seasons. A third line was added to
represent humanity as the synthesis of yin
and yang, heaven and earth, thereby creating
the eight elemental trigrams meant to rep¬
resent all the cosmic and physical conditions
on earth. The trigrams and their attributes are
The trigrams were used in early forms of
divination since they could easily be recog¬
nized and memorized. They represented, for
example, family members, parts of the body,
seasons, and many sets of ideas, as well as
more abstract attributes, so that they consti¬
tuted a very useful almanac for the ancient
Chinese to use in understanding the tenden¬
cies of change. The trigrams were also used
for divination in an arrangement of polar op¬
posites (e.g. , heaven across from earth, water
from fire). A later arrangement according to
periodicity is attributed to King Wen. Var¬
ious pairings of the trigrams by Chinese
scholars later led to the sixty-four hexagrams.
The union of the two trigrams represents the
dynamism of heaven and earth, their inter¬
action representing cosmic forces as they af¬
fect human affairs (Wing 1979, 14-15).
Carl G. Jung wrote an illuminating preface
to the English edition of Richard Wilhelm’s
translation. Blofeld praised Jung’s introduc¬
tion as a joy to read and declared that Jung
“courageously dared the scorn of his fellow
scientists by publicly asserting his belief in
the I C king's predictions. He even went so
far as to attempt to show why they are cor¬
rect” (1965, 25). Jung’s concepts of acau-
sality, synchronicity, and archetypes are es¬
sential to understanding the reliability of the
I Ching.
Concerning the causal view of the world,
Jung writes in Mysterium Coniunctionis
(1970, 464):
The causalism that underlies our scientific view
of the world breaks every thing down into in¬
dividual processes which it punctiliously tries
to isolate from all other parallel processes. This
tendency is absolutely necessary if we are to
gain reliable knowledge of the world, but phil¬
osophically it has the disadvantage of breaking
up, or obscuring, the universal interrelation¬
ships of events so that a recognition of the
greater relationship, i.e., of the unity of the
world, becomes more and more difficult.
Jung regarded this idea as a world view that
could be seen as valid, although very differ¬
ent from our Western perspective.
According to Marie Louise von Franz, the
Jungian author of On Divination and Syn¬
chronicity, synchronistic reasoning is the
classical Chinese way of thinking. The Chinese
think in terms of “fields” and know innately
that certain things “like” to happen together
in a meaningful way. In their thinking, no
differentiation has been made between “psy¬
chological” and “physical” facts. In syn¬
chronistic thinking, both inner and outer facts
can occur together. Causal thinking regards
time as linear and each moment as qualita¬
tively equal to any other; in acausal, syn¬
chronistic thinking, time is viewed as a qual¬
itative “field” in which groupings of events
typically occur. Thus in a certain moment in
time a complex of events made of inner (i.e. ,
158
Ancient Wisdom of the I Ching
thoughts, dreams) and outer (i.e., physical)
events constellate (1980b, 8).
Jung (1931, 85) stated in his commentary
on The Secret of the Golden Flower (which
Richard Wilhelm had translated from the
Chinese) that the Chinese developed intuition
to a very high degree. Because of this keenly
developed intuition, the Chinese were able
to recognize the polarity and paradox in what
is alive. Whether this gave them a greater
predisposition to comprehend the spiritual —
specifically the cosmos and the individual’s
right place in it as evidenced by the use of
the I Ching as a tool — is an interesting spec¬
ulation for the Western mind to ponder.
Blofeld also sheds some light on the es¬
sential difference between Eastern and West¬
ern thinking. “Asia’s thinkers,’’ he states,
“were chiefly occupied with the search for
life’s meaning (or at any rate, man’s true
goal) and for ways of utilizing that knowl¬
edge for self cultivation and self-conquest’’
(Blofeld 1965, 23). He felt the / Ching in¬
valuable as an aid to understanding life’s
rhythmic process with a view to bringing man
back into harmony with it.
The ancient Chinese perspective — almost
opposite to our Western view but possibly
complementary in its introverted, intuitive
way that takes into account the simultaneous
reality of spirit and matter — may hold some¬
thing valuable for us if we can be open to it.
In an essay on one of the hexagrams of
the / Ching , Rudolph Ritsema (1976, 191)
states that if the philosophical system and the
cosmological as well as social and historical
implications of the I Ching are left behind,
what remains is a book that contains a whole
web of interrelated archetypal images un¬
derlying our world. He views the I Ching as
a door to the archetypal realm, its position
between a dream and a mythology. Dreams,
he states, reveal the individual relevance of
an archetypal pattern or image, and mythol¬
ogy shows the archetypal patterns at work in
their own world. The I Ching enables one to
connect with the archetypal pattern under¬
lying the specific situation in time. The an¬
cient Chinese language lends a particular ad¬
vantage to this in that it allows images and
concepts to join in single words as well as
in sentences of the I Ching.
Just what are archetypes, and why is it
important to be in proper relationship to them?
According to Frieda Fordham (1957) in An
Introduction to Jung’s Psychology, arche¬
types are unconscious and can therefore only
be postulated, but we can become aware of
them through certain typical primordial im¬
ages. We may hazard a guess that these pri¬
mordial images or archetypes formed in the
unconscious during the thousands of years
when the human brain and human conscious¬
ness were emerging from the animal state,
and are modified or altered according to the
era in which they appear. They can be ex¬
perienced as emotions as well. When we en¬
counter a level of deep human experience
such as birth or death, triumph over ob¬
stacles, transitional stages of life, or extreme
danger, the personal level of experience taps
a deeper level. These “impressive’’ experi¬
ences break through into an old, previously
unconscious riverbed, and the experience is
extremely powerful.
According to Whitmont’s interpretation of
Jung, archetypes manifest in individuals as
automatic or instinctive emotions and drives.
The archetype appears as an experience of
fundamental importance and presents itself
as numinous. Its power can be either con¬
structive or destructive, depending on the form
of actualization and the attitude taken by con¬
sciousness (Whitmont 1969, 103f).
This does not mean that God is “nothing
but an archetype.’’ Rather, the transpersonal
power of archetypes that expresses itself to
us subjectively through psychological expe¬
rience as if it were personal guidance and
confronts us with meaning in our personal
lives and destinies is the transpersonal power
that has been called God, and this is one of
the ways we can experience Him. This may
shed some light on the nature of this ancient
book and how it was able to function as the
core of Chinese spirituality for so many years.
159
Wisconsin Academy of Sciences, Arts and Letters
Blofeld (1965, 31) quotes a Chinese friend
who had written a newspaper article in which
he explained:
The responses to be won from the Book of
Changes are sometimes of such tremendous im¬
port that they may save us from a lifetime of
folly, or even from premature death. It must
be treated with the deference due to its immense
antiquity and to the wealth of wisdom it con¬
tains. No living man can be worthy of equal
deference, for it is not less than a divine mirror
which reflects the processes of vast and never-
ending cosmic change, those endless chains of
actions and interaction which assemble and di¬
vide the myriad objects proceeding from and
flowing into T’ai Chi — the still reality under¬
lying the worlds of form, desire and formless¬
ness. It has the omniscience of a Buddha. It
speaks to the transient world as though from
the Womb of Change itself — Change, the one
constant factor amidst all the countless per¬
mutations and transformations of mental and
material objects which, when the eye of wis¬
dom is closed, appear to us as meaningless flux.
That their infinite number can be mirrored in
so small a compass is because they all proceed
according to adamantine laws and all are facets
of that spotless purity and stillness which some
men call T’ai Chi or the Tao and others the
Bhutatathata, the Womb of the Tathagatas
(Buddhas), the Source of All.
Jung felt that man needs to find a new
religious attitude, a new realization of our
dependence upon superior dominants (arche¬
types), that he is frequently operated on and
maneuvered by “archetypal forces” instead
of his “free will.” “He should learn that he
is not master in his own house and that he
should carefully study the other side of his
psychic world which seems to be the true
ruler of his fate.” Jung stated that “if the
archetype, which is universal, i.e., identical
with itself always and anywhere, is properly
dealt with in one place only, it is influenced
as a whole, i.e., simultaneously and every¬
where.” Paraphrasing Confucius’ commen¬
tary in the I Ching, Jung said, “The right
man sitting in his house and thinking the right
thought will be heard 100 miles away”
( Letters 2:594).
Since the I Ching is an ancient oracle, and
since the thesis of this paper is that the /
Ching can address our contemporary condi¬
tion with its wisdom, I asked I Ching how it
would like to be presented in this paper. The
divination procedure yielded hexagram num¬
ber 58, “The Joyous,” one of eight hexa¬
grams formed by doubling a single trigram,
“Tui,” the image of the “smiling lake” whose
attribute is joyousness (Wilhelm 1967, 223).
With each hexagram in the / Ching, the reader
finds a “judgment” and a statement of the
“image.” This is the judgment attached to
hexagram 58:
The joyous. Success.
Perseverance is favorable.
Wilhelm comments that “true joy, therefore,
rests on firmness and strength within, man¬
ifesting itself outwardly as yielding and
gentle” (1967, 224). In each trigram, a
“strong” or yang line (i.e., unbroken) lies
“within,” that is, flanked (“without”) by a
“weak” or yin (broken) line. Here is the
image accompanying hexagram 58:
Lakes resting one on the other:
The image of The Joyous.
Thus the superior man joins with friends
For discussion and practice.
Wilhelm (1967, 224f) interprets the image
in these words:
A lake evaporates upward and thus gradually
dries up; but when two lakes are joined they
do not dry up so readily, for one replenishes
the other. It is the same in the field of knowl¬
edge. Knowledge should be a refreshing and
vitalizing force. It becomes so only through
stimulating intercourse with congenial friends
with whom one holds discussion and practices
the application of the truths of life. In this way
learning becomes many-sided and takes on a
cheerful lightness, whereas there is always
something ponderous and one-sided about the
learning of the self-taught.
160
Ancient Wisdom of the I Ching
The enduring wisdom of the I Ching is man¬
ifest in this answer to the question. The joy
and success come through my inner enthu¬
siasm coupled with a genuine feeling of
wanting to share this information with others.
Use of the present paper would do well to
take the form of a discussion among friends
with whom one would ponder the truths of
life and their practical application.
The Western mind may agree or disagree
that the I Ching is a vehicle to tap the un¬
conscious for its guidance and perspective,
and that it has as much relevance for modem
Western man as for the ancient Chinese as
an alternate but valid spiritual world view.
But even without this perspective, the I Ching
exists like a venerable old Chinese master,
an example of the Chinese philosophical and
religious world view, with many secrets to
be explored and pondered.
Works Cited
Blofeld, J., trans. 1965. I Ching: The book of
changes. New York: E. P. Dutton.
Campbell, J. 1972. Myths to live by. New York:
Bantam.
Campbell, J., and B. Moyers. 1988. The power
of myth. New York: Doubleday.
Eliade, M. 1963. Myth and reality. New York:
Harper and Row.
- . 1969. The quest. History and meaning
in religion. Chicago: University of Chicago
Press.
Fordham, F. 1957. An introduction to Jung’s psy¬
chology. Harmonds worth: Penguin.
Gentzler, J. M. 1988. Confucianism. In Encyclo¬
pedia of Asian history. New York: Charles
Scribner’s Sons, 339-42.
Jung, C. G. [1931] 1962. Commentary on The
secret of the golden flower. New York: Har-
court, Brace and World.
- . 1970. Mysterium coniunctionis . Prince¬
ton: Princeton University Press.
- . 1974. Letters. Princeton: Princeton Uni¬
versity Press.
Legge, J., trans. [1899] 1963. The I Ching. New
York: Dover.
Ritsema, R. 1976. The quake and the bound. Spring
1976 191-212.
von Franz, M.-L. 1980a. Alchemy: An introduc¬
tion to the symbolism and the psychology . To¬
ronto: Inner City Books.
- . 1980b. On divination and synchronicity .
Toronto: Inner City Books.
Whitmont, E. C. 1969. The symbolic quest: Basic
concepts of analytical psychology. Princeton:
Princeton University Press.
Wilhelm, R., trans. 1986. The I Ching or book
of changes. Translated from the German by
Cary F. Baynes. Princeton: Princeton Univer¬
sity Press.
Wing, R. L. 1979. The I Ching workbook. New
York: Doubleday.
161
Distribution, Abundance, and Diversity
of Mollusks (Bivalvia: Unionidae)
from the Lower Chippewa River, Wisconsin
Terry Balding
Abstract. The lower Chippewa River, from the dam in Eau Claire to its mouth at Mississippi
River Mile 763.4, was divided into 37 sampling stations each about 2 km in length. During
the summers of 1986-1989, mollusks from each station were collected, mainly by wading.;
4,832 empty shells were identified to species and kept. In addition, 2,161 live shells were
identified as to species, measured, and returned to the river substrata. Twenty-six different
species were found, 24 having living representatives. The lower half of the river had signif¬
icantly fewer species and individuals (p < 0.01).
Freshwater mollusks or freshwater mus¬
sels of Wisconsin have been studied by
Baker (1928) and by Mathiak (1979). These
two studies were statewide in scope and
therefore did not give intensive coverage to
the mussels of the Chippewa River. Data of
an intensive scope are needed for the Chip¬
pewa because mussels are good ecological
indicators of water quality, and the expand¬
ing human population is placing an increas¬
ing burden on water quality. Freshwater mus¬
sel base-line data need to be obtained so that
they may be used as a biological monitor of
the environment or as a means of detecting
changes by a comparison to future studies.
Any plan to impound the lower Chippewa
River should refer to these data in order to
assess the detrimental effect on the mussels,
which are a river species. A recent Wisconsin
income tax checkoff for non-game species
has made money available and created in-
Terry Balding is Professor of Biology at the University
ofWisconsin-Eau Claire. In recent years he has studied
the red-shouldered hawk and the mussels of the Chip¬
pewa River.
terest in determining the presence and status
of species such as mussels.
Study area. According to Simons et al.
(1980), the Chippewa River begins in the
Upper Peninsula of Michigan, drains about
17% of Wisconsin, and empties at Missis¬
sippi River Mile 763.4. The Chippewa av¬
erages 243 m in width at its mouth and de¬
posits 553 million kg of sediment per year
into the Mississippi (recurrence interval, two
years). It is impounded many times, with the
last dam in Eau Claire, Wisconsin, where it
averages 158 m in width and discharges .028
cubic meters per second. The last 92 km of
the Chippewa River is free flowing from the
dam in Eau Claire downstream to its junction
with the Mississippi River. The 64-km por¬
tion from Eau Claire to Durand has a sin¬
uosity of 1 .49 and slope of 1 .5 feet per mile.
The substrata are primarily sand, gravel, and
glacial rocks. There is stratification of these
substrata as the channel meanders. Occa¬
sionally there are areas of silt deposits, but
in others the bottom is sandstone bedrock.
The last 27 km of river from Durand to the
Mississippi is less sinuous. Erosion from
163
Wisconsin Academy of Sciences, Arts and Letters
stream banks and islands results in a braided
channel with shifting sand sediments. In a
few places the channel cuts into large hills,
and sandstone bedrock breaks off into an¬
gular pieces and falls into the channel.
Methods
The first segment of river within the city
limits of Eau Claire was an 8-km stretch from
the dam to Interstate 94. This segment was
designated as an area for intensive study. All
shorelines were searched diligently to find
mussel concentrations. The remaining 84 km
of the Chippewa was divided into 36 seg¬
ments numbered from upstream to down¬
stream. Convenient landmarks such as islands,
small streams, bridges, and hills were used
to divide segments. Segments in this portion
of the river averaged about 2 km and were
searched by boat or by walking rapidly along
the shoreline. Once an area was found to have
mussels, it was searched more intensively
using a wading-pollywogging technique. This
technique was used because the upper part
of the study area has extensive amounts of
rocks, making the brailing method imprac¬
tical. A SCUBA technique, searching 1 m
on each side of a 30-m transect (60 m2), when
compared to the wading-pollywogging tech¬
nique showed no statistical difference. SCUBA
is a more expensive method of sampling and
is more important in deeper water. The shal¬
low nature of this river seemed to lend itself
to wading-pollywogging. The severe drought
of 1988 and a lesser drought in 1987 reduced
the water level and made the wading-
pollywogging technique more effective.
Lakes, side channels, and sloughs adjacent
to the main channel were not searched. In
some areas, which seemed likely to harbor
mudpuppies ( Necturus maculosus ), rocks were
turned over to search for the salamander mus¬
sel ( Simpsonaias ambigua).
Except for specimens that were kept as
vouchers, all live mussels were identified as
to species, enumerated, measured for length,
and returned to the river substrata. The length,
in millimeters, was the longest straight-line
distance from the anterior end of the shell to
the posterior end. The identification of the
voucher specimens was confirmed by Dr.
David Stansbery, and they were deposited
with him to be catalogued into the Ohio State
University Museum of Zoology.
All empty shells were collected and de¬
posited at the University of Wisconsin-Eau
Claire, where they were later identified.
However, two empty shell specimens, the
only representatives of their respective spe¬
cies, were sent as vouchers to Dr. Stansbery,
who confirmed their identifications.
Results and Discussion
Twenty-six species of mussels were iden¬
tified from the Chippewa River between Eau
Claire and its junction with the Mississippi
River (Table 1). Twenty-four species were
collected alive, whereas only empty shells
were found for Elliptio dilatata and Pleu-
robema sintoxia. The most abundant live
species were Fusconaia flava and Obovaria
olivaria, with other abundant species includ¬
ing Leptodea fragilis, Lampsilis ventricosa,
and Lasmigona complanata. However,
Lampsilis ventricosa and Obovaria olivaria
occurred more frequently than other species
(Table 1). One species that was found fre¬
quently yet not abundantly was Potamilus
alatus.
In Table 1 the mean length is given along
with the range length and the standard de¬
viation. The importance of these numbers can
be seen by noting Quadrula metanevra. The
mean length shell is large, but there is a small
range and standard deviation. This indicates
that only older shells exist and no recruitment
of new individuals for this species has oc¬
curred in several years. The small number of
shells and the observation that all live spec¬
imens had considerable erosion would sup¬
port the theory that this shell may soon be
extinct in the study area.
Simpsonaias ambigua can almost be con¬
sidered colonial, in that under a single rock
large numbers of specimens were found tightly
packed together. Total numbers of this spe¬
cies might not be a true reflection of its abun-
164
Mollusks of the Lower Chippewa River
dance relative to other species; therefore, to¬
tal numbers were not recorded. Of the three
sites where live salamander mussels were
found, one site was searched for a long time
before they were discovered; on the other two
sites they were found without much effort.
In contrast, a fourth site about 100 m long
was searched for four hours by two persons,
and while several mudpuppies were ob¬
served, no salamander mussels were found.
Since it was time-consuming to turn over
rocks in search of the salamander mussel,
only a few sites were searched for this spe¬
cies. Therefore, the 8.1% frequency found
in this study may not be a true representation
of the occurrence (Table 1).
Comparisons between the rank in abun¬
dance for species found alive versus dead
(Table 1) reflect only general agreement. This
could be because of collector bias or some
other cause. Nevertheless, finding empty shells
in pristine condition very likely indicates that
a species presently lives in the vicinity and
is a general indication of the relative pro¬
portions of each species.
Mussels do not occur uniformly dispersed
in a river, but rather large numbers may be
found clustered in one area and none in an¬
other. If a cluster is examined, several spe¬
cies will probably be found. The proportion¬
ate numbers of a species in a cluster may
differ from cluster to cluster. This needs to
Table 1. Data for freshwater mussel species found on the lower Chippewa River
*D = no live specimens found.
Continued on next page
165
Wisconsin Academy of Sciences, Arts and Letters
Table 1 — Continued
*D = no live specimens found.
be kept in mind when examining Table 1, as
the figures can be misleading. For example,
it would appear that Fusconaia flava (453
live specimens) might be very common.
However, the actual data show nearly 300
live shells taken in only two small areas of
the entire river.
The Wisconsin Department of Natural Re¬
sources has listed Plethohasus cyphyus as en¬
dangered and Quadrula metanevra, Simp-
sonaias ambigua, and Tritogonia verrucosa
as threatened. These species were found live
in the study area. Our general observations
and the abundance, frequency, and size data
indicate that all but Quadrula metanevra are
surviving and reproducing.
The number of species per station is shown
in Figure 1 . Data recorded for the number of
specimens per station, shown in Figure 2,
reflect a similar trend. A paired t-test (p < .01)
demonstrated that the number of species found
per station was significantly greater for the
first 18 stations than for the last 18 stations.
The difference is that at Station 19 the Red
Cedar River enters the Chippewa River, and
downstream there is a gradual loss of sin¬
uosity combined with more erosion of stream
banks and islands, which creates shifting sand
sediment. Observations revealed that stations
downstream of Station 19 with high species
diversity were related to areas where the
channel cut into sandstone hills and pieces
of bedrock were found in the channel. In a
few instances populations were found where
islands or peninsulas created pockets of water
that were protected from shifting sand sedi-
166
Mollusks of the Lower Chippewa River
20.
15_
1
; io.
5_
x = 9.24 species /station
1st 1 9 stations x = 1 3.05 species
2nd 18 stations x = 5.22 species
i
li
l l I l l l I l l l I l l I l I l l l I I I I I " I I I I I I I I I I I
★ 1 5 10 15 20 25 30 36
STATIONS
★ Intensive zone inside Eau Claire city limits
Entrance of Red Cedar River
4* Durand, Wise.
Figure 1. Number of species found at each station on the lower Chippewa River. Station 1 is
8 km downstream from Eau Claire; station 36 joins the Mississippi River at Mile 763.4.
ments. The only species found to inhabit the
shifting sand were Anodonta grandis, Lamp-
silis ventricosa, Leptodea fragilis , Obovaria
olivaria, Potamilus ohiensis, and Potamilus
alatus; most were younger specimens.
Besides the entrance of the Red Cedar,
another site where there were conspicuously
fewer mussels was the first 5 km downstream
of the Eau Claire dam. The author intends to
continue this study to the source of the Chip¬
pewa River and then its tributaries. It will be
interesting to see whether a trend exists re¬
garding the virtual absence of mussels below
a dam or entrance of a large tributary.
Mussels were found marooned in a small
pool (maximum depth 15 cm) at Station 1.
Since the pool would soon go dry, it was
decided to transplant the mussels to a nearby
suitable habitat with deeper water. Ninety-
seven live specimens (all that were visible)
were identified, measured, and transplanted
on 13 September 1988. On a return visit four
days later an additional 112 live specimens
were found and transplanted. A chi-square
test showed no significant difference in the
species composition between the two visits.
On 7 October 1988 a third visit proved not
all the living mussels on the first two visits
were found. Another 100 specimens were
discovered but not moved. There was a sig¬
nificant difference (chi-square test, p < .01)
between the species composition on the third
visit compared to either of the first two visits.
Clearly, mussels are not always detected be¬
cause they must totally bury themselves for
periods of time. Perhaps also some species
may remain buried longer than others. In fact,
on one occasion a live mussel was found
buried under at least 5 cm of sand and gravel
where there had been no water for several
hours. The mussel had good adductor re¬
flexes and seemed healthy.
Overall, the Chippewa River shells seem
to be in rather poor condition, especially the
167
Wisconsin Academy of Sciences, Arts and Letters
Figure 2. Number of specimens found at each station on the lower Chippewa River.
umbone area, probably because of the glacial
rocks rolling into them; then when the perios-
tracum is injured the acid water of the Chip¬
pewa River erodes the shell considerably.
There is good mussel species diversity on the
Chippewa, and some species seem to have
healthy populations. Some are apparently in
decline, and species represented by only a
few specimens do not give much indication
as to the health of their population. Densities
do not seem to be as high as in other rivers
I have visited such as the Mississippi, Na-
mekagon, and St. Croix. The highest density
on the Chippewa was about 30 mussels per
m2. This collecting location was markedly
better than any other site on the Chippewa.
Acknowledgments
Thanks are due to Pope and Talbot of Eau
Claire and the University of Wisconsin-Eau
Claire for funding student help. I am indebted
to Dr. David Stansbery for verification of the
mussel identifications and other helpful as¬
sistance with this study. I appreciate the help
in collecting and identifying mussels pro¬
vided by Leslie Scalzo; Andrea Pokrzywin-
ski; Marc and Sheri Harper; Rod, Jason, and
Brian Beheler; Shawn Balding; Lori Lyons;
Doug Stevens; and Linda Carls. A very spe¬
cial thanks is due to my wife Nancy for
spending many hours by my side collecting
and identifying mussels.
Works Cited
Baker, F. C. 1928. The freshwater Mollusca of
Wisconsin. Part II. Pelecypoda. Bull. Wis. Geol.
Nat. Hist. Surv. 70:1-495.
Mathiak, H. A. 1979. A river survey of the Unionid
mussels of Wisconsin, 1973-1977. Horicon,
Wis.: Sand Shell Press.
Simons, D. B., Y. H. Chen, J. Wedum, S. M.
Morrison, and H. C. Cochrane. 1980. Inves¬
tigation of the effects of Chippewa River ero¬
sion and silt reduction measures, Phase IIB. St.
Paul, Minn., U.S. Army Engineering District.
168
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The Wisconsin "Academy of Sciences, Arts and Letters was
chartered by the State. Legislature on March 16, 1870, as a
membership organization serving the people of Wisconsin by
encouraging investigation and dissemination of knowledge of
the sciences, arts and letters. For membership information,
please contact the office of the Wisconsin Academy.
TRANSACTIONS
of the Wisconsin Academy
of Sciences, Arts and Letters
Volume 81 • 1993
Editor
Managing Editor
Interns
William J. Urbrock
College of Letters and Science
University of Wisconsin Oshkosh
Oshkosh, Wisconsin 54901
Patricia Allen Duyfhuizen
328 West Grant Avenue
Eau Claire, Wisconsin 54701
Professor Wilma Clarke
Jan Kippenhan
Nathan Kailhofer
Valerie Nelson
1 ransactions welcomes articles that explore features of the State of
Wisconsin and its people. Articles written by Wisconsin authors on
topics other than Wisconsin sciences, arts and letters are occasionally
published. Manuscripts and queries should be addressed to the editor.
Submission requirements: Submit three copies of the manuscript,
double-spaced, to the editor. Abstracts are suggested for science/
technical articles. The style of the text and references may follow that
of scholarly writing in the author’s field, although author-year cita¬
tion format (CBE, APA style, etc.) is preferred for articles in the
sciences, author-page number format (MLA, The Chicago Manual of
Style ) for articles in the humanities. Please prepare figures with
reduction in mind.
© 1993 Wisconsin Academy of Sciences, Arts and Letters
All rights reserved
ISSN 0084-0505
Contents
TRANSACTIONS
Volume 81 • 1993
From the editor vi
The impact of chemical rehabilitation
on the parasitic fauna offish in a Wisconsin lake 1
Omar M. Amin, Colleen A. Dickey, and Alan R. Spallato
The application of rotenone at 1.5 ppm for chemical rehabilitation of Little Elkhart
Lake, Sheboygan County, Wisconsin, in 1981 caused the total elimination of
Proteocephalus ambloplitis, a damaging tape worm of bass, by 1 988. This finding, related
events, and control measures are discussed.
Lemanea (Rhodophyceae) in Wisconsin 7
John L. Blum
Populations of the red alga Lemanea (subg. Lemanea) are reported from streams in Iron
and Marinette counties, Wisconsin. Contrasts are drawn and explained between
members of this subgenus and subgenus Paralemanea, which is not known from
Wisconsin.
Plant ecology comes of age in the United States 1 2
Joshua C. Blumenfeld
This paper presents a history of the early twentieth-century work in plant ecology in the
United States and, in particular, looks at the work of Henry Cowles, Frederick
Clements, Henry Gleason, and William Cooper in the development of the basic
theories of plant ecology and plant community development.
The ejfects of ant mounds and animal trails
on vegetation pattern in calcareous fens 23
Quentin J. Carpenter and Calvin B. DeWitt
The effects of ant mounds and animal trails upon the distribution of plants on
calcareous fens was investigated at three sites in southeastern Wisconsin. One grass,
Muhlenbergia mexicana , was consistently associated with ant mounds, and several
species were significantly more common near animal trails.
Exhumed early Paleozoic landforms on the Baraboo Hills , Wisconsin 31
Lee Clayton and John W. Attig
Some landforms in the Baraboo Hills were formed in Precambrian or early Paleozoic
time, were then buried and preserved through Paleozoic time, were exhumed in
Mesozoic or Cenozoic time, and are preserved today in nearly their original form.
Hi
Notes on the biology of the American brook lamprey
(Lampetra appendix) in Wisconsin 39
Philip A. Cochran, Martin E. Sneen, and Alan P. Gripentrog
Collections of adult American brook lampreys from two localities revealed that
spawning occurred at temperatures cooler than previously reported and that males and
females differed in several morphological features. Also, analysis of distributional
records showed that American brook lampreys tend to occur upstream from northern
brook lampreys in those streams from which both have been collected.
Recent changes in the aquatic macrophyte
community of Lake Mendota 47
Elisabeth R. Deppe and Richard C. Lathrop
The authors summarize the results of aquatic macrophyte surveys conducted during
1 989—199 1 on Lake Mendota, a lake with a long-term record of macrophyte community
changes. Ceratophyllum demersum and Myriophyllum spicatum were the two most
abundant species, but changes in the entire macrophyte community occurred probably
as a response to unusually poor water clarity in 1990.
Were wild turkeys found historically in northwest Wisconsin ? 59
James O. Evrard
New evidence indicates the wild turkey was found in northwest Wisconsin outside
previously accepted historical range limits.
Creating the California Alps 65
Marguerite Helmers
Two articles by John Muir illustrate his use of eighteenth- and nineteenth-century
conventions of picturesque and sublime representation. While Muir employed these
established modes of representation, he also used them to satirize an audience of tourists
and leisure-class readers who expected that Nature conform to the rules of painting.
Blanchard's cricket frogs
(Acris crepitans blanchardi) in southwest Wisconsin 79
Robin E. Jung
State-endangered Blanchard’s cricket frogs were found in 19 of 40 sites which
historically had populations of this species. Of all habitat and water quality indices
measured, only water temperature differed significantly between sites with and without
Blanchard’s cricket frogs.
IV
A survey of the summer phytoplankton communities
of 579 Wisconsin lakes 89
Richard Lillie, Robert Last, Paul Garrison,
Paul Rasmussen, and John Mason
The results of a survey of phytoplankton communities from 579 Wisconsin lakes are
presented in this paper by a group of scientists from the Department of Natural
Resources. Blue-green algae (Cyanaopyceae) are shown to be, beyond any doubt, the
most common group of phytoplankton associated with blooms in Wisconsin lakes.
Discriminant analysis of geographic variation in long-tailed deer mice 1 07
from northern Wisconsin and Upper Michigan
Charles A. Long and John E. Long
The similar long-tailed deer mice of northern Wisconsin and Upper Michigan are
analyzed statistically to facilitate identification and appraise geographic variation.
Status and biology ofpaddlefish (Polyodon spathula) 1 23
in the Lower Wisconsin River
John Lyons
Paddlefish are currently rare in most of Wisconsin, but a large population persists below
the Prairie du Sac dam on the Wisconsin River. A recent study provides some of the first
biological data on the species in the state.
An “Education into Gladness”: Ron Wallace s The Makings of Happiness 137
and “The Mid-life Progress ” narrative
Bruce Taylor
This paper presents a critical appreciation of the work of Wisconsin poet Ron Wallace,
especially as represented in his latest book, The Makings of Happiness. Taylor discusses
this book both as a collection of individual poems and as a remarkable narrative about
mid-life progress.
V
From the editor
o n this beautiful May morning, awash (after many rains!)
in spring flowers and the first blossoms of fruit trees, several
stray visitors have found their way into my office through an
open window. I opened the window because the University’s
air conditioning has not been turned on for the season. The
fresh air, of course, is the welcomed visitor. The most obvious
unwelcomed guest is the sound of passing traffic on Algoma
Boulevard, but my sneezes and itchy eyes remind me that
pollen from the tulip beds beneath my windows has entered
the office, too.
By far the most interesting visitor, however, is a magnificent
queen bumblebee who has been trying for over two hours,
without success, to find her way back to the window by which
she entered! She has expended a good deal of energy attacking
a closed window and the fluorescent lighting fixtures and now
sits barely four feet away from me next to a heap of books,
resigned, evidently, to a temporary life of scholarly contempla¬
tion. Since the books are piled right next to my telephone, it
remains to be seen whether I will dare to answer if it rings!
By the time this 1 993 issue of Transactions appears, circum¬
stances in my office will have changed dramatically, I expect.
Windows will be locked shut against the fall or early winter
cold, the heating plant will be in action, pollen will have been
replaced by late molds, pumpkins and squashes will represent
the last hurrahs of the growing season, and according to my
entomologist colleague down the hall, the queen bee may have
settled into an abandoned mouse or bird’s nest for overwinter¬
ing — if she ever finds her way out of the office!
I have found my first year as editor of Transactions not
unlike the experience just described. Saying “Yes” to the
editorship has opened a window to visits by all sorts of new
experiences, expected and unexpected, but almost all of them
welcomed. Best among these has been the opportunity to make
the acquaintance of and to collaborate with the outstanding
array of authors, reviewers, and production staff represented
by this issue.
I have been extremely impressed, for example, by the
willingness of professional colleagues to spend much time,
care, and energy on their review of manuscripts. I have enjoyed
vi
telephone conversations with many of them
as well as the usual written correspondence.
Some are my own colleagues at UW Osh¬
kosh, whose expertise at reviewing I have
only just discovered. As we all know, the
work of reviewers often goes unheralded,
although it is a scholarly service absolutely
essential for ensuring that only articles wor¬
thy of publication and of further scholarly
citation are actually published. Without fear
of contradiction, I can speak for all the au¬
thors by acknowledging the debt they owe to
the constructive criticisms and suggestions
forwarded by their reviewers. All of us as
readers are indebted as well to these review¬
ers, whose only compensation for their
scholarly contribution is our “Thank You”
and their own satisfaction at having fur¬
thered the cause of responsible research and
of clear, accurate and reader- friendly written
communication.
Of course, I also opened my window to
the authors themselves, several of whom I
have had the great pleasure of meeting on
campuses and at workshops and conferences
around Wisconsin during the past year. Al¬
together, the variety of research they report
in this issue should be of interest to a wide
spectrum of readers and should contribute to
the related studies and research of many.
And for all of us who simply take delight in
learning more about the natural and human
history of Wisconsin, there is much here to
please: from a discussion of paleozoic land-
forms in the Baraboo Hills to an appreciation
of John Muir’s picturesque creation of the
California Alps; from endangered cricket
frogs in southwest Wisconsin, calcareous
fens in the southeast, and parasitic fauna in
eastern Elkhart Lake to deer mice and wild
turkeys in the north and northwest; from
paddlefish in the lower Wisconsin River to
American brook lamprey in Taylor and Jam-
bo creeks; from the red alga Lemanea in the
streams of Marinette and Iron counties to the
macrophyte community on Lake Mendota
and to the summer phytoplankton on 579
Wisconsin lakes; from a history of early twen¬
tieth-century work in plant ecology in
Wisconsin and the Midwest to a critical
appreciation of a late twentieth-century mid-
life progress as portrayed by one of
Wisconsin’s premier poets.
Readers may notice an updated look in the
design and layout of the 1993 Transactions ,
thanks to the artistic eye of our indefatigable
Managing Editor, Patricia Duyfhuizen. Her
contributions to the actual production of this
journal are too many to begin to enumerate.
Suffice it for me to express my gratitude to
Tricia for transforming a heap of individual
manuscripts into a finished publication of
which members and supporters of the Wis¬
consin Academy justifiably may be proud.
Thanks also to former editor Carl Haywood,
Wisconsin Academy Review editor faith
Miracle, Executive Director LeRoy Lee, and
my special on-site all-around advisor and
collaborator at UW Oshkosh, Neil Harri-
man, for their generous assistance in helping
me handle all the “visitors,” expected and
unexpected, who have come in through the
window during this first year of my editor¬
ship.
Bill Ur brock
P.S. I trapped the queen bee in a cup and
released her outside!
vii
Omar M. Amin, Colleen A. Dickey, and
Alan R. Spallato
The impact of chemical rehabilitation
on the parasitic fauna offish
in a Wisconsin lake
Abstract A 23-ha, meso-eutrophic lake in eastern Wisconsin was treated with rotenone
(1.5 ppm) in November, 1981, to kill its rough fish and restock with desirable
fish populations. Pre- and post-treatment samples of fish were examined for
parasites in September, 1981 and 1988, respectively. Moderately heavy infec¬
tions with Proteocephalus ambloplitis ( Cestoda) disappeared after the rotenone
treatment, which killed its fish hosts, largemouth bass, Micropterus salmoides,
and sunfish in 1981. Infections with Neoechinorhynchus cylindratus and
Leptorhynchoides thecatus (Acanthocephala) decreased and increased dra¬
matically, respectively, as a result of changes in their fish host populations
following rotenone use. Metacercariae of Posthodiplostomum minimum
(Trematoda) in sunfish and Hysterothylacium brachyurum (Nematoda) in
largemouth bass were also present before and after the chemical treatment,
respectively. The observed loss of P. ambloplitis after 1981 points to a possible
method for its control.
Efforts to establish desirable fish populations by rotenone
treatment of lakes dominated by winter-kill resistant spe¬
cies, e.g., bullhead, and restocking have recently been attempted
by the Wisconsin Department of Natural Resources, as well
as by other agencies throughout the United States since the
mid-1950s (Gilderhus et al. 1986). The impact of such chemi¬
cal treatments on invertebrate and fish populations and con¬
sequently on their parasite fauna is, however, not known. The
case of the recent chemical rehabilitation of Little Elkhart Lake
in eastern Wisconsin provided an opportunity to examine the
effect of chemically induced host population changes on the
composition and prevalence of parasite species. Findings not
only reflected answers to scientific curiosity, but also indicated
a possible new approach to the control of some injurious fish
parasites.
TRANSACTIONS Volume 81 (1993)
1
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Materials and Background
Little Elkhart Lake is a 23-ha glacial lake that
is 3 and 8 meters in average and maximum
depth and 1.3 years in average water resi¬
dence time, in Sheboygan County, eastern
Wisconsin. The meso-eutrophic lake is an
extension of hard ground water table with
moderate nutrient level, e.g., total phospho¬
rus, nitrogen, and alkalinity averaged 0.04,
1.0, and 124 mg/1, respectively, at various
water depths. Dominant species of rooted
aquatic vegetation included Myriophyllum
spicatum , Najas flexilis , Potamogeton illino-
ensis, and P. amplifolious. Algae were present
in low densities.
Before the chemical treatment in 1981,
the fish population was dominated by black
bullhead, Ictalurus melas, with a small num¬
ber of northern pike, Esox lucius; walleye,
Stizostedion vitreum; largemouth bass, Mi-
cropterus salmoides ; bluegill, Lepomis macro-
chirus; pumpkinseed, L. gibbosus\ black crap-
pie, Pomoxis nigromaculatus , yellow perch,
Perea flavescens; white sucker, Catostomus
commersoni', golden shiner, Notemigonus
crysoleucas ; and yellow bullhead, /. natalis. A
typical complex of lake invertebrates in¬
cluded abundant copepods and cladocerans
and rare daphnias. Shore terrestrial verte¬
brates that may be involved in the life cycle
of helminths infesting fish as larvae included
many bird species (mallard, blue-wing, teal,
wood duck, etc.) and mammals (mink, wea¬
sel, muskrat, raccoon, etc.); see Claggett
(1981) for more details.
Little Elkhart Lake has a history of
stunted sunfish and intermittent good large-
mouth bass populations. It is characterized
by infrequent winter kill (1932, 1939), a
chemical treatment and restocking with
northern pike, largemouth bass, walleye, yel¬
low perch, and golden shiner in 1961
(Schulz 1964, 1965), and subsequent acci¬
dental introduction of black bullhead,
pumpkinseed, and largemouth bass by 1971.
Black bullhead population increased as an¬
other winter kill in 1975 (Belonger 1976)
caused weeds and undesirable fish species to
become overabundant.
The lake was electroshocked to sample
pre-treatment fish on September 22 and 23,
1981. On November 17, 1981, 220 gallons
of rotenone were applied at a concentration
of 1.5 ppm. Except for bullhead, total kill
of all other fish species, including large¬
mouth bass and sunfish, was noted. The
crustacean and other invertebrate commu¬
nities, however, remained practically intact
(Nelson 1985a and pers. comm.). In May
and June, 1982, the lake was stocked with
10,000 fingerling largemouth bass from
Crystal and Gerber lakes, and 54,310 fin-
gerling hybrid sunfish (bluegill x green sun¬
fish) and bluegill from Beechwood Lake, as
well as with 118 big largemouth bass up to
37 cm long. Subsequent electroshocking ef¬
forts in 1983 and 1984 demonstrated good
survival of largemouth bass, sunfish hybrid,
bluegill, and black bullhead (Nelson 1985b).
On September 1, 1988, a post-treatment
sample of fish was similarly taken from the
lake. Both pre- and post-treatment fish
samples were promptly examined for para¬
sites after transfer to the lab on ice. Parasites
were systematically recovered and routinely
processed for microscopical examination.
Results
Two fish species were collected and exam¬
ined during each of the 1981 and 1988 sur¬
veys: largemouth bass (40 fish, 16-41 (mean
26) cm long in 1981 and 42, 17-28 (24)
cm long in 1988) and bluegill (9, 13-18
(15) cm long in 1981 and 12, 10-20 (15)
cm long in 1988). Largemouth bass and
bluegill, along with all other species, except
2
TRANSACTIONS
AMIN et al.: The impact of chemical rehabilitation on parasitic fauna of fish
black bullhead, were totally eliminated from
the lake as a result of the 1981 treatment.
The 1988 samples were from new fish in¬
troductions mostly during 1982. Four
pumpkinseed, 12 black crappie, and 13 yel¬
low perch were also examined in 1981, and
7 black bullhead were examined in 1988.
In the 1981 pre-treatment study, the
prevalence and intensity of Proteocephalus
ambloplitis (Leidy) (Cestoda) infections in
largemouth bass were moderate to heavy in
intestinal and body cavity (gut surface, liver,
spleen) locations (Table 1); only 27 mature
gravid adults were localized in the intestine.
The distribution of P. ambloplitis also ex¬
tended into the body cavity of other fish spe¬
cies, e.g., 14 worms in 4 of 9 bluegill (Table
1), 1 in 1 of 12 black crappie, 6 in 3 of 13
yellow perch, and 4 in 2 of 4 pumpkinseed.
In the 1988 post-treatment study, P. amblo¬
plitis was absent from all sites in the 3 fish
species examined then, including 42 large-
mouth bass, its major host.
The prevalence and intensity of Neoechi-
norhynchus cylindratus (Van Cleave) Van
Cleave (Acanthocephala) was relatively high
in largemouth bass before the chemical treat¬
ment but decreased by 1988. The opposite
trend was observed in the other acantho-
cephalan Leptorhynchoides thecatus (Linton)
Kostylev; its intensity of infection increased
from 2.12 to 64.24 per examined fish in gut
locations and from 0.37 to 0.93 in the body
cavity of largemouth bass. Bluegill were not
infected with either acanthocephalan species
in 1981 but were considerably more fre¬
quently and heavily infected with L. thecatus
than with N. cylindratus in 1988 (Table 1).
In addition, 1 76 Z. thecatus were also recov¬
ered from the gut of 5 of 7 black bullhead
(48 worms) and the body cavity of 2 other
bullheads (128) in 1988 .
In 1981, the sex ratio of N. cylindratus in
largemouth bass was 1 male : 2.31 females
(91% had eggs) and 1:1.44 in L. thecatus (all
females had eggs). In 1988, the sex ratio was
1:0.61 and 1:0.63, in the same order.
Gravid females of both acanthocephalan
species were considerably less frequent than
in 1981.
In addition, 33 Posthodiplostomum mini¬
mum (MacCallum) (Trematoda) were recov¬
ered from the body cavity of 3 of 4 pump¬
kinseed (32 metacercariae) and from the
body cavity of 1 of 9 bluegill (1) in 1981,
and 1 1 Hysterothylacium brachyurum (Ward
and Magath) (Nematoda) were recovered
from the body cavity (6) and intestine (3)
of 7 of 42 largemouth bass in 1988.
Discussion
Parasite community succession stabilization
was estimated to take at least 5 years from
the initiation of major ecological shifts, e.g.,
impoundment (Becker et al. 1978). In this
study, 8 years elapsed between the pre- and
the post-treatment studies. Brown and Ball
(1943), Sharma (1949), and Wright (1957)
indicated that populations of microcrusta¬
ceans, e.g., Daphnia and Cyclops , were vari¬
ably affected but not eliminated after treat¬
ment with rotenone. Major ecological shifts
were shown not to deplete crustacean popu¬
lations, e.g., copepods (Becker et al. 1978).
The half life of rotenone in warm (24 °C)
and cold water (0°C) ponds in Wisconsin
during September and March was 13.9 and
83.9 hr, respectively (Gilderhus et al. 1986).
Rotenone at a concentration of 1 ppm re¬
mained toxic to bluegills in wire cages for
7-18 days in ponds during an Alabama win¬
ter but caused no residual deleterious effects
upon subsequent bluegill production
(Wright 1957). Mortality of caged fathead
minnows, Pimephalus promelas, was 100%
48 hours after 0.15 mg/1 rotenone treatment
of a Wisconsin pond during September but
Volume 81 (1993)
3
Table 1. Pre- and post-treatment surveys of parasites
of Micropterus salmoides and Lepomis macrochirus in Little Elkhart Lake
CO c
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CL 03
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4
TRANSACTIONS
Leptorhynchoides Intestine 0/9 0 3/12 (25)
thecatus
Body cavity 0/9 0 6/12 (50)
AMIN et a I . : The impact of chemical rehabilitation on parasitic fauna of fish
declined to 0 after 72 hours (Gilderhus et
al. 1986). These data support findings about
the mortality of fishes and survival of Crus¬
tacea after the 1981 treatment of Little
Elkhart Lake with rotenone.
The complete disappearance of P amblo-
plitis from bass and sunfish is perhaps the
most dramatic outcome of the 1981 chemi¬
cal treatment. This clearly corresponded
with the mortality of the definitive host,
largemouth bass (where adult worms repro¬
duce), and the principal intermediate hosts,
e.g., bluegill and other sunfish species, and
probably resulted from it, following the ap¬
plication of rotenone.
The mortality of sunfish in 1981 was also
considered responsible for the dramatic de¬
cline in N. cylindratus population. The ma¬
jor intermediate host of this acanthoceph-
alan species is bluegill (see Becker et al.
1978); other sunfish species are important
paratenic (transport) hosts, e.g., pumpkin-
seed among others (Hoffman 1967). The
fact that one of the paratenic hosts of N.
cylindratus is black bullhead, which survived
the 1981 chemical treatment, might have
helped that acanthocephalan’s marginal sur¬
vival (Table 1).
The assumption that the loss of P. am-
bloplitis was related to changes in the fish
and not the crustacean (cladoceran, cope-
pod, or amphipod) host populations is based
on the following. The increase in L. thecatus
infections after the treatment must have
been related to the survival of its amphipod
intermediate hosts including Hyalella sp.,
e.g., H. knickerbockeri which is also an in¬
termediate host for P. ambloplitis. The mor¬
tality of the definitive and paratenic hosts of
L. thecatus , particularly largemouth bass and
sunfish in 1981, indicates that L. thecatus
could have only survived the chemical treat¬
ment in the crustacean host. The survival of
N. cylindratus was also possible because of
the survival of its ostracod intermediate host,
possibly Cypria sp.
The complete eradication of the bass
tapeworm, P. ambloplitis , by eliminating its
fish hosts in a closed system like Little
Elkhart Lake under the above conditions
points to a practical method for the control
of this injurious worm. This goal can be ac¬
complished at the same time the quality of
the fish fauna is being upgraded.
The loss of the light P. minimum infec¬
tions after the 1981 treatment might have
been affected by the mortality of its cen-
trarchid fish intermediate hosts. The new H.
brachyurum infections in bass after 1981
were probably introduced from stocking
sources. The significant changes in the sex
ratio and reproductive state of the two acan-
thocephalan species are not fully understood.
Acknowledgments
This work could not have been done with¬
out the help and cooperation of James
McNelly, Larry Claggett, and John Nelson,
Wisconsin Department of Natural Re¬
sources.
Works Cited
Becker, D. A., W. D. Carr, D. G. Cloutman,
W. A. Evans, R. G. Heard, P. D. Holmes,
M. D. Norman, and W. B. Owen, Jr. 1978.
Pre- and post-impoundment ichthyoparasite
succession in a new Arkansas reservoir. Ar¬
kansas water Resour. Res. Cent., Univ. Ar¬
kansas Pub. No. 54, 85 p.
Belonger, B. 1976. Little Elkhart Lake Fishery
Report. Wis. Dept. Nat. Resour. Intradept.
Memo.
Brown, C. J. D., and R. C. Ball. 1943. An ex¬
periment in the use of derris root (rotenone)
on the fish and fish-food organisms of Third
Sister Lake. Trans. Am. Fisher. Soc. 72: 267-
284.
Volume 81 (1993)
5
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Claggett, L. 1981. Little Elkhart Lake Fishery
Rehabilitation. Environ. Impact Assess.
Screen. Worksheet. Wis. Dept. Nat. Resour.
Gilderhus, P. A., J. L. Allen, and V. K. Dawson.
1986. Persistance of rotenone in ponds at dif¬
ferent temperatures. N. Am. J. Fisher. Man¬
age. 6: 129-30.
Hoffman, G. L. 1967. Parasites of North Ameri¬
can freshwater fishes. Berkeley and Los An¬
geles: University of California Press, 486 p.
Nelson, J. E. 1983a. Progress report - F264 -
Little Elkhart Lake - 1983 data. Wis. Dept.
Nat. Resour. Intradept. Memo.
- . 1985b. Progress report - F264 - Little
Elkhart Lake - 1984 data. Wis. Dept. Nat.
Resour. Intradept. Memo.
Schultz, P. T. 1964. Little Elkhart Lake - 1963
survey. Wis. Dept. Nat. Resour. Intradept.
Memo.
- . 1963. Little Elkhart Lake, Sheboygan
County- 1964 observations. Wis. Dept.
Nat. Resour. Intradept. Memo.
Sharma, S. 1949. Effect of partial poisoning of
ponds with rotenone on the abundance of
bottom organisms. MS Thesis, Alabama
Polytech. Inst. Auburn, Alabama.
Wright, T. W. 1957. The rates of dissipation of
certain rotenone preparations, their residual
effects on bluegill production, and their ef¬
fects on populations of fish-food organisms.
MS Thesis, Alabama Polytech. Inst. Auburn,
Alabama.
Omar M. Amin is the Director of the Institute of
Parasitic Diseases while in the United States. He
was formerly a Professor of Parasitology at the Uni¬
versity of Wisconsin since 1971. A major focus of
his work is the parasitology of wildlife, particularly
of fishes from Wisconsin, North America, and else¬
where in the world. He has an MS in Medical
Entomology from Cairo University and a PhD in
Parasitology from Arizona State University.
Colleen A. Dickey contributed to the post-treat¬
ment study while studying at the University of
Wisconsin where she received her BS in Biology in
1989. She currently holds a Masters of Education
from National-Louis University and teaches at the
Glenview District in Illinois.
Alan R. Spallato contributed to the pre-treatment
study while studying at the University of Wiscon¬
sin where he received his BS in Biology in 1983.
He worked at the Madison Hazleton Laboratories
before he became an Environmental Chemist at the
State Laboratory of Hygiene, Madison, Wisconsin,
in 1989.
6
John L. Blum
Lemanea (Rhodophyceae)
in Wisconsin
Zemanea Boiy is a relatively large and firm, attached fresh¬
water red alga. Seen underwater, it might easily be mis¬
taken for a small vascular plant. It grows most conspicuously
in and around waterfalls, thus in some of the more scenic and
recreational sites. In the northern states, it is generally repre¬
sented by a species of the subgenus Lemanea (formerly Sacheria
Sirodot). The other subgenus (. Paralemanea ) consists of algae
which are more complex in structure. Species of subgenus
Paralemanea are well represented in the southern states, the
Pacific states, and are especially abundant in the Ohio Valley,
but have never been reported in Wisconsin. The closest col¬
lection appears to be from the Greencastle, Indiana, area. Be¬
cause species of subgenus Paralemanea tend to occur where cer¬
tain types of limestone are abundant (Palmer 1933, 1940,
1941), rapids of streams in southeastern Wisconsin where lime¬
stone strata are exposed probably represent the best possible
places for finding members of Paralemanea within the state.
The earliest known Wisconsin collection of Lemanea subg.
Lemanea was made by L. S. Cheney in 1894 in the vicinity of
Stevens Point (precise location not identified) (MIN).* J. B.
Moyle found it in the Black River (Douglas County) in 1 940
(MIN), and J. W. Thomson also collected it there in 1942
(PH). The relative paucity of sites compared with other states
led to a search for other possible locales. The author has more
recently added other sites in the Pike (Dave’s Falls), Peme-
bonwon (Long Slide Falls), and Peshtigo (Strong Falls) rivers
(Marinette County), in the Potato (at Upson) and Montreal
^Herbarium specimens referenced are in the Field Museum (F), the Uni¬
versity of Minnesota Herbarium (MIN), and the Academy of Natural Sci¬
ences of Philadelphia (PH).
TRANSACTIONS Volume 81 (1993)
7
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 1. Distribution of Lemanea (subgenus Lemanea ) in the upper Great Lakes Region.
(near Hurley) rivers (Iron County), as well
as sites in northern Michigan (Fig. 1), in¬
cluding Jackson Creek (Gogebic County)
and the Ontonagon River (Ontonagon
County). Vouchers will be deposited in F
and elsewhere.
Lemanea appears to be abundant mostly
over short reaches of streams. It reproduces
sexually and produces large numbers of car-
pospores within the gametophyte thallus.
Stewart (1983) and Sheath (1984) indicate
that it can also grow as a perennial from at¬
tached basal filaments, and the author has
found it growing at numerous sites in Indi¬
ana where C. M. Palmer found it forty or
more years before.
Vis and Sheath (1992) have recently dis¬
cussed and revised the taxa of Lemanea in
North America. Their treatment permits
identification of Wisconsin and Upper Pen¬
insula material under the names of Euro¬
pean entities, e.g., Lemanea fluviatilis (L.) C.
Agardh and L. fucina Bory. In view of some
uncertainties in the applications of these
names, which I am unable to resolve, it
seems preferable not to attempt the identifi¬
cation to species of my materials at this time.
The gametophytic plant of Lemanea
bears, following fertilization, the carpo-
spores. Various characters of the gameto¬
phyte thallus constitute the principal basis
for differentiation of the two subgenera.
This differentiation can usually be made
with a hand lens. The thallus is nodose-
cylindric in both subgenera with the car-
pospores visibly borne en masse internally at
the “nodes” in subgenus Lemanea and at the
internodes in subgenus Paralemanea (Figs.
8
TRANSACTIONS
BLUM: Lemanea (Rhodophyceae) in Wisconsin
Figs. 2-6. Lemanea (subgenus Lemanea) spore-producing plants from Long Slide Falls,
Pemebonwon River, Marinette County, Wisconsin, collected 26 August 1985, J. Blum
#4764. X 10. Figs. 3-6 show points of differentiation between spore-bearing plants of
subgenus Lemanea and subgenus Paralemanea. Fig. 3. diagram shows relative posi¬
tion of the spore masses in relation to the “nodes.” Carpospore masses are shown by
stippling. Fig. 4. diagram contrasts the presence (subgenus Paralemanea with the ab¬
sence (subgenus Lemanea) of rhizoids surrounding the axial filament. Fig. 5. diagram
contrasts the position of the spermatangial areas (stipples). Fig. 6. diagram contrasts
the bases of plants.
Volume 81 (1993)
9
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
2, 3). This difference is less visible in cer¬
tain species and certain specimens than in
others. Sectioning the thallus reveals a single
central axial filament in subgenus Lemanea
whereas in subgenus Paralemanea numerous
internal rhizoidal filaments surround the
axial filament and constitute a strong cen¬
tral strand (Fig. 4) reminiscent of the stele
in dicot roots. Observation of the central
strand cannot easily be done with a hand
lens, but free-hand sections reveal the essen¬
tial point of distinction.
A third point of distinction is noted in
the spermatangial (male) areas of the thal¬
lus, which are external and found at the
nodes: subgenus Lemanea has protuberant
spots of spermatogenous tissue whereas in
subgenus Paralemanea the spermatangial areas
form a nearly complete ring or band encir¬
cling the nodal areas (Fig. 3). Thus in sub¬
genus Lemanea the gametangia of both sexes
are found in the nodal areas; in subgenus
Paralemanea they essentially alternate, with
the nodes bearing the male gametangia and
the internodes bearing the female gametan¬
gia. In early stages of growth, the minute
trichogynes, the receptive external female
structures, can be seen at 440X magnifica¬
tion. These trichogynes mark the areas where
the (internal) future carpospore masses will
develop; they appear on the flanks of the
enlarged “nodes” in subgenus Lemanea and
in the internodes in subgenus Paralemanea.
The trichogynes can be seen in edge view on
the silhouette of a thallus whole mount, as
well as in surface view of a sufficiently
cleared and stained specimen.
Lemanea carpospores are frequently ripe
in populations of the eastern United States
after approximately 1 July, and are dispersed
when the thallus decays or breaks apart.
The fourth point of distinction is a sim¬
pler but less dependable one: the base of the
gametophytic thallus in subgenus Parale-
manea is regularly and gradually narrowed to
its point of attachment, whereas in subge¬
nus Lemanea , at least in Wisconsin material,
there is frequently a distinct and non-nodose
stipe (Figs. 2, 6) which is sharply marked off
from an enlarged fertile upper part. Branch¬
ing of the spore-bearing thallus in subgenus
Lemanea is also more frequent than in sub¬
genus Paralemanea or at least in the Ameri¬
can species of Paralemanea.
All points of distinction can be misread:
(1) The single axial filament in subgenus
Lemanea is nearly approximated in some
California Paralemanea species which may
have only 2-4 internal rhizoids accompany¬
ing the axial filament. (2) The protuberant
male areas in subgenus Lemanea are occa¬
sionally confluent within the nodal area and
approximate the interrupted male “bands”
in some material of subgenus Paralemanea.
Additionally, this essential distinction is not
easy to make without prepared slides and
good microscope equipment. (3) The pres¬
ence of carpospores in either nodal or inter-
nodal areas is a seasonal phenomenon and
will not be observed in early spring or if fer¬
tilization has not occurred. The position of
the female apparatus and carpospore masses
is also considered a species character in sub¬
genus Paralemanea , and in both subgenera,
spores may escape from their usual position
so that individuals are commonly seen with
carpospores thoroughly scattered in the cavi¬
ties of the gametophyte thallus. (4) The con¬
trast between the gradually and the abruptly
narrowed basal segment is frequently not
pronounced.
The spore-producing haploid thallus al¬
ternates in its life history with a minute,
filamentous stage, the “Chantransia ” stage
(Sirodot 1872; Atkinson 1890), which is a
diploid plant. Meiosis occurs in apical cells
of tiny branches that grow from the diploid,
Chantransia stage (Magne 1967). The result-
10
TRANSACTIONS
BLUM: Lemanea (Rhodophyceae) in Wisconsin
ing haploid cells then develop into the mac¬
roscopic monoecious gametophytes; hence,
the gametophytes develop from and are at¬
tached to the Chantransia. The Chantransia
stage should not be disregarded by the col¬
lector. Bourrelly (1970) considers that the
Chantransia stage is probably necessary for
species determination in this genus. Summer
collections of the spore-bearing thallus un¬
fortunately are unlikely to have the Chan¬
transia stage attached; after about 30 April,
only fragments of it remain.
Summary
1. Lemanea (Rhodophyceae), erstwhile
considered to be rare in Wisconsin, is shown
to occur in streams of Iron and Marinette
counties.
2. Lemanea may have permanent sites of
residence in streams, since several places
where it was collected over the past century
have been successfully re-collected by the
author in the 1980s.
3. Points of distinction between the two
subgenera of Lemanea are summarized. Use¬
ful caveats in applying them are recom¬
mended and explained.
Acknowledgments
The author expresses his gratitude to Pro¬
fessors Michael Wynne and Neil Harriman
for their detailed comments on the manu¬
script and to Ronald Meister and Robert
Wakeman, who gave extensive help in field
work.
Works Cited
Atkinson, G. F. 1890. Monograph of the
Lemaneaceae of the United States. Ann. Bot.
(London) 4:177-229.
Bourrelly, P. 1970. Les Algues dEau Douce III.
514 + 2pp. (inch 134 plates.) Paris.
Magne, M. F. 1967. Sur le deroulement et le lieu
de la meiose chez les Lemaneacees (Rhodo-
phycees, Nemalionales). Compt. Rend. Hebd.
Seduces Acad. Sci. Ser. D. 265:670-73.
Palmer, C. M. 1933. Distribution of the alga,
Lemanea, in Indiana. Proc. Indiana Acad. Sci.
40:111-13.
Palmer, C. M. 1940. Development, Taxonomy
and Distribution of Lemanea in Indiana With
Notes on the Distribution of the Genus in North
America. Ph.D. Dissertation, 91 pp. Indiana
University, Bloomington.
Palmer, C. M. 1941. A study of Lemanea in In¬
diana with notes on its distribution in North
America. Butler Univ. Bot. Stud. 5:1-26.
Sheath, R. G. 1984. The biology of freshwater
red algae. In Progress in Phycological Research,
ed. F. E. Round and D. J. Chapman, 3:89-
158. Bristol: Biopress.
Sirodot, S. 1872. Etude anatomique, organo-
genique et physiologique sur les algues d’eau
douce de la famille des Lemaneacees. Ann.
Sci. Nat. Bot., Ser. 5, 16:1-95, 8 plates.
Stewart, Joan G. 1983. Lemanea (Rhodophyta)
in mountain streams of southern California.
Madrono 30: 255-56.
Vis, M. L. and R. G. Sheath. 1992. Systematics
of the freshwater red algal family Lema¬
neaceae in North America. Phycologia
31:164-79.
John L. Blum, B.S., M.S., Wisconsin, Ph.D.
Michigan 1953. Assistant Professor Biology,
Canisius College, 1946-1963. Associate Professor
and Professor ofBotony, University of Wisconsin-
Milwaukee 1963—1985. Associate Dean in Letters
and Sciences (Science), UW-Milwaukee 1967-
1969. Emeritus, 1985-date.
1 1
Volume 81 (1993)
Joshua C. Blumenfeld
Plant ecology comes of age
in the United States
Who were the first [ecologists] in America? Pound and Clements,
Cowles, Gleason, Harshberger, and Harper, in about that order, fol¬
lowed very closely by T ranseau and Shantz —
Henry A. Gleason, 1953
he final years of the nineteenth century and the first half
X of the twentieth century saw not only the establishment
of plant ecology as a discrete field of study, but the American
dominance in vegetation theory and research (Tobey 1981).
The later Wisconsin work of Curtis, McIntosh, and Cottam
was built on the groundwork laid by this early, and predomi¬
nantly Midwestern, group of ecologists. By the time Ernst
Haeckel’s term “oekologie” was officially changed to “ecology”
by the Madison Botanical Congress of 1893, university pro¬
grams in plant ecology were already well established, with the
first ecological doctorate awarded in 1879 (Arthur 1895;
Gleason 1953).
Two distinct schools of thought developed in America, both
centered in the Midwest: one, a holistic/organismic tradition
influenced primarily by Oscar Drude and the works of
Alexander von Humboldt and August Grisebach, the other, a
reductionist/individualistic tradition influenced by Johannes
Warming and Augustin Pyramus de Candolle (Cowles 1898;
Crawley 1986; Gleason 1953). The center of the holistic/or¬
ganismic school developed at the University of Nebraska (char¬
tered in 1869), while believers in the reductionist/individual-
istic school gravitated to the University of Chicago (founded
in 1891) (Kormondy 1965; McIntosh 1976, 1985).
1 2
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BLUMENFELD: Plant ecology comes of age in the United States
Tobey (1981) points out that one pos¬
sible reason for Chicago’s orientation was
the university’s policy of strictly pure re¬
search, as opposed to the land-grant college
orientation of practical science:
Chicago’s ecology was concerned less with
controlling than with preserving the natural
world. [Nebraskan] ecology grew out of and
returned for nourishment to the practical soil
of agriculture.
John Merle Coulter, a former student of
Asa Gray and, most recently, president of
both Indiana University and Lake Forest
College, arrived at the University of Chica¬
go in 1891 (Rodgers 1944; Sears 1969).
Coulter was but one member of an impres¬
sive faculty assembled to teach at the new
university and was given the task of estab¬
lishing the school’s Department of Botany
(McIntosh 1976).
Coulter was profoundly influenced by
Warming’s works and based a series of ini¬
tial graduate lectures on them, hampered by
the fact that, at the time, Warming’s works
were available only in the original Danish.
However, as Charles Chamberlain, a gradu¬
ate student in the class, recalled, “None of
us could read Danish except a Danish stu¬
dent, who would translate a couple of chap¬
ters, and the next day Coulter would give a
wonderful lecture on Ecology” (Chamber-
lain 1940).
One student in the lecture series grew
impatient with this slow translating process
and over time taught himself Danish, even¬
tually reading Warming’s works long before
an English translation was available (Cham¬
berlain 1940). This student was Henry
Chandler Cowles (1869-1939), who would
eventually become chairman of the depart¬
ment founded by Coulter.
Cowles received a fellowship to study ge¬
ology at the University of Chicago in 1893,
where he initially concentrated on landforms
and plant fossils and was introduced to the
glacial features and beach ridges of the Chi¬
cago area (Humphrey 1961). In addition,
through Coulter’s lectures, Cowles quickly
grasped the new ideas put forth by Warm¬
ing (Chamberlain 1940).
Coulter encouraged Cowles’s interest in
plant ecology, suggesting that Cowles com¬
bine his extensive knowledge of geomor¬
phology with plant ecology to see if Warm¬
ing’s studies of Danish sand dunes was ap¬
plicable to the dunes along Lake Michigan
(Cook 1980). Cowles eventually turned the
results of his findings into his Ph.D. disser¬
tation, An Ecological Study of the Sand Dune
Flora of Northern Indiana (1898).
By this time, Cowles was the recognized
expert on plant ecology at the university, and
Coulter called on him to teach most of the
ecology courses. Aside from his teaching
duties, Cowles took an active role in writ¬
ing for the Botanical Gazette , founded by
Coulter in 1873 and published by Coulter
at the university. The Gazette was an ideal
forum for the dissemination of Cowles’s re¬
search, which was published in the journal
between February and May, 1899 (Cowles
1899).
Through these papers, as well as two oth¬
ers published between 1899 and 1901,
Cowles not only established himself as one
of the pre-eminent plant ecologists of the
time, but also formulated two concepts
which, according to Cook (1980), form the
center of his theories of vegetation:
The paramount influence of the shape of the
land — the topography — on the composition
of plant communities, and the patterns of
change over time by which one plant com¬
munity succeeds another, leading gradually
to a climax formation.
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
In his studies of the dunes of Lake Michi¬
gan, Cowles noted several distinct commu¬
nities which replaced each other before ter¬
minating in a “mesophytic beech-maple de¬
ciduous forest,” which Cowles interpreted as
being the Final community type (Cowles
1899). Cowles noted that each successive
community altered the conditions of the
dune, transforming what was originally a dry
environment into a much more mesic envi¬
ronment (Cowles 1899).
In 1901, Cowles applied Warming’s
methods to the much broader area of “The
physiographic ecology of Chicago and vicin¬
ity; a study of the origin, development, and
classification of plant societies.” Published in
the Botanical Gazette and called “a landmark
in the developmental study of vegetation”
(Clements 1916), the article set forth
Cowles’s “physiographic theory” of vegeta¬
tion. This held that, along with moisture
content of the soil, topographic differences
were necessary to permit a variety of plants
to grow at similar moisture levels (Cowles
1901).
Cowles classified the landforms of Chi¬
cago and vicinity into different series and
proceeded to describe the general succession
pattern in each (Cook 1980; Cowles 1901).
Cowles considered different plant species
occurring on different landforms with an
identical degree of soil moisture to be rem¬
nants of previous successive stages in the
community development (Cowles 1901).
Stressing that the past history of a landform
had to be considered in any study of veg¬
etation, Cowles stated that “the laws that
govern changes in plant societies are mainly
physiographic; whether we have broad flood
plains, xerophytic hills, or undrained
swamps depends on the past and present of
the ever-changing topography. The idea of
constant change must be strongly empha¬
sized” (Cowles 1901).
14
After 1901, Cowles dedicated most of his
time to teaching, earning a full professorship
in 1915 and becoming chairman of the De¬
partment of Botany in 1925 (Humphrey
1961). As Gleason (1953) states, “[Cowles’s
students] were Cowles’s chief contribution
to ecology. Go through his printed works
and you will soon see that his only impor¬
tant scientific contributions were in his few
papers on succession.”
While Coulter and Cowles were shaping
plant ecology in Chicago at the turn of the
century, Charles Edwin Bessey (1845-1915)
was applying his ideas and experience in de¬
veloping the botany program at the Univer¬
sity of Nebraska.
Bessey began his studies in civil engineer¬
ing, but soon concentrated his efforts on
botany (Humphrey 1961). Like his contem¬
porary, Coulter, Bessey worked his way
West, serving on the faculty of Iowa’s Col¬
lege of Agriculture for fifteen years before
being invited to the University of Nebraska
to establish their Department of Botany
(McIntosh 1976). Bessey, too, was a student
of Asa Gray and received much of his ad¬
vanced botanical training from Gray at
Harvard (Rodgers 1944).
One of Bessey’ s key undertakings was the
formation of the Botanical Seminar (Wor-
ster 1977). Originally a rather loose associa¬
tion of undergraduate students, the seminar
became formalized by the early 1890s and
was given the task of cataloguing all the na¬
tive vegetation of the state before it fell to
the plow (McIntosh 1985). This became the
start of the Nebraska Survey, lauded by
George Vasey in 1893 as “setting an ex¬
ample, which if followed by other States, will
soon give us a complete botanical Survey of
the Country” (Rodgers 1944).
The students in the seminar were an in¬
credibly strong group, most of whom came
to botany from other fields and who would,
TRANSACTIONS
BLUMENFELD: Plant ecology comes of age in the United States
eventually, form the basis for one branch of
ecological thought. This school of thought
was characterized by a holistic/organismic
view of vegetation, much in the tradition of
Drude, and was influenced greatly by
Nebraska’s topography (or lack thereof) and
the nature of the land-grant college (McIn¬
tosh 1985). As Worster (1977) points out,
while Cowles and Gleason observed succes¬
sion on the individual sand dunes of Lake
Michigan, Pound and Clements concen¬
trated on the whole of Nebraska.
Roscoe Pound (1870-1964) was one of
Bessey’s students in the seminar. He began
his studies in law, took degrees in botany,
and ended up as dean of the Harvard School
of Law where he established a reputation as
a legal scholar (Pound 1954; Sears 1969).
Pound, along with J. G. Smith, started the
Botanical Survey of Nebraska and the Flora
of Nebraska, and Pound credits himself
with the idea of a Phytogeography of Ne¬
braska (Pound 1954). However, after 1901
and the publication of The Phytogeography
of Nebraska , Pound was forced to devote all
his time to law.
Another Bessey student was Frederick
Edward Clements (1874-1945), called “the
greatest individual creator of the modern sci¬
ence of vegetation” (Worster 1977) and “the
outstanding ecologist of his time and gen¬
eration” (Phillips 1954). Among his many
accomplishments, Clements is credited with
developing the classical theory of succession,
a view which dominated plant ecology well
into the twentieth century and which is still
used in vegetation classification today
(Barbour, et al. 1987; Braun 1958; Whit¬
taker 1962).
Frederick Clements was born in Lincoln,
Nebraska, just down the road from the new
University of Nebraska. Clements entered
the University of Nebraska, receiving his
B.S. in 1894, his A.M. in 1896, and his
Ph.D. in 1898 (his alma mater would later
confer the honorary degree of LL.D. in
1940) (Clements I960; Humphrey 1961).
Clements eventually rose to full Profes¬
sor of Botany at Nebraska, but resigned to
accept the position of Professor of Botany
and Head of the Department of Botany at
the University of Minnesota (1905), where
he remained until 1917. From 1917 until
his death in 1945, he was associated with the
Carnegie Institution of Washington, D.C.,
where he concentrated his work on soil con¬
servation and the ecology of the West
(Barbour et al. 1987; Pool 1954).
Clements’s first work in vegetation was as
a student in Bessey’s Botanical Seminar, to
which he was the first undergraduate admit¬
ted in 1892 (Tobey 1981). At that time,
Pound was one of the graduate leaders of the
seminar and was attracted to Clements “by
his zeal, ability and diligence,” recommend¬
ing him for admission and putting Clements
to work on the flora of Nebraska (Pound
1954).
By 1896, Pound and Clements were
working together on the fungi of Nebraska.
At this time, Pound got the idea of a phy¬
togeography of the state. The resulting work,
The Phytogeography of Nebraska, was pub¬
lished in 1898, partly as Clements’s Ph.D.
dissertation, and gained recognition as an
important new work in the field (Cowles
1898; McIntosh 1976). It is interesting to
note that one of the main reviews of the
book is by Cowles in Botanical Gazette
(1898, vol. 25), who considers the work “the
pioneer work of its kind in America.”
Cowles also brings up a subject that would
be a sticking point between plant ecologists
and Clements throughout his publications —
his creation of new terms and words “in
place of the simpler and more expressive
English equivalents” (Cowles 1898).
The Phytogeography of Nebraska deals
Volume 81 (1993)
15
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
with a study of plant formations (as opposed
to the Warming concept of plant societies),
which Pound and Clements describe as ei¬
ther primitive or recent (Pound and Cle¬
ments 1898). The recent classification is fur¬
ther divided into origin by nascence (occur¬
ring only on bare areas) and origin by modi¬
fication (occurring through changes in ex¬
isting communities). However, formation by
nascence always occurs after a plant forma¬
tion is destroyed “through the agency of
fires, floods, man, etc.” (Clements 1916).
Of particular importance is the introduc¬
tion of the quadrat method of vegetation
analysis, which Pound and Clements use to
determine the structure and development of
vegetation (Clements 1916; Pound and Cle¬
ments 1898). Basically, the quadrat method
is a means for determining the composition
of vegetation in a large area by looking at
the vegetation of limited, and representa¬
tive, smaller areas (Curtis 1959; Gleason
1920; Pound and Clements 1898).
While Pound left the seminar to pursue
law, Clements continued his ecological re¬
search. Plant Succession (1916), Clements’s
eighth book, not only introduced the con¬
cept of classical succession, but became one
of the most important works on plant ecol¬
ogy, establishing a variety of concepts and a
plethora of new terms and phrases. The
work is mainly a summation of Clements’s
earlier research into the ultimate end of plant
development (called the “climax”), first
treated in The Development and Structure of
Vegetation (1904) and again in Research
Methods in Ecology (1905).
Clements was very forthright as to the
purpose of the book, stating in the introduc¬
tion that “the earlier concept of the forma¬
tion as a complex organism with a charac¬
teristic development and structure in har¬
mony with a particular habitat is not only
fully justified, but that it also represents the
only complete and adequate view of vegeta¬
tion” (Clements 1916) [emphasis added].
Succession in general is concerned with
the process of vegetation change — the way
in which populations of a particular species
are gradually replaced by populations of
other species over time, usually making the
original site more fertile and mesic (Braun
1950; Curtis 1959). In the Clementsian
view of succession, a series of different plant
communities (called “seres”) will occupy a
given site in a set pattern based on the his¬
tory of the site (Pyne 1982). Knowing the
location of a particular site, it is possible, ac¬
cording to Clements, to predict the pattern
of change from pioneer species to the ulti¬
mate “climax” community, which is con¬
trolled by climate and is self-replicating
(Clements 1936; Kormondy 1965). This
optimum community is regarded as being a
mesic forest, although Clements makes pro¬
vision for deserts and aquatic communities
(Clements 1916).
The most important aspect of this pro¬
cess is Clements’s view of the climax com¬
munity as an organism (Clements 1905).
This “super-organism” view is not new, and
is seen in the writings of Plato (viz. Timaeus )
as well as in the concept of the Balance of
Nature. As Clements describes it:
The unit of vegetation, the climax formation,
is an organic entity. As an organism, the for¬
mation arises, grows, matures, and dies. . .The
climax formation is the adult organism, the
fully developed community, of which all ini¬
tial and medial stages are but stages of devel¬
opment (Clements 1936).
Like all of Clements’s earlier works, Plant
Succession introduced a multitude of new
terms and definitions. As Gleason (1953)
states, “To Clements, an association was
soon regarded as an organism, and he really
16
TRANSACTIONS
BLUMENFELD: Plant ecology comes of age in the United States
meant organism. You know Clements’s pas¬
sion for terminology: if he had not meant
an organism , he would have coined a differ¬
ent word.” As mentioned earlier, this em¬
phasis on new terminology would be a con¬
tinuing theme in the criticism of Clements’s
work, noted by, among others, Cowles
(1898), Tansley (1935), Egler (1951), Glea¬
son (1953), Phillips (1954), and Whittaker
(1962).
Clements continued to develop and re¬
fine his ideas of succession and the climax
community, culminating with “Nature and
structure of the climax,” published in the
Journal of Ecology (1936). In this encompass¬
ing article, Clements presents the crux of
what is considered classical or Clementsian
succession. This work was followed in 1939
by Bio-Ecology , co-authored with Victor
Shelford, in which the concept of the “bi¬
otic community” comprising both plants
and animals (the “biota”) was advanced. As
Worster (1977) points out, even at this late
date, Clements was insistent that it was the
vegetation which determined the animals in
a community, not the animals which deter¬
mined the vegetation. By its recognition of
animals as well as plants in the community,
Bio-Ecology was able to unite the two fields
of animal ecology and plant ecology and
foreshadowed the development of systems
ecology in the latter half of the century.
Whittaker (1962) points out three aspects
of the Clementsian system which drew criti¬
cism: the erection of a formal system of clas¬
sification based on hypothetical dynamic re¬
lations, the unlikely character of some of the
successional relations required of the climax
theory, and the inappropriateness of the sys¬
tem to the interpretation of natural commu¬
nities. Curtis, in his landmark work The Veg¬
etation of Wisconsin (1959), criticizes in par¬
ticular the delineation by Weaver and
Clements (1938) of the “Lake Forest” cli¬
max community to cover the vegetation of
the upper Great Lakes and St. Lawrence val¬
ley, commenting that this “is an indication
of the pitfalls that may be met when at¬
tempts are made to place vegetation into a
preconceived framework without supporting
quantitative evidence.”
The Clementsian view of succession was
not the only theory being developed at this
time, and alternative theories of vegetation
development were brought forth (i.e. Braun-
Blanquet 1913; Cooper 1926; Ramensky
1926). Of these, the most influential in the
U.S. was the individualistic concept of vege¬
tation development, which was used exten¬
sively in the work of Curtis and his students
at the University of Wisconsin and most
closely identified with the work of Henry A.
Gleason (1882-1975) (Curtis 1959; Whit¬
taker 1962).
Gleason was yet another Midwesterner,
growing up in Illinois and attending the
University of Illinois (B.S. 1901, M.A.
1904) and Columbia University (Ph.D.
1906). Gleason returned to the University
of Illinois; however, in 1910 he left for the
University of Michigan and in 1919 joined
the New York Botanical Garden (McIntosh
1975).
Gleason grew up at the prairie-forest edge
and as early as 1909 was aware that factors
other than climate were influencing the tran¬
sition of prairie into forest (McIntosh 1975,
1985). Further, he recognized that different
types of communities could, and did, invade
similar areas (Gleason 1909).
After the publication of Plant Succession ,
Gleason replied with an article entitled “The
structure and development of the plant as¬
sociation” (1917). Gleason presents four
main problems with Clements’s arguments:
(1) the view of the unit of vegetation as an
organism, (2) the inclusion not only of the
climax but of all stages leading to the climax
1 7
Volume 81 (1993)
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
as part of the vegetation unit, (3) a view of
vegetation so complex that it requires the
burdensome addition of many new terms,
and (4) the exclusion of exceptions to Cle¬
ments’s views of plant community develop¬
ment “by definition” (Gleason 1917).
Gleason then proceeds to list twenty-eight
points in sketching out what he calls the “in¬
dividualistic concept of ecology.” This the¬
sis holds that “the phenomena of vegetation
depend completely upon the phenomena of
the individual [plant]” and that “individu¬
als of the same species may occupy appar¬
ently different habitats and have different
associates in different localities.” Gleason
points out that while many seeds of a par¬
ticular species may enter an area through dis¬
persal, the germination of a particular seed
is dependent on the surrounding vegetation
and the environment: those species which
are best adapted to a particular combination
of vegetation and environment will be se¬
lected. As Gleason states, “the association
represents merely the coincidence of certain
plant individuals and is not an organic en¬
tity of itself.”
One important concept raised by Gleason
in his article is the idea of transition zones
between vegetation associations. These zones
are areas of species mixing where one asso¬
ciation ends and another begins, much like
the tension zone presented by Curtis for the
state of Wisconsin (Curtis 1939). This is in
contrast to the Clementsian view, which
holds that one would find no such mixing,
but, rather, a more abrupt shift between
communities as one climax turns into an¬
other (Curtis 1959; Gleason 1917; Whit¬
taker 1962).
Finally, Gleason challenges Clements on
the unidirectional trend of the climax, stat¬
ing that while forest may succeed prairie, the
opposite is also seen, termed retrogressive
succession by Gleason. Clements denies the
existence of these reversals and, according to
Gleason, excludes them from his system
through definition. Gleason formalized his
individualistic concept in his landmark ar¬
ticle “The individualistic concept of the
plant association” (1926).
Gleason states that the two factors which
most influence the structure of a plant asso¬
ciation are the environment and the sur¬
rounding vegetation. While the vegetation
of a particular area may show a great deal
of homogeneity (such as the Wisconsin por¬
tion of the Mississippi Valley), this unifor¬
mity is lost when one takes a larger geo¬
graphic view (such as the entire Mississippi
Valley).
It is this diversity in space which forms
the basis for the individualistic concept of
the plant association: “The plant individual
shows no physiological response to geo¬
graphical location or to surrounding vegeta¬
tion per se, but is limited to a particular com¬
plex of environmental conditions, which
may be [affected] by the vegetation” (Glea¬
son 1926). That is, a viable seed which mi¬
grates to an area of favorable environmental
and vegetation conditions will germinate,
while those falling into unfavorable condi¬
tions will not. To Gleason, chance, rather
than a predetermined series, plays the cru¬
cial role in the appearance of the commu¬
nity.
To account for succession, Gleason pre¬
sents a model where, because of changes in
the environment, older species find it in¬
creasingly difficult to propagate. At the same
time, seeds of outside species are constantly
entering. Eventually the environment will
pass the physiological limits of the old spe¬
cies and become favorable to the migrants,
which proceed to propagate rapidly and thus
change the form of the community. As
Gleason states, “the next vegetation will de¬
pend entirely on the nature of the immigra-
1 8
TRANSACTIONS
BLUMENFELD: Plant ecology comes of age in the United States
tion which takes place in the particular pe¬
riod when environmental change reaches the
critical stage” (Gleason 1926).
To say that Gleason’s concepts generated
criticism would be an understatement. As
Gleason recounts, his concept was debated
in 1926 at the International Botanical Con¬
gress and met with sharp disagreement and
ridicule (Gleason 1953). By the end of the
conference, the taxonomists still supported
Gleason, but the ecologists would have no
part of him, in effect ostracizing Gleason for
about ten years (Gleason 1953).
As McIntosh (1975) points out, until the
late 1940s, Gleason’s concepts were essen¬
tially ignored in the popular texts of the pe¬
riod, appearing in only one textbook, Plant
Ecology (McDougall 1927), and then barely
mentioned. However, in 1947, F. E. Egler,
S. A. Cain, and H. L. Mason all published
articles in Ecological Monographs strongly
supporting Gleason’s individualistic concept
of the plant association. It is interesting to
note that this revival of the individualistic
concept occurred only after the death of
Clements in 1945.
By the time the articles came out, Gleason
had stopped publishing articles on ecology
(his last strictly ecological article appeared in
1939) and devoted his efforts to taxonomy,
a field in which he excelled and found more
support (Steere 1958). McIntosh (1975)
notes that Gleason is probably the only per¬
son who is cited in bibliographies as both
the author of a major ecological concept and
the source of the plant nomenclature used.
The works of the European ecologists ex¬
erted great influence on the development on
the Clementsian and Gleasonian theories
and concepts. In fact, it appears that an in¬
dividualistic concept of the plant association
was put forth in a number of countries at
about the same time: Ramensky in Russia,
Negri in Italy, and Lenoble in France (Kor-
mondy 1965; McIntosh 1975). Ramensky’s
ideas are remarkably similar to those of
Gleason; he presents a concept of vegetation
continuity and species individuality in his
“Die grundgesetzmassigkeiten im aufbau der
vegetationskecke,” published in 1926, the
same year as Gleason’s article. Ramensky
also states that observation of communities
cannot be done by a reduction of the com¬
munity to small units. Rather, he suggests
the use of statistical surveys of greater areas
and the averaging of these surveys, which
would include such factors as frequency and
abundance, a method which bears striking
resemblance to that used by Curtis in The
Vegetation of Wisconsin (1959) (Kormondy
1965).
The Clementsian school also had its Eu¬
ropean supporters, including Sukachev (or
Sukatchew) in Russia, who likewise viewed
the community as a distinct entity. It ap¬
pears that the Ramensky/Sukachev differ¬
ence was essentially a Russian version of the
Gleason/Clements arguments, with Suka¬
chev taking a dominant role and the works
of Ramensky being suppressed for quite
some time after publication (Kormondy
1965; McIntosh 1975).
Clements and Gleason were not the only
ecologists in America making headway in
plant ecology during the first half of the cen¬
tury. As Gleason points out, a number of
ecologists contributed to the establishment
of plant ecology (Gleason 1953). Two who
made large contributions to plant ecology at
the time are John W. Harshberger and Wil¬
liam S. Cooper.
John W. Harshberger (1869-1929) spent
his academic career in Philadelphia and con¬
centrated his studies primarily on plant ge¬
ography (Humphrey 1961). By far his great¬
est contribution to plant geography is the
authorship of volume 13 of Vegetation der
Erde: Phytogeographic Survey of North Amer-
Volume 81 (1993)
19
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
ica (1911). While many considered this
work unreliable and inaccurate, Egler (1951)
points out that this work “presents an over¬
all picture of what vegetation is, rather than
what the successions and climax theoretically
are.”
Among Harshberger’s more regional
studies are “An ecological study of the New
Jersey strand flora” (1900) and “An ecologi¬
cal study of the flora of mountainous North
Carolina” (1903). Of particular interest in
the “North Carolina” article is the recogni¬
tion of shade tolerant and shade intolerant
trees, which Harshberger proceeds to clas¬
sify. Like Curtis (1959), Harshberger recog¬
nizes sugar maple as being the most shade
tolerant while white oak is placed near the
bottom of the tolerance list. Overall,
Harshberger recognizes that the vegetation
of an area becomes stratified due to chang¬
ing conditions of moisture and light.
Whittaker (1962) credits this article as be¬
ing one of the first to distinguish the for¬
mation and the association in their modern
senses.
William S. Cooper, a student of Cowles,
directed his research at the University of
Minnesota toward studying succession and
the mechanisms behind the climax commu¬
nity (McIntosh 1985). While Cooper stud¬
ied succession throughout the United States,
his very detailed studies on Isle Royale are
of particular note to the Midwest.
After studying trees and vegetation, Coo¬
per reached the conclusion that a linear suc¬
cession did not necessarily exist. Rather, veg¬
etation was in the form of a “flickering mo¬
saic” in a state of continuous change brought
on by small disturbances, especially windfalls
(Cooper 1913, McIntosh 1985).
Cooper’s ideas of vegetation change are
most succinctly stated in “The fundamen¬
tals of vegetative change” (Cooper 1926).
Cooper emphatically rejects the Clementsian
concept of the super-organism to represent
the development of the plant community,
stating that while the concept of an organ¬
ism is convenient in the descriptive sense, its
application to vegetation in a biotic sense is
totally unwarranted.
To describe the development of the plant
community, Cooper proposes a model of a
braided stream. Cooper begins with two pre¬
mises: (1) the universality of change, that is,
a study of vegetation must include not only
the present but also the past, and (2) the
field of study must include all types of veg¬
etation change. Cooper then likens the veg¬
etation of the earth to a flowing stream,
which is composed of braids and which has
its headwaters in the distant past. While
many small streams strive to take an indi¬
vidualistic path, the merging and simplifi¬
cation is balanced by a growing diversifica¬
tion in species. These changes may be fast
or slow, with the so-called climax commu¬
nity representing a stream which is chang¬
ing at an imperceptibly slow rate. As Coo¬
per summarizes, “Vegetational change is due
to the interaction of changing organisms and
changing environment, just as the contour
of a stream is continually modified by the
interaction of its changing current and
changing banks.”
This, then, was the state of plant ecology
at the end of the first quarter of the twenti¬
eth century. Establishment of the Ecologi¬
cal Society of America in 1915 did much to
advance the spread of research in the field
by bringing together both plant and animal
ecologists, while the journals of the Society,
Ecology and Ecological Monographs , served as
the American outlets for ecological research
(Sears 1969).
From its first formal beginnings at Chi¬
cago and Nebraska, plant ecology literally
burst upon the scientific consciousness in the
twentieth century, not only establishing it-
20
TRANSACTIONS
BLUMENFELD: Plant ecology comes of age in the United States
self as a formal and respected field of pro¬
fessional study, but supplying the initial
paradigms of thought — the Clementsian
and Gleasonian views of community devel¬
opment. Supplemented by the theories of
Harshberger and Cooper, among others,
plant ecology entered the next phase of its
development in the United States. The Uni¬
versity of Wisconsin played a key role in
this next phase, with the works of Curtis,
McIntosh, and Cottam helping to synthe¬
size the Clementsian and Gleasonian views
in the creation of a more modern view of
community development aided by math¬
ematical and statistical analysis.
Acknowledgments
The author wishes to express his apprecia¬
tion for the advice and input given by Dr.
Calvin DeWitt, Dr. Elliott Sober, and Dr.
Edward Beals (retired) of the University of
Wisconsin-Madison. Sincere thanks to Bar¬
bara Borns and the staff of the Institute for
Environmental Studies for their help in the
production of this project.
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Clements, E. S. 1960. Adventures in ecology. New
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Clements, F. E. 1904. The Development and
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vegetation on the sand dunes of Lake Michi¬
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281-308, 361-91.
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Curtis, J. T. 1959. The Vegetation of Wisconsin.
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Egler, F. E. 1947. Arid southeast Oahu vegeta¬
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435.
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plant ecology based on the textbooks of
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Gleason, H. A. 1909. Some unsolved problems
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284-307.
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Joshua C. Blumenfeld received his M.S. in Land
Resources from the Institute for Environmental
Studies at the University of Wisconsin-Madison.
His previous research has dealt with plant commu¬
nity ecology and the history of plant ecology in the
United States.
22
Quentin J. Carpenter and Calvin B. DeWitt
The effects of ant mounds
and animal trails on vegetation pattern
in calcareous fens
Plant communities are not distributed randomly over the
landscape, but are correlated with various climatic, topo¬
graphic, geologic and biotic factors, including many anthro¬
pogenic features (Jenny 1980). Similarly, most plant commu¬
nities are not homogenous, and the distribution of species
within a community is correlated with special niches
(Whittaker 1970). In calcareous fens, some previous authors
(e.g., Frederick 1974; Reed 1985; Boyer and Wheeler 1989)
have noted a two-part pattern in many (but not all) sites. This
pattern consists of a zone of taller herbaceous vegetation,
termed “Fen Meadow” (Frederick 1974), contrasted with a
short zone, described as “Marl Meadow” (Fredrick 1974) or
“Discharge Window” (Reed 1985). The “Marl Meadows” of¬
ten include many of the “Fen Meadow” species in depauper¬
ate form, but they also include other uncommon or rare spe¬
cies, not found generally distributed on the fen. Recent stud¬
ies suggest that this tall-short contrast may be due to geochemi¬
cal processes which affect the availability of phosphate and po¬
tassium to the plants (Boyer and Wheeler 1989; Wassen et al.
1990).
In observing a number of fen communities in southern Wis¬
consin, we have noticed two other possible sources of vegeta¬
tion heterogeneity, both of them biotic: ant mounds and mam¬
mal trails. Ant mounds, while not usually common on fens,
are very noticeable, often rising 1 0 to 30 cm above the general
surface and bristling with vegetation. A few ant mounds are a
meter in diameter, but most tend to be less than 0.5 m across.
Bruskewitz (1981) found that the locations of ant mounds were
positively correlated with the occurence of shrubs on the Wau-
besa Peat Mound Fen in southern Wisconsin, but did not ex-
TR ANSACTI ONS
Volume 81 (1993)
23
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
amine the relationships of other species to
ant mounds. The species of ant which builds
mounds on calcareous fens in this area is
Formica montana [= Formica cinerea mon-
tana ] , the same species which is responsible
for most mound building on Wisconsin
prairies (Greg Henderson, pers. comm.,
1988). Studies of this species at a wet prai¬
rie site in southwestern Wisconsin found
that hydraulic conductivity and nutrient
availability were higher in the ant mounds
than in the surrounding gleyed soil; vegeta-
tion-ant relationships were not studied
(Dening et al. 1977).
The mammal trails through the zone of
tall vegetation (“Fen Meadow”) appear to be
made by deer, muskrats, raccoons, and hu¬
mans. Our observations suggest that these
trails often contain species which we more
commonly find in the “Marl Meadows”
zone; in fact, animal trails are often the only
place where we noted these species in fens
where no clear “Marl Meadow” zone was
identifiable. The present study was designed
to test our perceptions by obtaining quanti¬
tative vegetation data to answer the follow¬
ing two questions:
1 . Is the vegetation found on or near fen
ant mounds significantly different from that
of the fen in general?
2. Is the vegetation found on or near trails
made by animals across fens significantly dif¬
ferent from that of the surrounding fen?
Methods
All ant mounds on three calcareous fens in
Walworth County, Wisconsin, were marked
while collecting general vegetation data from
these sites for a related study (Carpenter
1990 and unpublished). Nine ant mounds
were located on Bluff Springs Fen I (BSF-
I), 18 on Bluff Springs Fen II (BSF-II), and
15 on Clover Valley Fen (CVF) (Fig. 1).
During late summer or early fall, the vege¬
tation on and immediately surrounding each
of these ant mounds was surveyed using a
square 1 m2 quadrat frame centered on the
ant mound, while the general vegetation of
the fens was surveyed using the same appa¬
ratus randomly placed within cells of a grid
system which covered most of each fen (Car¬
penter 1990). Species abundance data were
recorded as per cent cover. After arc-sin
square root transformation, means for each
species found in quadrats centered on ant
mounds on a particular fen were compared
to means of the same species obtained from
the general survey of the same fen using an
unpaired t-test (Snedecor and Cochran
1980). Because the ant mound surveys and
the general vegetation surveys were not con¬
ducted at the same time (time differences
varied from two to four weeks depending on
the site), a conservative standard of differ¬
ence (p < .01) was adopted recognizing that
the relative abundances of many species
change gradually throughout the season.
To determine if the vegetation on or very
near animal trails differed from the adjacent
fen vegetation, 15 quadrat pairs were read
on BSF-I on 16 September 1988 and 13
quadrat pairs on BSF-II on 30 September
1988; a distinct animal trail of sufficient
length to provide at least 1 0 paired sampling
locations could not be located on CVF. The
1 m2 quadrat pairs shared an edge and were
spaced at 5 m intervals along a fisherpersons’
trail (BSF-I) and a deer trail (BSF-II) which
crossed the respective fens east-west. At each
sampling point, the quadrat frame was first
read centered on the trail. It was then flipped
over, to the south at the first sampling point,
to the north at the second, etc., such that
1 5 (or 1 3) vegetation samples were obtained
centered on the trail, and 15 (or 13) samples
were obtained centered 1 m off the trail in
24
TRANSACTIONS
CARPENTER and DeWITT: Effects of ant mounds on vegetation pattern
Fig. 1. Location of study site in Wisconsin and Walworth County. Source: State Cartog¬
raphers Office, Madison.
Volume 81 (1993)
25
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
nearly equal numbers on either side of the
trail. As in the ant mound survey, abundance
data were collected as per cent cover, then
arc-sin square root transformed for statisti¬
cal analysis. Because of the small sample size,
only those species which attained at least
0.5% mean cover were subjected to statisti¬
cal analysis. Means were compared using a
paired t-test (Snedecor and Cochran 1980).
Since the paired design of this part of the
study allowed better control of time and dis¬
tance disparities, a less rigorous standard of
difference (p < 0.05) was used.
Results
Table 1 summarizes the results of the gen¬
eral fen vegetation versus ant mound vegeta¬
tion study. Only one species, Muhlenbergia
mexicana , was found to be significantly more
common (at p < 0.01) on or near ant
mounds when all three fens were lumped.
Three other grasses, Andropogon gerardii,
Bromus ciliatus and Sorgastrum nutans, met
a lower standard (p < .05) on at least two of
the three fens, suggesting a possibly weaker
association with the ant mounds. No broad-
leaf herb or shrub met either standard.
Tables 2 and 3 present summaries of the
data from the mammal trail surveys for BSF-
I and BSF-II. Thirty six taxa met the 0.5%
mean cover criterion on at least one fen; fif¬
teen of these taxa were shared by both fens.
The data from Tables 2 and 3 suggest that
38% of the taxa considered from BSF-I and
44% of those considered from BSF-II were
affected significantly (p < 0.05) by the prox¬
imity of trails. Nevertheless, while each fen
contained many species affected by trails,
only three taxa ( Lobelia kalmii, Parnassia
glauca, and bare ground) were significantly
more common on trails in both fens; Par¬
nassia glauca was strongly favored (p < 0.01)
on both sites. Only detritus was negatively
associated (p < 0.05) with trails on both fens.
Disregarding significance, eleven of the fif¬
teen taxa in common varied monotonically
(i.e., the taxon increased or decreased on
both fens), while four species did not. No
taxon was significantly favored on one fen,
but significantly disfavored on the other.
Discussion and Conclusions
Muhlenbergia mexicana and perhaps some of
the prairie grasses are more common on and
Table 1 . Summary of distribution of Muhlenbergia mexicana on three fens in Wisconsin. Mean
cover on ant mounds was compared to that over the general fen using an unpaired t-test.
26
TRANSACTIONS
CARPENTER and DeWITT: Effects of ant mounds on vegetation pattern
Table 2. Summary of trail study on Bluff Springs Fen I, 26 August 1988.
Data are from 15 paired 1-m-square quadrats centered either on or 1 m from the center of a human trail.
After arc-sin square root transformation, differences in means were compared using a paired t-test. An
“F” indicates a taxon significantly more common on or near a trail (= Favored). A “D” indicates a taxon
significantly less common on or near the trail (= Disfavored).
around ant mounds than in the general fen.
From observations of the behavior of the
mound-building ant Formica montana on
the Whitewater area fens over several sea¬
sons, we suggest the following explanation
for this association: during April the ants
clear away all dead material on or near their
mound; during May and the early part of
June, they prune any green shoots which
erupt through the mound. During the sum¬
mer, however, shoots are allowed to grow
and often drape over the mound. We sus¬
pect the ants’ vegetation management is re¬
lated to thermo-regulation of the ant
mound, removing shading vegetation in the
cool spring, but allowing it to grow in the
hot summer. Whatever the reason for the
ant behavior, the repeated cutting of shoots
seems to favor late-season grasses, which of¬
ten do not emerge until June and which
flower in the late summer. Further, we have
observed that, of all the grasses found on
fens, Muhlenhergia mexicana is the last to
flower, sometimes as late as early October;
we suspect that the strong association be¬
tween this grass and the ant mounds is
Volume 81 (1993)
27
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Data are from 13 paired 1-m-square quadrats centered either on or 1 m from the center of a deer trail.
After arc-sin square root transformation, differences in means were compared using a paired t-test. An
“F” indicates a taxon significantly more common on or near the trail (= Favored). A “D” indicates a taxon
significantly less common on or near the trail (= Disfavored).
28
TRANSACTIONS
CARPENTER and DeWITT: Effects of ant mounds on vegetation pattern
somehow related to this trait. The increased
hydraulic conductivity and nutrient avail¬
ability found by Denning et al. (1977) may
also play a role in favoring grasses on or near
ant mounds.
