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SCIENCES, ARTS AND LETTERS
NATURALE SPECIES RAT«©QUE
Iosval »u?S
VOL. XXIV
MADISON, WISCONSIN
1929
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■£1K. 7 3
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TRANSACTIONS
OF THE
WISCONSIN ACADEMY
OF
SCIENCES, ARTS AND LETTERS
VOL. XXIV
MADISON, WISCONSIN
1929
Volume XXIV of the Transactions of the Wisconsin
Academy of Sciences, Arts and Letters is issued under the
editorial supervision of the Secretary.
Chancey Juday,
Secretary .
CONTENTS
Page
The Romanticism of Edward Young. Harry Hayden
Clark _ _ 1
Wisconsin Indians during the American Revolution.
Louise Phelps Kellogg _ 47
The Settlement and the Distribution of the Population
in Wisconsin. (With Plate 1.) Guy-Harold Smith 53
Glacial Geology of Part of Vilas County, Wisconsin.
F. T. Thwaites _ 109
Chemical Analyses of Lake Deposits. Charles Spur¬
geon Black _ 127
Notes on the Chemical Composition of Some of the
Larger Aquatic Plants of Lake Mendota. III.
Castalia odorata and Najas flexilis. Henry A.
Schuette and Hugo Alder _ _ 135
A Note on the Chemical Composition of Chara from
Green Lake, Wisconsin. Henry A. Schuette
and Hugo Alder _ 141
Life and Letters of Edward Lee Greene. Angie Kum-
lien Main _ 147
Microchemical Tests on the Cell Walls of Certain Fun¬
gi. Cellulose and Chitin. E. W. Hopkins _ 187
The Hepaticae of Wisconsin. George Hall Conklin 197
Preliminary Reports on the Flora of Wisconsin. I.
Norman C. Fassett _ 249
Preliminary Reports on the Flora of Wisconsin. II.
Ericaceae. Norman C. Fassett _ _ _ 257
Notes on Parasitic Fungi in Wisconsin. XV. J. J.
Davis _ 269
Notes on Parasitic Fungi in Wisconsin. XVI. J. J.
Davis _ 279
Notes on Parasitic Fungi in Wisconsin. XVII. J. J.
Davis _ 295
iv
Contents.
Page
A Cytological Study of Fertilization in Achlya hypo-
gyna Coker and Pemberton. (With Plate 2.)
George Olds Cooper _ 303
Cytological Studies on the Sporange Development and
Gametogenesis in Brevilegnia diclina Harvey
(With Plates 3, 4, 5.) George Olds Cooper _ 309
The Cytology and Morphology of Sordaria fimicola Ces.
and De Not. (With Plates 6, 7.) Addie Emma
Piehl _ 323
Some Notes on Allomyces arbuscula Butler (With
Plate 8.) Joseph Henry Lugg _ 343
Preliminary Reports on the Flora of Wisconsin. III.
Lobeliaceae, Campanulaceae, Cucurbitaceae. Ken¬
neth L. Mahoney _ 357
A Study of Two Limestone Quarry Pools. Edward
Joseph Wimmer _ 363
The Morphology and Developmental Stages of a New
Species of Piona. (With Plate 9.) Ruth Mar¬
shall _ 401
A Preliminary List of the Mollusca of Dane County,
Wisconsin. J. P. E. Morrison _ 405
The Hypodermal Glands of the Black Scale, Saissetia
oleae (Bernard) (With Plates 10, 11, 12.) Wm.
S. Marshall _ 427
Preliminary Studies of the Intracellular Symbionts of
Saissetia oleae (Bernard). (With Plates 13, 14,
15. ) A. A. Granovsky _ 445
The Birds of Dane County, Wisconsin. (With Plates
16, 17.) A. W. SCHORGER _ 457
Rainfall Maps of Wisconsin and Adjoining States.
Eric R. Miller _ _ 501
Transmission of Solar Radiation by the Waters of In¬
land Lakes. E. A. Birge and C. Juday _ 509
A Preliminary Report on the Growth of the Rock Bass,
Ambloplites rupestris (Rafinesque), in Two Lakes
of Northern Wisconsin. Stillman Wright _ 581
Proceedings of the Academy - 597
List of Officers and Members - 610
THE ROMANTICISM OF EDWARD YOUNG
Harry Hayden Clark
The object of the present study is to supply a concise and
reasonably comprehensive exposition of Edward Young's
kinship with the Romantic Movement. Although it has
been said that he is “scarcely to be reckoned among the ro¬
manticists/'1 that he made “no real contribution to the new
feeling for nature,"2 and that he belongs to the “traditional
school" of those who “reacted" against early romantic ten¬
dencies,3 recent studies of the rise of isolated romantic traits
suggest that Edward Young is perhaps worthy of renewed
study as an early romanticist. In particular, one striking
paradox appears to have escaped attention : what accounts
for the phenomenal popularity of this supposedly ultra-con¬
servative religious poet in an age of increasing radicalism?