The trail at the base of BSF-I is used pri¬
marily by persons trout fishing in Bluff
Creek and to a lesser extent by deer, rac¬
coons, and mink, while the trail on BSF-II
appears to be used mostly by deer. Most of
the species favored by the trails, such as
Carex leptalea and Lobelia kalmii , are also
small, and were classified as “competition-
intolerant species” by Zimmerman (1983).
If this classification is correct, they would be
expected to benefit from the trampling in
the center of a trail which might more ad¬
versely affect some of the larger competitors.
Others, however, such as Solidago ohioensis
and Muhlenbergia glomerata , are among the
tallest and most common herbs on these fens
(Carpenter 1990); thus, the single explana¬
tion of competition intolerance may not ap¬
ply to all species affected. We suggest that
the increased vigor of the apparently more
competitive species may be analogous to the
pattern one sees along a sheep or cattle trail
in a pasture where the grass grows tallest and
greenest just at the edge of a trail (personal
observation). The simplist explanation for
this observation is that the taller and greener
plants are far enough away from the center
of the trail to avoid trampling, yet close
enough to the trail to benefit from the ex¬
tra nutrients found in manure and urine.
The disfavored shrubs (Cornus stolonifera
on BSF-I and Potentilla fruticosa on BSF-II)
may be victims of browsing or are perhaps
simply killed by trampling when small. One
might speculate that the trails were simply
rerouted around shrubs; however, inspection
of several aerial photos of the sites (1947 to
1980) suggests that the trails are long estab¬
lished and relatively straight.
The idea of trails, especially human trails,
across natural areas is a complicated and of¬
ten emotional issue. On the one hand, the
evidence presented here suggests that trails
provide special habitat for some uncommon
competition-intolerant species; on the other
hand, trails provide access for humans to
damage the integrity of the natural area by
excessive trampling, flower picking and
inadvertant introduction of exotics such as
purple loosestrife ( Lythrum salicaria) and fen
buckthorn ( Rhamnus frangula) . Many dis¬
turbance or pioneer wetland species are com¬
monly associated with fens (Zimmerman
1983). Observations from fens in Wiscon¬
sin, Iowa, and Ohio (Zimmerman 1983;
Loeschke 1991; Denny 1991) which have
had their surfaces severely disturbed but have
maintained their former groundwater sup¬
plies suggest that many of the rare competi¬
tion-intolerant species such as Scleria ver-
ticillata, Parnassia glauca, and Rhynchospora
capillacea appear in great abundance after
disturbance. Thus, we suggest that all dis¬
turbance is not detrimental to fens and that
managers must judge specific types of dis¬
turbances on their individual ecological mer¬
its or dangers.
Works Cited
>
Boyer, M. L. H., and B. D. Wheeler. 1989. Veg¬
etation patterns of spring-fed calcareous fens:
Calcite precipitation and constraints on fer¬
tility./ Ecol. 77(2): 597-609.
Bruskewitz, James W. 1981. Wetland ants: in¬
ternal mound temperature and humidity
preferences; location and shape of mounds as
adaptations to a wetland environment. Trans.
Wise. Acad. Sci, Arts and Ltrs. 69:21-25.
Carpenter, Q. J. 1990. Hydrology and vegeta¬
tion of a calcareous peat mound fen in south¬
eastern Wisconsin. MS Thesis. University of
Wisconsin-Madison.
Dening, J. L., F. D. Hole, and J. Bouma. 1977.
Volume 81(1 993)
29
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Effects of Formica cinerea on a wetland soil
on West Blue Mound, Wisconsin. In Wet¬
lands ecology, values and impacts. Proceedings
of the Waubesa Conference on Wetlands, 2—5
June, 1977 \ ed. C. B. DeWitt, and E. Sola-
way, pp. 276— 89. Madison: Institute for En¬
vironmental Studies, University of Wiscon-
sin-Madison.
Denny, G. L. 1991. Personal Communication.
Director, Bureau of Endangered Resources,
Ohio Department of Natural Resources. Co¬
lumbus, Ohio.
Frederick, C. M. 1974. A natural history study
of the vascular flora of Cedar Bog, Cham¬
paign County, Ohio. Ohio Journal of Science.
74(2):65-ll6.
Henderson, G. 1988. Personal Communication,
Entomology Dept. UW-Madison.
Jenny, Hans. 1980. The Soil Resource. New York:
Springer- Verlag.
Loeschke, Mark. 1991. Personal Communica¬
tion. Botanist, Iowa Department of Natural
Resources. Des Moines, Iowa.
Reed, D. M. 1985. Calcareous fens of southeast¬
ern Wisconsin. MS Thesis. University of
Wisconsin-Milwaukee.
Reed, D. M. 1991. Personal Communication.
Chief Biologist, Southeast Wisconsin Re¬
gional Planning Commission. Waukesha,
Wisconsin.
Snedecor, G. W., and W. G. Cochran. 1980.
Statistical Methods, 7th ed. Ames, Iowa: The
Iowa State University Press.
Wassen, M. J., A. Barendregt, A. Palczynski, J.
T. Smidt, and H. de Mars. 1990. The rela¬
tionship between fen vegetation gradients and
groundwater flow and flooding in an un¬
drained valley mire at Biebrza, Poland. /.
Ecol. 78:1106-22.
Whittaker, R. H. 1970. Communities and Eco¬
systems. New York: Macmillan.
Zimmerman, J. H. 1983. The revegetation of a
small Yahara Valley prairie fen. Trans. Wise.
Acad. Sci, Arts, and Ltrs. 71(2): 87-102.
Quentin Carpenter is a dissertator in the Land
Resources Program of the Institute for Environmen¬
tal Studies, University ofWisconsin-Madison; his
research concerns the interaction of hydrology and
vegetation in calcareous fens of southern Wiscon¬
sin.
Calvin B. DeWitt is Professor of Environmental
Studies, Institute for Environmental Studies at the
University of Wisconsin-Madison. His teaching
and research concerns include wetland ecology, land
use ethics, and stewardship of the earth.
30
Lee Clayton and John W. Attig
Exhumed early Paleozoic landforms
on the Barahoo Hills, Wisconsin
Abstract The Baraboo Hills of southern Wisconsin consist of extremely durable Precam-
brian quartzite. Some of the well-preserved landforms seen there today were
formed during the early part of the Paleozoic, were then buried, and were sub¬
sequently exhumed in Mesozoic or Cenozoic time. Valleys in the western part
of the South Range were cut in Middle Cambrian time or earlier and buried
in Late Cambrian time. Subsummit benches and scarps were cut by marine
shore erosion in early Ordovician time and buried soon after. Summit pla¬
teaus were cut by subaerial or marine-shore processes, probably in Middle Or¬
dovician time, and buried in Late Ordovician time.
The Baraboo Hills of south-central Wisconsin (Fig. 1) con¬
tain some remarkably well-preserved Paleozoic landforms.
Late Cenozoic landforms are present, but in this paper we con¬
clude that many of the landforms were cut into the Baraboo
quartzite early in the Paleozoic, buried soon thereafter, and ex¬
humed in late Mesozoic or early Cenozoic times. These Pa¬
leozoic landforms include valleys, subsummit benches and
scarps, and summit plateaus.
Although some of these landforms were recognized more
than a century ago (Irving 1877, 504-05), they were poorly
known until studied by Thwaites (1931, 1935, 1958, 1960).
However, some elevations on the topographic maps available
to Thwaites are in error by more than 100 m, and some fea¬
tures are misplaced horizontally by as much as 1 km. As a re¬
sult, Thwaites was unable to adequately document the loca¬
tion and elevation of the landforms or to convincingly dem¬
onstrate the relationship between these landforms and the Pa¬
leozoic formations of the area.
Accurate topographic maps now exist, and the geology of
the region has been mapped in greater detail (Dalziel and Dott
1970; Clayton and Attig 1990; Attig and Clayton 1990; Attig
et al. 1990). As a result, we now are able to document the el-
TR A NS ACT I O NS Volume 81 (1993)
31
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 1 . Location of landforms of the South Range of the Baraboo Hills.
evation and the location of these landforms
and relate them more confidently to the Pa¬
leozoic stratigraphy of the region. In this
paper we reevaluate Thwaites’ speculations
about the age and origin of these landforms.
Description
The Baraboo Hills
The geology of the Baraboo Hills has been
outlined by Dalziel and Dott (1970) and
Clayton and Attig (1990). In the area sur¬
rounding the Baraboo Hills, Paleozoic rock
lies on a generally flat unconformity on Pre-
cambrian rock. Before the Paleozoic sedi¬
ment was deposited, the Baraboo Hills rose
more than 350 m above the surrounding
plain (Fig. 2). Then, as now, the hills were
made up of quartzite of the Baraboo Forma¬
tion, which was more than 1.5 km thick.
The quartzite consists of quartz sand that
underwent low-grade metamorphism to pro¬
duce a rock that is highly resistant to ero¬
sion. The Baraboo Formation was folded
into a doubly plunging syncline, resulting in
the oval pattern of hills shown in Figure lb.
The north and south halves of the oval are
called the North Range and the South
Range, respectively.
The Baraboo Hills then were buried with
quartz and lime sand during Late Cambrian
and Early Ordovician time. At least the top
of the South Range was reexposed by ero¬
sion during Middle Ordovician time, and
the hills again were buried during Late Or¬
dovician time, beginning with the quartz
sand of the St. Peter Formation. By Late
Paleozoic time, an additional few hundred
meters of marine sediment had probably
been deposited on top of the hills. Marine
deposition had ceased by late Pennsylvanian
time (Shaver et al. 1985). During the ensu¬
ing 200 million years, the land surface was
lowered to near the level of the top of the
Baraboo Hills.
The summit of South Range probably
was exposed again just before the fluvial
gravel of the “Windrow Formation” was de¬
posited on one of the highest parts of the
South Range, above the East Bluff of Dev¬
ils Lake (Fig. lc; Thwaites and Twenhofel
32
TRANSACTIONS
CLAYTON and ATTIG: Exhumed early Paleozoic landforms on the Baraboo Hills
Fig. 2. a: Composite profile of summit plateaus of the South Range (the thin, solid lines
at the top of the diagram), viewed from east to west. Vertical exaggeration x 10. The
position of the subsummit benches is shown with a heavy dashed line. The thin dashed
lines in the Baraboo quartzite indicate dip of the quartzite. The Precambrian, Cambrian,
and lowest Ordovician stratigraphy shown at the north and south flanks of the range is
based on local information, but the Platteville Formation has been projected from an
area 40 km farther south, b; Cross section through a representation plateau, with flank¬
ing valleys and valley fills (middle of the South Range; sec. 17, 20, and 28, Til N, R6E).
1921, 296-97). It is unknown when that
event occured, but guesses have generally
ranged from Early Cretaceous to Pliocene
(Thwaites and Twenhofel 1921, 307-10;
Andrews 1938; Anderson 1988, 233-36).
The landscape surrounding the Baraboo
Hills since then has been lowered about 60
m at the west end, 250 m at Devils Lake
(Fig. lc; WGNHS Geologic Logs Sk-17 and
Sk-39), and 300 m at Portage near the east
end of the hills (Fig. lb; WGNHS Geologic
Log Co-634). The stratigraphic relationships
summarized here and shown in Figure 2a in¬
dicate that the Baraboo Hills are much the
same shape today as they were in Middle
Cambrian time.
Valleys
The North and South Ranges of the Bara¬
boo Hills are irregular quartzite ridges cut
by gorges and valleys. The gorges have been
cut completely through the ranges. They had
a complex history, including considerable
Pleistocene erosion when they functioned as
spillways of glacial Lake Wisconsin (Clayton
and Attig 1989); they will not be further dis¬
cussed.
In contrast, the valleys head within the
ranges. The largest valleys in the unglaciated
part of the South Range are marked by As
in Figure lc. These are a few kilometers
long, about 1 km wide, and about 100 m
deep. They tend to be of uniform width and
abruptly terminate at broad, rounded valley
heads. The valleys are walled with Baraboo
quartzite, but their bottoms are generally
underlain by 20 to 60 m of Cambrian sand¬
stone and conglomerate (Fig. 2b).
Subsummit Benches and Scarps
Nearly flat benches have been cut into the
Baraboo quartzite on the sides of each of
these valleys (Fig. lc, 2a, and 2b). The
benches are typically a few tens of meters
wide. Above the bench is a scarp with a slope
of about 20°. Below the bench, the valley
Volume 81 (1993)
33
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
side typically slopes about 10° to 15°. The
benches are about 30 m below the edge of
the summit plateaus (Fig. 2a), and are at
nearly the same elevation throughout the
South Range, descending slightly to the
south at about 1 m/km; they are at an el¬
evation of about 402 m (1320 ft) on the
north side of the South Range and at about
393 m (1290 ft) on the south side.
The benches are generally covered by for¬
est, but, even so, in many places they are
obvious from a distance (Fig. 3a). In addi¬
tion, they are generally conspicuous in the
few places where crossed by roads, especially
when leaves are off the trees and patches of
snow remain on the bench after a thaw. The
benches can be seen where Freedom Road
descends from the summit plateau on the
south side of Fiappy Hill (4 km northeast
of the community of Denzer) and at the
junction of Tower and Denzer Roads (6 km
north of Denzer; Fig. 3b).
Summit Plateaus
The highest hill tops in the unglaciated part
of the South Range are remarkably flat (Fig.
2a, 2b, and 3c). These summit plateaus are
typically about 0.3 km wide, with a maxi¬
mum width of 1.3 km and a maximum
length of 7.5 km (Fig. 1 and 2). The middles
of the plateaus, at elevations between about
430 m (1410 ft) and 454 m (1490 ft), are
horizontal, with slopes increasing to several
degrees near the edge, at elevations between
about 421 m (1380 ft) and 433 m (1420 ft).
The plateaus are quartzite overlain by a few
meters of yellowish-brown silt and clay con¬
taining quartzite fragments. Thwaites (1935,
401; 1958, 147; 1960, 37-38) reported
scattered loose fragments of Paleozoic chert
on the plateau surface, as well as a few small
exposures of in-place lower-Paleozoic con¬
glomerate along the edge of the plateaus.
Age and Origin
Valleys
Most large valleys in the South Range are
known to have formed before Late Cam¬
brian time, because there is Late Cambrian
sandstone and conglomerate in the valley
bottoms (Fig. 2b; Dalziel and Dott 1970).
The valleys are known to have been at least
20 to 60 m deeper at the beginning of Late
Cambrian time than they are today, because
the Late Cambrian fill is that thick, and the
interfluves may have been considerably
higher, if the summit plateaus were eroded
in Ordovician time, as will be discussed.
Otherwise, the general shape of the valleys
probably has changed little, because thin
patches of Cambrian rock occur in a few
places high on the valley walls (Thwaites
1935, 401; Thwaites 1958, 147; Thwaites
1960, 38).
Most large valleys in the unglaciated part
of the South Range are shaped like typical
stream valleys. They require no explanation
other than hillslope and stream erosion
through Early and Middle Cambrian time
and perhaps also during latest Precambrian
time.
Sub summit Benches and Scarps
The subsummit benches on the South
Range are not structural terraces, because
they slope only about 1 m/km (less than
0.1°) but are cut in quartzite that dips 10°
to 40° (Fig. 2). These are not fluvial terraces
because they slope to the south, whereas the
valleys slope north and south on either side
of the range. Thwaites (1935, 401; 1958,
147-48; 1960, 38-39) concluded that these
benches and the scarps above were cut into
the quartzite by marine shore erosion. We
agree, because no other explanation seems
34
TRANSACTIONS
CLAYTON and ATTIG: Exhumed early Paleozoic landforms on the Baraboo Hills
Fig. 3. a: North edge (p) of the summit plateau northeast of Happy Hill (Fig. 1c), with
subsummit scarp (p-b) and bench (b); taken west-northwestward from middle of NE V*
sec. 22, T11N, R6E. b: West edge (p) of the summit plateau of Happy Hill, with
subsummit scarp (p-b) and bench (b); taken southward at southwest corner of SE !4
SW!4sec. 27, T11N, R5E. c: Summit plateau of Happy Hill; taken northward from the
south edge of the SE !4 SE 74 SE 74 sec. 26, T1 1 N, R5E.
plausible. However, the scarps clearly have
been rejuvenated in places by mass move¬
ment when permafrost was present during
the Pleistocene (Clayton and Attig 1990).
Although there is no direct stratigraphic
evidence, these shore benches were most
likely cut during the Ordovician. Patches of
early Paleozoic conglomerate occur locally
Volume 81 (1993)
35
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
on the slope just below the benches
(Thwaites 1958, 147; Thwaites 1960, 38).
The formations present on either flank of
the South Range can be projected into the
range (dotted lines in Fig. 2a), using the
stratigraphic and structural information of
Clayton and Attig (1990, Plates 1 and 2,
Fig. 13 and 14); the contact between the Jor¬
dan Formation (Late Cambrian) and Prai¬
rie du Chien Formation (Early Ordovician)
is about 30 m below the benches. The origi¬
nal thickness of the Prairie du Chien here is
unknown, but it is 60 m thick 40 km to the
south (WGNHS Geologic Log Dn-993) and
may have been that thick across the South
Range. If the Prairie du Chien dolomite was
deposited near sea level on the Baraboo
Hills, the benches would have been cut be¬
fore the upper Prairie du Chien was depos¬
ited, during Early Ordovician time, as sug¬
gested by Thwaites (1935, 401; 1958, 147;
1960, 38). This conclusion is corroborated,
in a general way, by the dip of the benches;
as shown in Figure 2a, the benches slope
southward about 1 m/km, about the same
as the regional dip of the Prairie du Chien
Formation in the Baraboo area (Clayton and
Attig 1990, Fig. 14).
If the benches were eroded at that time,
much eroded quartzite would be predicted
to occur in the upper part of the Prairie du
Chien Formation. However, we know of no
sedimentological evidence for increased ero¬
sion and deposition of quartzite around the
Baraboo Hills at that time, because appro¬
priate exposures are unavailable — no more
than about the lower 25 m of the Prairie du
Chien Formation is exposed in the Baraboo
region.
Other less conspicuous benches and
scarps occur at lower elevations on both the
North and South Ranges. Wanenmacher
(1932, 75-76) and Raasch (1958) inter¬
preted these as marine shore terraces formed
in Late Cambrian time, and we agree with
this interpretation.
Summit Plateaus
If the top of the South Range had been
rounded or marked by a series of hogbacks
over harder layers in the northward-dipping
quartzite, the shape of the range would re¬
quire little explanation other than normal
subaerial erosional processes operating over
a long period of time. However, the sum¬
mit plateaus, crosscutting the dipping
quartzite, require some special explanation.
Before Thwaites studied them, the summit
plateaus were considered the remnants of a
peneplain, or at least of a subaerial erosion
plain.
Martin (1916, 68) and Smith (1931,
128), and others, thought this was a pene¬
plain cut in Precambrian time. Thwaites
(1935, 398; 1958, 141; 1960, 37), however,
thought the plateaus were cut after Precam¬
brian time, doubting that the Precambrian
surface surrounding the Baraboo Hills could
have been lowered 350 m without destroy¬
ing the erosion-surface remnants on the hills.
Furthermore, the summit plateaus seem to
slope southward at about the same inclina¬
tion as the subsummit benches and the Prai¬
rie du Chien Formation (1 m/km; Fig. 2a);
if the summit plateaus were cut before the
Precambrian plain surrounding the Baraboo
Hills, they should slope south at least as
steeply as that plain (2 to 4 m/km; Thwaites
1957).
Others, such as Trowbridge (1917, 352-
53), suggested that the summit plateaus are
remnants of a plain (the “Dodgeville pene¬
plain”) cut by subaerial processes when the
South Range was being exhumed in Meso¬
zoic or Cenozoic time. Thwaites (1935, 403;
1958, 149; 1960, 37) argued that if subaerial
erosion was capable of planing the extremely
36
TRANSACTIONS
CLAYTON and ATTIG: Exhumed early Paleozoic landforms on the Baraboo Hills
resistant quartzite, erosion should also have
been capable of planing the much weaker
Paleozoic dolomite and shale of Blue
Mounds (40 km south of the Baraboo
Hills), one of which is 70 m higher than the
South Range. Thwaites argued further that
scattered loose blocks of Paleozoic chert on
the plateaus are an indication of much less
erosive activity than would have been re¬
quired to plane the quartzite from the top
of the South Range. The patches of lower-
Paleozoic conglomerate on the plateaus also
indicate they could not have been formed
in Mesozoic or Cenozoic time.
As indicated in the discussion of the sub¬
summit benches, if the formations on either
side of the South Range are projected into
the range (Fig. 2a), the position of the
middle Ordovician unconformity (at the
base of the St. Peter Formation) is unclear,
but it may be near the level of the edges of
the summit plateaus. This suggests the pos¬
sibility that the summit plateaus were cut by
subaerial erosion during this hiatus, which
lasted about 25 million years (Shaver et al.
1985). This possibility suffers from none of
the objections listed for Precambrian and
post-Paleozoic subaerial erosion plains, but
much less time was available. In addition,
the summit plateaus seem too flat to corre¬
spond to the middle Ordovician uncon¬
formity, which is known to have consider¬
able local relief near the Baraboo Hills; in
some places the unconformity is as low as
or even below the base of the Prairie du
Chien Formation (suggested in left-hand
side of Fig. 2a; Clayton and Attig 1990).
Thwaites (1931, 745; 1935, 401-02;
1958, 145-47; 1960, 36-38) suggested that
the summit plateaus on the South Range are
the result of marine shore erosion rather
than subaerial erosion. Thwaites favored this
interpretation for the following reasons. If
subaerial erosion is ruled out, marine ero¬
sion is the only reasonable alternative, and
marine shore erosion seems more capable of
eroding the quartzite than any other process.
Once the subsummit bench had been inter¬
preted to result from shore erosion, it was
reasonable to extend this interpretation to
the summit plateaus. However, a shore plain
might be expected to be even flatter than the
plateaus on the South Range, although
Thwaites suggested that sea level gradually
rose as the plain was cut.
The age of the summit plateaus, if in fact
they are marine shore terraces, is less clear
than that of the subsummit benches.
Thwaites (1935, 401; 1958, 147; 1960, 37)
suggested that if the Paleozoic formations
farther south are projected into the South
Range, the summit plateaus would coincide
with the base of the Platteville Formation
(Late Ordovician). Our projection (Fig. 2a)
shows this also, but because the original
thickness of the Ordovician units is uncer¬
tain here, this projection could be in error.
No sedimentological evidence is available for
increased Platteville erosion and deposition
around the Baraboo Hills, because there are
no Platteville outcrops in the area.
Conclusion
The Baraboo Hills retain early Paleozoic
landforms that have undergone little change
since they were exhumed in Mesozoic or
Cenozoic time. The large valleys in the
South Range (except Devils Lake gorge) are
normal stream valleys that formed before the
Late Cambrian. The subsummit benches
and scarps are almost certainly a marine
shore terrace, which probably formed dur¬
ing the Early Ordovician. The summit pla¬
teaus are remnants of a plain formed by ma¬
rine shoreline erosion or by subaerial ero¬
sion, possibly during either the Middle or
Late Ordovician.
Volume 81 (1993)
37
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Acknowledgments
We wish to thank David M. Mickelson, W. N.
Melhorn, and Thomas W. Gardner for review¬
ing an earlier version of this paper.
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Ninth Annual Field Conference Guide Book ,
ed. A. C. Trowbridge, 395-404.
- . 1957. Buried pre-Cambrian of Wiscon¬
sin. Wisconsin Geological and Natural His¬
tory Survey [Page-Size Map Series].
- . 1958. Land forms of the Baraboo dis¬
trict, Wisconsin. Transactions of the Wiscon¬
sin Academy of Sciences, Arts and Letters 47:
137-59.
- . I960. Evidences of dissected erosion
surfaces in the Driftless Area. Transactions of
the Wisconsin Academy of Sciences, Arts and
Letters 49: 17-49.
Thwaites, F. T., and W. H. Twenhofel. 1921.
Windrow formation; an upland gravel forma¬
tion of the Driftless and adjacent areas of the
upper Mississippi valley. Geological Society of
America Bulletin 32: 293-314.
Trowbridge, A. C. 1917. The history of Devils
Lake, Wisconsin. Journal of Geology 25: 344-
72.
Wanenmacher, J. M. 1932. Paleozoic strata of
the Baraboo region, Wisconsin. Ph.D. diss.,
University of Wisconsin, 105 p.
Lee Clayton is a Professor and John W. Attig is
an Associate Professor of the University of Wiscon¬
sin (Extension) at the Wisconsin Geological and
Natural History Survey in Madison.
38
Philip A. Cochran, Martin E. Sneen,
and Alan P. Gripentrog
Notes on the biology
of the American brook lamprey
(Lampetra appendix) in Wisconsin
Abstract American brook lampreys (Lampetra appendix) were collected from Taylor
Creek in Rock County and were documented for the first time from Jambo
Creek in Manitowoc County. Spawning at both sites occurred in early May at
lower water temperatures (12-14°C) than previously recorded in Wisconsin.
Although most spawning groups occurred in the open on gravel substrate, as is
typically reported of spawning by lampreys, some spawning groups were found
beneath cover. The sample of adult lampreys from Taylor Creek included a
statistically significant excess of males. Adult male lampreys had relatively larger
oral discs than females, whereas females displayed swelling along the leading
edge of the second dorsal fin. A review of previous studies indicated that mean
total lengths of adult males tend to be greater than those for females, although
differences between means are rarely statistically significant at individual sites.
Where the two species of nonparasitic lampreys have been collected from the
same stream systems in Wisconsin, American brook lampreys occur upstream
from northern brook lampreys (Ichthyomyzon fossor) significantly more of¬
ten than vice versa.
The American brook lamprey ( Lampetra appendix) is widely
distributed in eastern North America (Rohde 1980). Al¬
though it has been studied in other parts of its range (e.g., Hoff
1988; Lanteigne et al. 1981; Rohde et al. 1976; Seagle and
Nagel 1982), little information on this species has been col¬
lected in Wisconsin (Becker 1983). Since the time that Becker’s
(1983) account was prepared, additional references to Ameri¬
can brook lampreys in Wisconsin have been confined prima¬
rily to locality records (Cochran 1984; Fago 1982, 1983,
1984a, 1984b, 1983a, 1983b, 1986, 1992). The purpose of
this note is to report new data on the biology of the American
brook lamprey in Wisconsin, including several topics absent
from or incompletely considered in Becker’s (1983) account.
TRANSACTIONS Volume 81 (1993)
39
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Methods
We collected adult lampreys in breeding
condition at two sites. Taylor Creek is lo¬
cated in the Rock River basin (Mississippi
River drainage) in Rock County (T2N,
R10E, Sec. 30/31). At the site of capture,
the West Church Road crossing, stream
width was 4-3 m, the water was clear and
up to 1 m deep, and the bottom was pri¬
marily sand, except for a concentration of
rock slabs, cobble, and gravel beneath the
bridge. Jambo Creek is a tributary to the
East Twin River (Lake Michigan drainage)
in Manitowoc County (T21N, R23E, Sec.
26). Stream width was 4-5 m, depth was
15-100 cm, and the water was clear with a
slight reddish stain. Several gravel-bottomed
riffles were present. American brook lam¬
preys have been reported previously from
Taylor Creek (Fago 1982) and from the East
Twin River (Fago 1985b) but not from
Jambo Creek (Fago 1985b).
Samples of lampreys were taken to the
laboratory, where they were anesthetized
with tricaine methanesulfonate (MS-222),
measured for total length to the nearest mm,
and weighed to the nearest 0.01 g. Oral disc
lengths of lampreys from Jambo Creek were
measured by pressing them flat against a
transparent rule. Voucher specimens were
placed in the University of Wisconsin-Mad-
ison Zoology Museum (Taylor Creek:
UWZM 8432, Accession No. 84-77; Jambo
Creek: UWZM 9951, Accession No. 91-
176). Statistical analyses of morphological
data were conducted with MINITAB (Schae¬
fer and Anderson 1989).
It has been stated that American brook
lampreys tend to be found upstream from
northern brook lampreys (Ichthyomyzon
fossor) when the two species occur in the
same stream system (Morman 1979). We
tested the applicability of this conclusion to
Wisconsin waters with distribution maps
provided by Fago (1983, 1984a, 1984b,
1985a). By overlaying transparencies of the
maps for the two species, it was possible to
tally cases in which one species occurred up¬
stream of the other or vice versa. Deviations
from random were tested through use of the
binomial expansion (Sokal and Rohlf 1981).
Results
At Taylor Creek, 1 1 American brook lam¬
preys were collected on May 4, 1984, at a
water temperature of 14°C. All were cap¬
tured beneath the bridge; 8 of the 1 1 were
captured after a single individual was ob¬
served next to a rock slab and that slab was
overturned. Ten of 1 1 lampreys were male,
a result significantly different from expected
under the null hypothesis that both sexes are
equally abundant and equally vulnerable to
capture (binomial test, p = 0.012). The
males were readily made to express a rather
transparent fluid from their urogenital pa¬
pillae; the milt of spawning male American
brook lampreys was described by Dean and
Sumner (1897) as nearly colorless. Many
individuals displayed the sorts of abrasions
and other minor wounds that result from
spawning activity.
At Jambo Creek, we observed spawning
lampreys in 1988, 1989, and 1992. Several
spawning groups were detected on May 2,
1988, but only one lamprey was found on
May 4 at the same site. Water temperature
was not measured on May 2, but was 16°C
on May 4. In 1989, spawning groups were
not observed on April 22, April 26, or April
30, but were present on May 3 at a water
temperature of 13°C. All spawning groups
were located just above riffles. Five groups
were on open gravel substrate; individual
group sizes were 6, 6, 7-10, 10-15, and 20-
30. In addition, a group of ten lampreys was
40
TRANSACTIONS
COCHRAN et a I . : The biology of the American brook lamprey in Wisconsin
building a spawning depression beneath an
overhanging stump, and a group of un¬
known size was building a pit beneath a large
rock slab. Ten lampreys, five of each sex,
were collected from among the spawning
groups. On May 4, the lampreys had for the
most part dispersed from the area occupied
on the previous day; a single lamprey was
observed beneath the stump that had shel¬
tered a spawning group. Two spawning
groups were located much further down¬
stream. Water temperature remained at
13°C. In 1992, approximately 13 lampreys
were observed over approximately 130 m of
stream. All were in the open, and most were
isolated individuals, but two pairs and one
group of three were found in flat water just
above riffles. Six females and three males
were collected. Water temperature was 12°C.
The mean total length of the 30 Ameri¬
can brook lampreys collected during this
study was 160 mm (range: 139-187 mm).
Mean body mass was 6.61 g (range: 3.83-
1 1.27 g). Use of student’s t-tests revealed sig¬
nificant differences between lampreys col¬
lected at Jambo Creek in 1989 and 1992 in
mean total length (t = 4.43, p < 0.001) and
mean body mass (t = 4.52, p < 0.001) (Table
1). Differences in mean total length and
body mass between lampreys from Taylor
Creek and Jambo Creek (1989 and 1992
data pooled) were not significant. For each
of the three samples from Taylor and Jambo
creeks, mean total length and body mass of
male lampreys were greater than correspond¬
ing values for females (Table 1). When data
for the three samples were pooled, the sexes
were significantly different in both mean to¬
tal length (t = 2.81, p < 0.01) and mean
body mass (t = 2.77, p = 0.01).
The simple linear regression of the natu¬
ral logarithm of body mass in grams (InW)
on the natural logarithm of total length in
mm (InL) was:
(1) lnW = -16.9 + 3.70 InL
(r2 = 0.921, n = 30). Analysis of covariance
failed to reveal significant differences be¬
tween regression lines calculated separately
for the two sexes or for the two collection
sites.
We measured the oral disc length of each
lamprey from Jambo Creek and calculated
relative disc length as the ratio of disc length
to total length (expressed as a percentage).
Total length was positively correlated with
disc length (r = 0.617, d.f. = 17, p < 0.01).
Analysis of covariance, with total length as
the covariate, revealed a significant difference
in disc length between the two sexes (F 6 =
12.53, p < 0.005). This reflected a difference
in mean relative disc length of males
(5.83%, S.E. = 0.21%) and females (5.12%,
S.E. = 0.13%). In addition to having rela¬
tively smaller oral discs, the Jambo Creek
females displayed swelling along the leading
edge of the second dorsal fin.
At Taylor Creek, American brook lam¬
preys were collected with spotfin shiners
( Cyprinella spiloptera ), bluntnose minnows
( Pimephales notatus ), white suckers ( Cato -
stomus commersoni ), banded darters (. Etheo -
stoma zonale) , johnny darters ( Etheostoma
nigrum ), and fantail darters (. Etheostoma
flabellare ). At Jambo Creek, a designated
trout stream, electrofishing on October 21,
1991, yielded the following species: brown
trout ( Salmo trutta ), creek chub ( Semotilus
atromaculatus) , common shiner ( Luxilus
cornutus ), white sucker, black bullhead
( Ictalurus melas ), smallmouth bass ( Microp -
terus dolomieui ), green sunfish ( Lepomis
cyanellus ), and mottled sculpin ( Cottus
bairdi). Only sculpins and young-of-the-year
suckers were observed in large numbers.
We found 12 cases in which American
brook lampreys and northern brook lam¬
preys occurred together in the same stream
Volume 81 (1993)
41
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1. Mean total length in millimeters and mean body mass in grams of American brook
lampreys ( Lampetra appendix) collected at Taylor and Jambo creeks, Wisconsin. Standard
errors are in parentheses. All measurements were of living, anesthetized animals.
Sample
(Fago 1983, 1984a, 1984b, 1983a). The
American brook lamprey was reported fur¬
ther upstream in 10 of 12 streams. This re¬
sult is significantly different from expected
under the null hypothesis that the two spe¬
cies are equally likely to be found further
upstream (binomial test, p = 0.039).
Discussion
Some generalizations about the temperature
at which American brook lampreys spawn
in the spring appear to be inaccurate. Becker
(1983) stated that spawning in Wisconsin
may begin at water temperatures of “about
17.2°C (63°F).” Robison and Buchanan
(1988) cited Becker (1983) but inexplicably
raised the temperature to “about 63°F
(18°C) . . . .” In contrast, we observed
spawning groups at temperatures of 12—
14°C, and spawning at one site was appar¬
ently completed by the time water tempera¬
ture had reached 16°C. Moreover, Cochran
(1984) reported the occurrence of several
spawning groups in Waukesha County at a
water temperature of 15.4°C. While it is true
that spawning by American brook lampreys
in other parts of North America has been re¬
ported at temperatures as high as 20.6°C,
most published accounts place the onset of
spawning well below 15°C (Table 2).
Lampreys typically are reported to spawn
in open, shallow, gravel-bottomed habitats.
Cochran and Gripentrog (1992), however,
reported that several species in the genus
Ichthyomyzon aggregate beneath cover ob¬
jects and sometimes spawn beneath cover.
Our observations at both Taylor and Jambo
42
TRANSACTIONS
COCHRAN et al . : The biology of the American brook lamprey in Wisconsin
creeks show that American brook lampreys
also occasionally aggregate and spawn be¬
neath cover objects. Young and Cole (1900)
reported that nests may be situated beneath
overhanging banks or logs.
In one of our samples of spawning-phase
adults, males outnumbered females by a sig¬
nificant margin. Care must be taken when
interpreting the literature on this topic. For
example, Schuldt et al. (1987) cited Seagle
and Nagel (1982) among authors who re¬
ported an excess of males, but Seagle and
Nagel (1982) stated that the sex ratio was
not statistically different from 1:1. Floff
(1988) and Scott and Crossman (1973) cited
Young and Cole (1900) as reporting that
males outnumbered females by a ratio of 3: 1 ,
but it was Dean and Sumner (1897) who
reported that figure. Hoff (1988) reported
that females outnumbered males 3:2, but
with a sample size of 7, that result is not sig¬
nificantly different from 1 : 1 (binomial test,
p = 0.453). Generally, however, adult male
American brook lampreys outnumber fe¬
males in collections made during or just
prior to the spawning season (Dean and
Sumner 1897; Young and Cole 1900; Kott
1971; Schuldt et al. 1987). Presumably, the
sex ratio varies over time, since males are re¬
ported to precede females to the spawning
site (Young and Cole 1900; Okkelberg
1921; but see Kott 1971).
Becker (1983) listed several traits for
which American brook lampreys in Wiscon¬
sin are sexually dimorphic. Breeding males
each have a long, threadlike urogenital pa¬
pilla and relatively high dorsal fins separated
by a sharp notch. Breeding females each
have a prominent anal fin fold and relatively
low dorsal fins separated by a broad notch.
In addition, we report here that males have
relatively larger oral discs than females and
that the leading edge of the second dorsal
fin in females may be swollen. A difference
in disc size has been previously noted for
American brook lampreys in Quebec (Kott
Volume 81 (1993)
43
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
1974) and Delaware (Rohde et al. 1976); the
larger discs of male lampreys may reflect
their relatively greater role in nest construc¬
tion (Beamish 1982). Swelling along the an¬
terior margin of the female’s second dorsal
fin was mentioned by Gage (1928) and pre¬
viously reported in several other species of
Lampetra (Pletcher 1963; Larsen 1980;
Hardisty 1986a, 1986b). Perhaps the swell¬
ing provides support for the body of the
male, which during the spawning act fits
into the notch between the first and second
dorsal fins of the female (Breder and Rosen
1966).
Adult male American brook lampreys
tended to be larger than females in our
samples (Table 1). In each of five previous
studies (Okkelberg 1921; Hubbs 1923; Saw¬
yer I960; Kott 1974; Rohde et al. 1976),
mean total lengths of males were slightly but
not significantly greater than those for fe¬
males. However, if all paired samples for the
two sexes were drawn randomly from popu¬
lations with identical means, the probabil¬
ity of obtaining such a sequence of results is
very low (p = 0.0136, binomial test). The
apparently real trend for male American
brook lampreys to be on average slightly
larger than females is reinforced by the re¬
sults of Schuldt et al. (1987), who obtained
a mean total length for males slightly but sig¬
nificantly greater than that for females. Such
a tendency may reflect a balance of oppos¬
ing selective factors. Malmqvist (1983) and
Beamish and Neville (1992), respectively,
found that fertilization and spawning suc¬
cess declined as size differences between male
and female lampreys increased; fertilization
was most successful when the female/male
length ratio was 1.05-1.14 (Malmqvist
1983). In contrast, Malmqvist (1983) and
Becker (1983) reported behavior interpreted
as fighting between male brook lampreys.
The former results would select against
substantial sexual divergence in size; the lat¬
ter phenomenon would presumably favor
larger males.
The tendency for American brook lam¬
preys to be found upstream from northern
brook lampreys reflects their preference for
cooler temperatures (Scott and Crossman
1973), which are often associated with
spring-fed headwaters. This pattern is not
inviolate, however. Classical patterns of lon¬
gitudinal zonation of stream fishes may be
disrupted in drainages where springs empty
cold water into the mid-reaches of streams
(Swaidner and Berra 1979). Morman (1979)
provided examples of inverted distributions
of American and northern brook lampreys
that apparently were related to reversed
stream temperature gradients. An example of
this phenomenon in Wisconsin may occur
in the Mukwonago River (Cochran 1984).
Northern brook lampreys were collected not
far downstream from Eagle Springs Lake,
the surface of which presumably warms
quickly in the spring, whereas American
brook lampreys were collected further down¬
stream, below where a trout stream and nu¬
merous springs enter the river.
Wisconsin’s native lampreys are not well
understood by the general public (Cochran
1984) and may suffer through association
with the sea lamprey ( Petromyzon marinus ),
an exotic parasitic species that has caused
great destruction of valuable fish in Lake
Michigan and the other Great Lakes (Smith
1971). The American brook lamprey is
nonparasitic and does not harm other fishes.
Nevertheless, landowners along Jambo
Creek informed us that they had encouraged
a Cub Scout pack to catch and kill what they
mistakenly thought were sea lampreys
spawning on their property. (Spawning-
phase sea lampreys from Lake Michigan do
ascend the East Twin River to within 4 km
of its confluence with Jambo Creek, but they
44
TRANSACTIONS
COCHRAN et al . : The biology of the American brook lamprey in Wisconsin
are blocked by the Mishicot spillway from
ascending closer.) We hope that further re¬
search on Wisconsin’s native lampreys will
help to dispel this sort of misunderstanding.
Acknowledgments
We thank John Lyons, Tim Pettinelli, Joe
Marks, Doug Hasten, and Jennifer Cochran
for assistance in the field. John Lyons and
two anonymous reviewers provided valuable
comments on earlier drafts of this manu¬
script, and Marge Dollar and Kristen Lucier
contributed to its production.
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Philip A. Cochran is an Associate Professor of Bi¬
ology at St. Norbert College. He is especially inter¬
ested in the ecology and geographic distribution of
lampreys and other fishes.
Martin E. Sneen graduated from St. Norbert Col¬
lege in 1990 with a B.S. degree in Environmental
Science. He currently works in Madison as an en¬
vironmental consultant.
Alan P. Gripentrog completed his B.S. degree in
Environmental Science at St. Norbert College in
1988. He now works in Washington, D. C. for the
Central Lntelligence Agency.
46
Elisabeth R. Deppe and Richard C. Lathrop
Recent changes in the aquatic macrophyte
community of Lake Mendota
Abstract The aquatic macrophyte community of Lake Mendota was surveyed in the sum¬
mers of 1989, 1990, and 1991 using a technique based on plant recovery on a
rake. For comparison, more limited surveys were conducted during 1990 and
1991 on the other three lakes in the Yahara River Chain — Monona, Waubesa,
and Kegonsa. For Lake Mendota, presence, relative frequency, and density of
twelve species were determined. In 1989 and 1990, the Lake Mendota mac¬
rophyte community was dominated by Ceratophyllum demersum L. followed
by Myriophyllum spicatum L., plants with biomasses especially heavy near
the water surface. While the relative frequencies of these and other species re¬
mained nearly constant, almost all species decreased in density from 1989 to
1990. In 1991, Ceratophyllum declined again, and Myriophyllum became
the most dominant. This same pattern was observed in Lakes Monona and
Waubesa between 1990 and 1991. These density decreases were probably caused
by especially poor spring and/or summer water clarity in 1990 in all three lakes.
In Lake Mendota, this poor clarity resulted from an unusually long and dense
blue-green algal bloom during May and June. Areas of high plant density along
the west and southwest shorelines, including University Bay, showed the larg¬
est density decreases. This was probably due to an accumulation of algae in
these regions, blown in by northeast winds during the crucial spring growth
period. While Lake Mendota has been dominated by either Myriophyllum
spicatum or Ceratophyllum demersum since the 1960s, our survey results
indicate that plant densities can vary greatly in the short-term due to stochas¬
tic events.
The submersed aquatic macrophytes of Lake Mendota, a
calcareous 3,985-ha lake near Madison, Wisconsin, have
been of interest to ecologists and lake managers for decades.
Lake Mendota has undergone steady eutrophication since the
TRANSACTIONS Volume 81 (1993)
47
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
mid-1800s, when its watershed was first de¬
veloped (Lathrop 1992). Prior to the 1960s,
the lake was dominated by wild celery
( Vallisneria americana Michx.) and native
pondweeds ( Potamogeton spp.) (Lathrop
1989; Nichols et al. 1992). Since then, the
plant community became less diverse, being
dominated by Eurasian water milfoil
{Myriophyllum spicatum). Myriophyllum was
especially dense from the mid-1960s to the
mid-1970s, when densities declined in
Mendota and other Yahara River lakes
(Monona, Waubesa, and Kegonsa) and Lake
Wingra (Carpenter 1980; Lathrop 1989;
Nichols et al. 1992).
Local lake managers continuously devote
energy and resources to managing overabun¬
dant littoral biomasses of mainly Myrio¬
phyllum , particularly in Lakes Mendota,
Monona, and Waubesa (Lathrop 1989).
While providing important habitat for many
fish species, this plant can become a nuisance
to lakeside residents, boaters, and even an¬
glers when growth is unchecked. In Lake
Mendota, improved water clarity during
1986-88 caused ecologists and lake manag¬
ers additional concern that Myriophyllum
would increase in density and spread into
deeper water.
To address these concerns, and because
a systematic, detailed macrophyte survey had
not been conducted on Lake Mendota since
1920 (Rickett 1922), the Wisconsin Depart¬
ment of Natural Resources (WDNR) under¬
took full shoreline surveys in the summers
of 1989—91 to document the presence, rela¬
tive density, and maximum rooting depth.
The other three lakes in the Yahara River
chain were also surveyed in the summers of
1990 and 1991 for comparison. The survey
information should aid managers responsible
for controlling overabundant macrophytes
while indicating areas of native species de¬
serving protection.
Methods
Field Survey
Lake Mendota’s macrophytes were surveyed
during the last two weeks of July and the
first week of August in 1989—9 1 . Plants were
sampled along 47 transects positioned per¬
pendicular to the shoreline at approximately
750-m intervals around the lake (Fig. 1).
Transects were 300 m apart in University
Bay, where we wanted additional data for
historical comparison. Lakes Monona and
Waubesa were surveyed in late June of 1990
and 1991 and Lake Kegonsa in early July of
1990 and 1991. Macrophytes were sampled
along 1 3 evenly spaced transects in Monona
(plus one in Monona Bay), 10 in Waubesa,
and 8 in Kegonsa.
The surveys were conducted from a boat,
which was moved between sampling stations
designated at 0.5-m water depth intervals
(0.3 m, 1.0 m, 1.3 m, etc.) on each transect
until no vegetation occurred. Depths were
determined using a measured pole in waters
<3.0 m and a Lowrance FISH-LO-K-TOR
depth-finder in waters 3. 5-5. 5 m. Each sam¬
pling station was subdivided into four quad¬
rants, located off the front left, front right,
rear left, and rear right of the boat. Each
quadrant was then sampled by a single cast
with a double-headed, weighted garden rake
with a head width of 35 cm and 14 teeth,
each 5 cm long. The rake was thrown into
the water and dragged approximately 2 m
across the bottom by means of an attached
line. After the rake was pulled off the bot¬
tom it was flipped 180 degrees to ensure that
plants snagged from the bottom would re¬
main on the rake. This sampling technique
for dividing stations into quadrants and col¬
lecting plants by rake casts followed Jessen
and Lound (1962).
Plants collected on rake teeth were iden-
48
TRANSACTIONS
DEPPE and LATHROP: Aquatic macrophyte community of Lake Mendota
Lake Mendota
Fig. 1. Hydrographic map of Lake Mendota (with contours in meters) showing loca¬
tions of the 47 study area transects.
tifled following Fassett (1957) and Voss
(1972). Whereas Jessen and Lound rated
species on rake teeth only as present or ab¬
sent, we developed a measure of estimating
density as well. For each rake throw, a spe¬
cies was assigned a density rating from 0—5
based on the coverage of the upper rake head
(both number of teeth and length of teeth
covered). For example, species covering 1—
20% of the rake head were given a rating of
1, species covering 21^40% were rated 2,
etc. Ratings resulting from the four indi¬
vidual throws at each station were averaged
to determine a density rating (DR) for each
species. Further information on the rake sur¬
vey method was provided in Deppe and
Lathrop (1992). The simpler Jessen and
Lound method of presence/absence on each
rake was used for the 1990 surveys on
Monona, Waubesa, and Kegonsa.
Statistical Analysis
Relative frequencies, which describe each
species as constituting a certain percent of
the whole macrophyte community, were
computed for each lake. In addition, sepa¬
rate relative frequencies were computed for
University Bay (transects 38—42) of Lake
Mendota for the purpose of comparison
with previous surveys only conducted in that
area. Relative frequency for each species was
Volume 81 (1993)
49
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
calculated as the total number of rake casts
on which a species appeared, divided by the
total number of such encounters for all spe¬
cies. These calculations were based on data
for all stations. While frequency of occur¬
rence more accurately reflects a species dis¬
tribution in a lake, we present only relative
frequency information in this paper. Because
different plant sampling methodologies were
used in past surveys, only relative frequency
data are directly comparable. However, it
should be noted that relative frequency data
were found to parallel frequency of occur¬
rence data for Lake Mendota’s macrophytes.
For Lake Mendota, density of plants is
presented in two ways. Mean density ratings
(MDRs) were calculated for the abundant
and common species at each depth across all
transects. In order to characterize the mac¬
rophyte community with respect to shore¬
line areas, density ratings of each species
were summed for each transect and called
their additive density ratings (ADRs).
Results
Macrophyte Community Composition
A total of ten submersed and two floating¬
leaved species were found in Lake Mendota
during 1989—91 (Table 1). Ceratophyllum
demersum (coontail) and Myriophyllum
spicatum were the two most dominant spe¬
cies, each comprising 26-43% of the mac¬
rophyte community in 1989-91 (Table 1).
Four other species — Potamogeton pectinatus
L. (sago pondweed), Vallisneria americana
(wild celery), Heteranthera dubia (Jacq.)
MacM. (water stargrass), and Elodea cana¬
densis Michx. (American elodea) — were
common, each comprising 3—9% of Men¬
dota macrophytes in 1989—91. The other six
species were infrequently encountered (rela¬
tive frequencies of each generally <1% in
1989—91). Composition of the macrophyte
community in University Bay of Lake
Mendota was almost the same as for the
whole lake, dominated by Myriophyllum and
Ceratophyllum , and containing seven of the
other ten less abundant species found in
Lake Mendota (Tables 1 and 2).
The macrophyte community of Lake
Monona was very similar to Lake Men¬
dota’s, only lacking the two floating-leaved
species and one uncommon Potamogeton
(Table 1). Lakes Waubesa and Kegonsa were
much less diverse, harboring only three spe¬
cies besides Ceratophyllum and Myrio¬
phyllum.
Abundant Species
Our Lake Mendota surveys document a
transition from a Ceratophyllum-6.omm2a.QA
macrophyte community to one dominated
by Myriophyllum. Ceratophyllum was the
most abundant species in 1989 and 1990,
with higher relative frequencies and mean
density ratings (MDRs) than Myriophyllum
(Table 1 and Fig. 2). Both species experi¬
enced substantial decreases in density from
1989 to 1990 (Fig. 2). Densities of Cerato¬
phyllum declined again in 1991, while den¬
sities of Myriophyllum remained nearly con¬
stant from 1990 to 1991, making Myrio¬
phyllum the most abundant species in 1991.
For both Ceratophyllum and Myrio¬
phyllum, areas of greatest abundance in 1989
exhibited the most dramatic decreases in
density. In 1989, Ceratophyllum grew most
densely along Lake Mendota’s northwest
and west shorelines and in University Bay
(Fig. 3). Between 1989 and 1991, Cerato¬
phyllum densities declined along the north¬
west shoreline and in University Bay by 68%
and 93%, respectively. Relative frequencies
in University Bay also portrayed a dramatic
decrease in Ceratophyllum density from 1989
50
TRANSACTIONS
DEPPE and LATHROP: Aquatic macrophyte community of Lake Mendota
Table 1. Relative frequencies of macrophytes in Lake Mendota in 1989-91, and in Lakes
Monona, Waubesa, and Kegonsa in 1990-91.
to 1991 (Table 2). Along the south and
southeast shorelines, areas of moderate
Ceratophyllum density in 1989, densities
dropped by 80% from 1989 to 1991.
Myriophyllum , while fairly evenly distributed
around the lake in 1989, was slightly more
abundant in University Bay and along the
west shoreline (Fig. 3). After decreases in
nearly all regions from 1989 to 1990, sharp¬
est in University Bay and along the west
Volume 81 (1993)
51
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 2. Percent relative frequency of macrophyte species in University Bay in selected
years since 1966.*
'Data sources: 1966 (Lind and Cottam 1969); 1978-79 (raw data used by Andrews
1980); 1980 (Vander Zouwen 1982); 1984 (R. Lathrop, Wis. DNR, unpubl. data);
1989-91 (this study). All surveys were conducted during July-August.
52
TRANSACTIONS
DEPPE and LATHROP: Aquatic macrophyte community of Lake Mendota
Fig. 2. Mean density ratings with respect to depth for abundant and common species
in Lake Mendota, 1989-91. (Abundant species in top panels have scale 0-3.0; com¬
mon species in middle and bottom panels have scale 0-0.6; 1989 - solid line, 1990 -
dashed line, 1991 -dotted line.)
Volume 81 (1993)
53
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 3. Additive density ratings of Ceratophyllum demersum and Myriophyllum spicatum
in regions of Lake Mendota and the percent change in these ratings from 1989-91.
(University Bay = transects 38-41. See Figure 1 for other transect locations.)
shoreline, Myriophyllum densities almost re¬
gained their 1989 levels in 1991, unlike
Ceratophyllum. During 1989—91, Cerato¬
phyllum grew most densely between 2.0 m
and 3.0 m of water depth, and Myrophyllum
between 2.3 m and 3.0 m (Fig. 2).
In Lakes Monona and Waubesa, Cerato¬
phyllum dominated in 1990 but dropped
dramatically in relative frequency in 1991,
while Myriophyllum frequencies increased
(Table 1). Relative frequency for Lake
Kegonsa, a lake with much sparser macro¬
phytes, was almost totally dominated by
Myriophyllum in both 1990 and 1991.
Common Species
The four common submersed species — P.
pectinatus , Vallisneria , Heteranthera , and Elo-
dea-were. found less frequently and in
lower densities than Ceratophyllum and
Myriophyllum at all water depths, but grew
most abundantly in shallower depths (< 2.0
m) (Table 1 and Fig. 2). In 1989, Vallis¬
neria , Heteranthera , and Elodea had peak
MDRs at depths of 0. 3-1.0 m, 1.0-1. 5 m,
and 1.5— 2.0 m, respectively. Small differ¬
ences in MDRs occurred during the three
years for these four species, but no consis¬
tent trend was evident. They were generally
found at transects scattered all over the lake,
with some prevalence on certain shorelines.
Distribution of P. pectinatus , although fairly
uniform in 1989, seemed less random than
the others. It was entirely absent from Uni¬
versity Bay in 1990; however, its greatest
densities occurred there in 1991, reflected
by its relative frequency of 17.4% (Table 2).
54
TRANSACTIONS
DEPPE and LATHROP: Aquatic macrophyte community of Lake Mendota
Uncommon Species
In the three survey years, six species were
found very infrequently and at very low den¬
sities in Lake Mendota: Potamogeton crispus
L., P. richardsonii (A. Benn.) Rydb., P.
zosteriformis Fern., P. foliosis Raf., Nelumbo
lutea (Willd.) Pers., and Nymphaea tuberosa
Paine (Table 1). In 1989 and 1990, P.
crispus was found at low densities on
transects off the eastern and northern shore¬
lines, while in 1991 it was found almost ex¬
clusively along the west and south shorelines.
In all three years, P. richardsonii was found
in moderate abundance at transect 45 and
more sparsely at a few other transects dur¬
ing 1989-91. For all three years, the two
floating-leaved lily species Nelumbo lutea and
Nymphaea tuberosa grew densely at 1.0 m
in University Bay at transect 39, and else¬
where in the bay at varying densities from
1989-91. These species were also found
along the east shoreline at transect 47 in
1989. P. foliosis was found at scattered sites
in 1 990-9 1 , and P. zosteriformis was found
at only one station in 1989.
Depth Limit of Growth
The depth limit of plant growth in Lake
Mendota was somewhat variable between
transects but generally occurred between 3.0
and 4.0 m for 1989-91 (Table 3). Depth
limits shifted to slightly shallower ranges in
1990. but returned to near-1989 levels in
1991. Depth limit decreased by 0.5 m from
1989 to 1990 at all transects in University
Bay and along the northwest shoreline where
plant growth was densest.
Depth limit of plant growth in Lake
Monona generally occurred at 3.5 m in
1990, but at 2.5 m in 1991. Similarly, in
Lake Waubesa these figures went from 3.0
m in 1990 to 2.0— 3.0 m in 1991. In Lake
Kegonsa, depth limits increased from 2.0—
2.5 m in 1990 to 3.0 m in 1991.
Because most plant growth occurred at
3-4 m in Lake Mendota, it is noteworthy
that 10% of stations 3 m in 1989 were de¬
void of plants, while 20% had no vegetation
in 1990 and 1991. In 1990, 7% and 1% of
stations 3 m were without vegetation in
Monona and Waubesa, respectively, but this
increased to 30% and 48% for the two lakes
in 1991. In Lake Kegonsa, 58% of stations
3 m were without macrophytes in 1990,
but only 21% in 1991.
Discussion
Depth Distribution
In our 1989-91 surveys in Lake Mendota,
macrophytes were found almost entirely at
water depths between 0.5 and 3.5 m, while
certain depths favored growth of particular
species. Ceratophyllum and Myriophyllum spi-
catum , tall-growing plants with biomasses
heaviest near the water surface, grew most
densely between 2.0 and 3.0 m. The com¬
mon species, tending not to grow as tall,
were found largely between 0.5 and 2.0 m,
where they can receive adequate light. Their
infrequent occurrence in water depths >2.0
m suggests that they may be shaded by al¬
gal blooms and dense growths of Cerato¬
phyllum and Myriophyllum. Lack of macro¬
phyte growth at the 0.5 m contour is prob¬
ably due to one or more of the following rea¬
sons: rocky substrate, more pronounced
wave action, ice shifting in winter, and the
controlled lowering of the lake level over the
winter months.
Macrophyte Community of University Bay
Since the 1960s
Myriophyllum spicatum dominated the plant
community from its introduction in the
Volume 81 (1993)
55
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 3. Depth limits of macrophyte growth in Lake Mendota in 1989-91.
No. transects where plants ceased
Sampling -
Station (m) 1989 1990 1991
Transects 12-15 at the Yahara River inlet were excluded in all
years because of shallow maximum depths; transect 6 was also
excluded in 1989 because no plants were found there due to rocky substrate
early 1960s until the mid-1970s, while
Potamogeton spp. and Vallisneria declined
(Table 2). M. sibiricum Komarov (native
water milfoil, formerly called M. exalbescens
Fern.), apparently disappeared (Nichols
1975) and has not been recorded in surveys
through 1991. After the mid-1970s, relative
abundances of M. spicatum declined while
Ceratophyllum began to increase, with the
exception of 1980. This decline in M.
spicatum abundance following approxi¬
mately a 10-year period of domination has
been observed in other lakes including Lake
Wingra (Carpenter 1980), although the rea¬
sons for this are unclear. Transient increases
in narrow-leaved pondweeds such as P.
folio sis, P. filiformis and P. pectinatus oc¬
curred between 1978 and 1980 in Univer¬
sity Bay. Flowever, by the 1980s, several
pondweeds had either dropped dramatically
in relative frequency or totally disappeared
from the bay, indicating steadily declining
diversity.
Our surveys indicate the general continu¬
ation of Myriophylluml Ceratophyllum domi¬
nance, but the trend of increasing Cerato¬
phyllum relative frequency observed through
the 1980s was reversed in 1991. While both
species experienced reductions in area of
coverage, mean density, and maximum
depth distribution from 1989 to 1990,
Cerato-phyllum densities dropped again in
1991 (Figs. 2 and 3). Myriophyllum densi¬
ties, however, remained nearly constant
from 1990 to 1991. This reversal caused
Myriophyllum to once again dominate, but
at lower densities than were noted in the late
1960s or early 1970s.
56
TRANSACTIONS
DEPPE and LATHROP: Aquatic macrophyte community of Lake Mendota
Effect of Water Clarity
Water clarity can critically influence the den¬
sities, species composition, and maximum
rooting depth of submersed aquatic macro¬
phyte communities (Canfield et al. 1985;
Chambers and Kalff 1985). In our surveys,
it is most likely that the sudden shift to a
Myriophyllum- dominated community may
have been caused, at least in part, by a dif¬
ference in the responses of Ceratophyllum
and Myriophyllum to severely reduced light
conditions in the spring of 1990. This situ¬
ation was particularly pronounced in Uni¬
versity Bay. During the spring of that year,
an atypical heavy bloom of blue-green algae
occurred in Lake Mendota that resulted in
unusually poor water clarity when compared
to other spring periods (Lathrop 1992,
WDNR unpubl. data). Because prevailing
winds were predominantly from the north¬
east during May, rather than the more typi¬
cal southwestern winds for this season, the
buoyant blue-green algae accumulated in the
bay. Extremely poor water clarity resulted,
at a time of year when young plants require
adequate light to initiate growth from the
sediments. Water clarity was poorer than
normal for much of the remaining spring
and summer months of 1990. By August of
that year, the shore side of the sand bar in
University Bay was almost completely de¬
void of submersed vegetation, and the plant
densities of the entire bay were severely re¬
duced. In 1991, the bay still harbored only
sparse macrophyte growth, although P.
pectinatus was much more abundant than
usual (Table 2).
Poor spring water clarity in 1979 (Lath¬
rop 1 992) also affected the macrophyte com¬
munity in University Bay. Andrews (1980)
noted a decline in macrophyte abundance
(particularly Myriophyllum) from 1978 to
1979. In 1980, Vander Zouwen (1982)
found Ceratophyllum to be quite sparse when
compared to earlier and later surveys. Inter¬
estingly, P. pectinatus also exhibited in¬
creased abundances one year after poor wa¬
ter clarity in both 1980 and 1991 (Table 2).
Poor spring and summer water clarity in
1990 also had an impact on the macrophyte
communities of Lakes Monona and Wau-
besa. Both lakes exhibited huge drops in the
relative frequency of Ceratophyllum from
1990 to 1991 as had occurred in Lake
Mendota. Water clarity was typically poor
for both 1990 and 1991 in Lake Kegonsa,
where plants were sparse. Relative frequen¬
cies were similar in both years with Myrio¬
phyllum being the dominant plant.
While densities of the macrophyte com¬
munity in Lake Mendota as a whole drop¬
ped as a result of poor water clarity in both
1979 and 1990, it is unclear why only
Ceratophyllum declined a second year in a
row (1980 and 1991). Future summer sur¬
veys will help document the ongoing
changes that have occurred in Lake Men-
dota’s macrophytes, particularly since the
invasion of M. spicatum. It will be interest¬
ing whether the next few years will show a
resurgence of Ceratophyllum and/or Myrio¬
phyllum or an increase in diversity.
Acknowledgments
We are indebted to the many people who
assisted on this project. In particular, D. E.
Bergstrom, Jr., L. M. Hartman, and W. T.
Seybold assisted in field sampling, and P. W.
Rasmussen guided statistical analysis of the
data. In addition, C. R. Molter gave us ad¬
vice on field sampling techniques, and Dr.
J. D. Andrews graciously provided unpub¬
lished raw data used in his 1980 report. S.
B. Nehls made helpful suggestions on out¬
lines and drafts of this report, and J. S.
Winkelman provided helpful comments on
Volume 81 (1993)
57
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
the final draft. Finally, special thanks are
given to Dr. S. R. Carpenter and Dr. S. A.
Nichols for their critical review of an earlier
draft.
Works Cited
Andrews, J. H. 1980. Plant pathogens as agents
for biological and integrated control of
aquatic plants. Univ. Wis. -Madison, Water
Resour. Cent. Tech. Rep. No. 80-01. 36 pp.
Canfield, D. E., K. A. Langeland, S. B. Linda,
and W. T. Haller. 1983. Relations between
water transparency and maximum depth of
macrophyte colonization in lakes. J. Aquat.
Plant Manage. 23:23-28.
Carpenter, S. R. 1980. The decline of Myrio -
phyllum spicatum in a eutrophic Wisconsin
lake. Can.]. Bot. 58:527-35.
Chambers, P. A., and J. Kalff. 1985. Depth dis¬
tribution and biomass of submersed aquatic
macrophyte communities in relation to
Secchi depth. Can. ]. Fish. Aquat. Sci.
42:701-09.
Deppe, E. R., and R. C. Lathrop. 1992. A com¬
parison of two rake sampling techniques for
sampling aquatic macrophytes. Wis. Dep.
Nat. Resour. Res. Manage. Find. No. 32.
4pp.
Fassett, N. C. 1957. A manual of aquatic plants.
rev. ed. Madison: Univ. Wis. Press, 405 pp.
Jessen, R., and R. Lound. 1962. An evaluation
of a survey technique for submersed aquatic
plants. Minn. Dep. Conserv. Game Invest.
Rep. No. 6. 10 pp.
Lathrop, R. C. 1989. The abundance of aquatic
macrophytes in the Yahara lakes. Wis. Dep.
Nat. Resour. Res. Manage. Find. No. 22.
4pp.
Lathrop, R. C. 1992. Nutrient loadings, lake
nutrients, and water clarity, Chap. 6. In Food
web research and its application to lake man¬
agement: a case study of Lake Mendota, Wis¬
consin , ed. J. F. Kitchell. New York: Springer-
Verlag.
Lind, C. T., and G. Cottam. 1969. The sub¬
merged aquatics of University Bay: A study
in eutrophication. Am. Midi. Nat. 831:353—
69.
Nichols, S. A. 1975. Identification and manage¬
ment of Eurasian watermilfoil in Wisconsin.
Trans. Wis. Acad. Sci., Arts and Lett. 63:1 16—
28.
Nichols, S. A., R. C. Lathrop, and S. R. Car¬
penter. 1992. Long term vegetation trends:
a history, Chap. 9. In Food web research and
its application to lake management: A case study
of Lake Mendota, Wisconsin , ed. J. F. Kitchell.
New York: Springer-Verlag.
Rickett, W. H. 1922. A quantitative study of the
larger aquatic plants of Lake Mendota. Trans.
Wis. Acad. Sci., Arts and Lett. 20:501-27.
Vander Zouwen, W. J. 1982. Vegetational
change in University Bay from 1966 to 1980.
Trans. Wis. Acad. Sci., Arts and Lett. 70:42—
51.
Voss, E. G. 1972. Michigan flora: Part I. Gym-
nosperms and monocots. Cranbrook Inst. Sci.,
Bloomfield Hills, Mich. Bull. No. 55. 488
PP-
Elisabeth Deppe was a limited term biologist with
the Wisconsin DNR Bureau of Research, where she
conducted macrophyte surveys of the four Yahara
lakes in 1989, 1990, and 1991.
Richard Lathrop is a limnologist with the DNR
Bureau of Research. He has conducted research on
the Yahara lakes since 1976.
58
James O. Evrard
Were wild turkeys found historically
in northwest Wisconsin ?
The restoration of the wild turkey (Meleagris gallopavo) in
North America is one of wildlife management’s great suc¬
cess stories. Once near extinction, the wild turkey now num¬
bers approximately four million birds. The wild turkey’s res¬
toration in much of its historic range and successful introduc¬
tion in other areas were accomplished by transplanting wild
birds into suitable habitat.
With increasing interest in new environmental issues such
as biodiversity and restoration biology, the technique of releas¬
ing wildlife species outside their historical range is being in¬
creasingly questioned. It is, therefore, important to accurately
delineate the original distribution of wildlife species. This dis¬
tribution in North America is generally accepted as that exist¬
ing at the time of European exploration and settlement.
In Wisconsin, Schorger (1942) delineated the northern limit
of the historical range of the eastern wild turkey (M. g. silvestris)
as a line from Prairie du Chien to Green Bay (Fig. 1). Recent
turkey range distributions (Hewitt 1967; Kallman 1987) con¬
tinue to be based upon his work. In reaching his conclusions,
Schorger disregarded or discredited two observations of wild
turkeys near Lake Pepin, a natural widening of the Mississippi
River about 100 miles north of Prairie du Chien and at the
same latitude as Green Bay.
Schorger disregarded Father Hennepin’s report that his party
killed seven or eight large turkeys ( Cog dlnde) near Lake Pepin
in 1680. Hennepin also mentioned that Indians reported bus¬
tards or wild turkeys ( Outardes ou [or] Cogs dlnde) in that area.
In this instance, Outarde or bustard was synonymous with the
turkey.
TRANSACTIONS
Volume 81 (1993)
59
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 1 . Original range of the wild turkey delineated by Schorger (1942) and the Tension
Zone of Curtis (1959).
Schorger discredited Jonathan Carver’s
1776 observations of turkeys at Lake Pepin
because he concluded they were pilfered
from Hennepin’s writings. However, Parker
(1976) questioned the validity of the plagia¬
rism charges. Carver’s turkey observations
may, indeed, be more valid than Schorger
believed.
Schorger also attempted to make the case
that Outarde was the French word for Cana¬
da geese (Branta canadensis) by using two
observations east of Wisconsin, one from
Lake Champlain in 1683 and the other from
the mouth of the Cumberland River in
1793. However, he appeared to contradict
himself by stating:
Early explorers naturally would call the new
American animals by the names of creatures
in Europe that they resembled most closely.
The Outarde or Bustard is a large stocky bird.
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EVRARD: Were wild turkeys found historically in northwest Wisconsin?
The spreading of the tail and other phases of
courtship demeanor give it a decided resem¬
blance to the Turkey. Only speculation can
be offered for the synonymy of Outarde and
Canada Geese.
Two records not available to Schorger in
1 942 indicate the wild turkey may have ex¬
isted nearly 100 miles north of Lake Pepin
(Fig. 1).
An Outarde was provided to the fur
trader, John Sayer, by his Ojibway hunters
at the North West Company post on the
Snake River, near Pine City, Minnesota, on
September 18, 1804 (Gates 1965). The fur
trading post was located about 12 miles west
of the St. Croix River in Pine County. The
Outarde could not have been confused with
geese because Sayer’s journal also mentions
that he received geese from his hunters six
days later and on five subsequent occasions.
The second record consisted of a wild tur¬
key bone found in a refuse pit during an ar¬
cheological excavation of a combined North
West Company and XY Company fur trad¬
ing post on the Yellow River in Burnett
County, Wisconsin (Ewen 1983). The post
was occupied during the winters of 1802-
03 and 1804-05 and was located about 3
miles from the St. Croix River and about 28
miles east of the contemporary Snake River
post. Journal entries from both forts (Gates
1965; Thwaites 1911) indicated that hunt¬
ers took their game within 20-30 miles of
the posts, suggesting turkeys were present in
Pine and Burnett counties at that time.
Flow could the wild turkey have existed
nearly 200 miles north of the original range
outlined by Schorger?
A review of presettlement vegetation and
the location of the “tension zone” in Wis¬
consin, a transitional boundary between
northern and southern plant and animal
communities described by Curtis (1959),
supports the northern turkey records. Oak
forests and prairies — both turkey habitat —
were found south and west of the tension
zone (Fig. 1). Oak forests and remnant prai¬
ries still cloak the Mississippi River bluffs
from the Illinois border to the Minnesota
border and along the St. Croix River north
into Burnett and Pine counties.
If suitable turkey habitat existed in the
Mississippi and tributary river valleys north
of Prairie du Chien, why were there so few
historical turkey records for the area?
A potential answer to this question was
given by Schorger himself. He stated that the
northern range limit of the wild turkey, like
the bobwhite quail ( Colinus virginianus) and
prairie chicken (Tympanuchos pinnatus ), oc¬
curred in Wisconsin. He speculated that
prior to European settlement, these species
existed as members of Wisconsin’s fauna
only by periodic replenishment from Illinois.
The northern limit of their range varied in
response to the severity of winter weather,
moving northward during a succession of
mild winters and retreating southward fol¬
lowing severe winters.
Schorger reasoned that the scarcity of
wild turkey records in Wisconsin for the last
half of the nineteenth century was primarily
due to the severe winter of 1842—43 when
the species was nearly extirpated in the state.
Southern and western Wisconsin were not
settled by Europeans until the 1850s so there
were very few turkeys remaining for the set¬
tlers to see. Massive habitat destruction and
unregulated hunting that accompanied set¬
tlement sealed the fate of the few remain¬
ing turkeys.
Kumlien and Hollister in 1903 stated:
The Wild Turkey is to-day so rare in Wis¬
consin that it is safe to say that it is extinct.
Authentic references are meager and fragmen¬
tary. Dr. Hoy and others say it was abundant
in southern Wisconsin prior to 1840. Several
references, of which Hoy’s is one of the most
Volume 81 (1993)
61
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
reliable, state that the winter of 1842 was
practically fatal to them.
Shorger quoted Dr. Hoy concerning the
near extinction of the turkey in Wisconsin:
I am told by Dr. E. B. Wolcott that turkeys
were abundant in Wisconsin prior to the hard
winter of 1842—43, when snow was yet two
feet deep in March, with a firm crust, so that
the turkeys could not get to the ground; they
hence became so poor and weak that they
could not fly and so were an easy prey for the
wolves, wildcats, foxes and minks. The Doc¬
tor further stated that he saw but one single
turkey the next winter, and none since.
Shorger stated that the winter of 1842
was known as the “hard winter” for decades
afterward.
There were other hard winters that affec¬
ted wild turkeys, both historically and more
recently. Schorger quoted from the diary of
Marquette that the winter of 1674 was one
of intense cold and deep snow at the present
site of Chicago, Illinois. On December 12,
Marquette wrote:
We contented ourselves with killing three or
four turkeys out of many that came around
our cabin because they were almost dying of
hunger.
Schorger concluded that since these con¬
ditions existed at Chicago, it was probable
that most wild turkeys in Wisconsin per¬
ished that winter.
A more recent example is given by
Robbins (1991). Wild turkeys released in the
Meadow Valley area of central Wisconsin in
the mid-1950s increased to 2,500 birds a
decade later. However, the winter of 1 968-
69 was one of deep snow which took a heavy
toll of turkeys. By 1973, the estimated popu¬
lation was only 70 birds.
With this in mind, delineating the his¬
torical range of the wild turkey in Wiscon¬
sin would depend upon the time period ex¬
amined. Few turkey records for the north¬
ern edge of its range would be found for
years following an exceptionally severe win¬
ter. Conversely, one could expect to find
more wild turkey records during an extended
period of mild winters.
European settlement prevented the natu¬
ral ebb and flow of the northern limit of the
occupied wild turkey range and eliminated
the species in Wisconsin. But the conversion
of much of the state’s turkey habitat to ag¬
ricultural uses set the stage for the eventual
reestablishment of the wild turkey. The
availability of waste corn in harvested fields
and spread manure on today’s farms provide
winter food that was unavailable to turkeys
before European settlement. Wild turkeys
can now survive severe winters that would
have been impossible in the past.
The wild turkey has returned to the area
in east-central Minnesota and northwest
Wisconsin where it was found in 1804.
Birds released about ten miles south of Pine
City, Minnesota, in the 1980s reproduced
and spread northward and eastward, cross¬
ing the St. Croix River into Burnett and ad¬
joining Polk counties, Wisconsin. The tur¬
key population in this area of Wisconsin in
the spring of 1992 was estimated to be 100—
200 birds (Michael Johnson, Wisconsin De¬
partment of Natural Resources (WDNR),
pers. comm. 1992). These birds survived the
winter of 1991-92 despite snow depths of
nearly 20 inches covering the ground more
than five months.
It appears that the wild turkey is now a
resident of northwest Wisconsin after an
absence of nearly 200 years. The present
population was bolstered by a release of
wild-trapped turkeys from southern Wiscon¬
sin in early 1992. The WDNR plans an ad¬
ditional release of turkeys in the winter of
1992-93. Although severe winters may
cause large population fluctuations, food
62
TRANSACTIONS
EVRARD: Were wild turkeys found historically in northwest Wisconsin?
supplied by current agricultural practices
should ensure that scattered flocks will per¬
sist in this northern landscape.
Acknowledgments
I thank M. Johnson of the WDNR for the
discussion that generated the idea for this
paper. I also thank E. Oerichbauer of the
Burnett County Historical Society for guid¬
ance to historical records and J. Hoefler, M.
Johnson, J. Kubisiak, and W. Vander
Zouwen of the WNDNR and an anony¬
mous reviewer for critical review of the
manuscript. Partial funding for this study
was provided by the Federal Aid to Wild¬
life Restoration under Pittman-Robertson
Wis. Proj. W-141-R.
Works Cited
Curtis, J. T. 1959. The vegetation of Wiscon¬
sin. Madison: University of Wisconsin Press.
Ewen, C. R. 1983. Fur trade zooarcheology: A
faunal interpretation of two wintering posts
in northwestern Wisconsin. Master’s Thesis.
Tallahassee: Florida State University.
Gates, C. M. 1965. Five fur trader of the north¬
west. St. Paul: Minnesota Historical Society
Press.
Hewitt, O. H., ed. 1967. The wild turkey and
its management. Washington, D.C.: The
Wildlife Society.
Kallman, H. 1987. Restoring America’s wildlife—
1937-87. Washington, D.C.: U.S. Fish and
Wildlife Service.
Kumlien, L. and N. Hollister. 1903. The birds
of Wisconsin. Bull. Wis. Hist. Soc. 3(1-3): 1 —
143.
Parker, J., ed. 1976. The journals of Jonathan
Carver and related documents 1766-70. St.
Paul: Minnesota Historical Society Press.
Robbins, S. D., Jr. 1991. Wisconsin birdlife —
population and distribution — past and pre¬
sent. Madison: University of Wisconsin Press.
Schorger, A. W. 1942. The Wild Turkey in early
Wisconsin. Wilson Bull. 54(3):173-82.
Thwaites, R. G., ed. 1911. A Wisconsin fur-
trader’s journal, 1803-04. Wis. Hist. Coll.
20:396-471.
James O. Evrard is a wildlife research biologist for
the Wisconsin Department of Natural Resources at
Grantsburg.
Volume 81 (1993)
63
Marguerite H. Helmers
Creating the California Alps
John Muir, the nineteenth-century naturalist, was a prolific
and elegant writer whose articles and books numbered in
the hundreds through several revisions and editions. His na¬
ture writing combines the finest elements of scenic description
with exciting adventure stories, but several of his short essays
demonstrate much more: that certain styles of writing, like
styles of clothing, painting, and music, are historically deter¬
mined. Muir’s writing obeys the conventions of the typical re¬
sponse to nature in the eighteenth and nineteenth century —
appreciation of the sublime and the picturesque. This essay will
trace Muir’s uses of literary formulas of picturesque and sub¬
lime representation through two articles. These articles, “Snow-
Storm on Mount Shasta” (1877) and “In the Heart of the Cali¬
fornia Alps” (1880), appeared during what Muir’s literary bi¬
ographer Herbert F. Smith refers to as Muir’s “most fertile pe¬
riod,” a time of discovery, experimentation, environmental ac¬
tivism, and written expression. They were published initially
in Harper’s and Scribner’s magazines, although Muir later re¬
vised both for inclusion in his book The Mountains of Califor¬
nia (1894).
These two short essays reveal also that Muir’s writing is pro¬
foundly subversive. His efforts to incorporate description that
would meet the demands of his leisure-class readers carries with
it a touch of contempt at the passive mode of observation and
conformity to convention that fostered those readers’ demands.
While he paints beautiful verbal pictures for his audience, in¬
viting them to see the colors, shapes, and wonders of the moun¬
tains with him, he always leaves them behind as he moves, soli¬
tarily, into the frightful but enlightening adventures of the
“California Alps.” Literary biographers, including Herbert
Smith and Michael Cohen, have noted the antagonism between
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Volume 81 (1993)
65
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
John Muir and the public. In many ways,
this antagonism centered on a question of
access. A solitary wanderer, Muir was for¬
tunate to have access to the mountains,
which enabled him to learn to study the
peaks, in his words, “long and lovingly.” It
is such access that he wished to deny the
tourists, fearing, like later preservationists,
that sheer numbers of unappreciative hu¬
mans would destroy the ecological balance
and massive, expansive grandeur of the wild.
Access requires railroads, coach roads, tour
guides, walking paths, picnic grounds, in
short, an entire institutional structure to ac¬
commodate viewing. These two essays reveal
the conflict in his efforts to fashion the wil¬
derness verbally, to encourage his readers
that the wild was worth preserving, and yet
to bar access to the pristine mountains of
California. Muir’s writings are a tourist’s
guide through the California mountains,
through language and word-painting, but he
is also in opposition to the aims of tourists
and readers. He alone can find sublimity- —
a heightened experience — where others find
views. While he followed established “rules”
of scenic depiction in his writing, he used
those rules to ridicule an audience of tour¬
ists and leisure-class readers who sought
“fine scenes” in nature. My intent is to dem¬
onstrate the extent to which Muir employed
verbal and artistic conventions in his writ¬
ing and to show the degree to which these
conventions carried an intentional, dual in¬
tent.
Herbert Smith contends also that Muir
was a formula writer. Smith points to the
fact that Harper’s Magazine seemed to favor
highly informational articles about exotic
American subjects that could be illustrated
nicely (H. Smith 80), such as stories about
the wonders of nature. However, a more re¬
cent article by Michael Smith notes that
Muir, like his fellow naturalist and writer
Clarence King, was attempting to define a
new style of writing, linking the scientific
with the aesthetic (M. Smith 37). These two
assessments are not mutually exclusive: while
Muir sought to combine scientific writing,
adventure, and scenic description, he still
relied on specific aesthetic conventions in
recreating the grandeur of the California
Alps for his audience. Michael Smith’s judg¬
ment points to an interesting point, how¬
ever. Muir’s writing in “Snow-Storm on
Mount Shasta” and “In the Heart of the
California Alps” is rather rough and experi¬
mental, at times awkward, and it is perhaps
this roughness that has often caused biogra¬
phers, critics, and admirers to turn to the
revised versions in The Mountains of Cali¬
fornia. I believe, however, that these two
early pieces give us a fresher look at the art¬
ist trying to mold his writing to the expec¬
tations of his audience while attempting to
retain his own strong perspective.
The terms “picturesque” and “sublime”
derive from painterly conventions dating
from the eighteenth century. Literally, “pic¬
turesque” means bringing to nature the
qualities of a picture. One of the foremost
works of art criticism that deals with the pic¬
turesque is Christopher Hussey’s The Pictur¬
esque: Studies in a Point of View. In that
work, Hussey identifies the qualities selected
by painters as elements of the picturesque:
. . . roughness, lusciousness of texture, glint¬
ing, sparkling surfaces, the crumbling and de¬
cayed. These they found in the objects now
known as picturesque: sandy lanes, dock
leaves, gnarled trees, hovels, donkeys, and
ruins. Their brushes were attracted to the ren¬
dering of these qualities, because they were
well suited to paint. No moral feeling entered
into the business, though sentiment was at¬
tached to many of these objects, particularly
to rural scenes and ruins. (Hussey 246)
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HELMERS: Creating the California Alps
For painters (and, later, writers) who
adopted its techniques, the picturesque be¬
came a way to moderate, to order, and to
understand the wilderness of “the sublime.”
The sublime was, then, a rough and un¬
tamed version of wilderness, often frighten¬
ing in its aspects. The picturesque aestheti¬
cally cultivated the raw landscape. Actual
landscapes described as picturesque were
ones in which life imitated the regularity of
art, for the real was sought out for its most
painterly qualities.
Hussey points out that the picturesque
was “a practical aesthetic” prescribing rules
to painters for observation and re-creation.
The rules, which Muir applied to his writ¬
ing, include: a scene composed of fore¬
ground, containing stock features such as
rocks, cascades, broken ground, and ruins;
a middleground, containing meadows and
forests; off-skips, or side-screens, containing
valleys, woods, rivers; and a background,
consisting of perfectly pyramidal mountains
and placid lakes (Hussey 116). Particular
colors were in vogue. Greens were not al¬
lowed, while variations of brown were ex¬
pected. Brown was associated with romance
and sublimity, and because it is the season
of browns, autumn became the season fa¬
vored most by painters employing the pic¬
turesque (Hussey 43).
As Romantic poets turned to the pictur¬
esque and sublime for inspiration, certain
words were intended to infuse language with
the onomatopoetic sense of motion — of dis¬
order breaking in on order. These were
known as “words of high coloring” (Hussey
33). The presence of words like “shoot,”
“roll,” “dash,” “wrap,” “bend,” “rear,”
“stretch,” “nod,” “rage,” “gush,” “sweep,”
and “swell” became, like the color brown,
another test for the presence of the pictur¬
esque (Hussey 35). Norman Foerster points
out that Muir employed such onomato¬
poetic words frequently as a means of add¬
ing verisimilitude to what he described, and
Foerster labels this Muir’s “normal method”
of writing (260).
Muir, in fact, combines words that evoke
the sense of the picturesque and the sublime
with painterly conventions of scenic arrange¬
ment. He composes the descriptions in these
essays around orderly “scenes,” often paus¬
ing in his narrative to point out to his audi¬
ence the places where they may take in a pic¬
turesque view. It is not always easy to dis¬
cern the differences between artistic conven¬
tions and literary conventions in his work,
as there is a direct correspondence between
the two. Conventional words of high color¬
ing express the traditional tenets of pictur¬
esque and sublime painterly description. In
her study of the intersection between Ro¬
mantic poetry and conventions of represent¬
ing nature, Marjorie Hope Nicolson has
drawn attention to the ways in which Ro¬
mantic poets depended on and perpetuated
particular literary traditions of representing
mountains, just as Muir chose specific words
to evoke painter’s qualities of color and form
in California’s Yosemite. Nicolson’s work
helps to illuminate the repetitions of stan¬
dard scenic descriptions in Muir’s two es¬
says.
The progress of Muir’s movements
through the two narratives reveals an ascent
from the easily comprehensible world of the
picturesque to the awe-inspiring world of the
sublime. Aesthetically, the picturesque was
meant to have popular appeal. Virtually any¬
one could possess a “picturesque eye”
(Hussey 83-84). The sublime, however, was
a more exclusive experience, appealing to
those with a “Romantic mind.” While the
picturesque eye could be easily enthralled by
an arrangement of rocks and trees, the Ro¬
mantic mind sought the moral implications
of a scene. As Hussey writes, “The Roman-
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
tic mind, stirred by a view, begins to exam¬
ine itself, and to analyze the effects of the
scenery upon its emotions. The picturesque
eye, on the contrary, turns to the scene. ...”
(83-84).
In “Snow-Storm on Mount Shasta” and
“In the Heart of the California Alps” Muir
portrays himself as the possessor of a Ro¬
mantic mind. In both of the essays, Muir is
thrown by circumstances into dangerous
situations which offer him the opportunity
to reflect on the infinite. It is quite clearly
implied in the essays that he alone is able to
experience the sublime. While he often in¬
vites his readers to look upon a picturesque
scene with him, he alone is forced into ex¬
traordinary feats of exertion that leave him
pondering the workings of God in the
world. Surrounded by boiling mud and nox¬
ious gases, the peak of the adventure story
in “Snow-Storm,” Muir seems almost cheer¬
ful to have the opportunity to contemplate
“God’s Design.”' He also records in “In the
Heart of the California Alps” that when fac¬
ing death atop Mount Ritter, hanging from
a shard of rock, he is saved by a “Guardian
Angel.”
Muir constructs himself as the Romantic
type, an exemplary mountaineer, insightful,
sensitive, fearless, and self-sufficient. As
Muir reveals in “Snow-Storm on Mount
Shasta,” he climbed mountains in shirt
sleeves, packed no blankets, and, for suste¬
nance, carried only a crust of bread and a
tin cup with which to scoop water from
mountain streams. Although he never men¬
tions tourists in either of these articles, a
knowledge of Muir’s antagonism toward
popular travel as documented in his letters
and biographies sheds new light on his ethos,
the way in which he constructs himself and
reveals his attitude toward his audience.
Travelers from the East, overburdened with
clothing and provisions and blind to the true
significance and potential danger of the
wilderness around them, are conventional
enough in themselves today, but Muir was
working to fashion distinctions between
those with a shallow perception of nature
and those with a deeper understanding of
nature, between those who were interested
in the picturesque and those who were fas¬
cinated by the sublime. Fearful that the in¬
dustries of farming and tourism would re¬
sult in a desecration of wild spaces, Muir
sought to portray to his audience the need
to retain untouched spaces where the infi¬
nite might be experienced. In other words,
as many people as possible had to remain
behind on the path.
Muir’s extraordinary sensitivity to the
wilderness and his ability to experience the
sublime workings of nature are shown to
good advantage in “Snow-Storm on Mount
Shasta,” which is an unusual essay in that it
combines an introductory section of scien¬
tific explication with picturesque descrip¬
tions and an adventure story. In the essay,
Muir attends to the colors and contrasts of
the landscape, introduces natural ruins, re¬
flects with awe upon the powers of God,
and directly appeals to the reader to behold
scenes as picturesque. It is in “Snow-Storm
on Mount Shasta” that Muir’s efforts to
combine scientific observation with literary
writing seem most forced and awkward.
Early in the essay, he divides the mountain
into three botanic zones and enhances his
verbal description with an illustration of
three evenly concentric circles. He notes the
elevation of each region and its indigenous
flora and provides a list of all the conifer¬
ous trees he has discovered (Muir 323). The
outermost zone, at the base of the moun¬
tain, is the chaparral zone, covered in ever¬
greens and lilies; the inner zone, the fir zone,
is covered in silver-firs; and the uppermost
Alpine zone is covered in snow, dwarf pines,
68
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HELMERS: Creating the California Alps
and some flowering plants. Following this
catalogue, however, Muir shifts perspective,
moving from a focus on the natural sciences
to a focus on the aesthetic.
Muir opens the article by establishing the
differences in texture, temperature, and tone
between glacial ice and volcanic lava, con¬
trasts which will figure prominently in the
climax of the adventure story and which
demonstrate nicely the desire to emphasize
picturesque contrasts:
Mount Shasta, situated near the northern ex¬
tremity of the Sierra Nevada, rises in solitary
grandeur from a lightly sculpted lava plain,
and maintains a far more impressive and
commanding individuality than any other
mountain within the limits of California.
Go where you will within a radius of from
fifty to a hundred miles, there stands the co¬
lossal cone of Shasta, clad in perpetual snow,
the one grand landmark that never sets. . . .
During the glacial period Mount Shasta was
a center of dispersal for the glaciers of the
circumjacent region. The entire mountain
was then loaded with ice, which, ever de¬
scending, grooved its sides and broke up its
summit into a mass of ruins. (521)
Mount Shasta exhibits two qualities es¬
sential to sublime and picturesque descrip¬
tion. Primarily, the mountain is introduced
as a pyramid, part of the “backdrop” of the
picture. The mountain is also irregular, bro¬
ken into a mass of ruins. As Nicolson con¬
tends, irregularity began to replace the aes¬
thetic of regularity in the late eighteenth
century, when Addison praised the “rude¬
ness” of the gardens in France and Italy in
preference to the “regular,” landscaped gar¬
dens of England (317). Acceptance of ir¬
regularity as a valid aesthetic quality gave
new importance to the place of ruins in
scenes. Nicolson notes that ruins were at¬
tractive because of their “asymmetry” (336).
Muir describes the irregular summit of
Mount Shasta as a mass of crumbled rock.
It is, he writes, a natural “ruin.”
The ruins and irregularity have further
conventional associations, for Muir is care¬
ful to describe that they were formed by vol¬
canic eruptions. Volcanoes were important
to formulas of the sublime because the great
geologic upheavals that they caused epito¬
mized the ruining of nature: “The moun¬
tain bursts into flame, and man with all his
works lies buried in the Ruins of Nature”
(Nicolson 341). Ruins of nature reflected in
turn on the frailness of human existence,
which Nicolson refers to as the “Ruins of
Time.” Descriptive poets of the nineteenth
century, most notably Byron and Shelley,
shared the relish of earlier poets for thun¬
derstorm and tempest, earthquake and vol¬
canic eruption (Nicolson 380). Following
the conventional literary usage of words of
high coloring and the fascination with irre¬
gularity, ruins, volcanos, and natural vio¬
lence, Muir paints Shasta’s main summit.
Viewing it from the north, he describes
Shasta as:
... an irregular blunt peaklet about ten feet
high, fast disappearing before the stormy at¬
mospheric erosion to which it is subjected.
Hot sulphurous gases and vapors escape with
a loud hissing noise from fissures in the lava
near the base of the eastern ridge, opposite
the highest peaklet. Several of the vents cast
up a spray of clear bead-like drops of hot wa¬
ter, that ride repeatedly into the air and fall
back until worn into vapor. (Muir 522-23)
As he begins the narrative portion of the
essay — the story of his ascent of the moun¬
tain — he pauses to direct attention to the
view from Strawberry Valley. It is the first
direct appeal to the audience in the essay and
suggests Muir’s awareness of the leisured
reader’s demands for taking scenes. It is an
awkward and theatrical intrusion into the
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
narrative and an abrupt turning away from
the concern with botanical taxonomies.
With the potential for leisured readers to be¬
come inquisitive tourists, Cohen posits that
“all of Muir’s writings were for the tourists,
since they involved the question of how to
see. Most tourists did not want to hear phi¬
losophy, but wanted to know exactly where
to stop and look” (Cohen 207). Muir, per¬
haps experimenting with what he saw as a
necessity of the audience, caters to demands
by writing that:
... at Strawberry Valley there is a grand out-
opening of the forests, and Shasta stands re¬
vealed at just the distance to be seen most
comprehensively and impressively.
Looking at outlines, there, in the imme¬
diate foreground, is a smooth green meadow
with its crooked stream; then a zone of dark
forest, its countless spires of fir and pine ris¬
ing above one another higher and higher in
luxuriant ranks; and above all the great white
cone sweeping far into the cloudless blue. . . .
(524)
The obligatory picturesque elements of this
tri-level view are the meadow and (irregu¬
lar) “crooked stream” in the foreground, the
forest zone “rising above” the foreground as
all well-painted middlegrounds and side-
screens should, and the perfect pyramidal
cone of the mountain, the focal point of the
picture. Earlier in the essay, Muir had al¬
ready written that Shasta was perfectly
drawn, an exquisite pyramid: “[T]he regu¬
larity and symmetry of its outlines remain
unrivaled. The mountain begins to leave the
plain in slopes scarcely perceptible, measur¬
ing from two to three degrees. These are
continued by exquisitely drawn gradations”
to the surmounting crater (522).
Having painted the picture, he steps in
through the frame with his climbing part¬
ner Jerome Fay. “For him,” Cohen says
about Muir, “it was not as important to view
the scene as to be in it . . .” (241). Working
with Fay to record “barometrical observa¬
tions,” his close inspection of the jagged
cliffs and fumaroles of Mount Shasta is reg¬
istered with the knowledge that the tourists
to whom he writes will stay in the valley.
With his step inside the frame, the adven¬
ture story begins.
Fay and Muir make their climb three
months before the regular climbing season
begins and carry instruments designed to
study fluctuations in weather. After two
beautifully clear days, Muir and Fay are en¬
veloped by a violent snowstorm. To give the
reader an idea of the magnitude of the
storm, Muir employs another conventional
element of scenic description, the prospect
view, which establishes “the larger spatial
context in which . . . action takes place”
(Nevius 30). As the heavy clouds begin to
stir and brew, Muir looks about to see that:
[t]he black lava beds made famous by the
Modoc war; many a snow-laden peak far
north in Oregon; the Scott and Trinity
mountains; the blue Coast Range; Shasta Val¬
ley, dotted with volcanoes; the dark conifer¬
ous forests filling the valleys of the Upper
Sacramento — were all in turn obscured, leav¬
ing our own lofty cone solitary in the sun¬
shine, and contained between two skies — a
sky of spotless blue above, a sky of clouds be¬
neath. (Muir 526)
Here again are the contrasts essential to a
picturesque scene: the clouds are absent from
above, but obscure all below. And, as the
snow falls, Muir records that it touched
them “not a whit more harshly than warm
rain on the grass” (528). Muir’s experience
on the mountain would be a sharp contrast
to the experience of an ordinary tourist,
safely within reach of shelter below, for he
finds himself in a world where the trees are
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HELMERS: Creating the California Alps
“crushed by winter snow, and shorn off by
the icy winds,” a world of “frost wind” and
“scalding gas jets.” Muir appears to address
the absent readers as they sit comfortably
fireside with their copy of Harper's :
The ordinary sensations of cold give but faint
conceptions of that which comes on after
hard exercise, with want of food and sleep,
combined with wetness in a high frost wind.
(529)
To warm themselves, Muir and Fay find
their way to a small patch of volcanic earth
and are warmed by the thermal activity. The
two explorers spend seventeen hours on this
quarter acre of ground where contrasting
temperatures pose dangers to their lives.
From above they are threatened by the icy
snowstorm; from below noxious fumes
could poison them:
The acrid incrustations sublimed from the
escaping gases frequently gave way, opening
new vents, over which we were scalded; and
fearing that if at any time the wind should
fall, carbonic acid, which usually forms so
considerable a portion of the gaseous exhala¬
tions of volcanoes, might collect in sufficient
quantities to cause sleep and death, I warned
Jerome against forgetting himself for a single
moment. . . . (528-29)
They suffer “the pains of a Scandinavian
hell, at once frozen and burned” (Muir 529).
It is a truly sublime experience, filled with
terror, yet offering opportunities to reflect
on life, death, and the infinite, which Muir
and Fay (dutifully observing the conventions
of sublimity) do. Fay wonders if prayers
would help them, but Muir seems to dis¬
suade him from praying with a rather deter¬
ministic speech about “the unflinching fair
play of Nature.” Violent tempests that
threaten the lives of humans are all part of
the Design: “Life is ... a mere fire, that now
smoulders, now brightens, showing how eas¬
ily it may be quenched” (Muir 529).
Muir and Fay survive the night. “Snow-
Storm on Mount Shasta” ends abruptly as
they slide and shuffle into their camp, im¬
peded by frozen trousers and hunger. Their
descent is a descent from the sublime to the
picturesque, both scenically and intellectu¬
ally. For the sublime is the world of the Ro¬
mantic and the Romantic’s inward-turning
eye; it inspires awe because it is too large to
comprehend. But the picturesque can be ap¬
preciated by anyone who is familiar with
painting; the picturesque is simple, flat, con¬
tained. When Muir returns to the chaparral
zone he steps back through the picture
frame. From the safety and comfort of his
distant hotel room he may admire, for the
audience, the view in the frame: the next
morning “from the window I saw the great
white Shasta cone wearing its clouds and for¬
ests, and holding them loftily in the sky”
(Muir 530). In his assessment of Muir’s life
and work, Frederick Turner has noted that
the cheerful ending gives no clue that Muir
was suffering from severe frostbite (229).
“In the Fleart of the California Alps,” like
“Snow-Storm on Mount Shasta,” also places
an adventure story within an artistic frame.
Muir uses the elements of the picturesque
and the sublime more self-consciously than
in the earlier piece, often drawing attention
to the qualities of nature that are similar to
the elements of a painted landscape. For ex¬
ample, he sets his narrative in Indian sum¬
mer and opens by “painting” the scene of
the Tuolumne Valley with lavish detail:
The intense azure of the sky, the purplish
grays of the granite, the red and browns of
dry meadows, and the translucent purple and
crimson of huckleberry bogs; the flaming yel¬
low of aspen groves, the silvery flashing of the
streams, and the bright green and blue of the
glacier lakes. (Muir 346)
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The writing in this essay is less rough than
“Snow-Storm on Mount Shasta,” for Muir
dispenses with diagrams and scientific pre¬
tenses and draws attention to the artistic
qualities of the wilderness instead. He takes
as his companions two artists in search of “a
landscape suitable for a large painting”
(Muir 346). Despite the fact that one of the
artists (who remains nameless in the essay)
was a close friend of Muir’s named William
Keith (Turner 209), it is clear from the text
that the painters are constructed as posses¬
sors of picturesque eyes, interested only in
what they can paint and not in the finer, less
accessible, but more rewarding aspects of
sublime experience.
Symbolically, Muir’s ascent of Ritter rep¬
resents a passage out of the banal apprecia¬
tion of the picturesque to the greater under¬
standing of the sublime. At times, however,
Muir seems somewhat more sympathetic to
the picturesque qualities of the valley, at one
point remarking that the valley was waiting
for “the elected artist” — “I could not help
wishing that I were that artist,” he laments
(346). Nevertheless, although he is wont to
throw up his arms “to inclose [the Tuo¬
lumne Valley] as in a frame” (346) and to
wax eloquent on its majesty, it is significant
that he eventually leaves the painters behind
to sketch views while he alone accepts the
challenge of climbing to the summit of
Mount Ritter. Frederick Turner remarks, “as
far as anyone knew it had never been
climbed. Moreover, at this season (the last
days of August), though the weather was still
clear, there was the ever-present danger of
snow. For Muir, all the conditions were
right for climbing ...” (209). Ritter posed
a challenge for the individualist Muir.
At the start of the ascent to Mount Ritter,
Muir notices that the best view of the Cali¬
fornia Alps comes from the headwaters of
the Tuolumne River. Muir directs the
reader’s gaze to a scene he describes as “in a
high degree picturesque, and in all its main
features so regular and evenly balanced as
almost to appear conventional” (343). He
divides it into the traditional artistic fore¬
ground, middleground, and background.
The foreground is the “magnificent valley”
resplendent in autumn colors of brown,
purple, and gold. It is “smooth, meadowy”
and “level,” dotted with “dipping willows
and sedges” and “groves of arrowy pine.”
Through the foreground the waters of the
Tuolumne flow from their source in the
middleground, where it pours from “crystal
fountains” and leaps in “white cascades.”
The middleground also contains the restrict¬
ing “off-skips,” narrowing and focusing the
viewer’s eyes toward the background. The
off-skips in this painting are the “granite
bosses” and the walls of the valley, “beveled
away on both sides so as to embrace it all
without admitting anything not strictly be¬
longing to it.” The background is colored
in contrast to the foreground. The sky is co¬
balt blue, the glaciers are black, gray, and
“pure, spiritual white.” The focal point of
this picture is “one somber cluster of snow¬
laden peaks . . . surging free into the sky”
from the valley (343).
Curiously, while cataloguing the pictur¬
esque features of the scene from the valley,
Muir asserts that “[f]ew portions of the Cali¬
fornia Alps are, strictly speaking, pictur¬
esque” (345). He argues that making a pic¬
ture of these Alps would require separating
the magnificence of the range to allow the
discrete picturesque elements to be appreci¬
ated in isolation. However, he asserts, sepa¬
rating each mountain from the others so it
might serve as the focal point of a painting
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HELMERS: Creating the California Alps
would damage the incredible impression
made by the whole range:
The whole massive uplift of the range, four
hundred and fifty miles long, by about sev¬
enty wide, is one grand picture, not clearly
divisible into smaller ones; in this respect it
differs greatly from the older and riper moun¬
tains of the Coast range. . . . But all were not
brought forth simultaneously; and, in general,
the younger the mountain landscapes, the less
separable are they into artistic bits capable of
being made into warm, sympathetic, lovable
pictures. (345)
The artist companions, functioning as
tourist-surrogates, are searching for such “ar¬
tistic bits.” Early in their journey into the
mountains, they comment, “All this is sub¬
lime, but we see nothing as yet at all avail¬
able for effective pictures” (Muir 346). Yet,
although they are able to recognize the ele¬
ments of the sublime, they are without feel¬
ing for it. The sublime does not create in
them the appropriate response of fear and
wonder. They are innocents, unable to
wrestle with the implications of God in the
wilderness. Their Puritan notion of the wil¬
derness as something alien to humans, some¬
thing to be conquered, gives root to their
desire to tame the wild through pictorial rep¬
resentation — reducing it to a canvas three
feet by four feet, imposing composition on
the chaotic elements. The artists seek some¬
thing recognizable, something known, some¬
thing familiar, like the tourists on their ap¬
proved path, following a conventional guide¬
book.
In the earlier article, Muir positioned
himself in opposition to an assumed audi¬
ence of tourists. He was able to find sublim¬
ity in a landscape where others found only
views. It is evident in “In the California
Alps” that Muir values the primacy of his
own experience, and he hints that only he
has the strength and the understanding to
come to love the mountains, asserting that
it is only after the mountains have been stud¬
ied “one by one, long and lovingly,” that one
can begin to understand their full grandeur.
Ironically, “loving” the mountains embod¬
ies the very aesthetic that he opposes — fa¬
miliarizing the mountains into something to
be framed and placed over a mantlepiece,
destroying their ability to inspire fear and
awe. In admitting his own attachment to the
mountains, Muir deviates significantly from
the conventional sublime response to the
peaks — terror (Nicolson 356).
Loving, or “familiarizing,” according to
Walker Percy, is an inevitable part of being
a tourist. Percy notes that visual, touristic,
experience is bound by a “symbolic com¬
plex,” a formulation of expectations prefig¬
ured by textual treatments (55). Satisfaction
is measured by the way the tourist’s experi¬
ences conform to the symbolic complex. So
pervasive were the conventions of the pic¬
turesque in the nineteenth century (as John
Sears in his work on nineteenth-century
tourism and Nicolson have emphasized)
that Muir’s artist friends could not see be¬
yond their expectations. They instead seek
the familiar: a landscape suitable for paint¬
ing. With the popular European Alps etch¬
ing the scheme of mountains in their minds,
Muir’s artists eventually find a “landscape
suitable for a large painting,” and Muir
records that one of the artists “dashed ahead,
shouting and gesticulating and tossing his
arms in the air like a madman. Here, at last,
was a typical Alpine landscape” (346). With
this incident, Muir not only points out to
his readers that typical Alpine scenes are
available for viewing in California, he shows
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the extent to which searching for the typi¬
cal is ridiculous. So excited is this artist at
finding a landscape which matches his con¬
ception of the European Alps, that he is
blind to the authentic and particularly Cali¬
fornian charms of the Sierras. In this, as in
other passages in these two essays, Muir in¬
vites readers to partake of the scenery while
simultaneously drawing attention to the
shortcomings of prefigured vision.
While the artists decide to remain at their
mountain camp and sketch, Muir plans to
continue climbing to the summit of Mount
Ritter heretofore only visible as the focal
point of the background, the “grand mas¬
terpiece.” Again, as in “Snow-Storm on
Mount Shasta,” he steps through the fore¬
ground and middleground of the picture he
paints. As he crosses the foreground, he
traces “happy streams” to the “foot of a
white cascade,” which announces the begin¬
ning of the middleground, where he finds
“painted meadows” and silvery lakes. When
Muir begins to climb through the middle-
ground toward his destination of the back¬
drop, he is able to look back on the land he
has just traversed. It emerges as a prospect
view, the far reaches of which exist in Muir’s
imagination:
Over the summit, I saw the so-called Mono
desert lying dreamily silent in thick, purple
light — a desert of heavy sun-glare beheld
from a desert of ice-burnished granite. Here
the mountain waters divide, flowing east to
vanish in the volcanic sands and dry sky of
the Great Basin; west, to flow through the
Golden Gate to the sea. (347)
As Muir pushes further toward the sum¬
mit of Mount Ritter, he moves deeper into
the realm of the sublime. The landscape now
is full of “savage peaks,” “spurs,” and plung¬
ing “gorges”; it is sparse and bare. Such sur¬
roundings are the surroundings of the Ro¬
mantic and Muir writes:
In so wild and so beautiful a region my first
day was spent, every sight and sound novel
and inspiring, leading one far out of oneself,
yet feeling and building a strict individual¬
ity. (347)
Although the picturesque is still evident
here, in “little gardens” that decorate natu¬
ral streams and pools, for instance, Muir is
in the realm of the sublime. Sights and
sounds are “novel.” Like ruins atop hills, the
flat-topped spurs are “marked and adorned
with characteristic sculptures of the ancient
glaciers that swept over this entire region like
one vast ice-wind . . .” (347). When Muir
sets his camp, he is encircled by “somber
peaks, hacked and shattered . . . wearing a
most savage aspect.” A waterfall runs nearby,
and “scraggy pines” in “rock-fissures” were
“dwarfed and shorn” by wind (348).
Crossing mountains, Muir arrives at the
final summit before that of Mount Ritter:
There, immediately in front, loomed the ma¬
jestic mass of Mount Ritter, with a glacier
swooping down its face nearly to my feet,
then curving westward and pouring its fro¬
zen flood into a dark blue lake, whose shores
were bound with precipices of crystalline
snow; while a deep chasm drawn between the
divide and the glacier separated the massive
picture from everything else. (349)
Although he is now amidst the sublime,
the composition of the scene upon Muir’s
arrival atop the final summit before that of
Mount Ritter is again a basic three-part
structure recalling the picturesque. The fore¬
ground consists of the summit upon which
he stands, the middleground is a grouping
of a deep chasm, a glacier, and a lake toward
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HELMERS: Creating the California Alps
the left side of the side-screen, and the back¬
ground is the peak of Ritter itself. The at¬
mosphere is far from placid and ordered,
however. He faces steep gullies and vertical
cliffs, which only emphasize the size of the
mountain that lies ahead. Muir is atop an
adjoining summit, so his view is not from
the base of Mount Ritter, but from an el¬
evation equal to about a third to half of
Ritter. His vantage point does not come
close to equalizing the relationship between
his summit and the one that lies in front.
Instead “massive battlements” stand forth,
roughly hewn and shadowed. Even more
telling, “huge, crumbling buttresses” extend
to Muir’s right and to his left, as far as he
can see (Muir 349).
Suddenly, Muir’s descriptions take on
eery and unearthly overtones that seem to
correspond to his reach into the sublime at¬
mosphere that surrounds the summit of the
mountain. Muir begins to employ bodily
metaphors to emphasize the seemingly in¬
comprehensible size of Mount Ritter, lying
ahead; he defamiliarizes what had earlier
been familiar, “loved,” even understood.
Turner recalls that Muir, traveling in his
early years, had experienced a “dangerous
weakness” from lack of bread (189), and per¬
haps the visions that he experiences here
were induced by such deprivations. While
the “head of the glacier sends up a few fin¬
ger-like branches,” Muir picks his way
through numerous “narrow- throated gullies”
and “across the yawning chasm,” he hears
the “gurgling of small rills down in the veins
and crevasses,” and he discovers “the mouth
of a narrow avalanche gully.” These human¬
like metaphors serve to reduce the extraor¬
dinary size and distance which he confronts,
a technique employed, for example, by
painter Thomas Cole in his Snow Squally
Winter Landscape in the Catskills. In a book
that offers a psychoanalytical reading of
American art history, Bryan Jay Wolf com¬
ments that the viewer will relate to the fore¬
ground plane of Cole’s painting, a rocky
promontory which juts vertically into a va¬
cant chasm. A wolf and a few barren trees
are the visible inhabitants of that promon¬
tory. However,
[t]he wolf and trees that accommodate him,
however uncomfortably, to the promontory
world, individuating it and rendering it on
human scale, either disappear in the middle-
ground space or reappear in the case of the
background forests in such proportions that
their scale seems threatening and annihilat¬
ing. ... By traversing the valley and encoun¬
tering the change of scale it implies, the
viewer undergoes a process of “defamil¬
iarization.” The veil of familiarity is lifted
from the face of nature and an alien strange¬
ness left in its stead. (186)
Once looked upon “long and lovingly,” the
mountains are now something grotesque and
fearsome, underscoring the deathly situation
in which Muir soon finds himself: “at the
foot of a sheer drop in the bed of the ava¬
lanche channel” (330). Although he clings
to rocky footholds and handholds he fears
that his doom is fixed. “I must fall. There
would be a moment of bewilderment, and
then a lifeless rumble down the one general
precipice to the glacier below” (Muir 330).
If, as Hussey sets forth, the mountains are
“a memento mori on a gigantic scale” (Hussey
55), it is fitting that Muir should encounter
imminent death clinging to the face of the
most sublime of all peaks. It is significant,
too, that his faith in the infinite once again
allows him to survive:
I seemed suddenly to become possessed of a
new sense. The other self — the ghost of by¬
gone experiences, Instinct, or Guardian An¬
gel — call it what you will — came forward and
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assumed control. Then my trembling muscles
became firm again, every rift and flaw in the
rock was seen as through a microscope, and
my limbs moved with a positiveness and pre¬
cision with which I seemed to have nothing
at all to do. Had I been borne aloft upon
wings, my deliverance could not have been
more complete. (Muir 350)
Above the place where Muir faces his des¬
tiny, he describes the mountain as a place
“savagely hacked and torn,” chasms “yawn,”
“detached bowlders [sic]” fill the “crags.”
Nevertheless, befitting the mystical aesthetic
of the sublime, despite its potentially hor¬
rendous consequences, the peak and his ex¬
perience there is “glorious,” and Muir stands
suffused in “blessed light” (350).
Following his harrowing experience, Muir
descends toward the valley, returning to the
realms where the familiar scenic conventions
of the picturesque dominate. The landscape
becomes more composed. Mountains arch
into the background, their peaks “swelling
higher, higher as they sweep on southward”
(350). Natural ruins abound: Cathedral
Peak is “a temple of marvelous architecture,”
a mountain seems a “gigantic castle with tur¬
ret and battlement, the clusters of Alps are
“eloquent monuments of the ancient ice-riv¬
ers that brought them into relief’ (351).
Back at the camp, Muir finds the artists
worrying over his safety. They feared that he
had been overtaken by the elements, not
knowing that it was he who triumphed over
the elements. Muir comments that their
troubles seemed “curious.” After all, his jour¬
ney had been “only a matter of endurance
and ordinary mountain-craft,” and he had
been absent only three days when he had
warned that he might be absent for over a
week (Muir 352).
The artists pack “their precious sketches,”
concluding the “fine double excursion” of
artists and scientist into the picturesque and
the sublime.
Muir’s uses of the conventions of pictur¬
esque and sublime description were an in¬
evitable result of the literary climate in which
he worked. Yet, as I mentioned earlier, there
is also a strong subversive element in his
writing. Literary critic Marvin Fisher asserts
that Muir’s contemporary Herman Melville
was a subversive writer, forced to “go under¬
ground” in order to satirize the very audi¬
ence for whom he wrote. These two essays,
when examined in light of Muir’s own com¬
ments elsewhere and revelations of his pre¬
dispositions by his biographers, seem to in¬
dicate that Muir too sought to satirize and
patronize an audience upon whom he de¬
pended for financial support. Muir found
himself linked to an audience for whom he
held little respect. Fearful of the encroach¬
ments of technology, he was fighting to pre¬
serve the wilderness of the American West.
He realized that his stories of the mountains
would raise public awareness of western
beauty and would be instrumental in enjoin¬
ing Americans to protect the wild regions of
Yosemite from farming and industry. Cohen
points out that “Americans would need
some encouragement, would need to know
not only what they possessed, but why it was
worth protecting and how this could be
done” (205). Because the literary conven¬
tions of the sublime and the picturesque
were well worn and would be well recog¬
nized by his readers, their effect was to make
an unfamiliar landscape familiar to readers
living thousands of miles away, something
politically necessary.
Muir’s texts are problematic, revealing a
narrator in conflict about his role as preser¬
vationist and writer, for he knew his stories
would extend beyond the page to encour¬
age tourism, and he worried that the pris¬
tine land would be overrun by tourists in
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HELMERS: Creating the California Alps
search of the picturesque. In his biography
of Muir, Cohen notes that it was difficult
for Muir to hide his contempt of compla¬
cent tourists, eager only for a view and not
for knowledge about their surroundings
(129, 207). To illustrate his point, Cohen
draws from this letter written by Muir to his
friend Jeanne Carr. Muir wrote, “They
climb sprawlingly to their saddles like over¬
grown frogs pulling themselves up a stream
bank through the bent sedges, ride up the
Valley with about as much emotion as the
horses they ride upon — are comfortable
when they have ‘done it all’ and long for the
safety and flatness of their proper homes”
(Bade 220). His attitude toward the leisured
class takes on importance in a study of his
style, for it seems ironic that one so vehe¬
mently opposed to the domestication of the
wild would seek not only to entice visitors
forward, but would, though his very lan¬
guage, reduce Nature into a series of “fine
scenes” for the pleasure of the audience.
Yet Muir was prepared to pander to the
“frogs,” and he did so at least in part through
his application of the conventions of pictur¬
esque description. In his verbal landscapes,
mountains become frosty cones settling into
backdrops; sparkling lakes and twisted trees
dapple his foregrounds. And it is quite sur¬
prising that Muir often pauses in his narra¬
tion to point out “views.” Coupled with his
own stories of adventure and extreme expo¬
sure to inhospitable elements, the views seem
inconsequential, almost afterthoughts. At
times, especially after the more harrowing of
Muir’s adventures, his references to fine
scenes seem a harshly reductive view of na¬
ture. Like the artists who accompany him on
his journey into the heart of the California
Alps, Muir loved mountains. His prose
made them accessible, beautiful, and orderly,
rather than terrifying and forbidding. In
many ways, through his writing, Muir de¬
stroyed the aspects of the mountains that he
prized most: their seclusion, their remoteness
from human intervention, their inhospi-
tability. Described according to traditional
artistic and literary conventions, the Sierras
lost their uniqueness, becoming just another
typical example of an aesthetic norm deter¬
mined by the European Alps.
When the artists with whom he travels
into the California Alps pack their sketches,
they close the book on nature. Muir, how¬
ever, distinguishes himself as a privileged ob¬
server by finding the book continuously
open. He would return to read “the records
she has carved on the rocks, reconstruct . . .
the landscapes of the past” (Alps 351). While
Muir textually domesticated nature, reduc¬
ing it into a series of fine scenes for an ap¬
preciative audience, he preferred to leave the
reality wild, a setting for his many returns
to the summits and peaks.
Works Cited
Bade, William Frederick. The Life and Letters of
John Muir. Vol. I. Boston and New York:
Houghton Mifflin, 1924.
Cohen, Michael P. The Pathless Way: John Muir
and the American Wilderness. Madison: Univ.
of Wisconsin Press, 1984.
Fisher, Marvin. Going Under: Melville’s Short
Fiction and the American 1850’s. Baton
Rouge: Louisiana State Univ. Press, 1977.
Foerster, Norman. Nature in American Litera¬
ture. New York: Macmillan, 1923.
Hussey, Christopher. The Picturesque: Studies in
a Point of View. London: Archon Books,
1967.
Muir, John. “In the Heart of the California
Alps.” Scribner’s Monthly Magazine 20
(1880): 345-52.
- . “Snow-Storm on Mount Shasta.”
Harper’s New Monthly Magazine 55 (1877):
521-30.
Nevius, Blake. Cooper’s Landscapes: An Essay on
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
the Picturesque Vision. Berkeley and Los An¬
geles: Univ. of California Press, 1976.
Nicolson, Marjorie Hope. Mountain Gloom and
Mountain Glory: The Development of the Aes¬
thetics of the Infinite. Ithaca: Cornell Univ.
Press, 1959.
Percy, Walker. “The Loss of the Creature.” The
Message in the Bottle. New York: Farrar,
Straus and Giroux, 1975.
Sears, John F. Sacred Places: American Tourist
Attractions in the Nineteenth Century. New
York: Oxford Univ. Press, 1989.
Smith, Herbert F. John Muir. New York:
Twayne, 1965.
Smith, Michael L. “Clarence King and John
Muir: Ways of Seeing Mountains.” The Cali¬
fornians March-April 1990: 36-41.
Turner, Frederick. Rediscovering America: John
Muir in His Time and Ours. New York: Vi¬
king, 1985.
Wolf, Bryan Jay. Romantic Re-Vision: Culture
and Consciousness in Nineteenth-Century
American Painting and Literature. Chicago:
Univ. of Chicago Press, 1982.
Marguerite Helmers is Assistant Professor of En¬
glish at the University of Wisconsin Oshkosh.
78
Robin E. Jung
Blanchard's cricket frogs
(Acris crepitans blanchardi)
in southwest Wisconsin
Abstract State-endangered Blanchard's cricket frogs (Acris crepitans blanchardi) were
censused in southwest Wisconsin and found at 19 of 40 sites which historically
were known or were thought to have had populations of this species. Blanchard’s
cricket frogs were found most often at sites with mud and vegetation hanks
and shallow slopes leading to the water , although these trends were not statis¬
tically significant (P > 0.15). Neither habitat variables (e.g, water depth, bank
type) nor indices of water quality (pH, conductivity, oxidation-reduction po¬
tential, turbidity) differed significantly between sites with and without
Blanchard’s cricket frogs. However, water temperature was significantly greater
at sites with cricket frogs than without (P = 0.011). The total number of frog
and toad species at a site ( range = 0 —7) was positively correlated with water
temperature (P = 0.017) and negatively correlated with turbidity (P = 0.021).
In addition, significantly more anuran species were found at sites without ag¬
ricultural fields nearby (P = 0. 0026).
Recent reports of declining amphibian populations around
the world (Blaustein and Wake 1990; Lohmeier 1990;
but see Pechmann et ah 1991) have instigated research efforts
toward understanding potential causes (e.g., habitat loss, pes¬
ticides, acid rain, drought). In Wisconsin, the Blanchard’s
cricket frog (Acris crepitans blanchardi) has declined precipi¬
tously during the last two decades (Vogt 1981; Mossman and
Hine 1985) and has been on the State Endangered list since
1982 (Bureau of Endangered Resources 1989). Habitat loss,
drought, and polluted water are a few of the factors hypoth¬
esized to have caused Blanchard’s cricket frog population de¬
clines (Minton 1972; Oldham 1992; L. A. Wilsmann, pers.
comm.). Wisconsin represents the northern limit of the geo¬
graphic range of Blanchard’s cricket frogs (Conant and Collins
1991), and it is possible that routine physiological stresses (e.g.,
TRANSACTIONS Volume 81 (1993)
79
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during overwintering; cf. Bradford 1983)
may be relatively high. Thus, Wisconsin
populations of this species might be particu¬
larly sensitive to additional stressors, such as
those related to deteriorating water quality.
M. J. Mossman and R. L. Hine (Moss-
man and Hine 1984, 1983) of the Wiscon¬
sin Department of Natural Resources
(WDNR) initiated the Wisconsin Frog and
Toad Survey in 1981. Relatively few (2 of
63) of the early WDNR survey routes, how¬
ever, were located in southwest Wisconsin
(Grant, Iowa, and Lafayette counties), where
Blanchard’s cricket frogs still occur (Moss-
man and Hine 1983).
The purpose of this study was to survey
Blanchard’s cricket frogs in southwest Wis¬
consin, and to determine whether selected
habitat and water quality variables were re¬
lated to the presence or absence of Blan¬
chard’s cricket frogs or other frog and toad
species.
Methods
During the summer of 1991, I conducted a
census of the Blanchard’s cricket frog in
southwest Wisconsin (Dane, Grant, Iowa,
and Lafayette counties). I chose to study 40
sites which, according to records provided
by the WDNR (Wisconsin Natural Heritage
Inventory Program; Wisconsin Frog and
Toad Survey, M. J. Mossman, pers. comm.)
and the Milwaukee Public Museum (Wis¬
consin Herpetological Atlas Project, G. S.
Casper, in prep.), were historically reported
to support Blanchard’s cricket frog popula¬
tions. The sites were visited an average of 2.6
± 0.99 times (range = 1—4) between 1 1 May
and 17 July, during the peak chorusing sea¬
son for cricket frogs (Vogt 1981). Most sites
were visited at least once during the day and
once at night. If no chorusing was heard, a
tape recording (“Wisconsin frogs,” produced
by R. Anderson and D. Jansen, University
ofWisconsin-Stevens Point) of Blanchard’s
cricket frog vocalizations, which is thought
to elicit chorusing, was played for two min¬
utes (Mossman and Hine 1985). Frogs and
toads were identified by call, sight, and/or
catching animals by hand. Numbers of call¬
ing males were estimated by walking the
water’s edge, noting differences in directions
of calls, and listening for distinct vocaliza¬
tions. Spring peepers (. Hyla crucifer crucifer ),
western chorus frogs (. Pseudacris triseriata
triseriata ), northern leopard frogs ( Rana
pipiens pipiens) and pickerel frogs {Rana
palustris ) were not adequately censused as
they chorus primarily from March to May
(Vogt 1981). Seven of the sites I censused
were also covered in 1991 by the Wiscon¬
sin Frog and Toad Survey, so I incorporated
additional species-presence data from this
source as well.
Water samples were taken approximately
0.5 m from the water’s edge. Four indices
of water quality (temperature, pH, conduc¬
tivity, oxidation-reduction potential [ORP])
were measured using a Cole-Parmer Water
Test, Model 05556-00. Conductivity repre¬
sents the total concentration of electrolytes
in solution (higher conductivities corre¬
sponding to higher ion content, particularly
Ca++, Mg++, Na+, and K+) and is expressed
in micro-Siemens (Wetzel 1983). ORP,
measured in millivolts (mV), is proportional
to the equivalent free energy change per
mole of electrons associated with a given re¬
duction. Large, positive ORP values signify
strongly oxidizing waters. In addition, tur¬
bidity of a water sample in a beaker was es¬
timated by eye and scored on a scale of 0-
5, ranging from clear to muddy.
Habitat variables assessed included: (a)
whether the site was a lake/pond or stream/
river, (b) water depth 1 0 cm from the edge,
(c) percent cover of water area by rooted
80
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JUNG: Blanchard’s cricket frogs in southwest Wisconsin
aquatic plants, (d) percent cover of water
area by algae, (e) composition of water bot¬
tom (mud, mud and rock), (f) bank type
(mud, mud and vegetation, vegetation), (g)
bank slope (flat, flat and steep, steep), and
(h) whether the site was adjacent to pasture
land (primarily with cows) and/or agricul¬
tural land.
All statistical analyses were performed us¬
ing SPSS (Norusis 1988). Significance was
judged at P < 0.05. Yates’ correction was
used for chi-square tests of independence
when any expected frequencies were < 5. For
water quality analyses, I used mean values
for pH, water temperature, conductivity,
and ORP; I also computed a mean visita¬
tion date for each site to use as a covariate
in statistical analyses.
Results
Habitat data for 39 sites (excluding Iowa
County site 16) are presented in Table 1. No
habitat data were collected for Iowa County
site 1 6 because the lake had been completely
drained in the spring of 1991, and Blan¬
chard’s cricket frogs no longer occurred
there. Site locations are not presented here
(see Wisconsin State Statute 23.2 7, section
3, paragraph B; Casper 1989), but are avail¬
able from the WDNR Natural Heritage In¬
ventory Program upon request. Blan¬
chard’s cricket frogs were found at 19 of the
40 (48%) sites which historically were re¬
ported to have supported cricket frogs. I es¬
timated 125 cricket frog calling males at
18 sites (Table 1). Assuming equal sex ra¬
tios (see Pyburn 1958), the population esti¬
mate (males and females) for these sites is
250 frogs, yielding a mean ± standard de¬
viation (SD) of 14 ± 9.6 frogs per site.
Blanchard’s cricket frogs were heard cho¬
rusing during the day (between 1 1 a.m. and
6 p.m.) on seven occasions, but were most
often heard chorusing after 6 p.m. (18 oc¬
casions). Breeding choruses of the Blan¬
chard’s cricket frogs tended to be dense
(males within one meter of each other) and
localized in mudflat areas or in vegetated
shallow water habitat (see Perrill and Shep¬
herd 1989). Exact densities of frogs or in-
ter-male spacing were not recorded in order
to avoid disturbing frogs and habitat and
also because of the difficulty in delineating
how much area comprised suitable habitat.
Chi-square and Mann-Whitney U tests
indicated that the presence of Blanchard’s
cricket frogs was not related to the habitat
variables I recorded. Of the 19 sites where
Blanchard’s cricket frogs were present, 12
were categorized as stream/river habitats and
seven as pond/lake sites; cricket frogs were
no longer present at 1 5 stream/ river sites and
6 pond/lake sites (X2 = 0.311, d.f. = 1, P =
0.577).
Most Blanchard’s cricket frog populations
were found at sites with flat slopes leading
to the water’s edge (17 of 19) and mud and
vegetation banks (17 of 19; all with mud¬
flats). However, because most sites surveyed
had flat slopes (34 of 39) as well as mud and
vegetation banks (30 of 39), the distribution
of Blanchard’s cricket frogs did not differ
from random with respect to these habitat
variables (X2: P = 0.614 and 0.150, respec¬
tively). Only 6 of the 19 sites with Blan¬
chard’s cricket frogs were adjacent to agri¬
cultural (primarily corn) fields, yet a chi-
square test comparing presence and absence
of cricket frogs at sites adjacent versus not
adjacent to agricultural fields was not signifi¬
cant (X2 = 1.766, d.f. = 1, P = 0.184). No
relationships were found between the pres¬
ence versus absence of Blanchard’s cricket
frogs and pasture land, bottom type, water
depth, or percent cover of algae or rooted
aquatic plants. In summary, with respect to
the variables measured, Blanchard’s cricket
Volume 81 (1993)
81
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1 . Data for 39 sites in southwest Wisconsin.
Key: BC = Blanchard’s cricket frog (0 = absent, 1 = present); #BC = estimated # of BC frogs;
S = # of anuran species; H = Habitat (0 = stream/river; 1 = lake/pond); WD = Water Depth
(cm); T = water temperature (C); C = conductivity (uS); ORP = oxidation-reduction potential
(mV); Tu = Turbidity (0-5, clear-muddy); Aq = % rooted aquatic plants; Al = % algae; Be
(bottom composition): 1 = mud, 2 = mud/rock; Bt (bank type): 1 = mud, 2 = mud/vegetation, 3
= vegetation; SI (slope): 1 = flat, 2 = flat/steep, 3 = steep; Cow: 0 = absent, 1 = present; Cr: 0
= no agricultural fields nearby, 1 = fields nearby. -9 indicates missing data.
Site BC #BC S H WD pH T C ORP Tu Aq Al Be Bt SI Cow Cr
Dane
1 0 0 3 1 23 6.8 23.5 319 180 4 3 3 1 3 1 0 0
Lafayette
Iowa
Grant
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JUNG: Blanchard’s cricket frogs in southwest Wisconsin
frogs had disappeared from sites whose habi¬
tat characteristics were similar to sites where
they still occurred.
Using t-tests (Mann- Whitney U tests for
turbidity), only one water quality variable,
temperature, differed significantly between
sites with and without Blanchard’s cricket
frogs (t = 2.68, d.f. = 37, P = 0.011). The
mean (± SD) water temperature for sites
with Blanchard’s cricket frog was 27.22 C
(± 2.971), as compared to 24.03 C (± 4.297)
for sites without cricket frogs. Because wa¬
ter quality variables may vary with date, I
also performed analyses of covariance pre¬
dicting water quality variables with date and
presence/absence of cricket frogs (scored as
a 0-1 dummy variable). Both date (F =
4.991, P = 0.032) and presence/absence of
cricket frogs (F = 6.371, P = 0.013) were sig¬
nificant predictors of temperature. How¬
ever, presence/absence of cricket frogs was
not a significant predictor of pH (mean
range = 6.8- 8.8), conductivity (mean range
= 191-863), ORP (mean range = 41-191),
or turbidity.
Blanchard’s cricket frogs were found most
often with green frogs ( Rana clamitans
melanota\ 15 of 19 sites = 79%) and to a
lesser extent with eastern gray tree frogs
(. Hyla versicolor) and American toads ( Bufo
americanus) (63% and 53%, respectively).
Green frogs occurred most frequently, re¬
corded in 29 of the 39 sites.
Thirty-four of the 39 sites (87%) sup¬
ported anuran populations (Table 2). The
greatest number of species recorded at any
one site was seven (including data from the
Wisconsin Frog and Toad Survey). To de¬
termine whether the number of species
present at a site was related to the categori¬
cal habitat variables, I categorized sites as
having either few (0-2) or many (3-7) spe¬
cies. Eight of 12 pond/lake sites had many
species, whereas 21 of 27 stream/river sites
had few species (X2 = 5.33, d.f. = 1, P =
0.021). Also, 13 of 22 sites situated away
from agricultural fields had many species,
whereas 15 of 17 sites adjacent to agricul¬
tural fields had few species (X2 = 9.07, d.f.
= 1,P = 0.0026).
Species richness showed a significant posi¬
tive correlation with water temperature
(Pearson’s r = 0.381, n = 39, P = 0.017) and
a negative correlation with turbidity (Spear¬
man’s r = -0.395, n = 34, P = 0.021) (Table
3). In a multiple regression analysis, only
temperature was a significant predictor of
species richness. pH was significantly posi¬
tively correlated with temperature, and nega¬
tively correlated with both conductivity and
turbidity (Table 3).
Discussion
My estimates of Blanchard’s cricket frog
populations at each site are extremely con¬
servative, because I did not attempt to cap¬
ture all frogs, and frogs may not have been
chorusing during visits. Even if frogs were
chorusing, I may have underestimated the
number of males since some males assume
noncalling, or satellite, positions (see Perrill
and Magier 1988). The sampling method I
used represents only the greatest number of
calling males heard at a site during short vis¬
its, and therefore is tenuous. Indeed, at two
sites in Iowa County (sites 2 and 3) where I
could distinguish only 27 male Blanchard’s
cricket frogs, D. Nicolai (Mossman and
Hine 1985) counted over 220 frogs in the
early 1980s. This situation may very well be
the case at the other sites, a fact that points
to the need for intensive censusing (e.g.,
mark-recapture) efforts and long-term moni¬
toring if reliable population estimates are to
be obtained.
Previous studies indicate that Blanchard’s
cricket frogs prefer to breed in permanent
Volume 81 (1993)
83
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84
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JUNG: Blanchard’s cricket frogs in southwest Wisconsin
* Pairwise 2-tailed P < 0.05.
or semipermanent bodies of water (Pyburn
1958) with mudflats, shallow slopes toward
the water, and mud and vegetation banks
(Burkett 1984; Minton 1972). Most of the
40 sites studied, all of which were reported
to have had cricket frogs at one time, fit this
description.
Only one of the sites used by cricket frogs
in the past had been destroyed (drained).
However, this does not preclude the possi¬
bility that subtle site modifications (such as
aquatic vegetational changes) in the past may
have impacted cricket frog populations.
Blanchard’s cricket frogs were present at
sites with higher water temperatures com¬
pared to sites without cricket frogs, and
number of amphibian species was positively
correlated with water temperature. Sites with
higher water temperatures may be preferred
by anurans for breeding. On the other hand,
anurans may simply be more likely to be
observed on warm days. However, I do not
think this could account entirely for the re¬
lationships with temperature, in part because
most sites were visited more than once.
No sites contained pH levels considered
toxic to aquatic organisms (pH < 4.5 or >
9.5; Wetzel 1983), and I found no correla¬
tion between species richness and pH. Con¬
ductivity was within the range consid¬
ered normal for bicarbonate-dominated
lakes and streams (Wetzel 1983), but did
show considerable variability among sites
(coefficient of variation, CV = 30%). Oxi¬
dation-reduction potential (ORP) also var¬
ied (CV = 23%), but the mean value for all
sites was 136 ± 31.9 mV, which is substan¬
tially lower than 500 mV, a value expected
for neutral, fully oxygenated water equili¬
brated with air (Wetzel 1983). Turbidity
Volume 81 (1993)
85
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
was lower at sites with more species, which
may indicate preference for clearer waters.
Significantly fewer anuran species were
found at sites near corn and other crop
fields. This result suggests that fertilizer or
pesticide run-off from agricultural fields may
affect anuran populations. Herbicides and
insecticides still used on corn in Wisconsin
(Doersch et al. 1991) can be toxic to am¬
phibians and may have a significantly nega¬
tive impact on amphibian populations or
their prey base (atrazine, Hazelwood 1970;
Birge et al. 1980; malathion, paraquat, and
toxaphene, Sanders 1970; Hall and Kolbe
1980; Linder et al. 1990). Hylids in general,
such as cricket frogs, may be more suscep¬
tible to pesticides than other anuran species,
such as toads (Sanders 1970; Birge et al.
1979).
Other factors which may adversely affect
anurans in southwest Wisconsin are low dis¬
solved oxygen and high nitrogen concentra¬
tions. On the Little Platte River in southwest
Wisconsin, Holmstrom et al. (1988) re¬
ported dissolved oxygen concentrations as
low as 3.1 mg/1 in early July, which is below
the recommended 3 mg/1 dissolved 02 con¬
centration for housing amphibians (National
Academy of Sciences 1974). As well, nitrate/
nitrite and ammonia levels were as high as
3.1 mg/1 and 1.6 mg/1, respectively, during
the amphibian breeding season, which ex¬
ceed the 0.3 mg/1 nitrate/nitrite and 0.2 mg/
1 ammonia concentrations considered safe for
amphibians in the laboratory (National
Academy of Sciences 1974). On the other
hand, three of the sites included in the
present study were on the Little Platte River,
and all still had Blanchard’s cricket frogs.
Acknowledgments
I thank R. Hay, M. J. Mossman, B. Dhuey,
and C. Neumann from the Wisconsin De¬
partment of Natural Resources and G. S.
Casper from the Milwaukee Public Museum
for Blanchard’s cricket frog site records and
helpful discussion. Thanks also to S. M.
Conlin, T. Garland, Jr., C. T. Hannan, and
C. R. Vispo for field assistance, and to T.
Garland, Jr., R. Hay, and two anonymous
reviewers for improving the manuscript.
This research was sponsored by the Lois
Almon Fund and by the U.S. Department
of Energy, Office of Energy Research, En¬
vironmental Sciences Division, Office of
Health and Environmental Research, under
appointment to the Graduate Fellowships
for Global Change, administered by Oak
Ridge Institutes for Science and Education.
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J. E. Hudson. 1979. The effects of mercury
on reproduction of fish and amphibians. In
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1980. Effects of organic compounds on am¬
phibian reproduction. Lexington, KY: Uni¬
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WI: Wisconsin Department of Natural Re¬
sources, Bureau of Endangered Resources
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cricket frog, Acris crepitans. In Vertebrate ecol¬
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ed. R. A. Seigel, L. E. Hunt, J. L. Knight, L.
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The University of Kansas.
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locality data. Bull. Chic. Herpetol. Soc.
24:178.
Conant, R., and J. T. Collins. 1991. A field guide
to reptiles and amphibians. 3rd ed. Boston,
MA: Houghton Mifflin.
Doersch, R. E., J. D. Doll, R. A. Flashinski, R.
Grau, R. G. Harvey, J. L. Wedberg, G. L.
Worf. 1991. Corn pest management in Wis¬
consin. Madison, WI: University of Wiscon-
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Hall, R. J. and E. Kolbe. 1980. Bioconcentration
of organophosphorus pesticides to hazardous
levels by amphibians. J. Toxicol. Environ.
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Hazelwood, E. 1970. Frog pond contaminated.
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Holmstrom, B. K., P. A. Kammerer, Jr., and R.
M. Erickson. 1988. Water Resources Data
Wisconsin. Water Year 1988. Madison, WI:
U.S. Geological Survey Water-Data Report
WI-88-1.
Linder, G., J. Barbitta, and T. Kwaiser. 1990.
Short-term amphibian toxicity tests and
paraquat toxicity assessment. In Aquatic toxi¬
cology and risk assessment , Vol. 13, ed. W. G.
Landis and W. H. van der Schalie, 189-98.
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ing and Materials.
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Minton, S. A., Jr. 1972. Amphibians and rep¬
tiles of Indiana. Indiana Acad. Sci. Monogr.
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Wisconsin Frog and Toad Survey: establish¬
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WI: Wisconsin Department of Natural Re¬
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vey, 1984. Wisconsin Endangered Resources
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ton, D. C.: National Academy of Sciences.
Norusis, M. J. 1988. SPSS/PC+ Version 2.0 for
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Oldham, M. J. 1992. Declines in Blanchard’s
cricket frog in Ontario. In Declines in Cana¬
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tional monitoring strategy , Occasional Paper
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Ottawa, Ontario: Canadian Wildlife Service.
Pechmann, J. H. K., D. E. Scott, R. D.
Semlitsch, J. P. Caldwell, L. J. Vitt, and J.
W. Gibbons. 1991. Declining amphibian
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Robin E. Jung received her M.S. in Zoology in
1990 from the University of Wisconsin-Madison,
based on studies of individual variation in fruit
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111). She is currently working toward her Ph.D.
in the Department of Zoology, studying amphib¬
ian ecotoxicolgy.
Volume 81 (1993)
87
Richard A. Lillie, Robert Last, Paul Garrison,
Paul Rasmussen, and John W. Mason
A survey of the summer phytoplankton
communities of 579 Wisconsin lakes
Abstract Phytoplankton and associated limnological data were collected from surface
waters of a randomly stratified set of 579 Wisconsin lakes during the summer
of 1979. Frequency of occurrence and relative dominance of phytoplankton gen¬
era were determined from preserved samples.
Blue-greens were the most common and most frequently dominant taxa un¬
der all bloom conditions independent of sampling date (early to late summer).
The frequency of occurrence of blue-green dominance increased with bloom se¬
verity (chlorophyll a concentration). Blooms (defined as chlorophyll a concen¬
trations above 10 jig/L) occurred above a threshold of 20 pg/L total phospho¬
rus. Less than 7% of lakes that appeared blue or clear had any form of a bloom ,
while all lakes with severe or moderately severe blooms appeared either green,
brown, or turbid. Quantitative definitions are presented for classifying the se¬
verity of phytoplankton blooms in Wisconsin lakes based on relationships be¬
tween chlorophyll a and perceived water color.
Phytoplankton blooms cause taste and odor problems in
public water supplies and occasionally produce toxins that
may be harmful to humans or livestock (MacKenthun et al.
1945; Rohlich and Sarles 1949; Gorham 1965; Gilbert 1990).
Blooms are often comprised of algal species that are inedible,
unpalatable, or toxic to zooplankton, thus affecting zooplank¬
ton standing crop, productivity, or community size-structure.
Zooplankton, which utilize phytoplankton as a major food
source, are, in turn, important food resources for fish. There¬
fore, phytoplankton blooms may seriously interfere with the
efficient flow of nutrients and energy through the food chain.
Nighttime respiration or the sudden death and collapse of
blooms may cause total depletion of oxygen in the water col¬
umn, resulting in fish kills (Mackenthun et al. 1945, Barcia
1975). In addition to these biological responses, the changes
in water color that accompany the onset of a bloom (Lillie and
TRANSACTIONS Volume 81 (1993)
89
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Mason 1983) may directly influence lake
recreational use and affect lakeshore prop¬
erty values.
Direct control of phytoplankton blooms
in Wisconsin has been primarily limited to
the application of algicides containing cop¬
per. Many thousands of tons of copper sul¬
fate have been applied to Wisconsin lakes
since the early 1900s (Lueschow 1972).
Over 20 tons of copper sulfate were applied
to Lake Monona at Madison during 1933
alone (Domogolla 1933). The relative inef¬
ficiency of copper sulfate in controlling pe¬
lagic algae combined with environmental
concerns of copper toxicity has led the Wis¬
consin Department of Natural Resources to
ban whole-lake algae treatments (WDNR
1989) and has encouraged research and de¬
velopment of alternative management strat¬
egies to control blooms, including predator-
prey manipulations and nutrient reduction.
Blue-green algae have long been recog¬
nized as being a dominant component of
summer phytoplankton blooms in eutrophic
lakes (Reynolds and Walsby 1975; Wetzel
1975; Spodniewska 1986; Stewart and
Wetzel 1986; Canfield et al. 1989 among
many others). Numerous phytoplankton
studies (Sloey and Blum 1972; Bartell et al.
1978; Fallon and Brock 1980; Barko et al.
1984; Engel 1985; Narf 1985; Lathrop
1988; Barko et al. 1990; Klemer and Barko
1991) and general taxonomic surveys (Smith
1920; Smith 1924; Lackey 1945; Prescott
1970; USEPA 1978) of Wisconsin lakes
and reservoirs indicate the important con¬
tribution of blue-greens to blooms in Wis¬
consin. However, because much of the pre¬
vious work done in Wisconsin was restricted
to eutrophic bodies of water, there was some
concern that our perception of the problem
may be wrong. Indeed, not all blooms con¬
sist of blue-greens; diatoms, dinoflagellates,
and green algae were occasionally listed as
dominant in some of the aforementioned
studies. Consequently, in order to obtain a
more accurate assessment of the composition
of blooms on a statewide basis, we examined
phytoplankton from a set of samples that
had been collected in the summer of 1979
by the WDNR during an extensive limno¬
logical survey of 661 randomly selected Wis¬
consin lakes (Lillie and Mason 1983). Study
objectives were (1) to determine the struc¬
tural composition (occurrence and domi¬
nance) of summer phytoplankton commu¬
nities in Wisconsin lakes, (2) to establish the
relationship between bloom severity and
composition of phytoplankton communi¬
ties, and (3) to document the limnological
characteristics of lakes experiencing blooms.
Furthermore, because the lakes involved in
this study were sampled only once during
the summer, there was a legitimate concern
that normal seasonal phytoplankton succes¬
sion (per Stewart and Wetzel 1986; Bartell
et al. 1978; Dokulil and Skolaut 1991)
could invalidate our conclusions regarding
dominance during blooms (i.e. the compo¬
sition of blooms in lakes sampled early in
the summer may have differed from blooms
in lakes sampled later in the summer).
Therefore, we evaluated the relationships
between phytoplankton dominance and
bloom severity between lakes sampled early
in the summer versus lakes sampled late in
the summer. This paper summarizes the
conditions found during the summer 1979
survey and discusses the significance of these
findings to lake management.
Study Sites
A stratified random sub-sample was selected
from the approximately 2800 Wisconsin
lakes over 10 ha (25 acres) in size and greater
than 1.5 m (5 feet) deep. Lakes were strati¬
fied according to geographic distribution;
90
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LILLIE et al.: Survey of summer phytoplankton communities
Fig. 1 . Distribution of sampled lakes (N = 579) by water clarity (Secchi disc), total alka¬
linity, and total phosphorus, total nitrogen, and chlorophyll a concentrations, and per¬
ceived water color classification.
25% of all lakes present in each county were
randomly chosen. Phytoplankton samples
were available from only 579 of the 661
lakes in the survey. The sampled lakes rep¬
resented a broad range of morphometric
conditions and chemical compositions;
sampled lakes were nearly equally divided
between seepage (291) and drainage (287)
and between thermally stratified (262) and
mixed (256). Only 98 lakes were impound¬
ments. Most lakes generally had low nutri¬
ent (nitrogen and phosphorus) and chloro¬
phyll a concentrations and good water clar¬
ity (Fig. 1). Less than one third of the lakes
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were described as blue or clear. A more com¬
plete description of the limnological charac¬
teristics of the lakes in this study is provided
in Lillie and Mason (1983).
Methods
One sample was collected from each of the
379 lakes during the period 7 July - 7 Sep¬
tember, 1979. Samples were collected from
the lake surface above the point of maxi¬
mum depth (where known) or from a cen¬
tral basin location. All samples were pre¬
served immediately in Lugol’s solution.
Aliquots were settled using the Utermohl
settling technique (Lund et al. 1938) for a
minimum of 4 hrs per cm settling tube
height and analyzed at 1400 X and 560 X
magnification using a Wild inverted micro¬
scope. Taxa identifications (to genus and
species) were based on keys and descriptions
provided in Smith (1920, 1924, 1950),
Prescott (1970), and Weber (1971).
Dominance in algal communities may be
computed on the basis of numerical abun¬
dance, biomass, or biovolume. Because cell
sizes and chlorophyll content differ greatly
among the various phytoplankton species,
numerical dominance has less ecological sig¬
nificance than dominance based on either
biomass or biovolume. Because biomass was
not measured in this study, dominance of
phytoplankton communities was based on
estimates of relative biovolume. We devel¬
oped and applied a semi-quantitative ap¬
proach to evaluate cell biovolume dominance
because quantitative determinations of cell
biovolume of all 579 phytoplankton samples
were economically impractical. Assessment of
phytoplankton dominance was based on vi¬
sual comparisons of cell biovolumes in scans
of 40 randomly selected fields of view. Rat¬
ings of relative dominance and their respec¬
tive numerical weights were defined as fol-
Fig. 2. Flow chart describing criteria for
assessing phytoplankton dominance in
visual scans of 40 randomly selected
fields of view. Percentages (%)s refer to
typical percent of total biovolume in most
fields of view.
lows (see Fig. 2): absent (0) = not found; rare
(1) = occurring in less than 5 fields, or if oc¬
curring in more than 5 fields then never ac¬
counting for more than 10 percent of the
total biovolume in a field; present (3) = oc¬
curring in more than 5 fields but less than
10 fields and generally comprising 10-25
percent of the total biovolume in a field; im¬
portant (5) = occurring in more than 10
fields and generally comprising 10—25 per¬
cent of the total biovolume in a field (but
always secondary in rank to other taxa); co¬
dominant (7) = occurring in more than 5
fields and generally representing more than
25 percent of the total biovolume in a field
but sharing dominance with one or more
other taxa; and dominant (9) = occurring in
more than 5 fields and representing more
than 25 percent of the total biovolume and
clearly dominant over other taxa. Samples in
92
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LILLIE et al.: Survey of summer phytoplankton communities
which dominance could not be clearly deter¬
mined were identified as mixed assemblages.
Numerical weights (numbers given in paren¬
theses above) were averaged for all lakes
(statewide) and separately for only those lakes
in which the taxon occurred in determining
the relative importance of each genus.
The accuracy of this method of assessing
phytoplankton dominance was evaluated on
a subset of lakes prior to applying the
method to the entire set of samples. Inde¬
pendent assessments of cell biovolume domi¬
nance (by genus and class) were made by
four limnologists (with a combined 30 years
of experience) on a subset of 20 lakes selected
to represent a broad range of phytoplankton
communities. The 20 sets of assessments of
phytoplankton dominance (four replicates
for each lake) were compared with single as¬
sessments based on corresponding quantita¬
tive biovolume data. Cellular biovolumes
were computed from measurements of cell
dimensions taken from 10-23 randomly se¬
lected cells of each taxon in each sample.
Concurrence in ratings of the dominant taxa
was excellent among limnologists in samples
from lakes with blooms (100%) and only de¬
creased slightly (94%) in samples from lakes
with more diverse communities. Relative as¬
sessments of dominance agreed with quan¬
titative assessments of biovolume in 86 per¬
cent of the 80 determinations. The high de¬
gree of agreement justified the application of
the rapid semi-quantitative assessment to the
measurement of relative phytoplankton
dominance in all 379 samples. All analyses
were conducted by the same investigator (R.
Last) to minimize subjectivity.
Severity of phytoplankton blooms was
classed as mild, moderate, moderately severe,
or severe based on chlorophyll a concentra¬
tions above a threshold of 10 pg/L (Table
1). Breakpoints chosen to separate the four
categories were determined from linear re¬
gressions representing the interrelationships
among chlorophyll a concentrations, total
phosphorus concentrations, and water clar¬
ity measurements as reported by Lillie and
Mason (1983).
Limnological data collection procedures
and laboratory methods are detailed in Lillie
and Mason (1983). Chlorophyll data repre¬
sent trichromatic chlorophyll a (UNESCO
equation), uncorrected for pheophytin (0.45
um membrane fdters, homogenized 90%
acetone extract, modification of Strickland
and Parsons 1968). Total phosphorus con¬
centrations were measured using acid diges¬
tion-molybdate colorimetry (Eisenreich et al.
1975).
Data analysis was conducted using SAS
(SAS Institute, Inc. 1988). Values reported
represent means ± 1 standard error unless
otherwise stated. In order to evaluate
whether phytoplankton succession had a sig¬
nificant impact on our results (i.e., whether
phytoplankton dominance within specific
bloom categories was independent of sam¬
pling date), we compared dominance fre¬
quency plots of each phytoplankton group
for lakes sampled before and after August
8th (331 and 248 lakes, respectively). Lo¬
gistic regression was used to test for differ¬
ences in the relationship of phytoplankton
Table 1. Classification of phytoplankton
blooms based on chlorophyll a concen¬
trations.
Volume 81 (1993)
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Table 2. Taxonomic summary of algae by
order, family, and genera for 579 Wiscon¬
sin lakes.
dominance to chlorophyll concentration be¬
tween early and late sampling periods. Im¬
portance values (Appendix A) represent the
average of assigned weights given to relative
dominance ratings (i.e., 1-9).
Results
One hundred twelve phytoplankton genera,
representing 43 families and 19 orders, were
identified from the 379 lake samples (Table
2). A copy of the laboratory identification
sheet containing a complete list of genera
and species and their relative abundance for
each lake in the data set is available from the
authors (in microfiche) upon request. Green
algae (Chlorophyceae, including family
Desmidiaceae = desmids) and diatoms (Ba-
cillariophyceae) were represented by the
most genera. Only 17 blue-green (Cyan-
ophyceae) algal genera were recorded. Taxa
richness within individual lakes ranged from
2 to 24 genera. Most genera were uncom¬
mon: 16 genera were found in only one
lake, and 60 genera occurred in less than 3%
of the lakes (Appendix A). Only 20 genera
occurred in more than 25% of the lakes.
Cryptomonas , a cryptomonad (Crypto-
phyceae), was the most common alga, oc¬
curring in 80% of the lakes. Anahaena , a
filamentous blue-green alga, was the second
most common genus. Also found to be rela¬
tively common were two green alga genera,
Scenedesmus and Oocystis , a chrysophyte,
Dinobryon , another cryptomonad genus,
Chroomonas , two dinoflagellate genera,
Ceratium and Peridinium , a desmid genus,
Staurastrum , and another colonial blue-
green genus, Coelosphaerium. Fragilaria and
Melosira were the most common diatom
genera.
Collectively, blue-green algae were the
most frequently dominant taxonomic group,
with mixed assemblages, dinoflagellates, and
diatoms next in order of rank (Table 3). The
rank order based on the percentage of lakes
in which each of the various taxonomic
groups of algae were rated as at least impor¬
tant (includes important, co-dominant, and
dominant) did not differ substantially from
that of the dominant group. Seven of the 20
Table 3. Phytoplankton dominance by ma¬
jor taxa classification in all 579 Wisconsin
lakes. Numbers represent percent of lakes
in which each taxonomic group was domi¬
nant or at least important.
* includes lakes in which taxa were rated as
important, co-dominant, and dominant.
94
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LILLIE et al.: Survey of summer phytoplankton communities
* includes lakes in which genus was rated as important, co-dominant, and dominant,
t designates a tie.
most frequently dominant and important
genera were blue-greens (Table 4). Ana-
baena , Aphanizomenon , and Microcystis were
the most frequently dominant blue-greens;
Peridinium , Ceratium , and Glenodinium
were dominant dinoflagellates; and Melosira ,
Tabellaria , and Fragilaria were the most fre¬
quently dominant diatoms.
Composition of Blooms. Thirty-six percent
of the lakes in the survey had some form of
bloom present at the time of sample collec¬
tion (Table 5), and only 11% of the lakes
had severe or moderately severe blooms.
Sampling date (early versus late summer) did
not have a significant influence on phy¬
toplankton dominace. Logistic regression
indicated blue-green dominance increased (p
= 0.0001) with chlorophyll concentration
during both time periods. There was also an
interaction (p = 0.02) between chlorophyll
concentration and time period, because at
low chlorophyll levels, blue-green domi¬
nance was less frequent during the early than
the late sampling period, while at high chlo¬
rophyll levels blue-green dominance was
similar in both sampling periods. Neverthe¬
less, the overall trend for blue-green domi¬
nance to increase with chlorophyll concen¬
tration was present in both time periods.
Therefore, although the frequency of occur¬
rence of blooms was higher in lakes sampled
during late summer, the dominance struc¬
ture of blooms was not different from that
found in lakes sampled earlier in the sum¬
mer. Blue-greens were the most commonly
dominant group within all bloom categories
(Table 4), regardless of sampling date. The
Volume 81 (1993)
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Table 5. Phytoplankton dominance of major taxa relative to bloom condition (number of lakes
in each class shown in parentheses). Data represent % of lakes within each bloom category in
which taxa were dominant or co-dominant.
frequency of blue-green dominance in¬
creased from 37% during mild blooms to
70% during severe blooms (Fig. 3b). Di-
noflagellates were frequently dominant, and
chrysophytes were occasionally dominant
during mild and moderate blooms (Table 3),
but both declined in importance during se¬
vere blooms (Fig. 3g and 3c). Cryptophytes
tended to be more frequently dominant dur¬
ing severe blooms. Diatoms and greens
tended to be important or dominant during
moderately severe and severe blooms.
A core of 34 genera were commonly as¬
sociated with blooms. (Commonness here is
defined as those genera that were categorized
as present in at least 23% of the lakes or as
dominant in at least one lake experiencing
a bloom; see Table 6). Anabaena , Aphanizo-
menon , Aphanocapsa , Cryptomonas , Melosira ,
Oscillatoria , and Microcystis were common
dominants of severe and moderately severe
blooms. Aphanizomenon , Anabaena , and
Peridinium were common dominants of
mild and moderate blooms. While the fre¬
quency of occurrence or dominance of most
bloom genera did not appear to be directly
related to degree of bloom severity, some
genera, such as Anabaena , Aphanizomenon ,
Aphanocapsa , Aphanotheca , Melosira , Oscil¬
latoria :, Pediastrum , and Scenedesmus were
most often dominant during severe or mod¬
erately severe blooms. A few genera, includ¬
ing Peridinium , Synura , Tabellaria , and
Dinobryon , were more commonly dominant
during milder blooms.
Characteristics of lakes experiencing blooms.
Lakes experiencing phytoplankton blooms
tended to be larger and shallower than lakes
without blooms (Table 7). Lakes with
blooms also had shorter residence times and
had substantially higher nutrient concentra¬
tions and turbidities than lakes without
blooms. Of 71 lakes with total phosphorus
concentrations above 50 pg/L, 85% had
some form of bloom at the time of sampling
(54% severe or moderately severe). Only 9%
of the lakes with total phosphorus concen¬
trations less than 1 0 pg/L had a bloom (none
were severe or moderately severe). Lakes
with severe blooms had lower total nitrogen/
total phosphorus ratios (TN:TP) (low
TN:TP is an indication of possible nitrogen
limitation) than lakes without blooms.
96
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LILLIE et a I . : Survey of summer phytoplankton communities
Table 6. Frequency of occurrence of genera listed as dominant during blooms or occurring in
greater than 25% of lakes within a bloom class. Data represent percentages of lakes within
each bloom class in which genus was dominant (% frequency of occurrence based on pres¬
ence/absence shown in parentheses).
Genus _ Bloom Condition _
Non-bloom Mild Moderate Moder. -Severe Severe Bloom
Perceived water color was dramatically
impacted by blooms (Table 8). None of the
64 lakes with severe or moderately severe
blooms appeared blue or clear. Among lakes
with mild or moderate blooms, 30% ap¬
peared green, 48% appeared brown, 7% ap¬
peared as a mixture of colors, 7% were tur¬
bid, and only 8% were clear or blue in ap¬
pearance. While lakes without blooms also
were frequently classed as colored (13%
green, 33% brown, 3% mixed, and 2% tur¬
bid), 93% of the 176 lakes identified as blue
or clear did not have blooms.
Discussion
Blue-green algae were dominant or co-domi¬
nant in one-third to two-thirds of all lakes
Volume 81 (1993)
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0-10 10-15 15-30 30-50
0-10 10-15 15-30 30-50 >50
DINOFLAGELLATES
SEVERE
0-10 10-15 15-30 30-50 >50
CHLOROPHYLL a (ug/L)
CHLOROPHYLL a (ufl/L)
Figure 3. Relative dominance and importance of major taxa expressed as percentages
of lakes within each bloom category that taxa were dominant (o) or important (A). Per¬
cent important also includes lakes in which taxa were dominant or co-dominant.
98
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LILLIE et a I. : Survey of summer phytoplankton communities
Table 7. Limnological conditions associated with phytoplankton blooms in Wisconsin lakes.
Data represent means ± 1 SE.
Bloom Condition
* excludes 1 lake with very high chlorophyll concentration.
** from Lillie and Mason 1983.
p = partial data, information not available for all lakes.
with blooms and in almost one-quarter of
lakes without blooms. Collectively, blue-
green genera were dominant among all cat¬
egories of surface blooms. Blue-green domi¬
nance increased with the degree of bloom se¬
verity as measured by chlorophyll a concen¬
tration.
Although the frequency of blooms was
higher in late summer, the dominance struc¬
ture of phytoplankton communities did not
differ substantially within bloom categories
from early to late summer. Therefore, inter¬
pretation of relative phytoplankton domi¬
nance within specific bloom categories was
not biased by the 63-day time period during
which all 379 lakes were sampled. While the
severity of blooms in many of the lakes
sampled during the first half of the sum¬
mer may have been higher had they been
sampled during the later half of the sum¬
mer sampling period, the composition of
the blooms probably did not change sig¬
nificantly (at the group level). Such a con¬
clusion is not altogether unexpected, as
Bartell et al. (1978) reported relative con¬
stancy in species assemblages during the
summer in extensive studies in Lake
Wingra (Dane Co.). The frequency of oc¬
currence of blooms and severity of blooms
shown in Table 5 undoubtedly would have
been higher if all lakes had been sampled
during the later half of the sampling pe-
Volume 81 (1993)
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Table 8. Association between perceived color (visual appearance) and algal blooms in
579 Wisconsin lakes. Data represent numbers of lakes in each category.
* some field observers did not differentiate between clear-green, green and turbid, or green and brown
mixed.
riod. This, however, does not negate our
conclusions regarding compositions of
blooms.
These findings support conclusions re¬
garding phytoplankton dominance derived
from earlier cited studies of primarily
eutrophic Wisconsin lakes. Blue-green algae
are the single most important and dominant
taxonomic group of algae in most Wiscon¬
sin lakes during summer months regardless
of trophic state and, as such, should be tar¬
geted for management control.
The biomass of blue-greens in lakes has
been shown to be directly correlated with
TN:TP ratios under fixed light conditions
(Smith 1986). No significant relationship
was detected between TN:TP and the oc¬
currence of blooms in our study; however,
the ratio between mean TN and mean TP
was lower in lakes with severe blooms than
in lakes without blooms (Table 7). The pro¬
gressive increase in frequency of occurrence
of blue-green dominance with each level of
bloom severity corresponds with the progres¬
sive decrease in TN:TP and agrees with
Smith’s (1983) findings. Furthermore, our
data directly contradicts that of Canfield et
al. (1989), who reported a decrease in fre¬
quency of blue-greens at TN:TP < 29. Light,
temperature, inorganic nutrients, and zoo¬
plankton grazing can influence the relation¬
ship between blue-green dominance and
TN:TP within individual lakes (McQueen
and Lean 1987; Smith 1986; Spencer and
King 1987).
Blooms were more common in large,
shallow reservoirs or drainage lakes, in ac¬
cordance with findings of other studies (Fee
1979). The greater internal recycling of nu¬
trients, availability of sunlight, and thermal
homogeneity of these systems provide a
more optimum growth medium for blue-
greens than that offered by deeper, thermally
stratified lakes.
Limnological characteristics of the 379
lakes for which phytoplankton samples were
available did not differ significantly from the
characteristics of the entire 66 1 lakes in the
random survey (Table 7). Therefore, find¬
ings regarding phytoplankton dominance
reported in this survey and recommenda¬
tions based on these data should be appli¬
cable to all Wisconsin lakes of similar size
and depth. Based on the relationships be¬
tween bloom severity and nutrient concen¬
trations, as shown in Table 7, we propose
the following in-lake phosphorus concentra¬
tions be established as thresholds for con¬
trolling blooms in Wisconsin lakes. An in¬
lake summer phosphorus concentration of
1 00
TRANSACTIONS
LILLIE et al.: Survey of summer phytoplankton communities
20 pg/L is an appropriate level to assure non¬
bloom conditions, while 30 pg/L TP is a
more appropriate value to define the thresh¬
old between mild and moderate blooms.
The former value corresponds with estab¬
lished spring phosphorus standards for
southeastern Wisconsin lakes (SWRPC
1979) and with the results of the National
Eutrophication Survey, which indicated that
blooms did not occur in eastern United
States lakes with less than 19 pg/L mean to¬
tal phosphorus (Williams et al. 1977). Re¬
duction of in-lake phosphorus concentra¬
tions below a given threshold value does not
guarantee that a bloom will not occur
(Welch 1989), as some lakes with phospho¬
rus concentrations below the threshold will
have blooms and some lakes with phospho¬
rus concentrations above the threshold will
not have blooms. Other factors, including
differences in zooplankton grazing pressures,
temperature, toxins, and other growth-lim¬
iting elements may interfere with the rela¬
tionship. Threshold values may need to be
adjusted according to a lake’s geographic lo¬
cation to account for differences in phy¬
toplankton dominance and response to nu¬
trients that may occur among ecoregions (see
Heiskary et al. 1987). Irrespective of the cir¬
cumstances, reducing in-lake phosphorus
concentrations will certainly decrease the
likelihood that a bloom will occur or will re¬
duce its intensity and duration.
Welch (1989) proposed that the fraction
of total algae comprised of blue-greens is a
sensitive indicator of nutrient concentrations
and, thus, would make a good alternative
eutrophication index were it not for the lack
of agreed-upon trophic state threshold val¬
ues. Based on the data collected in this
study, a value of 23% blue-greens (by
biovolume) may be an appropriate thresh¬
old criterion. Blue-greens were dominant
(represented more than 23% of the total
biovolume) in more than 50% of the lakes
where chlorophyll a concentrations exceeded
30 pg/L. The 30 pg/L chlorophyll a concen¬
tration corresponds with our bloom classes
of moderately severe and severe and matches
the criterion for blooms established by
Walker (1985).
Perceived water color (or visual appear¬
ance) has particular significance to lake
managers in indicating blooms. In most
cases, lakes that appear blue or clear do not
have a bloom. Conversely, a green appear¬
ance to a lake does not necessarily signify a
bloom as some lakes have a natural clear-
green appearance. Likewise, while a brown
appearance often corresponds with a diatom
bloom, many lakes have a brown or yellow-
brown stained appearance due to high con¬
centrations of organic acids. However, if a
lake’s appearance changes from blue/clear to
green, brown, or turbid, there is a very good
probability that a phytoplankton bloom is
occurring and that chlorophyll a concentra¬
tions exceed 10 pg/L. These color observa¬
tions may not apply to other regions, but
similar relationships may be established
through direct observations. As such, gen¬
eral observations of water color can have a
very important role in monitoring the
trophic condition of a lake. Historical ob¬
servations of blue or clear water color may
be safely regarded as being indicative of good
water quality (chlorophyll less than 10 pg/L).
Lastly, the results of this survey demon¬
strate the utility of rapid subjective analysis
of summer phytoplankton community com¬
positions (in conjunction with other water
quality data) as an informative monitoring
tool in assessing water quality. Therefore,
monitoring phytoplankton community
compositions may be useful in measuring
the effectiveness of agency-mandated nutri¬
ent control programs and lake restoration
efforts.
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Appendix A. Frequency of occurrence (as %), number of occurrences by relative dominance
classification, and relative importance values for all phytoplankton genera found in 579 Wisconsin lakes
during the summer of 1979.
1 02
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LILLIE et al.: Survey of summer phytoplankton communities
**taxa believed to be primarily epiphytic and only incidently found in the plankton (i.e. tychoplankton).
Taxon codes: BACI=Bacillariophyceae, CHLO=Chlorophyceae, CHRY=Chrysophyceae,
CRYP=Cryptophyceae, CYAN=Cyanophyceae, DESM=Desrmidiaceae, DINO=Dinophyceae,
EUGL=Euglenophyceae, XANT=Xanthophyceae.
Volume 81 (1993)
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Acknowledgments
This study was conducted by staff of the Bu¬
reau of Research, Water Resources Research
Section, with the assistance of numerous in¬
dividuals who were acknowledged in Lillie
and Mason 1983. We wish again to express
our thanks to those individuals who con¬
ducted the field collections and completed
the laboratory analyses. Co-author Robert
Last diligently performed taxa identifications
and determined relative dominance on all
379 samples over a period of several years
and deserves most of the credit for seeing
this project through to the end. We also
thank I. Marx-Olson for assistance in com¬
puter data entry and D. Knauer for critical
review of the manuscript. Thanks are also
extended to W. Sloey, an anonymous re¬
viewer, and the Editor for their many use¬
ful suggestions.
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Lackey, J. B. 1945. Plankton productivity of cer¬
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Lathrop, R. C. 1988. Evaluation of whole-lake
104
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LILLIE et a I . : Survey of summer phytoplankton communities
nitrogen fertilization for controlling blue-
green algal blooms in a hypereutrophic lake.
Can. J. Fish. Aquat. Sci. 45:2061-2075.
Lillie, R. A., and J. W. Mason. 1983. Limno¬
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Wis. Dep. Nat. Resourc. Tech. Bull. No.
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Lueschow, L. A. 1972. Biology and control of
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Lund, J. W. G., C. Kipling, and E. D. LeCren.
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McQueen, D. J., and D. R. S. Lean. 1987. In¬
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Narf, R. P. 1985. Impact of phosphorus reduc¬
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
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Richard Lillie, Paul Garrison, John Mason ( re¬
tired) are Research Scientists with the Wisconsin
Department of Natural Resources, Bureau of Re¬
search, Water Resources Research Section, Monona,
Wisconsin.
Paul Rasmussen is a Biometrician with WDNR
Bureau of Research, Monona, Wisconsin.
Robert Last was a Natural Resources Specialist
with the Water Resources Research Section.
1 06
Charles A. Long and John E. Long
Discriminant analysis of geographic
variation in long-tailed deer mice from
northern Wisconsin and Upper Michigan
Abstract Identification of morphologically similar long-tailed mice (Peromyscus
spp.) from the forests of northern Wisconsin and Upper Michigan was
clarified by application of statistical analysis. Use of student t-tests,
bivariate plots. Dice squares, analysis of variance, and a stepwise discrimi¬
nant analysis revealed three distinctive populations. Previously, mice from
this region were identified asY . leucopus orY . maniculatus gracilis, and
neither was very clearly distinctive fom the other. The best characters used
to identify these mice were ear length, length of rostrum, and length of tail.
Other characters varied significantly fom group to group, but incisive
foramina length, cranial depth and cranial breadth (previously given
heavy weight by taxonomists) were the least reliable. Discriminant
coefficients were developed that may be used in future studies classifying
approximately 2,000 unidentified museum specimens, based upon the
canonical functions that segregate the three groups best. Predictability
based on probability of best fit, and next best fit, ranged fom 82 to 90
percent. The deer mice fom three isles in Lake Michigan closely resembled
typical P. maniculatus maniculatus from Labrador even more than they
resembled the northern Lake Superior mice formerly assigned to P. m.
gracilis. Northern specimens approach maniculatus but the southernmost
long-tailed deer mice are smaller. Six recently collected specimens fom
Washington Island, in Lake Michigan, resembleY . leucopus fom central
Wisconsin and are the first record of Y. leucopus on any island in Lake
Michigan.
Aside from the confusing and well-known resemblance of
the forest deer mouse ( Peromyscus maniculatus gracilis ) to
the common white-footed mouse {Peromyscus leucopus ), Long
(1978) noted slight clinal variation in P. m. gracilis from north
to south and documented differences (particularly in longer
TRANSACTIONS Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
rostrum) in P. m. gracilis on St. Martin
Island, Michigan. Cursory examination of
P. m. gracilis specimens from the Apostle
Islands revealed cranial differences from many
Wisconsin gracilis , especially the narrower
and shorter rostrum of the Apostle Islands
gracilis. This form resembled Peromyscus
maniculatus maniculatus from Ontario, as
figured by Osgood ( 1 909) in his classic taxo¬
nomic revision of Peromyscus. The difficulty
in segregating two or perhaps three similar
deer mice has hindered a proper classifica¬
tion (at the University of Wisconsin-Stevens
Point [UWSP] Museum of Natural His¬
tory) of nearly 2,000 specimens from Wis¬
consin and some from the Upper Peninsula
of Michigan. Over the last decade, students
who have regularly collected^raa/^ on Wash¬
ington Island began to catch some specimens
that appeared to be P. leucopus , hitherto
unknown on any island in Lake Michigan.
In southern Wisconsin, Stromberg (1979)
used discriminant analysis to distinguish P.
leucopus from P. m. bairdii , a small, short¬
tailed prairie form inhabiting sandy soils of
southern and western Wisconsin. However,
these two species are easily identified even as
young animals (Long 1968). Greater diffi¬
culty is encountered in separating P. m. gra¬
cilis from the Canadian race P. m. manicula¬
tus, and both of them from P. leucopus.
Why is it that in the past two subspecies of
long-tailed deer mice (P. maniculatus) have
been recognized in Michigan but only one
has been noted in Wisconsin? Baker (1983)
followed Burt’s ( 1 948) earlier classification of
Michigan mammals, and Burt, without com¬
ment, had followed Osgood’s (1909) mono¬
graphic revision of Peromyscus. Osgood re¬
ported P. m. maniculatus from Isle Royale
and P. m. gracilis from both Upper and Lower
Michigan. However, for nearby Wisconsin
mice, Osgood (1909) and Jackson (1961)
used only gracilis , and Minnesota workers,
without comment, used gracilis there (Gun¬
derson and Beer 1933; Hazard 1982) . We, on
the other hand, expected that P. m. manicu¬
latus ranged southward into Wisconsin.
The only samples available to Osgood
from northern Michigan and Wisconsin were
a large series (N=55) from Isle Royale (this
island lies very near the Ontario shore of
southern Canada, and these mice seem refer¬
able to Canadian maniculatus ) and a few
specimens from all of Michigan (13) and
Wisconsin (7). As classic and extensive as
Osgood’s (1909) revision was, his samples
were inadequate to classify and map the
detailed geographic distributions of these
closely similar mice in this region. In our
study we analyzed variation by rigorous sta¬
tistical tests of the long-tailed kinds present
in northern Wisconsin, the Upper Peninsula
of Michigan, and islands in Lake Superior
and Lake Michigan.
Taxonomic Characters
Typical P. leucopus may be distinguished
from P. m. gracilis by several visible charac¬
ters if one compares a series of specimens of
one mouse or the other, or if any given
individual possesses several trenchant char¬
acters. But if the collection is mixed — or if a
specimen is unusual or, as is more often the
case, subadult — then correct identification
may be impossible. Mismeasured specimens,
skins lacking skulls, or skulls lacking skins
complicate taxonomic identification.
The white-footed mouse ( P . leucopus) is
most recognizable by a short tail (both actual
and relative to body length) that is sparsely
haired. Vestigia of ancient reptilian scales are
commonly visible on it, and the sparse hairs
seem dingy. In those specimens having such
tails, the tail is not sharply bicolor nor does
the tip ever bear a pencillate tuft. In those P.
leucopus having bright-colored and well-
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TRANSACTIONS
LONG and LONG: Long-tailed deer mice of Wisconsin and Michigan
haired tails, the short length helps in identi¬
fication.
Peromyscus maniculatus gracilis, originally
described by its elongate tail and only tenta¬
tively ascribed to “Michigan,” has enormous
ears, prominent and elongated whiskers
(vibrissae), and a long snout. P. m. mani -
culatus , the nominate race of deer mouse
ranging across much of eastern Canada, and
reported by Osgood from Isle Royale in Lake
Superior, resembles gracilis in long, pencil-
late tail and long ears. Reportedly it is darker
in color. Owing to hybridization in southern
Ontario, any distinctive features there in P.
m. maniculatus reportedly are merged some¬
what with P. m. gracilis (Osgood 1909).
Peromyscus maniculatus maniculatus has
never been documented in Wisconsin, nor
has anyone since Osgood bothered to inves¬
tigate that possibility. However, skulls from
the Apostle Islands and from nearby Drum¬
mond, Wisconsin, appreciably differ from
those of P. m. gracilis from the Lake Michi¬
gan Isles (Washington Island, St. Martin
Island, and Rock Island, Long 1978). We
considered that the former sample might be
referable instead to nearby Canadian P. m.
maniculatus. The shape of their rostra, in¬
deed, resembles the figured rostrum chosen
for nominate P. m. maniculatus by Osgood
(1909).
Osgood (1909) selected representative
skulls to illustrate qualitative characters of
form. In P. leucopus the rostrum is short and
rather pinched anteriorly, and as a conse¬
quence the incisive foramina (Fig. 1) on the
hard palate are likewise constricted anteri¬
orly. In either P. m. maniculatus or P. m.
gracilis , or in their intergrades, the rostrum is
longer and the incisive foramina more nearly
parallel, i.e., straight sided. In P. m. gracilis ,
the rostrum is supposed to be quite elongate
and not so narrow as in P. leucopus or P. m.
maniculatus.
Fig. 1. Cranial measurements of Pero¬
myscus. RL, length of rostrum; TL, total
length or greatest length of skull; i, length
of incisive foramina. The cranial width is
measured across and including the
convex bulges of the cranium immediately
posterior to the zygomatic arches. The
cranial depth is measured from the top of
the braincase to its base, between (not
including) the auditory bullae.
Rostral shape and incisive foramen length
have been used by Hazard (1982) to distin¬
guish P. m. gracilis from P. leucopus. Overlap
of these characters between samples was con¬
siderable in Wisconsin. Burt ( 1 948) reported
the infraorbital canal as distinctive in form in
Michigan P. m. gracilis , but in Wisconsin no
difference in this character is apparent in
gracilis and leucopus (Jackson 1961; Long
1974). Choate (1973) found rostral breadth
useful in distinguishing these species, but in
our series there was no significant difference
in rostral breadth between gracilis and Por¬
tage County specimens of leucopus.
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Methods
In this article, the following terms refer to the
geographical areas where the large samples of
specimens were taken:
Lake Michigan Isles — Washington Island,
Rock Island, and St. Martin Island, all
found in Lake Michigan near the north¬
eastern Wisconsin shore.
Central Wisconsin — Portage County
Lake Superior — Outer Island and Stock-
ton Island (both members of the Apostle
Islands in Lake Superior near the northern
Wisconsin shore); and Drummond on the
mainland in northern Wisconsin.
Measurements used in this study included
the standard measurements of skin labels, of
which only the length of ear and tail proved
useful. Cranial measurements (Fig. 1) in¬
cluded total length of skull; length of rostrum
measured from the tips of the nasals to the
slight constriction anterior to the zygoma;
length of the incisive foramina; and in some
samples cranial width, measured immedi¬
ately posterior to the zygomata, and cranial
depth (not including the auditory bullae) . C.
A. Long made all the cranial measurements.
No size differences were noted between males
and females. Young animals were excluded.
These were recognized by juvenal pelage,
small weak skulls, unworn upper molars, and
an open basioccipital-basisphenoid suture.
Most of the specimens were from Wiscon¬
sin and Upper Michigan and are preserved in
the UWSP Museum of Natural History.
Eight specimens of typical P. m. maniculatus
from Labrador were borrowed from the
United States National Museum of Natural
History (USNM). Also borrowed were Jack¬
son’s (1961) two specimens of P. m. gracilis
caught in the Sheboygan Marsh in southern
Wisconsin, a population disjunct from the
geographic range of northern gracilis.
Skulls appreciably larger from the Lake
Michigan Isles were compared (student t-
tests of cranial width) with skulls of speci¬
mens from Outer Island, Lake Superior, and
from Stockton Island in Lake Superior and
nearby Drummond on mainland Wisconsin.
Means of cranial width were calculated to
compare the samples from Outer Island with
those from Stockton-Drummond, as well as
to compare Stockton mice with Drummond
mice. These comparisons were made to con¬
solidate the Lake Superior populations as a
homogeneous group, even though the group
includes insular and mainland populations.
Means and standard deviations were calcu¬
lated and compared from the Lake Superior
population, the Michigan Isles population,
numerous P. leucopus from Portage County
in central Wisconsin, and other appropriate
populations to study geographic variation
along a generally northwest to southeast di¬
mension or transect.
These three groups — the Lake Superior
group, the Lake Michigan Isles group, and
central Wisconsin leucopus — were analyzed
by stepwise discriminant function analysis,
using the aforementioned five linear vari¬
ables (length of ear, tail, skull, rostrum, and
incisive foramina) and ordering the char¬
acters by their apparent usefulness (F values)
in identification. All specimens that did not
fit well within their groups were flagged.
Allocation to a group was determined by the
probability of best fit, as well as by examin¬
ing the specimens that showed the second
highest probability for fit. Scatter plots were
mapped using two discriminant functions to
segregate the three groups. The data were
evaluated to determine the percentage that
fit in their appropriate groups, and discrimi¬
nant coefficients were calculated for use in
future testing. The three groups were used as
standards against which some small samples
were compared. One such sample reported
1 1 O
TRANSACTIONS
LONG and LONG: Long-tailed deer mice of Wisconsin and Michigan
herein was a collection of six mice that were
tentatively identified in our study as P.
leucopus, previously unknown in the fauna of
Washington Island, or on any island in Lake
Michigan or Lake Superior. The USNM
series of typical P. m. maniculatus from La¬
brador was assigned to one of the three
groups by use of the Fisher coefficients.
Results and Discussion
Measurements are given in Table 1 for the
three large populations or groups analyzed
(Lake Superior, Michigan Isles, and central
Wisconsin) . Although there is overlap in most
of the measurements, P. leucopus can be iden¬
tified usually by smaller dimensions, except
for length of skull and rostral width. When
length of the ear is plotted against length of
tail (Fig. 2), there is clear separation of the
Lake Michigan Isles mice from P. leucopus.
Some collections of Peromyscus from Wash¬
ington Island made in recent years scatter
across the graph, and some specimens sus¬
pected to be P. leucopus indeed fit among the
leucopus (central Wisconsin group).
The same results were found by plotting
incisive foramina length against length of
rostrum (Fig. 3). In this comparison numer¬
ous recently taken specimens fit among the
leucopus specimens; none of them fit in the
gracilis camp.
Transects of mice northwest to southeast
(Figs. 4-7) reveal overlap of all characters
chosen, except that the Lake Michigan Isles
gracilis tended toward high values and dif¬
fered most from P. leucopus. (Cranial width
was not measured in leucopus , Fig. 7.)
In a t-test the mice from the Lake Michi¬
gan Isles were broader across the cranium
than those in the Lake Superior group ( 12.32
± .43 versus 11.93 ± .24, p < 0.001). Five
specimens (of 22) in the Lake Michigan
Isles group had open sutures (indicating su¬
badult age), but their skulls were broader all
the same. We expected the Lake Superior
group to have the wider skulls. The two
populations did not differ in total length of
skull (26.3 versus 25.78 mm, p < 0.001) or
tail length (83.1 versus 83.6, p < 0.001).
Lake Michigan Isles mice also had wider
skulls than those mice combined from Stock-
ton Island and Drummond, but the latter
two samples were significantly wider than
the mice from nearby Outer Island. (The
Stockton Island and Drummond mice did
not differ significantly from one another.)
Therefore, in cranial width the Lake Supe¬
rior group was not strictly homogeneous.
Concerning cranial breadth or width, our
study only proves that mice on Outer Island
are narrow (though resembling our museum
specimens from southern Ontario; see Fig.
7). Osgood’s own figures and measurements
do not confirm his claim that P. m. mani¬
culatus has the wider skull, even though his
representative specimens were collected in
the eastern part of the range far from the zone
of intergradation of that race with P. m.
gracilis. The Labrador series we examined
averaged 12.3 in the seven adults, which is
but slightly wider than in Lake Superior
group specimens (Table 1). We found no
significant difference in tail length between
the groups from Lake Superior and Lake
Michigan (Table 1), or between Wisconsin
deer mice and the Michigan maniculatus
reported by Baker (1983).
In the analysis of variance, all five mea¬
sured variables showed geographic variation,
so that in the total analysis all three Wiscon¬
sin populations seemed distinct (Table 2).
Of the variables, ear length gave the high¬
est F -value among the three groups (Table 2) ,
but this field measurement is often slightly
made in error. Ffazard (1982) considered
smaller ears as characteristic of P. m. mani¬
culatus. However, the Labrador series of
Volume 81 (1993)
1 1 1
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
maniculatus averaged 19.7 mm. Short ear
length statistically separates P. leucopus from
the other two groups. Length of rostrum
separates all three populations evenly. Small
incisive foramina, short skull, and shorter tail
separate P. leucopus out fairly well, except for
the considerable overlap in all these variable
traits (Table 1).
In the stepwise discriminant analysis, the
variables were removed in the order listed in
Table 2. Wilk’s lambda varied from 0.54 to
0.31, all highly significant in length of ear
and the other variables through incisive fo¬
ramina.
The success in prediction of specimens for
all the three groups (Fig. 8) exceeded 80
percent. Of 60 specimens from the Lake
Michigan Isles, only 12 fit outside the group
1 sector, and eight fit better in the Lake
Superior sector. Of 30 leucopus from central
Wisconsin, 27 clumped together, and the
remainder fit with the Lake Superior group.
Of 20 specimens from the Lake Superior
group, 15 clumped together. Three fit with
the Lake Michigan specimens and two with
central Wisconsin leucopus.
The Lake Michigan Isles mice had a pre¬
dicted membership of 82 percent. Peromys-
cus leucopus , which is so difficult to identify
by conventional characters, predicted very
well with 90 percent strictly within its group.
Most of the specimens that did not fit their
appropriate group with highest priority did
so as the jmWhighest probability. Of the 1 2
in the Lake Michigan Isles group, six fit as a
second probability. Of the three central Wis¬
consin mice, two fit as a second probability.
The six specimens tentatively identified in
recent years as P. leucopus from Washington
Island (hitherto unknown on any islands in
Lake Michigan) identified closely with the
central Wisconsin group ( P . leucopus ) (Fig.
9). A small group consisting of Washington
Island mice not tentatively identified as P.
leucopus but from the same recently obtained
field collections (and including a few sub¬
adults) segregated poorly. The bivariate ana¬
lysis (Figs. 2- 3) also showed much overlap of
characters in these recently collected speci¬
mens.
Fisher’s Linear Discriminant Functions,
useful as classification coefficients, are given
in Table 3. The individual’s measures are
multiplied by the coefficients to weight them
for taxonomic discrimination. Use of Fisher’s
coefficients surprisingly classified the two dis¬
junct Sheboygan Marsh specimens with the
Lake Superior deer mice, although the two
skulls were so small and the ears so short they
closely resemble P. leucopus (which now seems
to be the only Peromyscus present in the She¬
boygan Marsh). One of the two is not quite
adult (USNM 227357) . These two small skulls
differ markedly from deer mice skulls of Lake
Michigan Isles, geographically much nearer
but isolated by water on islands of Lake Michi¬
gan. They are practically indistinguishable
from a large sample recently examined from
Lower Michigan (Long, unpublished).
The big surprise was the close resemblance
of the Lake Michigan Isles group to the
typical P. m. maniculatus from far away La¬
brador, rather than to mice nearer at Lake
Superior. The resemblance was also close in
color of the pelage. The Lake Superior group
differed from the Labrador P. m. maniculatus ,
which was unexpected because mice from
Ontario and nearby Isle Royale are referred
to as P. m. maniculatus. All seven adults in the
Labrador series fit with the Lake Michigan
Isles group. They resembled P. leucopus least,
which, of course, was to be expected.
Discriminant analysis weights the charac¬
ters in the optimal way to allow the groups to
segregate. Differences may or may not reflect
speciation, and they must be perceived with
caution. However, in Wisconsin, obvious
but overlapping differences in the three
1 1 2
TRANSACTIONS
LONG and LONG: Long-tailed deer mice of Wisconsin and Michigan
groups justified the analysis. If the variation
is spurious or drift-like, it is of interest all the
same, because the resemblance to Labrador
mice, which is very close, suggests that gracilis
is a synonum of P. m. maniculatus. Such
microgeographic variation may indeed be
part of the problem (Ledehrle et al. 1985).
On the other hand, the resemblance of Lake
Michigan and Labrador mice may arise from
effects of Pleistocene and early Holocene
climate on the zoogeography of Peromyscus
maniculatus.
The Canadian race P. m. maniculatus for¬
merly may have been widespread, with some
populations colonizing islands. Peripheral
mainland populations may have evolved new
characters, such as small size, leading per¬
haps to the geographic variation named gra¬
cilis by Osgood (1909) or described by us by
using Osgood’s name. Additional studies on
electrophoretic variation in proteins, such
as the work by Calhoun and Greenbaum
(1991), may clarify the observed differentia¬
tion.
LENGTH TAIL
Fig. 2. Bivariate plot of length of ear and length of tail.
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
a Rock and St. #
^ Martin Is. gracilis.
i ^ Wash. Is. gracilis r\
1975. ^
Wash. Is. gracilis
recent years.
Portage Co.
leucopus.
(D Wash. Is. "leucopus"
recent years.
Fig. 3. Bivariate plot of incisive foramina length and length of rostrum.
1 14
TRANSACTIONS
LONG and LONG: Long-tailed deer mice of Wisconsin and Michigan
Fig. 4. Transect of deer mice localities based on rostral length, northwest to southeast.
Modified Dice-squares (mean, the horizontal; N, sample size above the l-shaped vertical
for observed range; standard deviation, the boldfaced l-shaped vertical; twice the
standard error, the short thick, vertical. A, specimens from southwest Ontario; B, Outer
Island, Apostle Islands, Wisconsin; 0, Stockton Island, Apostle Islands; D, Drummond,
Wisconsin; E, Lake Michigan: St. Martin Island, Rock Island, Washington Island (early
years); F, Possible P. leucopus from Washington Island; G, P. leucopus from Portage
County, Wisconsin.
Fig. 5. Transect of deer mice localities based on total length of skull (see Fig. 4 for
explanation).
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 6. Transect of deer mice localities based on length of incisive foramina (see Fig. 4 for
explanation).
Fig. 7. Transect of deer mice localities based on cranial width (see Fig. 4 for explanation).
1 16
TRANSACTIONS
LONG and LONG: Long-tailed deer mice of Wisconsin and Michigan
LONGER ROSTRUM, EAR, ETC.
CANONICAL DISCBINIMANT FUNCTION 1
OUT -6.0 -4.0 -2.0 .0 2.0 4.0 6.0 OUT
Fig. 8. Scatter plots of Peromyscus groups 1-3.
1 = Lake Michigan Isles
2 = Central Wisconsin ( leucopus )
3 = Lake Superior
Values are canonical variates of discriminant functions. Asterisks are centroids.
Volume 81 (1993)
1 1 7
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
OUT
X-
OUT X
6.0
*6.0
•4.0
CANONICAL DISCRIMINANT FUNCTION 1
•2.0 .0 2.0 4.0
. . . . * . +-
6.0
OUT
•X
X
4.0
2.0
•2.0
-4.0
-6.0
OUT
X-
OUT
2
5
5 2
2
2 22 22
2 2 2 2
22*2 52
2 225222
2 52
2
2
2
-6.0
-4.0
-2.0
2.0
4.0
6.0
-X
OUT
Fig. 9. Scatter plot for P. leucopus. Values are canonical variates, P. leucopus of Portage
County, Wisconsin, comprising group 2. The six 5s were tentatively identified as P
leucopus from Washington Island, and this grouping confirms the identity.
1 1 8
TRANSACTIONS
Table 1. Mean lengths ± standard deviation of Peromyscus specimens from Wisconsin and Michigan
o -i;
O
co
a.
1 19
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 2. Analysis of variance of five measure¬
ments of Peromyscus spp.
Linear measurement F
* significant at p< 0.05
Acknowledgments
We wish to thank Drs. David Hillier and T.
A. Neuhauser for helping with the comput¬
ing. John Yost helped C. A. Long arrange
specimens of deer mice. The University of
Wisconsin-Stevens Point supported C. A.
Long’s sabbatical leave enjoyed at St. Olaf
College. Dr. Eric M. Anderson and Adrian
Wydeven gave appreciated advice on revising
the manuscript.
Literature Cited
Baker, R. H. 1983. Michigan mammals. East
Lansing: Michigan State Univ. Press.
Burt, W. H. 1948. The mammals of Michigan.
Ann Arbor: Univ. Michigan Press.
Calhoun, S. W. and I. R. Greenbaum. 1991.
Evolutionary implications of genic variation
among insular populations of Peromyscus
maniculatus and Peromyscus oreas.J. Mammal.
72:248-62.
Choate, J. R. 1973. Identification and recent
distribution of white footed mice (Peromyscus)
in New England. / Mammal. 54:41-49.
Gunderson, H. L. and J. R. Beer. 1953. The
Mammals of Minnesota. Minn. Mus. Nat.
Hist. Occasional Paper 6.
Hazard, E. B. 1982. The mammals of Minnesota.
Minneapolis: Univ. Minnesota Press.
Jackson, H. H. T. 1961. Mammals of Wisconsin.
Madison: Univ. Wisconsin Press.
Ledehrle, P. G. et al. 1985. Size variation in
Peromyscus maniculatus gracilis from the Bea¬
ver Islands. Jack-Pine Warbler 63: 107-10.
Long, C. A. 1 968 . Populations of small mammals
on railroad right-of-way in prairie of central
Table 3. Fisher’s Linear Discriminant Functions. These classification coefficients may be
applied to other samples.
Lake Michigan Isles Central Wisconsin Lake Superior
1 20
TRANSACTIONS
LONG and LONG: Long-tailed deer mice of Wisconsin and Michigan
Illinois. Trans. Illinois State Acad. Sci. 61:139—
45.
Long, C. A. 1 974. Mammals of the Lake Michigan
drainage basin. Environmental Status of the
Lake Michigan Region. Argonne, Illinois:
Argonne Nat. Lab.
Long, C. A. 1978. Mammals of the islands of
Green Bay, Lake Michigan. Jack-Pine Warbler
56:59-82.
Osgood, W. H. 1909. Revision of the mice of the
American genus Peromyscus. North Amer.
Fauna 28.
Stromberg, M. R. 1979. Field identification of
Peromyscus leucopus and P. maniculatus with
discriminant analysis. Trans. Wisconsin Acad.
Sci., Arts and Letters 67: 1 59-64.
Charles A. Long is Professor of Biology and Mu¬
seum Curator of Mammals at the University of
Wisconsin-Stevens Point.
John E. Long is a graduate student in the School of
Mathematics, University of Minnesota. C. Long
was a Visiting Professor and J. Long a math major
at St. Olaf College when the statistical study was
initiated.
Volume 81 (1993)
121
John Lyons
Status and biology of
Paddlefish (Polyodon spathula)
in the Lower Wisconsin River
A bstract The paddlefish (Polyodon spathula) is a Threatened Species in Wisconsin. His¬
torically, paddlefish occurred over 224 km of the Lower Wisconsin River, from
its mouth at the Mississippi River upstream to Wisconsin Dells. Paddlefish have
not been reported from the 76 km between Wisconsin Dells and the Prairie du
Sac Dam since the 1950s. A large population, possibly the largest remaining
in Wisconsin, exists below the Prairie du Sac Dam, but further downstream,
paddlefish are uncommon . From 1988 through 1990, several thousand paddle¬
fish occurred in the 6-km stretch below the Prairie du Sac Dam. Paddlefish
were most abundant in the 20-ha tailwater pool below the dam. Almost all
paddlefish observed below the dam were large, with mean sizes of 15 kg and
134 cm total length, and maxima of 29.5 kg and 168 cm. Mean total length
increased from 127 cm in 1988 to 139 cm in 1990. No young-of-year paddle¬
fish and only one paddlefish under 5 kg were among the 237 paddlefish cap¬
tured and weighed. The scarcity of small fish may have been merely a sam¬
pling bias or the result of poor spawning success and recruitment during the
mid to late 1980s. Most paddlefish had evidence of recent parasitism by silver
lampreys (Ichthyomyzon unicuspis), but lampreys probably did not cause sub¬
stantial paddlefish mortality. Collisions with boats and snagging by anglers had
injured many paddlefish. I recommend that management of paddlefish in the
Lower Wisconsin River focus on: 1) improved understanding of reproduction
and recruitment; 2) maintenance of a natural river flow regime; 3) continued
prevention of illegal harvest; 4) re-establishment of a population above the Prai¬
rie du Sac Dam.
The paddlefish (Polyodon spathula ), one of the largest and
most unusual of Wisconsin fishes, was once much more
widespread and abundant in Wisconsin than it is today (Becker
1983; Gengerke 1986). Before the beginning of intensive Eu-
TRANSACTI ONS
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
ropean settlement of the state 150 years ago,
paddlefish were common in the Mississippi
River and the lower reaches of its major
tributary basins, the St. Croix, Chippewa,
and Wisconsin. Small populations were also
present in Lakes Michigan and Superior.
Over the last 150 years, paddlefish have be¬
come far less abundant, disappearing from
the Great Lakes by the early 1900s, and de¬
clining greatly in the Mississippi and its
tributaries. Declines of paddlefish popula¬
tions in Wisconsin and elsewhere have been
attributed to habitat degradation, dam con¬
struction, water pollution, and possibly over¬
harvest (Becker 1983; Pasch and Alexander
1986; Sparrowe 1986; Unkenholz 1986).
The paddlefish is a Threatened Species in
Wisconsin.
Paddlefish persist at only a few locations
in Wisconsin (Becker 1983; Fago 1992).
What may be the largest population occurs
below the Prairie du Sac Dam on the Lower
Wisconsin River in southwestern Wisconsin.
Since 1988, I have studied the paddlefish
below the dam, with the goal of collecting
basic biological information necessary for
preserving and increasing the population. In
this paper, I report on the distribution and
population status of paddlefish in the Lower
Wisconsin River, and provide recommenda¬
tions for managing the population. The data
I present may serve as a baseline against
which to judge future efforts to rehabilitate
paddlefish populations throughout the state.
Study Area
The Wisconsin River originates at Lac Vieux
Desert on the Wisconsin-Michigan bound¬
ary and flows 684 km south and west to the
Mississippi River (Fig. 1). Since creation of
the Lower Wisconsin State Riverway in
1989, the Lower Wisconsin River has usu¬
ally been defined as the 148-km segment
between the Mississippi River and the Prai¬
rie du Sac Dam, the first dam encountered
upstream from the mouth of the Wisconsin
(WDNR 1988). However, for purposes of
this paper, I also include in my discussion
of the Lower Wisconsin River the 76-km
segment from the Prairie du Sac Dam up¬
stream to the next dam, the Kilbourn Dam
(river kilometer [RKM] 224) at Wisconsin
Dells (Fig. 1). Historically, the biota of this
segment were similar to those of the segment
downstream from the Prairie du Sac Dam.
The Prairie du Sac Dam and the Kilbourn
Dam are used for hydroelectric power. They
are both impassable to fish moving upstream,
although fish may be able to move down¬
stream through them. The Prairie du Sac
Dam has a head of 12.5 m and was con¬
structed during 1911-1914, and the Kil¬
bourn Dam has a head of 7.9 m and was
constructed during 1907-1910 (Wisconsin
Power and Light Corporation [WPLC] , un¬
published data). Presently, both dams are
operated on a “run-of-the-river” basis, and
water discharges through them are not regu¬
lated to any great extent. Dams further up¬
stream on the Wisconsin River regulate river
flows much more, and discharge patterns
from these upstream dams can influence dis¬
charge patterns through the Kilbourn and
Prairie du Sac dams.
The Lower Wisconsin River had mean
annual discharges of 192 m3/sec at the
Kilbourn Dam in 1934-1990 (Holmstrom
and Erickson 1990), 224 m3/sec at the Prai¬
rie du Sac Dam in 1950-1989 (WPLC, un¬
published data), and 246 m3/sec at the
Muscoda gaging station (RKM 71) in 1913-
1990 (Holmstrom and Erickson 1990).
Downstream from the Prairie du Sac Dam,
the river is generally wide (> 200 m) and
relatively shallow (< 3 m), with a primarily
shifting sand bottom. Unique conditions
exist immediately below the Prairie du Sac
1 24
TRANSACTIONS
LYONS: Status and biology of paddlefish in the Lower Wisconsin River
Fig. 1 . Map of the Lower Wisconsin River, showing locations mentioned in the text.
Dam, where scouring has created a 20-ha
tailwater pool that is over 12 m deep in
places. The bottom in and adjacent to this
pool contains relatively large amounts of
gravel and cobble, although sand still pre¬
dominates.
Methods
I obtained historical distribution and abun¬
dance data on paddlefish in the Lower Wis¬
consin River from published accounts, the
Wisconsin Department of Natural Re¬
sources (WDNR) Fish Distribution Survey
Database (Fago 1988), and unpublished
records associated with paddlefish specimens
preserved at the University of Wisconsin
Zoological Museum (UWZM) in Madison
and the University of Wisconsin Museum
of Natural History (UWSP) in Stevens
Point.
Current distribution and abundance data
came from recent WDNR electroshocking
surveys of the Lower Wisconsin River. The
electroshockers used were standard WDNR
two- or three-person, boat-mounted, pulsed
direct-current “boom shockers,” powered by
either a 2500 watt or 5000 watt generator
(Novotny and Priegel 1974). Between
March and November 1985-1990, one to
ten surveys per year were conducted in the
area immediately below the Prairie du Sac
Dam. Many paddlefish were observed dur¬
ing all of these surveys, but paddlefish were
collected only from 1988 to 1990. Addi¬
tional electroshocking surveys were con-
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
ducted at sites closer to the Mississippi River
on one date each in July 1985, April 1987,
and June 1991, and on eight dates in Octo¬
ber 1989.
During electroshocking surveys, efforts
were made to minimize mortality of cap¬
tured paddlefish. Paddlefish stunned by the
electroshocker were quickly removed from
the river and placed in a horse trough filled
with river water. There they were measured
to the nearest cm, weighed to the nearest 0.5
kg, checked for physical damage and lam¬
prey parasitism, and then allowed to recover
before being released. Normally, only one
paddlefish was captured and processed at a
time, and each captured fish was handled as
little and as gently as possible. Most fish re¬
covered from the electric shock within min¬
utes, swam away strongly when released,
and, presumably, survived. However, in each
survey, some paddlefish (5 to 15%) died as
a result of capture and handling. I took tis¬
sue and organ samples from 10 of the
paddlefish that died between late March and
early June 1990, and provided these samples
for use in a nationwide paddlefish genetics
study (Epifanio et al. 1990). I also examined
stomach contents from these 10 paddlefish.