In order to ascertain in what ways Young is romantic I pro¬
pose simply to discover which of his doctrines he held in
common with those later nineteenth century writers gen¬
erally referred to as romanticists. This method will be
safer, I think, than an attempt to measure him by any one
artificial definition of romanticism.4 Since it happens that
some of these doctrines are among the less admirable con¬
tributions of romanticism, occasional references to the
greater figures may seem a bit unsympathetic; it may not
be amiss to remark at the outset, therefore, that our concern
with them is limited to a consideration of points where their
thought chances to be tangent to that of Young, that it is
obviously impossible here to do justice to the many-sided
genius of such men as Wordsworth, Shelley and Keats.
1 H. A. Beers, A History of English Romanticism in the Eighteenth Cen¬
tury , N. Y., 1898, p. 163.
2 Myra Reynolds, The Treatment of Nature in English Poetry, Chicago,
1909, p. 121.
8 Ernest Bernbaum, English Poets of the Eighteenth Century, N. Y., 1918,
p. xxvi.
4 On the ambiguities of the term “romanticism” see A. J. Lovejoy, “The
Discriminations of Romanticisms,” PMLA, XXXIX, 299ff., and Paul Kauf¬
man, “Defining Romanticism : A Survey and a Program,” MLN, XL, No. 4.
2 Wisconsin Academy of Sciences, Arts, and Letters.
I
What first impresses the reader of Young's major work is
his dissatisfaction with actual life. We must remember
that his early life was thoroughly worldly. Although he
seems to have kept his Oxford chambers at All Souls until
he became the rector of Welwyn at the age of forty-seven,
he often attended Addison’s “little senate” at Button’s
coffee-house in Queen Anne’s London; one of the “Wits,”
he was the friend of Steele, Carey, Budgell, Tickell, Pope,
old Cibber and the profligate Wharton. All his early work,
culminating in the seven satires composing The Love of
Fame (1725), was dedicated, with fulsome flattery, to va¬
rious people of high station in the vain hope of obtaining
court preferment. As a dramatist, the author of the three
tragedies Busiris (1719), The Revenge (1721), and The
Brothers (written in 1726, but withheld until 1753), he had
been only moderately successful. He had been defeated as
a candidate for the House of Commons in 1722. 5 Having
exhausted the possibilities of worldly success, he took Holy
Orders in 1727, and in 1731 — after ceaseless efforts — he ac¬
cepted the living of a country clergyman at Welwyn. Am¬
bitious and proud, Young seems to have felt himself un¬
justly neglected; he became a man with a grievance. And
he apparently secured a sort of psychological revenge by
embracing what purports to be a religious other-worldli-
ness. Generally an extremist, he believed a choice must be
made between this world and the “next.”
“Religion’s all. Descending from the skies
To wretched man, the goddess in her left
Holds out this world, and in her right the next.”8
“The Visible and Present are for brutes,
A slender portion, and a narrow bound!
These Reason, with an energy divine,
O’erleaps, and claims the Future and Unseen;
The vast Unseen, the Future fathomless!7
6 H. C. Shelley, Life and Letters of Edward Young , London, 1914. p. 61.
0 Night Thoughts, (hereafter referred to as N. T.) IV, 550ff.
1 N. T., VI, 246-250.
Clark — Romanticism of Edward Young.
3
His attitude toward the here and now is clear; he calls it
this “miry vale”8; “this nest of pains”9; “this dark, incar¬
cerating colony”10; “this night of frailty, change, and
death”* 11; “this dismal scene”12; this “vapour”13; this “pri¬
son”14; this “pestilential earth”15. No wonder he longed to
escape to “that vast Unseen!”
This attitude toward the actual world was the ground¬
work of much of the later romantic idealism. It is inter¬
esting to find it predominating in Germany, — where
Young's influence was strongest— as illustrated in the phan¬
tasies of Tieck and Novalis. Shelley in his Dedication of
The Cenci tells us this “fearful and monstrous” story is
unique among his works in being “a sad reality,” while his
writings “hitherto published” are “dreams of what ought
to be, or what may be.” The arch-romantic American
James Branch Cabell— to whom “veracity is the one unpar¬
donable sin” — “perceives this race . . . to be beyond
all wording petty and ineffectual,” and he follows “ the in¬
stinct of any hurt animal to seek revenge. . . in the
field of imagination” by retreating to Poictesme, “that fair
country . . . which is bounded by Avalon and Phaea-
cia and Sea-coast Bohemia, and the contiguous forests of
Arden and Broceliande, and on the west of course by the
Hesperides.” Keats longs to escape “on the viewless wings
of Poesy” from
“The weariness, the fever and the fret
Here, where men sit and hear each other groan,”
to the
“Charmed magic casements, opening on the foam
Of perilous seas, in faery lands forlorn.”
All this obviously stands in sharp contrast to the vigorous
interest in contemporary problems manifested by the neo¬
classic Defoe, Swift, Addison and Steele. It also contrasts
8 N. T., IV, 537.
9 Ibid., Ill, 409.
wIbid., IV, 665.
11 Ibid., IV, 555.
™Ibid., Ill, 363.
13 Ibid., VIII, 138.
14 Ibid., IX, 1019.
™Ibid., IX, 1352.
4 Wisconsin Academy of Sciences, Arts, and Letters.
with the Greek objective which has been defined as “a seri¬
ous endeavor to understand the world and man, having for
its chief aim the discovery of the right way of life and the
conversion of people to it.”16 The heroes of antiquity were
essentially men of the senses and the understanding.