In previous studies, paddlefish size has
been characterized by different length mea¬
surements (Russell 1986). To allow Lower
Wisconsin River data to be compared with
results from these earlier studies, four length
measurements were made on most paddle¬
fish captured: body length — the distance
from the anterior edge of the eye to the cau¬
dal fork; mouth-fork length — the distance
from the anterior tip of the jaw to the cau¬
dal fork; rostrum-fork length — the distance
from the tip of the rostrum (paddle) to the
caudal fork; and total length — the distance
from the tip of the rostrum to the posterior
tip of the upper caudal lobe. I used linear
regression (SAS 1988) to develop equations
to convert one measurement to another. I
also used linear regression to develop quan¬
titative length-weight relationships. I com¬
pared paddlefish total length distributions
among different sampling periods with the
Kruskal- Wallis test (SAS 1988). I estimated
the approximate age of Wisconsin River
paddlefish based on the total length-age re¬
lationship developed for paddlefish in Pool
13 of the Mississippi River (Gengerke
1978). I did not determine age directly be¬
cause paddlefish must killed to be aged
(Russell 1986).
In October of 1988 and 1989, I made
mark-recapture estimates of the number of
paddlefish in the area below the Prairie du
Sac Dam. For two days each year, captured
paddlefish were marked with a length of
brightly colored plastic flagging tape that was
wrapped around the caudal peduncle (13 in
1988; 29 in 1989). Within 48 hours of
marking, the sampling crew made up to four
electroshocking recapture passes in the vicin¬
ity of the dam, counting, but not netting,
all paddlefish that surfaced between the elec¬
trodes (770 in 1988; 471 in 1989), and not¬
ing whether any of these paddlefish were
marked (2 in 1988; 2 in 1989). The bright
flagging tape was easily seen on paddlefish
in the water. I estimated population size us¬
ing the modified Petersen formula, with
asymmetric confidence intervals calculated
under the assumption that recapture prob¬
abilities for marked fish followed a Poisson
distribution (Ricker 1975).
In 1989, I began an annual paddlefish
abundance monitoring program. I estab¬
lished a standardized electroshocking circuit
for paddlefish around the tailwater pool be¬
low the Prairie du Sac Dam. The circuit was
about 1.6 km long and took up to 25 min¬
utes to shock. In October 1989 and Octo¬
ber 1990, the field crew shocked this circuit,
counting, but not netting, all paddlefish ob-
1 26
TRANSACTIONS
LYONS: Status and biology of paddlefish in the Lower Wisconsin River
served, and noting whether each one ap¬
peared to be more or less than 100 cm in
total length (a length at which paddlefish
weigh approximately 5 kg). I used the total
number observed in each circuit as an index
of paddlefish abundance.
Historical Distribution and
Abundance
The first reports of paddlefish from the
Lower Wisconsin River date from the early
1900s, the period when the first scientific
surveys of the fishes of the Wisconsin River
were undertaken. Greene (1935), Becker
(1966, 1983), and Fago (1992) summarized
these surveys, and recorded paddlefish from
the Wisconsin River at Prairie du Sac in
1924 and 1937 and at Wisconsin Dells in
1931. Wisconsin Dells probably was the his¬
toric upstream limit for paddlefish in the
Wisconsin River. No paddlefish have ever
been reported upstream from this point. Be¬
fore completion of the Kilbourn Dam in
1910, a large rapids occurred in the Wiscon¬
sin Dells gorge (Stark 1988), and this rap¬
ids may have been impassable to paddlefish.
Between the mid 1800s and 1910, a low-
head (2 to 3 m), wooden logging dam was
also present at Wisconsin Dells (Stark 1988),
further impeding paddlefish upstream move¬
ment.
The last record of paddlefish from the
stretch between the Prairie du Sac Dam and
Wisconsin Dells came in 1950 (Becker
1983), when a large individual was found in
the Baraboo River, a tributary that enters the
Wisconsin River at RKM 181, near the city
of Portage (Fig. 1). Fish surveys of this
stretch of the Wisconsin River during the
1970s, 1980s, and 1990s failed to find
paddlefish (Tim Larson, WDNR Fish Man¬
ager, Poynette, personal communication;
David Morrow, fisheries biologist, Mead and
Hunt, Inc., personal communication; per¬
sonal observations). Causes of the disappear¬
ance of paddlefish are unknown, although I
speculate that poor water quality in the river
from the 1940s through the 1970s may have
played a role. During this period, discharges
from upstream industries, particularly paper
mills, caused major declines in dissolved oxy¬
gen levels and were responsible for several
large fish kills in the Lower Wisconsin River
(Poff and Threinen 1965; WDNR 1976;
WDNR, unpublished data). By the late
1970s, pollution abatement had greatly re¬
duced water quality problems in the Lower
Wisconsin River, but by then paddlefish
were gone. The Prairie du Sac Dam prevents
paddlefish from moving upstream and re¬
colonizing this stretch of river.
Only eight confirmed records of paddle¬
fish, all large individuals, exist from the
Lower Wisconsin River below Sauk City
(RKM 142) (Becker 1966, 1983; Fago
1992; UWSP Specimens; WDNR Fish Dis¬
tribution Survey Database). Two records
were from the early 1960s, when two dead
paddlefish were found near Muscoda. The
remaining six records were from extensive
electroshocking and netting surveys that
were carried out during the 1970s. One of
these records (one fish) was from near the
mouth of the Wisconsin at Bridgeport
(RKM 10), and the other five (eight fish to¬
tal) were from between Spring Green and
Mazomanie (RKM 108 to 135) (Fig. 1).
During these same 1970s surveys, over 125
paddlefish were captured from the 6 km of
river between Sauk City and the Prairie du
Sac Dam. In the 1980s, more than 2000
paddlefish were captured or observed in 38
days of electroshocking in the three kilome¬
ters immediately below the dam. Conversely,
no paddlefish were observed in 1 1 days of
shocking areas further downstream — one
day upstream of Mazomanie (RKM 136-
Volume 81 (1993)
1 27
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
142), one day near Lone Rock (RKM 95),
and nine days from near the mouth up to
Boscobel (RKM 3 to 45) (Fig. 1).
Paddlefish have been common in the area
below the Prairie du Sac Dam for at least the
last 50 years. During March 1945, “over a
ton” of dead paddlefish was observed at Sauk
City, presumably killed by poor water qual¬
ity (UWZM, unpublished data). During the
1960s, large numbers of paddlefish were
regularly observed directly below the pow¬
erhouse of the Prairie du Sac Dam (Poff and
Threinen 1965; Becker 1966; UWZM, un¬
published data; Lyle Christenson, WDNR
Fisheries Research Biologist, Monona, per¬
sonal communication).
From 1985 through 1990, numbers of
paddlefish observed during surveys below
the Prairie du Sac dam were typically 10 to
30 times higher in and adjacent to the
tailwater pool than in areas further down¬
stream. However, groups of 1 0 to 20 paddle¬
fish were sometimes observed downstream
from the tailwater pool, usually in the vicin¬
ity of the State Highway 60 Bridge in Prai¬
rie du Sac (RKM 146) or the railroad bridge
in Sauk City (RKM 142).
Recent electroshocking surveys and dis¬
cussions with anglers and scuba divers indi¬
cated that large numbers of paddlefish were
present in the tailwater pool in all months
of the year. At least 75 paddlefish were ob¬
served during each monthly electroshocking
survey of the tailwater pool between March
and November 1989. Anglers who fished the
pool reported observing or accidently snag¬
ging paddlefish during all months of the
year. In February 1982, scuba divers in the
pool observed numerous large paddlefish
(Mike Talbot, WDNR Fisheries Manage¬
ment Biologist, Madison, personal commu¬
nication).
Although data are limited, there is no evi¬
dence that large numbers of paddlefish mi¬
grate to or from the area immediately below
the dam. There are few confirmed records
of paddlefish from the Lower Wisconsin
River below the Prairie du Sac Dam or from
Pool 10 of the Mississippi River at the
mouth of the Wisconsin River (Becker
1966, 1983; WDNR Fish Distribution Sur¬
vey Database), although a few commercial
fishermen say that they regularly catch
paddlefish in Pool 10 (Cecil Jennings, U.S.
Fish and Wildlife Service Fisheries Biologist,
LaCrosse, Wisconsin, personal communica¬
tion). Moreover, genetic differences exist
between the Prairie du Sac Dam paddlefish
population and Mississippi River paddlefish
populations (Epifanio et al. 1990), suggest¬
ing little mixing between paddlefish from
the Wisconsin and Mississippi rivers.
The tailwater pool area of the Prairie du
Sac Dam appears to have all the necessary
habitats for paddlefish to complete their life
cycle. Although paddlefish have not been
observed spawning in Wisconsin waters, the
gravel bars immediately below the tailwater
pool conform to descriptions of good pad¬
dlefish spawning habitat (Purkett 1961;
Pasch et al. 1980; Russell 1986; Crance
1987). The deep, slow-moving waters of the
tailwater pool itself appear to constitute ex¬
cellent summer feeding and winter resting
habitat for both juveniles and adults (based
on descriptions in Southall and Hubert
1984; Russell 1986; Crance 1987), and the
lentic environment above the dam provides
a source of crustacean zooplankton (personal
observations), a primary food of paddlefish
(Rosen and Hales 1981).
Abundance and Size Structure Below
the Prairie du Sac Dam
During the late 1980s, several thousand
paddlefish lived in the Prairie du Sac Dam
tailwater pool. In October 1988, the esti-
1 28
TRANSACTIONS
LYONS: Status and biology of paddlefish in the Lower Wisconsin River
mated population size was 3600 (93% con¬
fidence interval: 1320-9000), and in Octo¬
ber 1989 it was 4720 (93% confidence in¬
terval: 1730-11800). These may be overes¬
timates, because only two marked paddlefish
were recaptured in each month. Mark-recap¬
ture population estimates based on less than
three recaptures tend to be biased, typically
yielding estimates that are too high (Ricker
1975). Nonetheless, a population of several
thousand paddlefish seems reasonable.
Throughout the study period, it was normal
to observe hundreds of paddlefish during
two or three hours of shocking the tailwater
pool.
No population estimate was made in
1990, but the abundance of paddlefish may
have been less than in 1988 and 1989. The
number observed along the standardized
electroshocking circuit was 133 in 1989 and
78 in 1990. Generally, fewer paddlefish were
observed during 1990 surveys than during
1988 and 1989 surveys.
Almost all of the paddlefish observed or
captured were large. Of 237 paddlefish
weighed, only one was less than 5 kg (4 kg;
97 cm total length [TL] ) . Of the hundreds
of other paddlefish observed but not netted,
only four appeared to weigh less than 5 kg.
The mean size of captured paddlefish was 1 5
kg and 134 cm TL, and the maximum size
was 29.5 kg and 168 cm TL (36 paddlefish
with damaged upper caudal lobes or ros¬
trums were not included in total length sta¬
tistics). Nineteen percent of the captured
paddlefish weighed 20 kg or more.
Based on the distribution of total lengths,
most paddlefish were probably 8 to 14 years
old, with the smallest individuals aged 4 to
7 years and the largest individuals greater
than age 18 (Table 1). By age 8 to 12, all
male paddlefish and most female paddlefish
are likely to be mature (Gengerke 1978;
Russell 1986), hence mature adults domi¬
nated catches in the Lower Wisconsin River
from 1988 through 1990.
The average total length of captured
paddlefish increased from 1988 to 1990
(Table 1). Over the period October 1988
through April 1989, paddlefish mean TL
was 1 27 cm (I assumed no growth occurred
between October and April). By October
1990, mean TL was significantly greater at
139 cm (chi-square = 28.7; p - 0.0001).
During the period October 1988 through
April 1989, 36% of captured paddlefish
were less than 125 cm TL, and only 2%
were greater than 145 cm. From October
1989 through April 1990, 14% were less
than 125 cm and 34% were greater than 145
cm. By October 1990, only 3% were less
than 125 cm and 38% were greater than 145
cm.
Sampling bias might account for the near
absence of small paddlefish in electro-
shocking catches. In other river systems,
young-of-year paddlefish have usually prov¬
en more difficult to capture than larger in¬
dividuals (Purkett 1961; Pasch et al. 1980;
Russell 1986). Boom shockers effectively
sample only the top 2 m of the water col¬
umn, and if small paddlefish occupy deeper
water, they would not be vulnerable to cap¬
ture. However, little is known about habi¬
tat use by small paddlefish (Russell 1986).
Small paddlefish might occupy some other
part of the Lower Wisconsin River and only
move into the area below the Prairie du Sac
Dam after they grow to a relatively large size.
Although small paddlefish have never been
captured in surveys of the Lower Wiscon¬
sin River outside of the Prairie du Sac Dam
area, many kilometers of the river remain
unsampled.
Conversely, the scarcity of small paddle¬
fish in electroshocking samples might rep¬
resent a real scarcity in the Lower Wiscon¬
sin River population. The increasing aver-
Volume 81 (1993)
129
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1 . Numbers of paddlefish captured, by total length (TL) class, from below the Prairie du
Sac Dam, Lower Wisconsin River, 1988-1990. Only fish without damaged rostrums or upper
caudal lobes are included. Paddlefish captured between October and April are combined be¬
cause I assumed that no growth occurred during this time interval. Approximate age ranges
are based on data from Mississippi River paddlefish (Gengerke 1978).
October through April Only
aHigh flows prevented effective sampling for paddlefish in March & April 1991.
age lengths of captured paddlefish suggest
that there was little recruitment to the adult
population during the study period. Poor
reproductive success during the mid to late
1980s could explain the low numbers of
young paddlefish.
The time of paddlefish spawning in Wis¬
consin waters is unknown (Becker 1983),
but based on preferred water temperatures
for reproduction (13 to 22°C), and the
spawning season in states to the south (mid
to late April in Missouri and Tennessee —
Purkett 1961; Pasch et al. 1980; late April
to late May in Iowa — Southall and Hubert
1984), paddlefish in the Lower Wisconsin
River probably spawn in May. Water tem¬
peratures below the Prairie du Sac Dam
range from 1 1 to 17°C in early May and 18
to 22°C in late May (Don Fago, WDNR
Fisheries Research Biologist, Fitchburg, un¬
published data for 1987-1990). Paddlefish
reproductive success tends to be highest in
years when river flows are at or near flood
levels during and for at least a week after
spawning (Purkett 1961; Russell 1986). The
specific flows required to provide good
spawning conditions at a site depend on the
morphometry of the river channel; paddle¬
fish normally spawn on submerged gravel
bars with water velocities greater than 0.4 m/
sec and depths greater than 2 m (Crance
1987). Thus, river flows in May probably
determine paddlefish reproductive success in
the Lower Wisconsin River.
130
TRANSACTIONS
LYONS: Status and biology of paddlefish in the Lower Wisconsin River
I hypothesize that river flows in May at
the Prairie du Sac Dam might have been
below the optimum for paddlefish reproduc¬
tion during most of the 1980s. Based on my
extensive observations of the Lower Wiscon¬
sin River over a wide range of flows and an
examination of the discharge vs. water level
relationship below the dam, I have found
that large areas of submerged gravel bars
with depths deeper than 2 m and water ve¬
locities greater than 0.4 m/sec occur in the
vicinity of the dam only when river flows are
greater than 400 m3/sec. Mean flow in May
at the Prairie du Sac Dam was 306 m3/sec
in 1930-1990 (WPLC, unpublished data).
Since 1930, flows in May have exceeded 400
m3/sec for more than seven consecutive days
(and hence been best for paddlefish spawn¬
ing) in nine years: 1951, 1954, I960, 1965,
1972, 1973, 1975, 1979, and 1984 (WPLC,
unpublished data) . Thus, flows likely to pro¬
duce optimal spawning conditions occurred
only once during the 1980s. Seven of the
years in the 1980s had mean May flows be¬
low the long-term average. The 8- to 18-
year-old paddlefish that dominated catches
in 1988-1990 were hatched between 1970
and 1982, a period in which four years of
optimal flows occurred.
Miscellaneous Observations
The four length measurements made on
paddlefish from the Lower Wisconsin River
were strongly correlated with each other.
Regressions relating each measurement to
total length and body length, the most com¬
mon measurements in the literature, are as
follows:
Total Length - 20.384 + (1.271 x Body Length)
(N = 202; F= 1639; p = 0.0001; r2 = 0.89)
Total Length = 22.231 + (1.242 x Mouth-Fork
Length)
(N = 203; F = 1760; p = 0.0001; r2 = 0.90)
Total Length = 3.421 + (1.074 x Rostrum-Fork
Length)
(N = 203; T=3294; p = 0.0001; r2 = 0.94)
Body Length = 2.747 + (0.963 x Mouth-Fork
Length)
(N = 217; F= 10106; p = 0.0001; r2 = 0.98)
Body Length = -6.480 + (0.789 x Rostrum-Fork
Length)
(N = 205; F= 2537; p = 0.0001; r2 = 0.93)
Paddlefish weight was also strongly corre¬
lated with both total length and body length:
Loge (Weight) = -12.706 + (3.132 x Loge [Total
Length])
(N = 202; F= 484; p = 0.0001; r2 = 0.71)
Loge(Weight) = -10.408 + (2.902 x Loge [Body
Length])
(N = 213; F= 686; p = 0.0001; r2 = 0.76)
Nearly all paddlefish from below the Prai¬
rie du Sac Dam suffered from parasitism by
silver lampreys (Ichthyomyzon unicuspis). Of
240 paddlefish examined, 231 (96%) had
either an attached lamprey or a fresh lam¬
prey wound. Individual paddlefish had up
to 1 1 attached lampreys and 28 fresh
wounds; the mean number of lampreys at¬
tached was 1.2 and the mean number of
fresh wounds was 6.1. Most paddlefish also
had healed wounds from previous lamprey
attacks.
I do not believe that lampreys cause sub¬
stantial mortality of paddlefish in the Lower
Wisconsin River. Lamprey parasitism has
been common on Lower Wisconsin River
paddlefish since at least the 1940s (UWZM,
unpublished data), and yet a large paddle¬
fish population persists. The prevalence of
healed wounds on paddlefish shows that
paddlefish survive lamprey attacks. The ra¬
tio of paddlefish biomass to attached lam¬
prey biomass is typically much more than
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
50 to 1. In experiments involving sea lam¬
preys ( Petromyzon marinus) feeding on lake
trout (Salvelinus namaycush ) and rainbow
trout ( Oncorhynchus my kiss). Farmer et al.
(1975) found that lampreys caused direct
mortality of their host only when the ratio
of host biomass to lamprey biomass was less
than 40 to 1 .
Some paddlefish had injuries or tissue
damage not caused by lampreys. Twelve of
240 paddlefish (5%) had damaged rostrums,
and 55 (23%) had external damage to some
other part of the body. In some instances
paddlefish had lost most or all of their ros¬
trum. Much of the damage clearly had been
caused by fishing hooks and lines or by boat
motor propellers. Paddlefish often swam just
below the surface, and on several occasions
I saw motor boats collide with them. Al¬
though fishing for paddlefish is illegal, I
sometimes saw anglers deliberately trying to
snag them. When anglers did catch paddle¬
fish, they would often handle them roughly
and keep them out of the water for long pe¬
riods.
The frequency of tissue damage for
paddlefish from below the Prairie du Sac
Dam was similar to that observed elsewhere.
In Pool 13 of the Mississippi River, Gen-
gerke (1978) reported that 30 (5%) of 603
paddlefish had damaged rostrums. However,
only 9 1 of 1 543 paddlefish (6%) had dam¬
age from fishing hooks and lines or boat pro¬
pellers. In the Missouri River in South Da¬
kota, 46 of 458 paddlefish (10%) had dam¬
aged rostrums, and an additional 118 (26%)
had damage to other parts of their body
(Rosen and Hales 1980). Most of this dam¬
age was attributed to fishing hooks and lines
or to boat propellers.
Paddlefish appeared to travel in schools
below the Prairie du Sac Dam. Paddlefish
were usually observed in groups of 10 or
more. It was common to electroshock an area
of the tailwater pool and observe no paddle¬
fish, only to return several hours later and
observe many paddlefish. However, paddle¬
fish distribution in the tailwater pool was not
random. Certain areas, particularly eddies
adjacent to fast current, consistently had
higher numbers of paddlefish than elsewhere.
Based on limited data, it appeared that
paddlefish from below the Prairie du Sac
Dam fed largely on crustacean zooplankton.
All 1 0 of the paddlefish stomachs examined
between late March and early May 1990
contained primarily Daphnia , a crustacean
zooplankter that was present in high densi¬
ties in the tailwater pool during this period
(personal observations) . The amount of food
in the stomachs was impressive; several con¬
tained more than 1 kg (wet weight) of Daph¬
nia. Other studies have reported that crus¬
tacean zooplankton are a major food of
paddlefish (Wagner 1908; Rosen and Hales
1981; Becker 1983; Russell 1986).
Management Recommendations
I believe that a priority in management of
the paddlefish population of the Lower Wis¬
consin River should be to learn more about
the distribution and abundance of small (i.e.,
< 5 kg) paddlefish. Until the reason for the
scarcity of small paddlefish in electroshock-
ing samples can be explained, management
efforts will be hampered by uncertainty
about the size and abundance trends of the
population. I recommend a two-part study
to clarify the status of small paddlefish. The
first part should determine whether electro-
shocking is biased towards larger paddlefish.
A variety of methods that effectively sample
deep water, such as trammel and gill netting,
seining, and trawling, should be compared
with electroshocking in the tailwater pool
and in areas further downstream. If these
other techniques capture small paddlefish in
132
TRANSACTIONS
LYONS: Status and biology of paddlefish in the Lower Wisconsin River
good numbers, then they should be used to¬
gether with electroshocking in the annual
abundance monitoring program.
The second part of the study should de¬
termine where, when, and under what con¬
ditions paddlefish reproduce successfully in
the Lower Wisconsin River. Efforts to pre¬
serve and increase the paddlefish population
below the Prairie du Sac Dam would be en¬
hanced by a better understanding of factors
that dictate spawning success and subse¬
quent recruitment to the adult population.
Because optimal conditions for paddlefish
spawning might occur only a few times each
decade, this part of the study could take
many years to complete.
The likelihood that paddlefish need high
river flows for successful spawning suggests
that paddlefish reproduction might be en¬
hanced if the Prairie du Sac Dam and up¬
stream dams artificially increased flows dur¬
ing May. However, I recommend against
this for several reasons. First, there are no
actual data on paddlefish spawning in the
Lower Wisconsin River. As a result, it is im¬
possible to provide precise recommendations
as to when, how much, and for how long
flows should be augmented. The river flow
conditions that I have suggested were opti¬
mal for paddlefish reproduction represent a
hypothesis that needs to be confirmed with
field data before changes in dam operations
are considered. Second, neither the Prairie
du Sac Dam nor the Kilbourn Dam have
substantial water storage capacity, and it is
unlikely that they could be used to increase
flows without unacceptable declines in the
levels of the impoundments behind them.
Moreover, the Prairie du Sac Dam is in the
process of being licensed by the Federal En¬
ergy Regulatory Commission (FERC), and
a condition of the license will be that the
dam continue to operate in run-of-the-river
mode. The WDNR will also recommend
continued run-of-the-river operation of the
Kilbourn Dam if it is licensed by FERC
(Bob Hansis, WDNR Water Management
Specialist, Fitchburg, personal communica¬
tion). Several dams upstream from the
Kilbourn Dam have large storage capacity,
but using them to increase flows below the
Prairie du Sac Dam would be complicated.
Finally, although artificial high flows might
benefit paddlefish, they might harm other
species. The biotic community of the Lower
Wisconsin River is complex, and modifica¬
tion of natural flow patterns, however well
intentioned, might have unforeseen negative
consequences. I recommend taking a con¬
servative management tack and changing
natural river flows as little as possible.
Illegal harvest is a threat to the paddle¬
fish population in the Lower Wisconsin
River. Paddlefish eggs make excellent caviar
and fetch high prices. The potential to earn
large amounts of money by selling paddle¬
fish eggs has led to considerable organized
illegal harvest in some areas, and caused se¬
rious harm to paddlefish populations in
some waters (Missouri Department of Con¬
servation and U.S. Fish and Wildlife Service,
unpublished data) . Although there is no evi¬
dence of substantial illegal harvest in the
Lower Wisconsin River, the year-round con¬
centration of paddlefish below the Prairie du
Sac Dam makes the population vulnerable
to illegal snagging and netting. The area be¬
low the dam is regularly patrolled by
WDNR Law Enforcement personnel, but if
evidence of substantial illegal harvest comes
to light, the patrols should be increased.
Recreational use of the area below the
Prairie du Sac Dam clearly causes injuries to
many paddlefish, although the effect of these
injuries on the paddlefish population is un¬
certain. Two actions could be taken to re¬
duce the number of paddlefish injuries.
First, the tailwater pool area could be de-
Volume 81 (1993)
1 33
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
dared a “no wake” zone. Boaters would be
required to travel slowly when in the area,
allowing them and the paddlefish more time
to see each other and avoid collisions. Sec¬
ond, educational signs and pamphlets could
be developed to urge anglers to avoid trying
to snag paddlefish, and to release quickly
and carefully any paddlefish they caught ac¬
cidentally.
I recommend that an attempt be made to
re-establish paddlefish in the stretch of river
between the Prairie du Sac Dam and the
Kilbourn Dam. Water quality in the area
below the Kilbourn Dam has improved
markedly since the 1960s and 1970s, and
the habitat there appears to have depth, ve¬
locity, and substrate characteristics suitable
for paddlefish. The best source of paddlefish
for reintroduction would be from below the
Prairie du Sac Dam. The paddlefish from
here are probably part of the same stock that
once inhabited the area below the Kilbourn
Dam, and by using them, the potential for
introducing genetically unsuitable fish
would be minimized (Epifanio et al. 1990).
The population of paddlefish below the Prai¬
rie du Sac Dam is large enough that removal
of some individuals (< 250) for stocking up¬
stream probably would not be harmful.
Acknowledgments
I thank the many people who helped me
observe and collect paddlefish in the Lower
Wisconsin River. I am indebted to those bi¬
ologists who allowed me access to their un¬
published data and observations. Lyle M.
Christenson, Philip A. Cochran, and two
anonymous reviewers provided useful com¬
ments on an earlier draft of this paper.
Electroshocking surveys during which pad¬
dlefish were collected were funded in part
through the Sport Pish Restoration Program,
Project P-83-R, Study 044.
Works Cited
Becker, G. C. 1966. Fishes of southwestern Wis¬
consin. Transactions of the Wisconsin Academy
of Science, Arts, and Letters 53:87-1 17.
Becker, G. C. 1983. Fishes of Wisconsin. Madi¬
son: University of Wisconsin Press. 1052 pp.
Crance, J. H. 1987. Flabitat suitability index
curves for paddlefish, developed by the
Delphi Technique. North American Journal
of Fisheries Management 7: 123-30.
Epifanio, J., M. Nedbal, and D. P. Philipp.
1990. A population genetic analysis of
paddlefish (Polyodon spathula). Aquatic Biol¬
ogy Technical Report 90/13. Illinois Natu¬
ral History Survey, Champaign. 49 pp.
Fago, D. 1988. Retrieval and analysis used in
Wisconsin’s statewide fish distribution sur¬
vey, second edition. Research Report 148.
Wisconsin Department of Natural Resources,
Madison. 53 pp.
Fago, D. 1992. Distribution and relative abun¬
dance of fishes in Wisconsin. VIII. Final re¬
port. Technical Bulletin 175. Wisconsin De¬
partment of Natural Resources, Madison.
378 pp.
Farmer, G. J., F. W. H. Beamish, and G. A.
Robinson. 1975. Food consumption of the
adult landlocked sea lamprey Petromyzon
marinus L. Comparative Biochemistry and
Physiology 50A:753-57.
Gengerke, T. W. 1978. Paddlefish investiga¬
tions. Project Completion Report, Project
No. 2-255-R, Research Segment, Commer¬
cial Fisheries Investigations. Iowa Conserva¬
tion Commission, Des Moines. 86 pp.
Gengerke, T. W. 1986. Distribution and abun¬
dance of paddlefish in the United States, pp.
22-35 in J. G. Dillard, L. K. Graham, and
T. R. Russell (eds.), The paddlefish: status,
management and propagation. Special Pub¬
lication 7, North Central Division, American
Fisheries Society. 159 pp.
Greene, C. W. 1935. The distribution of Wis¬
consin fishes. Wisconsin Conservation De¬
partment, Madison. 235 pp.
Holmstrom, B. K., and R. M. Erickson. 1990.
Water resources data. Wisconsin. Water Year
134
TRANSACTIONS
LYONS: Status and biology of paddlefish in the Lower Wisconsin River
1989. Water-Data Report WI-89-1, USGS/
WRD/HD-90/298. U.S. Geological Survey,
Madison, Wisconsin. 436 pp.
Novotny, D. W., and G. R. Priegel. 1974.
Electrofishing boats. Improved designs and
operational guidelines to increase the effec¬
tiveness of boom shockers. Technical Bulle¬
tin 73. Wisconsin Department of Natural
Resources. 48 pp.
Pasch, R. W., and C. M. Alexander. 1986. Ef¬
fects of commercial fishing on paddlefish
populations, pp. 46-33 in J. G. Dillard, L.
K. Graham, and T. R. Russell (eds.), The
paddlefish: status, management and propaga¬
tion. Special Publication 7, North Central
Division, American Fisheries Society. 159 pp.
Pasch, R. W., P. A. Hackney, and J. A.
Holbrook. 1980. Ecology of paddlefish in
Old Hickory Reservoir, Tennessee, with em¬
phasis on first-year life history. Transactions
of the American Fisheries Society 109: 1 57—67.
Poff, R. J., and C. W. Threinen. 1965. Surface
water resources of Columbia County. Wis¬
consin Conservation Department, Madison.
Purkett, C. A. 1961. Reproduction and early
development of the paddlefish. Transactions
of the American Fisheries Society 90:125-29.
Ricker, W. E. 1975. Computation and interpre¬
tation of biological statistics of fish popula¬
tions. Bulletin 191. Fisheries Research Board
of Canada, Ottawa. 382 pp.
Rosen, R. A., and D. C. Hales. 1980. Occur¬
rence of scarred paddlefish in the Missouri
River, South Dakota - Nebraska. Progressive
Fish Culturist 40:82-85.
Rosen, R. A., and D. C. Hales. 1981. Feeding
of the paddlefish, Polyodon spathula. Copeia
1981:441-55.
Russell, T. R. 1986. Biology and life history of
the paddlefish - a review, pp. 2-20 in J. G.
Dillard, L. K. Graham, and T. R. Russell
(eds.), The paddlefish: status, management
and propagation. Special Publication 7,
North Central Division, American Fisheries
Society. 159 pp.
SAS (Statistical Analysis System). 1988. SAS/
STAT users guide, release 6.03 edition. SAS
Institute, Cary, North Carolina. 1028 pp.
Sparrowe, R. D. 1986. Threats to paddlefish
habitat, pp. 36-45 in J. G. Dillard, L. K.
Graham, and T. R. Russell (eds.), The
paddlefish: status, management and propaga¬
tion. Special Publication 7, North Central
Division, American Fisheries Society. 159 pp.
Southall, P. D., and W. A. Hubert. 1984. Habi¬
tat use by adult paddlefish in the upper Mis¬
sissippi River. Transactions of the American
Fisheries Society 113: 125-3 1 .
Stark, W. F. 1988. Wisconsin, river of history.
Published by the author, Thiensville, Wis¬
consin. 353 pp.
Unkenholz, D. G. 1986. Effects of dams and
other habitat alterations on paddlefish sport
fisheries, pp. 54-61 in J. G. Dillard, L. K.
Graham, and T. R. Russell (eds.), The pad¬
dlefish: status, management and propagation.
Special Publication 7, North Central Divi¬
sion, American Fisheries Society. 159 pp.
Wagner, G. 1908. Notes on the fish fauna of
Lake Pepin. Transactions of the Wisconsin
Academy of Sciences, Arts, & Letters 16:23-37.
WDNR (Wisconsin Department of Natural Re¬
sources). 1976. Wisconsin 1976 Water Qual¬
ity Inventory Report to Congress. Wiscon¬
sin Department of Natural Resources, Madi¬
son. 107 pp.
WDNR (Wisconsin Department of Natural Re¬
sources). 1988. Final environmental impact
statement. Proposed Lower Wisconsin State
Riverway. Wisconsin Department of Natu¬
ral Resources, Madison. 200 pp.
John Lyons is a fisheries research biologist with the
Wisconsin Department of Natural Resources. His
research focuses on the ecology and management of
fish in warmwater streams and rivers.
Volume 81 (1993)
135
Bruce T aylor
An “Education into Gladness
Ron Wallace's The Makings of Happiness
and “The Mid-life Progress” narrative
AS Director of Creative Writing at the University of Wis-
consin-Madison and Editor of the annual Bittingham
Award for poetry from the University of Wisconsin Press, Ron
Wallace, both as a practitioner and a promoter, has been for
many years a major voice of and for poetry in Wisconsin. As
author of three previous books of poetry, three books of criti¬
cism, and as editor of the anthology Vital Signs: Contemporary
American Poetry from the University Presses , Wallace’s national
reputation has been firmly established for quite some time.
However, with the publication of The Makings of Happiness,
Wallace has emerged squarely among the very best of the mid¬
generation poets in America.
The Makings of Happiness is a book of broad appeal that can
be enjoyed on many levels and from many different perspec¬
tives, a book for the general reader as well as the most seasoned
poetry aficionado. An hour, an afternoon, a day, or many days
can be spent wallowing in this book’s sumptuous pleasures. The
more a reader puts into it, the more he or she gets out of it — and
that, after all, is the hallmark of great literature.
To even the casual reader of Ron Wallace’s poems there is
much wit and music to be found, great humor and many words
at wonderful play. First, on this most available level, there is all
the beautiful noise, all the language which delights the reader by
delighting in itself.
TRANSACTIONS Volume 81 (1993)
137
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
I’m lifting the oysters
up from the ice chips,
scooping the slippery pap
loose with a spoon,
dripping the sliver of lemon, the ripe
island of tabasco, and then
flipping it all up to my lip and sipping
it in, the rough texture of shell
on incisor, the limp liquidy tongue
poised for the pleasure
of soft palate and swallow . . .
“Fresh Oysters & Beer”
Limp-wristed and slithery
she spins full around
and falls to the ground
dizzy, a fizzle.
“Wiffle Ball”
Breezy with bees
apple pulp rises under the grinder,
shreds of flesh and skin glistening,
the amber liquid dripping
into the tin acidic bucket.
“Apple Cider”
the tin stars pinned
in the tenpenny wind.
“February Thaw”
Even more simply, though no less ele¬
gantly, Wallace displays an impeccable dic¬
tion, illuminating again how the difference
between a word and the “right” word is, as
Mark Twain has always told us, the differ¬
ence between “lightning” and a “lightning-
bug.” Wallace gives us the “ rampantly adoles¬
cent daughter” of “Fresh Oysters & Beer”; a
“Puff nuthatch” and “roadside mud
quick with rivulets” in “February Thaw”; in
“Prayer,” “the wick of your own breath aflut¬
ter ',” as well as the exquisitely assonantial
Or the votive candles of snow
over which one crow,
cowled in its shadow, lengthens?1
Then there is the deft ability Wallace has
for reinventing the language with phrases to
which he has restored original meaning by
forcing them within the context of the poem
back through the colloquialisms, cliches and
euphemisms, the commonplaces of homely
discourse to reassume their original power as
metaphor. For example, in “Early Brass” the
brass section of the New York Cornet &
Sacbut Ensemble, “in long-tailed tuxedos /
rise to the bright occasion .” Marjorie, the
quadriplegic “information manager of a chin
operated wheelchair company,” composes in
“Fan Mail” a “chin-operated missive, / its five
good-tempered sentences / tapped out with
what intensive care . . . .”2
In “Basketball” the poet is shooting bas¬
kets with his six-year-old daughter who is
dawdling:
Hurry up, I shout.
We don’t have all day.
And we don’t.
The next time I look,
she’s sixteen . . .3
Wallace is capable of many wonderful
lines and of many master-crafted individual
poems. Yet, as enticing as it is to remain at
this level, if the reader considers The Makings
of Happiness merely as a collection and not
as a progression with a beginning, middle
and end, then that reader will miss the full
scope and profundity of what, when read in
sequence, reveals itself as a remarkable narra¬
tive — the slow, often sad, but ultimately
miraculous story of many of us.
With this book meticulously arranged in
three incremental sections, Wallace leads the
reader through a progression of poems that
ultimately achieves an equilibrium between
richness and loss, regret and ease. Through
the sequence of these wryly, often ironically
bemused, superbly wrought poems, Makings
1 38
TRANSACTIONS
TAYLOR: Ron Wallace and The Makings of Happiness
shares much with the recent American fic¬
tion described by Margaret Morganroth
Gullette in Safe at Last in the Middle Years as
“mid-life progress narratives ”(xiv). Gullette
voices the recognition that “rescuing them
[the protagonists/subjects] from . . . deple¬
tion seems to be one of the . . . unspoken
spiritual or ethical functions” (xiv); and that
these kinds of narratives utilize their “resis¬
tances, strengths, or sly timely weaknesses,
ingenious mental feints” to achieve a resolu¬
tion in which the author can begin even “to
think the kind of sentence” that would have
been impossible before (xiv.31). So, near the
end of Wallace’s book we find lines such as
How ease inhabits our lives.
“The Fox in the Berry Patch”
and in the same poem,
. . . O the gladness
that only a family understands . . .
“The Fat of the Land”
Standing as they do above, out of context
and in isolation, these lines could be dis¬
missed as glib and self-serving by a less than
attentive reader. But by the time the careful
reader gets to these lines, and the poems they
are part of, the poet has earned the right to
such statements. Though what damns us
saves us, and all that can save us from our¬
selves is time and change, here, according to
Gullette, there is “no necessary contradiction
between gladness and the self as it lives in
time.” Here then for both the poet and the
reader is an enticement into hopefulness, or
as Gullette calls it, an “education into glad¬
ness” (146).
If a man can’t be happy on a little farm
in Wisconsin,
he hasn’t the makings of happiness in
his soul.
Nick Englebert, artist-farmer,
1881-1962
The conditional “if’ of the title epigram,
with which Wallace’s book both begins and
ends, raises the central question of the collec¬
tion. The question is not, what is happiness,
not exactly; but more, of what and how is
happiness made? What, if you will, is the
recipe? What are the ingredients?
Early Brass
The poems of the first section, “Early Brass,”
are both strident and timid, full of failures
feared or perceived and all the initiatory
anxieties of youth “duped by dazzle and
subterfuge, / shimmer and flick” (“Blue-
gills”) where “desire and longing could in¬
spire / the unlikeliest situation” (“Love and
Sex”).4
Wallace seeks in the past (our memory,
our makings of the past) some distant proph¬
ecy of what the future has become. The
speaker, even in this first section, simulta¬
neously assumes the voice of both child and
father: the early awkward lover and practiced
husband, the tentative young poet looking
hopefully ahead and the master crafter look¬
ing back, however helplessly, towards the
past which in its own way seems as mysteri¬
ous now as the future did then.
Change and perspective are the major
ingredients introduced in the first section,
and time is the catalyst. The future from this
“early” view seems always “somewhere over¬
head, . . . / stretched out filmy and seductive”
(“Smoking”). The past, on the other hand,
can be realized only in retrospect as “a vast
doorway” (“Fan Mail”), “. . . one more /
phantasmagoric invention we use / to fool
ourselves into someone else’s shoes” (“Off
the Record”).
In “Speeding”:
Some damn fool kid
is racing his three-speed
down the hill in front of our house
Volume 81 (1993)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
and later,
spring flashing its bright green signals,
although, approaching fast from a
side street,
the future is looming,
preparing to barrel through.
Look out! I shout. But it’s too late.
The kid has been gone for years,
pedaling for all he was worth
into me, into you.
The events of these “early” poems, while
they were happening, seemed to be all
heading for the future / just as fast as the
mind can see” (“Camp Calvary”); yet the
same events seen in retrospect, the stance that
the mature artist is allowed by the art itself,
assume the slow motions of the inevitable, of
“. . . the ball slowly / rolling out the door / and
down these many years” (“Rebounding”).
The difference in perspective is the result of
the change time has brought to the poet, and
the attempt to fix the events themselves with¬
in the artistic frame of the poems.
Breakdown
The second section of the book, “Break¬
down,” is one of crises and passages, acci¬
dents and escapes, close calls and catastro¬
phes. The possibilities of luck and the fact of
evil itself must at least be “gotten by,” if not
“gotten through.”5 Again, it is the meliora¬
tions of art that make such a passage pos¬
sible.6 If “Turning Forty” (the title of one of
the poems) is “over the hill,” then being “At
Forty” (the title of another) is the hump; and
once again it is the act of the poems, the
learning of the makings of them, that allows
the “getting over.”7
“Breakdown,” the title and first poem of
this section, begins with that moment before
a seemingly inescapable accident which,
however, does not happen: “When you fi¬
nally know / you are not going to make it,”
and ends with an escape, with “. . . nothing
changed
except for this small uncertainty,
this fist in your gut, this sneer
twisting your lips, as if
you knew something true and awful,
something you could never, never
confess.
The accident which is ultimately avoided
is middle-age with its intimations of mortal¬
ity, its chances lost or not taken, with more
doors closed behind than ever able to be
opened ahead. As in the first section of this
book, the initial motion seems to be forward,
often out of control, as in “Hairpin”:
in the bright slow deceptions of the day,
you race toward that sudden
appointment
you never quite planned on,
the journey you didn’t mean to take.
Or, in “Headlines”:
. . . morning careening just around the
corner, your own spring flagging, the
drought burning on?
Yet, there is also a pause here, a slow
progression to the realization that we both
lose and gain as we grow, that, as is noted in
“Turning Forty”:
Time doesn’t speed up like a train
with somewhere to get to, railing
on that Augustinian straight line,
so much as it spirals, silent, circling
back on itself, an old dog, settling
on the same tired spot again . . .
1 40
TRANSACTIONS
TAYLOR: Ron Wallace and The Makings of Happiness
and that there is a certain joy to all this
when
In the country of stumble and drag
time settles, its bald tail a-wag.
In the first section of the book there were
poems such as “Early Brass,” “Rebounding,”
and, particularly, “” Birdsong Anyway” where
the subject was the poet early at his art, the
first few fumblings, the initial impulses which
brought the poet to that art at all. In the
second section, in poems such as the “Poetry
Report,” “State Poetry Day,” and “The Din¬
ner Party,” it is the distractions of the pseudo-
settled (the middle-aged, middle-class, and
middle-brow), the temptations of the hack
and the poetaster, that threaten the poet’s
progress towards the authentic.
In “State Poetry Day” (surely the most
cacophonous villanelle ever written), poetry
threatens not even to survive, as Auden once
demurred, “in the valley of its saying,” when,
“In the legislative chambers with their dactyls
and caesuras / the local poet laureates sing in
praise of cheese and beer,” and where
No one mentions Nicaragua, acid rain,
cocaine or Star Wars,
as the couplets and quatrains maintain
a pleasant atmosphere.
The mayor couldn’t be here, but he
sends his grand whereases.
Another year closes with a villanelle’s
razzmatazzes.
In the next poem, “The Dinner Party,” a
sestina, the poet, in another room, becomes
aware that
Everyone out in the living room’s
concerned
about groundwater and nuclear war,
their voices a warm glow in the dark,
the familiar country of engaging
party talk,
and I’m here in the kitchen with
my spinach
and my eggs, my vinegar and oil,
worrying
about how to make a salad. . . .
It seems, then, no accident that these
poems are followed by another pair, “The
Hell Mural: Panel I” and “The Hell Mural:
Panel II” which are also sestina and villanelle,
that profoundly and majestically address those
same subjects that the previous pair so trivi¬
ally if artfully avoid. The poet moves from
the light, almost parodic tone of the first pair
to the high seriousness, the elegiac thunder of
the second. (Both poems take as their subject
The Hell Mural, impressions of the Hiro¬
shima holocaust painted by Iri and Toshi
Maruki.)
Thus this section of the book ends with
two confrontations of an ultimate evil, each
in the strictest of forms, as if those restrictions
can contain the horror — the scope too large,
the cast too numerous in its sufferings, the
implications too gigantic to comprehend.
The Makings of Happiness
Is it escape then, that drives the poet to the
“small farm in Wisconsin” of the Engelbert
epigraph? Is it the world too much to handle,
the realization of the impossibility of hang¬
ing on to anything except yourself and your
own, to the few people that there is no doubt
you belong to, and to a place, however imper¬
fect, of your own making?
Yes, but not just. In this third and final
section, the poet, full in the flourish of his
talent, takes the way in to find the way out.
He attempts, as poets almost always have, the
ultimately universal through the deeply per¬
sonal, struggling though the only life each of
us has towards those other lives around us
Volume 81 (1993)
141
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
and toward the Life that surrounds and sup¬
ports us all: the life of nature and the life of
place, the life of family and work — which in
this case is the poet at his craft.
The place is, of course, essential to this last
section of the book. The farm, away from the
bustle of the city and distractions of a larger
society, allows the poet the time and space to
focus and concentrate — quite literally, to
see.
Far from the city with its bright
deceptions,
we lie down in the damp grass and
await whatever
blast of heat or radiance is on its way
to take us
out of ourselves into the future that
couldn’t be.
Tomorrow, the papers will claim
aurora borealis,
a rare display for those with dark
enough to see.
“Night in the Country”
The farmhouse itself, full of the immedi¬
ate family, when viewed from the outside on
a cold winter night, achieves the perspective
of rebirth and renewal:
. . . the warm house brightens
as if the click of the refrigerator
and the slow breathing of the children
could call up the whippoorwill,
cicada, spring peeper, and cricket,
could fill us beyond loss or doubt.
“February, Full Moon”
Family is key here. It is the “glue,” if you
will, that holds the place together. In “The
Fat of the Land” it is the extended family
gathered for a reunion primarily, it seems, to
eat, to partake of, not just food, but of each
other:
. . . one big happy family, back from
wherever we’ve spread ourselves too thin.
A cornucopia of cousins and uncles,
grand¬
parents and aunts, nieces and nephews,
expanding.
and later,
O the loveliness of so much loved flesh,
the litany of split seams and puffed
sleeves,
sack dresses and Sansabelt slacks,
dimpled knees and knuckles, the jiggle
of triple chins. O the gladness
that only a family understands . . .
and finally,
. . . huge and whole of this
simmering night,
battened against the small skinny
futures that must befall all of us,
the gray thin days and the
noncaloric dark.
Ultimately, however, it is the poet, and
the poet’s craft learned and practiced, that
provides the last ingredient for Wallace’s
recipe for happiness: to love and enjoy the
moment, the day, and the world, the people
who are yours in it, and to do good work.
In the first section of the book, in “Birdsong
Anyway,” the young poet attempts the meta¬
phor of poem as birdhouse and fails, at least
at the constructing of the house itself. In the
first poem of the third section, “Building an
Outhouse,” a wryly, nearly perfectly dis-
wrought sonnet which is also about “build¬
ing” poems, the poet can concluded with
confidence:
. . . it’s up! Functional. Tight as
a sonnet.
It will last forever ( or at least for awhile)
though the critics come sit on it, and
sit on it.
142
TRANSACTIONS
TAYLOR: Ron Wallace and The Makings of Happiness
How else to get through the day and the
life, both past and future, however tenuous,
that this day is a part of, but to bring it to the
poem and there allow the connections to be
made between the aging, changing self and
the family and place the self finds itself a part
of. The poem, of course, cannot capture the
moment, but can fix it and frame it, shape it
into one of those, as Wordsworth called
them, “spots in time,” or in Frost’s words,
“momentary stays against the confusion,” in
which the self, glad in the present, experi¬
ences grace.
This is, however, a grace more Frost than
Wordsworth, for it is a condition, though
perhaps given, only available to the poet
active and aroused at the makings of poetry.
The classic metaphor of Walt Whitman also
comes to mind here, of the “Noiseless Patient
Spider,” “isolated / Mark’d how to explore
the vacant vast surrounding, / It launched
forth filament, filament, filament, out of
itself’ compared to the Soul/Poet, “detached
in measureless oceans of space, / Ceaselessly
musing,
venturing, throwing, seeking the spheres
to connect them,
Till the bridge you will need be form’d,
till the ductile anchor hold,
Till the gossamer thread you fling
catch somewhere . . .
It is in the final, title poem, “The Makings
of Happiness,” that these connections are
most obviously made. The poem is based
upon a painting by Nick Englebert, “The
Photographer which also provides the epi¬
graph for this book. The literal subject of the
painting, as the title implies, is not primarily
a “small Wisconsin farm” but a visiting turn-
of-the-century French photographer taking
a picture of a Wisconsin farm family. This
was an event at that time of great wonder,
rarity and delight. The necessity of the role of
the artist then is inherent in the progressive
tense of the poem’s, and book’s, title, the
“Makings.”
It is not, as the poem tells us, “Until you
have looked at something so long / it grows so
familiar you can’t see it,” that you can “know
the soul’s work.” The expressive role of the
artist here — painter, photographer and
poet — is to reveal the miraculous in the daily,
homely and commonplace,
the alp that all but disappears in dailiness;
the sea that common routine conceals;
the little farm in Wisconsin that seems
painted in oil on your long picture
window . . .
and to create
the barn more like a hearth than a barn,
a mother, who could be your mother,
in the doorframe across the way,
bread in the oven and time on
her hands,
the little girl, who could be a boy,
roped to her calf, which could be a dog,
waving to her cat, which could be
a stoat,
apples in her cheeks and honey in
her hair,
the church in the permanent center,
the townspeople happy as larks . . .
What is important to keep in mind are the
layers of references the poem has established.
It is not the farm or the family itself, not the
photographer’s picture of it and them, not
even the painter’s picture of the photogra¬
pher taking his picture, but the poem of the
poet that pulls it all together and keeps
. . . the man floating, the girl
smiling, the calf changing, the cow
rolling
its eyes, the blue Frenchman tipping
his hat at you who live so far off
in the vanishing point of the future.
Volume 81 (1993)
143
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Notes
1 Italics in this paragraph are mine.
2 Italics in this paragraph are mine.
3 Italics are in the original.
4 “T o feel good about the middle years it is helpful
to have had a miserable young adulthood ...”
Gullette observes, while later adding that from
the advantage of the “later, . . . safer middle years,
. . . young adults [are regarded] with detached
sorrow, pity, and compassion — as if ... we now
understand how the young are obliged to navi¬
gate a perilous crossing with rudimentary equip¬
ment before getting to the other side” (6-7).
5 Suddenly we see what must be the basic psycho¬
logical situation, and it’s the same whether the
plot trouble is sex or parents or children or war.
Whatever it is, the characters fear that for some
reason they don’t understand, they can’t create a
self-chosen, more self-confident, happier future
— they can’t progress in the life course as they
must and want to.” For this reason, I believe,
dangerous-age novels contain more than their
share of shrieks, blows, accidents, and death”
(Gullette 15).
6“Meliorism is the narrative message of mid-life
Bildungsromane ” (Gullette 150).
7 “The ability to recognize that the middle years
can be welcomed as a relief by some people may
depend on our traversing the dangerous age imagi¬
natively, in however truncated and inevitably
detached away” (Gullette 18-19).
Works Cited
Gullette, Margaret Morganroth. Safe at Last in
the Middle Years: The Invention of the Mid-life
Progress Novel . Berkeley: University of Cali¬
fornia Press, 1988.
Wallace, Ronald. The Makings of Happiness. Pitts¬
burgh: University of Pittsburgh Press, 1991.
- . People and Dog in the Sun. Pittsburgh:
University of Pittsburgh Press, 1987.
- . God Be with the Clown : Humor in
American Poetry. Columbia: University of
Missouri Press, 1984.
- . Tunes for Bears to Dance To. Pittsburgh:
University of Pittsburgh Press, 1983.
- . Plums , Stones, Kisses, and Hooks: Poems.
Columbia: University of Missouri Press, 1981.
- . The Last Laugh: Form and Affirmation in
the Contemporary American Comic Novel.
Columbia: University of Missouri Press.
- . Henry James and the Comic Form. Ann
Arbor: University of Michigan Press, 1975.
Poet and Editor Bruce Taylor is Professor of
English at the University of Wisconsin-Eau
Claire. His most recent essays include “ The
Vietnam War Movie: Voyeur and Witness ”
and “Draft Dodger Rag” in The Highground,
published by the Wisconsin Vietnam Memorial
Project Lnc., and “Betrayed and Abandoned:
The Divorced Male in the Short Fiction of
John Updike ” in Wisconsin Dialogue.
144
Wisconsin Academy of Sciences, Arts and Letters
Executive Director LeRoy R. Lee
1993 Academy Council Officers
Daniel H. Neviaser, President, Madison
Robert P. Sorensen, President-Elect, Madison
Julie Stafford, Past- President, Chippewa Falls
FyOger Grothaus, Vice President-Sciences, Racine
- Gerard McKenna, Vice President-Arts, Stevens Point
Denise Sweet, Vice President-Letters, Green Bay
Gerd H. Zoller, Secretary/Treasurer, Madison
. i Councilots-dt-Large
LeefHalgren, Platteville
James S. Haney* Madison
L ~ Carl A. Weigell, Milwaukee
, Ody J. FishrPewaukee
Harry W. Fry, Bristol
Mildred Larson, Eau Claire
Judith L. Kuipers, La Crosse
Councilor-Jit-Large Emeritus
John Thomson, Mt. Horefcr
Your membership will encourage research, discussipn
and publication in the sciences, arts and letters of
' Wisconsin.
Wisconsin Academy of Sciences, Arts and Letters
1 922 University Avenue
Madison, Wisconsin 53705
Telephone (608) 263-1692
1 Wisconsin Academy of Sciences, A rts and Letters
1922 University Avenue
Madison, Wisconsin 53705
ISSN 0084-0505 Telephone (608) 263-1692
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of the Wisconsin Academy of Sciences, Arts and Letters
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Transactions of the Wisconsin Academy of Sciences, Arts and Letters
DEPPE and LATHROP: Aquatic macrophyte community of Lake Mendota
Uncommon Species
In the three survey years, six species were
found very infrequendy and at very low den¬
sities in Lake Mendota: Potamogeton crispus
L., P. richardsonii (A. Benn.) Rydb., P.
zosteriformis Fern., P. foliosis Raf., Nelumbo
lutea (Willd.) Pers., and Nymphaea tuberous
Paine (Table 1). In 1989 and 1990, P.
crispus was found at low densities on
transects off die eastern and northern shore¬
lines, while in 1991 it was found almost ex¬
clusively along the west and south shorelines.
In all three years, P. richardsonii was found
in moderate abundance at transect 45 and
more sparsely at a few other transects dur¬
ing 1989-91. For all three years, the two
floating-leaved lily species Nelumbo lutea and
Nymphaea tuberosa grew densely at^l .0 m
in University Bay at transect 39, and else¬
where in the bay at varying densities from
1989-91. These species were also found
along the east shoreline at transect 47 in
1989. P. foliosis was found at scattered sites
in 1990-91, and P. zosteriformis •wsls found
at only one station in 1989.
Depth Limit of Growth
The depth limit of plant growth in Lake
Mendota was somewhat variable between
transects but generally occurred between 3.0
and 4.0 m for 1989-91 (Table 3). Depth
limits shifted to slightly shallower ranges in
1990. but returned to ncar-1989 levels in
1991. Depth limit decreased by 0.5 m from
1989 to 1990 at all transects in University
Bay and along the northwest shoreline where
plant growth was densest.
Depth limit of plant growth in Lake
Monona generally occurred at 3.5 m in
1990, but at 2.5 m in 1991. Similarly, in
Lake Waubesa these figures went from 3.0
m in 1990 to 2. 0-3.0 m in 1991. In Lake
Kegonsa, depth limits increased from 2.0-
2.5 m in 1990 to 3.0 m in 1991.
Because most plant growth occurred at
£3-4 m in Lake Mendota, it is noteworthy^—
that 10% of stations £.3 m in 1989 were de-
void of plants, while 20% had no vegetation
in 1990 and 1991. In 1990, 7% and 1% of
stations £3 m were without vegetation in 4 —
Monona and Waubesa, respectively, but this
increased to 30% and 48% for the two lakes
in 1991. In Lake Kegonsa, 58% of stations
£3 m were without macrophytes in 1 990, ^ —
but only 21% in 1991.
Discussion
Depth Distribution
In our 1989-91 surveys in Lake Mendota,
macrophytes were found almost entirely at
water depths between 0.5 and 3.5 m, while
certain depths favored growth of particular
species. Ceratophyllum and Myriophyllum spi-
c a turn, tall -growing plants with biomasses
heaviest near the water surface, grew most
densely between 2.0 and 3.0 m. The com¬
mon species, tending not to grow as tall,
were found largely between 0.5 and 2.0 m,
where they can receive adequate light. Their
infrequent occurrence in water depths >2.0
m suggests that they may be shaded by al¬
gal blooms and dense growths of Cerato¬
phyllum and Myriophyllum. Lack of macro¬
phyte growth at the 0.5 m contour is prob¬
ably due to one or more of the following rea¬
sons: rocky substrate, more pronounced
wave action, ice shifting in winter, and the
controlled lowering of the lake level over the
winter months.
Macrophyte Community of University Bay
Since the 1960s
Myriophyllum spicatum dominated the plant
community from its introduction in the
Volume 81 (1993)
(over)
55
r ■ it\ a -i TP A/" — ^ | (t/atl TO of the Wisconsin Academy
J_ KAJN jAvo JL 1L/1N J of Sciences, Arts and Letters
Volume 82 9 1994
Editor
Managing Editor
Intern
William J.Urbrock
Department of Religious Studies
University of Wisconsin Oshkosh
Oshkosh, Wisconsin 54901
Patricia Allen Duyfhuizen
328 West Grant Avenue
Eau Claire, Wisconsin 54701
Christopher J. Solberg
1 ransactions welcomes articles that explore features of the State of
Wisconsin and its people. Articles written by Wisconsin authors on
topics other than Wisconsin sciences, arts and letters are occasionally
published. Manuscripts and queries should be addressed to the editor.
Submission requirements: Submit three copies of the manuscript,
double-spaced, to the editor. Abstracts are suggested for science/
technical articles. The style of the text and references may follow that
of scholarly writing in the author’s field, although author-year citation
format is preferred for articles in the sciences, author-page number
format for articles in the humanities. Please prepare figures with
reduction in mind.
© 1994 Wisconsin Academy of Sciences, Arts and Letters
All rights reserved
ISSN 0084-0505
For information on membership in the Academy,
call (608) 263-1692.
Contents
TRANSACTIONS
Volume 82 • 1994
From the editor v
Development of Brussels Hill Pit Cave , Door County , Wisconsin:
Evidence from flows tone and sediments 1
Jim Brozowski and Michael J. Day
Brussels Hill Pit Cave is the deepest vertical cave in Wisconsin and contains a valuable
suite of historic and prehistoric sediment, faunal, and floral assemblages. The sediments
at the -15 m level appear to be post-glacial in age.
Occurrence and significance of sea lamprey
(Petromyzon marinus) in the lower Fox River , Wisconsin 17
Philip A. Cochran
Improved water quality led to concern about the potential for sea lampreys to ascend the
Fox River and gain access to the Lake Winnebago watershed. The controversial closure
of the Rapide Croche lock in 1988 has been justified by subsequent collections of sea
lampreys in the lower river.
Observations on the white perch (Morone americana)
early in its invasion of Wisconsin 23
Philip A. Cochran and Peter J. Hesse
White perch are native to the Atlantic Coast but have recently invaded the Lake
Michigan drainage in Wisconsin. This paper documents the dramatic increase of white
perch in the Fox River and discusses various aspects of their biology.
History of the fishes of the Bois Brule River System , Wisconsin ,
with emphasis on the salmonids and their management 33
Robert B. DuBois and Dennis M. Pratt
The Bois Brule River has had a rich and colorful history from its early days as an
important water route to its present status as one of Wisconsin’s premier trout streams.
The authors present an historical sketch of the fishes of the river system along with the
factors that have affected them over the last two centuries.
Social behavior of adult jaguars (Panthera onca L.)
at the Milwaukee County Zoo 73
Thomas F. Grittinger and Deborah L. Schultz
Social behavior between two jaguars was described and analyzed in a four-year study at
the Milwaukee County Zoo.
Vesicular-arbuscular mycorrhizal fungi of Wisconsin s sandy soils 83
Richard E. Koske and Leonard L. Tews
Sandy soils of Wisconsin were surveyed for the first time for vesicular-arbuscular
mycorrhizal (VAM) fungi.
The plant communities of Nine-Mile Island — past and present 89
David J. Post
The article details a brief history of the vegetation and human impact upon Nine-Mile
Island since the original land survey of the 1840s and presents a phytosociological
analysis of the present vegetation patterns and community distribution. Current
successional trends are examined to predict possible future vegetation changes on the
island with a call to maintain the island in its present state for the enrichment of future
generations.
Analysis of black bear habitat in northeastern Wisconsin 109
Keith T. Weber
The article reports the results of field studies and roadside counts used to analyze black
bear population and habitat in northeastern Wisconsin. An earlier version of the paper
was awarded the Forest Stearns award for excellence in the biological sciences at the
WASAL annual conference in 1992.
Errata
Transactions of the Wisconsin Academy of Sciences, Arts and Letters
Volume 81, 1993
Because of a change in font made by the printer at the final typesetting, the < symbol
was lost, causing a major problem in data interpretation on page 5 5 of the article “Recent
changes in the aquatic macrophyte community of Lake Mendota,” by Elisabeth R.
Deppe and Richard C. Lathrop. For a copy of the corrected page, please contact the
Wisconsin Academy.
IV
From the editor
/\u of us who read Transactions , whether for the first time or
as old friends well acquainted with this journal, cannot help
but notice that the content of articles published year after year
leans heavily toward the natural sciences. While Transactions
extends a broad invitation welcoming scholarly articles “that
explore features of the State of Wisconsin and its people” and
while the Wisconsin Academy is devoted to arts and letters as
well as to the sciences, it is a fact that Transactions has been a
publishing vehicle of choice for those who write about the land
and waters of Wisconsin, its geology, geography, and the
diversity of its life forms. This is explainable in some measure
insofar as historians, artists, poets, and creative writers often
choose to write for our sister publication, the Wisconsin
Academy Review , a journal of Wisconsin culture.
This 1994 issue of Transactions is no exception to the rule.
Some of our articles feature studies of jaguars at the Milwau¬
kee Zoo and black bear habitat in northeastern Wisconsin,
the plant communities of Nine-Mile Island in the Chippewa
River and the mycorrhizal fungi of Wisconsin’s sandy soils,
and a look at the Brussels Hill pit cave in Door County.
Other articles explore the occurrence of sea lamprey in the
lower Fox River, the recent invasion of white perch into the
Fox River, and the history of the fishes of the Bois Brule River
system. Together these many fascinating studies alert us to
the changing natural environment of Wisconsin and to the
ongoing parade of “visitors” who keep coming to our state,
on their own (like the white perch and sea lamprey) or not
on their own (like the jaguars in the zoo).
As I write this column in mid-August, I am preparing to
teach an adult-education class next week at The Clearing, one
of my favorite Wisconsin institutions, located in a spectacu¬
lar natural setting on the bluffs above Ellison Bay. Currently
an affiliate of the Wisconsin Academy, The Clearing was
founded by the famous landscape architect and environmen¬
talist Jens Jensen back in 1935 as a sort of outdoors “school
of the soil.” Over the years it has evolved into an indepen¬
dent association, supported by its many members and friends.
During its summer residential program, The Clearing offers
one-week courses on nature, the arts and humanities.
v
My course will be on “The Poetry and
Message of the Psalms.” Among the Psalms
that will certainly feature prominently in our
discussions will be Psalms 8 and 104. I ex¬
pect some lively and interesting debate on
whether or not these ancient religious poems
have anything to say about current rela¬
tionships between humanity and the natural
environment. In my mind there is no doubt
that, if the writers of these two psalms could
be transported forward to our own day and
could read the sort of selections contained
in this current issue of Transactions, they
would be confirmed anew in their appre¬
ciation for the world of nature, their sense
of awe before its vast sweep and their feeling
of interconnectedness with all its creatures.
Perhaps, if he could also be educated into
the momentous ecological developments of
our time — the Biology Department at UW
Oshkosh teaches about “Ecosphere in
Crisis”! — the poet of Psalm 8 would wish
to reconsider or at least nuance the idea that
humans are to exercise “dominion” over all
other living things. Then again, if Hebrew
Bible scholar James Limburg is on the right
track in his reading of these two psalms,
their ancient composers might ask us with
deepened urgency, “Who cares for the
earth?” “Who loves and treats gently this
place that is home to so many relatives,
human and non-human?”
It is clear to me, through my correspon¬
dence with our authors in this issue of Trans¬
actions and with the many fine reviewers
who offered them valuable professional criti¬
cism and advice prior to publication, that
there are many who undertake their scien¬
tific research precisely because they care for
the earth. As a member of the Wisconsin
Academy who has spent a lifetime of study
and teaching in the humanities, I took great
interest in reading these several articles de¬
voted to the natural sciences and in discov¬
ering more about all the life around me in
Wisconsin. As a biblical scholar, I found ex¬
tra delight in playing off these articles against
the appreciation for nature evident in some
of the Psalms. I hope all of you, our read¬
ers, whether you lean more towards the sci¬
ences, arts or letters, will find similar enjoy¬
ment in reading the 1994 Transactions.
Now, as I look ahead to selecting manu¬
scripts for future issues of Transactions , it is
my pleasure to repeat the invitation for
scholarly research and criticism on all aspects
of science, arts and letters featuring the state
and people of Wisconsin.
Bill Urbrock
The Wisconsin Academy of Sciences, Arts and Letters was
chartered by the State Legislature on March 16, 1870, as a
membership organization serving the people of Wisconsin. Its
mission is to encourage investigation in the sciences, arts and
letters and to disseminate information and share knowledge.
vi
Jim Brozowski and Michael J. Day
Development of Brussels Hill Pit Cave,
Door County, Wisconsin: Evidence from
flowstone and sediment
Abstract Brussels Hill Pit Cave is a joint-controlled vertical cave developed
to a depth of 28 m in the Silurian- age Niagara Dolomite of the
Door Peninsula in northeastern Wisconsin. Sediments and flowstones
in the cave are post-glacial, with deposition beginning around the
end of the Greatlakean sub stage, approximately 10,000 B.P. The
cave sediments differ both physically and mineralogically from those
on the surface. The cave is potentially older than the sediment infill
and flowstones, possibly having formed in the late Ce no zoic. Dat¬
ing of cave flowstones using paleomagnetic and radioisotopic tech¬
niques suggests that initial deposition of these formations occurred
approximately 11, 000 B.P. From dissolution rate calculations
(Palmer 1980) and paleomagnetic evidence we infer that the cave
itself developed contemporaneously with sediment and flowstone
deposition. The cave also contains significant early Holocene mam¬
malian remains which are currently under investigation (Kox 1988).
The Door County karst landscape is one of the few gla¬
ciated karst areas in the United States. Brussels Hill Pit
Cave is a vertical cave formed along a dissolutionally widened
joint in an outlier of the Niagara Dolostone Escarpment (Figs,
la and lb). Such caves are an integral part of the karst terrain
of the Door Peninsula (Rosen 1984; Stieglitz 1984; Rosen et
al. 1989; Johnson and Stieglitz 1990; Rosen and Day 1990).
Recent paleontological work (Kox 1988; Robert Howe, pers.
comm. 1 992) has focused on a rich faunal suite within the cave.
Previously excavated organic sediments from 28 m depth have
been 14C dated at 671 and 1820 B.P. (Howe, unpubl. data).
TRANSACTIONS Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
The primary objective of this study was
to establish the chronology of sediment
deposition within the cave. Determining the
age of the flowstones and the source of the
sediments would also provide information
about the age and development of the cave
itself.
Clastic sedimentary deposits and flow-
stone in caves are a tool for reconstructing
the climatic history and geomorphic evolu¬
tion of karst terrains (Milske et al. 1983).
Lively (1983), Milske et al. (1983), and
Lively et al. (1984) have presented flowstone
and flowstone-sediment chronologies based
on Uranium-series disequilibrium dating
and have demonstrated that the flowstone
deposition rate was significantly reduced
during glacial periods in southeastern Min¬
nesota. Gascoyne (1977) determined that, in
general, speleothem deposits represent rela-
2
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BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
tively warmer paleoclimatic periods, al¬
though speleothem growth may not always
directly reflect surface conditions. Flowstone
dating allows for a minimum estimation of
cave age as the flowstone must be younger
than the cave itself.
Cave formation age can also be estimated
using carbonate dissolution rate calculations
(Palmer 1980). Maximum rates under
phreatic conditions are roughly 0.14 cm/yr
or about 3 m every 1000 years. Extrapolat¬
ing this rate calculation to vadose cave de¬
velopment, the 28 m depth of Brussels Hill
Pit Cave suggests a maximum age of about
10,000 B.P., which coincides approximately
with the deglaciation of the Door Peninsula.
The research presented here is used to test
a developmental hypothesis for Brussels Hill
Pit Cave and the associated sediment depo¬
sition. Based on Ford’s (1977) four-part
classification of glaciated karst areas, the cave
is glaciokarstic or more specifically, karsti-
glacial, i.e., a karst process that has accen¬
tuated jointing which originally had resulted
from glacial loading on the bedrock surface.
Glaciokarst reflects the cumulative effects of
karst formation and glacial activity (Ford
1977). We hypothesize that Brussels Hill Pit
Cave is karstiglacial, based on the glacial his¬
tory of the Door Peninsula (McCartney and
Mickelson 1982). Cave formation com¬
menced after glacial loading and unloading
had accentuated bedrock jointing. Subse¬
quent dissolution enlarged the joints and led
to sedimentation followed by speleothem
deposits.
Regional Geology and
Geomorphology
The Door Peninsula is primarily an upland
ridge with morphology controlled by the
Niagaran Dolomite cuesta (Sherrill 1978).
The bedrock geology of the Door Peninsula
is outlined by Chamberlin (1877), Thwaites
and Bertrand (1937), Klussendorf and
Mikulic (1989), and Stieglitz (1984, 1989).
The Niagaran Series is approximately 107 m
thick and includes, from oldest to youngest,
the Burnt Bluff Group, the Manistique Do¬
lomite, and the Engadine Dolomite (Sherrill
1978; Stieglitz 1989). The rocks are mainly
dolostones, thinly bedded in the lower for¬
mations and thinly to massively bedded in
the upper formations. The rocks are fossil-
iferous, medium to coarse grained, and
Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
BRUSSELS HILL PIT CAVE
ALONG JOINT AXIS
o ^ ( 5 ' t 10 t | | 15 | ^ 20
Meters
p||j Inferred Sediment Remaining □ Sediment Removed Inferred Boundary
Map by Norbert H. Kox, 1988.
Fig. 2. Planar view of Brussels Hill Pit Cave
4
TRANSACTIONS
BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
mostly buff gray colored. Dip of the bed¬
rock is less than one degree to the southeast.
The Niagaran rocks form the Niagara Es¬
carpment along the western edge of the
Door Peninsula in Door County. West of
the escarpment are several outliers of the
Niagara Formation including Brussels Hill
(Figs, la and lb). Brussels Hill is a glaciated
but erosionally resistant biohermal (reefal)
structure; erosional resistance is attributed to
the unstratified reef groundmass (Stieglitz
1984) . The base of Brussels Hill is at 215
m; the summit is at 260 m.
Glaciation of the Door Peninsula oc¬
curred most recently during the Wood-
fordian (22,000 to 13,000 B.P.) and the
Greatlakean (11,500 to 9500 B.P.) advances.
These were separated by the Twocreekan
interstadial between 13,000 and 1 1,500 B.P.
(McCartney and Mickelson 1982). De¬
glaciation of the Door Peninsula coincided
with the general retreat of the Lake Michi¬
gan and Green Bay lobes of the Laurentide
Ice Sheet (Bryson et al. 1969; Hansel et al.
1985) .
Although there were at least three major
stages of Pleistocene glaciation in Wiscon¬
sin, it is not known specifically how fre¬
quently and for what duration the Door
Peninsula was over-ridden by ice. Thus it is
not known how many times karst processes
on the Peninsula were disrupted.
Brussels Hill Pit Cave
The sinkhole entrance to Brussels Hill Pit
Cave is at approximately 256 m above mean
sea level and opens into a vertical drop of
28 m (Figs. 2 and 3). Brussels Hill Pit Cave
is the deepest cave in Wisconsin and is de¬
veloped along a joint oriented approximately
62 degrees east of north. Other prominent
joint sets on Brussels Hill are at 25 and 155
degrees east of north (Rosen 1984). On the
basis of altitude, location, and bedrock char¬
acter, the cave is probably developed in the
Manistique Dolomite.
In horizontal cross-section, the cave con¬
sists of two crude ellipsoids along the cave’s
vertical axis (Fig. 4). The three levels are ap¬
parently vadose, showing no evidence of
phreatic development. Cave walls at the
middle and lower levels are covered by flow-
stone drapery starting at bedding plane seep¬
ages 6 m below the cave entrance and ex¬
tending to -28 m. Drapery samples for
paleomagnetic determinations reported in
this paper were collected at -15 m (Fig. 2).
Prior to June 1986, only the cave above the
-15m level was known; excavations revealed
a lower cave that descended to -28 m. Al¬
though some sediments have been removed
from the lowest level, this study focuses on
sediments at the -15 m level.
Paleontological analyses of the faunal re¬
mains from the -15 m level have identified
short- tailed shrew ( Blarina brevicus ), com¬
mon shrew ( Sorex sp.), little brown bat
[My otis licifugus ), white-tailed deer ( Ode-
coileus virginiana ), black bear ( Ursus ameri-
canus ), beaver ( Castor canadensis ), muskrat
( Ondatra zibethicus) and otter ( Lutra
canadensis) (Howe, unpubl. data).
At -15 m, a flowstone ledge has formed
over clastic sediments (Fig. 5). The flow-
stone consists of several layers of calcite
mixed with bones, leaves, wood, fine organic
material, and clasts of varying size and lithol¬
ogy. The ledge is approximately 1 5 cm thick
and extends laterally for 150 cm to the
southwest. It dips and pinches out near the
lower room opening. The clastic sediments
beneath the composite layer consist of lay¬
ers of fine sand and silt about 25 cm thick.
Based on their graded deposition and mixed
igneous, metamorphic, and sedimentary
mineralogy, the sediments are probably re¬
worked surface glacial deposits. Samples for
Volume 82 (1994)
5
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
BRUSSELS HILL PIT CAVE
PERPENDICULAR TO CAVE AXIS
Meters
Map by Norbert H. Kox, 1988.
Fig. 3. Perpendicular cross-section of
Brussels Hill Pit Cave
particle size and paleomagnetic analyses were
removed from this relatively inorganic layer.
Although some of the fine sandy and silty
cave sediment is laminated, the laminae
probably result from sediment reworking
within the cave.
Below the fine sand layer is a variegated
layer of coarse sand and fine gravel also
about 25 cm thick (Fig. 5). This deposit in¬
cludes several layers of coarse sediment ce¬
mented by crystalline calcite precipitate. The
coarse basal sediments probably predate ini¬
tial calcite deposition.
Beneath the cemented coarse sediment is
a clay loam layer (Fig. 5) which contains sev¬
eral striated clasts and large (>30 cm) pieces
of dolostone breakdown. Several of the clasts
are well rounded, polished, and fluted indi¬
cating glacial transport.
In order to elucidate the relationship be¬
tween surface soils and cave sediments, two
soil pits approximately 15 m north and 15
m south of the cave entrance were opened,
sampled, and analyzed. The Namur silt loam
is the dominant regional soil (Link et al.
1978).
Methodology
Analytical Transmission Electron Micros¬
copy (TEM) and Energy Dispersive X-Ray
Analysis (EDXA) were used for visual and
elemental analysis of the soils and sediments.
Soil and cave clay mineralogy was deter¬
mined using standard X-ray diffraction at
the University of Wisconsin-Milwaukee.
Soil particle size determination and pH mea¬
surement of the cave sediments and surface
soils were performed using standard hy¬
drometer methods and a calibrated labora¬
tory pH probe.
Two flowstone cores were extracted for
analysis from drapery on the north wall of
the cave; each was physically abutting the
Niagaran bedrock and retained a trace of
dolostone patina when removed. Although
there is no way to obtain perfect rotational
alignment, every effort was made in the field
and laboratory to maintain the closest pos¬
sible alignment. Sediment cubes were ob¬
tained similarly.
6
TRANSACTIONS
BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
BRUSSELS HILL PIT CAVE
TOP VIEW
/
N
0 5 10 15 20
I _ I _ I _ I _ I _ I _ I _ I _ J _ I _ I _ I _ I _ _J _ I _ I _ I _ I _ 1 _ I _ I
Meters
Map by Norbert H. Kox, 1988.
Fig. 4. Vertical top-view of Brussels Hill Pit Cave
Volume 82 (1994)
7
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
8
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BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
Paleomagnetic analysis was performed on
a Superconducting Technology model Cl 02
two-axis cryogenic magnetometer. The natu¬
ral remnant magnetism (NRM) of each
sample was measured first, then each was
demagnetized using alternating fields
stepwise up to 100 millitesla (25 to 1000
oersted). Sample remanence was measured
after each demagnetization step; moment
directions and intensities were calculated uti¬
lizing Fischer (1953) statistics. The paleo¬
magnetic data for the cave were matched
with a master curve generated from data sets
for the Great Lakes area (Creer and Tu-
cholka 1982; Kean and Klebold 1981).
Results
The Namur silt loam horizons are composed
of the major elements aluminum, silicon,
potassium, and iron, and the minor elements
calcium, magnesium, and zinc. Clay miner¬
als include illite, smectite, and chlorite. Soils
sampled near the bedrock contact contain a
high percentage of sand-sized grains (Fig. 6).
The sample from 40 to 56 cm in Soil Pit
2 contained smectite (montmorillonite) clays.
Smectite forms in neutral to alkaline environ¬
ment containing relatively high concentra¬
tions of calcium and magnesium (Brady
1974; Birkeland 1984). The Niagaran Do¬
lomite, CaMg(C03)2, has sufficient calcium
and magnesium for smectite clay formation.
Cave clay loam analyses indicate the pres¬
ence of silicon, iron, potassium, aluminum,
calcium, magnesium, and zinc. Minerals in¬
clude calcite, dolomite, quartz, and the clay
minerals kaolinite and illite. Kaolinite is of
interest because it forms primarily by the
complete weathering of alkaline feldspars,
through hydration at low pH (Klein and
Hurlbut 1985). Generally, illites are the
most common clay minerals in caves (Bull
1983; Sweeting 1973); they are formed by
the loss of potassium and possibly aluminum
from the mineral layers through hydration.
The Namur soil pHs vary from slightly
acidic to neutral, pH 5.0 to 7.0, while the
cave clay-loam is relatively alkaline, at pH
8.3. This suggests that the solvent capacity
of percolating surface water, in contrast to
Barden (1980), is not effectively neutralized
by carbonate clasts, at least in this location.
The pH of each soil horizon is moderated
by proximity to the dolostone bedrock (Fig.
7). Contemporary dissolution may thus be
expected on dolomitic bedrock or within
caves beneath the Namur silt loam. No gla¬
cial striae were observed at either of the test
pit soil/bedrock contacts beneath the Namur
silt loam, and no striae were detected on any
bedrock exposures examined in the imme¬
diate area. A sample of seepage water from
the -15 m level had a hardness of 130 ppm
(mg/L) total calcium and magnesium.
Paleomagnetic analysis results were cor¬
related with the type-curve data provided by
Creer and Tucholka (1982). Flowstone core
inclination and declination values were 58°
and -60°, respectively, indicating deposition
between approximately 9830 and 11,260
B.P. For the cores, inclination correlations
occur about 9700, 10,200, and 1 1,000 B.P.
and declination correlations between 9000
and 11,260 B.P. (Figs. 8 and 9). Sediment
cube correlations (declination and inclina¬
tion values of 13° and 70°, respectively) oc¬
cur several times for declination at about
7000, 9000, and 10,000 B.P.; inclination
correlations occur at about 7000, 9000, and
9800 B.P.
Flowstone samples submitted to the Min¬
nesota Geological Survey for U-series dating
proved too porous and chalky for a reliable
analysis. During a previous flowstone dat¬
ing attempt the isotopic results showed evi¬
dence of post-depositional alteration in the
calcite. The uranium concentration was very
Volume 82 (1994)
9
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
low (0.08 ppm) and the 230Th/234U activity
ratio was greater than unity. Chemical re¬
covery of Th from the sample was less than
5% (Richard Lively, pers. comm. 1992).
Discussion
The area around Brussels Hill Pit Cave is a
glaciated karst terrain in which karstiglacial
or post-glacial development of karst, sedi¬
ment, and soil is related to the effects of
Wisconsinan glaciation. The cave serves as a
depository for allochthanous organic and
mineral debris brought in directly through
the cave opening and indirectly through per¬
colation.
There are few mineralogical similarities
between the surface soil horizons and the
cave clay-loam; minerals notably present in
the Namur soils and very limited in the cave
clay-loam are zinc and iron. Elements
present in the cave clay-loam and not in the
soil are calcium and magnesium. The mag¬
nesium and calcium in the cave clay loam
are probably derived by dolomite dissolu¬
tion. Chemical analyses of the dolostones ap¬
proximates a 1:1 (Ca:Mg) ratio (Johnson
and Stieglitz 1990). Although the specific
10
TRANSACTIONS
BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
Soil pH
Fig. 7. Soil pH values
mechanism of cave clay deposition is not
known, it may predate the surface soil for¬
mation, having been deposited either as loess
or in a slurry during deglaciation.
Mineralogically, illite in the cave and
smectite at the surface are not unexpected.
Illite is the dominant clay mineral in un¬
weathered till and loess as well as the most
common clay mineral in caves (Bull 1983;
Sweeting 1973); it forms under leaching
conditions with high K concentrations.
Small amounts of kaolinite and smectite may
also appear (Grim 1968). Smectite forma¬
tion may be related to tundra or boreal cli¬
mates in the Door Peninsula during glacial
recession.
1 1
Volume 82 (1994)
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Kaolinite and illite genesis are dissimilar,
the former being formed under acidic con¬
ditions and the latter under alkaline condi¬
tions. Kaolinite formation could not have
occurred in the cave under current pH con¬
ditions. The kaolinite may have been eluvi-
ated from the overlying Namur soil where
it developed at a lower pH. Alternatively,
kaolinite may have developed within an ear¬
lier pre-Late Wisconsinan soil profile, or it
may have been derived from a nearby bog,
marsh, or wetland. Illite probably developed
in situ by the alteration of alkaline feldspar
or mica under alkaline conditions.
In terms of particle-size distribution, only
the lowest horizon of Soil Pit 2 is similar to
the cave clay loam (Fig. 6). The increase in
grain size with depth in the lowest horizon
of Soil Pit 1 may be attributed to mechani¬
cal illuviation but is more likely to reflect
granular material incorporated from disso¬
lution of the dolomitic bedrock.
Since the cave clay loam is near the bot¬
tom of the cave sediment profile, it was ei¬
ther deposited before the rest of the sedi¬
ment in the profile or it represents fines
eluviated from overlying cave sediments.
The relocation of coarse sand and fine gravel
fractions probably occurred prior to the
flowstone deposition. The origin of the
coarse sand and fine gravel has not been de¬
termined, but these may have been intro¬
duced during deglaciation. Soil particles may
have moved from the surface into the cave
along joints and other fractures, with the
joint width acting as a natural sieve, screen¬
ing out larger particles. As Rosen (1984)
states, the thin surficial deposits facilitate
seepage concentration at or along joints and
joint intersections. Karstic development is
accelerated along the joints as is alloch-
thanous sediment movement into the sub¬
surface.
Based on paleomagnetic results, the flow-
stones were determined to have formed be¬
tween approximately 9000 and 1 1,000 B.P.
(Figs. 8 and 9). Although the established
date is relative, there is a clear correlation
with the results of previous Great Lakes
paleolimnetic studies (Vitorello and Van der
Voo 1977; Creer and Tucholka 1982) and
the deglaciation of the Door Peninsula
(Bryson et al. 1969).
Upper portions of the flowstone incorpo¬
rate numerous bones, while the wall flow-
stones contain either burned or chemically
reduced wood fragments. Howe (pers.
comm. 1992) suggests that the wood rem¬
nants may be from the 1871 Peshtigo Fire
which burned portions of the Door Penin¬
sula. Because the flowstone formation is
stratigraphically superior to the mineral sedi¬
ments, it is assumed that the latter were de¬
posited prior to flowstone deposition, dur¬
ing or immediately after glacial retreat. It is
not known whether the coarse sand and fine
gravel are an initial deposit or the result of
sediment reworking. The intrinsic variety
and morphology of the granular cave sedi¬
ments indicate reworking and relocation by
surface runoff. Whether this was because of
historical near-ice meltwater run-off or con¬
temporaneous temperate run-off is unclear;
however, the lack of organic material and the
flowstone shelf with subordinate striated
clasts strongly suggest the former. As the
flowstone is resting on the sediment and ap¬
pears to have remained physically and
stratigraphically stable and is probably less
than 10,000 years old, the underlying sedi¬
ment tentatively correlates to the Great-
lakean substage between 9000 and 10,000
B.P. Similar stratigraphic sedimentary pro¬
files have been described in Mystery Cave
in southeastern Minnesota (Milske et al.
1983).
1 2
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BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
Inclination
40° 60° 80°
14
Fig. 8. Inclination type curve for east-
central North America. Brackets indicate
inclination value of 58° for flowstone cores
which correlates to 9700 years B.P.
(Modified from Creer and Tucholka 1982)
Declination
50° W 0° 50° E
12
Fig. 9. Declination type curve for east-
central North America. Brackets indicate
inclination value of 58° for flowstone cores
which correlates to 9700 years BP.
(Modified from Creer and Tucholka 1982)
Volume 82 (1994)
13
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Conclusion
Based on our interpretation and following
Ford’s (1977) classification, Brussels Hill is
a karstiglacial topographic feature. We fur¬
ther interpret the topographic morphology
of the area as glacial in origin with subse¬
quent post-glacial karstic modification.
Jennings (1985, 239) defines karstiglacial
forms as “forms thought to be virtually en¬
tirely glacial but [upon which] karst drain¬
age characteristics have been superimposed.”
Brussels Hill may be a relict karstiglacial fea¬
ture, having been formed during or prior to
Wisconsinan glaciation with subsequent for¬
mation of the pit cave during interglacial
conditions.
Interglacial or post-glacial warming pro¬
duced meltwater which was responsible for
sediment transport into the cave. Increased
water flow would have enhanced dissolution
of the Niagaran Dolomite. Flowstone and
sediment ages in this report correlate with
the general time of Wisconsinan deglaciation
of the Door Peninsula but do not themselves
reveal the initial date of cave formation.
Brussels Hill Pit Cave sediments are of
post-glacial origin based on relative age de¬
terminations. Granular sediment in the cave
differs in texture, pH, and mineralogy from
surface materials. The cave itself pre-dates
the sediment fill, and the available evidence
and dissolution-rate calculations from
Palmer (1980) indicate that cave develop¬
ment and sediment deposition could have
begun at the end of the Greatlakean sub¬
stage, about 10,000 B.P. Further research
into the history of the lower cave levels will
continue at the conclusion of the faunal
studies.
Brussels Hill Pit Cave contains a post-gla¬
cial sediment sequence covered by later sedi¬
ments and organic material of Recent (Ho¬
locene) age. The coarse sands and gravel may
have been deposited during the initial stages
of glacial recession when water was available
to transport the coarser fractions; as water
volume decreased with ice retreat, transport
potential also decreased, depositing only
finer sands and silts. Collapse of the cave
sinkhole entrance probably occurred at a
relatively recent time.
Sediment variety poses interesting ques¬
tions about sediment deposition and min-
eralogical formation; physical and chemical
alteration of minerals through changes in
water and soil pH and possibly climatic tran¬
sition could be examined further as could
individual sediment particle mineralogy.
The record obtained from Brussels Hill
Pit Cave assists in the analysis of Wiscon¬
sin’s pre-settlement faunal assemblages
(Howe, pers. comm. 1992) and also pro¬
vides a valuable record of post-glacial
geomophological events in northeastern
Wisconsin.
Acknowledgments
Jim Brozowski wishes to thank the review
editors for their insight and constructive
comments. Thanks to Drs. W. F. Kean
and B. E. Brown, Dept, of Geoscience,
UW-Milwaukee, for their assistance
through the various analytical stages. Spe¬
cial thanks to Ms. C. A. Syrrakos for skill¬
ful cartographic and diagram preparation.
Works Cited
Barden, M. 1980. Caves of Door County. In
An introduction to caves of Minnesota, Iowa,
and Wisconsin, ed. E. C. Alexander, pp.
136-41 . National Speleological Society
Convention Guidebook 21.
Birkeland, P. W. 1984. Soils and geomorphol¬
ogy. New York: Oxford University Press.
Brady, N. C. 1974. The nature and properties
14
TRANSACTIONS
BROZOWSKI and DAY: Development of Brussels Hill Pit Cave
of soils. 8th ed. New York: Macmillan.
Bryson R. A., W. M. Wendland, J. D. Ives,
and J. T. Andrews. 1969. Radiocarbon iso¬
chrones on the disintegration of the
Laurentide ice sheet. Arctic and Alpine Re¬
search 1(1): 1-14.
Bull, P. A. 1983. Chemical sedimentation in
caves. In Chemical sediments and geomorphol¬
ogy: Precipitates and residua in the near-surface
environment , ed. A. S. Goudie and K. Pye,
pp. 301-19. London: Academic Press.
Chamberlin, T. C. 18 77. Geology of Eastern
Wisconsin. In Geology of Wisconsin: Wis¬
consin Geological Survey. 2:199-29 6.
Creer, K. M., and P. Tucholka. 1982. Con¬
struction of type curves of geomagnetic
secular variation for dating lake sediments
from east-central North America. Canadian
Journal of Earth Science 19:1 106-13.
Fischer, R. A. 1933. Dispersion on a sphere.
Proc. Royal Soc. London , Ser. A. v. 217.
295-305.
Ford, D. C. 1977. Genetic classification of
solutional cave systems. Proceedings: 7th In¬
ternational Speleological Congress, Sheffield,
England, pp. 211-13.
Gascoyne, M. 1977. Does the presence of sta¬
lagmites really indicate warm periods? New
evidence from Yorkshire and Canadian caves.
Proceedings: 7th International Speleological
Congress, Sheffield, England, pp. 208-09.
Grim, R. E. 1968. Clay Mineralogy. 2nd ed.
New York: McGraw-Hill.
Hansel, A. K, D. W. Mickelson, A. F.
Schneider, and C. E. Larsen. 1985. Late
Wisconsinan and Holocene history of the
Lake Michigan Basin. In Quaternary evolu¬
tion of the Great Lakes, ed. P. F. Karrow and
P. E. Calkin. Geol. Assoc. Canada Special
Paper 30.
Jennings, J. N. 1985. Karst Geomorphology.
Oxford, England: Basil Blackwell Ltd.
Johnson, S. B., and R. D. Stieglitz. 1990.
Karst features of a glaciated dolomite pen¬
insula, Door County, Wisconsin. Geomor¬
phology 4:37-54.
Kean, W. F., and T. E. Klebold. 1981. Paleo-
magnetism of sediment cores from Cedar-
burg Bog, Wisconsin, and a comparison
with cores from Lake Michigan. Journal of
Great Lakes Research 7(3): 276-85.
Klein, C., and C. S. Hurlbut, Jr. 1985.
Manual of Mineralogy (after James D.
Dana). 20th ed. New York: John Wiley and
Sons.
Klussendorf, J., and D. G. Mikulic. 1989.
Bedrock geology of the Door Peninsula. In
Wisconsin \ s Door Peninsula: A Natural His¬
tory, ed. J. C. Palmquist, pp. 12—31.
Appleton, Wisconsin: Perin Press.
Kox, N. H. 1988. Door County’s past locked
in Brussels Hill. The Wisconsin Caver 8(1):
10-11.
Link, E. G., S. L. Elmer, and S. A. Vander-
veen. 1978. Soil Survey of Door County,
Wisconsin. U. S. Dept. Ag. and Soil Cons.
Ser. 132 pp.
Lively, R. S. 1983. Late Quaternary U-series
speleothem growth record from southeast¬
ern Minnesota. Geology 11:259-62.
Lively, R. S., E. C. Alexander, and J. Milske.
1984. A late Pleistocene chronological
record in southeastern Minnesota. Proceed¬
ings: 8th. International Speleological Con¬
gress, pp. 623-26.
McCartney, M. C., and D. M. Mikelson.
1982. Late Woodfordian and Greatlakean
history of the Green Bay Lobe, Wisconsin.
Geological Society of America Bulletin.
93:297-301.
Milske, J. A., E. C. Alexander, Jr., and R. S.
Lively. 1983. Clastic sediments in Mystery
Cave, southeastern Minnesota. National
Speleological Society Bulletin. 45:55-75.
Palmer, A. N. 1980. Hydrochemical factors in
the origin of limestone caves. Proceedings:
8th International Speleological Congress, pp.
120-22.
Volume 82 (1994)
15
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Rosen, C. J. 1984. Karst Geomorphology of the
Door Peninsula, Wisconsin ., M.S. Thesis,
University of Wisconsin— Milwaukee. 119
pp.
Rosen, C. J., and M. J. Day. 1990. Glaciated
karst terrain in the Door Peninsula, Wis¬
consin. Transactions of the Wisconsin Acad¬
emy of Science, Arts, and Letters 78:39-44.
Sherrill, M. G. 1978. Geology and groundwa¬
ter in Door County, Wisconsin with em¬
phasis on contamination potential in the
Silurian dolomite. U.S. Geological Survey-
Water Supply Paper 2047. 38 pp.
Stieglitz, R. D. 1984. Karst landforms of east¬
ern Wisconsin. Geological Society of America
— North Central Section Abstracts 16(3):
200 pp.
- . 1990. The geologic foundations of
Wisconsin’s Door Peninsula. In Door County
and the Niagara Escarpment: Foundations for
the future , ed. K. E. Hersbell, pp. 3-14. Wis¬
consin Academy of Sciences, Arts and Letters
Conference Proceedings.
Sweeting, M. M. 1985. Karst Landforms. Lon¬
don: Macmillan.
Thwaites, F. T., and K. Bertrand. 1957. Pleis¬
tocene geology of the Door Peninsula, Wis¬
consin. Bulletin of the Geological Society of
America 68: 831-80.
Vitorello, I., and R. Van der Voo. 1977. Mag¬
netic stratigraphy of Lake Michigan sedi¬
ments obtained from cores of lacustrine clay.
Quaternary Research 7: 398-412.
Jim Brozowski is currently a research assistant for
the Soils and Physical Geography Laboratory at the
University of Wisconsin— Milwaukee. Address:
Dept, of Geography, University ofWisconsin-Mil-
waukee, P. O. Box 413, Milwaukee, WL 53201-
0413.
Michael J. Day is professor of geography at the
University of Wisconsin— Milwaukee; his research
includes tropical karst geomorphology and the geo¬
morphology of southwestern Wisconsin.
16
TRANSACTIONS
Philip A. Cochran
Occurrence and significance
of sea lamprey (Petromyzon marinus)
in the lower Fox Rivery Wisconsin
Abstract The Rapide Croche lock on the lower Fox River was sealed in the
winter of 1987-1988 to prevent upstream passage of spawning-
phase sea lamprey (Petromyzon marinus) into the Lake Winnebago
watershed. This action was taken because of concern that improve¬
ments in river water quality made colonization by lampreys in¬
evitable, even though sea lampreys had not been collected in the
Fox River prior to closure of the lock. The collection of sea lam¬
preys in the Fox River in 1991, 1992, and 1993 substantiates the
initial concerns and justifies the closure of the Rapide Croche lock.
The sea lamprey (Petromyzon marinus) had a devastating
impact on fish assemblages in the Great Lakes (Smith
1971; Smith and Tibbies 1980; Coble et al. 1990) prior to
initiation of effective chemical control in the early 1 960s. Sea
lamprey control remains largely predicated on chemical treat¬
ment of tributaries inhabited by ammocoetes, but the process
is costly and labor intensive. As control costs rise, it becomes
increasingly desirable to minimize the number and length of
streams that must be chemically treated.
Sea lampreys typically spawn in water of relatively high
quality. Although they have been known to ascend polluted
rivers to spawn in cleaner tributaries, there are known cases
in which lampreys have apparently been inhibited by poor
water quality from ascending polluted rivers. Ironically, one
result of pollution abatement and improved water quality has
been an increase in the number of streams used by spawn¬
ing-phase sea lampreys. For example, sea lampreys colonized
the Peshtigo River following pollution abatement in the early
TRANSACTIONS Volume 82 (1994)
17
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
1970s, leading to increased lamprey abun¬
dance in Green Bay (Moore and Lychwick
1980).
In the latter half of the 1970s, fisheries
biologists in the Wisconsin Department of
Natural Resources (DNR) and elsewhere be¬
came concerned about the possibility of sea
lampreys ascending the Fox River from
Green Bay (Kernen 1979; Smith and
Tibbies 1980). Although the river was his¬
torically subject to severe pollution, by the
1980s water quality had improved to the
point that a walleye ( Stizostedion vitreum)
fishery was reestablished below the De Pere
dam, spawning-phase silver lampreys (Ich-
thyomyzon unicuspis) ascended the Fox River
from Green Bay each year (Cochran and
Marks, in press), and several species of
salmonids were present in the lower river
during the spring when water temperatures
were favorable. Concerns about sea lampreys
in the Fox River reflect several possible sce¬
narios, which are not mutually exclusive.
First, if sea lampreys gained access to the ex¬
tensive high quality tributary systems in the
Fox River drainage above Lake Winnebago
(e.g., the Wolf and Embarrass rivers), spawn¬
ing would probably be successful. Lampreys
produced in these tributaries might pose a
threat not only to fisheries in Green Bay and
Lake Michigan, but also to Lake Winnebago
fisheries previously unexposed to sea lam¬
preys. The sea lamprey’s proclivity for size-
selective attack (Farmer and Beamish 1973;
Cochran 1983) would make Lake Winne¬
bago’s unique and valuable lake sturgeon
( Acipenser fulvescens ) population a likely first
target. Second, sea lampreys conceivably
might spawn in some reaches of the Fox
River proper, much as they are known to
spawn in the St. Mary’s River between lakes
Superior and Huron (Smith and Tibbies
1980). Regardless of where it occurred, re¬
production in the Fox River system would
greatly increase the cost and difficulty of
chemical control, or, if unchecked, endan¬
ger valuable fisheries.
In 1987, the Wisconsin DNR recom¬
mended that a lock on the lower Fox River
be sealed and the corresponding dam modi¬
fied to prevent upstream migration by sea
lampreys. This proposal drew protests, pri¬
marily from pleasure boaters accustomed to
traveling the Fox River between Lake
Winnebago and Green Bay. Nevertheless,
the recommendation was reaffirmed by the
Sea Lamprey Study Committee appointed
by the governor of Wisconsin (1988), and
the Rapide Croche lock at the third dam up¬
stream from Green Bay was sealed during
the winter of 1987-1988.
All of the previous developments oc¬
curred before any sea lampreys were col¬
lected in the Fox River proper. Beginning
in 1979, the U.S. Fish and Wildlife Service
had sponsored the placement and monitor¬
ing of a sea lamprey assessment trap below
the De Pere dam each spring during the lam¬
prey spawning season, but no sea lampreys
had ever been collected in the trap. Critics
of the closure of the Rapide Croche lock
pointed to the lack of evidence that sea lam¬
preys had ever entered the river. Proponents
of the closure argued that sea lampreys had
been taken in Green Bay at the mouth of
the Fox River and that by the time sea lam¬
preys were detected in the river itself, it
might be too late to prevent their spread into
the Lake Winnebago system.
In light of the controversy surrounding
the closure of the Rapide Croche lock, the
subsequent capture of sea lampreys in the
Fox River is noteworthy. The purpose of this
report is to describe the lamprey collections
and discuss their significance. Because of in¬
accuracies in accounts presented through the
popular media, it is important that an ac¬
curate record be provided.
18
TRANSACTIONS
COCHRAN: Sea lamprey in the lower Fox River, Wisconsin
Methods
A portable sea lamprey assessment trap
(Schuldt and Heinrich 1982) has been set
each year since 1979 below the east end of
the De Pere dam, approximately 12 km up¬
stream from Green Bay. Additionally, a trap
was set in 1988 and 1989 in the Osen mill-
race just east of the De Pere lock. Traps were
checked five days per week from early April
to mid-June. Measurements of total length
(TL) and body mass (BM) reported here
were collected from live animals anesthetized
with tricaine methanesulfonate or from dead
specimens prior to preservation.
Results and discussion
Lamprey assessment trapping and other fish
collecting during the years 1979-1990
yielded no sea lampreys, but five individuals
were collected from 1991 through 1993. In
1991, the trap was first lifted on April 3 af¬
ter being set the previous day, and it con¬
tained an adult male sea lamprey (University
of Wisconsin-Madison Zoology Museum,
UWZM 9973; TL - 587 mm, BM - 365 g).
Water temperature was 5°C. Subsequently in
1991, two additional specimens were col¬
lected by the Wisconsin DNR in fyke nets
set in the Fox River between the De Pere dam
and Green Bay. In 1992, an adult female sea
lamprey (TL - 600 mm, BM - 505 g) was
trapped on April 6 at a water temperature of
5°G. No sea lampreys were trapped in 1993,
but the Wisconsin DNR, while electrofishing
below the De Pere dam on October 12, col¬
lected a lake trout ( Salvelinus namaycush)
bearing a parasitic-phase individual (TL -
353 mm, BM - 84 g).
The relatively large sizes of the lampreys
captured in the trap is typical of sea lampreys
collected in tributaries to Green Bay (Johnson
1982). However, collection of upstream mi¬
grants in early April is relatively unusual in
northeastern Wisconsin. For example, during
1987-1989, the first sea lampreys were cap¬
tured in assessment traps in the East Twin
River, Manitowoc County, on April 23, May
7, and April 26. Johnson (1982) trapped
most of the lampreys he collected in the
Peshtigo River during the period May 1 6-3 1 .
Sea lampreys began entering the Peshtigo
River traps after the water temperature
reached 1 0°C, and peak catches occurred be¬
tween 15.6°C and 21.1°C.
Contrary to certain accounts in the popu¬
lar media, sea lampreys in the lower Fox
River do not constitute a threat to fisheries
in Lake Winnebago at this time, because the
Rapide Croche lock remains sealed to pre¬
vent their upstream passage. However, sen¬
timent to reopen the Fox River waterway to
unimpeded boat traffic persists in some
quarters, and documentation that sea lam¬
preys occur in the Fox River helps legitimize
opposition to that sentiment. Any future ar¬
rangement for boat passage at the Rapide
Croche dam must involve a boat lift or some
other terrestrial transport. This will prevent
upstream passage not only by sea lamprey,
but also by white perch ( Morone americana ),
an exotic species that was first captured in
Green Bay and the lower Fox River in 1988
(Cochran and Hesse 1994). Closure of the
Rapide Croche lock may fortuitously have
prevented the white perch from gaining ac¬
cess to Lake Winnebago.
It remains to be seen whether increasing
numbers of sea lamprey will ascend the
lower Fox River during subsequent spawn¬
ing seasons. Whether such an occurrence is
biologically significant depends on whether
spawning is successful and whether the bur¬
rowing ammocoetes can survive in the lower
river or Green Bay for the duration of the
larval phase (roughly four to five years). It
has been suggested that walleye reproduction
Volume 82 (1994)
19
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
in the lower Fox River is limited by the avail¬
ability of chemically suitable substrate (Auer
and Auer 1990), and it may be tempting to
extend that conclusion to sea lamprey. How¬
ever, although Auer and Auer (1990) cited
a lack of evidence of natural recruitment by
walleye in the Fox River, juvenile walleye of
yearling size are collected with regularity in
the sea lamprey trap in De Pere, despite the
fact that the Wisconsin DNR discontinued
stocking after 1984 (Schneider et al. 1991).
Moreover, the recent occurrence of adult
Hexagenia bilineata mayflies along the Fox
River at the De Pere dam (Cochran 1992)
indicates that conditions at the sediment-wa¬
ter interface have improved to the point that
Hexagenia naiads can once more complete
their development in a microhabitat similar
to that used by sea lamprey ammocoetes.
At this time, the significance of sea lam¬
preys in the lower Fox River is primarily
symbolic. They are symbolic, for example,
of the improvements in water quality that
first permitted them to become an issue.
More importantly, they provide an histori¬
cal footnote to a case in which foresight and
proactive measures prevented the contami¬
nation of a watershed by exotic species. Such
cases are all too rare and warrant documen¬
tation.
Acknowledgments
Sea lamprey assessment trapping below the
De Pere dam was supported by contracts
with the U.S. Fish and Wildlife Service.
Manuscript revision was completed while I
was funded by the National Oceanic and
Atmospheric Administration’s Office of Sea
Grant, Department of Commerce, through
an institutional grant to the University of
Wisconsin. I am grateful to the many un¬
dergraduate students from St. Norbert Col¬
lege who have assisted with the trapping over
the years and to Kristen Lucier for assistance
with manuscript preparation. I am also
thankful to Terry Lychwick, Dave Bougie,
and Lee Meyers of the Wisconsin Depart¬
ment of Natural Resources for sharing their
field data with me.
References
Auer, M. T., and N. A. Auer. 1990. Chemical
suitability of substrates for walleye egg devel¬
opment in the lower Fox River, Wisconsin.
Trans. Amer. Fish . Soc. 119:871-76.
Coble, D. W., R. E. Bruesewitz, T. W. Fratt,
and J. W. Scheirer. 1990. Lake trout, sea lam¬
preys, and overfishing in the upper Great
Lakes: a review and reanalysis. Trans. Amer.
Fish. Soc. 119:985-95.
Cochran, P. A. 1985. Size-selective attack by
parasitic lampreys: consideration of alternate
null hypotheses. Oecologia 67 : 1 37-4 1 .
- . 1992. The return of Hexagenia (Ephe-
meroptera: Ephemeridae) to the lower Fox
River, Wisconsin. Great Lakes Entomologist
25:79-81.
Cochran, P. A., and P. J. Hesse. 1994. Obser¬
vations on the white perch (Morone amer-
icana) early in its invasion of Wisconsin.
Trans. Wise. Acad. Sci. Arts Lett. 82:23-31.
Cochran, P. A., and J. E. Marks. In press. The
biology of the silver lamprey (Ichthy-omyzon
unicuspis) in Green Bay and the lower Fox
River, with a comparison to the sea lamprey
(Petromyzon marinus). Copeia.
Farmer, G. J., and F. W. H. Beamish. 1973. Sea
lamprey (Petromyzon marinus ) predation on
freshwater teleosts. J. Fish. Res. Board Can.
30:601-05.
Johnson, W. J. 1982. Body lengths, body
weights, and fecundity of sea lampreys
(Petromyzon marinus) from Green Bay, Lake
Michigan. Trans. Wise. Acad. Sci. Arts Lett.
70:73-77.
Kernen, L. 1979. Special report - sea lamprey.
20
TRANSACTIONS
COCHRAN: Sea lamprey in the lower Fox River, Wisconsin
Wisconsin Sportsman (May/June): 36.
Moore, J, D., and T. J. Lychwick. 1980.
Changes in mortality of lake trout (Salvelinus
namaycush) in relation to increased sea lam¬
prey (Petromyzon marinus) abundance in
Green Bay, 1974-1978. Can. J. Fish . Aquat.
Sci. 37:2052-56.
Schneider, J. C, T. J. Lychwick, E. J. Trimber-
ger, J. H. Peterson, R. O'Neal, and P. J.
Scheeberger. 1991. Walleye rehabilitation in
Lake Michigan, 1969-1989. In Status of wall¬
eye in the Great Lakes: case studies prepared for
the 1989 workshop , ed. P. J. Colby, C. A.
Lewis, and R. L. Eshenroder, pp. 23-61.
Great Lakes Fishery Commission Special
Publication No. 91-1.
Schuldt, R. J., and J. W. Heinrich. 1982. Por¬
table trap for collecting adult sea lampreys.
Prog. Fish -Cult. 44:220-21.
Sea Lamprey Study Committee. 1988. Report of
the sea lamprey study committee on closing the
Rapide Croche lock on the Fox River. 42 pp.
Smith, B. R. 1971. Sea lampreys in the Great
Lakes of North America. In The biology of
lampreys , Vol. 1, ed. M. W. Hardisty and I.
C. Potter, pp. 207-47. New York: Academic
Press.
Smith, B. R., and J. J. Tibbies. 1980. Sea lam¬
prey (Petromyzon marinus) in Lakes Huron,
Michigan, and Superior: history of invasion
and control, 1936-78. Can. J. Fish. Aquat.
Sci. 37:1780-1801.
Philip A. Cochran is an associate professor of
biology at St. Norbert College. Much of his re¬
cent research has involved exotic species in the
Fox River. Address: Division of Natural Sci¬
ences, St. Norbert College, De Pere, Wisconsin
54115
Volume 82 (1994)
21
>r\
Philip A. Cochran and Peter J. Hesse
Observations on the white perch
(Morone americana) early in
its invasion of Wisconsin
Abstract White perch (Morone americana) were first reported in the Green Bay/
Fox River system in Wisconsin in 1988. Total spring catches in sea
lamprey assessment traps below the De Pere dam increased until
1990 , decreased in 1991 , and then increased sharply in 1992 and
1993. These collections are believed to represent at least in part the
result of upstream movements related to spawning. White perch were
not trapped in large numbers each spring until water temperature
exceeded 17—18°C, and males were collected on average slightly ear¬
lier than females. Several age classes were present among fish cap¬
tured as early as 1989. Growth rates were relatively high and were
comparable to those in other locations in the Great Lakes where popu¬
lations are expanding. Growth declined, however, with increasing
age. Relationships between weight and length were similar between
the two sexes. Although the spread of the white perch up the Fox River
appears to have been blocked by the sealing of the Rapide Croche
lock in 1988, more information is needed to assess its impact in the
lower Fox River and Green Bay, and care must be taken to mini¬
mize its chances for invading Wisconsin s inland waters.
The white perch (. Morone americana) invaded the Great
Lakes from the Atlantic Coast in the 1940s and 1950s
by way of the Erie and Welland canals (Johnson and Evans
1990). It was first identified in the Lake Michigan drainage in
the lower Fox River of Wisconsin in May 1988 (Meyers 1988),
and in September 1988, it was collected in Belmont Harbor
in Chicago (Savitz et al. 1989). Collections from inland wa¬
ters in Illinois (Blodgett 1993), including the Illinois River,
indicate that it has spread from Lake Michigan through the
Upper Illinois Waterway, which connects the Great Lakes and
the Mississippi River drainages.
TRANSACTIONS Volume 82 (1994)
23
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Concern about the colonization of Wis¬
consin waters by white perch stems from ( 1 )
the possibility for it to compete with more
desirable native species (Schaeffer and
Margraf 1986a), (2) its potential impact as
a predator on the eggs of other species
(Schaeffer and Margraf 1987), and (3) the
potential for it to interbreed with the white
bass | M. chrysops). In this paper we present
data on the biology of the white perch early
in its invasion of Wisconsin waters. Our re¬
sults can serve as a basis for comparison with
future data collected after the white perch
presumably becomes fully established. Our
experience also provides insight into the ef¬
fectiveness of programs to monitor the
spread of other exotic species, such as the sea
lamprey (. Petromyzon marinus).
Methods
White perch were collected with other fishes
in portable sea lamprey assessment traps
(Schuldt and Heinrich 1982) set below the
De Pere dam on the Fox River, Brown
County, Wisconsin, 12 km upstream from
Green Bay. In 1988 and 1989, one trap was
set below the eastern end of the dam spill¬
way and another below a small spillway as¬
sociated with a hydroelectric generator in a
building east of the lock channel. The latter
spillway is situated at the head of a millrace
which enters the Fox River below the lock
channel. Data from the two traps were
pooled. From 1990 to 1993, a single trap
below the dam spillway was monitored.
Trapping was conducted for ten weeks from
early April to mid-June in all years accord¬
ing to a protocol dictated by contract with
the U.S. Fish and Wildlife Service. In addi¬
tion, we trapped during the periods Octo¬
ber 4—11, 1992, February 8-13, 1993, and
June 12-October 29, 1993. Traps were
emptied five days per week at intervals of no
greater than 48 hours. Water temperature
was recorded each time the traps were emp¬
tied. During the years 1989-1991, white
perch were enumerated and, on most days,
taken to the laboratory to be weighed and
sexed (1990 only) and measured for total
length (TL). In 1989 and 1990, scale
samples for age determination by the junior
author were collected from the upper left
side between the lateral line and the second
and third dorsal fin spines.
We occasionally collected fishes in the
Fox River upstream from the De Pere dam
by electrofishing with a boat-mounted gen¬
erator (pulsed DC current). Samples were
collected in 1988, 1989, 1992, and 1993,
most often in the vicinity of the St. Norbert
College campus.
Voucher specimens (UWZM 9726) have
been deposited in the University of Wiscon-
sin-Madison Zoology Museum.
Results
White perch were first detected in the Fox
River in 1988 during the interval that the
sea lamprey traps were operated (Meyers
1988), but no white perch were collected in
the traps that year. Twelve individuals were
captured in the two traps monitored in 1989
(Table 1). During the period 1990 to 1993,
when a single trap was operated, total spring
catch ranged from a low of 2 1 in 1991 to a
high of 1196 in 1992 (Table 1). In Octo¬
ber 1992, six trap days yielded a total of 20
white perch (22% of the combined catch of
all species). No white perch were collected
during four trap days in February 1993.
In most years trapping ended in mid-June,
but in 1993, it was extended through the
summer and into autumn. The high catches
of late May and early June (Fig. 1) declined
through late June and July. Monthly totals
for June and July were 463 and 60, respec-
24
TRANSACTIONS
COCHRAN AND HESSE: The white perch early in its invasion of Wisconsin
Table 1. Total yearly trap catch of white perch, percentage of total catch for all fish
combined, mean date of capture, and mean temperature of capture. Means are fol¬
lowed by standard errors in parentheses.
Number of Percentage of Mean Date Mean Temperature
Year White Perch Total Catch of Capture of Capture (°C)
tively. No white perch were collected from
August 1 through October 29.
Differences among years in the timing of
spring trap catches may have been related to
water temperature (Fig. 1). The dates of first
capture in 1989 (May 21) and 1991 (May
19) were similar, but the date of first cap¬
ture in 1990 (April 26) was much earlier. In
each of the three years, white perch did not
appear in the traps until after the water tem¬
perature first reached 18°C, but that oc¬
curred earlier in 1990 than in 1989 or 1991
(Fig. 1). In 1992, only 17 white perch were
collected prior to May 1 1 . On that date,
when water temperature first measured
17°C, 75 individuals were collected (Fig. 1).
In 1993, an unusually cool, wet spring with
high discharge, high catches were recorded
at cooler temperatures than in previous
years, but the highest daily catch (93) oc¬
curred on June 1 1 when the water tempera¬
ture first reached 19°C. Analysis of variance
revealed that weighted mean temperature of
capture (Table 1) differed significantly
among years (F42236 = 300.6, P = 0.000), as
did weighted mean date of capture (F4 2236 =
126.8, P - 0.000).
In 1990, sex was determined for 171 of
189 white perch. The numbers of males
(100) and females (71) were significantly
different from what would be expected un¬
der the null hypothesis that the two sexes
were equally abundant and equally suscep¬
tible to capture (normal approximation to
the binomial test, P < 0.05). Although there
was great overlap between the two sexes, the
mean date of capture for males (May 6) was
significantly different from that for females
(May 9) (t = -3.58, P = 0.0006), indicating
that males move upriver slightly prior to fe¬
males.
Ages were estimated from scales of 1 0 fish
collected in 1989 and 170 fish collected in
1990 (Table 2). In both years, several age
classes were present, but age IV+ fish were
most abundant. Based on the 1990 data,
there was no difference between sexes in the
relative numbers of individuals of different
ages (chi-squared test of independence, with
age II pooled with III and age V pooled with
VI to keep all expected values greater than
five, x2= 0.769).
The size distributions of white perch col¬
lected in the traps differed among years (Fig.
2). In 1989, one fish with a TL of 1 15 mm
was collected, and nine were in the TL range
of 192-230 mm. In 1990, TL ranged from
162 mm to 242 mm, but most individuals
fell within 180-210 mm (body mass in
1990 ranged from 53 g to 246 g). In 1991,
Volume 82 (1994)
25
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
25 150
-20 120
- 15 90
10 60
15
C
10 70
- 5
DAY OF YEAR
Fig. 1. Number of white perch collected in sea lamprey assessment traps and wa¬
ter temperature versus day of year during 1989-1993. Note that the left vertical scale
for the years 1989-1991 is different from that for 1992-1993.
26
TRANSACTIONS
COCHRAN AND HESSE: The white perch early in its invasion of Wisconsin
Table 2. Ages of ten of twelve white perch collected in 1989 and 170 of 189 white
perch collected in 1990, followed by mean total length in millimeters and mean body
mass in grams. Means are followed by standard errors (SE) in parentheses and as¬
sociated sample sizes. 1990 data are partitioned by sex. Not all fish were measured
for both length and body mass.
Age
1989
1990
Linear regressions of the natural loga¬
rithm of body mass (LNWT) on the natu¬
ral logarithm of total length (LNTL) were
calculated with 1990 data for both sexes
pooled and for each sex individually. For all
fish, LNWT = -10.7 + 2.92 LNTL (r2 =
0.866, n = 1 16). For males, LNWT = -10.3
above the De Pere dam throughout the
course of this study, we collected no white
perch above the dam until September 23,
1993, when two small individuals (TL: 64
and 72 mm) were captured along the St.
Norbert College shoreline. These were prob¬
ably young-of-the-year.
Volume 82 (1994)
27
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Discussion
If the catch of white perch in the lamprey
assessment trap is a reliable index of their
abundance, then the white perch population
increased dramatically from 1988, when
they were first discovered in the Green Bay/
Fox River system, until 1993, when they
represented 24% of the total spring Fish
catch (Table 1). In at least one location in
the Great Lakes, the Bay of Quinte of Lake
Ontario, white perch rapidly became a
dominant component of the fish assemblage
within a few years of their invasion, only to
undergo a dramatic decline attributed either
to severe winter weather or to increased
piscivore abundance (Hurley 1986). In Lake
Erie, however, white perch first invaded in
the 1930s but did not increase in abundance
substantially until the 1970s (Schaeffer and
Margraf 1986b). Johnson and Evans (1990)
suggested that the Great Lakes distribution
of white perch is limited by low tolerance
of cold temperatures. They pointed out that
the current distributional limit approximates
the -5°C winter air isotherm. Since Green
Bay lies slightly outside that isotherm, its
white perch populations might be expected
to fluctuate in response to year-to-year cli¬
matic variability.
We interpret at least part of our spring
catch of white perch to represent the result
of an upstream spawning migration, al¬
though our samples and reports by local an¬
glers indicate that at least some white perch
are present in the river outside of the spawn¬
ing season. Sea lamprey assessment traps are
positioned to capture fish whose movement
upstream has been blocked by a dam or
other barrier. At least some of the white
perch we captured in this manner were in
spawning condition (i.e., milt was freely ex¬
pressed by some males), and they displayed
the bluish cast reported on the lower jaws
TOTAL LENGTH (mm)
Fig. 2. Frequency histograms of total
length of white perch collected in (a)
1989 (n= 1 0), (b) 1990 (n=176), and (c)
1991 (n=21). Note that the vertical scale
for the 1990 histogram is different from
that for 1989 and 1991.
28
TRANSACTIONS
COCHRAN AND HESSE: The white perch early in its invasion of Wisconsin
Fig. 3. Total length versus estimated age
of white perch collected in 1989 and
1990
of spawning adults (Scott and Crossman
197; Smith 1985). Moreover, both Smith
(1985) and Schaeffer and Margraf (1987) in¬
dicated that spawning peaked when water
temperature reached 18°C, the approximate
temperature at which white perch first ap¬
peared in abundance in our traps. Most of
the fish we captured were above the size and
age of maturity reported for white perch in
Lake Erie (Schaeffer and Margraf 1986b).
The modal group of smaller fish in 1991
(Fig. 2) may have been prereproductive, but
in 1992 we noted fish as small as 112 mm
from which milt was freely expressed.
Our interpretations of age and growth
must be accepted with caution until the use
of scales for aging white perch in this sys¬
tem has been validated. Nevertheless, our
results indicated that white perch in the
Green Bay system grow relatively quickly,
especially early in their life, and that growth
is comparable to that by white perch in Lake
Erie (Schaeffer and Margraf 1986b). How¬
ever, this species tends in freshwater habi¬
tats to become overpopulated, resulting in
slower growth and an abundance of stunted
individuals (Scott and Crossman 1973).
The presence of several age classes of
white perch in the Fox River in 1989 and
1990 (Table 2) suggests either that white
perch were present in the area and reproduc¬
ing for several years prior to their discovery
in 1988 or that their initial colonization in¬
volved large numbers of individuals, perhaps
by multiple introductions (e.g., bilgewater
release by freighters). In either case, the lam¬
prey assessment trap did not capture white
perch until a year after they were known to
be present in the Fox River. This lapse be¬
tween the occurrence and detection of an
exotic species is especially relevant to the
operation of the lamprey trap in the Fox
River. The trap has been monitored each
spring since 1979 because of concern that
water quality improvements might be fol¬
lowed by the movement of spawning-phase
sea lampreys up into the river. Indeed, a sea
lamprey was collected in the trap for the first
time in 1991 (Cochran 1994), but in light
of our experience with white perch, it is
quite possible that sea lampreys occurred in
the river in prior years.
Although white perch were present in the
lower Fox River as early as 1988, we did not
collect them above the De Pere dam until
1993. During part of the spring spawning
period, the De Pere lock channel is not yet
open to boat traffic (it typically is opened
at the end of May), and this may have de¬
layed dispersal past the dam. We have, how¬
ever, trapped white perch below the dam on
dates after the locks were in operation.
Spawning white perch from Lake Erie move
at least 45 km up into tributary streams
(Schaeffer and Margraf 1987), and it would
seem inevitable that white perch would
eventually traverse the 48 km between Green
Bay and Lake Winnebago, where their pres-
Volume 82 (1994)
29
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
ence could have substantial effects on impor¬
tant fisheries. Fortuitously, however, up¬
stream movement by white perch toward
Lake Winnebago has been blocked by the
sealing of the Rapide Croche lock at the
third dam upstream from Green Bay. This
action was somewhat controversial when it
was undertaken early in 1988 in anticipation
of an invasion by sea lampreys. In retrospect,
it may have been just in time to stop white
perch.
Dams may eventually play an important
role in limiting dispersal by white perch into
other parts of Wisconsin. Now that they
have invaded the Mississippi River in Illinois
(Blodgett 1993), white perch can be ex¬
pected to re-enter Wisconsin through the
Mississippi River and its tributaries, some of
which are impounded by impassable dams.
For example, upstream dispersal in the Wis¬
consin River will be blocked by the Prairie
du Sac dam, preventing access to a large area
in north central Wisconsin.
At present, white perch in Wisconsin are
apparently concentrated in the lower Fox
River and southern Green Bay. They have
not been collected in sea lamprey assessment
traps operated in Green Bay tributaries other
than the Fox River (i.e., the Menominee,
Peshtigo, and Oconto rivers). Moreover, al¬
though they have been collected in Lake
Michigan near Chicago (Savitz et al. 1989),
they have not been taken in lamprey traps
in the East Twin River, a tributary to Lake
Michigan in Manitowoc County.
More information is needed to fully
evaluate the ecological impact of white perch
on the Green Bay/Fox River system, includ¬
ing their effects on the recently revitalized
walleye ( Stizostedion vitreum) and yellow
perch {Perea flavescens) fisheries. In particu¬
lar, the extent to which white perch feed on
walleye eggs should be assessed, because they
are known to feed on walleye eggs in the
Lake Erie basin (Schaeffer and Margraf
1987). In addition, the extent to which wall¬
eye and other piscivores use white perch as
forage should be investigated. Finally, efforts
should be made to minimize the spread of
this exotic species within Wisconsin’s inland
waters.
Acknowledgments
Our yearly spring trapping was funded by
contracts with the U.S. Fish and Wildlife
Service. Additional trapping, manuscript re¬
vision, and analysis of data for 1992 and
1993 were enabled by support from the Na¬
tional Oceanic and Atmospheric Admin¬
istration’s Office of Sea Grant, Department
of Commerce, through an institutional grant
to the University of Wisconsin. We thank
Doug Basten, Joe Cochran, Eric Golden,
Brian Jones, Adam Leisten, Joe Marks,
Marty Sneen, and Rick Wagner for assis¬
tance with the collection of white perch, and
James Hodgson for aid in the preparation
of the figures. We appreciate the support of
St. Norbert College for undergraduate re¬
search.
Works Cited
Blodgett, D. 1993. Exotic species collected from
Illinois River. River Almanac (U.S. Fish and
Wildlife Service, Environmental Manage¬
ment Technical Center, Onalaska, Wiscon¬
sin), May 1993, pp. 1, 3.
Cochran, P. A. 1994. Occurrence and signifi¬
cance of sea lamprey {Petromyzon marinus) in
the lower Fox River, Wisconsin. Trans. Wise.
Acad. Sci., Arts Lett. 82:17-21.
Hurley, D. A. 1986. Fish populations in the Bay
of Quinte, Lake Ontario, before and after
phosphorus control. Canadian Special Publi¬
cation of Fisheries and Aquatic Sciences
86:201-14.
30
TRANSACTIONS
COCHRAN AND HESSE: The white perch early in its invasion of Wisconsin
Johnson, T. B., and D. O. Evans. 1990. Size-
dependent winter mortality of young-of-the-
year white perch: climate warning and inva¬
sion of the Laurentian Great Lakes. Trans.
Amer . Fish. Soc. 119:301-13.
Meyers, L. 1988. 1988 sea lamprey sampling ef¬
fort, Fox River, Brown County. Wisconsin
Department of Natural Resources Correspon¬
dence/Memorandum, File Ref. 3600, June
23.
Savitz, J., C. Aiello, and L. G. Bardygula. 1989.
The first record of the white perch ( Morone
americana) in Illinois waters of Lake Michi¬
gan. Trans. Illinois Acad. Sci. 82:37-38.
Schaeffer, J. S., and E. J. Margraf. 1986a. Food
of white perch (Morone americana ) and po¬
tential for competition with yellow perch
(Perea flavescens ) in Lake Erie. Ohio J. Sci.
86:26-29.
- . 1986b. Population characteristics of
the invading white perch (Morone americana)
in western Lake Erie. J. Great Lakes Res.
12:127-31.
- - 1987. Predation on fish eggs by white
perch, Morone americana , in western Lake
Erie. Env. Biol. Fishes 18:77-80.
Schuldt, R. J., and J. W. Heinrich. 1982. Por¬
table trap for collecting adult sea lampreys.
Prog. Fish-Cult. 44:220-21.
Scott, W. B., and E. J. Crossman. 1973. The
freshwater fishes of Canada. Fish Res. Board
Can. Bull. 184. 966 pp.
Smith, C. L. 1985. The inland fishes of New York
state. Albany: N.Y. State Department of Env.
Cons., 522 pp.
Philip A. Cochran is an associate professor of biol¬
ogy at St. Norbert College. Much of his recent re¬
search has involved exotic species in the Fox River.
Address: Division of Natural Sciences , St. Norbert
College , De Pere, Wisconsin 54115
Peter J. Hesse completed his B.S. degree in biol¬
ogy at St. Norbert College in 1990. He currently
works as an environmental consultant.
Volume 82 (1994)
31
Robert B. DuBois and Dennis M. Pratt
History of the fishes of the Bois Brule
River System , Wisconsin, with emphasis
on the salmonids and their management
Abstract The Bois Brule River in Douglas County is one of Wisconsin's larg¬
est, best known, and most intensively studied trout streams. A di¬
verse fish fauna of at least 63 species (11 of which are exotic, plus
one cultured hybrid) has been collected from the watershed. How¬
ever, only 21 are coldwater, riverine species with viable populations;
the remainder are either lentic, warmwater forms found in lakes
Minnesuing and Nebagamon, are Lake Superior species that only
occasionally enter the lower river, or are locally rare. The fish fauna
has been profoundly altered by species introduced both intentionally
and accidentally, and by control efforts directed at the exotic sea lam¬
prey (Petromyzon marinus), which have caused severe density and dis¬
tribution reductions in two species of native lampreys. Once sustain¬
ing a population of native brook trout (Salvelinus fontinalis) as the
only angling target, the river now provides angling opportunities for
four species of exotic salmonids as well. Declines in several of the
salmonid populations, especially during the last two decades, may be
attributable to over-exploitation; consequently, fishery regulations
have become increasingly restrictive. Top priority, however, will con¬
tinue to be maintenance of excellent habitat and water quality. With
fine riparian stewardship practiced by private landowners, coupled
with the state stewardship land acquisition program within state for¬
est boundaries, this focus appears to be sustainable. Other habitat
management efforts have included in-stream habitat enhancement
techniques, beaver (Castor canadensis) control and dam removal,
dredging projects, and bank stabilization efforts in red clay areas
prone to slippage.
TRANSACTIONS Volume 82 (1994)
33
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
The Bois Brule River (hereafter referred
to as the Brule River) in Douglas
County is perhaps the most famous trout
stream in Wisconsin (O’Donnell 1944). Its
fame is related both to its rich history as an
important water route, linking Lake Supe¬
rior and the upper Mississippi River drain¬
age, and its historic reputation for excellent
trout fishing (O’Donnell 1944; Marshall
1954). Additionally, the river has long been
renowned for its beauty and an intangible
mystique that repeatedly draws anglers and
canoeists alike back to its waters. Today the
Brule River provides habitat for a diverse ar¬
ray of coldwater organisms in addition to the
salmonids. As one of the larger spring-fed
streams in the Midwest, it is a regionally
unique resource deserving the highest level
of protection.
The original renowned trout fishery of
the Brule River bore little resemblance to the
current fishery for five salmonid species.
During the middle and late 1800s the river
was widely acclaimed for its native brook
trout ( Salvelinus fontinalis) fishing; this fish¬
ery was comprised of both stream-resident
and anadromous, or coaster1, components.
Unfortunately, this fishery had begun to de¬
cline sharply by the turn of the century
(Jerrard 1956), probably from a combina¬
tion of over-exploitation, habitat loss, and
logging dam effects (O’Donnell 1944).
However, subsequent introductions of steel-
head (anadromous rainbow trout, Onco-
rhynchus my kiss), brown trout ( Salmo trutta ),
and various brook trout strains, in combi¬
nation with increasingly restrictive angling
regulations and termination of extensive log¬
ging, helped bolster the fishery back into
prominence. High quality trophy fisheries
for anadromous brown trout, steelhead, and
more recently, coho salmon ( Oncorhynchus
kisutch) and chinook salmon ( Oncorhynchus
tshawytscha ) have since developed and are
augmented by challenging upper river and
tributary fisheries for stream-resident brook
trout and brown trout.
Throughout the recorded history of this
river, anglers have often voiced complaints
related to perceived dips in the quality of
fishing. In 1983 an advisory task force (the
Brule River Committee) was formed in re¬
sponse to perceived declines in populations
of both steelhead and brown trout. The com¬
mittee was formed of Wisconsin Department
of Natural Resources (WDNR) personnel
and concerned citizens representing area
sports clubs. Objectives for the committee
were to identify and prioritize the most press¬
ing fishery problems and formulate sugges¬
tions for remedial actions. Most of the prob¬
lems identified pointed to a common need:
to promptly initiate a long-term, comprehen¬
sive research project to provide quantitative
data about the salmonid populations. Al¬
though the river had been the focus of much
investigation since the 1 940s (see section on
ecological investigations), the descriptive in¬
formation obtained was of limited value for
guiding management actions. Late in 1983
the WDNR initiated a broadly based re¬
search initiative to provide the information
on salmonid population dynamics needed to
optimize management of the fishery.
This report grew out of the research ef¬
fort initiated in 1983 and presents an his¬
torical sketch of the fish populations of the
Brule River system along with the factors
that have affected them over the last two
centuries. Our emphasis regarding these ob¬
jectives is focused primarily on the salmo¬
nids and their management, and secondarily
on the exotic sea lamprey ( Petromyzon
marinus ), because these species have received
the most management and research atten¬
tion. However, because these species are just
part of a diverse fish community, we also
summarize the information available, both
34
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
historical and recent, for all fish species
within the river system.
Methods
This report represents a compilation of in¬
formation taken from a variety of sources in¬
cluding data collected by the authors during
various phases of a multifaceted salmonid
research project on the Brule River during
1983-93, file data from the WDNR Brule
Area and Superior offices, and numerous
published sources. Works by O’Donnell
(1944), Holbrook (1949), Marshall (1934),
and Jerrard (1936) were instrumental in pro¬
viding information about human develop¬
ment within the Brule River Valley and the
historic trout fishery. Physical and chemical
information for the mainstem2 and tributar¬
ies were summarized from several sources.
Fish distribution information was obtained
either during WDNR fishery surveys over
the last decade or from the sources acknowl¬
edged in Table 1 . Both WDNR file data and
published accounts of species distributions
were used except in a few cases where they
were clearly inconsistent with established in¬
formation on statewide distributions (Becker
1983; Fago 1992). Collections from 44 sam¬
pling stations located throughout lotic areas
of the river system during 1987-91 (DuBois
et al. in press) targeted juvenile and stream-
resident salmonids and were made with stan¬
dard WDNR stream electrofishing units us¬
ing 220 volt direct-current generators;
electrofishing surveys prior to about 1980
used less efficient (on salmonids) alternating-
current generators. Spawning runs of anadro-
mous salmonids were examined with a view¬
ing window and sea lamprey trap at the sea
lamprey barrier/fishway (hereafter referred to
as the barrier/ fishway — Fig. 1). Smolts were
studied with an inclined-screen trap (DuBois
et al. 1991) below2 the barrier/ fishway.
Sport fishery statistics were summarized
from WDNR creel surveys conducted in
1973, 1978-79, 1984, and 1986 on the
mainstem from Stone’s Bridge to the mouth,
in 1990 on the lower river2 only, and in
1992 on the upper river2 only. Random,
stratified, timed-interval designs — also
known as bus route designs (Jones and
Robson 1991) — were used to obtain com¬
pleted trip interviews at major access points
in 1986, 1990, and 1992; earlier access-
point surveys were not stratified according
to anticipated angling pressure. Information
requested from each angler included the
length of time fished, fishing methods used,
data about the catch, and perceptions of the
fishing experience. Creeled salmonids were
measured to the nearest 0.1 inch and scale-
sampled as needed for age analysis.
The Physical Setting
The 47-mile-long Brule River drains a wa¬
tershed of about 130 square miles and flows
north into Lake Superior (Fig. 1). The av¬
erage discharge near the WDNR Brule Area
Headquarters on the river’s midsection is
169 ft3/sec with extremes ranging from 67
to 1,520 ft3/sec (Niemuth 1967); this flow
regime is relatively stable for a large stream
in Wisconsin (Bean and Thomson 1944;
Sather and Johannes 1973). The upper sec¬
tions of river originate in, and flow through,
a large conifer bog surrounded by a sandy
outwash plain known as the “pine barrens.”
This area acts as a sponge by absorbing a
high percentage of the rainfall entering the
region, and then delivering it to the stream
through numerous springs at a uniform rate
(Bean and Thomson 1944). The high input
of spring flow is the defining feature of the
river system. This uniform source of abun¬
dant ground water creates stable flows and
a moderated thermal regime, which is cooler
Volume 82 (1994)
35
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1 . Relative abundance and distribution of fish species of the Brule River system
Common Name Scientific Name Origin Relative Information
Abundance 1 sources2
PETROMYZONTIDAE
Silver Lamprey Ichthyomyzon unicuspis Native R
Northern Brook Lamprey Ichthyomyzon fossor Native R
AU; SG; LB; MC; US
CH; OD; US
Sea Lamprey
Petromyzon marinus
Exotic C AU; LB; MC; MO; Wl
LEPISOSTEIDAE
Longnose Gar Lepisosteus osseus Native
ANGUILLIDAE
American Eel Anguilla rostrata Exotic
CYPRINIDAE
Common Carp Cyprinus carpio Exotic
Golden Shiner Notemigonus crysoleucas Native
Creek Chub Semotilus atromaculatus Native
Pearl Dace Margariscus margarita Native
Finescale Dace Phoxinus neogaeus Native
Northern Red belly Dace Phoxinus eos Native
Lake Chub Couesius plumbeus Native
Blacknose Dace Rhinichthys atratulus Native
Longnose Dace Rhinichthys cataractae Native
Hornyhead Chub Nocomis biguttatus Native
Common Shiner Luxilus cornutus Native
Emerald Shiner Notropis atherinoides Native
Spottail Shiner Notropis hudsonius Native
Mimic Shiner Notropis volucellus Native
Blacknose Shiner Notropis heterolepis Native
Brassy Minnow Hybognathus hankinsoni Native
Bluntnose Minnow Pimephales notatus Native
Fathead Minnow Pimephales promelas Native
CATOSTOMIDAE3
Silver Redhorse
Shorthead Redhorse
White Sucker
Longnose Sucker
Moxostoma anisurum Native
Moxostoma macrolepidotum Native
Catostomus commersoni Native
Catostomus catostomus Native
R
R
R
O
C
U
R
O
C
C
A
C
C
R
O
U
R
U
O
U
O
O
A
C
MO; Wl
AU; FB
LB
AU; FB; GR; LB; MC; MO; Wl
AU; FB; GR; LB; MC; MO; OD; Wl
AU; FB; MO; Wl
MO; Wl
AU; FB; MO; OD; Wl
MC, MO, OD; Wl
AU; FB; GR; LB; MO; OD; Wl
AU; FB; GR; LB; MC; MO; OD; Wl
MC; LB; MO; Wl
AU; FB; LB; MC; MO; OD; Wl
FB; LB; MO; OD; Wl
AU; GR; LB; MC; MO; Wl
AU
FB; MO; Wl
AU; FB; MO; Wl
FB; GR; MO; Wl
AU; FB; GR; MO; Wl
AU; MO; OD; Wl
AU; LB; MC; MO; OD; Wl
AU; FB; GR; LB; MC; MO; OD; Wl
AU; LB; MC; MO; OD; Wl
36
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
(Taxonomic names and order of families follows Robins et al. [1991].)
Main Areas of Distribution
lowest mile of the lower river; greatly reduced by lampricide treatments that began in 1959
lower two thirds of the mainstem and several tributaries prior to lampricide treatments that began in 1959;
now Minnesuing and upper Nebagamon creeks
much of the mainstem and larger tributaries until restricted by the lamprey barrier in 1986; now below
the barrier
one specimen reported from the lowest mile of the lower river
specimens reported from the estuary and Lake Nebagamon
lower river up to the lamprey barrier
the lowest few miles of the mainstem and Lake Minnesuing
scattered throughout the lower river; most common in slow water in the larger tributaries
lower river, Casey, Blueberry, and Wilson creeks, West Fork, and Lake Minnesuing
the lowest several miles of the lower river
lower river and most of the tributaries
lowest mile of the lower river
entire mainstem and Trask, Casey, and Blueberry creeks
riffle areas throughout the mainstem and in the larger tributaries
lowest mile of the lower river
lower river up to the lamprey barrier and Trask Creek
lower river up to the lamprey barrier and Lake Minnesuing
lower river up to the lamprey barrier
estuary
lowest mile of the lower river and Lake Minnesuing
lower river and larger tributaries
lowest mile of the lower river, Blueberry Creek, and Lake Minnesuing
lowest several miles of the lower river, Casey and Wilson creeks, Rocky Run, and the East Fork
lowest several miles of the lower river
lowest several miles of the lower river
entire mainstem and the larger tributaries
most common in the lowest several miles of the lower river but occasionally as far upstream as
Winneboujou
slow, deep sections of the mainstem, Nebagamon Creek, and lakes Nebagamon and Minnesuing
lowest mile of the lower river
lowest mile of the lower river and Lake Minnesuing
one specimen reported from the lower river at McNeil’s Bridge
lower river below the lamprey barrier
lakes Nebagamon and Minnesuing, uncommonly reported from scattered lotic sections
midsection and lower river except for extreme lowermost section and most of the tributaries
lowest mile of the lower river
occurrence/extent of reproduction in the Brule River system is unknown; probably strays from others waters
the larger tributaries and most of the mainstem, except for the extreme uppermost and lowermost sections
midsection of the mainstem and Nebagamon and Blueberry creeks
upper river mostly above Stone’s Bridge, Rocky Run, Blueberry Creek, Jerseth Creek, East Fork
entire mainstem (but less common in the extreme uppermost and lowermost sections) and the larger
tributaries
Volume 82 (1994)
37
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
'A = abundant - species often collected in large numbers.
C = common - species often collected in moderate numbers.
0 = occasional - species occasionally collected in moderate or small numbers.
U = uncommon - species infrequently collected in small numbers.
R = rare - species collected at rare intervals in very small numbers.
2AU - authors collections; CH - Churchill 1 945; FB - WDNR Fisheries Management file data (Brule
Area Office); GR - Greene 1 935; LB - WDNR lamprey barrier trap 1 986 - 1 993; MO - Moore and
Braem 1965; MC - McLain et al. 1965; PR - Pratt et al. 1992; OD - O’Donnell and Churchill 1954;
SC - Scholl et al. 1984; SG - Schuldt and Goold 1980; US - USFWS data files (J. Heinrich, pers.
comm.); Wl - Wisconsin Fish Distribution Study (cited by Fago 1992).
30’Donnell and Churchill (1954) reported the golden redhorse ( Moxostoma erythrurum ) to be
common in the estuary; based on current distribution information this is likely to have been a
misidentification — their specimens probably were shorthead redhorse or silver redhorse.
38
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
Main Areas of Distribution
not consistently reproducing in the Brule River system since 1979; rarely strays from other waters
upper river and most of the tributaries
transient from Lake Superior into the lower river up to the lamprey barrier
transient from stocking programs elsewhere in Lake Superior
lowest mile of the lower river
lowest several miles of the lower river
lower river, becoming increasingly common towards the mouth
lower river below the lamprey barrier
scattered throughout the mainstem and most of the tributaries in weedy, slow-water areas
lowest mile of the lower river
entire mainstem but least common in cold, headwater areas; Trask, Blueberry, and Nebagamon creeks
colder tributaries and headwater areas; present but less common throughout most of the mainstem
lakes Nebagamon and Minnesuing; rarely reported from scattered mainstem and tributary areas
lakes Nebagamon and Minnesuing; rarely reported from scattered mainstem areas
lakes Nebagamon and Minnesuing; rarely reported from scattered mainstem and tributary areas
lakes Nebagamon and Minnesuing, and Nebagamon Creek; rarely reported from scattered mainstem areas
lakes Nebagamon and Minnesuing, and Nebagamon Creek; rarely reported from scattered mainstem areas
lakes Nebagamon and Minnesuing; rarely reported from scattered mainstem and tributary areas
lakes Nebagamon and Minnesuing; uncommonly reported from the lower river below the lamprey barrier
lakes Nebagamon and Minnesuing, and Nebagamon Creek; rarely reported from the lower river below the
lamprey barrier
lower river up tp the lamprey barrier
entire mainstem, Trask and Blueberry creeks, East Fork, West Fork, and Lake Minnesuing
lowest mile of the lower river, Blueberry Creek, and Lake Minnesuing
lower river below the lamprey barrier; most common in the estuary
4ln the early 1900s, some specimens of grass pickerel ( Esox americanus) were reported from
Lake Nebagamon (Fago 1992). These reports were probably erroneous because Lake
Nebagamon is well outside of the known range of this species, and in early years northern pike
were sometimes referred to as grass pickerel and walleye were often called pickerel. It is
possible that true grass pickerel were introduced into Lake Nebagamon, but that a viable
population failed to become established.
5 There appear to be overlapping distributions of mottled and slimy sculpins throughout the
mainstem and in the tributaries, with mottled sculpins predominating in the mainstem and the
warmer tributaries and slimy sculpins predominating in the colder tributaries. Although the
limited data collected are consistent with this pattern, the conclusion is tentative because
separation of these species in the field is difficult and specimens from relatively few sites were
examined in the laboratory.
Volume 82 (1994)
39
40
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
in summer and warmer in winter than most
streams its size (both characteristics are most
evident in the upper reaches). The lower
river flows through a region of red clay that
contributes high runoff and associated tur¬
bidity and siltation to this section during
pluvial periods. A longitudinal gradient of
water temperature consists of less thermal
moderation as the river proceeds to its
mouth. In winter, the lower river is heavily
ice-covered, and during warm summers, wa¬
ter temperature in the lowest section is mar¬
ginal for salmonids. Patterns of spring flow
and water temperature probably existed his¬
torically much as they do today. The physi¬
cal and chemical characteristics of the river
system are summarized in Table 2; more de¬
tailed descriptions of the geology and topog¬
raphy of the watershed (Bean and Thomson
1944; Dickas and Tychsen 1969), the for¬
est cover types within the Brule Valley
(Fassett 1944; Thomson 1945), and the
chemical aspects of the river system (Bahnick
et ah 1969) are also available.
History of Human Activity Within the
Brule River Watershed as Related to
the Fish Community
The Brule River is renowned for its relatively
undisturbed natural setting, and indeed it
has weathered human encroachment with
less disturbance than most large streams in
Wisconsin. Nonetheless, changes to its
physical setting have occurred during the last
century that may have affected the fish com¬
munity. Early accounts of explorers’ jour¬
neys up the Brule River mention the pres¬
ence of one hundred or more beaver ( Castor
canadensis) dams that had to be broken
through (Marshall 1954). Locations of these
dams are not given, but they were probably
most abundant in the upper river above the
Cedar Island estate area (Fig. 1). Beaver were
heavily trapped in 1803-04, and the dams
were subsequently removed by the military
to facilitate use of the river as a water route
between the Great Lakes and the Mississippi
River prior to the development of a military
road network (Marshall 1954). These dams
may have hindered brook trout movement
to spawning areas (Marshall 1954), and ac¬
counts reporting excellent trout fishing are
commonly found only after removal of most
dams in about 1830 (O’Donnell 1944;
Marshall 1954; Jerrard 1956).
Aboriginal fish harvest from the Brule
River for centuries had undoubtedly been
modest; recent research indicates that the
Chippewa who settled along the southern
littoral of Lake Superior did not rely on fish
for their primary subsistence (Kaups 1984).
Access to the river for European settlers prior
to about 1850 was by water along Lake Su¬
perior, mostly from the city of Superior
(Marshall 1954). Travel upriver was time-
consuming and arduous, and the bark ca¬
noes used required skillful handling and fre¬
quent repair from damage caused by striking
rocks. Hence, the brunt of early fishing
probably took place near the mouth of the
river. The first record of fish caught from
the Brule River comes from the journal of
Michel Curot who set two gill nets near the
mouth of the river in 1804 and caught eight
unidentified fish (Wisconsin Historical So¬
ciety 1911).
Increased angling pressure on other sec¬
tions of river began when overland access
improved following the cutting of a crude
wagon trail from St. Paul to Bayfield in 1850
(Marshall 1954). This trail passed through
the town of Gordon (then called Amik), and
an early road cut from Gordon to the Brule
River about two miles south of Cedar Island
soon followed (Marshall 1954). As early as
1855, canoes and Mackinaw boats complete
with crews for fishing were advertised for
Volume 82 (1994)
41
Table 2. Mean physical and chemical characteristics of the lower Brule River1, the upper Brule River2, and 14 tributaries for
which reliable fish distribution information exists (compiled from data collected by the authors, from Fisheries Management
files [Brule Area and Superior Offices], or from Sather and Johannes 1973; n/a means no data available).
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42
TRANSACTIONS
^ower river refers to the stretch of river from U. S. Highway 2 north to Lake Superior
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
hire in the Superior newspaper ( Superior
Chronicle, August 28, 1855). A fishing ex¬
cursion on the Brule River in 1862 was re¬
ported to have caught “a lot of trout weigh¬
ing from four to five pounds each” ( Superior
Chronicle , August 23, 1862).
The period from 1870 through 1890 is
noteworthy in the history of the Brule River
because means of transportation for reach¬
ing the river improved dramatically, the
countryside around the river was rapidly
“filling up with immigrants” (Marshall
1954), and recreational use of the river
steadily increased from that time on (Hol¬
brook 1949; Marshall 1954). In 1870, the
Bayfield trail was cut from Superior to
Bayfield, and it quickly became an impor¬
tant artery (Marshall 1954; Jerrard 1956).
Near the Brule River the Bayfield trail fol¬
lowed the Copper Range and crossed the
river about two miles south of the present
County Highway FF bridge. At first, this
trail was usable by wagon only in winter un¬
til it was improved in 1876. During the early
1870s, Alexander McDougall caught bush¬
els of trout through the ice from the Cedar
Island spring ponds and shipped them by
dogsled to the Bayfield trail, then by horse
team to markets in Duluth (Marshall 1954).
Several articles in the Superior Times during
the mid- 1870s indicated that angling par¬
ties were making fine sport catches from the
river. During the 1880s another wagon trail
was cut from the town of Solon Springs
(then called White Birch) to the Blue Spring
area of the upper river just south of Stone’s
Bridge. Access to the river was eased further
by the development of a railway system. The
Northern Pacific rail line from Duluth to
Ashland was laid in 1883; this train crossed
the river at the newly established town of
Brule (Marshall 1954). In 1892, the laying
of the Duluth and South Shore Railroad
crossed the river at Station Rapids just south
of the present County Highway B bridge
(Marshall 1954). By 1884, Joe Lucius was
operating a guiding service for anglers on the
river, and by 1900, the river had been “well
discovered by anglers” (Marshall 1954).
Roads in the Brule area were first passable
by automobile in 1914.
Much of the virgin timber within the
Brule River valley was clear-cut beginning in
the early 1890s; this activity ushered in a new
era of human perturbation and development
in the region (Jerrard 1956). Two logging
dams (also called splash dams), one near the
mouth of the river and the other about two
miles north of the town of Brule (near the
present Boxcar Hole), were built to facilitate
movement of logs downriver (Marshall 1954;
Jerrard 1956). These dams, although thought
to be short-term, appear to have blocked the
migration route of coaster brook trout at
critical times and contributed to decline of
the fishery (O’Donnell 1944). Also signifi¬
cant was damage caused to the streambed
and shoreline areas when the dams were
breached and large numbers of logs were run
swiftly downstream (Marshall 1954). The
extent of siltation, erosion, and subsequent
flooding caused by timber cutting in ripar¬
ian areas is uncertain but likely was substan¬
tial. Clearly, the logging dams and lumber¬
ing operations negatively affected the fishing,
and fishing improved when these activities
were terminated (O’Donnell 1944; Jerrard
1956). The late 1800s and early 1900s also
saw increasing human activities regarding ag¬
riculture, road construction, and delivery of
utilities within the watershed; the extent to
which these activities may have harmed the
fish populations is unknown.
The written record of human dwellings in
the Brule River Valley begins with a Chip¬
pewa village at the mouth of the river dur¬
ing the late- 1850s that exported large quan¬
tities of Lake Superior fish by sailing sloop
Volume 82 (1994)
43
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
(Marshall 1954). At about the same time,
several commercial fishers of European de¬
scent also maintained their fishing stations
there, and exported large quantities of white-
fish ( Coregonus clupeaformis), sisco wet (a “fat”
morph of lake trout, Salvelinus namaycush ),
and lake trout ( Superior Chronicle , May 1,
1859; also Superior Times, July 15, 1875). In
1880, Samuel Budgett established the town
of Clevedon at the same location, but this
community persisted for only about five
years. Land on the present Cedar Island Es¬
tate was purchased in 1877 by two Minne¬
sota men for the purpose of using the series
of spring ponds on the property to commer¬
cially raise brook trout (, Superior Times , Janu¬
ary 27, 1877). In the late 1870s Frank Bow¬
man built a cabin on this property (Marshall
1954). During the early 1880s, Henry Clay
Pierce added to the now extensive Cedar Is¬
land Estate, and the first of the Winneboujou
Club cabins were built. These were the first
of the long-term dwellings that sprang up
along the banks of the Brule River during the
1880s (Marshall 1954; The Winneboujou
Club 1990). By the early 1900s, most of the
permanent dwellings that now exist along the
upper river had been completed. On the en¬
tire length of river, about four dozen perma¬
nent dwellings can now be found; this num¬
ber is slowly shrinking as properties become
available for public ownership through the
state’s land acquisition program.
In 1905, Pierce created a large fish hatch¬
ery at Cedar Island by blocking off the ex¬
tensive system of interconnected spring
ponds from the river; this action greatly re¬
duced the amount of spawning area for the
upper river brook trout population. These
spring ponds had gravel bottoms with areas
of upwelling ground water that provided ex¬
cellent spawning habitat. Early accounts
(summarized by O’Donnell 1944) affirm
that these spring ponds were the primary
spawning grounds of the original brook
trout population. A major decline in the
brook trout fishing apparently started about
1910, about five years after this spawning
area was separated from the river proper
(Jerrard 1956). Early reports by WDNR
fishery workers (O’Donnell 1944; O’Don¬
nell and Churchill 1954) recognized the tre¬
mendous spawning potential represented by
the Cedar Island spring ponds and recom¬
mended state acquisition of the ponds to
make them accessible once again as natural
spawning grounds.
The State of Wisconsin built a fish hatch¬
ery on the Little Brule River, at about its
midpoint (Fig. 1), in 1928. Still operating,
this hatchery has always been used to meet
statewide demands for domestic salmonids.
It uses the entire flow of the Little Brule
River for its water supply and creates a com¬
plete barrier to upstream fish movement.
Run-of-the-river fish hatcheries like the ones
on the Little Brule River and at Cedar Is¬
land, in addition to the positive effect of pro¬
ducing fish, have the potential to affect eco¬
system health in negative ways as well. They
can cause localized habitat destruction by
their placement, downstream water pollu¬
tion by their operation, and present a risk
for disease outbreaks that can spread to ad¬
jacent wild populations. Although they have
reduced critical habitats, there is no evidence
to suggest that either hatchery within the
Brule River system has negatively affected
ecosystem health through disease outbreaks
or substantial pollution.
The Brule River ecosystem is subject to
considerable recreational activity apart from
fishing, the byproducts of which could con¬
ceivably impact the fish community. Recre¬
ational canoe and kayak use is seasonally
heavy, averaging about 13,000 people annu¬
ally over the last ten years on the upper river
alone, and use on the lower river is prob-
44
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
ably similar in magnitude (C. Zosel,
WDNR, pers. comm.). The relatively pris¬
tine setting of the upper river, having two
stretches exceeding eight miles in length
without road crossings, provides a rare ca¬
noeing experience. Litter resulting inadvert¬
ently from spills is substantial given the vol¬
ume of boat traffic. Tubing was a popular
activity on the Brule River before it was
banned in 1981 because of conflicts between
tubers and other river users, and a myriad
of other concerns. Issues of crowding and
potential conflicts between anglers and rec¬
reational canoers have frequently surfaced
and may ultimately need to be addressed by
managers through some sort of usage allot¬
ment system as increases in both activities
continue. Campgrounds on the middle and
lower sections of river also draw numbers of
recreationists into the watershed.
Ecological Investigations on the Brule
River Watershed
While the Brule River has received fame as
a quality trout stream, its history has been
interspersed with perceived declines in fish¬
ing quality (Schneberger and Hasler 1944).
Consequently, the river has been the focus
of many studies to preserve and restore the
fishery. These efforts have compiled a wealth
of information about physical, chemical, and
biological aspects of the river.
In the early 1940s, the Brule River and
its watershed were subjected to one of the
most exhaustive interdisciplinary studies ever
done on a Wisconsin stream. The intent was
to evaluate the physical, biological, and
chemical characteristics of the watershed so
that an efficient and well-balanced manage¬
ment plan could be developed by the
WDNR (then the Wisconsin Conservation
Department, WCC). Eleven technical pa¬
pers were subsequently published in the
Transactions of the Wisconsin Academy of Sci¬
ences, Arts and Letters’, these were later reis¬
sued as one collection (Wisconsin Conser¬
vation Department 1954). Topics covered
included: topography and geology of the
Lake Superior basin (Bean and Thomson
1944); vegetation of the watershed (Fassett
1944; Thomson 1945); a history of fishing
(O’Donnell 1944); a survey of the aquatic
plants and bank flora (Thomson 1944);
parasites found on fishes (Fischthal 1945);
results of a four-year creel census (O’Don¬
nell 1945); bottom sediments (Evans 1945);
biology of the northern brook lamprey,
Ichthyomyzon fossor (Churchill 1945); and
physical, chemical, and biological attributes
of the river as habitat for trout (O’Donnell
and Churchill 1954). In 1946, a brief sum¬
mary of fishery recommendations emerged
which constituted the first WDNR manage¬
ment plan for the river. Recommendations
included stocking guidelines, public acqui¬
sition of the Cedar Island spring ponds, an
extended autumn season on the lower river,
initiation of creel and trout population sur¬
veys, and several riparian protection and ero¬
sion control measures.
Early efforts by the WDNR to sample
fishes using weirs occurred at Stone’s Bridge
and near the WDNR Ranger Station in 1943
(O’Donnell and Churchill 1954) and again
at Stone’s Bridge in 1958-60 (Fallis and
Niemuth 1962). During 1961-64, intensive
investigations of the anadromous brown
trout and steelhead populations by the
WDNR (Niemuth 1967, 1970) provided the
first substantial data set from which manage¬
ment applications could be drawn. Field
work included operation of fish weirs with
two-way traps during much of the open-wa¬
ter seasons, electrofishing sampling of some
mainstem reaches using mark/ recapture tech¬
niques to obtain population estimates, and
documenting upper river spawning sites. The
Volume 82 (1994)
45
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
weir locations were initially below Winne-
boujou and later just north of U.S. Highway
2. A similar investigation was repeated in
1978-79 (Scholl et al. 1984). This study also
included electrofishing surveys of several
Brule River tributaries. From 1957 to 1979
the U.S. Fish and Wildlife Service (USFWS)
operated an electrical weir one mile above the
mouth of the river in an effort to control
spawning of sea lamprey. However, in the
later years of its use, the TFM (3-tri-
fluoromethyl-4-nitrophenol) lampricide pro¬
gram had been developed, and this weir
served primarily to monitor the effectiveness
of chemical control. Some data on anadro-
mous salmonid populations were obtained
from the operation of this weir (Scholl et al.
1984); however, its operation caused an in¬
creased incidence of spinal deformities to the
salmonid populations as the downstream-mi¬
grating smolts passed through the electric
field (Devore and Eaton 1983) and may have
contributed to mortality of adult spawners
as well. Unfortunately, reliable quantitative
information about the salmonid populations
was difficult to obtain from any of the fish
weirs because of malfunctions caused by high
water or vandalism during portions of vir¬
tually every operating season.
Studies by two University of Wisconsin-
Madison graduate students also added to
knowledge about Brule River trout. Hunt
(1965) studied the importance of surface-
drift insects in the trout diet, and Salli (1962,
1974) reported on the early life history of
trout species in the lakes sector3 (Fig. 1).
Efforts to assess the sport fishery were
made in 1936, 1940, and in 1943-44
(O’Donnell 1945); in 1948-49 (Brasch
1950); in 1954 (Daly 1954); in 1962-64
(Niemuth 1970); in 1973 (Swanson 1974);
in 1978-79 (Scholl et al. 1984); and in
1984, 1986, 1990, and 1992 (this report).
Some of these were partial surveys with lim¬
ited objectives, while others were intended
to be comprehensive surveys of the river’s
mainstem. However, the size of the river,
difficulty of access to some areas, and mul¬
tiple uses by the public (which can render
car and canoe counts unreliable as indica¬
tors of pressure), have created difficulties in
obtaining reliable estimates of total angling
pressure and harvest. Furthermore, compari¬
sons among surveys are difficult because of
changing open seasons, daily bag limits, size
limits, stocking policies, and survey tech¬
niques over time. Results show that angling
pressure on the upper river has remained
fairly stable during recent years, but pressure
has decreased on the lower river since the
late 1970s (Table 3). On the upper river,
catch rates have increased in recent years,
while harvest has declined because of more
restrictive regulations and an increased ten¬
dency by fly-fishers to voluntary release their
catch (Table 3). On the lower river, catch
rates have remained fairly stable or declined,
while harvest of steelhead and brown trout
have declined substantially (Table 3).
The WDNR Bureau of Research initiated
a two-year pilot research study in 1983,
which was followed by a long-term research
initiative (1986-93). Both were carried out
jointly with the WDNR Bureau of Fisheries
Management. Topics addressed during this
research, and the reporting status of the
studies, are described in Table 4. This
cooperative effort, in conjunction with
direction supplied by the Brule River
Committee, has contributed to improved
management of the Brule River ecosystem.
History of the Fish Community
At least 63 fish species, including 1 1 exotic
species plus one cultured hybrid, have been
collected from the Brule River system (Table
1). Important shifts in the fish community
46
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
from the historical condition have unques¬
tionably occurred. These shifts appear to be
primarily related to establishment of exotic
species (additions of exotics or reductions in
populations of native species attributable to
sea lamprey control), although the loss of
coaster brook trout could be attributed to
over-exploitation or blockage of migration
routes. Many species have not experienced
demonstrable changes in their populations
over the last 50 years, suggesting little (if
any) change in habitat conditions. These
conclusions were reached through a com¬
parison of the fish fauna of 1987—91 with
that described from surveys done during the
mid- 1 940s (O’Donnell and Churchill
1954). However, the comparison is valid
only for common species because of differ¬
ences in equipment, survey techniques, and
survey effort between the two time periods.
Early surveys were less intensive than surveys
done from 1987-91, and early electrofishing
gear was less efficient. Hence, we have as¬
sumed that uncommon species not reported
by O’Donnell and Churchill (1954) were
missed by their sampling (as opposed to not
being present).
Among the lampreys (Petromyzontidae)
the differences between time periods are
striking, with two species of native lampreys
suffering from greatly reduced distributions
and population densities. Northern brook
lamprey were abundant in the 1940s
(Churchill 1945), but we found no speci¬
mens during electrofishing surveys in the
Brule River system in recent years. Silver
lamprey ( Ichthyomyzon unicuspis ) were once
plentiful in the lower river (McLain et al.
1965), but only a few specimens were found
there in the 1970s (Schuldt and Goold
1980). The only silver lamprey we have seen
in the Brule River was a specimen attached
to a migratory brown trout that could have
originated elsewhere. Using specialized gear
and techniques developed for sampling lam¬
preys, USFWS personnel have collected
three adult northern brook lampreys from
one mainstem area and Minnesuing Creek
over the last thirty years (J. Heinrich,
USFWS, pers. comm.). They have also col¬
lected moderate numbers of ammocoetes of
Ichthyomyzon from Minnesuing Creek and
the upper section of Nebagamon Creek, as
well as small numbers of specimens from
scattered mainstem areas. These specimens
could be northern brook lamprey, silver lam¬
prey, or some combination of the two. (It
is not presently possible to identify Ich¬
thyomyzon ammocoetes to species.) These
species were seriously impacted by 1am-
pricide treatments aimed at controlling the
sea lamprey (Table 1 - see also section on
the sea lamprey and the effects of control ef¬
forts). Sea lamprey are now common down¬
stream of the barrier/fishway, but were not
yet established in the 1940s.
The most common minnow species
(Cyprinidae) do not appear to have changed
in either relative abundances or distributions
since the 1940s. A number of occasional,
uncommon, or rare species reported here
(Table 1) were not reported by O’Donnell
and Churchill (1954), but this difference is
attributed to less efficient sampling during
the 1940s. Shifts in population distributions
of two less-common minnow species may
have occurred. The northern redbelly dace
( Phoxinus eos) occurred in the upper part of
the mainstem in the 1940s, but in 1987-
9 1 was confined to the lower river and tribu¬
taries. Similarly, the common shiner (Luxilus
cornutus) was found in deeper sections of the
upper river in the 1940s, but we found it
only in the lower river below the barrier/fish-
way.
Among the salmonids no major changes
in distributions of brook trout or steelhead
between time periods were apparent. Brown
Volume 82 (1994)
47
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 3. Angling pressure, catch, and harvest statistics for upper and lower sections
meaningful comparisons). Steelhead throughout the river system and brown trout from
marily of anadromous adults, whereas the < 13" categories are primarily stream-resi-
(reported in the “all brown trout” category only).
Lower River
'See section on regulations for daily bag and size limits in effect during each survey year;
sampling periods for the surveys included the entire regular open seasons on the upper river
in 1973, 1984, 1986, and 1992, and the regular open seasons plus extended early and late
seasons on the lower river in 1973, 1984, 1986. In 1990 the survey period on the lower river
coincided with the spring and autumn anadromous salmonid runs and was not extended
through the summer; also sampled was the time interval 1 July 1978-30 June 1979 during the
regular and extended seasons on the upper and lower sections of river.
48
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
of the Bois Brule River since 19731 (n/a = data not available or insufficient to provide
the lower river are reported in two categories; the >13" categories are comprised pri-
dent or juvenile anadromous forms. This distinction is invalid for upper river brown trout
Upper River
Volume 82 (1994)
49
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 4. Topics addressed during the 1984-93 research initiative
Investigation of density, biomass, age and growth, and DuBois et al. in press
species composition of stream-resident and juvenile
anadromous salmonids within discrete habitat zones
Experiment to determine the effectiveness of planting file data in the WDNR Superior Office
hatchery-reared steelhead smolts of Brule River origin (D. Pratt contact), file report forthcoming
The effects of lampricide treatments on the salmonids DuBois and Plaster 1993; DuBois and Blust
and their aquatic invertebrate food source 1994
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
species gained access to the river uninten¬
tionally through a variety of avenues includ¬
ing the opening of the St. Lawrence Seaway
via the Welland Canal (sea lamprey, Ameri¬
can eel [Anguilla rostrata ]), population ex¬
pansion from introductions to connecting
waters of the Great Lakes (rainbow smelt
[Osmerus mordax\ , common carp [ Cyprinus
carpio]), or from the release of ballast water
from transoceanic vessels in the Duluth/Su¬
perior harbor (ruffe [Gymnocephalus cer-
nuus\ , Pratt et ah 1992). The establishment
of exotic species in the Brule River has had
mixed effects that have sometimes been dif¬
ficult to assess, with some additions regarded
positively (the salmonids), but others caus¬
ing concern. For example, the sea lamprey
has been the focus of expensive control ef¬
forts in Lake Superior tributaries for over
thirty years, and the ruffe, while viewed as
a limited threat to the Brule River ecosys¬
tem, may ultimately warrant control efforts
in some areas of Lake Superior.
Of 52 native species, only 21 (40%) are
primary riverine species with viable popula¬
tions in lotic areas. The remainder are spe¬
cies that either are primarily found in the len-
tic habitats of Lakes Nebagamon and
Minnesuing, are residents of Lake Superior
that occasionally move into the lowest sec¬
tion of the lower river, or are locally rare
forms that stray into the river. The longnose
gar ( Lepisosteus osseus ), collected once from
the river (Table 1), is at the northern periph¬
ery of its range in Lake Superior. Lake stur¬
geon (Acipenser fulvescens ) have not been re¬
ported from the Brule River, but may have
entered it historically since they are present
in western Lake Superior and are known
from nearby rivers. Arctic grayling (Thy-
mallus arcticus) were not mentioned in any
of the early accounts about the Brule River,
but a population (now extinct) existed in
Michigan waters of Lakes Superior, Michi¬
gan, and Huron (Scott and Crossman 1973).
Therefore, Arctic grayling could have strayed
into the Brule River, and they are mentioned
along with brook trout in regulations per¬
taining to the river in the early 1890s.
Because the salmonids and the sea lam¬
prey have received the majority of research
and management attention, their life histo¬
ries and interactions in the Brule River eco¬
system are described in more detail in the
remainder of this section.
Brook Trout
Reports of tremendous fishing for brook
trout in the Brule River abound, particularly
for the period 1830-1900 (O’Donnell 1944;
Holbrook 1949; Marshall 1954). Historical
records of angling catches can be unreliable,
and must therefore be interpreted cautiously,
but the consistency of the early angling
records pertaining to brook trout is impres¬
sive (summarized by O’Donnell 1944). For
example, a U.S. Infantry Lieutenant wrote
in 1831 that “the river is exceedingly clear
and cold and is filled with thousands of real
mountain brook trout.” And in 1846, a ge¬
ologist charting the region in the interest of
mining companies wrote, “It surpasses all
other streams in its brook trout, some of
them, . . . weighing ten pounds. Its waters
colder and clearer, if possible than any other
river.”
The only salmonid native to the Brule
River, brook trout sometimes grew to a large
size with reports of 6- to 10-pound fish not
uncommon (O’Donnell 1944). That a part
of the original brook trout population was
of the coaster variety is virtually certain
(O’Donnell 1944). This conclusion is con¬
sistent with historical information about the
life history of brook trout in other Lake Su¬
perior tributaries (Bullen 1988). Coasters
appear to have been common along the
Volume 82 (1994)
51
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
south shore of Lake Superior during the
1800s (Shiras 1935), and the Brule River
may have been a major producer.
However, by the 1870s evidence of pub¬
lic concern about overharvest of brook trout
in the region was beginning to surface. A
quote from the Superior Times (February 24,
1876) is illustrative: “while our legislature is
devoting time and money to the propaga¬
tion of fish within the state it is a pity they
do not stop the wholesale slaughter of brook
trout through the ice in the Lake Superior
counties.” By the early 1890s, the Brule
River brook trout fishery had begun to de¬
cline (O’Donnell 1944). Excessive angling
catches undoubtedly occurred frequently
during the late 1800s and early 1900s as
regulation of recreational angling was very
liberal with no daily bag in effect until 1905
(see section on regulation of the fishery).
This history of substantial and at times ex¬
cessive harvest has continued to the present
as increases in angling pressure have accom¬
panied increasing harvest restrictions.
Hence, over-exploitation is a major factor
implicated in the reduction of the brook
trout population.
Lumber interests started cutting the vir¬
gin timber in the Brule Valley in 1892, and
the logging dams they built allegedly dam¬
aged the coaster population by blocking
their migration route at critical times
(O’Donnell 1944). A law was on the books
at that time requiring a fishway at any dam
or obstruction on the Brule River (Chapter
251, Laws of 1891). Unfortunately, non-
compliance was rife, and a 1906 article
(quoted by O’Donnell 1944) explicitly states
that there were no fishways at the logging
dams on the Brule River. Siltation associated
with poor forestry practices likely also con¬
tributed to the decline of the fishery
(Holbrook 1949; Marshall 1954). Interspe¬
cific competition with heavily stocked
brown trout and rainbow trout during the
1920s and 1930s may have also negatively
affected the brook trout population. Coaster
brook trout were apparently extirpated from
the river by the mid- 1940s, with the latest
reliable record being of a 24-inch fish ob¬
served spawning in the upper river in 1944
(O’Donnell and Churchill 1954).
Efforts to bolster the sagging brook trout
fishery included supplemental stocking of
domestic strains, which began in 1894 and
continued steadily for over 80 years, and in¬
troductions of exotic species (early plantings
summarized by O’Donnell 1945). Stocking
of brook trout was terminated in 1979 be¬
cause of emerging evidence from the fisher¬
ies literature, now even more firmly estab¬
lished, that stocking domestic strains of trout
on top of healthy wild populations often has
more negative than positive effects (White
1989; Goodman 1990). This represented a
major change in management policy which
may have contributed to some recent recov¬
ery of the brook trout population.
Early records about the strains of brook
trout stocked in the Brule River are sketchy,
and undocumented plantings by wealthy
private citizens or citizen groups may have
occurred. Domestic brook trout strains,
which have been systematically selected from
fast-growing, early-maturing brood stocks,
can have a significant reproductive advan¬
tage over wild fish. Gene flow from these
strains may have altered, to an unknown de¬
gree, the genetic structure of the original
brook trout population, which was uniquely
adapted to the physical setting of the river.
The present brook trout population is
largely confined to the upper (southern) half
of the river and most of the tributaries (Fig.
2) where ice-free conditions for long
stretches during winter provide evidence of
abundant spring flow. They exist sympatri-
cally with populations of exotic salmonids
52
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
Fig. 2. Present distribution of juvenile coho salmon and brook trout of all age classes
in the Bois Brule River system showing areas of highest population density
in sufficient numbers and sizes to provide
acceptable fishing. The Brule River contains
the largest population of brook trout of all
Wisconsin streams draining into Lake Supe¬
rior. However, they are the least abundant
of the three primary salmonid species in the
Brule River system (DuBois et ah in press).
Brook trout spawn in slower-flow areas, of¬
ten near springs where small-sized gravel and
the upwelling ground water conditions they
require are suitable. Spawning areas in the
Brule River have not been well documented,
but are probably scattered widely through¬
out the upper river and several tributaries in
spring pond areas and other areas of reduced
flow. Spawning likely could be enhanced by
dredging spring pond areas adjacent to the
main channel along the upper river (Carline
1980).
Steelhead
Steelhead were first introduced to the Brule
River in 1892 (O’Donnell 1944), and stock¬
ing of a variety of Pacific Coast strains con¬
tinued periodically through 1981 (see
Volume 82 (1994)
53
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 3. Present distribution of juvenile steelhead and brown trout of all age classes
in the Bois Brule River system showing areas of highest population density
MacCrimmon and Gots 1972, and Krueger
and May 1987a). Steelhead have become the
most abundant salmonid in the river system
(DuBois et al. in press). This species has a
strong migratory tendency, and it appears
that the entire Brule River population is
anadromous, although this apparently was
not the case originally (O’Donnell 19 44).
Steelhead inhabit most of the river system
as juveniles (Fig. 3), but descend into Lake
Superior as smolts after one, two (usually),
or three summers in the river. Once in the
lake, these fish grow to a large size and then
return to the river to spawn after one, two,
or three more years. Upon reaching matu¬
rity they spawn annually (Swanson 1983),
with a few spawning every other year
(Seelbach 1993).
About 13 major spawning areas used by
steelhead were identified by O’Donnell and
Churchill (1954) and Niemuth (1967, 1970)
in riffle areas between Stone’s Bridge and
Winneboujou. Spawning in the lower river
is even more significant, but it is more diffi-
54
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
Fig. 4. Five-year (1989-93) mean weekly distribution of anadromous salmonids as¬
cending the Bois Brule River on annual spawning runs (mean number per year for
each species during 1989-93 in parentheses)
cult to document the specific locations used
and hatching success because of higher tur¬
bidity and deeper water. Year-class strength
of juvenile steelhead has been stable in the
upper river, but highly variable in the lower
river (DuBois et al. in press). This variation
has apparently resulted largely from environ¬
mental factors and may not be closely related
to numbers of spawning adults. The lower
river is less stable than the upper river in
terms of both its flow and temperature re¬
gimes. We speculate that a critical factor as¬
sociated with year-class strength in the lower
river pertains to spring flooding, which may
have a devastating effect on eggs and newly
hatched fry (Seegrist and Gard 1972). The
limited data available suggest that when there
is little flooding in the spring when the
young-of-the-year are small (less than about
2 inches), survival is high. Also, during warm
springs, growth is faster, allowing young fish
to grow more quickly through the critical
“window of time” when they are vulnerable
to spring flooding.
Though all steelhead spawning apparently
occurs in the spring (some autumn spawn¬
ing is possible but unlikely), two distinct
migration patterns have emerged: a larger
autumn run of fish ascends the river from
September through December and overwin¬
ters in deeper holes throughout the main-
stem of the river, and a smaller spring run
begins to ascend the river in February or
early March and continues through May
Volume 82 (1994)
55
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
(Fig. 4). A study to investigate the possible
genetic distinctness of these runs did not
show significant differences (Krueger and
May 1987a). It is therefore probable that the
autumn and spring spawning runs are actu¬
ally one extended run interrupted by a tem¬
perature-related cessation of migration cues
during winter. The large autumn run on the
Brule River is unique among Wisconsin’s
tributaries to Lake Superior, the others hav¬
ing substantial spring runs and only small
autumn runs (if any). The reason(s) for this
difference among streams is not clearly un¬
derstood, but the availability of suitable
overwinter holding areas in the Brule River,
the relative seasonal stability of its flow, and
the moderated thermal regime that contrib¬
utes to its becoming ice-free earlier in the
spring than most other streams, may all be
important factors.
Evidence mounted during the early and
mid-1980s that the steelhead population of
the Brule River had declined disturbingly
since the late 1970s and may now be at only
a small fraction of its former abundance. For
example, the estimated steelhead harvest
during 1978-79 (about 7,000) was roughly
similar in magnitude to the estimates of the
entire run sizes in recent years (Fig. 4).
Other tributaries along Wisconsin’s Lake
Superior shoreline showed a similar pattern
of declining runs since the 1970s suggest¬
ing that the western Lake Superior steelhead
stocks were collectively declining and may
have been approaching a collapse (B.
Swanson, WDNR, pers. comm.). Causes of
this decline (which unfortunately coincided
with a year-round open season on the Brule
River during 1983—85) were unknown, but
overharvest in Lake Superior as well as in the
streams was strongly suspected.
In response to this perceived threat to the
steelhead stocks, the WDNR enacted more
restrictive size and bag limit regulations in
1989, and embarked on a limited-term steel¬
head stocking experiment for the Brule River
to bolster its population. The stocking plan
called for an annual egg-take operation
(about 100,000 eggs annually) from wild
Brule River steelhead and subsequently rais¬
ing these juveniles to smolt size (approach¬
ing or exceeding 7lh inches). The goal of
the stocking program was to release 50,000
functional smolts at various locations on the
lower river each May until the program was
no longer needed. Anglers have also re¬
sponded by practicing more catch-and-re-
lease angling for steelhead on a voluntary
basis (D. Pratt, unpubl. data). In 1993, the
minimum legal size was further increased to
26 inches to allow maiden spawners to
spawn at least once before entering the har¬
vest (see section on regulations for complete
description of steelhead sport harvest restric¬
tions) . At least several years will be required
to assess the effectiveness of these measures.
The wisest course of action for steelhead is
to manage with conservative regulations;
however, highly variable year-class strength
related to environmental factors outside of
management control will lead to large varia¬
tions in annual angling quality, regardless of
best management practices. Additional de¬
scriptive aspects of the Brule River steelhead
population have been reported by Niemuth
(1970) and Scholl et al. (1984).
Brown Trout
Brown trout were introduced to the Brule
River by the WCC in 1920 (O’Donnell
1945). However, they may have already be¬
come established in the river from stockings
elsewhere in the Lake Superior basin
(Krueger and May 1987b). Many domestic
strains of brown trout were undoubtedly
stocked before stocking of this species was
terminated in 1974. A self-sustaining popu-
56
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
lation of brown trout developed early on,
and they are now common throughout
much of the river system (Fig. 3).
Two ecologically distinct groups of au¬
tumn-spawning brown trout coexist: a
stream-resident component provides a chal¬
lenging upper-river fishery and an anadro-
mous (lake-run) component exhibits a life
history strategy similar to steelhead. Anadro-
mous brown trout spawning runs begin in
July, peak in August, and extend into Oc¬
tober (Fig. 4). Juveniles reside in the river
for one (usually), two, or rarely three years
before smoking in the spring or autumn.
They then usually spend two years in the
lake before returning to the river to spawn
at age three or four. Although spawning
need not result in the death of brown trout
in an obligatory sense, as is the case with
Pacific salmon, repeat spawning is relatively
uncommon, and high natural mortality ap¬
pears to be associated with spawning.
Brown trout sampled from western Lake
Superior tributaries were found to differ ge¬
netically among drainages, between anadro-
mous and stream-resident life histories, and
among locations within the Brule River
drainage (Krueger and May 1987b). Ana-
dromous brown trout provide a rare trophy
fishery, but have been generally less popu¬
lar than steelhead because they are often
harder to catch. Additionally, they are sus¬
ceptible to mortality from furunculosis,
caused by the bacterium Aeromonas sal-
monicida. Some fish carry the disease with¬
out symptoms, but outbreaks of furunculo¬
sis have sometimes reached epidemic
proportions in the Brule River. Although fu¬
runculosis was observed prior to intensive
weir study (Niemuth 1967), stress associated
with handling large numbers of brown trout
at weirs may have aggravated the prevalence
of the disease. Warm river temperatures in
August when most anadromous brown trout
ascend the river have also been implicated
in activating symptoms.
Resident and anadromous brown trout
spawn in many of the same mainstem areas
used by steelhead, but are more spatially re¬
stricted than steelhead in the lower river and
the tributaries. This restriction may be re¬
lated to the difficulty posed for overwinter¬
ing eggs by anchor ice in the less-thermally-
moderated lower river and by typically lower
water levels in the tributaries during their
autumn-spawning period. Information on
size and age structures and other descriptive
aspects of the anadromous brown trout
stocks have been reported by Niemuth
(1967) and Scholl et al. (1984).
Pacific Salmon
Three species of Pacific salmon have been
found in the Brule River in recent years; all
are strays from stockings by neighboring
states and the Province of Ontario. Their
sudden appearance serves as a reminder that
the Brule River is not an isolated system, but
rather, is intimately tied to the ecology of
the surrounding region. Scott and Crossman
(1973) described the life histories of Pacific
salmon in the Great Lakes, and Scholl et al.
(1984) provided early information about the
population characteristics of these autumn¬
spawning species in the Brule River.
First documented in the Brule River in
1973, coho salmon have established a viable
though widely fluctuating population. Their
life history strategy involves one year of
stream residency for juveniles, outmigration
as smolts in May, and growth in the lake for
one or two years before returning to the
stream to spawn and die. Adults have con¬
tributed significantly to the sport catch in
recent years. Coho salmon have so far shown
a three-year cycle of abundance in the order
of a small-run year, an intermediate year, and
Volume 82 (1994)
57
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
a large-run year. Numbers of spawners and
the resulting young are strongly correlated.
Spawning runs have peaked in mid to late
September, but substantial movement has
occurred throughout autumn and extended
into winter (Fig. 4). Coho salmon spawn
successfully throughout the upper river and
tributaries, favoring smaller riffles near head¬
water areas (Fig. 2). Juveniles fare well in the
slow, deep, alder-choked sections of the up¬
per river south of Stone’s Bridge; because
these areas are extensive, the Brule River will
likely remain a strong coho salmon producer.
Establishment of a viable chinook salmon
population has developed more slowly than
that of the coho salmon, although the first
adults were also documented in 1973. Dur¬
ing the late 1970s and early 1980s, small
numbers of juveniles were found only in
Blueberry and Nebagamon creeks and in
mainstem riffles close to the confluence with
Nebagamon Creek (Fig. 1). Since 1988,
chinook salmon have spawned over a slightly
wider range of locations, although still cen¬
tered in the same general area. Year-class
strength has also shown modest increases
(DuBois et al. in press). Chinook salmon in
the Brule River smolt mostly as young-of-
the-year in May and June, with the remain¬
der smoking during autumn or the follow¬
ing April/May. After smoking, chinook
salmon spend up to five years in the lake
(four years is most common) before return¬
ing to the river to spawn. Spawning runs,
which have peaked from mid-August
through September (Fig. 4), have contained
modest numbers of spawners; the extent to
which their population size or distribution
may change is unknown.
Pink salmon have also been found in the
Brule River, although not in appreciable
numbers since 1979 (Scholl et al. 1984).
Their potential for establishment in the
Brule River appears limited.
The Sea Lamprey and Effects of
Control Programs on Other Fishes
Sea lamprey were introduced to the upper
Great Lakes through the opening of the
Welland Canal and were first reported in
Lake Superior in 1938 (Becker 1983). Al¬
though early records of their spawning in the
Brule River are sketchy, they had developed
a viable population in western Lake Supe¬
rior by the mid-1950s. Primary spawning
areas in the Brule River are not well known
but likely included both mainstem and
tributary riffles near silt beds for ammocoete
habitat. Sea lampreys quickly caused serious
damage to the salmonid populations of Lake
Superior (National Research Council of
Canada 1985). The Brule River was one of
the largest tributary producers of sea lam¬
preys in the Lake Superior basin, yielding
approximately one-third of the entire catch
from Lake Superior streams (McLain et al.
1965). Beginning in the mid-1950s, control
programs initiated by the USFWS began to
dramatically reduce sea lamprey recruitment
to Lake Superior by disrupting their spawn¬
ing and larval phases through use of me¬
chanical weirs, electrical weirs, and later, se¬
lective lampricide treatments (Smith and
Tibbies 1980). In the Brule River, sea lam¬
prey control has included an electrical weir
one mile upstream from the mouth of the
river from 1957 through 1979, selective
lampricide treatments using TFM at three-
year intervals in the entire mainstem and
throughout most of the tributaries from
1959 through 1986, and a mechanical bar¬
rier used in conjunction with chemical treat¬
ment only below the barrier since 1986.
Although TFM treatments of the Brule
River successfully reduced sea lamprey re¬
cruitment, deleterious effects on some non¬
target organisms were observed, and the
monetary cost of treatments was high
58
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
(Gilderhus and Johnson 1980). Concern
about these negative aspects led to the con¬
struction in 1984 of a sea lamprey barrier,
located about seven miles upstream from the
mouth of the river (Fig. 1). Initially, a low-
head dam with jumping pools to allow mi¬
gratory salmonids to pass upriver was built.
However, this structure did not pass salmo¬
nids during all water conditions, and plans
for remodeling were formulated. A recon¬
struction, including an effective fishway, was
begun in 1985 and completed in March of
1986. The fishway included a viewing win¬
dow which has proven to be a valuable re¬
search tool to obtain data on salmonid run
numbers and other population statistics.
This barrier system functions via use of a
low-head dam with an overhanging metal lip
within the fishway of a height surmountable
by leaping salmonids but insurmountable to
sea lampreys. The primary barrier which
crosses the entire width of the river is higher
than the fishway barrier and is impassable
by all sea lampreys and most migratory
salmonids. During autumn when no sea
lamprey movement occurs, the fishway bar¬
rier is removed, allowing free upstream ac¬
cess to all fish species. The USFWS will pe¬
riodically monitor the entire river and
tributaries for presence of ammocoetes to de¬
termine any need for further treatment of
upriver areas. Because the barrier appears to
stop all sea lamprey movement, routine
TFM treatments will now be made only
downstream of it. Sections of the Brule River
above the sea lamprey barrier were last
treated in 1986 to kill ammocoetes produced
before its completion.
All species of lampreys are highly sensi¬
tive to TFM, and populations of three spe¬
cies of endemic lampreys have been greatly
reduced or eliminated from treated streams
within the Lake Superior basin (Schuldt and
Goold 1980). In the Brule River, the silver
lamprey and the northern brook lamprey
were once abundant, and their populations
have been greatly reduced by repeated treat¬
ments (Table 1). It is not known if silver
lamprey were historically indigenous to sec¬
tions of the Brule River above the sea lam¬
prey barrier.
Other groups of aquatic organisms are af¬
fected to different degrees by TFM. Lam-
pricide treatments usually have substantial
negative effects on a relatively few forms of
invertebrate life (Gilderhus and Johnson
1980; Dermott and Spence 1984; Mac-
Mahon et al. 1987), and they do not usu¬
ally have severe direct effects on salmonid
populations (Dahl and McDonald 1980).
Secondary effects on salmonids due to a re¬
duced invertebrate food supply also are un¬
likely to be severe (Merna 1985; DuBois and
Blust 1994). However, other families of
fishes, particularly ictalurids (catfishes) and
catostomids (suckers), are quite sensitive to
TFM. Stonecats ( Noturus flavus) in the
lower Brule River and in other western Lake
Superior tributaries were so severely affected
by TFM treatments that it was initially
feared that they might have been eliminated
from Lake Superior (Dahl and McDonald
1980). Fortunately, untreated refugia appar¬
ently existed for a portion of the stonecat
population. Populations of other fishes in¬
digenous to the Brule River may have been
reduced because of TFM treatments. Dahl
and McDonald (1980) provide a thorough
discussion of the known effects of sea lam¬
prey control on non-target fishes in the
Great Lakes.
Control of sea lamprey spawning in the
Brule River will remain an important fish¬
eries management priority. Although re¬
search on alternative methods of sea lamprey
control is ongoing, mechanical barriers and
chemical treatments remain the most suc¬
cessful of the practical options.
Volume 82 (1994)
59
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Habitat Management
The history of the Brule River is dotted with
numerous efforts to preserve the integrity of
the physical habitat, enhance habitat for
salmonids, and stabilize riparian areas. These
efforts fall into five categories: (1) preserva¬
tion of riparian areas, (2) in-stream habitat
enhancements, (3) beaver control and dam
removal, (4) bank stabilization in red clay
areas subject to slippage, and (3) dredging
projects.
Preservation Efforts
A major factor in the preservation of the
Brule River ecosystem has been the protec¬
tion afforded by state stewardship acquisition
of land bordering the river. The Brule River
State Forest was established in 1907 when
Frederick Weyerhauser deeded 4,320 acres
to the state. Land acquisition since has added
to that total as funds have allowed. In 1939,
the boundaries of the Brule Forest were ex¬
tended to include the entire Brule River cor¬
ridor. Presently about 40,000 acres of land
are under state ownership, which represents
about 80% of the total acreage within the
boundary of the Brule River State Forest and
includes about 50% of the river frontage (C.
Zosel, WDNR, pers. comm.).
Another major factor contributing to
preservation of the Brule River ecosystem
has been the excellent stewardship practiced
by private riparian landowners over many
years (Holbrook 1949). Brule River Preser¬
vation, Inc., is a public nonprofit corpora¬
tion including over 20 landowners from the
upper river dedicated to preserving the Brule
River and fostering sound ecological man¬
agement for its use. The Nature Conser¬
vancy (TNC), an international organization
dedicated to preserving unique natural areas,
has a Conservation Easement Grant Pro¬
gram in effect on much of the upper river
between Blue Spring and the WDNR Brule
Area Headquarters (Fig. 1). This program
features agreements between individual land-
owners and TNC whereby the landowners
voluntarily restrict certain rights of use and
development on their lands in perpetuity in
order to ensure that these lands are protected
against unwise commercial development and
ecological degradation.
In-Stream Habitat Enhancement
The WDNR has been at the forefront nation¬
ally in the development of in-stream habitat
enhancement techniques for salmonids. Con¬
sequently, much is now known about the
identification of environmental deficiencies
and the application of appropriate structural
remedies to Wisconsin’s streams (White and
Brynildson 1967; Hunt 1988, 1993). Tech¬
niques used in the Brule River have included
wing deflectors and bank covers, debrushing
and installation of brush bundles, and re¬
moval of downed trees and other debris. Un¬
fortunately, most of these efforts were under¬
taken before knowledge about effective
techniques had been refined. A project of
“stream improvement” was started in 1936
using Works Progress Administration labor.
A total of 286 structures were installed in the
river including deflectors, bank covers, and
other stream enhancement devices, many of
dubious value for creating trout habitat
(O’Donnell 1944; Holbrook 1949; Marshall
1954). This effort appears to have been fo¬
cused on making the river easier to canoe
(O’Donnell 1944). Some of these structures
still exist, either complete or as remnants, be¬
low County Highway P, below Stone’s
Bridge, in the Winneboujou area, and near
the WDNR Area Headquarters. Additionally,
the Civilian Conservation Corps installed
structures, planted willows, and “clean [ed]
60
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
out large amounts of down trees and other
materials” at about the same time (O’Don¬
nell 1944, p.29). The beneficial role of large,
downed timber in shaping stream morphol¬
ogy and creating salmonid habitat has only
been realized in recent decades (Harmon et
al. 1986; Bisson et al. 1987). Current think¬
ing now favors adding large woody debris to
the stream to compensate for wood removed
by earlier enhancement efforts or lost for
other reasons4. During the 1960s, a series of
rock deflectors was installed below the State
Highway 13 bridge by the Brule River
Sportsmen’s Club, Inc. Also in the 1960s, the
Douglas County fish and Game League (un¬
der cooperative agreement with the WDNR)
constructed rock wing deflectors in several
lower river locations to provide cover for
salmonids and deflect current away from red
clay banks. No follow-up evaluations were
made of any of these early efforts.
Habitat enhancement efforts by the
WDNR within the Brule River watershed
since the 1960s have focused on riparian
debrushing and installation of brush bundles
on inside bends in some tributary areas
choked with speckled alders ( Alnus rugosa).
Riparian debrushing lets sunlight into the
stream for aquatic plant growth and encour¬
ages physical improvements in the stream
channel, while brush bundles provide cover
for trout fry and accelerate a favorable chan¬
nel-constriction process (Hunt 1979). Such
efforts on the East and West forks of the
Brule River appear to have improved trout
habitat, but were not evaluated to document
their impact on trout populations. During
1978-91, a project on the Little Brule River
below the state trout hatchery to remove all
beaver dams and riparian alders and install
brush bundles was conducted and evaluated
(DuBois and Schram 1993). Natural repro¬
duction in both treatment and control sec¬
tions improved during the post-treatment pe¬
riod. However, numbers of legal-size brown
trout declined markedly in both treatment
and control sections following treatment, a
result that could have been due to increased
fishing pressure brought on by publicity sur¬
rounding the project, the improved fishability
of the debrushed stream segment, or move¬
ment of large brown trout out of the stream.
Beaver Control and Dam Removal
Effects of beaver dams on trout habitat have
generally been regarded as negative in Wis¬
consin although they are considered benefi¬
cial in small, high-gradient streams. Negative
effects are most likely to occur on streams of
low-to-moderate gradient where dams may
contribute to warming of water, hinder
salmonid movement and spawning, cause
silting-in of gravel areas important for pro¬
ducing insects, and produce poor channel
characteristics (summarized by Avery 1983).
Traditionally, beaver have been regulated by
trapping because of the value of their fur. For
example, heavy trapping of beaver on the
Brule River in 1803—04 drastically reduced
their numbers (Marshall 1954). However,
beavers are prolific, and animals from sur¬
rounding areas tend to recolonize trapped-
out areas, creating an unceasing cycle. Fur¬
thermore, intensity of trapping effort
fluctuates because of unstable fur prices. Re¬
cent low fur prices and excellent habitat have
resulted in a large beaver population in
northern Wisconsin. The Brule River and
tributaries have been on various special ex¬
tended beaver trapping seasons since the
early 1960s, including a liberal open season
since the mid-1980s. The Brule River water¬
shed was included in a WDNR beaver sub¬
sidy program from 1986-88 that provided
a financial incentive for trappers to control
populations in designated areas. Since then,
a trapper from the Animal Plant Health In-
Volume 82 (1994)
61
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
spection Service of the U.S. Department of
Agriculture has worked under WDNR direc¬
tion to remove beaver from within the Brule
River watershed and other salmonid tribu¬
taries to Lake Superior.
Beaver dams on the Brule River generally
occur only in the upper river above Stone’s
Bridge and in several tributary areas. Al¬
though historically these were probably also
the areas of heaviest beaver activity, dams
may have occurred further downstream as
well. Numerous recent excursions to remove
beaver dams from the upper Brule River have
been made by WDNR workers, members of
area sports clubs, and other citizens, but re¬
sults have been short-lived, especially if bea¬
ver were not also removed. A recent habitat
development project on the Little Brule
River evaluated the effects of beaver dam re¬
moval and riparian debrushing on the physi¬
cal conditions of the stream channel and on
the salmonid populations (DuBois and
Schram 1993). Although physical changes in
channel morphometry following these ma¬
nipulations were impressive, salmonid popu¬
lation responses were mixed, and the bene¬
ficial aspects could not be attributed solely
to dam removal.
Bank Stabilization
The clay soils in the Brule River watershed
appear to be geologically young and under¬
going a high rate of natural erosion. When
Europeans settled in this area, their lumber¬
ing, road construction, and agricultural ac¬
tivities removed the established mixed-coni¬
fer forest cover type and altered drainage
patterns in ways that accelerated this pattern
of erosion. Present-day activities, although
more carefully controlled, continue to aggra¬
vate the erosion process.
Erosion of the red clay soils of the lower
Brule River Valley has the potential to nega¬
tively affect salmonid populations. The po¬
tentially most damaging effect is from sedi¬
mentation, which can inhibit aquatic inver¬
tebrate life and reduce salmonid spawning
success by causing high egg and larval mor¬
tality. In extreme cases, turbidity can also re¬
duce feeding success of visually feeding
salmonids and inhibit proper gill function
(Berg and Northcote 1983). Llowever, tur¬
bidity is probably not an important limiting
factor for salmonids in the lower Brule River
because other tributaries to Lake Superior in
Wisconsin are known to have longer-term
turbidity episodes yet have contained robust
salmonid populations in recent decades.
A red clay interagency committee was
formed of state and federal agencies in 1955
to investigate land-use problems on the red
clay soils of northwestern Wisconsin. The
goals of this committee were to determine
the causes of red clay sedimentation in area
lakes and streams and to study means of ero¬
sion and sedimentation control. Experimen¬
tal work to reduce clay erosion was done on
the Brule River in a few areas using gabions
and riprap to stabilize bank slippage. Some
successes were achieved in areas of less ex¬
treme slippage, but the efforts were expen¬
sive and the results obtrusive in a natural set¬
ting. Various mulchings and plantings were
also tried in localized areas. State forest man¬
agement goals now call for specialized tim¬
ber management in steeply sloped red clay
areas, with the long-range objective of re¬
turning the area to mixed-conifer forest. This
plan may eventually lead to reduced bank
erosion and slippage, but decades will be re¬
quired to assess the results.
Dredging Projects
Dredging of spring ponds in Wisconsin can
benefit brook trout spawning (Carline
1980), but indiscriminate dredging of
62
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
streams can produce harmful physical
changes such as degradation of the stream-
bed (headcutting) and bank erosion (Kanehl
and Lyons 1992). Several dredging projects
have been carried out on the Brule River
with the goal of enhancing trout habitat by
creating deeper pools and exposing gravel
substrates for spawning and increased inver¬
tebrate production. In the late 1920s, the
WDNR dredged the east side of Big Lake.
Blue Spring and a short stretch of river above
Stone’s Bridge were dredged by a private in¬
terest in the late 1960s. These projects were
never evaluated to document benefits that
may have accrued. In 1967, Douglas County
and the Douglas County Fish and Game
League tried to deepen and straighten the
mouth of the Brule River by dredging. The
attempt was futile, however, as the river
quickly reverted back to its original form.
Regulation of the Fishery
Restrictions on fishing on the Brule River
have usually followed the regular statewide
trout and salmon regulations, with various
extended special seasons on the lower river.
In the early 1900s the four northern coun¬
ties bordering Lake Superior were sometimes
subjected to different open seasons on trout
than the rest of the state. The many changes
occurring over the history of regulation in
open season dates, daily bag limits, and
minimum sizes are described below in chro¬
nological order.
Open Seasons
Regular Season. The first restriction on
trout fishing enacted in Wisconsin was a re¬
duction in the length of the statewide open
season from 12 months to 8 months in
1858. In 1878, the open season was further
reduced to 5 months. From 1891 through
1898 the open season on the Brule River was
greatly reduced to just 26 days in August
(Chapter 138, Laws of 1891). Reasons for
this restriction were not stated, but we note
its concurrence with the era of intensive log¬
ging and use of logging dams to transport
logs downriver en masse; release of these
dams would have created serious hazards for
anglers downstream. The length of the open
season has varied since, but has usually been
between 4 and 5 months. The season opener
has varied between mid-April and mid-May;
the closing date for the regular season has
also been variable, occurring sometime be¬
tween August 20 and September 30.
Special Extended Seasons. Various ex¬
tended seasons have been enacted for the
lower river between U.S. Highway 2 and
Lake Superior to allow increased angling op¬
portunities during anadromous salmonid
spawning runs. These seasons have included
a special early season, starting in spring some¬
time prior to the regular season and running
to the regular season opener, and a special late
season, starting in autumn after the regular
open season and extending for various peri¬
ods of time. Season extensions began in 1935
with a special early season starting on May 1
(at that time the regular season opener was
on May 15). The starting day for the special
early season has since varied, but has most
often been the Saturday nearest April 1 . Spe¬
cial regulations for the autumn season began
in 1954 with an extension through Novem¬
ber 1 5 and a further extension to December
31 in 1974. In 1983, a year-round open sea¬
son was created from U.S. Highway 2 to Lake
Superior to spread out fishing pressure and
create additional angling opportunities. This
year-round season was rescinded after the
1985 season because of public dissatisfaction
and indications of excessive harvest. Present
season extensions run from the Saturday
nearest April 1 through November 15.
Volume 82 (1994)
63
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Daily Bag and Minimum Size
Restrictions Prior to 1989
There was no statewide restriction on the
daily bag for trout until 1905 when it was
set at not more than 10 pounds. In 1909,
the daily bag was changed to 45 trout; it was
subsequently reduced to 35 trout in 1917,
25 trout in 1923, 15 trout in 1929, and 10
trout in 1949. From 1949 to 1989 the daily
bag during the regular season remained at
ten trout or salmon, sometimes with the
stipulation that only five of those could be
steelhead, or steelhead and brown trout in
aggregate. Daily bag limits for the Brule
River during the special extended seasons
were more restrictive than during the regu¬
lar seasons. From 1962 to 1989, the daily
bag during the extended seasons was five
trout or salmon in aggregate, with the stipu¬
lation for 1 6 of those years that only two of
the five could be steelhead.
The first size limit enacted for trout in
Wisconsin was a 6-inch minimum in 1905.
In 1915 the statewide minimum length was
increased to 7 inches; it was set back to 6
inches again in 1950, where it remained un¬
til 1989. The special extended seasons have
been subject to higher minimum lengths,
beginning with a 13-inch minimum for the
late season in 1954. In 1970, the minimum
length limit during the early and late seasons
was reduced to 10 inches, where it remained
until 1989.