Dante, on the other hand, records the passing of man from
the world of sense to that of spirit ; in other words his moral
conversion. And in Cervantes there follows what was al¬
most inevitable, — the perpetual contradiction between the
spiritual and the natural man in actual life. Conscious, as
they were, of the gap between the ideal and the real, many
of the romanticists turned their back upon the real and, like
Young, became enamoured of 'The vast Unseen, the Future
fathomless.”
In the case of Young, we are led to seek the cause of his
contempt for the world. In his personal life he met with
considerable grief and disillusionment ; his wife died in the
prime of life, and “he never obtained the preferment of
which he thought himself entitled.”17 However, I am in¬
clined to think that the real source of his disdain for the
“Visible and Present” is to be found in his conception of
original genius, emphasizing, as it does, the quest of novelty.
He has been expostulating with Lorenzo upon the shocking
thought of a continued existence on earth, when he con¬
cludes :
“ . . . With laboring step
To tread our former footsteps? pace the round
Eternal? to climb life’s worn, heavy wheel,
Which draws up nothing new? to beat, and beat
The beaten track? to bid each wretched day
The former mock? to surfeit on the same,
And yawn our joys? or thank a misery
For change, though sad?”18
It is evident here as elsewhere that ennui resulting from
an unsuccessful quest for novelty tends to mould his view
of life : he longs for what is new and distinctive. These
are a few of the reasons why he sought consolation in soli¬
tude.
10 See J. Burnet’s The Legacy of Greece p. 58, where this is said to be the
dominating spirit of Greek philosophy.
17 Sir Leslie Stephen, Dictionary of National Biography , XXI, 1285.
» N. T., Ill, 329-36.
Clark — Romanticism of Edward Young.
5
II
The importance of solitude in the romantic movement has
been stressed many times, and it has been brilliantly dem¬
onstrated that solitude is not far from the core of the whole
romantic philosophy.19 Speaking in social terms, the most
classic person is the Greek citizen in the agora merging
his personality into the life of the state ; the most romantic,
the hermit. It is quite fitting that Petrarch, usually rec¬
ognized as the first modern man, should have been a lover
and praiser of solitude. Wordsworth “wandered lonely as
a cloud” amid the silent scenes of nature ; solitude made him
a poet. The disillusioned author of Alastor fled away from
man and all his works ; he found a symbolic meaning in the
desolate mountain peaks and the open unoared sea. The
insolence of egotism isolated Childe Harold from his fel¬
lows, while across the channel his brother Rene bemoaned
the “moral solitude” so characteristic of romanticism.
In the case of Young’s later life this isolation is marked
indeed. When he praises night he means solitude, for in
his nocturnal habits he finds a double solitude, — both physi¬
cal and moral. The darkness, of course, separates him
from his fellows, but probably he enjoyed much more than
this the thought which he expresses as follows:
“Let Indians and the gay, like Indians, fond
Of feathered fopperies, the sun adore;
Darkness has more divinity for me.”20
He loved to think that there was something distinctive —
and therefore superior— in his midnight watches. Doubt¬
less his desire to be distinctive in this way sprang from his
theory of original genius, again, and suggested his love of
solitude. Furthermore, in his praise of night we have an
example of his choice of melancholy suggestion, — he pre¬
ferred night to day because, among other advantages, it
was more melancholy,
19 Cf. Professor Odell Shepard’s doctoral dissertation on Solitude consid¬
ered as One Phase of the Romantic Movement , Harvard, 1916, to which I
am indebted for several suggestions.
20 N. T.y V, 126-128.
6 Wisconsin Academy of Sciences , Arts, and Letters .
“How like a widow in her weeds, the night,
Amid her glimmering tapers, silent sits!
How sorrowful, how desolate, she weeps
Perpetual dews, and saddens nature's scenes!”21
Two other reasons may be given to account for his love
of solitude, both of which carried much weight with the
later romanticists. First, Young thinks that solitude nour¬
ishes virtue.
“Virtue, for ever frail as fair below,
Her tender nature suffers in the crowd,
Nor touches in the world without a stain;
The world's infectious.”22
Here we have the germs of a doctrine which was later to
deny the struggle between good and evil within the breast
of the individual and assert that every evil thing comes from
without, from social institutions. Young, however, did not
accept any such fallacy. In his love of solitude, however,
he is certainly gravitating toward the later romantic ex¬
treme. The second reason why he is so “studious of se¬
questered scenes” is that he thinks solitude inspires wisdom,
--wisdom of the unearthly, ecstatic sort which ends in mel¬
ancholy.
“0 sacred solitude! divine retreat!
Choice of the prudent! Envy of the great!
By thy pure stream, or in the wavy shade,
We court fair wisdom, that celestial maid . . . ”23
This conception is clearly related to his nature cult which
we shall have occasion to study later. On close examina¬
tion, however, it is evident that this solitary communion
with nature is but a disguised form of his disdain of this
world and an excuse to escape on the viewless wings of his
imagination
“ s . . I bless Night's consecrating shades,
Which to a temple turn a universe,
Fill us with great ideas full of heaven,
And antidote this pestilential earth.”24
81 N. T.j IX, 1978-81.
V, 139-142.