Recent Changes in Daily Bag and
Minimum Size Limits
Although anglers were increasingly practic¬
ing voluntary catch-and-release during the
1980s (Table 3), by 1989 it became appar¬
ent that additional restrictions on harvest of
both adult spawners and presmolts of steel¬
head and brown trout were necessary to pro¬
tect the fishery. Excessive harvest was
strongly implicated in the declining numbers
of spawning steelhead. Concern had also
mounted that harvest of large, resident
brown trout may have been dangerously
high, especially during a popular, early-sum-
mer mayfly hatch ( Hexagenia limbata) when
trout are particularly vulnerable. An addi¬
tional concern surfaced that harvest of
presmolt steelhead and brown trout 6 to 10
inches in length may have been substantially
reducing run sizes of adult spawners. Stud¬
ies had shown smolt size to be positively cor¬
related with survival to the maiden spawner
stage (e.g. Ward and Slaney 1988), and that
if steelhead survived their presmolt winter,
they had an excellent chance to grow to tro¬
phy size (Seelbach 1987). Hence, larger
presmolts were valuable and required pro¬
tection from harvest.
New regulations in 1989 for the entire
open season therefore included a reduced
daily bag limit of five salmonids in total of
which only one could be a steelhead 12
inches or larger and only two could be
brown trout larger than 15 inches. Mini¬
mum sizes were increased to 10 inches on
brown trout and 12 inches on steelhead and
Pacific salmon to protect anadromous
presmolts. The minimum size limit on
brook trout was also increased to 8 inches
to allow more fish to grow into a desirable
size range. In 1993, the minimum size limit
on steelhead was further increased to 26
inches for the Brule River and all other Wis¬
consin tributaries to Lake Superior to ensure
that young adults would have the opportu¬
nity to spawn at least once before entering
into the harvest. More time is needed to de¬
termine the extent to which these changes
may have benefited the fishery.
Concurrent with the decline in the Brule
River steelhead run in the mid and late
1980s were indications of reduced steelhead
64
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
runs in other Wisconsin tributaries to Lake
Superior (B. Swanson, WDNR, pers.
comm.). Increasing harvest of steelhead in
Lake Superior by anglers in charter and pri¬
vate boats was suspected of contributing sub¬
stantially to this disturbing decline of steel-
head in western Lake Superior. Hence, in
1990, regulations for the Wisconsin waters
of Lake Superior were changed to allow daily
harvest of only one steelhead over 28 inches
in length.
Other Restrictions on Fishing
Gear restriction proposals for reducing har¬
vest and post-release mortality of salmonids
in sections of the upper river have periodi¬
cally been voiced. However, the only gear
restriction ever enacted was for “fly fishing
only” for the stretch of river from Stone’s
Bridge to Winneboujou in 1938 in response
to a petition by landowners. This restriction
was rescinded shortly thereafter, following a
storm of public protest that may have been
politically generated (Holbrook 1949).
Other restrictions include no fishing from
1/2 hour after sunset to 1/2 hour before sun¬
rise during the extended seasons on the lower
river, and the establishment of small refuge
areas closed to fishing where migrating fish
tended to congregate and illegal snagging was
known to have occurred (below the sea lam¬
prey barrier, the Boxcar Hole and the Skid
Mays area within the Ledges section5) .
Voluntary catch-and-release is being
practiced with increasing frequency, espe¬
cially on the upper river, as the angling
community re-evaluates the value it places on
wild salmonids. Brule River Preservation,
Inc., the Brule River Sportsmen’s Club, Inc.,
and Trout Unlimited of Wisconsin have
undertaken a collaborative signage project on
the upper river that suggests the practice of
voluntary catch-and-release to preserve good
fishing. These types of voluntary initiatives
have been shown to effectively shape angler
behavior because many anglers are influenced
by the ethics of their peers. Also, anglers are
concerned about the future of their sport and
typically respond well to education.
Management Implications
1. Continue the focus on riparian pro¬
tection. A strong posture by the WDNR
and private interests on protecting riparian
areas has existed for decades and should con¬
tinue. A healthy riparian zone is instrumen¬
tal for maintaining water quality and
instream habitat diversity, which are in turn
critical for the continuing support of a di¬
verse array of coldwater life. The state should
continue its policy of land acquisition within
State Forest boundaries from willing sellers
where feasible. An enduring focus on coop¬
erative ecosystem stewardship by riparian
landowners, the Brule River Preservation,
Inc., the Brule River Sportsmen’s Club, Inc.,
and a broad range of WDNR functions will
remain invaluable.
2. Focus for instream habitat mainte¬
nance and enhancement. Habitat mainte¬
nance and enhancement considerations for
the river system should include beaver con¬
trol and dam removal to maintain an unim¬
peded migration route, dredging of spring
pond areas to increase brook trout spawn¬
ing potential (however, dredging of the
mainstem is not recommended), and addi¬
tion of downed conifers, rootwads, and
other forms of large woody debris through¬
out the system to compensate for wood re¬
moved since the late 1800s. Existing
instream habitat improvement structures
that seem to have been useful should be re¬
furbished. Public acquisition of any part of
the Cedar Island spring pond area to be
again made accessible for wild brook trout
Volume 82 (1994)
65
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
spawning should be a high priority if that
property becomes available; a substantial
boost to the brook trout population would
likely result given the apparent paucity of
spawning areas for that species elsewhere
throughout the upper river.
3. Continue the trend for increasingly
restrictive angling regulations. Increasingly
restrictive regulations have contributed to a
brighter future for the salmonid fisheries,
especially since over-exploitation was iden¬
tified as one of their major threats. This
trend should be maintained, and gear restric¬
tions for sections of the upper river should
be considered pending determination of the
effects of the 1989 regulatory changes on the
salmonid populations.
4. Further monitoring of the stream
biota. The recent establishment of numer¬
ous exotic species serves to remind us that
the Brule River is part of a larger ecosystem
that will remain continually at risk from dis¬
tant occurrences. Given the climate of un¬
certainty under which the system must be
managed, there will be a need for periodic
monitoring of the fish and aquatic insect
communities to test for impacts of exotic
animals on native species. Additionally, an
abbreviated sampling schedule of the quan¬
titative investigations into salmonid popu¬
lation dynamics initiated during the last de¬
cade should continue, as much as eco¬
nomically feasible, to ensure benefits to
future fisheries management.
3. Continue the research focus. Many in¬
dications from the public including the ad¬
mirable work of the Brule River Commit¬
tee and the committee’s strong support of
the research efforts of the last decade sug¬
gest that status quo management of this river
system is not acceptable. The public has a
right to expect state-of-the-art management
on a resource as valuable as the Brule River.
Resource management policies have come
under increased scrutiny from special inter¬
est groups and this trend will likely continue.
Continuing research will be needed to sat¬
isfy the demand for sound management.
Acknowledgments
We extend special thanks to our technicians
W. Blust, S. Plaster, and F. Stoll, without
whose able assistance in the field this re¬
search would not have been successful, and
to the many limited-term employees who
contributed to the research in a variety of
ways. We also thank R. Hunt and S. Schram
for their roles in fostering this research. Re¬
view comments from M. Holbrook, R.
Hunt, J. Lyons, M. Jennings, P. Seelbach,
M. Staggs, W. Weiher, C. Zosel, and an
anonymous referee improved the manu¬
script. M. Jesko and C. Olson assisted with
the figures. L. Claggett, R. Lee, and person¬
nel from the State Historical Society of Wis¬
consin helped unearth information about
early regulation of the fishery. This research
was supported in part with funds authorized
by the Anadromous Fish Conservation Act
(project WI-AFS-16), the Federal Aid in
Sport Fish Restoration Act (projects F-83-
R and F-95-P, study 413), and the Wiscon¬
sin Department of Natural Resources.
Endnotes
1 Coaster brook trout apparently exhibited an
anadromous life history strategy similar to
that of the anadromous brown trout. Little
scientific information on coaster brook trout
in Great Lakes tributaries is available because
most populations were extirpated before sci¬
entific data were collected. Bullen (1988) de¬
scribes a remnant population in a Lake Su¬
perior tributary.
2 Upper river refers to the river reach from U.S.
Highway 2 south (upstream) to the con-
66
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
fluence of the East and West forks. Lower
river refers to the river reach from U. S. High¬
way 2 north (downstream) to Lake Superior.
Upper or above always refers to an upstream
direction; lower or below always refers to a
downstream direction. Mainstem refers to the
main thread of the Brule River proper with¬
out the tributaries.
3 The lakes section is composed of four wide-
spreads of the Brule River (Sucker, Big,
Lucius, and Spring lakes) located just south
of the river’s midsection (Fig. 1).
4 There is likely much less large woody debris
in the Brule River (and other rivers and
streams in northern Wisconsin) than there
was historically for several reasons in addition
to the removal efforts of early fisheries work¬
ers. Woody debris has been systematically re¬
moved from our rivers and streams for more
than a century to maintain open channels for
human navigation, and during the intensive
logging era of the late 1800s removal efforts
were intensified to maintain smooth channels
for the downriver transport of logs. Addition¬
ally, the clear-cutting of our northern forests
at that time (which included steamside areas)
temporarily interrupted the continual natu¬
ral process of dying streamside trees falling
into waterways.
5 The ledges section is the reach of river having
maximum gradient where it crosses the
Copper Range (Fig. 1). The river descends 80
ft in 274 miles at that point (Bean and
Thompson 1944).
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Jerrard, L. P. 1956. The Brule River of Wiscon¬
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Jones, C. M. and D. S. Robson. 1991. Improv¬
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70
TRANSACTIONS
DUBOIS and PRATT: History of the fishes of the Bois Brule River System
Aquat. Sci. 45:1 1 10-22.
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transition from hatcheries to habitat. Trout ,
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Madison: Wisconsin Historical Society.
Robert B. DuBois is a research scientist with the
Rivers and Streams Research Group of the Wiscon¬
sin Department of Natural Resources at Brule.
From 1983-94 he was project leader for a research
study to obtain quantitative population statistics
about the salmonids of the Bois Brule River.
Address: Wisconsin Department of Natural Re¬
sources , 6250 S. Ranger Rd, P. O. Box 125 , Brule,
WI 54820
Dennis M. Pratt is Western Lake Superior Fish¬
eries Manager with the Wisconsin Department of
Natural Resources in Superior. He has functioned
as fisheries manager for the Bois Brule River eco¬
system since 1985. Address: Wisconsin Department
of Natural Resources, 1705 Tower Avenue, Supe¬
rior, WI 54880
Volume 82 (1994)
71
Thomas F. Grittinger and Deborah L. Schultz
Social behavior of adult jaguars
(Panthera onca L.) at the
Milwaukee County Zoo
Abstract The purpose of this study was to describe and analyze social behav¬
ior between two captive jaguars (Panthera onca Li), a male and a
female, at the Milwaukee County Zoo. Some one hundred eighty-
nine bouts were recorded and analyzed over four years. These bouts
consisted of numerous acts, some of which appeared to be sex-specific
or at least individual- specific. The bouts varied considerably in du¬
ration; the mean duration of the bouts initiated by the male was
significantly longer than that for bouts initiated by the female. Two-
act sequences for each animal revealed a large number of complex
grooming and wrestling acts, with much switching back and forth.
Apparent sex differences were revealed in the two -act sequences of
complex grooming and clasping, clasping and lying on, and wres¬
tling and lying on. As the study progressed, the male spent a signifi¬
cantly greater percentage of bout time in sexual behavior during the
bouts.
The jaguar ( Panthera onca L.) is one of the least studied
large cats in the world (Rabinowitz and Nottingham
1986). Until recently, the only published information avail¬
able on the jaguar came from anecdotal reports of explorer-
naturalists, hunters, and surveyors (Crawshaw and Quigley
1991). The recent scientific papers focus on various aspects of
jaguar biology, such as home ranges, movements, and daily ac¬
tivity patterns (Crawshaw and Quigley 1991; Rabinowitz and
Nottingham 1986; Schaller and Crawshaw 1980). Other pub¬
lished works emphasize various aspects of their predatory be¬
havior (Emmons 1986; Mondolfi and Hoogesteijn 1986;
Rabinowitz 1986b; Schaller and Vasconcelos 1978). Despite
this, little is known regarding jaguar social interaction patterns
(Mondolfi and Hoogesteijn 1986). The purpose of this paper
TRANSACTIONS Volume 82 (1994)
73
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
is to provide an ethogram (“a set of com¬
prehensive descriptions of the characteristic
behavior patterns of species” [Brown 1975]),
in this case limited to social behaviors. This
paper describes social interactions between
two captive jaguars (a male and a female) at
the Milwaukee County Zoo over a four-year
period. In addition to the description of in¬
dividual acts, the duration of interactions
was examined, the sequences of acts was ana¬
lyzed, and variation over the study period
was monitored.
Methods
The subjects of this study were both captive
born (male: February 10, 1983; female: Sep¬
tember 22, 1982) and had been housed to¬
gether since September 15, 1983. The pe¬
riod of observations extended from October
22, 1985, to July 17, 1990. During this time
the male was sexually unaltered while the
female was maintained on Melangesterol (a
progesterone derivative) implants until July
1990, when an ovariohysterectomy (spay)
was performed. The study was terminated
at that time.
Observations were made two to three
times per week during the snow-free months
of the year in Milwaukee (from May to early
December) when the animals were released
into the outside enclosure, an area 40 x 60
ft (12.2 x 18.3 m) (Fig. 1). Observations
lasted from 30 to 60 min, depending on the
level of activity. Social interactions were re¬
corded by means of a video camera with a
built-in timer and were analyzed later. The
interactions consisted of bouts, “relatively
stereotyped sequence [s] of behaviors that oc¬
cur in a burst” (Lehner 1979). A bout ex¬
tended from first contact (an obvious act of
initiation of social interaction such as a fixed
gaze) to termination (when one or both ani¬
mals withdrew and did not reunite within
30 s). The initiator of the bout was the in¬
dividual that approached the other (receiver)
or initiated first contact.
The bouts were recorded, and the se¬
quences of motor acts or patterns within the
bouts were obtained by replaying the video
tapes on slow speed. Accurate analysis of rap¬
idly changing events was facilitated by the
fact that the male is black and the female
spotted. Two-act sequences (Latour 1981)
were presented to contrast the behavior of
the male to that of the female, both as ini¬
tiator and as recipient. Changes in relative
frequencies of some of the more common
acts were examined for the study period.
Results and Discussion
An ethogram is the starting point in any
ethological research (Lehner 1979). Al¬
though the behaviors of various cats have
been documented (e.g. Schaller’s [1972]
work on the lion [P. leo L.] Wasser’s [1976]
thesis on tiger [ P . tigris L.] play, and
Leyhausen’s [1956] and West’s [1974] pa¬
pers on the domestic cat [Felts catus L.]),
there are no reports describing jaguar social
behavior.
In order to construct the ethogram, sev¬
eral hundred contact bouts were recorded,
one hundred eighty-nine of which were
unobstructed enough for analysis. The
bouts were described by the acts they con¬
tained.
Non-Contact Acts and No Activity
Non-Contact Acts:
• Fixed gaze (FG) - The animal initiated the
bout with its head and neck held low and
oriented toward the recipient animal. The
animal rapidly shifted weight from one front
leg to another.
74
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GRITTI NGER and SCHULTZ: Social behavior of adult jaguars
Fig. 1 . The female jaguar (right) and the male jaguar in their outdoor enclosure
• Stalking (St) - The initiator moved slowly
with a fixed gaze toward the recipient. The
body was tense and held in a lowered man¬
ner; the forequarters were usually held lower
than the hindquarters.
• Rushing (Rus) - One animal ran toward
the other, who did not flee.
• Chasing (Ch) — One animal ran toward
the other, who did flee.
• Approaching (Ap) - One animal simply
walked toward the other.
• Facing off (FO) - The initiator’s head was
oriented toward the other animal, who
would or would not do likewise. The ani¬
mals were in close proximity to each other,
usually about a body length or less apart.
• Walk/run away (WA/RA) - The animal
walked or ran away, usually ending the bout.
No Activity (NA):
This category was probably more artificial
than the rest. It ranged from the relaxed
mode of behavior seen when one animal re¬
mained motionless while being licked by the
other, to a lack of discernible activity on the
part of both animals. This category involves
no reciprocity of actions whatsoever.
Contact Acts
Playing Behavior:
• Rubbing (Rub) - Rubbing often was the
first actual physical contact between the two
animals. One animal would rub the other
using its head or body.
• Holding (Ho) - Holding was accom¬
plished by throwing either one front leg over
Volume 82 (1994)
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the neck or shoulder of the subject, or by
placing both front legs around the other’s
neck. In some cases it appeared to restrain,
while in others it was less forceful.
• Grabbing (Gr) — Grabbing was done by
the rapid thrust of one of the front paws out
and around the leg of a fleeing animal.
• Jumping on Back (JOB) - The animal
placed its forequarters on the other’s back.
The front legs were usually straddled over
the subject, and the hind legs may or may
not have been on the ground.
• Pawing (Pa) - The subject was struck with
the forepaw. The intensity of pawing varied
greatly.
• Biting (Bi) - Biting covered a range in ac¬
tivity from a brief mouthing to a much more
sustained biting and pulling.
• Wrestling (W) — Wrestling ranged from a
close embrace with combinations of pawing,
biting, and kicking with the hind legs, to a
loose relationship with one animal lying on
its back and the other animal standing over
or near it. The animal lying on its back
would often paw or reach with its front feet
while treading with its hind legs; the mouth
was held open and the teeth were exposed.
The standing animal’s mouth was also open
while it bit and pawed at the lying animal.
The belly-up and stand-up positions were
sometimes interchanged and often mixed
with close contact wrestling. The intensity
of wrestling varied considerably.
Grooming behavior:
• Simple licking (SL) - In this situation one
animal simply licked the other, from either
a standing or lying position. The recipient
of the simple licking did nothing (listed as
NA). Both animals did this.
• Complex Grooming (CG) - In this com¬
plicated act, the female straddled the male
who was belly side up, and she would lick
him. This behavior was more protracted
than simple grooming, and the animals were
always in the straddle-belly up positions.
This was maintained as long as she contin¬
ued to lick him. This belly-up position dif¬
fers from the NA behavior insofar as he ac¬
tively participated in the sequence of events.
When she stopped, he would start wrestling
and either she would resume grooming and
he would again relax, or she would wrestle
with him.
Sexual behavior:
• Clasping (Cl) - The male hugged the fe¬
male closely while both were lying on their
sides. The male would pull the female against
his chest and pull her down toward his stom¬
ach while treading with the hind feet.
• Lying on (LO) - With her head at or be¬
low the level of his stomach, the male rested
his body on the female. While doing this,
he would often move back and forth over
the recumbent female, sometimes displaying
a partial erection.
• Mounting (Mo) - The male straddled the
female who was sitting or lying on her belly.
The results of this study demonstrate that
within the conditions of captivity jaguars
socially interact in bouts. The data support
the idea that playing, grooming, and sexual
behaviors often are found in the same bout.
Bout duration varied considerably, with a
range of 1 to 1204 s (Fig. 2). The majority
(57%) of the bouts were less than 1 min
long. These short bouts often consisted of
one of the approach acts, followed by one
of the contact acts such as biting or pawing,
and usually ended with a walk-away or run-
76
TRANSACTIONS
GRITTINGER and SCHULTZ: Social behavior of adult jaguars
Number of Bouts
Bout Length (in 60 second intervals)
Fig. 2. Distribution of bouts by length
away. The longer bouts began and ended
much as the shorter ones did, but contained
a reciprocal switching of contact acts. The
male initiated 106 bouts and the female ini¬
tiated 83 bouts, with no significant differ¬
ence between initiators (test concerning pro¬
portions, z = ±1.67, P > .03). The mean
duration of the bouts initiated by the male
(174 s) was significantly longer than that for
the bouts initiated by the female (99 s) (t-
test, t = 2.31, P < 0.01).
Bouts were each analyzed as being com¬
posed of a series of two-act sequences of suc¬
cessive acts by the same animal. The inte¬
gration of individual acts and the differences
between the two animals is seen in these
two-act sequences (Figs. 3-6). The area of
each box in Figures 3—6 is proportional to
the total number of acts, and the arrow
width is proportional to the total number of
transitions between the acts. For the sake of
simplicity, only those transitions of ten or
more were included.
Complex grooming and wrestling were
the most common acts seen in both animals,
both as initiator and as receiver. As the ar¬
rows suggest, these two acts often switch
back and forth during a bout. It is of inter¬
est that prolonged wrestling, a common play
act seen here, did not occur in wild adult
lions (Schaller 1972).
While most of the acts were performed
by both animals, there were a few that ap¬
peared to be individual-specific or sex-spe¬
cific. Complex grooming varied between the
animals, with the female always straddling
the belly-up male. The male’s position was
maintained as long as she continued to lick
Volume 82 (1994)
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Fig. 3. Two-act sequences for male as
initiator
NA
Fig. 4. Two-act sequences for male as
receiver
Wrestle
NA WA/RA
Fig. 5. Two-act sequences for female as
initiator
him. If she stopped, he would start wres¬
tling, and either she would resume the com¬
plex grooming and he would again relax or
she would wrestle with him. A similar reac¬
tion was seen in the bite-lick-bite sequence
in tigers (Wasser 1978) and in the use of
social grooming as an interruption of
playfight (Fagen 1981).
Only the male engaged in clasping, lying
on, or mounting (all sexual) behaviors. As
the study progressed, a shift was apparent
Wrestle
Hold WA/RA
Fig. 6. Two-act sequences for female as
receiver
(Fig. 7). There was a significant change in
the proportion of time engaged in sexual
behavior during the four years (the differ¬
ence between proportions test was, for
1987-88, z = -6.72; for 1988-89, z = -13.15;
and for 1989-90, z = -6.82, P < 0.001). His
two-act sequences often included wrestle-lie
on, lie on-wrestle, wrestle-clasp, and clasp-
lie on; clasp appeared to be transitional be¬
tween wrestling and lying on. When the
male would either clasp or lie on the female,
78
TRANSACTIONS
GRITTINGER and SCHULTZ: Social behavior of adult jaguars
she would lie still (NA) or wrestle. Mount¬
ing was too infrequent to warrant placing
in the two-act sequences; the female usu¬
ally walked away from him as he attempted
to do this. While Wasser’s (1978) tiger
bouts included contact behaviors such as
paw, bite, wrestle, and lick, they lacked any
sexual activity, possibly because his tigers in¬
cluded only one adult male and five of its
offspring (one sub-adult and four cubs).
The progressive increase through the four
years in the percentage of sexual behavior
cannot be attributed to the development of
sexual maturity. The animals were adults
when the study started. He was 4.5 years and
she was 5 years old, and they had produced
a cub before the observations started.
Rabinowitz (1986a) considered jaguars to be
subadults at 2-3 years and mature adults at
4-10 years. Mondolfi and Hoogesteijn
(1986) gave 2—2.5 years for sexual maturity
in the female jaguar and 3-4 years for the
male jaguar.
Unlike the complex grooming and the
sexual behaviors, the acts that constitute play
appeared not to show individual or sex dif¬
ferences. This is in agreement with Fagen’s
(1981) prediction that in carnivores, where
both sexes display similar fighting skills, no
differences should exist in play. In both ani¬
mals the play demonstrated some accepted
characteristics: exaggerated and repeated acts
(Loizos 1966; Fagen 1981). Among the
many functions attributed to play, expend¬
ing excess energy (Bekoff 1976; Schaller
1972) and strengthening social bonds
(Schaller 1972) were two that may have
merit here. In confinement, male and female
Volume 82 (1994)
79
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
felids often live together in relative peace and
may frequently play with each other, phe¬
nomena seldom recorded in the wild (Fagen
1981). In the wild, all felids, with the ex¬
ception of the lion and the cheetah (. Aci -
nonyx jubatus Schreber), live solitary lives
(Bekoff 1989). Hemmer (1978) suggested
that carnivores are capable of a much greater
plasticity in social behavior in zoos than they
usually are in nature. He proposed that there
are three factors that affect sociality in the
pantherines: (1) environment, (2) relative
brain size, and (3) temperament. He be¬
lieved that the jaguar and the leopard (P.
pardus L.) are capable of group living but,
like the tiger, are forced by habitat condi¬
tions in the wild to forgo this.
Detailed longitudinal field observations of
identified individual felids are still lacking
(Bekoff 1989), and the evidence presented
here is limited to only two captive jaguars;
nevertheless, this study, which covers four
years in the lives of two readily identified
individuals, supports Ewer’s (1973) conten¬
tion that felids are not as asocial as com¬
monly believed. The observations indicate
that social behavior is organized into bouts
of varying length and complexity and that
the contact portion of the bout is dominated
by play and complex grooming activity.
Acknowledgments
Support for this study was provided by a
University of Wisconsin Centers sabbatical
leave. We would like to thank Dr. Millicent
Ficken for her ideas, Dr. Jack Hailman for
suggestions on the manuscript and the treat¬
ment of data, and the Milwaukee County
Zoo, especially Ralph Konrath and Valerie
Werner, for their assistance throughout the
project.
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1(1): 1 02 — 05
Latour, P. B. 1981. Interactions between free-
ranging, adult male polar bears ( Ursus
maritimus Phipps): a case of adult social play.
Can.]. Zoo. 59:1775-83.
Lehner, P. N. 1979. Handbook of ethological
methods. New York: Garland STPM Press,
403 pp.
Leyhausen, P. 1956. Verhaltensstudien an
Katzen. Z. Tierpsychol. Beiheft 2:1-120.
Loizos, C. 1966. Play in animals. Symposium of
the Zoological Society of London 18:1-9.
Mondolfi, E. and R. Hoogesteijn. 1986. Notes
on the biology and status of the jaguar in Ven¬
ezuela. In Cats of the world: biology, conserva¬
tion, and management , eds. S. D. Miller and
80
TRANSACTIONS
GRITTINGER and SCHULTZ: Social behavior of adult jaguars
D. D. Everett, pp. 85-123. Washington,
D.C.: National Wildlife Federation.
Perry, R. 1970. The world of the jaguar. New
York: Taplinger Publishing Co, 168 pp.
Rabinowitz, A. R. 1986a .Jaguar. New York: Ar¬
bor House, 368 pp.
Rabinowitz, A. R. 1986b. Jaguar predation on
domestic livestock in Belize. Wildl. Soc. Bull.
14:170-74.
Rabinowitz, A. R. and Nottingham, B. G., Jr.
1986. Ecology and behavior of the Jaguar
( Panthera onca) in Belize, Central America. J.
Zool. Lond. (A) 210:149-59.
Schaller, G. B. 1972. The Serengeti lion. Chicago:
Univ. of Chicago Press. 480 pp.
Schaller, G. B. and P. G. Crawshaw, Jr. 1980.
Movement patterns of jaguar. Biotropica
12:161-68.
Schaller, G. B. and J. M. C. Vasconcelos. 1978.
Jaguar predation on capybara. Z. Sauge-
tierkunde 43:296-30 1 .
Wasser, S. K. 1976. Play in felidae: a relation of
structure and function. M.S. Thesis, Univer¬
sity of Wisconsin— Milwaukee.
- . 1978. Structure and function of play
in the tiger. Carnivore I (3):27-40.
West, M. 1974. Social play in the domestic cat.
Amer. Zool. 14:427-3 6.
Thomas F. Grittinger is a professor of biology at
the University of Wisconsin Center— Sheboygan
County. Address : Dept. Biological Sciences , UW
Center— Sheboygan County , Sheboygan, WI 53081 -
4789
Deborah L. Schultz, a former student at the Uni¬
versity of Wisconsin Center-Sheboygan County, is
a graduate of the Medical College of Wisconsin and
is completing her residency program.
Volume 82 (1994)
81
Richard E. Koske and Leonard L. Tews
Vesicular- arbuscular mycorrhizal fungi
of Wisconsin s sandy soils
Abstract The root zones of beach grass, Ammophila breviligulata, of Wisconsin s
Great Lake’s dunes, and of other plants of sandy soils of the state were
surveyed for the first time for vesicular-arbuscular mycorrhizal (VAM)
fungi. The most frequently obtained were Glomus etunicatum,
Gigaspora rosea, Glomus geosporum, and Glomus macrocarpum. Taxo¬
nomic characteristics of the obtained fungi are similar, in most cases,
to those of VAM fungi found elsewhere.
The soils of Wisconsin heretofore have been unexamined
for vesicular-arbuscular mycorrhizal (VAM) fungi. More¬
over, there have been only a few reports of VAM fungi being
isolated from the lacustrine dunes of the Great Lakes region
of North America (e.g., Koske et al. 1975; Koske 1985). Be
that as it may, there have been numerous reports of VAM fungi
from North America’s maritime dune systems (e.g., for New
England, Koske and Halverson 1981; Koske 1981; and for the
Northwest, Gerdemann and Trappe 1974). It is of ecologic
and taxonomic interest to know the VAM fungi of Wisconsin’s
soils, since they have potential in the improvement of crop pro¬
duction. Moreover, it is worthwhile to compare the VAM flora
of the Great Lakes dunes to those of New England’s dunes,
especially since the higher plant flora, and therefore the ecol¬
ogy itself, is so similar.
Procedures
VAM fungi were collected from the root zones of plants at
53 sites at 21 different geographic locations (described with
their sampling dates in Table 1). The sites included dunes of
TRANSACTIONS Volume 82 (1994)
83
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1. Collected species found in association with VAM fungi
Site
Species Association
Date Collected
1 . Cornucopia
(Bayfield Co.)
2. Rock Island
(Door Co.)
3. Washington Island
(Door Co.)
4. Newport State Park
(Door Co.)
5. Bailey’s Harbor
(Door Co.)
6. Jacksonport
(Door Co.)
7. Whitefish Dunes
State Park (Door Co.)
8. Kewaunee
(Kewaunee Co.)
9. Two Rivers
(Manitowoc Co.)
10. Kohler-Andrae
(Sheboygan Co.)
1 1 . Kohler-Andrae
(Sheboygan Co.)
12. Harrington Beach
(Ozaukee Co.)
13. Sauk City
(Columbia Co.)
14. Lake Wisconsin
(Columbia Co.)
15. Plainfield
(Waushara Co.)
16. New Hope
(Waupaca Co.)
Ammophila breviligulata Fern, growing on July 3, 1984
Lake Superior dunes at Cornucopia’s public beach
Agropyron dasystachum (Hook) Scribn. growing July 22, 1984
on Lake Michigan dunes
A. breviligulata growing on Lake Michigan dunes July 22, 1984
Calamovilfa longifolia (Hook) Scribn. growing July 22, 1984
on Lake Michigan dunes
A. breviligulata from Lake Michigan dunes July 21, 1984
at The Ridges beach (remnant arboreal forest)
A. breviligulata growing on Lake Michigan dunes July 21, 1984
at Jacksonport’s public beach
A. breviligulata growing on Lake Michigan dunes July 21, 1984
A. breviligulata growing on dunes at Kewaunee June 16, 1985
Pioneer Park
In the root zone of A. breviligulata growing on June 16, 1986
dunes at Two River’s public beach
A. breviligulata growing on Lake Michigan July 21, 1984
dunes at Kohler-Andrae State Park
A. breviligulata, Lathyrus maritimus (L.) Bigelow, October 9, 1983
C. longifolia, and Rosa sp. growing on Lake
Michigan dunes at Kohler-Andrae State Park
L. maritimus and A. breviligulata growing on Lake July 11, 1984
Michigan dunes at Harrington Beach State Park
In the root zone of A. breviligulata growing on June 21 , 1985
a bank of the Wisconsin River in Sauk City
Sedges and Lythrum salicaria L. growing on June 21 , 1985
a sandbar in Lake Wisconsin
Associated with the roots of Z mays L. near June 27, 1985
an irrigated field near Plainfield
From an unirrigated oat field near New Hope June 27, 1985
84
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KOSKE and TEWS: Vesicular-arbuscular mycorrhizal fungi of Wisconsin
Lake Michigan and Lake Superior, sandbars
and banks of the Wisconsin River, and ir¬
rigated and unirrigated sandy soils of cen¬
tral Wisconsin. Sand-root samples were
placed in plastic bags and stored at about
6°C until processed. Field notes regarding
position, date, and vegetational cover were
recorded.
Spores of the VAM fungi were separated
from the soil by a technique of flotation and
filtration (Koske and Halvorson 1981). Ap¬
proximately 35 g of sand were added to
about 500 ml of tap water in a 1 liter bea¬
ker and agitated vigorously. The suspension
was filtered immediately, before settling,
through a .5 mm soil sieve with the spores
being collected on a No. 4 Whatman filter
paper placed in a Buchner funnel with
vacuum pressure. The filter paper was re¬
moved from the funnel and placed under a
dissecting microscope where spores of the
fungi were isolated to a drop of water on a
glass slide with the aid of a fine forceps. A
cover slip was placed atop the spores, and
with gentle pressure, the spores were burst
for better examination of spore wall struc¬
ture. Spores were sometimes stained with
Melzer’s Reagent for diagnostic purposes. If
a permanent slide was desired, spores were
placed in a drop of polyvinyl alcohol solu¬
tion and sealed with clear fingernail polish.
Results
Fifteen species of VAM fungi were collected
from the sandy soils of Wisconsin (Table
2). All species are new records for Wiscon¬
sin. The fungus most frequently isolated
from sand dunes was Glomus etunicatum.
Other commonly obtained fungi are
Gigaspora rosea , Gl. geosporum and Gl.
macrocarpum.
The VAM fungi of Wisconsin’s dunes
were compared to those of other soils using
a coefficient of similarity, C (Bray and Curtis
1957) (Table 3). The equation is: C = 2w/a
+ b where w = the number of species
common to both floras, a = the number of
species of one flora, and b = the number of
species of the other flora. The VAM fungi
of Wisconsin’s dunes are most similar to
those of Wisconsin’s non-dune soils, and
somewhat more similar to those of Iowa than
they are to those of the maritime dunes of
Rhode Island or the prairie soils of Illinois.
One of the obvious differences between
the VAM floras of Iowa, Wisconsin, and
Rhode Island is that Gigaspora-Scutellospora
Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 2. VAM fungi collected from Wisconsin’s Great Lake dunes and other sandy
soils
Species
Sites from which species were isolated *
Acaulospora scrobiculata Trappe
A. spinosa Walker & Trappe
Gigaspora rosea Schenck & Smith
Gi. gigantea (Nicol. & Gerd.) Gerd. & Trappe
Glomus aggregatum Schenck & Smith
GI. caledonicum (Nicol. & Gerd.) Gerd. & Trappe
GI. etunicatum Becker & Gerd.
GI. geosporum (Nicol. & Gerd.) Walker
GI. macrocarpum Tul. & Tul.
GI. microaggregatum Koske, Gemma & Olexia
GI. mosseae (Nicol. & Gerd.)
GI. lamellosum Dalpe, Koske & Tews
Glomus sp. B.
Scutellospora calospora (Nicol. & Gerd.) Walker & Sanders
S. dipapillosa (Walker & Koske) Walker & Sanders.
9,13
11,14,16
9.10.11
9,13,14
4.11
5
3, 4, 5, 6, 7, 9,11,12,14,17,19
9,11,18, 20, 21
11,18
1,8
15, 20, 21
5.10.11
10.11
1,11
15
*Numbers indicate the site from Table 1 from which the fungi were collected.
complex seems to be a more important
component of Rhode Island’s flora. (These
genera are combined on Table 4 since the
separation of Scutellospora from Gigaspora
occurred after most of these studies were
reported). In the Rhode Island dunes, 44%
of the collected species are of the genera
Gigaspora-Scutellospora whereas in the dunes
of Iowa and Wisconsin this figure is much
lower (Table 3). The relative importance of
Acaulospora also differs. In the Rhode island
dunes, only one species, A. scrobiculatum,
was obtained. In the midwestern soils of
Iowa and Wisconsin, 20 to 23% of the
species belonged to the genus Acaulospora.
In addition, the genus Glomus appears to be
more important in the midwestern soils than
it is in Rhode Island dunes.
Discussion
For the most part, VAM fungi collected from
Wisconsin resemble those collected else¬
where. Nevertheless, there are some excep¬
tions. The azygospores of Acaulospora spinosa
are small, but fall within the range of Walker
and Trappe’s (1981) description. The spores
and suspensors of Scutellospora calospora are
smaller than those reported by Gerdemann
and Trappe (1974), but probably this is not
enough of a difference to warrant a new spe¬
cies. The walls of Gi aggregatum did not ex¬
hibit the greenish tint previously reported as
sometimes present (Koske 1983). The
chlamydospores of GL caldeonicum fall be¬
low the range of those of Gerdemann and
Trappe (1974). The walls are thicker than
86
TRANSACTIONS
KOSKE and TEWS: Vesicular-arbuscular mycorrhizal fungi of Wisconsin
Table 3. The VAM fungi of Wisconsin’s Great Lakes Dunes compared to those of
other soils using a coefficient of similarity, C
C
Wisconsin non-sandy soils .84
Iowa soils (Walker et al. 1982) .44
Rhode Island maritime dunes (Tews and Koske 1986) .30
Illinois prairie soil (Anderson and Liberta 1989) .25
Coefficient of similarity, C is calculated by using the fomula C = 2W/a + b, where w is the number
of species common to both populations, a is the number of species in one population, and b
is the the number of species in the other population.
those reported earlier, and they are some¬
times tinged with a pale pink, which may be
the result of a bacterial invasion carrying the
pigment. Again, these differences are not im¬
portant enough to warrant naming a new
species.
The chlamydospores “Gl. tortuosum” are
about twice the size of these of Schenck and
Smith (1982). The spore wall is about three
times thicker. They reported a single lami¬
nate wall; four were observed here. The
width of the attachment is three times that
reported by Schenck and Smith.
The higher plant flora of the lacustrine
dunes of the Great Lakes are similar to those
of the maritime dunes of New England, but
their VAM fungal flora vary greatly. In a
study of Moonstone Beach (Tews and Koske
1986), a barrier dune in Rhode Island, only
two of the nine species isolated, S. calospora
and Gl. aggregatum , were similar to those of
the present study. Three species in the
Moonstone study had a frequency of over
50%: A. scrobiculata , Gi. gigantea , and S.
persica. None of these appears in the present
study. The most commonly isolated fungi in
the present study are GL etunicatum , Gl.
macrocarpum , and Gl. aggregatum; only the
latter was isolated from Moonstone Beach.
The fungal flora of Wisconsin’s Great
Lakes dunes are at least as similar to that of
the disturbed stream and river bank soils of
Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
central Iowa (Walker et al. 1982) as they are
to the dunes of Rhode Island. There were
three species in common with the Iowa
study: A. spinosa , S. calospora, and Gl.
geosporum. Nevertheless, the higher plant
flora of the Iowa study (. Fraxinus americana
L., Bromus inermis Leyss, and Setaria spp.)
differed greatly from that of the Great Lakes
dunes. Furthermore, at one site the soils
were not sandy but were composed of silt-
loam.
If the flora of Wisconsin sandy soils are
compared to that of a sandy prairie of Illi¬
nois (Anderson and Liberta 1989), we find
that there are three species in common: Gl.
geosporum , Gl. fasciculatum, and Gi. gigantea.
This prairie soil was dominated by little
blue-stem grass ( Schizachyrium scoparium
[Michx.] Nash).
Acknowledgments
The authors wish to express their apprecia¬
tion to Dr. Neil Harriman for his assistance
in identifying the higher plants and to Ms.
Jackie Bestler, Mr. Nathan Ihrcke, and
Ms. Vicki Verbrick for their help in the
laboratory.
Works Cited
Anderson, C. A. and A. E. Liberta. 1989.
Growth of little bluestem ( Schizachyrium
scoparium ) (Poaceae) in fumigated and non-
fumigated soils under various inorganic nu¬
trient conditions. Amer. J. Bot. 76:95—104.
Bray, J. R, and J. T. Curtis. 1957. An ordina¬
tion of the upland forest communities of
southern Wisconsin. Ecol. Monographs
27:325-49.
Gerdemann, J. W., and J. M. Trappe. 1974. The
Endogonaceae in the Pacific Northwest.
Mycologia Memoir No. 5. The New York
Botanical Garden. 76 pp.
Koske, R. E. 1981. A preliminary study of in¬
teractions between species of vesicular-arbus-
cular fungi in a sand dune. Trans. Brit. My col.
Soc. 76:411-16.
- . 1985. Glomus aggregatum: a distinct
taxon in the Glomus fasciculatum complex.
Mycologia 77:61 9-3 0 .
Koske, R. E., and W. L. Halvorson. 1981. Eco¬
logical studies of vesicular-arbuscular mycor-
rhizae in a barrier sand dune. Can. J. Bot.
59:1413-22.
Koske, R. E., J. C. Sutton, and B. R. Sheppard.
1975. Ecology of Endogone in Lake Huron
sand dunes. Can. J. Bot. 53:87-93.
Tews, L. L., and R. E. Koske. 1986. Toward a
sampling strategy for vesicular-arbuscular
mycorrhizas. Trans. Br. My col. Soc. 87:353-
58.
Schenck, N. C., and G. S. Smith. 1982. Addi¬
tional new and unreported species of mycor-
rhizal fungi (Endogonaceae) from Florida.
Mycologia 74:77-92.
Walker, C., C. W. Mize, and H. S. McNabb Jr.
1982. Populations of endogenous fungi at
two locations in central Iowa. Can. J. Bot.
60:2518-29.
Walker, C., and J. M. Trappe. 1981 . Acaulospora
spinosa sp. nov. with a key to the species of
Acaulospora. My cotaxon 12:51 5—2 1 .
Richard E. Koske is a professor of botany at the
University of Rhode Island. He has done a great
deal of research on the vesicular-arbuscular fungi ,
especially those growing on the dunes of New En¬
gland. Address: Botany Dept., University of Rhode
Island, Kingston, Rhode Island 02881.
Leonard L. Tews is a professor of biology at the
University of Wisconsin at Oshkosh. He has done
most of his research in the area of fungal ecology.
Address : Dept, of Biology and Microbiology, Uni¬
versity of Wisconsin Oshkosh, Oshkosh, WI 54901.
88
TRANSACTIONS
David J. Post
The plant communities of Nine-Mile
Island — past and present
Abstract Nine-Mile Island , in the Chippewa River , west-central Wisconsin ,
w currently under consideration for purchase as a natural area by
the Wisconsin Department of Natural Resources. Logging and agri¬
cultural use would be limited while occasional controlled burning
would be utilized in certain areas to maintain the prairies. Hunt¬
ing and other public uses of the land for enjoyment would still be
permitted. A description of the vegetation of Nine-Mile Island is pre¬
sented. Land Office survey records from 1848-49 indicate at that
time the island was about 35% floodplain forest, 25% oak open¬
ings, and 40% mesic forest. Settlement was limited to a single small
farm. 1990 quadrat sample data were used to trace changes that oc¬
curred since that time. Little change was found to have occurred in
the extent and composition of the forests, and the overall vegetation
pattern now present appears to be similar to that found on the is¬
land in the 1800s. Size class distribution suggests little change in
the composition of vegetation for the foreseeable future.
Nine-Mile Island of the Chippewa River in west-cen¬
tral Wisconsin provided an opportunity to investigate the
interactions between the vegetation and environmental factors
in a relatively natural and undisturbed setting. Although hu¬
man activities have exerted and continue to exert some influ¬
ence on parts of the island, much of the original vegetation
consisting of oak openings, oak forest, floodplain forest, and
prairie is present at this time.
In 1991, the Wisconsin Department of Natural Resources
(WDNR) acquired a 23.5 ha (63 acre) tract on the island,
which is to be designated a natural area. This area is to be pre-
TRANSACTIONS
Volume 82 (1 994)
89
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
served intact as an example of lowland for¬
est habitat. In addition, the WDNR has pro¬
posed the purchase of about 2145.75 ha
(5300 acres), including the remainder of the
island and some of the adjacent floodplain.
This natural area will preserve the plant
communities as well as bird and reptile spe¬
cies that are considered rare in Wisconsin.
In the establishment of a natural area it
is important to determine “baseline” condi¬
tions and to integrate current data in de¬
scribing the extent, distribution, and condi¬
tion of existing vegetation types (Noss
1989). The purpose of this study was to de¬
scribe the plant communities of the island
and to identify environmental factors re¬
sponsible for their presence. The original
survey records of 1 848—49 were used to re¬
construct the vegetation of that era. The
present vegetation was sampled using quad¬
rats. Standard phytosociological analytical
techniques were used to compare present
community types with those of the past and
to draw inferences about future composi¬
tional changes.
Study Site
Nine-Mile Island is located in the Chippewa
River in southern Dunn and northern Pepin
counties, Wisconsin. The island is located
in sections 35 and 36 of Township 26 N and
Range 13 W in Dunn County and sections
1, 2, and 3 of Township 25 N, Range 13
W in Pepin County (Fig. 1). It is about 56.5
km (35 mi) upstream from the Mississippi
River, approximately 6.5 km (4 mi) north
of the City of Durand, Wisconsin. The is¬
land is about 4.3 km (3 mi) long and 4.0
km (2V2 mi) at the widest place, with a to¬
tal area of about 1012 ha (2500 acres). The
topography is basically flat with elevations
ranging from 1.2 to 3 m (4 to 10 ft) above
the normal level of the river. The channel is
characterized by eroding banks of sand and
gravel. The island is situated about 1.6 km
(1 mi) below the confluence of the Chip¬
pewa and Red Cedar rivers and was formed
by the presence of a back channel commonly
referred to as Nine-Mile Slough. The back
channel carries less water than the main
channel and may be nearly dry when the
river level is low, leaving much of the sand
and gravel bed exposed. Changes have taken
place in the river channel surrounding the
island since about 1965. The meander on
the southwestern part of the island was cut
off, decreasing the size of the main island
(Fig. 1). Also, the meander at the northeast¬
ern corner of the island was cut off at about
the same time, causing another change in the
configuration of the island. Figures depict¬
ing aspects of the island’s vegetation prior
to the changes in the river channel include
the areas formerly considered a part of Nine-
Mile Island.
Loose, river-deposited sand and gravel
called “riverwash” occurs along the banks
and shorelines (Fig. 1). The vegetation is
sparse because the water level changes fre¬
quently and often floods the shoreline. At
higher elevations the island is mostly sandy
alluvial land with an isolated area of loamy
alluvial land. The soils are nearly level sandy
loams to silt loams and in places are nearly
level and poorly drained with a silty clay-
loam subsoil (USDA 1964, 1975). Frequent
flooding occurs on the western one-third of
the island where the average elevation is
about 1.2 m (4 ft) above the normal level
of the river. The normal level of the river
was determined as the line of demarcation
where terrestrial vegetation began (Peterson
and Gamble 1968). Swamp hardwood veg¬
etation and low land (below 4 ft above the
normal level of the river) are prevalent on
the western one-third of the island. The re¬
mainder of the island has an average eleva-
90
TRANSACTIONS
POST: The plant communities of Nine-Mile Island — past and present
Fig. 1. Nine-Mile Island in its geographic setting on the Chippewa River, west-cen¬
tral Wisconsin. Soil types: Ae, alluvial land, sandy; Lw, loamy alluvial land, wet; Sa,
sandy alluvial land; Re, riverwash. The dashed lines show elevation contours.
tion of about 1.8 to 3 m (6 to 10 ft) above
the normal level of the river. Oak forests,
scattered oak openings interspersed with
remnants of prairie vegetation, and some
mesic forest vegetation occur on these soils.
Flood frequency data for this region of
the Chippewa River was obtained from an
analysis that used a United States Geologi¬
cal Survey (USGS) method (USDA 1975).
The mean annual flood is defined by the
USGS as a flood having a recurrence inter¬
val of 2.33 years and occurring at an eleva¬
tion of .9 m on this island. An elevation of
1.2 m can be expected to flood every five
years; at 2.1 m, a flood can be expected ev¬
ery ten years. The entire island may be in¬
undated once in about every 60 to 80 years.
About 60% of the floods occur during the
months of March and April (Barnes 1989).
History of Settlement
The original land survey of Nine-Mile Island
and the surrounding area was conducted in
1848-49 prior to settlement. The surveyor
described the island as level with second-rate
sandy soils. The timber was described as
black, white, bur, or red oak on the level and
as maple, ash, elm, birch, aspen, and lind in
the bottoms. Much of the vegetation was
described as either timber, scattered timber,
or simply as “bottoms.”
Volume 82 (1994)
91
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
The first settlers came to the island dur¬
ing or soon after the original survey was
completed. During the late 1 840s and as late
as the 1850s there was a small wood-cutting
operation located on the central part of the
island. It provided fuel for the steamboats
that periodically traveled on the Chippewa
River between the Mississippi River and Eau
Claire. Although there is no record of exten¬
sive use of the land for crops, hay and grasses
were raised for forage, especially during the
depression years (1930s) when farmers used
all available land. The hay was often hauled
across the ice during the winter. Also, cattle
grazed on the island in a fenced pasture that
encompassed at least 202 ha (500 acres) dur¬
ing the depression and in the years follow¬
ing. Remnants of the old fence as well as old
mowing machinery are still present. There
are accounts of an old ferry measuring about
4 x 10 m, connected to the island by ropes
and cables, that was in service as late as the
1940s. This ferry was used to transport a
team and wagon to the island to haul hay
or to transport livestock. The only dwelling
presently on the island is an old hunting
cabin. On the same site are the remnants of
an old barn foundation purported to date
back to the 1850s as part of a small farm¬
stead. This small farm was apparently occu¬
pied until the 1940s and was used to raise
crops (Hubbard 1991). Finally, sportsmen
use the island today to hunt an evidently
ample deer population and other small
game.
Original Vegetation
Data from microfilm copies of the original
field notebooks were used to determine the
approximate distribution and composition
of plant communities present at the time of
the original survey. Corner points were es¬
tablished at .8 km (.5 mi) intervals where
east-west and north-south transects inter¬
sected (corners). A post was driven into the
ground at each corner, and the distance and
bearing to the four nearest trees was recorded
along with their common names and diam¬
eters. If no trees were nearby, the surveyors
constructed a mound of earth as a corner ref¬
erence and so recorded it (Barnes 1989). A
chain was used to measure the distance be¬
tween section corners; about 80 chains is
equivalent to 1.7 km (1 mi). The distances
to the trees nearest the corner point were re¬
corded in links (0.66 ft in a link).
Areas in which the trees were calculated
to be more than 15 m apart were mapped
as oak openings (<65 m) or prairie (>65 m)
(Curtis 1959), while the remaining areas
were mapped as either bottomland hard¬
woods or mesic hardwood forest (Fig. 2).
Survey data used were from twenty corners
that lie on Nine-Mile Island.
Eighteen of the twenty corner points were
ascertained by the original surveyors to have
trees less than 15 m (45 ft) apart, indicat¬
ing that the island was about 93% forested
in 1848. Three basic community types were
present on the island as interpreted from the
surveyors records: floodplain forest, mesic
hardwood forest, and oak openings within
which some small prairie occurred.
Floodplain forest comprised what was
then about 33% of the entire island. Silver
maple {Acer sac ch annum), basswood ( Tilia
americana ), and green ash {Fraxinus penn-
sylvanica ) were the most common species,
each occurring in 50% of the corner points.
Also abundant in the floodplain forest were
American elm ( Ulmus americana) and black
willow {Salix nigra) (Table 1). Butternut
( Juglans cinerea) was also recorded in parts
of the floodplain forest.
Oak openings occupied about 20% of the
island at the time of the 1848-49 survey.
(This includes land area that is no longer
92
TRANSACTIONS
POST: The plant communities of Nine-Mile Island — past and present
Fig. 2. The vegetation of Nine-Mile Island in 1848 as interpreted from the original
survey records
part of the island proper.) “Black Oak,” as
listed in the survey records, was the most
common species occurring in the oak open¬
ing community, being witnessed at 60% of
the corner points. This may have been red
oak ( Quercus borealis) or Hill’s oak ( Quercus
ellipsoidalis) . White Oak (most likely
Quercus alba) was also common here, occur¬
ring at 40% of the corner points. Other spe¬
cies present in the oak openings were Ameri¬
can elm, hackberry ( Celtis occidentalis ),
Populus sp., and river birch ( Betula nigra)
(Table 2). At one point along a line in this
area of the island (between sections one and
two in Pepin County) the surveyor recorded
“entering prairie” but did not list prairie
plant species. It may be possible to make an
inference about the species that occurred in
this area from Buss’s (1956) study on the
nearby Meridean prairie. He found that the
most abundant plants occurring on undis¬
turbed sites were big bluestem {Andropogon
gerardi ), spiderwort ( Tradescantia ohiensis ),
and flowering spurge ( Euphorbia corollata ).
Little bluestem ( Andropogon scoparius) ,
indian grass ( Sorgloastrum nutans ), and
purple prairie clover {Petalostemum pur-
pureuni) are species that Buss listed as being
scattered.
A “mesic hardwood” forest, as inferred
from the survey records, comprised about
45-50% of what remains the main part of
the island. Here, the number of individuals
of each species was more equal, although
Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1. Frequency (%F), average diameter (xdbh), and relative density (%RD)* of
trees of the Floodplain Forest at the time of the original land survey (1848-1849)
(5 corners, 12 trees)
Table 3. Frequency (%F), average diameter (xdbh), and relative density (%RD) of
trees of the Mesic Flardwood Forest at the time of the original land survey ( 1 848—
1849) (9 corners, 12 trees)
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POST: The plant communities of Nine-Mile Island — past and present
basswood, typically listed as “lind,” was the
most abundant at 44.4% of the corner
points. Other species that occurred fre¬
quently in the mesic hardwood forest were
American elm at 33.3% of the corner points,
maple (presumably silver maple) at 33.3%
of the corners, and hackberry at 22% of the
corners. In addition, some green ash, white
oak, and bur oak ( Quercus macrocarpa) were
witnessed (Table 3).
The land throughout the island was de¬
scribed in the survey records as “surface
level” with “second rate” soils. The vegeta¬
tion pattern coincided with the present de¬
scription in most respects except that there
seemed to be less open land around the oak
openings in 1848.
Present Vegetation
The vegetation was sampled during the au¬
tumn of 1990 using 1/100 ha (1/40 acre)
circular quadrats to sample trees and 1/40
ha (1/1/00 acre) circular quadrats to sample
herbaceous vegetation. One hundred seventy
(1/100 ha) quadrats were placed along
north-south transects established by a com¬
pass bearing at .32 km (.2 mi) intervals along
the entire width of the island. Adjacent
transects were spaced .32 km apart, so that
the entire island was divided into a nearly
regular grid pattern of quadrats. Tree diam¬
eters greater than 10 cm (4 in) at diameter
breast height (dbh) were recorded using a
diametric caliper, and sapling species were
recorded as present or absent. Herbaceous
species were also recorded as present or ab¬
sent within the quadrats. Soil samples were
collected at every tenth quadrat beginning
with the fifth and continuing throughout
the entire island. A consistent practice of col¬
lecting a sample of the top six inches of soil,
after removing the leaves and other duff, was
followed. The soil samples were analyzed for
percentages of water-retaining capacity,
sand, and organic matter. Elevation above
the normal water level was estimated using
a level and stadia rod.
Simpson’s Diversity Index was used to
calculate the degree of species diversity
within each community. This index assigns
a value between zero and one to a commu¬
nity; the closer the number is to one, the
higher the diversity. A definition of biologi¬
cal diversity is “the variety and variability
among living organisms and the ecological
complexes in which they occur” (Noss
1989).
A two-dimensional polar ordination was
performed using the importance values of
the tree species to determine the difference
in composition between the communities.
The degree of compositional dissimilarity
between each of the community pairs was
determined using the 1 - (2w/a + b) index.
Beal’s (1960) geometric method was used to
position the five communities in the plane
defined by the first two axes.
A histogram was constructed for the ma¬
jor species in each community type to reveal
their distribution of size classes. Trees were
divided into size classes ranging from 1 0 cm
(4 in) dbh to 30 cm (12 in) dbh.
A relative peak value method was used to
infer possible successional changes in the
mesic forest and floodplain forest. Succes¬
sion may be defined as the replacement of
some species by others through time. The
end result of succession is a community in
which the member species perpetuate them¬
selves through reproduction, are in a dy¬
namic balance with one another, and are in
equilibrium with the prevailing environmen¬
tal conditions (Buckholz and Pickering
1978). This technique plots the relative
number of stems of all important species in
a community against relative size classes.
Using relative numbers removes the effect of
Volume 82 (1994)
95
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
numerous species having higher peaks. Us¬
ing relative size classes (i.e., each size class is
expressed as a percentage of the largest size
class for the species) reduces the risk of mis¬
interpretation because different species attain
different maximum heights. The resultant
line graphs show the peak number of a given
species in each size class. The position of a
peak number for a species relative to other
species allows inferences to be made about
possible successional trends in a stand.
Young species will peak on the left side of
the graph while older species will peak on
the right side of the graph. A species with a
peak value on the right side of the graph in
a large size class is likely to be eventually re¬
placed by a species with a peak value closer
to the left side of the graph in a smaller size
class.
Community Descriptions
Five community types were identified on the
island: two oak openings, an oak forest, a
mesic forest, and a floodplain forest (Fig. 3).
Hill’s oak and red cedar (Juniperus vir-
giniana) were the most abundant trees of the
seven species in Oak Opening I, which oc¬
curs at an average elevation of about 2.7 m
(8 ft) above the normal level of the river
(Table 4). This community occurs on sandy
soil on the eastern end of the island and is
mostly inland. White oak was also prevalent
in this community, and basswood and green
ash were the most common associates of the
oak species. This oak opening had a density
of 83.3 trees/ha, a compositional index of
397, and a Simpson’s Diversity Index of
.827. Silver maple and American elm trees
and saplings were common, but were re¬
stricted to the low areas closer to the river.
Green ash and basswood saplings were com¬
mon throughout the area. In Oak Opening
I, white oak was present primarily in the 23
to 30 cm (12 to 13 in) size range (Fig. 4).
There were no small individuals and few sap¬
lings of white oak; most of the saplings and
small trees were basswood and green ash,
suggesting a change in future dominance
and possibly a change from oak opening to
forest. Fire was responsible for maintaining
the oak openings in Wisconsin (Curtis
1959) and may be necessary to maintain this
community type on Nine-Mile Island.
Table 4. Frequency (%F), mean diameter (xdbh), relative density (%RD), and
importance values (IV)* of trees in Oak Opening I (26 quadrats, 22 trees)
importance value (IV) is the number assigned to a tree species in a stand relative to other trees in that
stand. It is an average of the relative density (how many trees are present), the relative dominance
(size of the trees present), and the relative frequency (how often a tree occurs).
96
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POST: The plant communities of Nine-Mile Island — past and present
Fig. 3. The present vegetation of Nine-Mile Island as interpreted from sampling with
quadrats and aerial photographs (Oak Opening I, 0.0. 1.; Oak Opening II, 0.0. II.;
Oak Forest, O.F.; Mesic Forest, M.F.; Floodplain Forest, F.P.F.). The dashed lines
indicate north-south transects along which 1/40 acre circular quadrats were estab¬
lished at .32 km (.2 mi) intervals.
Shrubs present included prickly ash {Zan-
thoxylum americanum ), prickly gooseberry
( Rihes Cynosbati ), and grey dogwood ( Cornus
racemosa ), while staghorn sumac ( Rhus
typhina ) was abundant in the open areas.
Both big bluestem and little bluestem domi¬
nated in the oak openings along with abun¬
dant prairie smoke ( Geum riflorum ) and in¬
dividuals of several species of goldenrod
{Solidago spp) .
Red oak, swamp white oak ( Quercus bi¬
color ), Hill’s oak, and American elm were the
most abundant of the eight species of trees
in Oak Opening II (Table 5). This commu¬
nity is also located on the eastern end of the
island and occurs at an average elevation of
2.4 m (8 ft) above the normal level of the
river on sandy and poorly drained soils.
White oak was also abundant in this com¬
munity, and hackberry and green ash were
common associates of the oak species. This
community had an estimated density of 1 80
trees/ha, a compositional index of 506, and
a Simpson’s Diversity Index of .895. Green
ash and hackberry saplings were relatively
common as were pin oak ( Quercus ellip-
soidalis) and white oak saplings. The com¬
parative size class distribution indicates a
stable population of the dominant species
with an abundance of red and white oak sap¬
lings as well as older, mature trees (Fig. 5).
Prickly ash, grey dogwood, and staghorn
Volume 82 (1994)
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| Quercus
alba
Quercus
ellipsoidalis
Y/A Fraxinus
perinsylvanica
10 15 20 25 30 32.5+
Size Classes (cm)
Fig. 4. Size class distribution of three important tree species in Oak Opening I
sumac shrubs were abundant in the open
areas. Big bluestem, little bluestem, and rye
grass {Elymus canadensis) were prevalent in
this community along with abundant indi¬
viduals of goldenrod. The two oak openings
were distinguished by dominant oak species.
Pin oak was dominant in the first oak open¬
ing while red oak and swamp white oak were
dominant in Oak Opening II.
White oak was the largest and most abun¬
dant of the fourteen species of trees in the
oak forest (Table 6). This community oc¬
curs toward the east-central part of the is¬
land at an average elevation of about 2.4 m
(7.9 ft) above the normal level of the river,
somewhat lower than the elevation of the
oak openings. Red oak was also abundant,
and to a lesser extent bur oak and Hill’s oak
were also present.
Yellowbud hickory {Cary a cordiformis)
and green ash were the most common asso¬
ciates of the oak species. The oak forest had
an estimated density of 421 trees/ha, a com¬
positional index of 543, and a Simpson’s
Diversity Index of .826. Hackberry, bass¬
wood, and black cherry ( Prunus serotina)
were abundant, although black cherry was
found most often as a sapling. There were
scattered individuals of large, older white
pines near the north end of the island, and
American elm, red cedar, and silver maple
were common. A few individuals of sugar
maple {Acer saccharum) were also found in
the oak forest at the lower elevations. Yel¬
low bud hickory saplings were abundant,
and hackberry, cherry, and basswood sap¬
lings were also common. Red oak was the
most common of the oak saplings present
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POST: The plant communities of Nine-Mile Island — past and present
Percent
of Total
10
15
20
25
30
Size Classes (cm)
| Quercus
elllipsoidalis
IHI Quercus
borealis
Quercus
bicolor
ypp\ Ulmus
americana
32.5+
Fig. 5. Size class distribution of four important tree species in Oak Opening II
with fewer white oak and bur oak saplings.
The comparative size distribution suggests
continuation of the oak forest, but as shown
by the numbers of young trees present, it ap¬
pears that red oak may continue to domi¬
nate in the future (Fig. 6), and white oak
and basswood populations appear stable.
Prickly ash, prickly gooseberry, grey dog¬
wood, and red osier dogwood ( Cornus
stolonifera) were common shrubs in the un¬
derstory. Staghorn sumac was also present,
although less abundant than in the oak
openings. Rye grass and bottlebrush grass
( Hystrix patula ) as well as woodland nettle
(. Laportea canadensis) , Aster sp., and wild ge¬
ranium ( Geranium maculatum) were abun¬
dant throughout the oak forests.
Hackberry and green ash were the larg¬
est and two of the most abundant of the fif¬
teen species of trees in the mesic forest and
had as close associates basswood (the most
abundant tree species) and yellowbud
hickory (Table 7). Curtis (1959) described
basswood and red oak as dominant species
of mesic forests. This community occurs
over much of the central part of the island
at an elevation of 1.5 to 2.1 m (4.9 to 6.5
ft) above the normal level of the river. There
are some places where the land is low and
contains marshes. The soil is sandy loam,
with an isolated patch of silty loam, and has
a high percentage of organic matter and a
relatively high water-retaining capacity. This
community had an estimated density of 338
trees/ha, a compositional index of 659, and
a Simpson’s Diversity Index of .828. Ameri¬
can elm and white oak were also common.
Hop hornbeam ( Ostrya virginiana) occasion-
Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Percent
of Total
10 15 20 25 30 32.5+
EH
S22
Quercus
borealis
Quercus
alba
Quercus
macrocarpa
Tilia
americana
Size Classes (cm)
Fig. 6. Size class distribution of four important tree species in the Oak Forest
Table 5. Frequency (%F), mean diameter (xdbh), relative density (%RD), and
importance values (IV) of trees in Oak Opening II (1 1 quadrats, 20 trees)
100
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POST: The plant communities of Nine-Mile Island — past and present
Volume 82 (1994)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
ally occurred as saplings or small trees, and
silver maple was often found in low lying ar¬
eas. Yellowbud hickory saplings occurred in
almost every quadrat, while hackberry and
basswood saplings were also common. In the
mesic forest, it appears that green ash and
basswood may replace silver maple (Fig. 7).
Also, white oak is an important species, and
while some of the older trees may be re¬
placed, there are many white oak saplings
present to suggest that white oak will remain
an important species. Shrubs included
prickly ash, prickly gooseberry, red and grey
dogwood, and an occasional thornapple
(< Crataegus sp.). Blackberry (. Rubus alleghe-
niensis), raspberry ( Rubus sp.), and the vining
and climbing species Virginia creeper ( Par -
thenocissus quinquefolia ) and grape (Vitis
riparia) were abundant in the forested areas.
Rye grasses were also common in the less
shaded areas of the mesic forest.
Silver maple was the largest and most
abundant of the nine species of trees in the
floodplain forest (Table 8). Its major asso¬
ciates were green ash and American elm.
Basswood was often abundant on higher
ground while clumps of river birch were
found in the lowest areas. Silver maple,
American elm, or ash are often dominant in
floodplain forests in Wisconsin (Curtis
1959).
The floodplain forest occurs primarily on
the western end of the island with an iso¬
lated patch in the central part. The average
elevation is from .9-1.2 m (2. 9-3. 9 ft) above
the normal level of the river, which makes
this community prone to frequent flooding.
The soil is silty, poorly drained, and some-
Fig. 7. Relative peak values of four important species of the Mesic Forest
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TRANSACTIONS
POST: The plant communities of Nine-Mile Island — past and present
what lower in organic matter than the mesic
forest. These soils are higher in water-retain¬
ing capacity than the soils of other commu¬
nity types on the island. The floodplain for¬
est had a lower number of tree species than
the mesic forest and had an estimated den¬
sity of 325 trees/ha, a compositional index
of 507, and the lowest Simpson’s Diversity
Index of .712. It had a high estimated basal
area per ha of 47,267 cm3 because of the
many large, apparently older trees. Hack-
berry and slippery elm ( Ulmus rubrci) were
common, and occasional, mature individu¬
als of bur oak were also present. Hackberry
and basswood saplings occurred in almost
every quadrat, while green ash and Ameri¬
can elm saplings were also common.
In the floodplain forest, it appears that
green ash may eventually be replaced by
American elm, although that seems unlikely
because of Dutch elm disease, and some of
the large, older silver maples may be re¬
placed by green ash (Fig. 8). Prickly ash was
present throughout the area, although gen¬
erally the floodplain forest was more open
and had few shrubs. Rye grass was abundant
in less shaded areas, and Virginia creeper,
wild yam (Dioscorea villosa ), and grape were
common vines.
Community-Environment Relations
The result of the ordination shows the dis¬
tribution of the communities in a two-di¬
mensional space (Fig. 9). The distance be¬
tween the positions of communities along the
x-y axes represents the degree of composi¬
tional difference between them. The Five
Percent
of Total
Ulmus
americana
• • • • Fraxinus
Pennsylvania
Acer
saccharinum
25 37.5 50 62.5 75 87.5 100
Size Classes (cm)
Fig. 8. Relative peak values of three important species of the Floodplain Forest
Volume 82 (1994)
103
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
communities form two distinct groups, the
oak openings and oak forest in one group and
the mesic forest and floodplain forest in a sec¬
ond. The grouping appears to be related to
several environmental factors. The oak open¬
ings and oak forest occur at the higher eleva¬
tion where there is a lower water-retaining ca¬
pacity and higher sand content, while the
mesic forest and floodplain forest occur at the
lower elevation where there is a high water-
retaining capacity and lower sand content.
The communities at higher elevations
have lower water-retaining capacities because
of higher percent sand and lower organic
matter content, while the mesic forest and
floodplain forest at lower elevations have
higher water-retaining capacities because of
higher organic matter content and lower per¬
cent sand. The evident interrelationships
between the environmental factors are
shown in Table 9. The percentage of sand
is positively correlated to elevation, and there
is also a strong positive correlation between
percent of organic matter and water-retain¬
ing capacity (Table 10). Elevation and per¬
cent organic matter do not appear to be cor¬
related, while there is a negative correlation
between increasing elevation and water-re¬
taining capacity. It should be noted that
there were only five sets of data, and with
the resultant low number of degrees of free¬
dom, none of the values obtained were
found to be statistically significant above a
90% level of confidence.
Fig. 9. A two-dimensional polar ordination of the five communities by importance
values
104
TRANSACTIONS
POST: The plant communities of Nine-Mile Island — past and present
Table 10. Correlation coefficient values (r) for physical environmental factors exam¬
ined in the study
Volume 82 (1994)
105
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Discussion
The original survey records indicate that the
island was about 93% forested prior to
settlement (Fig. 10). About 30% of the area
was floodplain forest, 50% was mesic hard¬
wood forest, and 20% was oak openings.
Early settlement and the limited use of
the island for agriculture had little apparent
impact on the forests. Human activities were
limited to a small woodcutting operation on
the central part of the island and use of the
open prairie area for agriculture. There was
a small farmstead on the eastern end of the
island in the oak opening/prairie area on
which crops were raised. This farm dated
back to the late 1800s and was in use until
the 1940s according to a personal account
(Hubbard, pers. comm., 1991). Hay and
grasses were raised for forage in the open ar¬
eas, especially during the depression years
and early 1940s. No other evidence suggests
use of the open areas for crops although
cattle grazed in a fenced pasture of about
202 ha (500 acres) on the prairie. Remnants
of the old fence are still visible. Pasturing
may account for what seems to be an in¬
crease in the extent of the oak openings/prai¬
rie area since the 1940s.
The present composition of the forests is
similar to that of 1848. The same overall
configuration of plant community types was
observed on the island in the autumn of
1 990 as were described in the original land
survey records. The only change is that there
appears to be more open land occupied by
prairie vegetation. This may be a result of
the use of the land for pasture or simply the
lack of precision when working with the
1848-49 data.
Percent
Cover
YEAR H Forest
I I Open
Fig. 10. Changes in vegetation cover from 1848 to 1980
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POST: The plant communities of Nine-Mile Island - past and present
Elevation appears to be the major influ¬
ence on the development of plant commu¬
nity types on this island as it is related to
flooding frequency and to soil texture as well
as to organic matter content and water-re¬
taining capacity of the soil. Major changes
in the topography of the island are not likely
to occur; thus, significant changes in the veg¬
etation are unlikely in the future. Also, a
high frequency of saplings and small trees of
the most abundant species in the oak open¬
ings and oak forest suggests little probable
change in their tree composition in the fore¬
seeable future. In the mesic forest and flood-
plain forest relative peak value summaries of
the dominant species suggest that there may
be some successional changes. However,
such changes are likely to be small and
gradual and not have a pronounced effect on
the overall type and extent of these forests.
There is a note of urgency to the
WDNR’s current process of acquiring the
island as a State Natural Area. Parts of the
original floodplain forest remain threatened
by logging operations. In the past decade
about 202 ha (500 acres) of floodplain for¬
est timber currently owned by the Schlosser
Lumber Company have been either select-
cut or clear-cut, removing about 40-60%
of the canopy cover (Epstein, pers. comm.,
1990).
Most studies of vegetation history in Wis¬
consin have examined upland areas and have
reported drastic changes in the vegetation as
a result of man’s activities (Gleason 1913;
Curtis 1959; Stroessner and Habeck 1966;
Barnes 1974). However, extensive changes
in the nature of the vegetation of Nine-Mile
Island have not occurred. The only effect of
human impact in the past has been in the
form of minor woodcutting for fuel (with
available equipment, it is probable that only
trees 20 cm dbh or smaller were harvested),
pasture for dairy cattle in the open areas of
the island, and crops raised on a small farm
in the same area. Only the barn foundation
of the old farmstead remains. The only hu¬
man habitation on the entire island is a
hunting cabin on the site of the old farm¬
stead. Isolation on this island with its low
elevation and undeveloped soil types have
maintained these communities in their origi¬
nal state. Nine-Mile Island is a unique eco¬
logical resource containing rare plant and
animal species and is a good example of a
lowland forest habitat which should be pre¬
served in its nearly pristine state for the en¬
richment of future generations.
Acknowledgments
I wish to thank James A. Reinartz, Resident
Biologist and Manager of the University of
Wisconsin-Milwaukee Field Station in
Saukville, who reviewed the manuscript and
offered valuable advice for its improvement.
I also wish to thank Dr. William J. Barnes,
Ecologist and Professor of Biology at the
University of Wisconsin-Eau Claire, who
provided technical assistance, comments,
and suggestions for revision of the manu¬
script.
Works Cited
Barnes, W. J. 1974. A history of the vegetation
of Eau Claire County, Wisconsin. Trans. Wis.
Acad. Sci., Arts and Lett. 62:357-75.
- . 1989. A case history of vegetation
changes on the Meridean Islands of west-cen¬
tral Wisconsin, USA. Biol. Conserv. 49:1—16.
Beals, E. W. 1960. Forest bird communities in
the Apostle Islands, Wisconsin. Wilson Bull.
72:156-81.
Buckholz, K., and J. L. Pickering. 1978. DBH-
distribution analysis: an alternative to stand-
age analysis. Bulletin of the Torrey Botanical
Club 105:282-85.
Volume 82 (1994)
107
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Buss, I. O. 1956. Plant succession on a sand
plain, northwest Wisconsin. Trans. Wis. Acad.
Sci. Arts and Lett. 45:11-19.
Curtis, J. T. 1959. The vegetation of Wisconsin.
Madison: University of Wisconsin Press.
Epstein, E. 1990. Personal communication.
(DNR employee).
Gleason, H. A. 1913. The relation of forest dis¬
tribution and prairie fires in the Middle West.
Torreya 13:173-81.
Hubbard, E. 1991. Personal communication.
(Local landowner).
Noss, R. F. 1989. Indicators for monitoring
biodiversity: a hierarchial approach. Conser¬
vation Biology 4(4): 355-63.
Peterson, J. L., and C. R. Gamble. 1968. Mag¬
nitude and frequency of floods in the United
States, part 5. Hudson Bay and Upper Mis¬
sissippi River basins, USGS Water Supply
Paper, No. 1678.
Stroessner, W. J., and J. R. Habeck. 1966. The
presettlement vegetation of Iowa County,
Wisconsin. Trans. Wis. Acad. Sci., Arts and
Lett. 55:167-79.
United States Department of Agriculture, Soil
Conservation Service. 1975. Soil survey of
Dunn County, Ed. Gordon Wing.
United States Department of Agriculture, Soil
Conservation Service. 1964. Soil Survey of
Pepin County, Wisconsin. Series 1958, No.
29, Ed. Gordon Wing.
David Post teaches life science and earth science
at Greenwood Junior/Senior High School, Green¬
wood, Wisconsin. He holds bachelor's and master's
degrees in biology and education from the Uni¬
versity of Wisconsin— Eau Claire. Completing his
eighth year in education, David has taught high
school biology at Barron and Bruce schools and has
taught plant and animal biology courses in the
Biology Department at the University of Wis¬
consin— Eau Claire. Address: N7996 St. Hwy. 73,
Greenwood, WI 54437
108
TRANSACTIONS
Keith T. Weber
Analysis of black bear habitat
in northeastern Wisconsin
Abstract A study was performed in northeastern Wisconsin from June through
December 1991 to analyze the regional habitat and density of black
bear (Ursus americanus). Previously reported habitat requirements of
the black bear were used to establish areas that might be successfully
exploited by bears within a proposed 259.2 km2 ( 100 mi2) study site.
The field study area contained moderate to good black bear habitat
estimated to carry a bear population density of one bear/6. 1 km2 based
on roadside counts.
Black bear ( Ursus americanus) research by the Wisconsin De¬
partment of Natural Resources (WDNR) and the Univer¬
sity of Wisconsin— Stevens Point has focused recently in the
north-central and northwestern portion of the state (Kohn
1982; Anderson, pers. comm.). My study in northeastern Wis¬
consin was initiated to determine the present population of
black bear and to assess habitat quality. The results of this study
will aid in the understanding and management of the bear.
The study began on 1 June 1991 and terminated on 2 De¬
cember 1991. Known black bear habitats with varying popu¬
lation densities from around the United States and Canada re¬
vealed several key features, including soil characteristics, forest
type and age of stand, altitude, and the effects of timber har¬
vest, agricultural activity, hunting pressure, human population
density, and the road-to-forest ratio. Generally, black bears re¬
quire habitat containing a soft-mast food source (e.g., berries)
and a hard-mast food source (e.g., nuts and acorns) (Rogers
and Allen 1987) accompanied by forested cover (Unsworth et
TRANSACTIONS
Volume 82 (1994)
109
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 1. Comparison of various black bear population studies
Human population density figures were taken from the U.S. Census Bureau reports and reflect the
population at the time of the bear study. Bear densities are the mean of the estimated population range
given in the study (e.g., a study performed by Rogers [1987] reported a black bear density of one bear /
4. 1-6.3 km2. The value used above was the mean of 4.1 + 6.3 [x=5.2]).
al. 1989). Bear populations range from one
bear/ 1.3 km2 in California (Piekielek and
Burton 1973) to one bear/ 17.1 km2 in New
York (McCaffrey et al. 1974) (Table 1).
My research was conducted to ( 1 ) deter¬
mine the population of black bear in a local
study of northeast Wisconsin, (2) assess
habitat quality, and (3) compare the study
site with the findings from other black bear
studies.
Study Area
Field work was conducted in a 239 km2 (100
mi2) region in northeast Wisconsin, located
in the southwest corner of Marinette
County. Throughout this paper the study
site will be called the Marinette County
Study Area (MCSA) (Fig. 1). Elevation
ranges from 4l2m(1350 ft) above sea level
to 229 m (750 ft), with the mean elevation
extending from 274-305 m (900-1000 ft).
Soils in this region are primarily sand, with
the Menagha association (70%) and the
Mancelona-Emmet Menagha associations
(15%) being most prevalent. Seven percent
of the region is a Seelyville Markey associa¬
tion with 2.5% each of the Sarona- Kewee¬
naw and Ishpeming-Michigamee associa¬
tions. The majority of these soils were
created from glacial outwash and till
(87.5%) (Lorenz 1991). The remaining 3%
of soils is flooded by the High Falls reser¬
voir and Thunder Lake. Several streams,
small lakes, and ponds are not mentioned or
included in the above data.
The rock outcroppings in the Ishpeming-
Michigamee soil association can provide
good den sites for the black bear (Lorenz
and Thrall 1991), but bear usage of these
rock outcroppings, while not common, is
more typical of pregnant sows than boars
(Jackson 1961; Fair 1990). Much of this
area is wooded, providing the black bear
with forage, the bulk of which can be found
on well-drained uplands containing oak, ha¬
zelnut, and berries ( Quercus spp., Corylus
spp., and Rubus spp., respectively). The
MCSA consists of 90% well-drained soils,
of which 87% is forested uplands (Populus
spp., Pinus spp., and Quercus spp.), 7% is
lowlands ( Fraxinus nigra , Acer rubrum,
Thuja occidentals, and Picea mariana) , and
3% is rock outcroppings (Couvillion 1990)
(Figs. 2 and 3).
Forest composition in the MCSA is simi¬
lar to that published for the property within
the MCSA owned by the regional utility
company, Wisconsin Public Service Corpo¬
ration (WPSC). The WPSC property adja-
1 10
TRANSACTIONS
WEBER: Analysis of black bear habitat in northeastern Wisconsin
Fig. 1. The location of the Marinette
County Study Area (MCSA) in northeast
Wisconsin
cent to the High Falls reservoir is composed
of 86.8% uplands and 6.7% lowlands; the
balance is unforested (WPSC 1991). The
MCSA exhibits homogeneity in soil features
and forest composition.
Privately owned land accounts for 65%
of the study area. Public land comprises
25% while the remaining 10% belongs to
WPSC (Land Atlas and Plat Book 1985).
Forest inventory maps were made available
by the WDNR and WPSC for all public
and utility land, respectively. Forest compo¬
sition of the entire MCSA was approximated
by correlating forest inventory maps with
the Marinette County soils maps and topo¬
graphic maps.
Agricultural land comprises 19% of the
county (Decker, pers. comm.). A few large
tracts of forest exist, but forest edge is ex¬
tensive in the region because of substantial
logging. Studies performed in Idaho suggest
that black bears prefer 20-40-year-old stands
of timber that have not been previously clear
cut. Unsworth et al. (1989) and Fair (1990)
found that bears tend to avoid those areas
that have sustained a clear cut of 20 or more
acres. The age of the stand was considered
when evaluating habitat quality.
The use of sanitary landfills by bears has
been noted in other papers (Kohn 1982;
Rogers 1987), but such areas are absent in
the MCSA. An intensive study zone of 30.72
km2 (12.0 mi2) was established within the
MCSA to provide a better perspective of the
study area. The intensive study zone is de¬
lineated in Figures 2 and 3. The majority of
field research was conducted on public land.
Nevertheless, I shall assume that the results
found in the intensive study zone are appli¬
cable to the entire MCSA due to the homo¬
geneous nature of the soil and forest features
found throughout the region.
Materials and Methods
The research conducted in this study per¬
tains to the americanus subspecies of the
black bear ( Ursus americanus) which occurs
throughout northern Wisconsin and is typi¬
cally of the black color morph (98%)
(Rounds 1987). For this paper, adult bear
refers to a bear at least four years old, sub¬
adult to one between the ages one and four,
and cub to one less than one year old
(Rogers 1987). Henceforth, a female will be
referred to as a sow and a male as a boar.
Information on forest inventories was
provided by the WDNR (mapped by Cou-
villion) and the WPSC. Supplementary for¬
est surveys were conducted to provide an en¬
hanced picture of the study area and to
provide ground truth with the WDNR-
Couvillion forest inventory. I used the point-
quarter method (Smith 1974) to establish an
independent description of vegetation which
Volume 82 (1994)
1 1 1
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
R 18 E R IRE
Fig. 2. Map of soils found within the MCSA and the delineation of the Intensive Study
Zone
1 12
TRANSACTIONS
WEBER: Analysis of black bear habitat in northeastern Wisconsin
Fig. 3. Map of forest composition found in the Intensive Study Zone and Forest In¬
ventory Table for the MCSA
Volume 82 (1994)
1 13
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
could then be compared with WDNR for¬
est inventory maps; the results showed simi¬
larities between the two vegetation descrip¬
tions. Eighty quadrats were sampled along
two transects, resulting in a 2.5 m (8.2 ft)
mean point-to-plant distance and a mean
area per individual of 6.25 m2 (67.24 ft2)
(Fig. 3).
Bear populations were estimated by road¬
side count of tracks, droppings, and visual
observation. Note that roadside counts do
not generally represent actual populations
but are an index to the population and are
indicative of population trends. However,
because of the characteristic low population
density and sexual dimorphism of the black
bear and the relationship between bear
weight and track size (demonstrated by
Piekielek and Burton [1975]), an estimate
of the bear population in the intensive study
zone could be made with ample time spent
in the field. Counts were performed by ve¬
hicle once per week for twelve weeks over a
26 km route (n= 12, 25.9 km/ week [10 mi/
week]). The same route was driven on each
occasion, although minor deviations were
allowed and were sometimes necessary based
on road conditions. The number of distinct
bear sign observed was divided by the dis¬
tance traveled. Only those tracks unques¬
tionably made by a black bear were counted.
When a set of tracks was found, the most
distinct front and rear prints were chosen as
samples, and measurements were taken of
pad length, pad width, and width across the
toes. The sum of the six values is the com¬
posite foot measurement and can be used to
determine the approximate live weight of a
bear (Piekielek and Burton 1975; Kohn
1982). Additional data collected included di¬
rection of travel, the soil type in which the
track was made, the time when the track was
found, and the precise location of the prints
(Smith 1974). I estimated the age of the
tracks, the distance the animal traveled on
the forest road, and its speed of travel. The
location of each bear was determined in the
field and later plotted on a topographic map
to help distinguish individuals and to bet¬
ter understand the home range of bears in
the MCSA.
Results
The study of tracks located during the road¬
side count suggested that five bears (three
adults/sub-adults, two cubs) inhabit the in¬
tensive study zone of the MCSA. This is the
known minimum population (P ), deter¬
mined by comparing track size and weight
estimates derived by using composite foot
measurements. Individuals were recognized
by variations in track size that could not be
attributed to speed of travel or the soil me¬
dium. The Pk of five bears/30.7 km2 yields
a density of one bear/6. 1 km2 for the MCSA.
Standard deviation of the road side counts
was determined according to the formula
7
where x. represents the number of bears lo¬
cated in the intensive study zone during one
time period, x is the mean number of bears
located during the weekly count periods, and
n is the total number of weekly roadside
counts (x = 3.67, s = .58, n = 12).
Comparison of the Pk and the WDNR-
estimated black bear population density for
Wisconsin’s bear range (P = 1/ 3.8 km2 [1/
1.5 m2]) revealed a slightly lower population
in the MCSA. The WDNR has created
three black bear management zones within
the bear’s range across the northern third of
the state (i.e., zones A, B, and C). The
MCSA lies in management zone B, which
the WDNR estimates to have a population
density of P . The basis of the variation can
1 14
TRANSACTIONS
WEBER: Analysis of black bear habitat in northeastern Wisconsin
be illustrated by examining several factors:
(1) The MCSA is a relatively small region
within bear management zone B. Therefore,
various population density estimates should
be anticipated when sampling subsets of a
larger region (Amundsen, pers. comm.). (2)
A higher human population weighs heavily
against bear numbers, as a negative correla¬
tion seems to exist between the two factors
(Kohn, pers. comm.). The general agree¬
ment between these patterns and the con¬
ditions in my study area supports the results
obtained by this study.
Initially the results of the weekly counts
(x.) were alike, but as the autumn proceeded,
bear sign diminished. This probably was
due, at least in part, to the pressure exerted
on the bears by hunting, perhaps causing
them to avoid roads during the bear hunt¬
ing season (September-October). The re¬
duction could also be related to a diminished
amount of available food and the onset of
hibernation. A distinct drop in bear sign was
evident on 14 October; this trend contin¬
ued until the roadside counts were termi¬
nated on 28 October. For this reason, I have
limited my calculations from roadside
counts to the period prior to 14 October
1991. The roadside count index for 1991 is
.217 bears per kilometer of road traveled.
At least one boar is known to inhabit the
intensive study zone. A set of large tracks
(length of hind print = 267 mm [10.5 in])
was discovered in mid-September and, due
to size alone, the possibility of the tracks be¬
ing made by a sow was eliminated. Research
indicates that the hind print of a sow rarely
exceeds 240 mm (9.5 in) (Trauba, pers.
comm.; Jackson 1961). These tracks were no
more than a few hours old (a strong down¬
pour had occurred earlier that morning) and
made in soft sand. The weight of the boar
was approximated by employing the method
detailed by Piekielek and Burton (1975) us¬
ing composite foot measurement (800 mm
[31.5 in]) to determine the live weight of the
animal. The boar’s weight is approximated
at 147 kg (327 lb).
Discussion
Bears use forest roads as a means of travel.
The observed tracks indicate that bears tend
to follow the road for some distance (in one
case for over 200 m) rather than simply cross
the road. The bears that used the roads
nearly always traveled on the road edge and
in the same direction as vehicular traffic
flow. This appears to contradict the findings
of Unsworth et al. (1989) who reported that
black bears rarely used roads as travel routes.
However, since an estimate of the number
of bears that avoided roads was not known,
I cannot infer that all black bears in the
study area regularly use roads.
Two possible den sites were located in the
intensive study zone, both beneath the roots
of upturned trees ( Tsuga canadensis and
Thuja occidentals L.). One site was neatly
cleaned and contained the tracks of a sow
with one cub, believed to be the same fam¬
ily indicated earlier, although the tracks from
her second cub were not found (Fig. 4).
Black bears in the MCSA presumably use
ground-level dens because the mean Janu¬
ary temperature is -1 1.4°C (1 1.4°F) (Lorenz
1991) and excavated black bear dens occur
less frequently unless the mean January tem¬
perature falls below -20°C (-4°F) (Tietje and
Ruff 1980).
To evaluate the vitality of the bear popu¬
lation in the MCSA, data regarding repro¬
duction was necessary. As mentioned previ¬
ously, one sow is known to have given birth
to two cubs that were still alive in early Oc¬
tober 1991. Results of the 1991 mating sea¬
son are unknown. However, signs of activ¬
ity by area boars, including the presence of
Volume 82 (1994)
1 15
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 4. The author is shown investigating an overturned cedar where the tracks of a
sow and one cub were found. It is not certain if bears will use this site for denning
purposes as it is susceptible to spring flooding.
“bear trees,” indicates that some competition
for mates may have occurred.
Black bears, especially adult boars (85%),
use trees as marking posts during the June
mating season, presumably to communicate
their presence to other males (Laycock 1988;
Rogers 1987). Further, Rogers found that
utility poles were also used for marking pur¬
poses. The power line which cuts through
the intensive study zone was closely exam¬
ined for bear sign. All utility poles examined
(n = 30) were constructed of large southern
yellow pine (Pinus palustris ), and one was
positively identified as having been marked
by a bear (Fig. 5). This determination is
based on (1) the manner in which it is
marked, which corresponds well with typi¬
cal marking behavior (predominately open¬
ings or edges with markings toward the
opening), (2) five individual claw marks evi¬
dent in each stroke (width =133 mm [5.25
in]) reaching a height of 2.12 m (83.25 in),
and (3) six black hairs (length = 20-65 mm)
removed from the pole between a height of
.61-1.27 m (24-50 in). Because these hairs
suffered little bleaching from the sun, one
may conclude that this pole was marked dur¬
ing the 1991 mating season (Rogers 1987).
The availability of forage for bears in the
MCSA was studied in the field to help de¬
termine the quality of black bear habitat pro¬
vided by this area. Whereas an abundance
of soft-mast food was available, oak mast
(acorn) production for fall 1991 failed
throughout the county, perhaps because of
stress from previous years of drought, dam¬
age caused by the tent caterpillar (. Mala -
cosoma constrictum) , and injury from oak
1 16
TRANSACTIONS
WEBER: Analysis of black bear habitat in northeastern Wisconsin
wilt. The effect of oak mast failure is not
known at this time, but the abundance of
berries should alleviate this stress. A study
by Elowe and Dodge (1989) indicates that
if sows become nutritionally stressed, >66%
of potential mothers could reabsorb the blas¬
tocyst and come into estrus again the follow¬
ing season. Another study (Fair 1991) sug¬
gests that reproductive synchrony could
result from mast crop failure.
The mean age of the black bear popula¬
tion is a concern in bear management. Stud¬
ies in Montana by Jonkel and Cowan (1971)
reveal that the black bear does not reach
sexual maturity until it is 3.5 years old. This
is further supported by the findings of
Rogers (pers. comm.) and Anderson (pers.
comm.) in Minnesota and north-central
Wisconsin, respectively. Furthermore, a sow
may not breed until it is four years old and,
in some cases, the sow will not successfully
raise a litter until it is more than six years
old. Therefore, a healthy bear population is
expected to have a mean age of 4-5 years
(Laycock 1988). While a relatively young
population (mean age 4.1 years) presently
exists in the MCSA (Kohn 1982), the num¬
ber of sub-adults is estimated to be high, and
these bears are projected to carry the popu¬
lation to an older mean age (Amundsen,
pers. comm.). Thus, the age structure of the
bear population in the MCSA appears to be
healthy.
Bear populations can be affected by nu¬
merous factors such as development and
road construction, which tend to fragment
the population and produce “islands” of bear
habitat isolated from one another by regions
of unsuitable habitat. Research indicates that
the “black bear cannot function as a popu¬
lation” when its density drops below one
bear/25.9 km2 (1/10 mi2) (Fair 1990). The
bear population density in the MCSA (one
bear/6.1 km2) is much higher than this
Fig. 5. This photograph shows a utility
pole which was used by a boar, presum¬
ably to signal his presence to other
bears. The author is shown here to bet¬
ter illustrate the height of the marks.
“threshold level,” indicating that, currently,
the black bear population in the MCSA is
healthy and that the quality of habitat is ad¬
equate to support this population into the
foreseeable future. This is based on the fol¬
lowing evidence: (1) the bears are known to
be reproductively active, (2) various forage
exists in the region offering alternative food
sources in the event that one source should
fail, (3) the mean age of the population is
4. 1 years (black bears in northeast Wiscon¬
sin are sexually mature at 3.5 years), and (4)
estimated population densities are high
enough to sustain a viable population as de¬
fined by researchers elsewhere (Fair 1990;
Faycock 1988).
Volume 82 (1994)
1 1 7
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
The population estimate of one bear/6.1
km2 is lower than the WDNR estimate (one
bear/3.8 km2), probably because of the fol¬
lowing factors rather than a decline in bear
population or discrepancy in population es¬
timates: (1) the MCSA is only a portion of
bear management zone B, (2) different
methods were used to arrive at the two
population estimates, and (3) a higher hu¬
man population exists in the MCSA
(weighing against bear population density
[Kohn, pers. comm.]) than in northwest
Wisconsin.
Acknowledgments
I am grateful to W. J. Johnson, project ad¬
visor, UW Center-Marinette, R. Amund¬
sen, J. Beecham, G. Egtvedt, B. E. Kohn,
W. Kowalski, A. G. MacHutchon, D.
Schad, S. Stubenval, S. D. Trauba, and the
UW-Marinette County Extension Office
for sharing their data and supporting this
project. R. Howe and an anonymous re¬
viewer aided greatly in the development of
this paper. Field assistance was provided by
S. Y. Weber. To all those involved in this
project, I thank you.
Works Cited
Beecham, J. 1983. Population characteristics of
black bears in west central Idaho. Journal of
Wildlife Management 47(2):405-12.
Bureau of the Census. 1982. General Population
Characteristics— WI, 1980. U.S. Dept, of
Commerce, Bureau of the Census. Washing¬
ton, D.C., 51-12.
Couvillion, C. 1990. Wisconsin Department of
Natural Resources (WDNR) Forest Inven¬
tory. Manual code 8625.2. Madison. 15 pp.
Elowe, K. D., and W. D. Dodge. 1989. Factors
affecting black bear reproductive success and
cub survival. Journal of Wildlife Management
53(4):962-68.
Fair, J. 1990. The great American hear. Minoc-
qua, WI: Northwords Press. 192 pp.
Jackson, H. H. T. 1961. Mammals of Wiscon¬
sin. Madison: University of Wisconsin Press.
504 pp.
Jonkel, C. J., and I. McT. Cowan. 1971. The
black bear in the spruce-fir forest. Wildlife
Monographs, No. 27. 57 pp.
Kohn, B. E. 1982. Status and management of
black bears in Wisconsin. Wisconsin Depart¬
ment of Natural Resources Technical Bulle¬
tin No. 129. Madison: WDNR. 32 pp.
Land Atlas and Plat Book, Marinette County, WI.
1985. Rockford IL: Rockford Map Publish¬
ing.
Laycock, G. 1988. The wild bears. New York:
Outdoor Life Books. 272 pp.
Lorenz, H. E., and T. P. Thrall. 1991. Soil Sur¬
vey of Marinette County, Wisconsin.
McCaffrey, E. R., G. B. Will, and A. S. Berg¬
strom 1974. Preliminary management impli¬
cations for black bears ( U. americanus) in the
Catskill region of New York State as the re¬
sult of an ecological study. Bears — Their Bi¬
ology and Management. Paper presented at the
International Conference on Bear Research
and Management. IUCN No. 40., pp. 235-
45.
Piekielek, W., and T. S. Burton. 1975. A black
bear population study in northern California.
California Fish and Game 61(1 ) :4 — 25.
Raybourne, J. W ., G. L. Alt, and M. R. Pelton.
1987. Restoring America's wildlife. U. S. De¬
partment of the Interior, Fish and Wildlife
Service. Washington, D.C.: GPO, 394 pp.
Rogers, L. L. 1987. Effects of food supply and kin¬
ship on social behavior, movements, and popu¬
lation growth of black bears in northeast Min¬
nesota. Wildlife Monographs, No. 97. 64 pp.
Rogers, L. L., and A. W. Allen. 1987. Habitat
suitability index models: black bear, upper
great lakes region. Biological Report 82
(10.144) Sep. U.S. Department of the
1 18
TRANSACTIONS
WEBER: Analysis of black bear habitat in northeastern Wisconsin
Interior, Fish and Wildlife Service.
Rounds, R. C. 1987. Distribution and analysis
of colourmorphs of the black bear ( Ursus
americanus ) . Journal of Biogeography 14:521—
38.
Smith, R.L. 1974. Ecology and field biology. 2d
ed. New York: Harper and Row. 832 pp.
Tietje, W. D., and R. L. Ruff 1980. Denning
behavior of black bears in boreal forest of
Alberta. Journal of Wildlife Management
44(4):858-70.
Unsworth, J. W., J. J. Beecham, and I. R. Irby.
1989. Female black bear habitat use in west
central Idaho. Journal of Wildlife Management
53(3):668-74.
Wisconsin Public Service Corporation. 1991.
Comprehensive Land Management Plan for
Caldron Falls Project.
Keith T. Weber has been studying the black bear
in northeast Wisconsin since 1991, beginning un¬
der the supervision ofW. Johnson at UW Center—
Marinette. Weber continued his bear research while
at UW-Green Bay, advised by H. J. Harris and
R. Howe. He is currently studying elk at the Uni¬
versity of Montana Missoula under Dr. C. Les
Marcum. Address : 1910 Scott #4, Missoula, MT
59802
Volume 82 (1994)
1 19
Wisconsin Academy of Sciences, Arts and Letters
Executive Director
1994 Academy Council
LeRoy R. Lee
Officers
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Gerd H. Zoller, Secretary/Treasurer, Madison
Councilors-at-Large
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Linda Stewart, Milwaukee
Carl A. Weigell, Milwaukee
Councilor-at-Large Emeritus
John Thomson, Mt. Horeb
Your membership will encourage research, discussion and
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Wisconsin Academy of Sciences, Arts and Letters
1922 University Avenue
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Telephone (608) 263-1692
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Wisconsin Academy of Sciences , Arts and Letters
1922 University Avenue
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Telephone (608) 263-1692
ISSN 0084-0505
Volume 83 • 1995
125th Anniversary Issue
TRANSACTIONS
of the Wisconsin Academy of Sciences, Arts and Letters
Volume 83 • 1995
125th Anniversary Issue
Editor William J. Urbrock
Department of Religious Studies
University of Wisconsin Oshkosh
Oshkosh, Wisconsin 54901
Managing Editor Patricia Allen Duyfhuizen
328 West Grant Avenue
Eau Claire, Wisconsin 54701
Interns Cynthia Barber
Jennifer Fandel
Sheri Jackson
Gretchen Toth
1 ransactions welcomes articles that explore features of the State of
Wisconsin and its people. Articles written by Wisconsin authors on
topics other than Wisconsin sciences, arts and letters are occasionally
published. Manuscripts and queries should be addressed to the editor.
Submission requirements: Submit three copies of the manuscript,
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© 1995 Wisconsin Academy of Sciences, Arts and Letters
All rights reserved
ISSN 0084-0505
For information on membership in the Academy,
call (608) 263-1692.
Contents
TRANSACTIONS
Volume 83 • 1995
From the editor v
Part One: 125th Anniversary Articles
The Age of the Quartzites, Schists and Conglomerates of Sauk Co., Wis. 3
Roland Duer Irving
First appeared in Transactions 1 (1870-72) 128-137
Oconomowoc Lake and Other Small Lakes of Wisconsin,
Considered with Reference to Their Capacity for Fish-Production 1 1
Increase Allen Lapham
First appeared in Transactions 3 (1875-76) 30-36
Copper Tools Found in the State of Wisconsin 1 8
James Davie Butler
First appeared in Transactions 3 (1875-76) 99-104
United States Sovereignty — Whence Derived, and Where Vested 25
William Francis Allen
First appeared in Transactions 3 (1875-76) 125-132
On the Extent and Significance of the Wisconsin Kettle Moraine 33
Thomas Chrowder Chamberlin
First appeared in Transactions A (1876-77) 200-234
The Larger Wild Animals That Have Become Extinct in Wisconsin 65
Philo Romayne Hoy
First appeared in Transactions 5 (1882) 255-257
Some Personal Recollections of Abraham Lincoln 69
John Wesley Hoyt
First appeared in Transactions 16 (1909-10) 1305-1309
Hi
Part Two: Current Articles
Dairying in an urban environment : The Milwaukee metropolitan area 75
John A. Cross
Urban expansion has displaced dairy farmers from the Milwaukee metropolitan area,
with some farmers now relocating to other parts of Wisconsin.