23 Love of Fame, Satire V, 255-58.
™N. T., IX. 1349-53.
Clark — Romanticism of Edward Young.
7
III
Perhaps Young’s conception of the imagination is his most
fruitful contribution to the romantic movement. If one
distinction between the classic and romantic is to be found
in the different role of the imagination in each, then we are
justified in an attempt to determine the precise quality of
Young’s imagination.
In an endeavor to do what is almost impossible, to explain
the classic conception of the imagination, one may refer to
Plato’s somewhat enigmatic theory of the “Ideas”. The
“Idea” was a mental concept which embodied the essence of
the class of sense-objects under consideration; it was the
ultimate reality; it was the perfection, of which all things
perceived by the senses were but imperfect copies. The
object of classic art was to reproduce this “Idea”, to derive
the universal from the particular. It should be observed
that this is a process essentially creative and imaginative;
it is based on actual reality, but is selects only the repre¬
sentative aspects of that reality. The classic artist ex¬
amined many models, selected the one or two more perfect
fragments of each, and then by imagination assembled these
perfect fragments into a perfect whole, a masterpiece of
his imagination ; then, holding this vision of perfection, this
“Idea”, before his mind’s eye, he imitated it with his pen
or brush. A program such as this involves the very highest
exercise of imagination and imitation. It is the imitation
of what has no visible existence ; it is something like trying
to retell our purest dreams. Classic art is ideal; accepting
the view that sense-objects are but faulty copies of the
“Idea”, the artist seeks to correct this imperfection. Some¬
one has said that nature is a great artist whose hand
trembles; classic art, as the neo-classic Sir Joshua Reynolds
puts it, “strives to supply the natural imperfection of
things, and often to gratify the mind by realizing and
embodying what never existed but in the imagination.”25
Of romanticism, one must remember, there are at least
three varieties. The first variety of romanticism- — a roman¬
ticism of action, the Quixotic type — germinated in the
35 Fifteen Discourses on Art.
8 Wisconsin Academy of Sciences, Arts, and Letters.
medieval romances; the imagination sought fictitious ad¬
venture of an extravagant sort, as in Aucassin and Nic-
olette. The Renaissance strove to recover the classical
imagination from the oblivion into which it had fallen, but
the pendulum soon swung again to the second variety of
romanticism, a romanticism of thought, usually associated
with Donne and the “metaphysical school”. Poetry pur¬
sued thought down all its fantastic paths to its furthest
ramifications ; the milestones which marked the limits of the
normal and typical were coolly ignored. This quest of the
singular and novel brought forth from Doctor Johnson the
charge that the “grandeur of generality” was lost, and
straightway the “understanding age” ostracized imagina¬
tion as being germane to the metaphysical malady and
deified reason in the Temple of Taste. The delicate balance
which Aristotle had maintained between imagination and
reason was upset ; his theory of imitation became misinter¬
preted to mean imitation of outer models, and allegiance to
an elaborate system of rules based on these models was
required of all artists. The imagination was discredited
from all sides : Descartes and Bacon had attacked it in the
name of logic ; Spinoza in the name of Stoical reason ; Locke,
Hartley, and Hume in the name of analysis; and Boileau
and Pope attack in the name of good senses. Hobbes
identifies the imagination with the memory of actual outer
images, and so calls it “decaying sense”. An index to the
situation is given us by Addison, in his papers on The
Pleasures of the Imagination when he limits his discussion
to images produced by the sense of sight alone. The point
to observe is that the imagination of the neo-classic period
always clung close to reality — what had existed, or at least
what might exist. Thus there came a revolt — not against
Aristotle and true classicism — but against the false inter¬
preters of classicism. Human nature could not brook hav¬
ing emotion and imagination in such servile bondage to
reason, and a revolt began in what we know as the third va¬
riety of romanticism,— a romanticism of feeling. All three
varieties, let us notice, have a common denominator: the
quest of the strange, the remote, the unique, as opposed to
the normal, the immediate, the representative.
Clark — Romanticism of Edward Young.
9
Among those who helped to liberate the imagination, chief
mention should be made of Akenside whose Pleasures of the
Imagination (begun in 1738, although the first versions
were not published until 1744) supplied aesthetic theory
for romantic practice. Here we find a disdain for the
familiar and the commonplace, in connection with an in¬
definite, aimless, expansive longing, a homesickness for the
infinite.
“Where does the soul
Consent her soaring fancy28 to restrain . . . ?
. . . The rich earth
Cannot contain her; nor the ambient air . . .
. . . The highborn soul
Disdains to rest her heaven-aspiring wing
Beneath its native quarry. Tired of earth
And this diurnal scene, she springs aloft
Through fields of air . . .
Even on the barriers of the world, untired
She meditates the eternal depth below.”
As the eighteenth century wore on even the exponents of
academic culture exalted imagination above reason. As
early as 1756 Joseph Warton startled the strongholds of
criticism with the declaration that the work of Pope was
“not of the most poetic species of poetry . . . because his
imagination was not his predominant talent”. And it is a
striking fact that the first use of the phrase “creative ima¬
gination” occurs in the same author's dedication to the
Essay on the Genius and Writings of Pope.27 The function
of the imagination had changed from that of mental photog¬
raphy based on actual reality to that of creation. And this
early exaltation of the creation imagination was strongly
advocated by Young in 1759. The imagination was fast
drifting away from all reality of the earth, earthy.