Small mammal distribution associated with commercial cranberry production 87
Eric E. Jorgensen and Lyle E. Nauman
Small mammals are unevenly distributed in wetlands associated with commercial
cranberry production.
The effect of manure management on phosophorus and suspended solids 91
in the Lake Tainter, Wisconsin, watershed
Ken Parejko and Douglas Wikum
Fields near streams in Barron and Dunn Counties, Wisconsin, were spread with turkey
litter before and after snowmelt in spring, 1993. Monitoring runoff by sampling the
streams upstream and downstream from the fields did not detect significant loading of
phosphorus or sediments.
The effect of picnic beetles on European corn borer larval mortality 1 05
Kamela K. Schell and John L. Wedberg
The picnic beetle is often found in the corn agroecosystem in close association with the
European corn borer. This study demonstrates the reductive effects picnic beetles have
on corn borer populations.
IV
From the editor
And girlish April went ahead of them.
The music of her trailing garment's hem
Seemed scarce a league ahead. A little speed
Might yet almost surprise her in the deed
Of sorcery; for ever as they strove ;
A gray-green smudge in every poplar grove
Proclaimed the recent kindling.
he lines are from the opening portion of John
JL Neihardt’s epic poem “The Song of Three Friends.”
They evoke the spring of 1822 when the famous band of
one hundred trappers, the Ashley-Henry men, set out from
St. Louis to the beaver country of the upper Missouri River.
A sense of adventure, of new beginnings, and of still-unfold¬
ing springtime pervades Neihardt’s lyric depiction of the
scene.
In February, 1870, Dr. John Wesley Hoyt of Madison
addressed several hundred people gathered in the Assembly
Chamber of the Wisconsin State Legislature to consider the
feasibility of forming a Wisconsin Academy of Sciences, Arts
and Letters. As he describes the occasion in a letter of remi¬
niscence written forty years later in February, 1911, this un¬
dertaking also was begun with a sense of new beginnings and
of high adventure. Indeed, the embarkation was not with¬
out obstacles, especially the opposition of those who, while
admitting that such an institution might be very useful, felt
that the attempt to organize such a comprehensive society
was quite premature at such an early stage in the history of
the State. Undaunted, and lured, as it were, by “girlish April
. . . scarce a league ahead,” Hoyt presented a plea of such
conviction, invoking the spirits of Christ and Emerson, that
his dream for the immediate formation of the Academy car¬
ried the day. Accordingly, just a month later, on March 16,
1870, the Academy was formally chartered and incorporated
by “the people of the State of Wisconsin, represented in Sen¬
ate and Assembly.” Hoyt was himself elected to serve as the
Academy’s First president from 1870-1875.
The Act of Incorporation charged the Academy with en¬
couraging research in “the material, metaphysical, ethical,
v
ethnological and social sciences,” with un¬
dertaking “a progressive and thorough sci¬
entific survey of the State,” with advancing
both the fine and useful arts, including origi¬
nal invention, with fostering philological
and historical research, with the formation
of appropriate libraries and museums (!),
and, finally, with “the diffusion of knowl¬
edge by the publication of original contri¬
butions to science, literature and the arts.”
Just two years after the Academy’s incorpo¬
ration the first volume of Transactions ap¬
peared. It reported on Academy proceedings
for 1870-1872 and carried a lengthy presi¬
dential report on the founding of the Acad¬
emy, lists of officers and members, the
Academy’s charter and constitution, and an
impressive number of original scholarly ar¬
ticles.
In a commemorative address to those
gathered at the Fiftieth Anniversary Meeting
of the Academy in 1920, Professor Thomas
Chrowder Chamberlin looked back at the
Academy’s heady beginnings. Then at the
University of Chicago, Chamberlin had
been another of the original incorporators of
the Academy and had served as President of
the University of Wisconsin and as president
of the Academy from 1885-1887. In this
address, Chamberlin noted how the
Academy was formally inaugurated largely
upon the ideals, fond hopes, and aspirations
of those who planned it, and especially on
the tenacious vision of Dr. Hoyt. “Scarcely
a dozen of those who signed the call for the
convention,” he observed, “were productive
workers in any of the fields embraced within
the purposes of the Academy.”
Among these dozen or so early contribu¬
tors who were to set the tone for the future
of the Academy, Chamberlin singled out Dr.
Increase Allen Lap ham, the first General
Secretary of the Academy and editor of
Transactions , who was “at once a botanist, a
zoologist, an archeologist, a geologist, and a
meteorologist. He was a distinguished ex¬
ample of the best order of the old school of
all-around students of natural science.”
Lapham served as Wisconsin’s state geolo¬
gist; he collected what was to become the
nucleus of the University of Wisconsin’s
enormous herbarium; and he was largely re¬
sponsible for the establishment of the U.S.
Weather Service.
Chamberlin also took special notice of
such lights as Lapham’s friend and co¬
worker, Dr. Philo Romayne Hoy, a person
who “bubbled over with enthusiasm” for
naturalistic investigations, and Dr. William
Francis Allen, whose papers “set a high stan¬
dard of true original investigation in human¬
istic lines”; of “the sprightly literary contri¬
butions of the inimitable Dr. James Davie
Buder, and of the notable contributions of
the Academy’s fourth president, Professor
Roland Duer Irving, who “took an active
part in leading geological inquiry along
sound scientific lines.”
Upon completion of this anniversary ad¬
dress, Chamberlin himself was presented the
honorary degree of Doctor of Science by
President Birge of the University of Wiscon¬
sin. The citation noted Chamberlin’s half-
century of contributions to the Academy
and to the State of Wisconsin, including his
direction of the Wisconsin Geological Sur¬
vey and praised the “spirit and temper of sci¬
ence” which he had embodied in his life and
work.
1995 marks yet another significant mile¬
stone for the Academy, the 125th Anniver¬
sary of its founding. To commemorate the
occasion, I have selected for this special an¬
niversary issue of Transactions a colorful
sampling of articles written by the illustri¬
ous founders and scholars identified above.
All of the articles, except for Hoyt’s remi¬
niscence on Abraham Lincoln, appeared in
vi
the first five volumes of Transactions. Al¬
though necessarily limited in number, they
represent some of the diversity of interest
and expertise of the early contributing mem¬
bers of the Academy. They still make mar¬
velously engrossing reading, whether for
their pioneering scientific observations and
insights (viz., the articles by Irving, Lapham,
and Chamberlin), or their zoological (Hoy),
anthropological (Butler), and political (Allen
and Hoyt) interests. Like Neihardt’s “gray-
green smudge in every poplar grove,” these
articles proclaim “the recent kindling” of
scholarly productivity by these and many
other members in the earliest “Aprils” of the
Academy. Moreover, the contribution by the
ever-productive Lapham on Oconomowoc
Lake seems peculiarly noteworthy, since its
writing was completed during the afternoon
of September 14, 1875, on the very evening
on which Lapham died.
Also in this issue of Transactions we in¬
clude four new contributions, all of which
relate in one way or another to Wisconsin
dairying and agriculture. John Cross writes
about an “endangered species,” dairy farm¬
ing in the expanding Milwaukee metropoli¬
tan area. Kamela Schell and John Wedberg
discuss the interactive effects of picnic
beetles and corn borers on sweet corn, while
Eric Jorgensen and Lyle Nauman document
small mammal distribution associated with
cranberry production in south-central Wis¬
consin. Finally, Ken Parejko and Douglas
Wikum present results from their monitor¬
ing of manure runoff, resulting from two
different management strategies, at five sites
in the Lake Tainter watershed in Dunn
County.
Special issues involve the special efforts of
many. For helping to make this issue of
Transactions possible, I wish to acknowledge
especially the extra efforts of managing edi¬
tor Patricia Duyfhuizen and her student in¬
terns Cynthia Barber, Jennifer Fandel, Sheri
Jackson, and Gretchen Toth, and the val¬
ued advice and collaboration of Faith
Miracle, editor of the Wisconsin Academy
Review. Very special thanks are also extended
to Academy staff members Gaile Burchill,
Robert Lovely, Jean Sebranek, and Helen
Vukelich, all of whom spent many hours
word-processing the entire articles by Irving,
Lapham, Butler, Allen, Chamberlin, Hoy,
and Hoyt from the early issues of Transac¬
tions so they could appear in print once
again. Finally, a word of appreciation to the
State Historical Society for providing some
of the photographs that accompany the re¬
produced articles.
Now let me join all of those taking part
in the Wisconsin Academy’s many anniver¬
sary celebrations this year by offering a toast
to the next 125 years and to many more
springtimes, new beginnings, and adventures
for the Academy and for Transactions. May
“girlish April” always beckon with “the mu¬
sic of her trailing garment’s hem,” and may
our collective efforts, year after year, con¬
tinue to “proclaim the recent kindling.”
Bill Urbrock
The Wisconsin Academy of Sciences, Arts and Letters was
chartered by the State Legislature on March 16, 1870, as a
membership organization serving the people of Wisconsin. Its
mission is to encourage investigation in the sciences, arts and
letters and to disseminate information and share knowledge.
Part One:
125th Anniversary Articles
Managing Editor's note : We have prepared the anniversary articles to recreate the look of the originals, preserving
their “old style” margins, headings, and punctuation. — T.A.D.
Roland Duer Irving, E.M., Ph.D.
(1847-1888)
Professor of geology, mineralogy, and metallurgy at the
University of Wisconsin; comissioned geologist for Wisconsin
Geological Survey and United States Geological Survey; fourth
president of the Wisconsin Academy, 1882-1884
i
Wisconsin Academy of Sciences, Arts, and Letters
2
Age of the Quartzites, Schists, Etc., of Sauk Co.
THE AGE OF THE QUARTZITES, SCHISTS AND
CONGLOMERATES OF SAUK CO., WIS.*
BY ROLAND IRVING, E M.
Professor of Geology, Mining and Metallurgy at the University of Wisconsin
Through the central portion of the county of Sauk, Wisconsin, run
two ranges of hills or ridges, having an east and west trend, and a
height varying from a mere rise above the general prairie to an alti¬
tude of five hundred feet. The width from north to south never ex¬
ceeds three or four miles, and in places is much less than one mile.
The total lengths from east to west, or rather, the exact points at which
the peculiar rocks which make up the ridges give place to the ordi¬
nary country rock, are not as yet accurately known. These lengths,
however, seem to be from fifteen to twenty miles.
The rock material of the ridges is mainly a hard dark-colored quartz¬
ite; with this in some places are siliceous and talco-siliceous schists, and
two or three kinds of conglomerate. The dip of the strata, which, though
in some places obscure, is in others very marked — and can everywhere
be determined by careful observation — is uniformly toward the north. The
angle varies from 20 deg. to 25 deg. in the south range, to 75 deg. to 80
deg. in the north.
The occurrence of these bold ridges in the midst of a prairie coun¬
try, together with the marked contrast between their upturned and
metamorphosed layers and the entirely undisturbed strata of the
Potsdam and Calciferous epochs, which for miles around form the
country rock, has caused much speculation and discussion. From time
to time, during the past twenty years, brief notices have appeared in
various journals and reports, but no careful investigation of the lo¬
calities in question seems ever to have been attempted. In most of
these notices, or rather in most of those that are not absurdly inaccu¬
rate in their statements and wild in their ideas, the main point under
discussion has been the relative age of the metamorphic strata. Do
they, or do they not, antedate the Potsdam period? Are they the re¬
sults of local metamorphism on the Potsdam sandstones, or are they
the remnants of pre-existing rocks? The advocates of the former
theory have had the last word in the discussion.
* This paper has already been published, with some slight differences, in the Ameri¬
can Journal of Science and Art for February, 1 872.
3
Wisconsin Academy of Sciences, Arts, and Letters
The facts recorded in the present article are the results of a series
of visits made to the localities by the writer, during the months of Sep¬
tember, October and November of this year (1871), and they will, I
think, be seen to prove beyond all doubt or cavil, that the quartzites
and schists antedate entirely the Potsdam epoch, i.e., are either
Huronian or Laurentian in age.
Of all of the notices mentioned, none are more than brief mentions
and only a few seem to have any value at all. Dr. Shummard, in Owen’s
report on Wisconsin, Iowa and Minnesota, makes the first mention of
the quartzite. He gives no opinion. Dr. James G. Perceval, in the re¬
port of progress of the Wisconsin survey for 1856, refers again to the
quartzites, calling them merely “metamorphic sandstones,” but intimat¬
ing that they result from a change on the Potsdam sandstones. Mr.
James Hall, in his report of progress to the Governor of Wisconsin for
1860, gives by far the most accurate description I have been able to
find. He refers the quartzites unhesitatingly to the Huronian — but gives
no proofs whatever. His pamphlet did not fall into my hands until af¬
ter my own investigations were entirely completed. In the first vol¬
ume of his final report, Mr. Hall again mentions the quartzites, but still
more briefly, expressing the same opinion as before, and still giving
no proofs. In 1864 there appeared in the American Journal of Science
and Art (II, vol. xxxvn, p. 226) an article by Mr. Alexander Winchell
of Michigan, in which he describes, among others, some fossils from
the conglomerates overlying the quartzites; and upon them bases his
claim that the quartzites are a downward continuation of the Potsdam
sandstones. He himself never visited the localities. Finally, Mr. James
H. Eaton of Beloit College, in a paper read before the Wisconsin Acad¬
emy of Science, in February, 1871, expresses the same opinion though
on somewhat different grounds. The foregoing list includes everything
of any value that has been published on the subject.
The accompanying map includes those portions of the two ridges
where most of my observations have been made.
I. The South Range, to which my attention was first directed, pre¬
sents, on approaching it from Sauk Prairie on the south, a bold, and,
in places, precipitous rise from the plain of from 350-450 feet. The
northern side of this ridge has however, in all places as yet studied, a
much more gradual slope down to the valley of the Baraboo river, this
slope being in many places determined by the northward dip. Run¬
ning entirely through this ridge is a deeply cut valley, which has at
first for about two miles, a direction slightly north of west, and then
turns due north quite abruptly. This northern end holds the Devil’s
4
Age of the Quartzites, Schists, Etc., of Sauk Co.
Lake, which entirely fills the valley from side to side. Throughout
its whole length the sides of this cleft are precipitous masses of quartz¬
ite rising everywhere more than four hundred feet above the bottom,
and reaching at the lake an altitude of 501 feet above its level, and of
1,474 feet above the sea. The bottom of the valley is covered with a
heavy mass of Drift material, and the lake is held in its position by
low Drift hills at its northern and southeastern extremities. The bot¬
tom of the lake itself seems to be in a Drift sand, and is over most of
its area about thirty feet below the surface of the water. The lake has
no outlet; but draining as it does a very small amount of surface, the
extraordinary evaporation caused by reflection from the cliffs above,
together with the high winds of Wisconsin, is quite sufficient to ac¬
count for its maintenance of level; whilst the character of the surround¬
ing rock shows readily the reason for its not becoming saline.
The great exposures of cliff at this locality, and the deep rock cut¬
tings on the newly-opened railroad, afford most excellent opportuni¬
ties for study. The change of direction, too, of the valley, gives fa¬
cilities for approaching the rocks from different sides, not elsewhere
easily obtainable.
The rock here is mainly a hard, dark-colored, very compact quartz¬
ite, though the colors vary from a very light grey in places to deep
brownish-red. The bedding joints of the quartzite are in some places
rather obscure, but the railroad cuttings have so far exposed them, that
with a little care I was able readily to ascertain the dip. This on both
sides, and throughout the whole length, of the valley, is uniformly
about 20 to 25 degrees a little west of north. Some of the writers men¬
tioned, and notably Winchell, have described this valley as correspond¬
ing to an old anticlinal axis, but the uniform dip of the strata through¬
out its length proves, of course, that this is not the case.
Section 1. — North and south through the south range on section line 1 of map. A,
quartzites; A' quartzites with some schists; C, conglomerate; S.P., Sauk Prairie; B.V.,
Baraboo Valley; L, level of lake.
The quartzite, although often looking massive, shows in many
places, on weathered surfaces, the lamination and cross-lamination of
more modern sandstones. Many of the fallen masses show, too, on
5
Wisconsin Academy of Sciences, Arts, and Letters
exposed surfaces of lamination, the most distinct ripple markings I
have ever seen. On the shallow sandy bottom at the north end of the
lake below, may be found their very counterparts. Between the beds
of quartzite, in many places, are thin layers of a schist principally sili¬
ceous, but having always some talcose material. These correspond
apparently to the clayey or shaly layers between the beds of sand now
represented by the quartzite. In some places these layers seem to be
merely a thinly laminated quartzite, with talcose films covering the
laminae; in others, the talcose material pervades and gives character
to the whole mass, the siliceous material, however, always being
present.
The most remarkable feature of this locality is, however, the very
striking system of vertical joints which everywhere intersect the
quartzite. The bearing of these joints, taken in some fifty or sixty
different localities, I found to be uniformly N.E. and S.W. and S.E.
and N.W., the variations in a few places being evidently due to local
displacement. On the cliff sides, and more especially about the lake,
these joints, together with the bedding joints, have so cut the rock into
separate blocks, that these have from time to time been thrown down
the bluff by frost and atmospheric agencies in huge rectangular masses,
weighing by calculation from seventy-five to two hundred tons apiece.
In many places along the north flank of this ridge and lying always
above the quartzite, are outcrops of a conglomerate, containing pebbles
unmistakably from the quartzite below, always rounded, and in size
varying from a few lines to four or five inches in diameter. In some
few places there seems to be a second conglomerate in which the sandy
cement itself appears altered to a quartzite. This is a point, however,
deserving of further investigation. There are also places where dis¬
tinct layers of coarse and fine conglomerate occur, the latter always
above and graduating into a simple sandstone.
In this conglomerate are found in one locality just northeast of the
lake, the Potsdam fossils described by Mr. Winchell in the article re¬
ferred to, viz: Scolithus linearis Hall, Orthis Barabuensis Hall,
Delphinocephalus Minnesotensis Owen, etc. I have examined a col¬
lection of these fossils from the above locality, in the possession of
Dr. Lapham, of Milwaukee, and have seen the fossils and quartzite
pebbles in the same fragments side by side.
II. The observations on the North Range were made about the
Lower Narrows of the Baraboo river and westward from there about
half a mile. This north range seems to be less continuous both as to
elevation and as to the character of its rock material. I am told by
6
Age of the Quartzites, Schists, Etc., of Sauk Co.
Dr. Lapham that it seems rather to be made up of detached masses of
metamorphic rocks. The rising ground, however, never entirely dis¬
appears, and the quartzite seems to be found as far to the east and
west as in the south range. At the Baraboo Narrows the metamorphic
rocks are in great force, the cliffs on either side the river, which here
makes a direct cut through the range from south to north, being as
much as four hundred feet in height. The body of the bluff on the
Section 2. — Through North range at W. Bluff of Baraboo Narrows. A, thick-bed¬
ded dark colored quartzites, with some talco siliceous schist; B, siliceous schist; C,
horizontal sandstone; B. V., Baraboo Valley.
west side is made up of heavy beds of quartzite, with, in places, in¬
tercalated beds of metamorphic conglomerate, and of a talcose schist
like that in the south range. These beds all stand at a very high angle,
between 75 and 80 degs. from the horizontal, the dip being north, with
possibly a slight inclination to the east. At the bottom of the hill on
the south side is an exposure of a peculiar light-colored siliceous
schist, entirely different from any of the other rocks of the series. An
old shaft sunk some thirty feet on the schist, affords most excellent
opportunity for examination. The total thickness seen was about
twelve feet, the layers varying in thickness from a few lines to four
or five inches. Very thin films of a talcose material sometimes ap¬
pear between the layers. Directly above this schist, I found a hori¬
zontal undisturbed sandstone, laid open for some distance by quarry¬
ing. The beds are generally a foot or two in thickness. In the loose
pieces near by is found Scolithus linearis. The sandstone is, of course,
the Potsdam of the surrounding valleys. Section 2 will serve to give
a clear idea of the structure of this bluff.
The narrow detached ridge just to the westward, represented on the map,
is also made up of horizontal Potsdam sandstone. There are many other
such detached ridges along the Baraboo valley, bearing the same relation
to the quartzite ranges, and showing the same horizontality of strata.
7
Wisconsin Academy of Sciences, Arts, and Letters
The following arguments in favor of the priority of these rocks to the
Potsdam period will, I think, after what has been said, be admitted as
valid. I give them in the order in which they became apparent to me.
1st. The limited area of disturbance, the undisturbed Potsdam and
Calciferous strata being found north, south, and between the ridges,
and in close proximity to them.
2d. The absence of any anticlinal axes. Dipping as the rocks do
uniformly to the north, in order to place them in the Potsdam category,
we must imagine a metamorphism of the strata, accompanied by a great
fault, having on one side the unchanged sandstones, and on the other
the tilted quartzites and schists, an idea new, I think, to geology.
3d. The occurrence of rounded pebbles of quartzite in the conglom¬
erate on the south side of the south range. To suppose this conglom¬
erate. which by its fossils is unmistakably Potsdam, to be of the same
period as the quartzites below, we must suppose that period to have
lasted long enough to cover the deposition of the quartzites as sand¬
stones, their metamorphism, and the rounding of the pebbles by beach
action, before the formation of the conglomerate; not to speak of the
time sufficient to erase all signs of an anticlinal.
4th. The occurrence of horizontal sandstones resting uncon-
formably on the flanks of the tilted strata. This last is, of course, ab¬
solutely conclusive as to the north range , but lest it might be claimed
that the two are independent, I have given the others.
Mr. Winchell argues that, since Mr. Hall states that the fossils I have
mentioned as occurring in the conglomerate are restricted to the Middle
Potsdam, either this statement must be untrue or the quartzite must be the
downward continuation of this formation. This argument, however, loses
all force when we regard these ranges as high ridges in the Potsdam seas,
never having been entirely covered by these seas, but having merely had
the new sandstones and conglomerates deposited about their flanks. The
place where these fossils were found must be at least 200 feet above the
base of the sandstones of the surrounding country. A single glance at Dr.
Lapham’s geological map of Wisconsin will show this. The conglomer¬
ate is by no means necessarily the base of the Potsdam because it rests
immediately on Huronian or Laurentian rocks.
In the final report of Mr. Hall already referred to, he mentions a
low hill north of Baraboo, in which the middle of the hill is quartzite,
and the flanks conglomerate and sandstones graduating upward into
calcareo-sandy layers, without giving any further explanation. This
statement, before somewhat unintelligible to me, now throws further
light on my own results.
8
Age of the Quartzites, Schists, Etc., of Sauk Co.
To my mind these ridges were unquestionably islands in the
Potsdam sea, and a more beautiful illustration than is furnished by the
sandstones and conglomerates of wave action on a rocky coast, can
hardly be imagined.
There are many very interesting details of structure in these ridges
which would repay thorough study. The points presented in the present
paper are only those necessary to show the age of the rocks.
There are several more of these scattered quartzite ranges in Wis¬
consin, all but one of them occurring within the Potsdam and Calcif-
erous areas. During the coming season I hope to be able to make a
connected study of them.
University of Wisconsin, November 18, 1871.
9
Wisconsin Academy of Sciences, Arts, and Letters
Increase Allen Lapham, LL.D.
(1811-1875)
Civil engineer; all-around student of natural science; author of
first hardcover book published in Wisconsin, A Geographic and
Topographical Description of Wisconsin, etc., 1844; founder of
the herbarium at the University of Wisconsin, 1849; founding
member and president, State Historical Society of Wisconsin;
charter member and first elected secretary of the Wisconsin
Academy, and first editor of Transactions
10
Courtesy The State Historical Society of Wisconsin WHi(x22)1 1
Oconomowoc Lake, Etc.
OCONOMOWOC LAKE AND OTHER SMALL LAKES OF
WISCONSIN, CONSIDERED WITH REFERENCE TO
THEIR CAPACITY FOR FISH-PRODUCTION
BY I. A. LAPHAM
The Oconomowoc Lake in Waukesha county, on the line of the Chi¬
cago, Milwaukee and St. Paul Railway, is one of those beautiful sheets
of clear, cold water that may be taken as a type or representative of
hundreds of others within the State of Wisconsin. A few facts and
observations in regard to this lake may therefore be of interest to the
Fish Commissioners, and to all who desire to encourage the increase
of fish-production.
As shown upon the plats of the government land surveys, it has a
length of two miles; breadth, three fourths of a mile; a shore-line of
six and a half miles; covering an area of 830 acres, or one and three-
tenths square miles.
Its elevation above Lake Michigan, as ascertained many years ago,
in making the survey of the Milwaukee and Rock River Canal, is two
hundred and eighty-two feet. Its irregular form can best be seen by
reference to the accompanying chart.
The Oconomowoc River, a small stream which is the outlet of sev¬
eral other lakes, enters it on the north shore and leaves it at the north¬
west corner. So irregular is the shape of this lake that it might be
taken to illustrate geographical terms, as gulf, bay, point, cape, prom¬
ontory, peninsula; it has also straits, channels, bars, shoals and its
coast-line.
The banks of the lake consist mostly of high grounds which are
selected as sites of beautiful, often costly residences, which, especially
when duplicated by reflection from the smooth surface of the water,
form landscapes worthy of the pencil of the painter.
The lines of figures on the accompanying chart show the depth of
the water as measured in 1875. They indicate three principal depres¬
sions, the deepest being 66 feet,* the mean of all the soundings is 39
feet.
*The greatest depths measured in other lakes in the vicinity were:
1 1
Wisconsin Academy of Sciences, Arts, and Letters
12
one m i/e
Oconomowoc Lake, Etc.
There are several shoals with from two to six feet depth of water.
There is no deposit of mud or sand brought into the lake by the
river; the water supply both from the river and from the numerous
springs on the shore, being always clean and pure. One of these
springs on the south shore, known by its Indian name Minnewoc,
(place of waters) has been analyzed by Mr. G. Bode, of Milwaukee,
Chemist of the Wisconsin Geological Survey, with the following re¬
sult:
Chloride of sodium . . . . . . . . . . 0.129
Sulphate of soda . . . 0.627
Bicarbonate of soda . . . . . 1.041
Bicarbonate of lime . . . . . . . . . . . . 9.638
Bicarbonate of magnesia . . . . . 6.138
Bicarbonate of iron . . . 0.129
Alumina . . . . . . . . . . . . 0.067
Silica . . . . . . . . . . . 0.879
Total (grains in one gallon) . . . 18.648
It will be seen that the chief ingredients, as in most Wisconsin wa¬
ters, are lime or magnesia, derived doubtless directly from the mag¬
nesian limestone rocks and pebbles buried beneath the soil. This
analysis also shows that the water does not differ essentially from those
having great reputation for their medicinal virtues.
The lime from the springs is deposited, under favorable circum¬
stances, upon the bottom of the lake forming beds of pure white marl;
a process which is materially assisted by the secretions of mollusks
and aquatic plants, especially the chara and algae.
The temperature of the water, being an important item in fish cul¬
ture, was taken at different times near the surface, where it had con¬
siderable depth, with the following result:
In May . . . . . . . . . . . . . 41°Fahr.
In June . . . . . . . . 63
In July . . . . . . . . . 72 “
In August . . . . . . . . . . . 72
In September . . . . . . . 72 “
In October, 1874 . . . . . . . . . 53 “
An attempt was made to find the temperature at the bottom in
deep water and resulted in showing at some times no differences,
at other times one or two degrees warmer or colder; though the
13
Wisconsin Academy of Sciences, Arts, and Letters
deep water is popularly believed to be much colder than that at
the surface.
wind
The strong wind blowing over the lake causes a surface current
which must be balanced by a counter current below, and thus by
a constant interchange of water equalizes the temperature. If the
day is warm with but little wind, the surface water will become
the warmest; at night the surface cools down so that in the early
morning it is colder than at the bottom.
The deep-water fishes do not, therefore, seek that locality on ac¬
count of diminished temperature.
One lake is said to have remained open nearly all winter; the
cold weather having been accompanied by high wind, which pre¬
vented the water from freezing.
When the surface is once covered with ice the currents cease, and
ice is formed of great depth and of crystal transparency and purity.
The temperature of the spring-water along the shores remain [sic]
nearly uniform throughout the year, varying from 47 to 49 degrees,
which is not far from the mean temperature of this locality.
The currents caused by the wind blowing over the surface of the
lake, act upon the bottom and shores, causing abrasions at some places
and accumulations at others, very much as by the larger currents of
the ocean. This is quite apparent at two points on the channel be¬
tween the lake and the large bay at the northeast angle. The current
flowing into the bay from the lake causes an eddy at these points from
which are deposited long narrow bars projecting from the shore. This
channel it will be seen is quite narrow and the water in it shallow.
These currents also cause accumulations of beach sand and gravel
at certain points along the shore; separating and assorting the mate¬
rial upon a small scale, precisely as is done on a larger scale by the
currents in the great lakes, and in the ocean.
While white shell marl is accumulating in some portions of the lake,
soft muck resulting from the annual decay of aquatic vegetation is ac¬
cumulating in others. Some of the lakes, especially those not con¬
nected with a stream of running water, are thus becoming rapidly filled
with marl and peat, causing changes that become apparent after long
14
Oconomowoc Lake, Etc.
intervals of time. Some small shallow lakes have thus been changed
to meadows within the recollection of the first settlers of the county
only 38 years ago.
The government plats represent some lakes in 1835, which are now
only known as marshes or wet meadows. One called “Soft Water
Lake,” was a clean sheet of water only four years ago, but is now
nearly covered with the leaves of the yellow pond lily (Nuphar) and
other water plants. Soon it will cease to be known as a lake.
There are also some changes of the level of some of these lakes,
indicating a less amount of water than formerly. Sand bars formerly
covered with water are now dry, and in one case the bar extends quite
across the lake, thus dividing it into two. Another proof of a dimin¬
ished supply of water is afforded by the occurrence of ancient beaver
dams in places where no pond could be formed at the present time,
for want of running water.
The time may come when by the use of some simple, easily worked
dredge, the marl, and muck may be removed from the bottom of some
of the more important of these lakes, to be used as a fertilizer of the
neighboring farms; especially as the beauty of the lakes would be in¬
creased by deepening the water, and by the consequent removal of the
unsightly vegetable growth along their shallow margins.
Ice ridges are formed at certain places around the shore, some of
them double, or triple, and varying in height up to ten feet. These
ridges are formed by the expansion of the ice during the winter, push¬
ing the materials of the beach in-land. They consist of sand, gravel,
or boulders; in the latter case they constitute the so called “walled
lakes.” If the banks are high and steep at the edge of the water, no
ridge can be formed, but wherever low grounds or marshes approach
the lake, they may be looked for. Where springs enter the lake, no
ridges are formed, the water remaining above the freezing point all
winter. Trees are often found with their roots crowded inland by the
ice expansion; their tops leaning over the water. These ridges make
excellent road-beds, and are often used for that purpose.
The ancient mound-builders, that mysterious people who pre¬
ceded the present Indian races, once occupied the banks of these
lakes as is clearly shown by their numerous works; and they prob¬
ably derived no inconsiderable portion of their subsistence from fish.
No shell-heaps have been found to indicate their use of the abun¬
dance of Unio and Anodons found in these lakes. The works of
the mound-builders are rapidly disappearing, being levelled by the
plow of the farmer.
15
Wisconsin Academy of Sciences, Arts, and Letters
Besides the Unios, these lakes abound in other bivalve and univalve
mollusks; crustaceans and worms, and the larvae of insects appear in
wonderful numbers. These, with the innumerable minnows found in
shallow waters, afford at all times an abundant supply of food for the
larger fishes. Loons, geese, ducks, gulls, plover, and many other birds
swim upon the waters or wade along the margin.
Among the fishes to be found are the following:
Perch, Perea flavescens, Cuvier.
Wall-eyed Pike, Lucoperca americana.
Striped Bass, Roccus chrysops, Girary.
Rock Bass.
Stone-Roller, Etheostonia.
Black Bass, Micropluas nigricans, Agassiz.
Sun-Fish, Pomotis.
PUMPKINSEED.
Shiner.
Sheephead, Haploidonotus grunnieus.
Stickle-back, Applissinconstans, Kirtland.
Pickerel, Esox, Lesueur.
Sisco, Argysosomus sisco, Jordan Am. Nat., 1875, p. 135, Ind.
Geol. rep. 1875, p. 190.
Sucker, Catastomus.
Red-Horse, Plychostonus.
Cat-fish, Amiurus catus, Cuvier.
Bull-Head.
Bill-Fish, Lepidosteus oxyurus, Rafinesque.
The Salmon and Brook-trout are reared artificially, and have been
introduced into some of the lakes.
Young salmon (Salmo salar) and the brook-trout (S. fontinalis) have
been introduced into this lake, but so far as known they have not in¬
creased.
From the data given above one will be able to decide whether it
would be advisable for the State to attempt to stock this lake with fish;
and if so, the kinds best adapted to the conditions named.
The natural supply of fish has been drawn upon so heavily that the
present yield is quite small, compared with what it was a dozen or
more years ago; and hence the necessity of some effort for the resto¬
ration of the supply of the better kinds.
16
James Davie Butler, LL.D.
(1815-1905)
Student of theology and Congregational minister; seasoned
worldwide traveller and popular travel lecturer; professor of
Greek at Wabash College; professor of ancient languages and
literature at the University of Wisconsin; curator of the Wisconsin
Historical Society
17
Wisconsin Academy of Sciences, Arts, and Letters
COPPER TOOLS FOUND IN THE STATE OF WISCONSIN
BY PROF. J. D. BUTLER, LL.D.
Implements of unalloyed copper are among the most rare and curi¬
ous of archaeological findings. The exhibit of these articles now made
at the Philadelphia Centennial comprises the largest collection ever
brought together. The copper age proper, in distinction from the age
of bronze, forms a link in the chain of human development which ac¬
cording to Sir John Lubbock, “is scarcely traceable in Europe.” The
only European museum known to that distinguished archaeologist
which contains any copper tools is the royal Academy at Dublin. The
number there was thirty till within a year or two, when five were re¬
ceived from Gunjera — a province in India north of Bombay.
The articles now on view at the Centennial are as follows: In the
Government building, from the Smithsonian Institution, seventeen real
tools, besides casts of several others, and various copper trinkets. In
the same building two articles, much corroded, owned in the State of
Vermont.
In the mineral annex. From Ohio eight implements; from Michi¬
gan nineteen, and from Wisconsin, one hundred and sixty four. The
whole number from all quarters is two hundred and ten.
I made notes regarding all the exhibits, but having lost them; can
only describe the show from Wisconsin. But the coppers from that
State are nearly four times as many as all the rest of the world has
sent to Philadelphia, and they surpass others in size, variety, and per¬
fection of preservation, as much as in number. The only instrument
from any other source, not represented among Wisconsin Coppers, is
a crescent about six inches long — perhaps intended for a knife, though
it has no handle.
Among the varieties in the Wisconsin exhibit — which is made by
the State Historical Society — are the following:
Ninety-five spear-heads. Of these the larger number are what some
antiquarians called “winged,” that is the sides of the base are rolled
up towards each other so as to form a socket to receive a shaft. Some
of these sockets are quite perfect, and all are ingeniously swaged. Six¬
teen of them are punched each with a hole, round, square, or oblong,
for a pin to fasten the shaft, and one of the copper pins still sticks
fast in its place. Twenty-three of the spear-blades swell on one side
something like bayonets, the rest are flat. Three are marked with seven
1 8
Copper Tools Found in Wisconsin
dents apiece, and one with nine; indentations which have been fan¬
cied to indicate the number of beasts, or men, the weapons had killed.
Nine spear-heads have round tangs which are so long, smooth and
sharp, that they may well have been used as awls and gimlets. The
blades of these nine spears swell in the middle of each side. Their
shape is a beautiful oval. The largest specimen of this class is about
a foot in length. In the middle of its blade there is a hole as large as
a pipe-stem, which may have been drilled for putting in a cord to re¬
cover the spear when it had been thrown into the water. One spear
has a unilateral barb. This, meeting with unequal resistance, will not
go straight in water, so we think it of an absurd pattern. But the truth
is that if aimed at a fish where he looks to be, it will hit him where he
is — though, owing to the refraction of light in water, he is not where
he looks to be. One barb is then better than two, and we are the fools
after all. Spears of a similar pattern, though of other material have
been exhumed in France and California, and are still used in Terra
del Fuego. Specimens in bone from Santa Barbara may be seen in
the Smithsonian exhibit. Thirteen spears have flat tangs to thrust into
shafts. Six of these tangs are serrated or notched like the necks of
flint weapons for binding about with sinews. They seem to mark the
very point of transition from one material to another — from mineral
to metal.
There are fifteen knives. Most of these were intended to be stuck
in handles, but one of them has a handle rolled out of the same piece
of copper with its blade. Another has its copper handle bent into a
hook. There are several gads, or wedges, to be driven. There are three
adzes — tools beveled only on one side of their edges, and with broad
sockets for handles. There are eleven chisels, some as heavy as those
we now use. There are twelve axes, one weighing three and three-
quarter pounds in exactly the weight of those common among Wis¬
consin lumbermen to-day. Another, which is a pound heavier, is the
largest specimen of wrought copper that has even been brought to light.
There is one hook, and a square rod. There are more than half a dozen
borers of various sizes. One may be called an auger, being sixteen
inches long and three in circumference. There is a dagger ten inches
long with a blade an inch wide. These, with various anomalous ar¬
ticles, complete the catalogue.
For the conservation and display of this unique copper treasure the
State of Wisconsin has set apart one of the towers of the Capitol in Madi¬
son. There they will be daily open for inspection, and will no doubt be a
magnet attracting to themselves other curiosities of like nature.
Wisconsin Academy of Sciences, Arts, and Letters
The question is always asked, “Where did these coppers come
from?” It cannot be so definitely answered as is to be desired. Nev¬
ertheless something is known in respect to the finding of them. They
were all discovered within the limits of Wisconsin — while the
Smithsonian specimens — less than one eighth as many, were gleaned
from eight different States. Nearly all of them have come to light in
eleven southeastern counties of Wisconsin. Only in those counties
has much search been made.
Most of the Wisconsin coppers were brought together into one col¬
lection by the zeal and perseverance of one single man, Frederick S.
Perkins of Racine county. Five years ago this gentleman, though he
had long been forming a museum of stone implements, had never seen
one of copper. On the 25th of November, 1871, he was first shown
such an antique. It was a large spear-head that had been exhumed
three miles north of his residence in Burlington, Wisconsin. That No¬
vember date marks the birthday of his interest in copper — or his tran¬
sition from the stone to the copper age. His enthusiasm which had
been great for the former became greater for the latter. He had lei¬
sure — or he made it, to ride over county after county on every road,
waylaying every pedlar [sic], calling at every school, every store, at
almost every house. He advertised in newspapers, he threw tempting
baits abroad on all waters. He found what he sought, where no one
else would have looked for such a prize, and where many proved to
him that it could not be found. He has recorded the name and resi¬
dent, by county and town, of one hundred and twenty-one persons from
whom he obtained pre-historic coppers, as well as of three hundred
and twenty-five others who furnished him stone antiques, but had no
coppers to furnish. This record shows how thorough and wide-spread
were his researches. Indeed, although the Wisconsin Historical Soci¬
ety has bought the bulk of his findings, some of them are scattered
far and wide. Five of them are in the Central Park museum, others in
the Metropolitan in New York, others I think have enriched the
Smithsonian. A further question which must occur to every investi¬
gator, is, where were these implements obtained by those from whom
Mr. Perkins obtained them? On this point my information is more
scanty than it would be were not Mr. Perkins now in Europe, and than
it will be on his return. Large numbers of the tools were turned up in
plowing or hoeing. Others at greater depth in digging foundations of
houses or sinking wells. Not a few have come to light in burial
mounds close by skeletons. In one such mound at Prairie du Chien
an axe weighing two and seven-sixteenths pounds and eight inches
20
Copper Tools Found in Wisconsin
long was discovered lying on a large flint spade, fourteen feet below
the top of the mound, and seven feet below the level of the earth
around, and among human bones. Another axe, with other coppers,
was taken from a similar mound in Barron county. The only socket
spear-head which shows its rivet still in place, was found on a knoll
in plowed land by James Driscoll in May, 1874, at Lake Five,
Waukesha county. One knife was dug out of a mound by a dog while
hunting, in 1860, in Troy, Waukesha county. One chisel was met with
ten feet below the surface in cutting a road through a bluff at
Cedarburg, Ozaukee county, in 1871. One of the most remarkable
articles, a sort of copper pike, was dug up three feet under ground on
the bank of Pike Lake, Hartford, Washington county, by Samuel
Mowry in 1865. One massive celt, at first turned up in Merton,
Waukesha county, a pedlar [sic] had preserved for twenty years. Sev¬
eral knives and other implements found near lakes and rivers appear
to have been washed out of their banks. A lance-head found at
Rubicon, Dodge county, in 1869, has a lump or stud of silver on one
side of it.
But we cannot fail to ask, “who made these copper instruments?
was it Indians or some pre-Indian race?” It has been argued that they
are of pre-Indian origin because the skeletons with which they are dis¬
covered in burial mounds are not of the Indian type, but of a very dif¬
ferent cranial development. Again, as the mounds, multitudinous and
often of vast size, are beyond Indian industry, so the tools seem be¬
yond Indian ingenuity. Most of them indeed, are hammered, and so
show copper used rather as a mineral than as a metal. Others of the
coppers betray no marks of hammering, no laminations or flaws. Prac¬
tical foundrymen detect on them mould-marks where the halves of a
flask united, and so declare them smelted. Others they hold were run
in a sand-mold. These indications of casting are plainest on the larg¬
est piercer and on one of the chisels, except perhaps on certain imple¬
ments which Mr. Perkins has carried abroad for the conversion to his
views of trans-Atlantic skeptics regarding our pre-historic metallurgy.
All proofs that our coppers were cast, tend to show that they are not
the handiwork of Indians.
Our early annals indicate that our copper implements were a pre-
Indian manufacture. They testify that the earliest travelers in Wis¬
consin found the Indians using copper, if at all, only for trinkets and
totems, but not for implements either of war or of peace. Thus La
Salle on his last expedition through this region, well nigh two centu¬
ries ago, says of the Indians: “The extremity of their arrows is armed,
21
Wisconsin Academy of Sciences, Arts, and Letters
instead of iron, with a sharp stone or the tooth of some animal. Their
buffalo-arrow is nothing else but a stone or bone, or sometimes a piece
of very hard wood” Charlevoix, writing about 1720, mentions Indian
“hatchets of flint which take a great deal of time to sharpen, as the
only mode of cutting down trees.” “To fix them in the handle,” says
he, “they cut off the head of a young tree, and make a notch in it in
which they thrust the head of a young tree, and make a notch in it in
which they thrust the head of the hatchet. After some time the tree
by growing together keeps the hatchet so fixed that it cannot come
out. They then cut the tree to such a length as they would have the
handle.” “Both their arrows and javelins,” he adds, “are armed with
a point of bone wrought in different shapes.” According to Hennepin
about 1680, (2.103) “the Indians, instead of hatchets and knives, made
use of sharp stones which they fastened in a cleft piece of wood with
leather thongs, and instead of awls they made a certain sharp bone to
serve.” The Jesuit Father Allouez, writing about 1660, says “I have
seen in the hands of the savages, pieces of copper weighing from ten
to twenty pounds. They esteem them as divinities or as presents made
them by the gods. For this reason they preserve them wrapped up
with the most precious things, and have sometimes kept them time
out of mind.” In none of these or other early chronicles do I find any
mention of any copper tool whatever. Pre-historic mines about Lake
Superior are a proof that our copper implements are not Indian work.
No tradition of such mines was brought to light by early adventurers
among Indians. But if excavated by them to such an extent as we see
them, and for ages, how could they have been given up and even for¬
gotten? On the whole the evidence now before us tends to show that
our copper tools are the work of some pre-Indian race. The success
of Mr. Perkins in unearthing coppers in unlooked for numbers should
raise up a legion of copper-hunters. For encouraging such investiga¬
tors still more, my last words shall be regarding the greater harvest
than has crowned his labors which seems to me ripe for their sickles.
Indications are not wanting that our past prizes in copper hunts,
are all as nothing to what is in store for us. Pre-historic mining-pits
honeycomb Isle Royal all over. Along the south shore of Lake Supe¬
rior they are frequent for a hundred miles. They were every one rich
pockets. Their yield of copper must have been many times enough for
sheathing the British navy. What has become of this copper? It can¬
not have vanished like iron in oxidizing rust. It must still exist, and
lurk all around us. At Assouan the quarries prove to a stranger that
Egypt must be rich in granitic monoliths, for there we see the rock
22
Copper Tools Found in Wisconsin
whence they were hewn. Spanish treasure-ships sunk in the Carib¬
bean ages ago, still teach divers where to ply their sub-marine ma¬
chinery for richest spoils. In Greece, the Styx, and other catabothra,
or lost rivers — emptying into subterranean abysses, suggested to the
ancients streams that girdled the whole under world. So our mining
shafts sunk time out of mind are a prophecy and an assurance of cop¬
per bonanzas for explorers in the future so vast as will make us ut¬
terly forget whatever has been discovered. All hail such a ressurrection
[sic] of the copper age. The longer it has been lost the more wel¬
come will it be when found again.
23
Wisconsin Academy of Sciences, Arts, and Letters
William Francis Allen, A.M.
(1830-1889)
Expert in Roman history, voluminous author, and editor of the
classics; professor of ancient languages and history (later his¬
tory and Latin) at the University of Wisconsin; sixth president
of the Wisconsin Academy, 1888-89
24
United States Sovereignty
DEPARTMENT
OF SOCIAL AND POLITICAL SCIENCE
UNITED STATES SOVEREIGNTY— WHENCE DERIVED,
AND WHERE VESTED
BY W. F. ALLEN, A. M.,
Professor of History and Latin in the University of Wisconsin
The late war brought to an end the long and fierce controversy as
to the nature of the Federal Union. What argument had not been able
to decide, was decided by arms; and the United States are recognized
as a Nation, possessed of sovereignty. With the determination of this
controversy, however, another question has come into prominence, as
to the origin of this sovereignty. Before the war it was commonly
held that the act which severed the colonies from the mother country
had as its effect the creation of thirteen independent and sovereign
States; and that it was not until the formation of the Federal Constitu¬
tion that sovereignty was conferred upon the central government. This
doctrine, however, of the original sovereignty of the States, has been
thought to afford some foundation for the doctrine of Secession. Some
of the most ardent advocates, therefore, of the national and sovereign
character of our Union, have, since the war, brought into great promi¬
nence the theory that the Nation was not created by the States, but the
States by the Nation; that the States were never, in any true sense of
the term, sovereign, but that the act of independence created at once a
sovereign Nation. This view has been most fully elaborated in a se¬
ries of articles in the first volume (1865) of the Nation, by Hon. Geo.
P. Marsh, United States minister to Italy; it is presented also by Pro¬
fessor Pomeroy in his “Introduction to Constitutional Law.” In this
work the authority of Hamilton, Jay, Marshall, Story and Webster is
claimed for this theory. I do not think, however, that Marshall and
Webster can fairly be cited as its adherents. Mr. Pomeroy has given
no citations in support of his view, and on the other hand both these
jurists have expressed themselves unequivocally in favor of the origi¬
nal sovereignty of the States. Webster says, of the Confederation: “it
25
Wisconsin Academy of Sciences, Arts, and Letters
was a league, and nothing but a league.” Chief Justice Marshall’s lan¬
guage is: “it has been said, that they [the States under the Confedera¬
tion] were sovereign, were completely independent, and were con¬
nected with each other only by a league. This is true.”1
Admitting, therefore, that the one theory has in its behalf the au¬
thority of Jay, Hamilton, Story and Kent, the other has the equally
high authority of Marshall, Madison and Webster. We may, there¬
fore, where authorities disagree, proceed to examine the arguments
with perfect freedom from bias. The question is eminently an his¬
torical one — that is, a question of facts, not of theory. Sovereignty
being the supreme power to command, it is simply a question of fact
what organization was found in possession of this power, when it
ceased to be exercised by Great Britain.
It requires no argument to show that before the Revolution the colo¬
nies were absolutely dependent upon Great Britain; whatever powers
of government they severally possessed was in virtue purely of suf¬
ferance or explicit grant, on the part of the mother country. It is
equally clear that the colonies were connected with one another by
no organic bond. There was no government of the united colonies;
each colony had its own government; and if sometimes, for the con¬
venience of administration, two or more colonies were united under
the same royal governor, this was simply an administrative union —
one official managing two independent governments at a time, not a
single government resulting from the fusion or union of two individual
ones. There were thirteen organized communities, standing in a con¬
dition of coequal dependence upon the government of Great Britain.
This tie of dependence was severed by the Declaration of Indepen¬
dence, July 4, 1776, sustained, as this act was, by armed force.
Two points fall here under consideration: first, the power which sev¬
ered the tie; second, the logical effects of the act of severance.
First, the power that performed the act of severance was the Con¬
tinental Congress. But by what authority, and in virtue of what del¬
egation of power did the Continental Congress act? Was the Con¬
gress the organ of the several States, or of the “people at large” (to
use Mr. Marsh’s expression)?* To answer this question, which rests
at the bottom of the argument, we must trace briefly the history of
this Congress.
* Speech on “The Constitution not a Compact,” Works, iii. 454.
f Ogden vs. Gibbons, 9 Wheaton, 187.
* The Nation, No. 23.
26
United States Sovereignty
In the year 1764, upon motion of James Otis, the General Court of
Massachusetts passed a resolution proposing to the other colonies to form
a union for the purpose of resisting the acts of the British government.
This proposition was accepted, first by Virginia, then by the other colo¬
nies. The Congress met the next year (1765), and shortly afterward, as a
result of the spirit thus manifested, the Stamp Act was repealed. The
Second Continental Congress met in 1774, called in a precisely similar
manner. In both cases the members of the Congress were elected by the
several colonies, and in both cases it was only a portion of the colonies —
nine the first time, twelve the second — that were represented. Now so
long as Georgia staid [sic] away, it is clear that not “the people at large
of the United States,” but only the people of twelve colonies, were en¬
gaged in formal acts of resistance. In the assembly thus composed of
delegates from the several colonies, the colonies voted as such; no mea¬
sure was adopted by a majority of votes, as would have been the case if
they had been considered to represent the people at large; a majority of
the colonies must always decide. It was by colonies that the Declaration
of Independence was passed, and in this document the several colonies
are declared to be “free and independent States.”
Let us pause a moment upon this word “State,” which thus makes
its appearance in our political vocabulary. The great convenience of
having a different term to denote the units which compose our fed¬
eral government from that which designates the federal government
itself, has established, in American constitutional law, a fundamental
difference in the meaning of the respective terms. By State we un¬
derstand a political organization inferior to the Nation. But this dis¬
tinction is peculiar to American public law. The two terms are origi¬
nally identical in meaning, or rather in application; being applied in¬
differently to the same object, but from different points of view. A
State is, in public law, a Nation, regarded from the point of view of
its organization; a Nation is a State, regarded from the point of view
of its individuality. We must not, therefore, suppose that when the
colonies, in 1876, declared themselves to be free and independent
States, they attributed to the word State the same inferiority which
we now associate with the word. They understood by it, a sovereign
political organization. That they selected this term, rather than Na¬
tion, is no doubt partly due to its expressing more distinctly the idea
of organization; partly, I am ready to admit, to the feeling that Nation
was a larger term, and that a higher organization, which should em¬
brace all these individuals in one whole, was destined to result. Nay,
we meet the term Nation very early, as applied to the united body.
27
Wisconsin Academy of Sciences, Arts, and Letters
That the Congress considered itself as acting as the organ of the
colonies or States, and not of the people at large, appears manifest
from the language habitually used. On the tenth of May, 1776 Con¬
gress resolved to “recommend” to the “respective assemblies and con¬
ventions of the United Colonies,” to form permanent governments.
August 21, of the same year, it made use of the expression: “All per¬
sons not members of, nor owing allegiance to any of the United States
of America,” — showing that allegiance was regarded as due to the sev¬
eral States. Its constant title for itself was “the United States in Con¬
gress assembled” — a term which plainly recognizes that the United
States, as an organized body, has no existence except in the Congress,
which Congress, as we have seen, acted purely as the organ of the
several States.
I pass now to the nature and effect of the act of severance. This
act was in the first place purely negative in its intrinsic character. It
simply put an end to a certain previously existing relation — that by
which the colonies individually depended upon the British sovereignty.
The relations of the several colonies to one another could not be af¬
fected by it. If before the act they formed a united, organized body,
this united body, in virtue of the act of independence, succeeded to
the sovereignty surrendered by the mother country; if they were indi¬
vidual and disconnected before, they remained so after the act, and
each individual passed into the full enjoyment of sovereignty.
Now I have shown first, that before the revolution the colonies had
no organic connection with one another, but only with the mother
country; second, that the union which they formed for purposes of re¬
sistance professed to be nothing but a voluntary, incomplete and tem¬
porary association, with only limited and temporary aims, possessing
none of the essentials of a permanent government, capable, it is true,
of developing into a complete sovereignty, but in all its acts and words
appearing as not itself an organic body, but the representative of cer¬
tain organic bodies. “The United States in Congress assembled,” made
no claim to individual or independent existence, but acted avowedly
as a mere intermediary or instrument of joint action for organisms
which did possess individual existence. And this practical indepen¬
dence accrued to the several colonies simply from the fact that, upon
the severance of the tie which connected them severally to the mother
country, each was left standing legally alone; and, standing alone, hav¬
ing no legal superior, but possessing a complete and adequate organi¬
zation of its own, each colony passed into the undisputed enjoyment
of sovereignty.
28
United States Sovereignty
Neither before nor after the commencement of the revolution, there¬
fore, did there exist any united organic body which could supersede
the several colonies, and assert a claim to the lapsed sovereignty of
Great Britain. And if this is true for the period of inchoate national¬
ity which intervened between the first acts of resistance and the prac¬
tical establishment of independence, still more is it true for the ensu¬
ing period of the Confederation. It needs no argument to show that
the States were at this time recognized as fully and exclusively sov¬
ereign; its Articles explicitly provide “that each state retains its sov¬
ereignty and power which is not by this Confederation expressly del¬
egated to the United States in Congress assembled.” All that can be
said in opposition to this view is that this was a “palpable usurpation,”
set on foot during this “embryonic or inchoate period”*; and their ar¬
guments plainly imply that they understand the Articles of Confed¬
eration to represent a different phase of national life from the Decla¬
ration of Independence, and as requiring therefore to be construed from
a different point of view; they were adopted by Congress sixteen
months later than the other act, (Nov. 15, 1777.) and in this period of
time, it is hinted, the “flow of enthusiasm,” under which the united
act of independence had been accomplished, “receded,” and selfish
and local prejudices took its place. Now, if the Articles of Confed¬
eration were really drawn up a year and a half after the Declaration
of Independence, this reasoning would have much weight. But the
date here given is only that of the adoption of the articles by Con¬
gress. They were reported to Congress July 12, 1776, just a week af¬
ter the Declaration — the preliminary steps, indeed, were taken in June,
before the passage of the act of independence. It is therefore perfectly
legitimate to interpret the act of independence in the light of the gov¬
ernment which was established after it. The two acts were to all in¬
tents and purposes parts of one and the same act. In the very act of
declaring their independence, the States formed themselves into a Fed¬
eral Union; and in this Union the several States were explicitly de¬
clared to be independent and sovereign; from which it necessarily fol¬
lows that the Union thus formed, was, in Webster’s words, “a league
and nothing but a league.”
It will be seen that the whole controversy turns upon the period
between the suspension of the royal authority and the establishment
of the confederation. While the royal authority continued to be rec-
* Pomeroy, p. 48.
+ Mr. Marsh, in the Nation, No. 1.
29
Wisconsin Academy of Sciences, Arts, and Letters
ognized, sovereignty of course belonged to Great Britain; after the es¬
tablishment of the Confederation, it as manifestly belonged to the sev¬
eral States. Was there an interval during which it was possessed by
the United Colonies? Mr. Marsh says:* “it was not for a moment
imagined that the sovereignty was in the interim lodged anywhere ex¬
cept in the whole people of the United Colonies.” But he brings no
facts to prove this assertion.
At the beginning of this discussion it was remarked that the ques¬
tion was essentially an historical one, and must find its decision in
historical facts — that is, in the series of events by which the sover¬
eignty was transferred from Great Britain to the United States; and I
think I have shown that, as a matter of fact, this transfer was not made
at one stroke, but that the sovereignty was actually possessed for a
while by the several States, before it was transferred by a deliberate
act to the nation. There remain, however, some theoretical objections
to this view, which it will be necessary to consider.
Mr. Pomeroy states these theoretical objections in the following
strong terms: “Grant that in the beginning the several states were, in
any true sense independent sovereignties, and I see no escape from
the extreme positions reached by Mr. Calhoun. ”+ No arguments are
presented in support of this startling assertion, except the doctrine that
among the attributes of sovereignty, “the one which underlies all oth¬
ers, and is, in fact, necessarily implied in the very conception of sepa¬
rate nationality, is that of supreme continued self-existence. This in¬
herent right can only be destroyed by overwhelming opposing force;
it cannot be permanently parted with by any constitution, treaty,
league, or bargain, which shall forever completely resign or essen¬
tially limit their sovereignty, and restrain the people from asserting
it.” There is no attempt made to prove this doctrine; it rests simply
upon Mr. Pomeroy’s assertion, backed by references to the works of
half a dozen European publicsts [sic]. According to this doctrine Texas
was never annexed; if the United States had conquered her, and forced
her into the Union, her status would have been a legal one; but as she
came in voluntarily, surrendering her sovereignty and individual ex¬
istence, the act was null and void. According to this doctrine the act
of union by which, in 1706, England and Scotland surrendered their
individual sovereignty, and united into the new sovereignty of Great
Britain, was an impossible act; and Scotland might now, if she chose,
* The Nation. No. 21.
fp. 39.
30
United States Sovereignty
re-establish her Parliament at Edinburgh, and crown a Presbyterian
King at Scone. Again; on this theory, what are we to do with Rhode
Island and North Carolina in the interval between the establishment
of the Federal Government, and their accession to it? They were cer¬
tainly not members of the new Union; which made no claim to extend
its power over them. The Confederation of which they had been mem¬
bers, no longer existed. There is but one answer to this question. They
were independent, sovereign States, as independent and as sovereign
as Costa Rica, or San Marino, or the Free City of Hamburg.
In arguing for the original sovereignty of the States, I would not
be understood to advocate the modern doctrine of State Rights. I hold
with Marshall, Webster and Story, with Mr. Marsh and Mr. Pomeroy,
that the United States form a nation, and possess full powors [sic] of
sovereignty. But I hold that this sovereignty was formally and vol¬
untarily conferred upon them by the States in the act of forming the
Federal Constitution. The doctrine advanced by Mr. Pomeroy* as to
the relation of the States to the United States, which is essentially that
of Mr. Austin, I fully accept. “The people of the United States, as a
nation, is the ultimate source of all power, both that conferred upon
the General Government, that conferred upon each State as a separate
political society, and that retained by themselves.” Only, by “ulti¬
mate source,” I do not understand historical filiation, but legal author¬
ity, under the constitution; the States — meaning by that the people of
the several States — formed themselves, by this act, into “the People
of the United States;” and this sovereign people, as organized in States,
exercises its sovereign powers by the two-fold instrumentality of the
National Government and the States’ Governments, distributing these
powers between these two instrumentalities as seems most expedient.
Thus the States are as much sovereign as the nation; but in truth nei¬
ther is sovereign, but each is an organization for the exercise of a cer¬
tain definite portion of the powers of government. The sovereignty
is not divided between States and Nation, because sovereignty is in¬
divisible and absolute; but the functions of government, in which con¬
sists the exercise of the powers of sovereignty, can be divided, and
are divided between these two organizations.
* Page 23
31
Wisconsin Academy of Sciences, Arts, and Letters
Thomas Chrowder Chamberlin, Ph.D., Sc.D., LL.D.
(1843-1928)
Wisconsin State Geologist; professor of geology at Beloit
College; charter member and fifth president of the Wisconsin
Academy, 1885-87; president of the University of Wisconsin,
1887-92; professor at the University of Chicago
32
Courtesy The State Historical Society of Wisconsin Whi(x3)33074
Wisconsin Kettle Moraine
DEPARTMENT
OF THE MATHEMATICAL AND PHYSICAL SCIENCES
ON THE EXTENT AND SIGNIFICANCE OF THE
WISCONSIN KETTLE MORAINE
BY T. C. CHAMBERLIN, A M.,
State Geologist, and Professor of Geology in Beloit College1
At the meeting of the Academy, three years since, I took the lib¬
erty of occupying the attention of the members by the presentation of
some observations and conclusions in reference to a peculiar series
of drift hills and ridges in eastern Wisconsin, known as the Kettle
range, and the views then advanced afterwards found a place in my
report on the geology of eastern Wisconsin.2 Similar observations
were subsequently made by Professor Roland D. Irving, of the Wis¬
consin survey, and his conclusions are in perfect agreement with my
own.3
In neither case, however, was any attempt made to show the full
extent of the formation outside of the districts reported upon, or to
point out its theoretical significance, the chapters being intended only
as contributions to local geology, made under somewhat severe limi¬
tations as to space.
It is not now possible to map, or even safely conjecture, the complete
extent and limitations of the formation; but it is the purpose of this ar¬
ticle to add such trustworthy observations as have since been made, and
to gather such evidence as may justify a provisional mapping of the range,
where it has not been actually traced. A portion of the paper will, there¬
fore, relate to well ascertained facts, while other portions will be in vari¬
ous degrees hypothetical. If care is taken to distinguish between these
portions, no harm can arise from their association; while the provisional
’l have taken advantage of the interval between the date of reading and the print¬
ing to introduce new matter. T.C.C.
2Geology of Wis., Vol. II, 1877 (revised edition 1878), pp. 205-215.
3Geology of Wis., Vol. II, 1877 (revised edition 1878), pp. 608-635.
33
Wisconsin Academy of Sciences, Arts, and Letters
34
Wisconsin Kettle Moraine
mapping will, it is hoped, prove of service in both stimulating and guid¬
ing further investigation. The extent of the range is likely to prove too
great for the immediate time and means of a single observer; while the
broad and irregular, and sometimes obscure, character of the belt is such
that it is likely to be overlooked, as a continuous range, as experience
has shown, unless attention be called to it, or the observer be keenly alive
to distinctions in drift topography. It is believed, therefore, that the pre¬
sentation of some things that are only probably, not certain, will not be
without value.
It will be advisable to consider first, somewhat critically, the char¬
acter of the formation. The following description, which is based upon
careful observation, relates more specifically to the moraine in Wis¬
consin, where it is usually well developed, and may require some
modification in its application to the range where sub-aqueous deposits
overlap or encroach upon it, and in other special situations.
Surface Features. — The superficial aspect of the formation is that
of an irregular, intricate series of drift ridges and hills of rapidly, but
often very gracefully, undulating contour, consisting of rounded
domes, conical peaks, winding and, occasionally, geniculated ridges,
short, sharp spurs, mounds, knolls and hummocks, promiscuously ar¬
ranged, accompanied by corresponding depressions that are even more
striking in character. These depressions, which, to casual observa¬
tion, constitute the most peculiar and obtrusive feature of the range,
and give rise to its descriptive name in Wisconsin, are variously known
as “Potash kettles,” “Pot holes,” “Pots and kettles,” “Sinks,” etc.
Those that have most arrested popular attention are circular in out¬
line and symmetrical in form, not unlike the homely utensils that have
given them names. But it is important to observe that the most of
these depressions are not so symmetrical as to merit the application
of these terms. Occasionally, they approach the form of a funnel, or
of an inverted bell, while the shallow ones are mere saucer-like hol¬
lows, and others are rudely oval, oblong, elliptical, or are extended
into trough-like, or even winding hollows, while irregular departures
from all these forms are most common. In depth, these cavities vary
from the merest indentation of the surface to bowls sixty feet or more
deep, while in the irregular forms the descent is not unfrequently one
hundred feet or more. The slope of the sides varies greatly, but in
the deeper ones it very often reaches an angle of 30° or 35° with the
horizon, or, in other words, is about as steep as the material will lie.
In horizontal dimensions, those that are popularly recognized as
“kettles” seldom exceed 500 feet in diameter, but, structurally con-
35
Wisconsin Academy of Sciences, Arts, and Letters
sidered, they cannot be limited to this dimension, and it may be diffi¬
cult to assign definite limits to them. One of the peculiarities of the
range is the large number of small lakes, without inlet or outlet, that
dot its course. Some of these are mere ponds of water at the bottom
of typical kettles, and, from this, they graduate by imperceptible de¬
grees into lakes of two or three miles in diameter. These are simply
kettles on a large scale.
Next to the depressions themselves, the most striking feature of this
singular formation is their counterpart in the form of rounded hills and
hillocks that may, not inaptly, be styled inverted kettles. These give to
the surface an irregularity sometimes fittingly designated “knobby drift.”
The trough-like, winding hollows have their correlatives in sharp serpen¬
tine ridges. The combined effect of these elevations and depressions is
to give to the surface an entirely distinctive character.
These features may be regarded, however, as subordinate elements
of the main range, since these hillocks and hollows are variously dis¬
tributed over its surface. They are usually most abundant upon the
more abrupt face of the range, but occur, in greater or less degree, on
all sides of it, and in various situations. Not unfrequently, they occur
distributed over comparatively level areas, adjacent to the range.
Sometimes the kettles prevail in the valleys, the adjacent ridges be¬
ing free from them; and, again, the reverse is the case, or they are
promiscuously distributed over both. These facts are important in con¬
sidering the question of their origin.
The range itself is of composite character, being made up of a se¬
ries of rudely parallel ridges, that unite, interlock, separate, appear and
disappear in an eccentric and intricate manner. Several of these sub¬
ordinate ridges are often clearly discernible. It is usually between the
component ridges, and occupying depressions, evidently caused by
their divergence, that most of the larger lakes associated with the range
are found. Ridges, running across the trend of the range, as well as
traverse spurs extending out from it, are not uncommon features. The
component ridges are themselves exceedingly irregular in height and
breadth, being often much broken and interrupted. The united effect
of all the foregoing features is to give to the formation a strikingly
irregular and complicated aspect.
This peculiar topography, however, finds a miniature representa¬
tive in the terminal moraines of certain Alpine glaciers. Most of the
glaciers of Switzerland, at present, terminate in narrow valleys, on very
steep slopes, and leave their debris in the form of lateral ridges, or a
torrentially washed valley deposit. A portion of them, however, in
36
Wisconsin Kettle Moraine
their recently advanced state, descended into comparatively open val¬
leys of gentle decline, and left typical, terminal moraines, formed from
the ground moraines of the glaciers, and only slightly obscured by
the medial and lateral morainic products, which have little or no rep¬
resentative in the Quaternary formations. The Rhone glacier has left
three such ridges, separated by a few rods interval, that are strikingly
similar in topographical eccentricities to the formation under discus¬
sion, save in their diminutive size. The two outer ones have been
modified by the action of the elements, and covered by grass and
shrubs, while the inner one remains still largely bare, and, as they have
been cut across by the outflowing glacial streams, they are exceed¬
ingly instructive as to glacial action under these circumstances. The
inner one graduates in an interesting way into the widespread ground
moraine, which occupies the interval between it and the retreating gla¬
cier, where not swept by floods, and which presents a different sur¬
face contour, illustrative of Till topography. The two Grindelwald
glaciers have left similar moraines; those of the upper one, being the
more massive, and being driven closer together, present an almost per¬
fect analogy to the Kettle ranges. The Glacier du Bois, the terminal
portion of the Mer de Glace, the Argentiere, and, less obviously, the
Findelen, and others, so far as their situation favored, have developed
similar moraines, and indicate that this is the usual method of deposit
under these conditions. Reference is here made only to the terminal
deposit of the ground moraine , eliminating, as it is quite possible to
do, for the most part, the material borne on the surface of the glacier.
The Material of the Formation . — This topic, which is one of pri¬
mary importance in determining the origin of the deposit, readily di¬
vides itself into three subordinate ones, all of which need discrimina¬
tive attention; (1) the form of the constituents, (2) their arrangement
as deposited, and (3) their source.
(1) Premising that the Karnes, and those deposits which have been as¬
sociated with them in the literature of the subject, are described as com¬
posed mainly of sand and gravel, it is to be remarked, in distinction, that
all the four forms of material common to drift, viz.: clay, sand, gravel,
and boulders, enter largely into the constitution of the Kettle range, in its
typical development. Of these, gravel is the most conspicuous element,
exposed to observation. This qualification is an important one in form¬
ing an adequate conception of the true structure of the formation. It is to
be noticed that the belt, at many points, exhibits two distinct formations.
The uppermost — but not occupying the heights of the range — consists al¬
most wholly of sand and gravel, and lies, like an irregular, undulating
37
Wisconsin Academy of Sciences, Arts, and Letters
sheet, over portions of the true original deposit. This superficial forma¬
tion is confined mainly to the slopes and flanks of the range, and to de¬
pressed areas between its constituent ridges; though, when the whole belt
is low, it often spreads extensively over it, so as sometimes to be quite
deceptive. But, where the range is developed in force, this superficial
deposit is so limited and interrupted, as to be quite insignificant, and not
at all misleading; and, at some points, where it is more widely developed,
excavations reveal unequivocally its relationship to the subjacent accu¬
mulations. In such cases, the lower formation shows a more uneven sur¬
face than the upper one, indicating that the effect of the latter is to mask
the irregular contour of the lower and main formation. Notwithstanding
this, the upper sands and gravels are often undulatory, and even strongly
billowy, and the bowls and basins in it commonly have more than usual
symmetry. A not uncommon arrangement of this stratum is found in an
undulating margin on the flank of a ridge of the main formation, from
which it stretches away into a sand flat or a gravel plain.
Setting aside this, which is manifestly a secondary formation, it is
still true that gravel forms a large constituent of the formation. Some
of the minor knolls and ridges are almost wholly composed of sand
and gravel, the elements of which are usually very irregular in size,
frequently including many boulders. But, notwithstanding these quali¬
fications, the great core of the range , as shown by the deeper excava¬
tions, and by the prominent hills and ridges, that have not been masked
by superficial modifications, consists of a confused commingling of
clay, sand, gravel, and boulders, of the most pronounced type. There
is every gradation of material, from boulders several feet in diameter,
down to the finest rock flour. The erratics present all degrees of an¬
gularity, from those that are scarcely abraded at all, to thoroughly
rounded boulders. The cobble stones are spherically rounded, rather
than flat, as is common with beach gravel, where the attrition is pro¬
duced largely by sliding, rather than rolling.
Stratification. — As indicated above, the heart of the range is es¬
sentially unstratified. There is, however, much stratified material in¬
timately associated with it, a part of which, if my discriminations are
correct, was formed simultaneously with the production of the
unstratified portion, and the rest is due to subsequent modification.
The local overlying beds, previously mentioned, are obviously strati¬
fied, the bedding lines being often inclined, rather than horizontal, and
frequently discordant, undulatory or irregular.
The Source of the Material. — This, so far as the range in Wis¬
consin is concerned, admits of the most unequivocal demonstration.
38
Wisconsin Kettle Moraine
The large amount of coarse rock present renders identification easy,
and the average abrasion that has been suffered indicates, measurably,
the relative distance that has been traveled. The range winds over
the rock formations in a peculiar manner, so as to furnish fine oppor¬
tunities for decisive investigation. Of the many details collected, there
is room here for a single illustrative case only. The Green Bay loop
of the range surrounds on all sides, save the north, several scattered
knobs of quartzite, porphyry and granite, that protrude through the pre¬
vailing limestones and sandstones of the region. These make their
several contributions to the material of the range, but only to a lim¬
ited section of it, and that invariably in the direction of glacial stria-
tion. Any given segment of the range shows a notable proportion of
material derived from formation adjacent to it, in the direction of stria-
tion; and a less proportion, generally speaking, from the succeeding
formations that lie beyond it, backward along the line of glacial move¬
ment for three hundred miles or more. It is undeniable, that the
agency, which produced the range, gathered its material all along its
course for at least three hundred miles to the northward, and its larg¬
est accumulations were in the immediate vicinity of the deposit. For
this reason, as the range is traced along its course, its material is found
to change, both lithologically and physically, corresponding to the for¬
mation from which it was derived.
These facts find ample parallel in the moraines of Switzerland. The
marginal portion of the great moraine of the ancient expanded gla¬
ciers, on the flanks of the Juras, is composed, very largely, of boul¬
der clay, derived from the limestones that lie in its vicinity, while the
quantity of material derived from the more distant formations of the
Alps is quite subordinate. Of the more recently formed moraines,
those derived from the Bois, Viesch, Rhone, Aar, and other glaciers,
which pass over granitic rocks, consist quite largely of sand, gravel,
and boulders, clay being subordinate, while those glaciers of the
Zermatt region, that pass mainly over schistose rocks, and the
Grindelwald glaciers, that, in the lower part of their course, traverse
limestone, give rise to a decided amount of clay. The moraines, pre¬
viously referred to as miniature kettle ridges, are composed of com¬
mingled unstratified debris, in the main, but there are instances of as¬
sorted and stratified material. The inner moraine of the upper
Grindelwald glacier presents much fine assorted gravel and coarse
sand, heaped up, very curiously, into peaks and ridges, in various at¬
titudes on the summit and sides of the moraine.
Relations to Drift Movements. — This is manifestly of most vital
39
Wisconsin Academy of Sciences, Arts, and Letters
consideration. The course of drift movement may be determined, (1)
by the grooving of the rock surface, (2) by the direction in which the
material has been transported, (3) by the abrasion which rock promi¬
nence have suffered, (4) by the trend of elongated domes of polished
rock, and, (5) less decisively, by the arrangement of the deposited
material and the resulting topography. Recourse has been had to all
these means of determination, in that portion of the range that has been
carefully investigated, and their individual testimony is entirely har¬
monious, and their combined force is overwhelming. Exceptional op¬
portunity for positive determination is afforded by the protruding
knobs of Archaean rocks before alluded to, from which trains of er¬
ratics stretch away in definite lines, continuous with the striation on
the parent knobs, and parallel to that of the region, as well as concor¬
dant with the general system. The united import of all observations,
in eastern Wisconsin, testifies to the following remarkable movements,
which may be taken as typical, and which are here given, because they
have been determined with much care. Between Lake Michigan and
the adjacent Kettle range, the direction was obliquely up the slope, as
now situated, southwestward, towards the range. On the opposite side,
between the Green Bay valley and the range, the course was, after sur¬
mounting the cliff bordering the valley, obliquely down the slope,
southeastward, toward the range. In the Green Bay trough, the ice
stream moved up the valley to its watershed, and then descended
divergingly the Rock River valley. Between the Green Bay valley
and the Kettle belt on the west, the course was up the slope, west¬
ward, or southwestward, according to position. These movements,
which are imperfectly shown on the diagram, exhibit a remarkable di¬
vergence from the main channel toward the margin of the striated area,
marked by the Kettle range.
Much of the data relating to the movements, outside of Wisconsin,
has been derived from a study of publications relating to the geology
of the several states, to whose authors I am indebted, but who should
not be held responsible for the special collocation presented in the ac¬
companying diagram, which, in some of its details, may prudently be
held as somewhat tentative, until more rigorously verified. But the
grand features of these movements, which may be confidently ac¬
cepted, are very striking, and are very singularly related to the great
basins of the lake region. The three main channels were the troughs
of the great lakes, Superior, Michigan, and the couplet, Erie and
Ontario, while between these lay three subordinate ones in the basins
of the great bays, Saginaw, Green and Keweenaw.
40
Wisconsin Kettle Moraine
41
Wisconsin Academy of Sciences, Arts, and Letters
The divergence of the striations from the main channels toward the
range, in the case of the Green Bay valley, and, so far as the evidence
goes, in other troughs, was an unexpected result, developed by com¬
bining individual observations; but, when the method of wasting and
disappearance of a glacier is studiously considered, appears not only
intelligible, but a necessary result, and one which finds partial illus¬
tration among existing glaciers.
Topographical Relations and Distribution. — The topographical
relations of the formation are an essential consideration, but may be
best apprehended in connection with its geographical extension, which
now claims our attention. If we start with the northern extremity of
the long known Potash Kettle Range, in Wisconsin, we find ourselves
about midway between the southern extremity of Green Bay and Lake
Michigan, and on an eastward sloping, rocky incline. The base of the
range is here less than 200 feet above Lake Michigan, and is flanked
on either side by the lacustrine red clays of the region; and seems, in
some measure, to be obscured by them. From this point, it stretches
away in a general south-southwestward direction, for about 135 miles,
ascending gradually, and obliquely, the rocky slope, until it rests di¬
rectly on its crest.
When within about twenty miles of the Illinois line, it divides , one
portion passing southward into that state, and the other, which we will
follow, curves to the westward, and crosses the Rock river valley. A
profile of the rock surface across this valley, beneath the range, would
show a downward curve of more than 300 feet. The range should not,
perhaps, be regarded as sagging more than half that amount, however,
in crossing the valley, as the canon-like channel of the pre-glacial river,
seems to have been filled without much affecting the surface contour
of the drift. But the fact of undulation to conform to an irregular sur¬
face, produced by erosion, and not by flexure of the strata, is a point
to be noted, as it is a serious obstacle in the way of any explanation
that is only applicable on the supposition that the formation was in a
horizontal position when formed, as the view that it was produced by
beach action, or the stranding of icebergs.
After crossing Rock river, the range curves gradually to the northward,
passing over the watershed between the Rock and Wisconsin rivers, “de¬
scends abruptly 200 feet into the low ground of the valley of the Wis¬
consin,”1 crosses the great bend of the river, sweeping directly over the
quartzite ranges, according to Prof. Irving, with a vertical undulation of
1 Prof. Irving, Geol. of Wis., Vol. II, 1877, page 616.
42
Wisconsin Kettle Moraine
over 700 feet, after which it gradually ascends the watershed between
the Mississippi and St. Lawrence drainage systems, until its base reaches
an estimated elevation of 700 to 800 feet above Lake Michigan. From
thence it has been traced across the headwaters of the Wisconsin river,
by Mr. A. Clark, under my direction.1
Within the Chippewa valley, it has been observed by Prof. F. H.
King, of the Wisconsin Survey, and I have observed it in the vicinity
of the Wisconsin Central railroad. This region is covered by an im¬
mense forest, mainly unsettled and untraversed, even by foot paths,
so that geological exploration is difficult and expensive, and, as no
industrial importance attaches to it, and the rock below is deeply con¬
cealed by it, I have not deemed it sufficiently important to trace the
belt continuously to justify the large expenditure of time and means
requisite, especially as I entertain no serious doubts as to its continu¬
ity and general position. The observations made, indicate that it de¬
scends obliquely the eastern slope of the Chippewa valley, and crosses
the river below the great bend (T 32, R. 6 and 7), near which the Flam¬
beau, Jump, and several smaller streams gather themselves together,
in a manner very similar to that of the branches of the Rock and Up¬
per Wisconsin rivers, just above the point where they are crossed by
the range. From this point the belt appears to curve rapidly to the
northward, forming the western watershed of the Chippewa. It is
joined in eastern Burnett county by a portion of the range coming up
from the southwest, the two uniting to form a common range, analo¬
gous to that of eastern Wisconsin. The conjoint range thus formed,
extends along the watershed of the Chippewa and Nemakagon rivers,
to the vicinity of Long and Nemakagon lakes, on the watershed of Lake
Superior. This part is given mainly on the authority of Mr. D. A.
Caneday, who visited a portion of the formation with me, and whose
discrimination can, I think, be trusted. Mr. E. T. Sweet, of the Wis¬
consin Survey, describes2 a kettle range as lying along the axis of the
1 To the eastward of the range, as thus traced. Col. Whittlesey describes
(Smithsonian Contributions, 1866) a similar formation in Oconto County. I have ob¬
served the same at several points. Mr. E. E. Breed informs me that it occurs on the
watershed between the Wolf and Oconto rivers, but it has not yet been traced through
the wilderness, to any connection with the main range, and it is uncertain whether it
is so connected or constitutes a later formation, as such later moraines have been ob¬
served at other points.
2 Manuscript report on Douglas and Bayfield counties, to form a part of Vol. Ill,
Geol. of Wis.
43
Wisconsin Academy of Sciences, Arts, and Letters
Bayfield peninsula, but it has not been ascertained that this is con¬
nected with the belt under consideration.
Returning to the junction of the two ranges in eastern Burnett
county, I have traced the belt thence southwestward through Polk and
St. Croix counties to St. Croix lake, on the boundary of the state. The
lower portion of this has also been studied by Prof. L. C. Wooster, of
the Wisconsin Survey. The southeastern range of the belt may be con¬
veniently seen on the North Wisconsin railroad, near Deer Park, and
on the Chicago, St. Paul & Minneapolis line, to the west of the sta¬
tion Turner, but only in moderate force.
If a good surface map of Minnesota be consulted, it will be seen
that there lies along the watershed, between the Upper Mississippi and
the conjoint valleys of the Minnesota and Red rivers, a remarkable
curving belt of small lakes. Along this line, lies a chain of drift hills,
known in its northwestern extension as the Leaf hills. In the Sixth
Annual Report of the Geological Survey of Minnesota, received just
as this article is going to the printer, Prof. N. H. Winchell, speaking
of the great moraines of the northwest, says: “There are two such that
cross Minnesota, the older being the Coteau and the younger, the Leaf
hills. Corresponding to the latter, the Kettle Range in Wisconsin seems
a parallel phenomenon.”1 I have seen this belt, west of Minneapolis,
and concur in Prof. Winchell’s opinion. I have also observed, hast¬
ily, what I regard as portions of it — dissevered by the river channels —
on the peninsula formed by the bend of the Mississippi and the Min¬
nesota, south of St. Paul, and on the similar peninsula between the
Mississippi and Lake St. Croix; and this seems to be the line of con¬
nection between the Wisconsin and Minnesota ranges. It appears to
me, therefore, well nigh certain, that the Leaf hills of Minnesota are
not only analogous to the Wisconsin Kettle range, but are portions of
the same linear formation.
The multitude of small lakes, found in Wisconsin, lie almost ex¬
clusively either along the Kettle belt itself, or in the area within, or
north of it. The surface outside has a much more perfect system of
drainage, and is almost entirely free from lakelets. The Kettle range
constitutes the margin of the lake district. But in Minnesota, south of
the Leaf hills, there is an extensive lake region stretching southward
in a broad tongue, nearly to the center of Iowa, though the lakes are
1 Sixth Annual Rept. Geol. & Nat. Hist. Sur. Minn., p. 106. The R. R. profiles
crossing this belt furnish valuable data. See Ann. Rept. for 1872, pp. 53 and 57, and
Sixth Ann. Rept., pp. 47 and 156.
44
Wisconsin Kettle Moraine
not very numerous in the latter state. The question naturally arises,
whether this lake district is likewise bordered by similar drift accu¬
mulations, and this question, though not essential to the present dis¬
cussion, has much interest in connection with it. In respect to this, I
can only give some detached observations and quotations. As already
stated, accumulations of this character occur south of St. Paul. Still
further to the southward, in the town of Aurora, Steel county, there is
a moderate exhibition of gravelly boulder-bearing hillocks and ridges,
accompanied by shallow basins and irregular marshes, much after the
manner of the formation in question. From the descriptions of Prof.
Harrington,1 these features appear to characterize the county some¬
what widely, especially in the southern part. Near Albert Lea, in the
adjoining county, on the south, and only a few miles from the Iowa
line, there is a more prominent development of similar features, the
ridges having a southwestward trend. Dr. C. A. White, in the Geol¬
ogy of Iowa, describes a terrace in the northern part of the state, which,
in its eastern extension, “becomes broken up into a well marked strip
of ‘knobby country.’ Here it consists of elevated knobs and short
ridges, wholly composed of drift, and usually containing more than
an average proportion of gravel and boulders. Interspersed among
these knobs and ridges, are many of the peat marshes of the region.”2
One knob he estimates as rising 300 feet above the stream at its base.
This area lies in the line of the preceding localities, and near the Min¬
nesota border. Between the “knobby country” and the Algoma branch
of the C., M. & St. P. R. R., and stretching southwestward from the
latter, there is a broad belt of low mounds and ridges, some of which
show the structure and composition common to the Kettle moraine,
while others present externally only a pebble clay, similar to that which
characterizes the level country to the west of it. The whole presents
the appearance of a low range modified by lacustrine deposits.
Near the center of the state, Dr. White describes a second range under
the name of “Mineral Ridge,”3 as consisting, “to a considerable extent,
of a collection of slightly raised ridges and knolls, sometimes interspersed
with small, shallow ponds, the whole having an elevation, probably, no¬
where exceeding 50 feet above the general surface, but, being in an open
prairie region, it attracts attention at a considerable distance.” Both of
these ridges, Dr. White classes as probable moraines.
1 Geol. and Nat. Hist. Sur. Minn., Ann. Rept. 1875, pp. 108 et seq.
2 Geol. of Iowa, 1870, p. 99.
3 Loc. cit.
45
Wisconsin Academy of Sciences, Arts, and Letters
This Mineral ridge lies south of the lake district, and may be re¬
garded as forming its margin in that direction. On the western bor¬
der, Dr. White describes “knobby drift,” in Dickinson county, which,
however, is “without perceptible order or system of arrangement.”1
To the northwest from this, we soon encounter the morainic accumu¬
lations of the “Coteac de Prairie,”2 and the “Cobble Knolls” and “An¬
telope Hills.”
These observations do not indicate a continuous, well defined range,
but seem rather to point to a half-buried moraine, that only here and
there, along its course, protrudes conspicuously, and this is the im¬
pression gained from an inspection of the formation. It is to be noted,
as supporting this view, that, at least so far as the eastern side is con¬
cerned, this supposed moraine is flanked on the exterior by level
plains, of smooth surface, often underlaid by sand and gravel, that
seemingly owe their origin to broad rivers or lakes that fringed the
border of the glacier, in its advanced state, when it probably discharged
its waters over the moraine at numerous points, rather than at one, or
a few, selected points, as would more likely be the case during its re¬
treat, when accumulations of water could gather along its foot, within
the moraine, and large areas be discharged at some single favorable
point. But on the inner side of the moraine, the surface, although
nearly level, in its general aspect, undulates in minor swells and sags,
and the drainage is imperfect. The substratum, instead of being gravel,
sand, or laminated clay, is generally a pebble or boulder clay. Out¬
side of the moraine, the existing surface contour was formed in the
presence, and, to some extent, under the modifying influence, of a
fairly established drainage system. But on the interior , the drainage
system has not, even yet, become fully established, much less im¬
pressed itself upon the surface configuration, except in the vicinity of
the main rivers.
The terrace-like ridge mentioned by Dr. White, and some of the
lines of hills described by Prof. Winchell in Minnesota, as running in
a similar direction, may be perhaps regarded as minor morainic lines,
stretching across the glacial pathway and marking oscillations in its
retreat, analogous to some quite clearly made out in Wisconsin.3
1 Geol. of Iowa, Vol. II, p. 221.
2 See note of Prof. Mather, Nat. Hist. Sur. 1st Dist., N.Y., p. 193. See also 2d
Annual Report Geol. and Nat. His. Sur. Minnesota, by N. H. Winchell, pp. 193 to 195;
also loc cit., ante.
3 Geol. of Wis., Col. II, 1876, p. 215 et seq.
46
Wisconsin Kettle Moraine
This southern morainic loop is, of course, presumed to be older than
the Kettle range, and is here discussed because of the interesting way
in which it is associated with the latter formation, and the suggestions
it may contribute to the final solution of the main problem, to which
the special one under discussion is only a tributary, viz.: the definite
history of the Quatenary formations.
Returning to the branching of the range in southeastern Wiscon¬
sin, we find the left arm, or that nearest Lake Michigan, striking south¬
ward into Illinois. If we lay before us Prof. Worthen’s geological map
of that state, and attentively observe its topographical features and its
drainage systems, it will be observed that nearly all the lakelets, the
greater part of the marshes, and most of the region of abnormal drain¬
age may be included in a curving line, rudely concentric with the shore
of Lake Michigan, starting near the center of McHenry county, on the
Wisconsin line, and ending in Vermillion county, on the Indiana bor¬
der. It may also be observed, on a similar inspection of Indiana, that
nearly all the lake district lies north of the Wabash.
In Wisconsin, as already stated, we have found this area bordered
by the Kettle range, which is itself notably lake-bearing. The range
continues to sustain this relationship in Illinois, so far as I know it to
be directly continuous. It exhibits a progressive broadening, and flat¬
tening, as it enters upon the level county that encompasses the head
of Lake Michigan. The pebble clay deposit — not coarse boulder
clay — that characterizes the flat country, and which, to the north, has
been separated from the range by a belt of coarse boulder clay, here
approaches, and appears, to some extent, to overlap the range, and to
be one cause of its less conspicuous character. From what I have seen
of the region south of Lake Michigan, and from all I can find in geo¬
logical reports relating to the region, I gather that the range, so far as
it escaped the destructive action of the floods issuing from the Lake
Michigan basin, both while occupied by ice, and subsequently, is, to
a large extent, buried beneath later deposits, or so modified as to be
inconspicuous. Whatever the correct interpretation, it remains a fact
beyond question, that the belt becomes very obscure, compared with
its development to the northward. Dr. E. Andrews says: “As we trace
it southward, the material becomes finer, and the hills lower, until they
shade off imperceptibly into the drift clay, of the Illinois prairies.”1
The members of the geological corps of Illinois did not recognize it
distinctively, in the sense in which it is now considered, but Dr. Ban-
1On Western Boulder Drift, Am. Jour. Sci., Sept. 1869, p. 176.
47
Wisconsin Academy of Sciences, Arts, and Letters
nister, in his report on Lake county, says: “In the western part of the
county, near the Fox river, we find the ridges, in some places, to be
largely composed of rolled limestone boulders. The same character
has been observed further south along the same stream and remarked
upon in the chapter on Cook county.”1 In respect to McHenry county
he says: “In the vicinity of the Fox river, the same kind of gravel ridges
are met with as those which have been described as occurring in the
western part of Lake county.”2 This lies in the belt identified by me,
from personal observation, as belonging to the Kettle range.
Concerning the district farther south, he says: “Boulders of gran¬
ite, quartzite, greenstone, and various other rocks are abundant in vari¬
ous localities on the surface of the ground, and are frequently met with
in excavations for wells, etc., and large deposits of rolled boulders,
chiefly of limestone from the underlying Niagara beds, similar to those
already described in the report on Cook county, occur in the drift de¬
posits of the adjoining portions of Kane and Du Page counties.”3 Con¬
cerning the topography, the same writer says: “Along some of the prin¬
cipal streams, and especially the Fox river in Kane county, the coun¬
try is more roughly broken, and can, in some parts, even be called
hilly, although the more abrupt elevations seldom exceed eighty or
one hundred feet above their immediate base.”4 This broken country,
if we may judge from what is true of the rough country along the same
river to the north of this, is not due so much to the drainage erosion
of the river as to the original deposition of the drift. The same fea¬
tures are said to continue into Kendall county, next south, which brings
us to the vicinity of the ancient outlet of Lake Michigan, where, of
course, the moraine is locally swept away. Still farther south, in
Livingston county, Mr. H. C. Freeman mentions a ridge running south¬
easterly from a point in La Salle county, to near Chatsworth, a dis¬
tance of about forty miles. “This is gravelly and sandy, giving it a
distinctive character as compared with the adjacent prairie.”5 This is
quite too meager to base an identification upon, but I have thought it
worthy of quotation here. At Odell, which lies near this ridge, the
drift is said to be 350 feet deep.6
1 Geol. Sur. of Ill., Vol.I V, p. 130.
2 Loc. cit., p. 131 .
3 Geol. Surv. of Ill., Part IV, p. 1 13.
4 Geol. Surv. of Ill., Part IV, p. 113.
5 Geol. Surv. of Ill.,Vol. IV, p. 227.
6 Geol. Surv. of Ill., Vol. VI, p. 237.
48
Wisconsin Kettle Moraine
On the railroad line from Chicago to Kankakee, there is no recog¬
nizable indication of the formation under consideration. Southwest-
ward from Kankakee, on the line to La Fayette, Ind., there are a few
mounds and ridges that bear a somewhat morainic aspect, but they
are isolated in a generally level tract of lacustrine, rather than glacial,
topography. They are, perhaps, remnants of a formation that has been
largely eroded or buried. Near Fowler, in Benton county, Indiana,
there is a belt of low mounds and ridges, accompanied by shallow de¬
pressions, that quite closely resemble the Kettle range in its more
modified phases. Boulders appear upon the surface, and, in the more
immediate vicinity of the village, are large and numerous. This is
probably a portion of the “stream of boulders two miles wide,” which
Mr. F. H. Bradley mentions as extending through the eastern part of
Iroquois county, Illinois, and the central part of Benton county, Indi¬
ana,1 and which he attributes to floating ice. He does not, however,
mention the associated topography or underlying drift formation.
South of this low range, the country again becomes level, or gently
undulating, as far as the Wabash.
The Indiana geologists have not yet critically examined the heavy
drift region in the northern part of the state, through which the mo¬
raine might be supposed to pass, but in such preliminary inspection
as has been made, they have not recognized any prominent moraine¬
like accumulation. The superficial expression of the region is quite
monotonous, and presents to view deposits of sand, gravel, lacustrine
or pebble clays, but more rarely the coarse boulder clay or mixed ma¬
terial, that I regard as the unmodified ground moraine. The modify¬
ing agencies which produced this phase of the deposits, would be an¬
tagonistic to ridge-like morainic accumulations, and their presence,
in sharp outline, is not to be expected. In the vicinity of Ligonier, in
Noble county, there is a feeble, but somewhat characteristic develop¬
ment of some of the features of the formation. So also, in the vicin¬
ity of Rome and La Grange to the northeast. Between La Port and
Otis there is a kindred, though somewhat peculiar formation, but I am
in doubt as to its true character.
On entering Michigan, we find the formation more unequivocally
developed. Just north of Sturgis, which is near the southern line of
the state, the formation appears in marked development. It does not
attain a great altitude, but presents the peculiar strongly undulating
and hummocky contour, and the coarse, mingled material, character-
1 Geol. Surv. of Ill., Vol. VI, p. 236.
49
Wisconsin Academy of Sciences, Arts, and Letters
istic of the deposit. It may be seen to advantage on the line of the
Grand Rapids & Indiana R. R. To the northeast in the vicinity of
Albion, it may be seen from Springport on the north, to Condit on the
south. It is here broad and flat, and superficially composed of gravel,
for the greater part, but some of the deeper excavations reveal the char¬
acteristic coarser material. On the Michigan Central R. R., the for¬
mation may be observed between Jackson and Dexter, the most promi¬
nent portion being between the stations Francisco and Chelsea. It is
not very prominent on the immediate line of the road, which was
doubtless selected to avoid it, but in the vicinity it rises into promi¬
nent hills and ridges. Some of these, on the north, are conspicuous
objects at considerable distances. Still farther to the northeast, my
friend, Dr. D. F. Boughton, whose identifications I have elsewhere
verified, informs me that the range is well developed in Oakland
county, and is finely exhibited near the line of the Flint & Pere
Marquette R. R., between Plymouth and Holly. Still farther to the
northeast, it may be seen at great convenience and advantage, along
the Detroit & Milwaukee R. R. from Birmingham, below Pontiac, to
Holly. On the flanks, its features are subdued, the hills and ridges
being rather low, with more or less level surface between them, and
the superficial sands and gravels are prevalent; but from Waterford
to beyond Clarkston, the range has a fine, though irregular develop¬
ment. The hills rise with characteristic contours, to an estimated alti¬
tude of 200 feet or more above the surface of the beautiful lakelets
embosomed at their base. The deep cuts near the latter station, am¬
ply exhibit the coarse, commingled material, characteristic of the core
of the range.
Putting the foregoing observations together, they seem to estab¬
lish beyond reasonable doubt the existence of a broad, massive belt
stretching northeastward on the highland between the Saginaw and
Erie basins.
If we return again to the southwestern part of the state, we are in¬
formed by Dr. Boughton that we shall find a similar accumulation at,
and in the vicinity of, Kalamazoo. To the north-northeast, in Barry
county, the Thorn Apple river cuts across this range between Sheridan
and Middleville. This belt here, though broad, presents a more promi¬
nent and ridge-like aspect, with better defined limits than elsewhere
observed in Michigan. To the north of this, opposite Saginaw bay,
there occurs, near Farwell, broken, rough country and abundant coarse
drift, that probably belongs to the belt in question, but my opportu¬
nity for observation was unsatisfactory. Beyond this point, I have no
50
Wisconsin Kettle Moraine
definite information, but I deem it highly probable that the moraine
will be found extending some distance farther, on the highlands of the
Peninsula.
The lake survey charts show that Grand Traverse bay has the re¬
markable depth of over 600 feet. This great depth, together with its
linear character, and the form and arrangement of the associated in¬
lets and lakes, has suggested that it may have been the channel of a
separate minor glacier, analogous to that of Green Bay on the oppo¬
site side of the great lake, but I have no direct evidence that such was
the fact.
In the reports of the geological survey of Ohio, a formation of nearly,
or quite, identical characteristics is carefully described by the several writ¬
ers whose districts embraced it. In the second volume,1 Dr. Newberry
gives, under the name of “Karnes,” an excellent summary of its leading
features. These harmonize very nearly with those of the Kettle belt. The
main points of difference are the less conspicuous character and massive¬
ness of the Ohio range, and the greater prevalence of assorted and strati¬
fied material; in other words, its features are the same that the Kettle range
presents in its more subdued aspects, especially where it is formed in a
comparatively smooth country, and is flanked by pebble clays, with level
surface, instead of coarse boulder clay, with ridged, or mammillary, con¬
tour. I cannot turn aside, here, to define, with sufficient circumspection,
the distinction between these clays, further than to indicate my belief that
the former are sub-aqueous, and the latter sub-aerial, or, if you please,
sub-glacial, deposits.2
Where I have seen the Ohio formation, it presents almost precisely
the characteristics that are exhibited by the Kettle range in northern
Illinois, where it is similarly related to plane topography and pebble
clays, and it is also very similar to the same formation opposite Green
Bay, where it is bordered on both sides by red lacustrine clays of later
date. Dr. Newberry quite clearly recognizes the parallelism, but per¬
haps not the identity, of the formations.3 Col. C. Whittlesey, in his
1 Pages 41-47. See also “Surface Geology of Northwestern Ohio,” Proc. Am.
Assoc. Ad. Sci. 1872, by Prof. N. H. Winchell, under heads of St.Johns and Wabash
Ridges.
2 I have mapped these formations separately in eastern Wisconsin. See Atlas ac¬
companying Vol. II, Geol. of Wis. 1877, Plate III, Map of Quaternary formations. See
also, p. 225 of the volume.
3 Geol. Surv. of Ohio, Vol. II, pp. 4, 5, and 453. Dr. Newberry’s views as to the
origin of the Ohio “Kame” belt are at variance with those here presented.
51
Wisconsin Academy of Sciences, Arts, and Letters
article on the “Fresh Water Glacial Drift of the Northwestern States,”1
classes the formations together as identical in character, though he does
not seem to have considered them members of a continuous forma¬
tion, and could not well do so with the prevalent view, which he some¬
what emphasizes, that it is peculiarly a summit formation. It very of¬
ten does occupy the summit of a rock terrane, and it sometimes forms
a watershed by its own massiveness, but it likewise occupies slopes
and crosses valleys, as shown in detail in the Wisconsin report. Prof.
Andrews of the Ohio survey, in a personal communication, adds his
conviction that the Ohio and Wisconsin deposits are parallel forma¬
tions. It would seem, then, that the only question relates to the conti¬
nuity of the belts. Unfortunately there intervenes the Wabash valley,
the ancient drainage channel of the Erie basin. Absolute continuity
undoubtedly does not exist. If my views are correct, this was the
great — not exclusive — channel of discharge of the glacial floods, at
the very time the moraine was being formed, where it could be formed,
and, for that reason, the debris was swept away or leveled. In addi¬
tion to this, the region has been subjected to the vicissitudes of ero¬
sion, of a reversal of drainage systems, and of lacustrine and fluvia-
tile accumulation. It is to be presumed, therefore, that a portion of
the range, where once formed, has been lost, leveled, or buried. Some
remnant indications of the range, on the upper slopes, might, how¬
ever, rationally be presumed to exist. But, awaiting a critical exami¬
nation of the region, we must confess a want of direct evidence. The
belt stretches entirely across Ohio and enters Indiana, but has not been
traced farther.
In the line of indirect testimony, however, some facts may be no¬
ticed. Prof. N. H. Winchell describes in the Ohio reports2 six ridges
running parallel to Lake Erie, and Mr. G. K. Gilbert has described
that portion of these which lie in the more immediate Maumee val¬
ley.3 Two of the inner ones are conceded to be lake beaches. The
two outer ones are members of the “Kame,” or Kettle belt, according
to Dr. Newberry.4 The one next within, the St. Mary’s ridge, Prof.