“So boundless are the bold excursions of the human mind, that in
the vast void beyond real existence, it can call forth shadowy being
26 Fancy and imagination were used interchangeably in the early eigh¬
teenth century. Addison, Spectator No. 416, remarks, “I shall use them
promiscuously.”
37 ”... A clear head and acute understanding are not sufficient
alone to make a poet ; it is a creative and glowing imagination, acer spiritus
ac vis , and that alone, that can stamp a writer with this exalted and very
uncommon character . . .” Warton’s Essay was dedicated to Young
whose Conjectures were written in 1756.
10 Wisconsin Academy of Sciences , Arts , cmd Letters .
and unknown worlds, as numerous, as bright, and perhaps as last¬
ing, as the stars; such quite-original beauties we may call para¬
disaical.”25
Dull reason was left to crawl abjectly upon the ground;
imagination became a power to soar among the stars, to
create what was “beyond real existence”. If Young’s lit¬
erary dwelling-place was on a water-shed between the coun¬
try of the neo-classic and that of the romantic, certainly in
his conception of the imagination we have a stream of in¬
fluence which flows down into the land of romance, — let us
say of Shelley, or of Tieck and Novalis.29
IV
We now return to the paradox already mentioned, the pa¬
radox regarding the extreme popularity of the reactionary
religious poet in a radical age. Sir Leslie Stephen points
out Young’s conservative trend: “The Night Thoughts , as
he tells us, was intended to supply an omission in Pope’s
Essay on Man . Pope’s deistical position excluded any ref¬
erence to revealed religion, to posthumous rewards and
penalties, and expressed an optimistic philosophy which
ignored the corruption of human nature. Young represents
a partial revolt against the domination of the Pope circle.
He had always been an outsider, and his life at Oxford had,
you may perhaps hope, preserved his orthodoxy.”30 Mr.
H. C. Shelley regards Young, as the author of The Centaur
not Fabulous (1754), as a comrade of both Wesley and
Whiston “in fighting the common enemy of deism,”31 as
represented by the work of Bolingbroke. If there is no
question that he was ostensibly reactionary, there can be no
28 Conjectures on Original Composition, (1759, London, 2nd edition), pp.
70-71.
29 George Brandes ( The Romantic School in Germany , London, 1902, p.
229) writes: “The task of literature in all ages is to give a condensed rep¬
resentation of the life of a people and an age. Romanticism contemptu¬
ously refused this task. Novalis in Germany and Shack Staffeldt in Den¬
mark present the most typical examples of manner in which it turned its
back on reality, to create a poetico-philosophic system out of the mind and
the poetical longing of the author."
80 English Literature and Society in the Eighteenth Century, London, 1904,
p. 152.
81 Life and Letters of Edward Young, London, 1914. P. 234.
Clark — Romanticism of Edward Young .
11
question of his immense popularity. M. Thomas, at the
end of his exhaustive study,32 lists nearly forty editions in
England alone before 1792 containing all or parts of Night
Thoughts . Professor J. L. Kind’s Edward Young in Ger¬
many contains a bibliography covering forty-five pages.
Mr. Shelley says that Night Thoughts “was more frequently
reprinted than probably any other book of the eighteenth
century. It was to be translated into French, and Spanish,
and Italian, and German, and Portuguese, and Swedish, and
Magyar. . .”33
One may explain the popularity of this religious reaction¬
ary in a radical age in at least two ways. Either the age
may have had a deep religious instinct, or his religion may
have contained an alloy which the increasingly radical age
found acceptable. The first alternative appears untenable
when one considers the wholesale lack of spirituality in the
pictures of the later eighteenth century found in Chester¬
field’s Letters to his Son (1773), or in Horace Walpole’s let¬
ters. “There was a revolt,” wrote J. R. Green, the histo¬
rian, “against religion and against the churches in both the
extremes of English society. In the higher circles of society
‘everyone laughs/ said Montesquieu on his visit to England,
‘if one talks of religion.’ Of the prominent clergymen of
the time the greater part were unbelievers in any form of
Christianity, and distinguished for the grossness and im¬
mortality of their lives ... At the other end of the social
scale lay the masses of the poor. They were ignorant and
brutal to a degree which it is hard to conceive.” If we turn
to the second alternative, have we any right to question the
sincerity, or at least the quality of the aspiration, of Young’s
religion? Consider his life in London and his motives for
entering the church. We recall that he did not take Holy
Orders until the age of forty-four, after he was not- — accord¬
ing to Croft — “an ornament to religion and morality.” Mr.
Shelley says of the prime of Young’s life in London, “Noth¬
ing was more improbable at this time than that he should
have become a minister of religion. His chief companions
were the ‘Wits’ of the coffee-houses, the managers and
82 Le PoSte Edward Young , Paris, 1901.
88 Op. cit. p. 198.
12 Wisconsin Academy of Sciences, Arts, and Letters.
players of Drury Lane, and ambitious politicians such as
Wharton and Stanhope.”34 His pleasures were those of “the
Thames and Vauxhall.” Mr. Shelley, who defends Young
at every step, is obliged to admit that Young’s “chief reason
for deciding to enter the clerical profession was a desire
for ... a settled income . . ..”35 After assuming
his sacred responsibilities, he is capable of courting royal
favor through a most surprising letter to the King’s mis¬
tress, Mrs. Howard, in which he parades his “Abilities,”
“Good Manners,” “Service,” “Age,” “Want,” “Suffering,
and Zeal for his Majesty.”36 “There is as little of really
deep sentiment as of sincerity,” concludes Sir Leslie
Stephen ; “for, in fact, Young’s hatred of the world revealed
the disappointed patronage-hunter, rather than the relig¬
ious enthusiast.”37
Not to rely too much on biographical evidence, let us com¬
pare Young’s aspiration with that of other Christian writ¬
ers. St. Augustine testifies that man is restless until he
finds his rest in God. The exiled, aspiring Dante finds that
“In His will is our peace.” Milton, suffering the misery of
blindness alone in the fallen world of Restoration London,
voices not despair but “timeless peace” in that last superb
chorus of Samson Agonistes:
“His servants He, with new acquist
Of true experience from this great event,
With peace and consolation hath dismissed,
And calm of mind, all passion spent.”
The mystic Vaughan aspires to “retreat” from a “heaven
in sense” to “the way, which from this dead and dark abode
leads up to God,” a retreat which gave him peace and calm :
“I saw Eternity the other night,
Like a great ring of pure and endless light,
All calm, as it was bright.”
True Christian aspiration, one may conclude, is definite
and focused: it is based on the imitation of Christ as he
appears revealed in the Scriptures. Instead of exalting
34 Op. cit. P. 62.
36 Op. cit. P. 93.
86 Op. cit . P. 100.
37 English Thought in the Eighteenth Century, London, Vol. II, p. 363.
Clark — Romanticism of Edward Young .
13
desire, it progressively quenches it and leads to “the peace
that passes understanding/' Now let us examine Young's
aspiration in contrast. Is his aspiration definite and fo¬
cused, or is it indefinite and expansive?
“Extended views a narrow mind extend;
Push out its corrugate, expansive make,
Which, ere long, more planets shall embrace.”38
“The soul of man was made to walk the skies;
Delightful outlet to her prison here!
There, disencumber’d from her chains, the ties
Of toys terrestial, she can rove at large;
There freely can respire, dilate, extend,
In full proportion let loose all her powers,
And undeluded, grasp at something great.”39
We shall see later that Young is hostile toward the princi¬
ple of imitation— “that meddling ape Imitation"— which
gives direction and purpose to Christian aspiration. The
next question is whether or not Young's aspiration leads
to calm and peace, or to ennui and restless melancholy. He
envisages “endless joys"—
“Joy breaks, shines, triumphs; ’tis eternal day.
Shall that which rises out of nought complain
Of few evils, paid with endless joys?40
But he has told us earlier of the reaction which is caused by
his celestial reveries :
“Or is it feeble Nature calls me back,
And breaks my spirit into grief again ?
Is it a Stygian vapour in my blood,
A cold, slow puddle creeping through my veins ?
Or is it thus with all men? — Thus with all.
What are we? how unequal! now we soar,
And now we sink. To be the same, transcends
Our present prowess.”41
Is there a relation between his inability to mediate between
extremes and his dejection? We are reminded not so much
of true Christian aspiration as of Shelley's plaintive ques¬
tion addressed to the benign “Spirit of Beauty" :
88 N. T., IX.
39 Ibid., IX, 1018ff.
40 Ibid., IX, 2379ff.
« Ibid., V, 216ff.
2
14 Wisconsin Academy of Sciences , Arts , and Letters .
“Why dost thou pass away and leave our state,
This dim vast vale of tears, vacant and desolate?”
He asks
. . . “why man has such a scope
For love and hate, despondency and hope.”
Thus, whereas true Christian aspiration is definite and fo¬
cused, Young’s is vague and expansive; whereas the Chris¬
tian stresses the principle of imitation, Young rejects it;
and whereas Christian aspiration generally leads to peace,
Young’s led to ennui and despair as we shall see more fully
later.
Perhaps the precise quality of his aspiration will be some¬
what more clearly perceived if we examine it in relation to
the doctrines in the Conjectures on Original Composition.
For, in many ways, the Conjectures is simply a prose am¬
plification of doctrines set forth in Night Thoughts . In
describing his heavenly flights Young exclaims,
“Far from my native element I roam,
In quest of New and Wonderful to man.”42
In the Conjectures his yearning for novelty, expansion, in¬
finitude finds expression through the glorification of an
imagination emancipated from all purpose, restraint, and
reality :
“In the fairy land of fancy genius may wander wild; there it has a
creative power, and may reign arbitrarily over its own empire of
chimeras. The wide field of nature also lies open before it, where it
may range unconfined, make what discoveries it can, and sport with
its infinite objects uncontrolled, painting them as wantonly as it
will.”43
Is this attitude very dissimilar from his so-called religious
aspiration to “disencumber” his soul from toys terrestrial
in order that she may “rove at large” and “grasp at some¬
thing great”? The question now presents itself: Is not
Young’s religious aspiration akin to the later romantic
nostalgia, the quest of Novalis for the Blue Flower, the
quest of Shelley for “something afar from the sphere of our
sorrow”? Dare we suggest that it is the presence of this
43 N. T., IX, 1758-59.
a Conjectures, p. 30.