Newberry distinguishes, apparently, with justness, from both the other
classes. Mr. Gilbert gives a clear and discriminating description of
this, and expresses the conviction that it is “the superficial represen-
1 Smithsonian Contributions, 1866.
2 See also Proc. Am. Assoc. Ad. Sci., 1872.
3 Geol. Sur. Ohio, Vol. II, pp. 56 and 57.
4Geol. Surv. Ohio, Vol. I, pp. 537 et seq.
52
Wisconsin Kettle Moraine
tation of a terminal glacial moraine, that rests directly on the rock bed
and is covered by a heavy sheet of Erie clay, a subsequent aqueous
and iceberg deposit.”1 The views of Professors Newberry and
Winchell, while they each differ somewhat, agree with this in the only
point essential to the present discussion, viz: that this ridge represents
the margin of the glacier at the time it was formed. This shows the
glacier to have been a tongue or lobe of ice, differentiated from the
supposed continental glacier, and having its axis coincident with the
Maumee valley, and, withal, capable of forming a morainic accumu¬
lation on both sides. The St. Mary’s ridge crosses the Maumee-
Wabash valley — the glacial trough — and, recurving upon itself, bears
away to the northeast, approximately parallel to the Kettle belt already
described in southeastern Michigan. This wing of the St. Mary’s ridge
bears the same relation to the Kettle belt bordering the Erie basin on
the Michigan side, that the opposite wing does to the “Kame” belt on
the south side. The force of this relationship is not easily escaped.
If my views are correct, that this Michigan belt was formed along
the right hand margin of the Erie glacier (conjointly with the Saginaw
glacier), just as the “Kame” belt was formed on the left hand margin,
then its composition should give evidence of the fact. In the case of
the Green Bay glacier, I have shown that the lines of striation and
transportation diverge from the main axis toward the margin,2 and, so
far as the paths of other glaciers lie within Wisconsin, the observa¬
tions made upon them, imply the same method of movement, and this
habit finds partial exemplification among the glaciers of the Alps —
partial, because their contracted valleys and steep slopes afford little
opportunity to deploy in this fashion. If this manner of movement
holds true with the Erie glacier, material from its trough will be found
to have been transported westward and northwestward toward the mo¬
raine. Thirteen years ago, in an article in the American Journal of
Science, entitled, “Some Indications of a Northward Transportation
of Drift Material in the Lower Peninsular of Michigan,”3 Professor
Alexander Winchell called attention, with much detail and precision,
to a large mass of evidence, which finds, for the first time, so far as I
am aware, satisfactory explanation in the view now presented, and,
in return, has the force of confirmatory evidence. It appears that im¬
mense, and often but slightly eroded masses of Corniferous limestone,
1 Loc. cit.
2 Geol. of Wis., Vol. II, pp. 199 et seq.
3 Am. Jour, of Sci.,Vol. XL, Nov., 1865.
53
Wisconsin Academy of Sciences, Arts, and Letters
have been borne in the direction indicated, and scattered over the ar¬
eas of the Hamilton group, the Marshall sandstone, and the
Subcarboniferous limestone; that similar blocks of Hamilton rock have
been deposited over the two last named formations and even beyond;
that the Marshall sandstone has likewise been borne on to the Car¬
boniferous limestone, and that this transportation has been from lower
to higher levels, as the strata now lie, and are presumed to have lain,
since the basin is one of excavation and not of flexure. These phe¬
nomena, in all their details, are precisely what we should expect from
the action of a glacier advancing through the Erie valley, and moving
in a manner analogous to that of the Green Bay glacier. That a gla¬
cier moved through this valley has been abundantly shown by the Ohio
geologists. The only labor of this article is to show that it was an
individualized stream, forming the Ohio “Kame” belt on one side, and
the Michigan on the other, simultaneously, and that they are collat¬
eral members of a common moraine.
Eastward from Ohio, there has been, so far as I am aware, no definite
attempt to trace out the extent of the belt. In western New York, Prof.
Hall mentions, as one of the three general aspects of the superficial de¬
posits, a surface “broken into irregular hills or ridges, with deep bowl¬
shaped depressions, or long valleys, which often communicate in more
extensive ones, or are enclosed on all sides by drift,”1 but he does not
definitely locate the formation, or indicate whether it assumes the form
of a belt, or otherwise. In central New York, Prof. Vanuxem says: “There
is another class of deposits, well defined as to position, but irregular as
to composition, which are worthy of note. They occur in the north and
south valleys, which are on the south of the Mohawk river, or the great
level.” “The whole of these deposits have a common character. They
are in short hills, quite high for their base and are usually in considerable
numbers.” “They consist of gravel, stones, of stones also of greater size,
sand and earth.”2 These, he says, greatly resemble the “deluvial eleva¬
tions” noticed in the survey of Massachusetts,3 the description of which
is perfectly applicable to the formation under consideration. Furthermore,
Prof. F. H. King, of the Wisconsin survey, has examined the same de¬
posits in the vicinity of Ithaca, and recognizes their identity in kind. Nei¬
ther of these observers, however, discern a definite belt, although Prof.
Vanuxem destroys the force of his apparent limitation of the formation
1 Nat. Hist. Surv. 4th Dist., Geol., Pt. IV, pp. 320, 321.
2 Nat. Hist. Surv. N.Y., 3d Dist., p. 218,
3 Geol. of Mass., E. Hitchcock, 1833, p. 144.
54
Wisconsin Kettle Moraine
to the valleys, by stating that there are numerous points where it has
formed over the hill sides, and by associating in mention with it accumu¬
lations on the “heights, apparently in no regular order.”1 As these are
deep, canon-like valleys, they would probably modify in some degree,
the comparatively thin margin of the glacier, giving it a somewhat digi¬
tate outline, and the greatest accumulations would take place near the ex¬
tremities of the tongues, in the valleys, so far as drainage permitted; while
the connecting chains would form retreating lines, and be less conspicu¬
ous, and might, therefore, escape observation not definitely turned to the
subject. This, at least, is suggested by some observations of my own in
similar situations. Such valley accumulations, however, do occur at the
extremities of linear glacial lakes that are unconnected with a definite
belt, as in the case of Green Lake, Wisconsin.2
On the line of the Erie R. R., along the small tributary of the Dela¬
ware river that is followed up, westward, from Deposit, I have ob¬
served winding Osar-like ridges, parallel to the valley, and Kame-like
hills upon the slope, up to the watershed of the Delaware and
Susquehanna; likewise in the valley of the latter, at and near the vil¬
lage of Susquehanna, but I have no knowledge of their intimate struc¬
ture, extent or relations.
In the southeastern district of New York, Prof. Mather recognizes
the distinctive aspect of this class of accumulations.3 He cites sev¬
eral instances of its occurrence on the east side of the Hudson, leav¬
ing the impression that they are local features. But on Long Island, it
forms “an elevated ridge, called by some ‘Green Mountains,’ and by
others, the ‘Backbone’ of the island.4
This he describes in detail and maps, showing that it branches at
the east, one chain extending along the southern peninsula to Montauk
Point, and the other, along the northern to its extremity, and, theo¬
retically, to the islands beyond.
Professors Cook and Smock have recently examined this, and have
shown its connection with a similar moraine, that stretches across the
northern part of New Jersey, from Perth Amboy to the Delaware river,
below Belvidere.5 The descriptions of this range tally quite perfectly
with that of the Kettle moraine. This range, however, lies on the mar-
1 Loc. cit., p. 219.
2 Nat. Hist. Surv. N.Y. 1st Dist., Pt. IV, p. 212.
3 Geol. of Wis., 1877, Vol. II, p. 138.
4 Loc. cit., p. 161.
5 Ann. Rept. of State Geologist, N.J., 1877, pp. 9 et seq.
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Wisconsin Academy of Sciences, Arts, and Letters
gin of the area of northern drift, while the western one is medial in
position, and at some points is quite distant from the margin. It will
be observed, nevertheless, that this distance is greatest, in general, at
the west, and that in Ohio it becomes very greatly reduced, so that
the fact of coincidence on the Atlantic coast, presents no reason for
supposing the ranges to be distinct. But, whether distinct or not, is a
matter to be settled by observation, and it is to be hoped that it will
not long remain undecided for want of it. The extension of the New
Jersey moraine westward has not, so far as I can learn, yet been traced,
but the survey of Pennsylvania, in progress, will, doubtless, soon leave
nothing to be desired, so far as that State is involved.
To the eastward, Mr. Warren Upham has recently been engaged in
studying its probable continuation in southeastern Massachusetts. In
a personal communication he writes: “A very clear line of terminal
moraine extends along the chain of the Elizabeth islands southeast of
Buzzard’s Bay; thence it bends to the northeast and north as far as to
North Sandwich, when it turns at a right angle to the east, and ex¬
tends through Barnstable and other towns to Orleans, running along
the east and west portion of Cape Cod, and terminating at its east
shore.” “This terminal moraine, like the ‘Kettle moraine’, is not at
the outmost limit reached by the ice-sheet; for hills, in series nearly
parallel to the moraine already described, and similarly composed of
glacial drift with many boulders, occur on Martha’s Vineyard and Nan¬
tucket islands, corresponding, perhaps, to the terminal moraine which
forms the ‘backbone’ of Long Island. * * The moraine of the Eliza¬
beth islands and Cape Cod has a length of about 65 miles.” It may be
suggested that the range along the Elizabeth islands may correspond
to the northern branch of the Long Island moraine described by Prof.
Mather, and that, as Mr. Upham suggests, that of Martha’s Vineyard
and Nantucket corresponds to the southern.
Dr. E. Hitchcock refers to these accumulations in his report on the
geology of Massachusetts,1 and classes with them “diluvial elevations
and depressions,” occurring at other points in that and adjoining States.
It would appear, from the geological reports of the Eastern States that
analogous, though not certainly identical formations, occur locally,
more frequently than in the interior, and this, from the mountainous
nature of the country, is not strange; but no continuous massive range
seems to have been discerned, except the southern one already de¬
scribed.
1 Geol. of Mass., 1883, pp. 144 et seq.
56
Wisconsin Kettle Moraine
In the interior, so far as yet ascertained, the drift limit is not marked
by any such persistent ridge-like accumulation, but gradually dies
away or is buried by later deposits, so that the precise limit of glacial
advance is not easily determined. The only approach to an exception
to this, known to me, is the case of the Kettle moraine in Central Wis¬
consin, where it lies near the border of the driftless area. Elsewhere
around that area, the drift thins out very gradually, so as to render the
mapping of its margin a work of close inspection; and, as the region
presents no evidence of subsequent submersion, or any other special
modifying agency, except the usual meteorological forces, this would
seem to represent approximately the original form of deposit.
It is evident from the foregoing sketch that much observation re¬
mains to be made before the complete geography of this formation is
determined. The conjectural lines on the map are only theoretical sug¬
gestions, preliminary to observation.
Summary. — It may be helpful at this point to summarize, and bring
into close juxtaposition, in thought, the leading characteristics of this
remarkable formation.
1. Its linear extent is very great, whatever its final limits may be
found to be.
2. It has a width of from one to thirty miles.
3. Its average vertical thickness can only be very roughly estimated,
but may, very prudently, be placed at 200 or 300 feet.
4. Its surface configuration is peculiarly irregular, and denotes an
extraordinary origin.
5. It is a complex range, the component ridges being often arranged
in rude parallelism.
6. A distinction is usually to be observed between the superficial
and lateral portions of the deposit on the one hand, and the central,
underlying one on the other, the former being chiefly sand and gravel,
the latter complex commingled debris.
7. The superficial sands and gravels are usually stratified in vari¬
ous attitudes, but the core of the range is mainly unstratified.
8. The irregularities of the range are most conspicuous where the
superficial sands and gravels are least abundant.
9. The material was derived, in part, conspicuously so, from the
vicinity of the range, and, in part, from the formations lying back¬
ward along the line of drift movement for at least 300 miles.
10. A portion of the material is spherically rounded, a part is
scratched and polished, and some is little affected, though sometimes
soft or friable, the latter being usually from adjacent formations.
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Wisconsin Academy of Sciences, Arts, and Letters
1 1. The range is tortuous in its course, but sustains a remarkable
and significant relationship to the great lake basins.
12. It undulates over the face of the country, varying at least 800
feet in its vertical oscillations.
13. It does not sustain any uniform relation to present, or what are
presumed to have been, preglacial drainage systems in their details.
In some portions, it occupies water-partings; in others, lies on slopes;
and in still others, stretches across valleys.
14. It crosses, in its course, all the indurated formations, from the
Laurentian to the Coal measures, but exhibits no specific relation to
their strike or dip.
15. It sustains a definite and most important relationship to the
lines of general drift movement.
16. The range is frequently flanked on its southern, or outer edge,
by level areas of sand and gravel, of greater or less extent. These
also occur between the component ridges of the belt, and on the inner
flank, but less frequently.
17. The surface contour of the adjacent region within, or north of,
the belt, usually, though not invariably, has a less perfect drainage
system, and exhibits less noticeably the effects of superficial modifi¬
cation, than the outer side.
Origin. — Waiving, for the present, some further generalizations,
it is thought that the foregoing phenomena present a specific combi¬
nation which points unequivocally to a morainic origin. To the writer,
familiar with the multitudinous details, that cannot here find a place,
and having studied recent moraines with special reference to this for¬
mation, they have a force little less than demonstrative. The range is
confidently regarded as a moraine formed at the margin of a group of
glaciers — which may be regarded as a single lobate one — and mark¬
ing a definite stage of their history. A more vivid and graphic view
of the outline and movements of these glaciers, than can be given in
words, may be obtained from the accompanying map, from which it
will appear that through each of the great lake troughs there poured
an ice stream, attended by minor currents through the lesser channels.
Its Medial Position. — It has already been remarked that, in the inte¬
rior, this moraine does not mark the extreme limit of glacial advance.
Numerous striations, and other evidence of glaciation, occur on the south
side of it. A line has been drawn on the map intended to indicate the
approximate limit of northern drift, based on several authorities.1 How
1 Tesley, Newberry, Cox, and assistants, Worthen, Swallow, and Mudge.
58
Wisconsin Kettle Moraine
nearly this shows the limit of actual glacial progress, in distinction from
other means of transportation, is not, I think, as yet definitely ascertained,
but the general fact of progress, to a considerable distance beyond the
Kettle moraine, is sufficiently established. The moraine was, therefore,
formed after the retreat of the glacier had commenced, and marks a cer¬
tain stage of its subsequent history.
Glacial Movements before the Formation of the Moraine. — It be¬
comes an interesting question to ascertain whether the glacial move¬
ments were the same before the formation of the moraine, as after¬
wards. Fortunately, in southern Wisconsin, we have very definite and
specific evidence bearing on this question. In the towns of Portland
and Waterloo, which lie within the area of the Green Bay glacier, and
from twenty-five to thirty miles distant from the moraine, there are
several domes of quartzite that rise through the horizontal sandstones
and limestones, which occupy the surrounding region. These domes
are glacially abraded and grooved in a direction S. 30° W., and trains
of quartzite boulders stretch away in that direction to the moraine, and,
mingling with it, pass onward to an equal distance beyond. At the
same time there is abundant evidence from the material of the drift,
from the surface contour and from striation, recently observed by Mr.
I. M. Buell, that the westerly movement of the Lake Michigan gla¬
cier, near the Illinois line, extended to the west side of Rock River,
and that the line of junction of the two glaciers was on the west side
of that stream. It appears then, that in this region, the movements
were in the same general direction before and after the formation of
the moraine, but that there were changes in the details, and that the
relative size and position of the glaciers were somewhat different, the
Green Bay glacier being relatively smaller in the earlier epoch. Tes¬
timony of similar general import, but less specific, may be gleamed
[sic] from the reports of the other states involved.
Method of Formation. — If, then, the glacial movements were the
same, in general, before and after the formation of the moraine, and
yet the minor movements and relative size of the glaciers somewhat
different, how was the moraine formed? A halt in the retreat of the
glaciers, by which their confluent margin should remain stationary for
a period, would doubtless cause an unusual accumulation of debris,
but this would fail to account for the varying width or irregularities
of the moraine. The structure of the range seems to indicate an alter¬
nating retreat and advance of the ice mass. During the former, debris
was thrust out at the foot of the melting mass, which, when the gla¬
cier advanced, was plowed up into immense ridges. If this process
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Wisconsin Academy of Sciences, Arts, and Letters
be repeated several times parallel ranges will be accounted for, and
the irregularities incident to such advance and retreat will explain the
complexity of the range. Where the later advances were equal to the
earlier ones, the accumulation of drift material would be forced into
a single massive ridge. Where any advance failed to equal a former
one, an interval between the accumulations of the two would result,
giving rise to a depression whose form would depend upon the rela¬
tions of the two accumulations, but would in general be more or less
trough-like in character. Where tongues of ice were thrust into the
accumulated material an irregular or broken outline would be the re¬
sult. If masses of the ice became incorporated in the drift, as has been
suggested, their melting would give rise to depressions, constituting
one form of the kettles that characterize the range. The suggestion
just made, with reference to the irregular advance of the ice mass, ac¬
counts for other forms, and, at the same time, for the irregular hills,
mounds, and hillocks. Certain of the kettles may be due to
underdrainage, through the action of strong underground streams that
occasionally flow, as full brooklets, from its base. The drainage of
the glacier, while it was advancing and pushing the debris before it,
was probably quite general and promiscuous over the moraine, and
this would give rise to the stratified sands or gravels, and other evi¬
dences of the action of water, among which may perhaps, be reck¬
oned some of the minor mounds, ridges and depressions. The chang¬
ing attitudes, which the debris would be likely to assume, as it was
forced along, would, perhaps, give peculiar force to torrential effects.
The gaps in the range, attended by plains, or long streams of gravel
and sand, appear to represent the more considerable points of discharge
of the glacial floods. When the surface about the margin of the gla¬
cier permitted the accumulation of water, the moraine would doubt¬
less be much modified by it and present a subdued aspect.
The Alpine moraines, above referred to, are regarded as miniature
exemplifications of the process by which the Kettle moraine was
formed.
But, in addition to the structure of the range, the change in the rela¬
tive position of the Green Bay and Lake Michigan glaciers, already
alluded to, affords evidence of an exceedingly interesting character,
which has a significance much beyond what can be here indicated. It
appears that the junction between the Green Bay and Lake Michigan
glaciers at the last observable stage, preceding the formation of the
Kettle moraine, was about twenty-five miles farther west, than at the
time of the latter’s formation, or in other words, there is an abrubt
60
Wisconsin Kettle Moraine
easterly shift of the line of junction. It appears, also, that the width of
the ante-morainic Green Bay glacier, measured just south of the Kettle
moraine, was only half that of the post-morainic glacier, north of it,
measured at a distance just far enough to escape the terminal curva¬
ture. An inspection of the outline of the Green Bay glacier shows
that this eastward shift of the junction of the two glaciers was not due
simply to encroachment on the Lake Michigan stream, nor to a com¬
mon movement of both in that direction, for the opposite margin of
the Green Bay glacier lay close upon the borders of the driftless re¬
gion, demonstrating that there was no eastward swaying on that side.
Indeed, the indenture of the outline of the driftless area strongly sug¬
gests actual encroachment on that side also, and this view is not with¬
out independent support.
In harmony with these phenomena are the fiords of the Green Bay
peninsula, which indicate that the Green Bay ice stream overflowed
into the basin of Lake Michigan. These facts, taken altogether, seem
to warrant the belief that both glaciers retreated sufficiently far to the
northward, and within their respective basins, to allow time and op¬
portunity for the change in the relative size and position of the two
ice streams, and that, under slightly changed conditions that favored
the Green Bay glacier, they advanced to the position of the Kettle
moraine, and, after a series of oscillations, retreated permanently. This
view seems also to be demanded by certain details in the distribution
of the drift material that are otherwise enigmatical, but whose discus¬
sion would too much extend this article.
Significance . — As forty-five years have passed since Dr.
Hitchcock called attention to some of the phenomena under consider¬
ation, or, at least, to some distinctly related to it, and yet, the matter
has received so little consideration, that our present knowledge is lim¬
ited to such a degree, that I lay myself liable to the charge of undue
temerity in attempting to correlate the observations, I may be pardoned
in attempting to indicate, briefly, something of the significance and
importance the foregoing conclusions, if sustained, have in relation
to the Quaternary history of the region involved. The moraine con¬
stitutes a definite historical datum line , in the midst of the glacial ep¬
och, and becomes a basis of reference and correlation for adjacent for¬
mations. It is an historical rampart, outlining the great dynamic agency
of the period, at an important stage of its activity, and separating the
formations on either hand by a chronological barrier. It is manifest
that the true Boulder Clay, or ground moraine, south of the belt, must
have been formed earlier than that north of it, and that the two por-
61
Wisconsin Academy of Sciences, Arts, and Letters
tions are not at all synchronous. In sedimentary formations synchro¬
nism is found in horizontal strata, but in glacial deposits it is to be
sought in linear belts, concentric with the margin of the glacier. This
fact finds illustration, and emphasis, in the demarcation introduced by
this singular corrugation of the wide-spread glacial sheet. It is diffi¬
cult to limit the value of such a determinate line, in the midst of the
complex drift formations, if fully established, and should similar belts
be found to mark other stages of glaciation, there would be opened a
definite line of investigation that promises much assistance in unrav¬
eling the gnarled skein of Quaternary history.
While it does not follow, necessarily, that all formations overlay¬
ing the true glacial clay, south of the Kettle moraine, are older than
those occupying similar relations to the newer Till, north of it, it is
clear, that similarity of stratigraphical sequence is not, by any means,
sufficient ground for assuming chronological equivalence. It is evi¬
dent, that all endeavors at correlation between the superficial depos¬
its, on the opposite sides of the moraine, should be attempted with
much circumspection.
These suggestions have especial application to the discussion of the
vegetal deposits, so frequently found in the later Quaternary forma¬
tions. By many writers, the various deposits of this kind, in the Mis¬
sissippi basin, have been, very naturally, in the present state of our
knowledge, grouped together without reference to the necessary dis¬
criminations above indicated, and, as a result, beds of diverse age are
referred to a common stratum. A general discussion of these depos¬
its is not sufficiently germane to our subject to be fittingly introduced
here, but it is appropriate to point out the fact that some of the veg¬
etal strata sustain such a relation to the Kettle moraine, that they must
be widely separated from others, in the date of their accumulation and
burial. Some of these organic strata lie at the immediate foot of the
moraine, beneath fluviatile and lacustrine deposits that, I am confi¬
dent, began to be accumulated during the accumulation of the moraine,
and through the agency of glacial floods; while it is even more cer¬
tain, that other vegetal deposits accumulated much subsequently, as
those found in the red clays of Wisconsin, which are lacustrine de¬
posits of the great lakes formed after the recession of the glacier. It
would be too much to assume that all plant remains, found south of
the moraine, antedate its formation, but it is safe to affirm that, with
only phenomenal exceptions, e.g., such as escaped glacial abrasion,
all north of it are more recent.
The bearing of these definite determinations of the glacial outlines
62
Wisconsin Kettle Moraine
and movements upon the question of the origin of the remarkable
driftless area of Wisconsin, Minnesota, Iowa and Illinois (see map)
was early perceived, and it was clearly foreseen that this line of in¬
vestigation promised a demonstrative solution of the problem. The
driftless area manifestly owes its origin to the divergence of the gla¬
ciers through the Lake Superior channel, on the one hand, and that of
Green Bay and Lake Michigan, on the other, and to the obstacle pre¬
sented by the highlands of northern Wisconsin and Michigan. This
obstacle the glacier surmounted, and passed some distance down the
southern slope, but apparently not in sufficient thickness to overcome
the melting and wasting to which it was subjected, and so it termi¬
nated midway the slope. But the deep, massive ice currents of the
great channels pushed far on to the south, converging toward each
other; and, if they did not actually unite, at least commingled their
debris south of the driftless area.1 An instance closely similar to this,
considered from a dynamical point of view, may be seen, at the present
termination of the Viesch glacier, and illustrations of the general prin¬
ciples involved in the explanation may be seen in connection with sev¬
eral other Alpine glaciers.
If the evidence adduced to show that the Kettle moraine was due
to an advance of the glaciers be trustworthy, then, to the extent of that
advance, whether much or little, the moraine marks a secondary pe¬
riod of glaciation, with an interval of deglaciation between it and the
epoch of extreme advance. Its great extent indicates that whatever
agency caused the advance was very wide spread, if not continental
in its influence. The moraine, therefore, may be worthy of study in
its bearings upon the interesting question of glacial and interglacial
periods.
It will also furnish definite data bearing upon the somewhat mooted
question of the origin of the Great Lakes, as well as other questions
involving both perglacial [sic] and postglacial topography.
1 Compare N. H. Winchell in An. Rep., Geol. of Minn., 1876, and R. D. Irving,
Geol. of Wis., Vol. II, 1877, whose views are closely analogous to the above and each
to the other but are not strictly identical. See, also, J. D. Dana, Am. Jour. Sci., April
1878.
63
Wisconsin Academy of Sciences, Arts, and Letters
Philo Romayne Hoy, M.D.
(1816-1892)
Physician, surgeon, and scientist; collector of plants, fossils, and
relics of aboriginal life; charter member and second president
of the Wisconsin Academy, 1876-78
64
Extinct Wild Animals in Wisconsin
THE LARGER WILD ANIMALS THAT HAVE BECOME
EXTINCT IN WISCONSIN
(Read at the Racine meeting)
BY DR. P. R. HOY
A record of the date and order in which native animals become
extinct within the bounds of any country is of present interest, and in
the future may be perused with redoubled satisfaction.
Fifty years ago the territory now included in the state of Wisconsin
was nearly in its primitive condition. Then many of the larger wild
animals were abundant. Now all has changed; the ax and plow, gun
and dog, railway and telegraph, have completely metamorphosed the
face of nature. Not a few of the large quadrupeds and birds have been
exterminated or have hid themselves away in the wilderness of
northern Wisconsin.
There was a time, away back in the dim past, when the mastodon,
ox, elephant, tapir, peccary, and musk-ox roamed over the ancient
prairies of Wisconsin, but now only their bones, from time to time,
are exhumed and thus exposed to the wondering gaze of the ignorant
many and the trained eye of the wiser few. We shall at this time,
however, confine our attention to the historic period.
The antelope, Antilocarpa Americana , now found only on the
western plains, did, two hundred years ago, inhabit Wisconsin as far
east as Lake Michigan. In October, 1679, Father Hennepin, with La
Salle and party, in four canoes, coasted along the western shore of
Lake Michigan. In Hennepin’s narrative he says: “The oldest of them”
[the Indians] “came to us the next morning, with their calumets of
peace, and brought some wild goats.” This was at or near Milwaukee.
“Being in sore distress, we saw upon the coast a great many ravens
and eagles, from whence we conjectured there was some prey, and
having landed on that spot we found above the half of a fat wild goat
which the wolves had strangled. This provision was very acceptable
to us, and the rudest of our men could not but praise the Divine
Providence which took so particular care of them.” This was,
undoubtedly, near Racine. “On the 16th” [October 16, 1679] “we met
with abundance of game; a savage we had with us killed several stags
and wild goats , and our men a great many turkey, very fat and big.”
This last point was between Kenosha and Racine. Hennepin’s goats
were without doubt antelopes. Father Joliet, a little earlier, mentions
65
Wisconsin Academy of Sciences, Arts, and Letters
that “on the Wisconsin there are plenty of turkey cocks, parrots, quails,
wild oxen, stags and wild goats.” All species of the deer family were
called stags by the early travelers. Schoolcraft mentions antelopes as
occurring in the Northwestern Territory, and as late as 1850.
Antelopes were not uncommon in southern Minnesota, only forty miles
west of the Mississippi river. It is evident, then, that antelopes have
retired quite leisurely.
When the last buffalo, Bos. Americana , crossed the Mississippi is
not precisely known. Governor Dodge told me that buffalo were killed
on the Wisconsin side of the St. Croix river the next year after the
close of the Blackhawk war, which would be 1833. So Wisconsin had
the last buffaloes east of the Mississippi river.
The Woodland Caribon, Rangifer Caribou , were probably never
numerous within the limits of the state. A few, however, were seen
near La Point in 1840; none since.
Elk, Cervus Canadensis , were on Hay river in 1863, and I have
but little doubt that a few still linger with us. The next to follow the
buffalo, antelope and reindeer.
Moose, Alee Americanus , continue to inhabit the northern part of
the state, where they still range in spite of persecution. A fine cow
moose was shot near the line of the Wisconsin Central Railway in
December, 1877.
A few panthers, Felis Concolor , are yet with us; a straggler is
occasionally seen. Benjamin Bones of Racine shot one on the head¬
waters of Black river, December, 1863.
Wolverines, Gulo luscus, are occasionally taken in the timber; one
was taken in La Crosse County in 1870.
Of beaver, Castor Canadensis , a few still continue to inhabit some
of the small lakes situated in Lincoln and adjacent counties.
The badger, Taxidea Americana, is now nearly extinct in Wisconsin.
In a few years the only badger found in the state will be the one on
the coat of arms.
The opossum, Didelphis Virginiana, were not uncommon in Racine
and Walworth counties as late as 1848. They have been caught as far
north as Waukesha, and one near Madison in 1872, since which time
I have not heard of any being taken. I am told that a few are still
found in Grant county. They will soon be exterminated, no doubt.
The last wild turkeys, Meleagris Gallopavo, in the eastern part of the
state, was [sic] in the fall of 1846, at which time a few were discovered
near Racine. They were hunted with such vigor that the entire number
were shot, “The last of the Mohicans.” I am told, by Dr. E. B. Wolcott,
66
Extinct Wild Animals in Wisconsin
that turkeys were abundant in Wisconsin previous to the hard winter
of 1842-3, when snow was yet two feet deep in March, with a firm
crust, so that the turkeys could not get to the ground; they hence
became so poor and weak that they could not fly and so were an easy
prey for the wolves, wildcats, foxes and minks. The Doctor further
stated that he saw but one single turkey the next winter, and none
since. One was shot in Grant county in the fall of 1872. Possibly
there are a few yet to be found in this large southwestern county; if
not, then wild turkeys are exterminated in the state of Wisconsin.
67
Wisconsin Academy of Sciences, Arts, and Letters
John Wesley Hoyt, M.D., LL.D.
(1831-1912)
Professor of chemistry and natural history at Antioch College;
secretary of the Wisconsin State Agricultural Society; editor of
the Wisconsin Farmer ; charter member and first president of the
Wisconsin Academy, 1870-75; governor of the Wyoming
Territory and first president of the University of Wyoming
68
Courtesy The State Historical Society of Wisconsin WHi(x3)1759
Personal Recollections of Abraham Lincoln
SOME PERSONAL RECOLLECTIONS OF
ABRAHAM LINCOLN
BY JOHN WESLEY HOYT
My deep interest in Mr. Lincoln came, first, of his manifestations
of opposition to any further extension of slavery over the territories
of the United States — an opposition in which I believe I shared as sin¬
cerely as any American; for, while a student and medical professor in
Cincinnati, in the early fifties of the last century, I had ofttimes looked
across the Ohio River to the shadows of the Kentucky side, and now
and then, by sympathy, felt the smart of a driver’s lash on Freedom’s
shore; there, too, had earnest part in forming the great political party
solemnly sworn to resist extension of the damning curse of human
bondage, and thence had gone out, as one of Freedom’s advocates on
more than a hundred ‘stumps,’ in Ohio, Indiana, Illinois, and Wiscon¬
sin.
Meanwhile, I had, with profound interest, so watched the masterly
discussions of Mr. Lincoln with Douglas, in northern Illinois, and so
marked him for his destiny, that, in the winter of 1858-9, being then
in command of agricultural affairs in Wisconsin, I went down to Chi¬
cago to congratulate him and, if possible, secure him for delivery of
the annual address at the next state fair, to be held at Milwaukee in
September, 1859.
We spent half the night together, in his chamber, reviewing the past
and outlining a possible, even probable future — an evening so deeply
interesting that, after fifty years, the discussions and incidents are still
almost fresh enough for recital in detail. Even then the dark clouds
of a coming conflict hovered near enough to make one anxious; but
in the minds of both, even civil war, with carnage widespread and fear¬
ful, seemed not so dreadful as a further extension of human slavery
over half a continent by consent of possessors whose immediate an¬
cestors had themselves been freed from British oppression, not half
so terrible, at great cost of blood and treasure. There was yet hope
that the resolute champions of the curse would stay their demands,
but the prospect was sadly faint, for even then the need of preparing
for the worst was painfully felt.
I need hardly say that my conviction of the greatness of Mr. Lin¬
coln, already gained by a reading of his discussions of the all-engross¬
ing questions of the time, was yet further deepened by that night’s
69
Wisconsin Academy of Sciences, Arts, and Letters
experience and study of the homely, robust statesman before me, and
that, with a glad heart I bore away, at midnight, his promise to be with
us, in Milwaukee, at the appointed time.
When, at the moment of departure, he was asked to let me know
the time of his leaving Chicago, so that I could meet him on his ar¬
rival in Milwaukee, he merely said, with his characteristic simplicity:
“Oh, don’t trouble yourself on my account; I’ll be at the Newhall in
good time, all right.” And so he was, some eight months later.
But it so happened that his actual arrival was at midnight, and that
the room intended to be reserved for him had, by the blunder of a clerk,
been given to a man and his wife who were already in bed and asleep.
There was no remaining vacant room in the house, and the clerk, hav¬
ing been stoutly arraigned by the landlord, was in distress of mind;
seeing which, Mr. Lincoln, with a smiling countenance and comfort¬
ing words, said: “Oh, my dear sir, don’t be unhappy on my account.
I see there is vacant space enough right here, at the end of the counter.
Just bring a cot and clothes-rack, with sheet for a screen, and I’ll sleep
like a top.” The thing was done, and the distinguished guest, after a
cheerful and hearty “Goodnight, gentlemen,” handsomely retired.
Of course I was prompt to fulfill my promise to come down in good
time to breakfast with him, but he was a little tardy, so that when,
having heard a little stir behind the screen, I ventured to tap gently
on the frame, word came out at once, “Come in!” But, on passing
‘round, I found him not only half dressed, but shaving himself, and
so encumbered that, instead of moving his chair for a greeting of his
visitor, having recognized my voice, he turned his head squarely back
and saw me, with his lathered face inverted and considerably broad¬
ened by a smile. Of course I was quick to retire and wait.
The breakfast disposed of, we were soon on our way to the Fair
grounds, for Mr. Lincoln said he wanted to see what sort of farmers,
gardeners, and mechanics the Badgers made.
The address was to be at 1 1:00, and meanwhile we made ourselves
very busy, going the rounds of all the departments. It soon became
apparent that, notwithstanding his modest disclaimer of knowing much
of practical affairs beside wood-chopping and rail-splitting, he did
know much of many things in country life; that he was in fact capable
of critical judgment of horses, cattle, sheep, and other domestic ani¬
mals, as well as of most products of the soil.
The address was listened to by many thousands, some say thirty
thousand, not a few of whom had made special efforts and sacrifices
that they might see and hear the man who, from the depths of poverty
70
Personal Recollections of Abraham Lincoln
and laborious service in wood and field, had risen to a foremost place
in the legal profession and in statesmanship. Perhaps no address more
practical, useful, and entertaining was ever delivered on any such oc¬
casion. It dealt with the necessary relation between education and la¬
bor, as well as with the economy of thorough work in farming espe¬
cially, and was so enlivened by humorous hits that it was at once
highly entertaining and of enduring value. It was in fact so admirable,
and so deepened my conviction of his eminent fitness for leadership,
that then and there I began to speak of him as the man for the next
President of the United States — fit for a superior service in statesman¬
ship at any time, but pre-eminently fit for such a crisis as then seemed
surely very near — in due time I went to Chicago, to help nominate
him, and thereafter gave myself to platform service in many of the
Northern states, and to the end of the campaign.
How nobly, now grandly he transcended the highest expectations
of his most sanguine admirers is too well known for historic proof.
No greater demand for a national guide and guardian was ever made,
or more nobly and wonderfully met in any part of the world. It is
certain that, for measure of endowment and balance of powers, the
supreme founder and father of the Republic alone can be compared
with Lincoln, its preserver and the emancipator of millions of a down¬
trodden and most wretched race.
Intellectually, Mr. Lincoln was remarkable for the habit of close
and critical attention to whatever engaged his thought; for such power
of discrimination and comparison as made him clear-headed; such
power of logical analysis as made him quick to detect a flaw and ex¬
pose a fallacy, on which account his opponent in debate ofttimes found
himself floundering ere he knew he was on the wrong side, and pain¬
fully subject to such withering sarcasm, if he deserved it, as Mr. Lin¬
coln knew so well how to use; remarkable also for such readiness to
discover the relations of things as made him far-sighted and hence ei¬
ther courageous, even bold and daring, or prudent, as the occasion
might justify or demand.
On the side of the sensibilities I was happy to find, after a further
acquaintance, that I had myself underrated him. His rugged, stalwart
frame was at first suggestive of a probable sternness of spirit and man¬
ner. But, as I came nearer, I was charmed by the delicacy, even ten¬
derness, and all-abounding sympathy of a great and beautiful soul —
qualities that made him a lover of the beautiful in nature; that prompted
him, on entering the great round rent at the Wisconsin State Fair, with
its magnificent display of fruits and flowers, to take off his hat, for a
71
Wisconsin Academy of Sciences, Arts, and Letters
salute, with a grace that won the hearts of all who were present, say¬
ing: “How beautiful! Eden transferred!;” that made him too glad for
utterance when he signed the immortal Emancipation Proclamation and
saw the shackles fall from millions of his fellow-men, and again when,
after one of the most fearful conflicts in human history, he knew the
Republic saved and foresaw a Union grander and more glorious than
had been dreamed of in all the past, a thing of destiny; qualities, too,
that made him so impressionable by others, so sensitive in soul, that
he almost never failed to judge rightly the men with whom he had to
do, and enabled him to draw into the service of his country so great a
galaxy of men of genius, devotion, and heroic virtue.
Morally, Mr. Lincoln was nothing less than an embodiment of vir¬
tue, truth, and justice. Those who knew him best believed him inca¬
pable of wilful wrong. He so loved truth that he was ever in earnest
search of it, and anxious to make it known; and it was the cherishing
of a profound love of justice, and his exalted aims and aspirations that
made him every ready, even glad, to do and die for his country.
As for the will, he was resolution itself — never halting or hesitat¬
ing in his course. Because he felt himself right, and knew the right
must win, there was fixedness of purpose. He never just hoped for a
final victory; he saw it coming, and though deeply sad over the dread¬
ful fate of so many martyrs, yet, after all, whenever the future of the
Republic was referred to, his noble face was illumined. It was this
high assurance of a determined soul that made it easy for him to say
to me, one dark morning, when I had gone to the White House, with
anxious sympathy, because great armies of Confederate troops had
boldly crowded into Pennsylvania and were threatening both Harris¬
burg and Philadelphia, “Never mind, Dr. Hoyt, you may be sure we’ll
trot them out of there very soon and make them glad to get home
again.”
It was this fixedness of purpose and his unfailing confidence that
enabled him to preserve his calmness, so that he was rarely disturbed
in spirit and never really agitated. His face and voice and daily life
were ever giving expression to an unwavering trust in God.
And thus it is that we are amply justified in pronouncing Abraham
Lincoln one of the very noblest and grandest of men in all human his¬
tory.
Washington, D.C.
72
Part Two:
Current Articles
John A. Cross
Dairying in an urban environment:
The Milwaukee metropolitan area
Abstract Urban expansion in the greater Milwaukee area has displaced
dairy farming. Southeast Wisconsin now accounts for only three
percent of the state’s milk production. The spatial pattern of de¬
cline in dairy production in southeastern Wisconsin between 1989
and 1994 is examined at the civil town level, considering farm
entry and exit. Farm relocation from the region is also explored,
and the stress experienced by the region ’s farmers is surveyed.
Urban expansion in the greater Milwaukee metropoli¬
tan area has significantly altered the pattern of dairy farm¬
ing in the southeastern Wisconsin agricultural reporting dis¬
trict, whose boundaries roughly coincide with the Census
Bureau-defined Milwaukee, Racine, and Kenosha metropoli¬
tan areas. Early this century Waukesha County was considered
one of the state’s leading milk producers, having more pure
blood dairy cows than any other similar area in the nation
(Whitbeck 1921). Half a century ago 50 percent of Milwau¬
kee County was in agricultural production, with 440 dairy
farms in operation (Durand 1962). Durand (1962, 1963) has
described in detail the steep decline of dairying in Milwaukee
County in the two decades following the outbreak of World
War II. By the beginning of the 1960s approximately 50 dairy
farms remained in Milwaukee County and dairying had re¬
treated in Waukesha and Ozaukee Counties. The zone of ag¬
ricultural demise that Durand described is now overwhelming
the surrounding counties. Four dairy herds remain in Milwau¬
kee County, and only two are on commercial farms. One of
the other two herds is at the Milwaukee County Zoo Heri¬
tage Farm, while the second is at the Wisconsin State Fair Park.
These herds are but small reminders of dairying in a county
that once had been “near the top of the leading dairy counties
of the nation” (Durand 1962).
TRANSACTIONS Volume 83 (1995)
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This paper explores the spatial pattern of
decline in dairy production in the metropoli¬
tan counties of southeastern Wisconsin. By
spring 1994 the region had only 925 herds
(Table 1), with a total of 63,100 cows
counted the previous year (Wisconsin Agri¬
cultural Statistics Service 1994a). Twenty
years earlier Milwaukee County already had
dropped to 14 dairy herds, but southeastern
Wisconsin still had 2,050 herds, with a to¬
tal of 89,400 cows (Wisconsin Statistical Re¬
porting Service 1975). The entire region lost
41.3 percent of its herds over the past de¬
cade, with a decline of 23.7 percent between
1989 and 1994.
The decline in number of milk cows in
southeastern Wisconsin has only slightly
trailed the shrinkage in number of dairy farm
herds. While the region lost 21.1 percent of
its milk cows since 1983, most of the loss was
in the past half decade, inasmuch as the num¬
ber of cows within the area fell by only 2,700
between 1983 and 1988. Between 1988 and
1993 the region lost 9,300 dairy cows, a loss
of 18.4 percent, and milk production de¬
creased by 14.2 percent (Wisconsin Agricul¬
ture Reporting Service, 1984; Wisconsin Ag¬
ricultural Statistics Service 1989, 1994a).
Spatial Patterns of Decline
Over the past half decade, at the county level
the greatest proportional declines in dairy¬
ing in southeastern Wisconsin occurred in
Waukesha and Kenosha Counties, both of
which lost more than one quarter of their
herds (Table 1). Wisconsin Department of
Agriculture lists of dairy herds that have un¬
dergone the Brucellosis Ring Test (required
for commercial milk sales) reveal changes in
dairy operations at the civil town (Table 2)
and section level (see Cross 1994a). The en¬
try and exit behavior of dairy farmers at the
civil town level provides a clearer picture of
the expanding ring of dairy abandonment,
a ring corresponding to the perimetropolitan
bow wave that Hart (1991) has so eloquently
described. Dairy farming, of all types of ag¬
riculture, is among the most vulnerable to
urban development, because of a variety of
conflicts that arise with non-farm neighbors
(Hirschl and Long 1993). Such conflicts in¬
clude vandalism and trespassing, complaints
about farm odors and farm equipment mov¬
ing over suburban roads, and inevitable in¬
creases in taxes and land use controls.
Only four towns in the southeast Wiscon-
Table 1 . Decline in number of dairy herds in southeast Wisconsin
Data sources: U.S. Census of Agriculture, 1947 (number of farms reporting milk cows in 1940), Wisconsin
Agriculture Reporting Service 1984; Wisconsin Agricultural Statistics Service 1994 (number of herds in late
March or early April of each year tested for Brucellosis — required for milk sales).
76
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CROSS: Dairying in an urban environment
Number
of Dairy Herds
in 1994
□ 0 to 2
H 3 to 10
□ 1 1 to 20
■ 21 to 57
WASHINGTON
COUNTY
City of Milwaukee
COUNTY
Fig. 1. Number of dairy herds is greatest in towns most distant from Milwaukee. Data
source: See Table 2.
sin agricultural reporting district had as
many as one dairy herd per square mile by
early 1994 (Figure 1). One of these was in
northern Ozaukee County, and the other
three were in western Washington County.
Closer to Milwaukee County, three towns
within eastern Waukesha County (Brook¬
field, Menomonee Falls, and New Berlin)
had two or fewer herds. Brookfield had
none. None of the towns along Lake Michi¬
gan south of Milwaukee to the Illinois state
line had more than five herds. The greatest
proportional decreases in dairy herds since
1989 were in a swath of towns extending
from Menomonee Falls in northeast Wau¬
kesha County southwest to Mukwonago in
Waukesha County and three towns in
northern Walworth County. All lost at least
ten percent of their herds annually since
1989 (Figure 2). Because many of these
towns already had relatively few surviving
dairy farms, the greatest numerical losses
were in towns that were farther from Mil¬
waukee (Figure 3). For example, four towns
north and west of the Menomonee Falls-
Pewaukee areas all lost more than six herds.
Four towns in northern Ozaukee and Wash¬
ington Counties and several towns within
Walworth County had similar losses.
Entry and Exit Activity
One measure of the declining viability of
dairying is the ratio of new dairy operators
to farmers leaving. The number of farmers
quitting dairying outnumbered the number
Volume 83 (1995)
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CROSS: Dairying in an urban environment
Data sources: Computer tapes listing dairy farms having had the Brucellosis Ring Test (required of all
commercial dairy herds), which provided the dairy farmers’ names, mailing addresses, grade and farm
location (by county, civil town, and section). These tapes, produced in early April 1 989, early April 1 990, early
April 1991, late March 1992, late March 1993, and April 4, 1994 (the last four dates coordinated with the
published statistics of the Wisconsin Agricultural Statistics Service) were obtained from the Wisconsin
Department of Agriculture, Trade, and Consumer Protection, Madison.
NOTE: Farmers who had ceased operations within any given civil town between any two consecutive years
were considered to have exited. Farmers who had begun operations within any given civil town between any
two consecutive years were considered to have entered. Because many farmers moved their operations during
a year, these figures will overestimate the actual number of farmers who have abandoned dairy operations
entirely, regardless of location. Farmers moving within a town had moved their operations within the same civil
town to a noncontiguous section, thus a distance of at least one mile, and are listed separately, but not included
within the totals of farmers entering and exiting dairying.
Volume 83 (1995)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Fig. 2. Percentage decrease in dairy herds is typically greatest in towns experiencing
urban expansion, as in Waukesha County. Data source: See Table 2.
of farmers entering in the metropolitan
counties of southeastern Wisconsin (exclud¬
ing Walworth County) by a ratio of 4.2 to
1.0, far surpassing the overall statewide ra¬
tio of 2.0 to 1.0. Although dairying is still
viewed as viable within some towns, such as
Farmington in northeastern Washington
County, where eight new operators replaced
ten who ceased operations between 1989
and 1994, only a few other towns have ra¬
tios close to the state average. These towns
are around the northern and western perim¬
eter of the southeast Wisconsin agricultural
reporting district. Many towns have no en¬
tering dairy farmers, and former dairy farms
are becoming the sites of shopping centers
and residential subdivisions.
Farmers exiting dairying in southeastern
Wisconsin are not necessarily leaving agri¬
culture. Over half of the land area in five of
the region’s seven counties is still classified
as “land in farms” (Table 3), and 63.1 per¬
cent of Racine County is considered agricul¬
tural. Conversely, farmland now comprises
less than ten percent of Milwaukee County,
but the 130 remaining farmers received av¬
erage earnings of $1,727 per acre in 1992,
a figure far higher than the statewide aver¬
age of $318 (calculated from Wisconsin Ag¬
ricultural Statistics Service 1994b). Wiscon¬
sin farmers as a whole rely on dairying to
produce over half of their cash receipts, but
farmers in several metropolitan counties
have concentrated upon other agricultural
80
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CROSS: Dairying in an urban environment
Fig. 3. Largest losses in number of dairy herds often occur in towns with the largest
number of herds. Data source: See Table 2.
Table 3. Agricultural production in southeastern Wisconsin
Data source: Wisconsin Agricultural Statistics Service 1993a and 1994b
Volume 83 (1995)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
activities. Milwaukee County leads the state
in its cash receipts for “other crops,” a cat¬
egory that includes greenhouse and nursery
crops together with fruit and miscellaneous
specialty crops. Racine County is the state’s
second largest producer of eggs and poultry
(Wisconsin Agricultural Statistics Service
1993). Moreover, as Hart (1991) notes in
his discussion of the bow-wave process,
“[s]ome . . . keep their land and continue
to farm it less intensively.” The low cash re¬
ceipts per acre for Kenosha and Waukesha
Counties, where only one in seven farmers
have dairy herds, lend credence to such ar¬
guments.
Emigration of Dairy Farmers
Some farmers remain in dairying by fleeing
the advance of urbanization. Undoubtedly,
some dairy farmers may have moved to lo¬
cations in northwestern Illinois, eastern Iowa
or southern Minnesota, although the avail¬
able data does not permit those movements
to be traced. Statistics do show that between
1989 and 1994 fifty-two southeastern Wis¬
consin dairy farmers moved their operations
to other locations in the state. Some of these
moves were local: thirteen were to another
town within the same county, eight others
were to non-contiguous sections within the
same town, and two were to another county
within southeastern Wisconsin. However,
29 dairy farmers moved from southeast Wis¬
consin to other areas of the state (Figure 4).
For example, dairy farmers moved from
Racine County to Lafayette, Sauk, Buffalo,
Manitowoc, and Sheboygan Counties.
Waukesha County lost dairy farmers to
Dodge, Jefferson, Waupaca, Green, and
Richland Counties.
Neighbors or kin sometimes move in uni¬
son. Clark County received three incoming
operators from a small area of far western
Kenosha and adjacent Walworth Counties.
Two farmers living six miles apart in Wash¬
ington County relocated to towns in east¬
ern Taylor County. Eight of the 29 farmers
relocating from southeastern Wisconsin
moved more than 100 miles.
Stresses of Urbanization
Stresses of urbanization are clearly recognized
by dairy farmers in southeastern Wisconsin
(Table 4). Although survey respondents
(Cross 1994b) in southeastern Wisconsin did
not differ significantly from dairy farmers
elsewhere in the state in their evaluation of a
variety of potential problems facing them —
including wholesale milk prices, hay and feed
prices and shortages, labor availability, farm
debt and interest rates, various climatic
hazards, and government regulations — their
evaluations of property taxes and local urban
expansion were significantly different
statistically. Indeed, half of the surveyed dairy
farmers in southeastern Wisconsin (33
percent, excluding Walworth County) ranked
“local urban expansion” as a “major
problem,” compared with just 6.1 percent of
dairy farmers elsewhere in the state.
Nevertheless, only 13.2 percent of the farmers
in this region wish to sell their farms,
compared with 26.8 percent of the farmers
elsewhere in the state, a statistical difference
significant at the 0.10 level.
Ties to the land are strong, with dairy
farms in southeastern Wisconsin more likely
to have been owned by a family member
since the 1800s than in any other region of
the state. Twenty-four percent of the dairy
farmers surveyed by the author in the region
indicate that their family had operated their
farm since the last century, compared with
14 percent elsewhere in Wisconsin. South¬
eastern Wisconsin dairy farmers were less
likely to submit bids to participate in the
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CROSS: Dairying in an urban environment
Dairy Farmers Relocating
Fig. 4. Twenty-nine dairy farmers moved their herds from southeastern Wisconsin to
other parts of the state between 1989 and 1994. Data source: See Table 2.
Volume 83 (1995)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
Table 4. Perception of problems facing dairy farmers: southeast Wisconsin versus other
Wisconsin farmers
Data source: Survey of Wisconsin dairy farmers conducted spring 1993. See Cross 1994b.
Dairy Termination Program in 1986 than
elsewhere in the state, although their bid ac¬
ceptance rates exceeded the state average
(Cross 1989). On the other hand, southeast¬
ern Wisconsin dairy farmers are slightly less
likely (36.8 versus 42.7 percent, not statis¬
tically significant) to “expect that a son, a
daughter, or another relative will operate
their dairy farm after [they] retire.”
Agricultural land sales prices in southeast
Wisconsin are the highest in the state, aver¬
aging $1,929 per acre if the land remained
in agriculture and $3,679 if the land was di¬
verted to other uses. These 1992 figures were
actual decreases from 1991 values (Wiscon¬
sin Agricultural Statistics Service 1994b).
Prices averaged $3,539 and $3,348 per acre
for land remaining in agriculture in Milwau¬
kee and Waukesha Counties. Such high land
prices burden farmers with high taxes, make
farm expansion prohibitively expensive, and
provide strong incentives for farmers wish¬
ing to sell. In contrast, agricultural land sales
averaged less than $1,000 per acre in 54 of
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CROSS: Dairying in an urban environment
Wisconsin’s 72 counties and below $750 in
38 of the state’s counties.
The median acreage owned by southeast¬
ern Wisconsin dairy farmers responding to
the 1993 survey was 130 acres, contrasting
with a statewide median of 180 acres. (Ac¬
tual median farm size was 255 acres, because
many farmers rented additional acreage.)
Furthermore, land fragmentation is a prob¬
lem. In no other region of Wisconsin did a
smaller proportion (13 percent) of dairy
farmers report that their “farm fields were
located all together,” not even separated by
a road. One third of the southeastern Wis¬
consin dairy farmers, the highest proportion
in the state, reported that their fields were
separated by at least two miles.
The average dairy farmer in southeastern
Wisconsin grows over 95 percent of the hay
and feed grains fed to his herd, the highest
average among the state’s nine agricultural
reporting districts. Nearly three-quarters of
the dairy farmers in this area report that they
normally grow all of the feedstuffs for their
cows. Unlike their counterparts in other ur¬
banizing areas, whose dairy farms survive by
importing large quantities of feed, the south¬
eastern Wisconsin dairy farmer relies upon
production from his increasingly expensive
farmlands. Nevertheless, average herd size in
southeastern Wisconsin exceeds that of all
the state’s other agricultural districts.
Conclusions
Three broad dairying zones can be identi¬
fied in southeastern Wisconsin: (1) a zone
in which dairying has ceased to be impor¬
tant, with only a few solitary holdouts who
have resisted the pressures to sell-out; (2) a
zone of rapid decline, in which large num¬
bers of farmers are abandoning dairying at
rates considerably in excess of average rates
statewide; and (3) a peripheral zone in which
dairying remains important. Although the
impacts of urban expansion upon dairying
may be most conspicuous in the greater Mil¬
waukee metropolitan area, fingers of decline
have spread towards other areas of Wiscon¬
sin, including a zone extending west to
Madison and north along the Fox River Val¬
ley, particularly between Fond du Lac and
Green Bay. What we have seen surround¬
ing Milwaukee is not unique. The same pro¬
cess has been documented around Chicago
(Berry 1979), but it is radically different
from the process in southern California (Gil¬
bert and Akor 1988), which has resulted in
highly capitalized corporate dairy farms that
took away Wisconsin’s number one rank as
a milk producing state during August 1993.
Wisconsin remains a state of family farms,
but dairy farmers are rapidly leaving the
southeastern metropolitan area of the state.
Acknowledgments
I wish to thank two anonymous reviewers
for their valuable comments upon an earlier
version of this manuscript. Funding for the
survey that provided data for part of this pa¬
per was provided by the University of Wis¬
consin Oshkosh Faculty Development Re¬
search Board.
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urbanization: a study of Northeastern Illinois.
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
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sin Department of Agriculture, Trade and
Consumer Protection.
- . 1993. Wisconsin agricultural statistics
— 1993. Madison: Wisconsin Department of
Agriculture, Trade and Consumer Protec¬
tion.
- . 1994a. Wisconsin 1994 dairy facts.
Madison: Wisconsin Department of Agricul¬
ture, Trade and Consumer Protection.
- . 1994b. Wisconsin agricultural statistics
- 1994. Madison: Wisconsin Department of
Agriculture, Trade and Consumer Protection.
John A. Cross is professor of geography and chair
of the Geography Department at the University
of Wisconsin Oshkosh. He has published several
papers on Wisconsin agriculture and on natural
hazards. Address: Dept, of Geography, UW
Oshkosh, 800 Algoma Blvd., Oshkosh, WI
54901-8642
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Eric E. Jorgensen and Lyle E. Nauman
Small mammal distribution associated
with commercial cranberry production
Abstract We documented the distribution of small mammals in association
with commercial cranberry production in south-central Wiscon¬
sin. Small mammals were captured with snap traps, in and adja¬
cent to cranberry beds. Fewer small mammals were present in cran¬
berry beds compared to habitat directly adjacent them (P <
0.0214, 2 dfi. This may have been due to cultural influences,
structural diversity, or predation. A range extension for the arctic
shrew (Sorex arcticus) was documented.
ommercial cranberries are extensively cultivated in Wis-
V^/consin. Cultivation is typified by intensive management
of discrete crop producing complexes within a wetland matrix
(U.S. Army Corps of Engineers 1991). Cranberry growing is
unique, compared to other agricultural practices, because it is
practiced in modified wetlands. Only one study has investi¬
gated small mammals in this unique setting. IEP (1990) stud¬
ied three cranberry production facilities. They used snap traps
but only had five captures (three meadow voles [. Microtus
pennsylvanicus] , and one each of white-footed mouse
[Peromyscus leucopus\ and meadow jumping mouse \Zapus
hudsonius\ , with about 1% trap success.
Small mammal distributions in association with agricultural
practices have seldom been the subject of research. This re¬
search documented the distribution and diversity of mammals
near commercial cranberry beds.
Description of the Study Areas
Five commercial cranberry production facilities in Wood
County (Township of Babcock Sec. 32, T22N, R4E; Township
of Vesper Sec. 13, T22N, R4E; Township of City Point Sec.
19, T21N, R2E), Juneau County (Township of Shennington
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Sec. 17, T18N, R2E), and Portage County
(Township of Dancy Sec. 17, T25N, R7E)
were studied in south-central Wisconsin.
Different wetland habitat types were asso¬
ciated with these facilities including shallow
open water communities, sedge (Carex spp.)
meadows, and sphagnum [Sphagnum spp.)
bogs. All of the commercial cranberry
wetlands studied in this research were
classified as palustrine (Cowardin et al.
1979).
Trapping was conducted in commercial
cranberry beds and adjacent wetlands. Ad¬
jacent wetlands were composed of sedge
meadows and mats, sphagnum communi¬
ties, wet meadows, and lowland forest. A
detailed description of the study areas is
found in Jorgensen (1992).
Methods
Each of the five facilities was sampled over
two periods of two nights each from May
through August, 1991. Each day, 100 snap
traps baited with peanut butter were placed
in 25 identical clusters of four traps
consisting of two Museum Special and two
Victor mouse traps (Call 1986). Clusters
were located using a stratified random
sampling method. Stratification was in three
distance classes, relative to the cranberry bed
matrix: clusters were placed in the cranberry
beds, within 50 m of the cranberry beds and
greater than 100 m from the cranberry bed
matrix. The response variable was the
number of small mammals caught per
cluster per night. Data were analyzed with
a Friedman Test because of non-normality
and heteroscedasticity (unequal variances).
Facilities were blocks and distance classes
were treatments. There were 40 cluster
nights in the 50 m treatment and 30 cluster
nights in each of the other treatments per
facility. Because each cluster night was a
sample and not an experimental unit, this
imbalance only affected the results to the
extent that experimental error was increased,
biasing our results toward nonsignificance.
Means of ranked data within blocks were
separated by the Tukey-Kramer method
(Sokal and Rohlf 1981) for unequal sample
sizes when treatment effects were observed.
Results
Eight species were trapped, and each
facility’s (block) small mammal assemblage
appeared unique (Table 1). Species level
comparisons were not made because sample
sizes were too small to detect differences.
Table 1. Small mammals snap-trapped in and adjacent to commercial cranberry pro¬
duction beds in south-central Wisconsin during 500 trap cluster nights, 1991
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JORGENSEN and NAUMAN: Small mammals and cranberry production
Table 2. Catches of small mammals snap-trapped in and adjacent to commercial cran¬
berry beds in south-central Wisconsin, 1991
aMean separation (a = 0.05, Tukey-Kramertest) on ranks indicates that the number of small mammals captured
was unique at each distance class.
The meadow vole (M. pennsylvanicus) was
the most frequently caught and observed
small mammal. At Wood County no. 2 the
prairie deer mice (P. maniculatus) were of
the subspecies P. m. bairdii. Small mammal
populations were nonrandomly distributed
with respect to distance class (P = 0.0214,
Friedman’s Test, 2 df). Subsequent mean
separation (a = 0.05) indicated that the
number of small mammals caught at each
distance class was distinct, with the fewest
animals captured in the cranberry beds (0.06
catches/cluster night) and the most captured
(0.27 catches/cluster night) within 50 m of
the beds (Table 2). The mean separation test
was exceptionally powerful because of con¬
sistent results in the ranked data across treat¬
ments, though large differences in the
unranked data were observable only with re¬
spect to the cranberry beds themselves
(Table 2).
Catches within the beds totaled six M.
pennsylvanicus , three P. maniculatus , and two
masked shrews (So rex cinereus). We consider
the number of captures too few to allow ac¬
curate analyses of diversity.
Arctic shrews ( S . arcticus) were caught at
Juneau County no. 1 (Township of Shen-
nington, Sec. 17, T18N, R2E) and Wood
County no. 3 (Township of City Point, Sec.
19, T21N, R2E). These records are outside
of the previously documented range for the
arctic shrew in Wisconsin (Jackson 1961; C.
A. Long, pers. comm.), and two specimens
have been placed in the mammal collection
of the Museum of Natural History at the
University of Wisconsin-Stevens Point
(catalogue numbers 7100 and 7101).
Discussion
Small mammals were present in greater
numbers (P = 0.0214) in the semi-natural
habitat outside of the cranberry beds com¬
pared to the beds themselves (Table 2). We
call these areas semi-natural because al¬
though they are not directly modified to
cranberry beds, their close proximity prob¬
ably affects plant distribution (Jorgensen
1992; Jorgensen and Nauman 1994) and
bird distribution (Jorgensen and Nauman
1993).
There were three factors that could con¬
tribute to the distribution of small mammals
we measured. The first factor was the con¬
tinual disturbance that is present in the beds.
Disturbance included various human intru¬
sions and pesticide applications. The second
factor may have been a relative lack of cover
(Wrigley et al. 1979; Reich 1981). There ap¬
peared to be less vertical cover in the beds,
which were maintained as monocultures,
than in the adjacent habitat. Harriers ( Cir¬
cus cyaneus) hunt over the beds. A lack of
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cover might have contributed to increased
predation, or otherwise caused a lack of suit¬
able habitat. The third factor might have
been a lack of diversity in both vegetative
structure and plant species within the beds.
The vegetation is essentially monotypic, and
the insect populations, when they were not
being controlled, were probably monotypic
also and present in low numbers. Interspe¬
cific competition has also been implicated as
a factor affecting small mammal distribution
(Buckner 1966) in wetlands. This study was
not designed to detect this type of interac¬
tion.
Acknowledgments
We thank the Wisconsin State Cranberry
Growers Association and the Natural Re¬
sources Foundation of Wisconsin, Inc. for
funding this research. Thanks to Dr. Charles
Long for verifying identification of the small
mammals.
Works Cited
Buckner, C. H. 1966. Populations and ecologi¬
cal relationships of shrews in tamarack bogs
of southeastern Manitoba. J. Mammal.
47:181-194.
Call, M. W. 1986. Rodents and insectivores. In
Inventory and monitoring of wildlife habitat ,
ed. A. Y. Cooperrider, J. J. Boyd, and H. R.
Stuart, 429-452. U.S. Dept, of the Interior,
Bureau of Land Management.
Cowardin, L. M., V. Carter, F. Golet, and E. T.
LaRue. 1979. Classification of wetlands and
deepwater habitats of the United States. U.S.
Dept, of the Interior, Fish and Wildlife Ser¬
vice. Washington, D.C.: GPO. 131 pp.
Jorgensen, E. E. 1992. Wildlife diversity and
habitat associated with commercial cranberry
production in Wisconsin. MS Thesis, Univ.
of Wisconsin-Stevens Point. 232 pp.
Jorgensen, E. E., and L. E. Nauman. 1993. Bird
distribution in wetlands associated with com¬
mercial cranberry production. Pass. Pigeon
55:289-298.
- . 1994. Disturbance gradients in wet¬
lands associated with commercial cranberry
( Vaccinium macrocarpon ) production in Wis¬
consin. Am. Midi. Nat. 132:152-158.
IEP. 1990. Wildlife utilization and ecological
functions of commercial cranberry wetland
ecosystems. New Hampshire: IEP, Inc. 23
PP-
Jackson, H. T. 1961. Mammals of Wisconsin.
Madison: Univ. ofWisconsin Press. 504 pp.
Reich, L. M. 1981. Microtus pennsylvanicus.
Mammalian Species. Publ. by Am. Soc.
Mammal. 159:1-8.
Sokal, R. R., and F. J. Rohlf. 1981. Biometry.
2nd ed. New York: W. H. Freeman and
Company. 859 pp.
U.S. Army Corps of Engineers. 1991. Draft; St.
Paul district analysis regarding section 404
review of commercial cranberry operations.
St. Paul, MN: U.S. Army Corps of Engineers.
34 pp.
Wrigley, R. E„ J. E. Dubois, and H. W. R.
Copland. 1979. Habitat, abundance, and dis¬
tribution of six species of shrews in Manitoba.
/. Mammal. 60:505-520.
Eric E. Jorgensen is a research assistant at Texas
Tech University. He is investigating small mam¬
mal and reptile associations in the Chihuahuan
Desert. Address : Dept, of Range and Wildlife
Management, Texas Tech Univ., Lubbock, TX
79409
Lyle E. Nauman is a professor of wildlife man¬
agement at the University of Wisconsin-Stevens
Point. In addition to teaching wildlife manage¬
ment courses he advises a number of graduate
students on wetland-related projects.
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Ken Parejko and Douglas Wikum
The effect of manure management on
phosophorus and suspended solids in
the Lake Tainted Wisconsin, watershed
Abstract Phosphorus , suspended solids , and conductivity were measured
during the spring of 1993 at five sites in the Lake Tainter (Dunn
County , Wisconsin) watershed upstream and downstream from
fields spread with turkey litter. Three sites were spread with litter
during the winter , and two sites were spread in the spring after
the ground was thawed . There was no significant difference in
most sites in the phosphorus , suspended solids , and conductivity
upstream versus downstream. Increased solids were detected down¬
stream from two sites during spring-spreading and spring tillage.
Phosphorus concentration showed a highly significant relationship
to suspended solids but was inversely related to conductivity.
Human effects on aquatic ecosystems stem primarily from
point-source and nonpoint-source effluents. Point
sources such as industries and municipal wastewater facilities
are regulated through the permitting process. Nonpoint inputs
are more difficult to regulate or control. Yet in 75% of the
nation’s lakes, measureable improvement in water quality will
only come with control of nonpoint-source impacts (Commit¬
tee on Restoration et al. 1992.)
Tainter Lake, in Dunn County, Wisconsin, is an impound¬
ment of the Red Cedar and Hay Rivers, with a large, predomi¬
nately agricultural watershed of over one million acres. This
lake is experiencing significant cultural eutrophication, prima¬
rily due to phosphorus inputs that contribute to nuisance al¬
gal blooms and consequent undesireable effects. With relatively
large watersheds and a high potential for nutrient loading in
comparison to natural lakes, reservoirs such as Tainter Lake
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are considered especially susceptible to cul¬
tural eutrophication (Baxter 1977; Thorn¬
ton 1984). Schreiber (1992) estimated that
at least 90% of the phosphorus entering
Tainter Lake is nonpoint source in origin.
More than half of the total annual phospho¬
rus loading into Tainter Lake arises during
spring snowmelt runoff (U.S. Geological
Survey 1990). Spring snowmelt also con¬
tributes more than half the total suspended
solids loading into the lake.
Runoff from agricultural sources is sus¬
pected as the major source of phosphorus
loading into the Tainter Lake watershed
(Mechelke et al. 1992). Phosphorus enters
the watershed from barnyard runoff, decay¬
ing vegetation, and runoff of commercial
and manure fertilizers. Although consider¬
able interannual variation may occur due to
the amount of precipitation, the variation in
frost depth, and the timing of manure ap¬
plication relative to snow and rain events,
nutrient runoff studies indicate that total
phosphorus runoff is significantly affected by
several factors: the amount of phosphorus
applied as fertilizer (Coote et al. 1979), the
presence or absence of vegetated buffers be¬
tween the field and the waterway (Thomp¬
son et al. 1979), soil texture and type, slope
(Magette 1988), and winter-spreading ver¬
sus spring-spreading manures (Khaleel et al.
1980.) Though there is little information on
actual nutrient inputs into streams from in¬
dividual fields, studies of intercepted runoff
from experimental plots suggest that manure
applied to frozen ground may have much
more runoff potential than manure spread
on thawed ground and incorporated in a
timely manner (Converse et al. 1975;
Klausner et al. 1976; Minshall et al. 1970;
Mueller et al. 1984; Young and Mutchler
1976.)
Best management agricultural practices,
based on these experimental studies, call for
soil testing for phosphorus and limiting ap¬
plication to necessary amounts; applying
manures in the spring after thaw and incor¬
porating them as soon as possible; and re¬
stricting application of manures to fields
with minimal slopes (Magette 1988). For
example, the spreading of turkey litter from
large turkey farms in the Tainter Lake wa¬
tershed falls under Wisconsin Pollutant Dis¬
charge Elimination System (WPDES) per¬
mits that use these best management prac¬
tices as guidelines for spreading. In a recently
granted permit for one large turkey farm,
farmers who purchase turkey litter from
Jerome Foods of Barron, Wisconsin, are pro¬
hibited from winter-spreading that litter in
several townships in the Tainter Lake water¬
shed.
This study had two purposes. The first
was to test the ability to detect phosphorus
and erosional sediment inputs into water¬
ways from individual fields that have been
winter-spread and spring-spread with turkey
litter. The second was to test for significant
differences in phosphorus and sediment in¬
puts into the rivers, comparing the two ma¬
nure management strategies.
Study Sites and Methods
With the cooperation of Jerome Foods,
Barron, Wisconsin, two fields adjacent to
streams within the watershed that were
spring-spread with turkey litter (sites A and
B) and three fields that had been winter-
spread (sites C, D and E) were selected (Fig.
1). Turkey litter applied at spring-spread
sites was incorporated within three days of
spreading. Sites are characterized as to run¬
off potential in Table 1. Sampling locations
were established just upstream and just
downstream from the field sites. Sampling
began on March 13 and ended on May 6,
1993. During snowmelt runoff and for
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PAREJKO and WIKUM: Manure management in the Lake Tainter Watershed
Fig. 1 . Sampling sites for manure management study
Volume 83 (1995)
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aSoil types: I = Urne Silt Loam; 2 = Plainfield Sandy Loam and Brems Sandy Loam; 3 = Antigo Silt Loam;
4 = Otterholt Silt Loam.
bStreambank = approximate footage of streambank along the field.
cfor Phosphorus Runoff Potential calculation, see text, pp. 95-96.
about one week bracketing the spreading
and incorporation of litter at spring-spread
sites, sampling was done on an approxi¬
mately daily basis. At other times sampling
was done approximately two to three times
per week. For technical reasons, we were
unable to coordinate sampling dates with
precipitation events. Precipitation amounts
at Cedar Falls Dam on the Red Cedar River,
supplied by the Wisconsin State Climatolo¬
gist, and snow depth (measured at each site
on each sampling date and averaged over all
sites) are shown in Figure 2. Dates are con¬
verted to consecutive numerals: March 1 =
1 and May 6 = 67.
At each upstream or downstream location
for each site, samples were taken as a single
grab, about six feet from the stream bank
and just beneath the water’s surface. Aliquots
(100 ml) were acidified, refrigerated up to
28 days, and analyzed for total phosphorus
by the Colfax Commercial Testing Lab,
Colfax, Wisconsin. Phosphorus concentra¬
tions are means of duplicate analyses. The
limit of detection was 0.04 mg/liter total
phosphorus. Suspended solids were deter¬
mined from a one liter sample taken in the
same manner as the phosphorus sample. Sol¬
ids were filtered through a preweighed
Gelman type A/E membrane, dried to con¬
stant weight and reweighed, according to
Standard Methods (APHA 1985). In-lab
measurement of conductivity (umhos/cm)
was performed using a YSI model 33 con-
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PAREJKO and WIKUM: Manure management in the Lake Tainter Watershed
Fig. 2. Snowdepth (cm) and precipitation (in.) vs date
ductivity meter on samples refrigerated at
4°C. All conductivity values were standard¬
ized to 25°C.
Staff gages were installed in the rivers at
downstream sampling locations before the
study. Discharge at staff gages was measured
on four to seven dates during the study with
use of a Marsh McBurney Model 201
flowmeter. Stream cross-section was deter¬
mined by measuring stream depth at 1 m
intervals. Flow was measured at 0.6 x depth
at midpoint of the 1 m intervals, and total
discharge summed over the entire stream
cross-section. A gage versus discharge curve
was developed by plotting log gage depth
versus, log discharge (Chow 1964). Dis¬
charges for dates on which flows were not
measured were interpolated from the gage/
discharge curve. For sites B, C, D, and E
there were several dates when staff gages had
been uprooted by ice or high water and not
yet replaced. For those dates, flags on the
streambank were used to measure stream
height. For a few dates, it was necessary to
interpolate river depths from data from
other sites. Stream discharges are shown in
Figure 3.
To compare sites in their potential for
phosphorus loading, we created a Phospho¬
rus Runoff Potential (PRP). This parameter
for sample sites was calculated as follows:
PRP = kilograms phosphorus applied x ap¬
plication date factor x slope factor x
streambank factor x vegetative buffer factor
x crop factor x tillage factor, where
Application date factor = 1 for manure ap¬
plied under snow, 0.3 for manure applied
on top of snow, and 0.23 for spring-
spread manure (see Thompson et al.
1979);
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Slope factor = 0.25 for slope < 2%, 0.5 for
slope 2%, 0.75 for slope 2-4%, 1 for
slope 4-6%;
Soil factor, all = 1 for soils encountered in
this study;
Streambank factor = number of feet of
streambank/ 2400;
Vegetative buffer factor is an estimate, in¬
terpolated from Thompson et al. 1979,
of the percent of phosphorus reaching the
streambank with various widths of veg¬
etative buffers. For 5 meters, buffer =
35%, for 8 m = 27%, for 12 m = 18%;
Crop factor estimates runoff potential from
different crops onto which manure is ap¬
plied (Thompson et al. 1979). For corn
or soybeans, = 1; for non-alfalfa hay =
0.5;
Tillage factor estimates the effect of tillage
on runoff (from Mueller et al. 1984). For
no tillage = 1; for chisel plow =0.5.
For example, for site A, PRP = 4590 x 0.25
x 0.5 x 1 x 0.42 x 0.35 x 1 x 1 = 84.
The Phosphorus Runoff Potential should
not be viewed as a quantitative estimate of
the amount of phosphorus potentially enter¬
ing the stream. It is instead a numerical
method of comparing the study sites A
through E. Sites with higher PRPs would be
expected to have more nutrient runoff po¬
tential than sites with lower PRPs. Mean
PRP for spring-spread sites is 44; for win¬
ter-spread sites it is 69. These can be con¬
sidered approximately equivalent.
Most-probable-number fecal coliform
analyses were done in the UW-Stout micro¬
biology lab on stream samples taken on one
date during snowmelt runoff and one date
during spring-spread runoff, for all sites. To
estimate what portion of total phosphorus
was attached to particulates, on several dates
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PAREJKO and WIKUM: Manure management in the Lake Tainter Watershed
total phosphorus analyses were done on
samples of unfiltered stream water and on
fdtrates of the same sample that had passed
through a Gelman A/E 1 micron cutoff
membrane.
Statistical analyses were done with
Minitab release 7. 1 . Data were checked for
normality by correlating n scores with raw
data. Correlations routinely fell below criti¬
cal values due to a few very high values
(phosphorus, suspended solids) or low val¬
ues (conductivity) during runoff events. Log
transformation of data gave very high n score
correlations, indicating normality. Thus, we
normalized data using log transformation.
Results
Data for suspended solids, conductivity,
phosphorus, and flow discharge are shown
in Appendices 1—4. In general, we found no
obvious water quality patterns as a result of
runoff from winter-spread versus spring-
spread fields. Using paired t-tests, sites A
through E were tested for a significant dif¬
ference in suspended solids at upstream ver¬
sus downstream locations. Over all 22 sam¬
pling dates, suspended solids were signifi¬
cantly higher downstream than upstream
only at site B (p < .01) and higher down¬
stream at both sites B and E for dates corre¬
sponding to spring-spreading (April 20-May
6). For all other dates and sites, there was
no significant difference (p > .10) in sus¬
pended solids upstream versus downstream
of each field.
Conductivity was lower downstream than
upstream at site B (p <. 001) and higher
downstream than upstream at site E (.01 <
p < .05), over all dates. For dates corre¬
sponding to snowmelt runoff (March 26-
April 17), site B showed lower conductivity
downstream than upstream (.02 < p < .05),
and site E showed higher conductivity
downstream than upstream (.05 < p < .10).
For dates corresponding to spring-spreading,
only site B showed a significant difference
in conductivity, with downstream sites lower
than upstream (p < .01).
Phosphorus concentrations at all sites ex¬
cept site E were not significantly different
between downstream and upstream sam¬
pling locations for all dates, whether winter-
spread dates or spring-spread dates. Site E
had significantly lower phosphorus down¬
stream than upstream (.02 < p < .05), for
snowmelt runoff dates only.
Separate single-factor regressions relating
log-transformed phosphorus concentrations,
conductivity and levels of suspended solids
for each site/sampling location (five sites x
two sampling locations per site) indicated a
significant (p < .01) positive relationship be¬
tween phosphorus concentration and sus¬
pended solids for all ten site locations. Phos¬
phorus was significantly negatively related to
conductivity for site A only (p < .01.) Con¬
ductivity was significantly negatively related
to suspended solids at all sites except site D,
downstream data (all p < .10).
When all sites were regressed together,
phosphorus was significantly positively cor¬
related with suspended solids (p < .001) and
negatively correlated with conductivity (p <
.001.) Conductivity was negatively related to
suspended solids (p < .001.)
Table 2 shows the results from two sites,
on two dates, of phosphorus analyses on fil¬
tered and unfiltered samples. Total soluble
phosphorus in these samples accounted for
44-100% of the total phosphorus in the
sample.
Fecal coliform analysis of stream samples
taken on March 31 (during snowmelt run¬
off) resulted in all ten samples having 100
or more bacteria per ml, with site D, up¬
stream, showing highest coliform levels
(14,000/ml) and site B showing lowest
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coliform (100 and 200 per ml, downstream
and upstream.) Coliform estimates on
samples taken on April 27 (spring-spread)
resulted in site A, upstream, having 100 bac¬
teria/ml; all other sites had less than 100/ml.
Discussion
Highest stream discharges coincided with
the period of most rapid snow melt, with
peak discharges occurring around March 26
(see Figs. 2 and 3). A second discharge peak
occurred on April 8, at a time of relatively
heavy rains. A heavy rainfall on April 30,
however, did not result in significant in¬
creases in stream discharge. Snow depth and
frost depths for the 1992-93 winter were be¬
low normal compared to averages for the
past 20 years, as was precipitation for March
(pers. comm., Wisconsin State Climatolo¬
gist). Precipitation for April and May were
above normal. Total runoff potential for the
spring of 1993, which is directly related to
snow depth and precipitation and inversely
related to frost depth, should therefore be
considered about normal compared to the
past 20 years.
A single sample of surface water runoff,
taken in a part of the field containing tur¬
key litter at site C during snowmelt, was
measured for total phosphorus as 32 mg/1
(data not shown). However, these very high
nutrient levels on-site do not translate into
statistically detectable solids or phosphorus
inputs from individual fields during spring
snowmelt runoff. Snowmelt is a period of
very high discharge and relatively high phos¬
phorus concentrations and levels of sus¬
pended solids. These high levels within the
streams result from the cumulative contri¬
butions of many fields, barnyards, and for¬
ests within the watershed. The incremental
load from an individual field appears to rep¬
resent too small an input, under these con¬
ditions, to be detectable. Fields used in this
study had phosphorus and sediment runoff
ameliorated by relatively gentle slopes, sandy
soils, and the presence of vegetative buffers.
It is possible that under conditions of more
potential runoff, such as fields with higher
slope, more clay, and no buffers, or during
years of abnormally high runoff, increments
from individual fields may be detectable.
During spring-spreading of litter, and dur¬
ing spring tillage, sediment inputs were de¬
tected at two of the sites. At this time, while
fields were being tilled, streams had low dis¬
charge and low sediment loads. Under these
conditions, it was possible to detect sediment
loading from some of the fields.
Conductivity (dissolved ions) was gener¬
ally not significantly different upstream from
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PAREJKO and WIKUM: Manure management in the Lake Tainter Watershed
downstream. One site (site B) showed lower
conductivity downstream than upstream,
and one site (site E) showed higher conduc¬
tivity downstream than upstream, during
snowmelt runoff.
Janzen et al. (1974) measured nutrient
concentrations in small streams in South
Carolina, adjacent to fields spread with dairy
manure. They noted somewhat higher phos¬
phorus concentrations in the streams directly
adjacent to but not 50-600 meters down¬
stream from the fields. In our study, it was
generally not possible to detect phosphorus
inputs from individual fields. At site E and
site C during snowmelt runoff, phosphorus
concentrations were significantly lower
downstream than upstream.
In addition to nutrient input into
streams, animal manure has the potential to
degrade water quality from the input of
coliform bacteria. Robbins et al. (1971)
noted a significant increase in the numbers
of coliforms in water adjacent to fields
spread with animal manure. Janzen et al.
(1974) found similar results. Although we
were not able to detect significant incre¬
ments of bacteria from individual fields, our
results indicate that bacteria from the ma¬
nure are in fact making their way into the
streams. This was especially notable during
snowmelt runoff. Numbers of coliform
dropped below limits of detection after
snowmelt runoff, when suspended solids and
phosphorus concentrations also declined.
Phosphorus entered the study streams in
both the soluble and insoluble forms. An
average of 65% of measured phosphorus re¬
mained in the filtrate as soluble phosphorus
for two dates, shown in Table 2. The por¬
tion of phosphorus as soluble phosphorus
appears, from these results, to vary from site
to site and date to date. The significant posi¬
tive regression between phosphorus and sus¬
pended solids for all sites over all dates
suggests that in general increased erosional
input into streams will increase the input of
phosphorus. Conductivity is determined by
the concentration of total cations and anions
in the stream, which may show a different
mobility in the soil, compared to phospho¬
rus itself (Rodhe 1949). Previous studies
have also demonstrated a negative relation¬
ship between phosphorus and conductivity
(Mueller et al. 1984).
Although this study was not able to pin¬
point phosphorus loading from particular
fields into nearby streams, levels of this nu¬
trient and of erosional sediments within the
streams were seen to increase drastically dur¬
ing snowmelt runoff. The incremental load¬
ing of nutrients from various sources into
the nation’s waterways provides an input
into lakes and reservoirs that causes a signifi¬
cant impact on water quality. Monitoring
streams during snowmelt runoff and during
low-flow, background loading both for to¬
tal phosphorus and total dissolved phospho¬
rus has the potential to provide information
useful in improving nutrient management.
In addition, monitoring phosphorus concen¬
trations in streams during snowmelt can help
identify those subwatersheds that contribute
the most phosphorus to particular lakes or
reservoirs.
Volume 83 (1995)
99
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For dates, 1 = March 1 ; 32 = April 1 ; 62 = May 1 .
100
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PAREJKO and WIKUM: Manure management in the Lake Tainter Watershed
For dates, 1 = March 1; 32 = April 1; 62 = May 1.
Volume 83 (1995)
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Acknowledgments
The authors wish to acknowledge the Uni¬
versity of Wisconsin— Stout Alumni Founda¬
tion for a significant portion of funding of
this study. Additional funds were provided
by the Dunn County Land Conservation
Department through a Lake Improvement
Planning Grant. We gratefully acknowledge
the cooperation and technical assistance pro¬
vided by Jerome Foods, Inc., Barron, Wis¬
consin, and the West Central Office of the
Wisconsin Department of Natural Re¬
sources. Ed Ramsaroop competently and
enthusiastically performed field sampling.
We thank George Nelson, UW-Stout Biol¬
ogy Department, for fecal coliform analyses.
Bill James, Eau Galle Limnological Labora¬
tory, Spring Valley, Wisconsin, and Mike
Nelms, Jerome Foods, made helpful com¬
ments on this manuscript.
Works Cited
American Public Health Association. 1985.
Standard methods for the examination of
water and wastewater. 16th ed. Washing¬
ton, D.C.: APHA.
Baxter, R. M. 1977. Environmental effects
of dams and impoundments. Ann. Rev.
Ecology and Systematics 8:255-283.
Chow, V. 1964. Handbook of applied hydrol¬
ogy. New York: McGraw-Hill.
Committee on Restoration of Aquatic Eco-
systesm, Water Science and Technology
Board, Commission on Geosciences, En¬
vironment and Resources, and National
Research Council. 1992. Restoration of
aquatic ecosystems: Science , technology and
public policy. Washington, D.C.: National
Academy Press.
Converse, J. C., G. D. Bubenzer, and W.
H. Paulson. 1975. Nutrient losses in sur¬
face runoff from winter-spread manure.
Paper No. 75-2035, Am. Soc. Ag. Eng.,
St. Joseph, MI.
Coote, D. R., E. M. MacDonald, and R.
DeHaan. 1979. Relationships between
agricultural land and water quality. In
Best management practices for agriculture
and silviculture , R. C. Loehr et al. eds.
Ann Arbor, MI: Ann Arbor Science.
Janzen, J. J., A. B. Bodine, and L. J. Luszcz.
1974. A survey of effects of animal waste
on stream pollution from selected dairy
farms. ]. Dairy Science 57:260-263.
Khaleel, R., K. R. Reddy, and M. R. Over¬
cast. 1980. Transport of potential pollut¬
ants in runoff water from land areas re¬
ceiving animal wastes: A review. Water
Research 14:421—436.
Klausner, S. D., P. J. Zwerman, and D. F.
Ellis. 1976. Nitrogen and phosphorus
losses from winter disposal of dairy ma¬
nure. ]. Environ. Qual. 5(l):47-49.
Magette, W. L. 1988. Runoff potential from
poultry manure applications. In National
poultry waste management symposium ,
U.S.D.A. and Ohio State Univ., April
18-19, 1988, Columbus, Ohio.
Mechelke, R. 1992. Lake management study
of non-point sources of phosphorus to
Tainter Lake. Dunn Co. Land Conserva¬
tion Dept.
Minitab. 1990. Release 7.1. Minitab, Inc.
3081 Enterprise Dr., State College, PA.
16801.
Minshall, N. E., S. A. Witzel, and M. S.
Nichols. 1970. Stream enrichment from
farm operations. In J. San. Eng. Div .,
Proc. Am. Soc. Civil Eng. April 1970, pp.
513-524.
Mueller, D.H., R.C. Wendt, and T.C.
Daniel. 1984. Phosphorus losses as af¬
fected by tillage and manure application.
Soil Sci. Soc. Am. J. 48: 901-905.
Robbins, J. W. D., G. J. Kriz, and D. H.
Howells. 1971. Quality of effluent from
102
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PAREJKO and WIKUM: Manure management in the Lake Tainter Watershed
farm animal production sites. In Livestock
waste management and pollution abate¬
ment, Proceedings of the International
Symposium on Livestock Wastes. Ameri¬
can Society of Agricultural Engineers
Pub. Proc. 271.
Rodhe, W. 1949. The ionic composition of
lake waters. Verb. Internat. Verein.
Limnol. 10:377-386.
Schreiber, K. 1992. Red Cedar/Tainter Lake
Phosphorus Assessment, Wisconsin DNR
Western District.
Thompson, D. B., T. L. Loudon and J. B.
Gerrish. 1979. Animal manure move¬
ment in winter runoff for different sur¬
face conditions. In Best management prac¬
tices for agriculture and silviculture , R. C.
Loehr et al. eds. Ann Arbor, MI: Ann Ar¬
bor Science.
Thornton, K. W. 1984. Regional compari¬
sons of lakes and reservoirs: geology, cli¬
matology, and morphology. In Lake and
Reservoir Management, Proceedings of the
3rd annual conference North American
Lake Management Society, 18-20 Oct.,
1983, Knoxville, Tennessee. Washington,
D.C.: U. S. Environmental Protection
Agency. EPA 440/3-821-001.
U.S. Geological Survey Water Data Report
WI-90-1. 1990.
Young, R. A. and C. K. Mutchler. 1976.
Pollution potential of manure spread on
frozen ground. J. Environ. Qual. 5(2):
174-179.
Ken Parejko is an Assistant Professor of Biology
at UW— Stout. His research interests include the
effects of human activities on water quality and
genetic diversity of aquatic organisms. Address:
Department of Biology, University of Wisconsin-
Stout, Menomonie, WI 54751
Doug Wikum is an Emeritus Professor of Biol¬
ogy at UW-Stout. He has had a long career of
research, teaching, and service, particularly in
areas related to the impacts of humans on natu¬
ral systems.
Volume 83 (1995)
103
■
Kamela K. Schell and John L. Wedberg
The effect of picnic beetles
(Glischrochilus quadrisignatus)
on European corn borer (Ostrinia
nubilalis) larval mortality
Abstract A study was conducted to ascertain the effects of the picnic beetle
(Glischrochilus quadrisignatus [Coleoptera: Nitidulidae]) on
European corn borer (Ostrinia nubilalis [Lepidoptera: Pyralidae])
larval mortality. The experiment consisted of two treatments: corn
plants with a European corn borer larva only and corn plants with
a European corn borer larva and picnic beetles. Significantly more
corn borer larvae survived in the control than in the treatment
receiving picnic beetles. Picnic beetles caused a 17.5% increase in
corn borer mortality.
The picnic beetle ( Glischrochilus quadrisignatus [Say]),
which is distributed throughout the northern United
States (Luckman 1963), is associated with a variety of foods
including plant sap, fungi, fruits, and vegetables and is often
found in the corn agroecosystem (Luckman 1963; Foott and
Timmins 1971). As McCoy and Brindley noted (1961), the
introduction of the European corn borer (Ostrinia nubilalis
[Hiibner]) to the Midwest provided the picnic beetle with an
additional food source in the form of injured corn plants and
European core borer (hereinafter ECB) frass. The picnic beetle
is primarily saprophagous and feeds on a variety of ferment¬
ing and decomposing plant material. However, on the basis
of early reports by Everly (1938) and Barber and Dicke (1944)
that picnic beetles may be ECB predators, McCoy and Brindley
(1961) investigated the possible reductive effects picnic beetles
have on ECB populations. They reported that during the pe¬
riod of peak beetle populations, the number of dead ECBs and
empty tunnels increased. McCoy and Brindley also reported
that picnic beetles do not actively prey on ECB larvae in the
TRANSACTIONS Volume 83 (1995)
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TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
confines of an ECB tunnel. Rather, it ap¬
peared that the activity of picnic beetles may
cause accidental injury to the ECB larvae,
which the beetles subsequently fed on.
Empty tunnels are a result of picnic beetle
movement irritating ECB larvae so as to
drive them from their tunnels (McCoy and
Brindley 1961). When Carlson and Chiang
(1973) assessed the role of sucrose sprays on
concentrating predatory insects against the
ECB, they found a negative correlation be¬
tween the number of beetles per plant and
number of ECBs per tunnel. They con¬
cluded that picnic beetles reduce ECB popu¬
lations after the larvae have entered the stalk.
The literature supports the idea that picnic
beetles can have a deleterious effect on ECB
populations; however, the evidence has been
circumstantial, and no rigorous study has
been conducted to confirm these effects. The
purpose of this study was to evaluate the ef¬
fects picnic beetles have on a known popu¬
lation of ECBs.
Methods and Materials
This experiment was conducted at the Uni¬
versity of Wisconsin Arlington Agricultural
Research Station in Columbia County dur¬
ing the summer of 1994. The experiment
was a completely randomized design with
two treatments and six replicates. Each plot
consisted of ten corn plants. The Kansas pi¬
pette tip procedure (Higgins, described by
Bode and Calvin 1990) was used to intro¬
duce one-fourth instar ECB larvae into the
stalks of the corn plants on 26 August 1994.
A single larva was placed in the third inter¬
node above the brace roots. The larvae used
in this study were reared from eggs bought
from Dekalb Genetics. When the treatments
were applied, the corn (Pioneer 3731) was
in approximately the R3 milk stage of de¬
velopment, the kernels had yellowed, and
the silks were brown and dry (Ritchie et al.
1982). To facilitate entry of the 1 ml pipette
tip (Fisher Scientific, Reference Tip) into the
corn stalk, a 20 d common framing nail was
driven through a block of wood and used
as a hole punch. The nail was forced into
the stalk to a depth of 2 cm, making it easier
to insert the pipette tips. All plants were
punched, and the stalks were given 1 5 min¬
utes for the flow of plant sap to diminish
before the pipette tips containing ECB lar¬
vae were inserted. This delay was necessary
since it was known from prior experience
that pipettes inserted immediately after the
stalk was punched resulted in high ECB
mortality: the larvae drowned in plant sap
collecting in the pipette tip. The larvae were
brought to the field in pipette tips. The wide
end of the pipette was plugged with clay,
and the tip was plugged with cotton to pre¬
vent the ECB larvae from escaping. The cot¬
ton was removed from the pipette tip when
it was time to insert the ECB larvae into the
corn plants. The larvae were allowed three
days to burrow into the stalks. On 29 Au¬
gust one-half of the stalks were infested with
three picnic beetles per plant via pipette tips.
The picnic beetles used in this study were
collected in banana-baited Lindgren funnels
at the Arlington Agricultural Research Sta¬
tion. Forty-eight hours after the picnic
beetles were introduced into the corn plants,
the stalks were split open and the condition
of the ECB larvae was recorded. The per¬
cent of ECB larvae alive, dead, or missing
was calculated for each treatment, and a chi-
square analysis was used to analyze the data.
Results and Discussion
A significant difference was found between
corn plants containing an ECB larva only
and corn plants containing both an ECB
larva and picnic beetles (X2 = 15.715, P <
106
TRANSACTIONS
SCHELL and WEDBERG: Picnic beetles’ effect on European corn borer larvae
Fig. 1. Percent of European corn borer larvae missing, dead, and alive in cavities con¬
taining European corn borer larvae and picnic beetles and in cavities containing Euro¬
pean corn borer larvae only.
0.001). Of the plants infested with picnic
beetles, 61.4% of the ECB larvae were either
dead (17.5%) or missing (43.9%); by com¬
parison, plants in the control group con¬
tained no dead larvae and 33.3% missing lar¬
vae (Fig. 1). Significantly more ECB larvae
(66.7%) survived in the control than in the
treatment receiving picnic beetles (38.6%).
These data quantify the direct effects picnic
beetles can have on ECB mortality and con¬
firm the earlier suspicions of McCoy and
Brindley (1961) and Carlson and Chiang
(1973). It appears that the death of 17.5%
of the larvae in this experiment probably re¬
sulted from mechanical injury of larvae by
picnic beetles. The inability to observe events
within the plant, however, makes it impos¬
sible to know whether nitidulids are actually
preying on ECB larvae or injuring them via
accidental mechanical damage. Although on
two occasions picnic beetles feeding on ECB
larvae were observed in the field, it is pos¬
sible that the larvae were injured or weak¬
ened prior to picnic beetle feeding. Gener¬
ally, picnic beetles appear to be disinterested
in ECB larvae when the two are placed to¬
gether in petri dishes.
Picnic beetles have been shown to reduce
ECB populations after the larvae have en¬
tered the stalk. However, this reductive ef¬
fect may be more pronounced during the
first generation than the second generation.
Picnic beetles are attracted to and feed on
corn pollen, and second generation ECBs lay
eggs on corn plants that have tasseled and
are near the pollen-shedding stage. Once
pollination begins, picnic beetles may con¬
centrate more heavily where pollen collects
than where ECB larvae and frass collect,
which reduces opportunities for contact with
Volume 83 (1995)
107
TRANSACTIONS of the Wisconsin Academy of Sciences, Arts and Letters
second generation ECB larvae and reduces
their value as a predator.
Although picnic beetles have a reductive
effect on ECB populations, this positive re¬
sult should be weighed against the possible
deleterious effects picnic beetles may have on
the corn system itself. Picnic beetles have
been implicated in the transmission of the
corn fungal pathogens Fusarium spp. (Win-
dels, Windels, and Kommedahl 1976) and
Gibberella zea (Attwater and Busch 1983).
The plant injuries (i.e., ECB cavities and
ECB ear damage) that attract picnic beetles
are excellent entrance points for plant patho¬
gens. The spread of stalk and ear rots may
be exacerbated by movement of picnic
beetles from plant to plant.
Moreover, picnic beetles themselves may
be considered corn pests if their numbers be¬
come heavy. Picnic beetles are considered sec¬
ondary invaders of injured or over-ripe fruits
and vegetables, and a buildup of picnic
beetles may occur anywhere a plant has been
damaged. In the case of silking corn ears,
Luckman (1963) found picnic beetles to be
primary invaders, possibly attracted to pol¬
len fermenting in the silks. A large picnic
beetle population in sweet corn may warrant
control measures where contamination of
processed corn with insect body parts occurs.
Works Cited
Attwater, W. A., and L. V. Busch. 1983. Role
of sap beetle Glischrochilus quadrisignatus in
the epidemiology of giberella corn ear rot.
Can. J. Plant Path. 5:158-163.
Barber, G. W., and F. F. Dicke, 1944. Obser¬
vations on beetles of the family Nitidulidae
in corn plants during 1944. USDA, Bureau
of Entomology and Plant Quarantine, Cereal
and Forage Division R-122, Toledo, Ohio.
Bode, W. M., and D. D. Calvin. 1990. Yield-
loss relationships and economic injury levels
for European corn borer (Lepidoptera:
Pyralidae) populations infesting Pennsylvania
field corn. J. Econ. Entomol. 83:1595-1603.
Carlson, R. E., and H. C. Chiang. 1973. Reduc¬
tion of an Ostrinia nubilalis population by
predatory insects attracted by sucrose sprays.
Entomophaga. 18(2): 205-21 1.
Everly, R. T. 1938. Spiders and insects found
associated with sweet corn with notes on the
food and habits of some species. Ohio Jour.
Sci. 38: 136-148.
Foott, W. FI., and P. R. Timmins. 1971. Im¬
portance of field corn as a reproductive site
for Glischrochilus quadrisignatus (Say) (Co-
leoptera: Nitidulidae). Proc. ent. Soc. Ont.
101:73-75.
Luckman, W. H. 1963. Observations on the bi¬
ology and control of Glischrochilus qua-
drisignatus. J. Econ. Entomol. 56: 681-686.
McCoy, C. E., and T. A. Brindley. 1961. Biol¬
ogy of the fourspotted fungus beetle, Glis¬
chrochilus q. quadrisignatus, and its effect on
European corn borer populations. J. Econ.
Entomol. 54(4): 713-717.
Ritchie, S. W., and J. J. Flanway. 1982. How a
corn plant develops. Special Report No. 48.
Iowa Cooperative Extension Service, Ames,
Iowa.
Windels, C. E., M. B. Windels, and T. Kom¬
medahl. 1976. Association of Fusarium spe¬
cies with picnic beetles on corn ears. Phyto¬
pathology 66:328-331 .
Kamela Schell received her M.S. degree from UW—
Madison, Dept, of Entomology in December 1994.
Address: 59 Manor Lane, East Hampton, NY
11937
John Wedberg is a professor in the Dept, of En¬
tomology, UW— Madison and is currently chair of
the department. His research programs deal with
the development and implementation of pest man¬
agement systems for insect pests.
108
TRANSACTIONS
Wisconsin Academy of Sciences, Arts and Letters
Executive Director
1994 Academy Council
LeRoy R. Lee
Officers
Robert P. Sorensen, President, Madison
OdyJ. Fish, President-Elect, Pewaukee
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Rolf Wegenke, Vice President-Letters, Madison
Gerd H. Zoller, Secretary/Treasurer, Madison
Coun ci lo rs-at-Large
Mary Lynn Donohue, Sheboygan
DeEtte Beilfuss Eager, Evansville
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Judith L. Kuipers, La Crosse
Mildred N. Larson, Eau Claire
Howard Ross, Janesville
Linda Stewart, Milwaukee
Carl A. Weigell, Milwaukee
Councilor- at-Large Emeritus
John W. Thomson, Mt. Horeb
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