Clark — Romanticism of Edward Young .
15
indefinite, aimless, infinite expansiveness in the religious
poetry of Young which accounts for his extreme popularity
in an age of increasing radicalism?44
V
At any rate, Young’s conception of the imagination
helped advance the romantic “play-theory” of art. Plato
and Aristotle had exalted the disciplined imagination as a
means of penetrating into the heart of things, a means of
visualizing the eternal realities of life. Dante had used
the imagination to envisage the allotted destiny of human¬
ity. It had enabled Milton “to justify the way of God to
men.” With the incursion of modern science, however, a
different attitude developed : if to Aristotle poetry was the
expression of the highest reality, to Lord Bacon it was an
agreeable lie. And it is not without significance that
Young should have chosen to derive his theories from the
father of modern science, — “Bacon, under the shadow of
whose great name I would shelter my present attempt in
favor of Originals”.45 To Young art became a “consola¬
tion” for the “ tedium vitae Instead of being the su¬
preme work of the human mind, art became the most civil¬
ized form of play.46 The aesthetic imagination was to
“wander wild in its own empire of chimeras,” to “range
unconfined,” to “sport with its infinite objects uncon-
trouled.”
Young’s conception of the imagination passed over to
Germany and quickly merged into the nascent romanticism
of that country. In his Critique of Judgment (1790),
44 The age is not without other examples of expansiveness masquerading
as religion. Akenside remarks, for example, “The sovereign maker
said . . . the soul should . . . enlarge her view/Till every bound
should disappear.” I find that the attitude suggested above has the partial
support of as shrewd a student of religion as George Eliot; see her essay
on “Worldliness and Otherworldliness : the Poet Young,” p. 64, (in Leaves
in a Note-Book, London, 1888) where she scores Young’s so-called religious
quest for “the remote, the vague and the unknown.”
46 Conjectures p. 69.
46 It is true that Young once said that “Judgment . . . should bear the
supreme sway; and a beautiful imagination, as its mistress, should be sub¬
dued to its dominion,” but this was in the essay On Lyric Poetry (1728),
thirty-one years before his final pronouncement in the Conjectures.
16 Wisconsin Academy of Sciences , Arts, and Letters .
Kant47 distinguished between two kinds of imagination:
the scientific, sternly disciplined to reality; and the aesthe¬
tic, emancipated from all restraint whatever. And by free¬
ing speculation from the subjective “forms” of time and
place, he opened up limitless worlds for the conquest of the
imagination. Schiller's novitiate in the school of Kant in¬
spired him with the crusader's zeal to deliver art from the
foils of the scientist ; unwittingly, he betrayed the very cita¬
del of art. In the Letters upon the Aesthetic Education of
Man (1795), Schiller tried to solve the problem of shattered
harmony by saying that only in play is man truly man ;
therefore art must be absolutely free and undisciplined to
reality. Oscar Wilde is only a far-off echo of Schiller’s
Gods of Greece when he declares that the only beautiful
things are those which do not concern us. Here we have
the imagination straining away from a disagreeable reality
towards something which never existed and never can ex¬
ist, — the infinite, aimless longing of the romantic heart; as
Pater was to say, “the Sangreal of an endless pilgrimage”.
I have said that certain of Young's doctrines feed those
streams which ripple most gaily in the sunlight of romance ;
and it is the play genre of imagination which made possible
the most charming of the phantasies of Novalis and the
most exquisite of the fairy tales of Tieck. Nevertheless, it
is a theory which gave presage of a decadent aestheticism,
— of that poisonous heresy “art for art's sake”, the divorce
of art from life. The acme of this heresy is perhaps best
seen in Schopenhauer, by whom, as Dr. Harald Hoffding
says, “the opposition between art and life posited by Kant
and Schiller was ridden to death”. “The value of art would
ultimately disappear”, Dr. Hoffding reminds us, “if there
were really no value in life.48 And of this schism, by virtue
of his disdain for life, his longing for death, his expansive
imagination, and his view of “composition” as a “consola¬
tion” from the ennui of life, Young deserves the somewhat
unenviable honor of being the partial source.
47 For a discussion of the similarities of the views of Young, Herder &
Kant, see Kind, Edward Young in Germany, N. Y., 1906. p. 54.
48 History of Modern Philosophy, London, 1920, Vol. II, p. 234. George
Brandes, (op. cit., p. 75ff) has a whole chapter entitled “Romantic Purpose¬
lessness.”
Clark — Romanticism of Edward Young.
17
VI
An imagination which sports with infinitude must al¬
most necessarily chafe against restriction, and in Young's
revolt against the rules we have an important corollary of
his program of expansion. The school of Pope had deteri¬
orated into the pseudo-classic emphasis on empty forms
and rules; the thoughts and feelings of normal, healthy
men which were not concurrent with these rules were bot¬
tled-up and denied expression in legitimate literature. In¬
deed, we may think of the whole romantic movement as of
a great flood dammed up in the mountains, daily rising and
chafing at its artificial barriers, until, with a mighty surge,
it swept down walls and barricades and fences and rushed
in turmoil to the sea. Thus Young not only marshalled his
invective against imitation and rhyme — that “Gothic de¬
mon" — but also against the rules, which, “like crutches, are
a needful aid to the lame, tho' an impediment to the
strong."49 To Voltaire Shakespeare had been a “drunken
barbarian" chiefly because he ignored the rules; to Young
he was an “adult genius", and ne plus ultra , because, among
other things, he transcended the rules. Inasmuch as the
domain of the rules was to be the battle-ground of the ro¬
manticists for some time, it is interesting to find Young
among the first to challenge them. The most devasting at¬
tack upon the rules, curiously enough, was launched by that
otherwise “eighteen-century" essayist, Macaulay.50 And
Keats used no equivocal terms when he cursed the age
which had been wedded
“To musty laws lined out with wretched rule
And compass vile.”51
VII
The love of infinitude almost inevitably leads to a dis¬
trust of the comprehensive powers of the intellect; limit¬
less worlds are opened which transcend rational knowledge.
48 Conjectures, p. 28. Young’s adoption of blank verse as opposed to the
neo-classic heroic couplet exerted wide influence.
60 In the essay on The Doctrine of “Correctness.”
61 Sleep and Poetry.
18 Wisconsin Academy of Sciences , Arts, and Letters.
As a complement to Young's liberation of the imagination,
it is quite appropriate that he should manifest a notable
tendency, characteristic of the romantic movement gener¬
ally, to disparage learning and logic in the interest of spon¬
taneity and the teaching of nature. Burns, the peasant
poet, exclaims,
“Gie me ae spark o’ nature’s fire!
That’s a’ the learning I desire.”
And Blake was always hostile to any interference with the
natural development of the individual genius. '‘There is
no use in education," he told Crabb Robinson. “I hold it
wrong. It is the great Sin." Rousseau's naturalistic edu¬
cational program in Emile is of course well known. There
is considerable controversy regarding Wordsworth's many
slighting remarks about second-hand learning and the in¬
tellect — “that false secondary power by which we multiply
distinctions" — but it seems reasonable to suppose that he,
like Emerson, preferred to enjoy an “original relation to
the universe." Of course this attitude was more or less of
a revolt from the Cartesian and rationalistic endeavor to
reduce everything to logical formulae. Kant, Hegel, Car¬
lyle, Coleridge, and Emerson — not to mention lesser figures
— exalted Vernunft, while they poured fine scorn upon the
less inspired and more logical faculty, the Ver stand, which
John Locke placed at the head of his corner. And of this
attitude it is interesting to find that Young was one of the
earliest champions, half a century before the great roman¬
tic revolt came into full sway.
Inasmuch as his position has been stated in somewhat
equivocal terms, there is need of making careful distinc¬
tions. Professor M. W. Steinke asserts that “his concep¬
tion of God and man is not mystic, but rationalistic".52
This assertion may be true so far as Young chooses reason
as opposed to sense, since, as he believed, “one, in their
eternal war, must bleed";53 here once more, he is at the
mercy of his fatal inability to mediate between extremes,
to blend and harmonize the physical and the intellectual as
Ha Edward Young’s “ Conjectures on Original Composition ” in England and
Germany (N. V. 1917), p. 13.
»N. T., VIII, 856-62.
Clark — Romanticism of Edward Young.
19
illustrated in the classical exaltation of Mens sana in cor¬
pora sano.
Young’s reliance upon what he calls reason is not so much
a recrudescence as a late survival of the “understanding
age” of Addison and Pope. However, when we subject his
ideas to careful discrimination, it becomes apparent that
they represent a strange fusion — one might better say con¬
fusion — of somewhat incompatible views. In the first
place, if we ask the motives for his choice of reason over
sense, we find that he disowns sense not because of any
Christian hatred of the flesh as the seducer of the spirit
but because the sense can yield only “leaden iteration”,54
or in other words, because it can offer so little novelty of
experience. Once more we have a hint of that ennui so
common in later romanticism; witness Byron and Cha¬
teaubriand. The second motive for his homage to reason
is even more enlightening : he scorns sense because, like all
things terrestrial, it restricts and it confines; he exalts
reason because
“The Present is the scanty realm of Sense;
The Future, Reason’s empire unconfined.”55
Now it is not altogether clear that there is anything very
dissimilar between this “empire unconfined” which is sup¬
posed to be the domain of reason, and his imaginative “em¬
pire of chimeras” where “genius may wander wild” and
“sport with its infinite objects uncontrouled”. Young pro¬
poses reason as a means of restraint for the expansive de¬
sires ; however, it is open to question whether reason, as he
conceives it, is not itself expansive. Certainly these two,
at least, of the motives which determine his choice of rea¬
son over sense are not usually thought of as influencing a
true rationalist. It is clear, I think, that we need to be on
our guard about the way Young uses the word reason, for
his age has a mischievous way of using old words with new
twists in their meanings.
Furthermore, if we examine the quality of his reason
more in detail, we are not without a feeling of paradox ; if
he is a rationalist, it is but fair to add that he disdains all
6*N. T.j III, 373-375.
65 Ibid., VII, 1432-34.
20 Wisconsin Academy of Sciences , Arts, and Letters.
the means by which true rationality is usually acquired.
“Genius needs not go to school.”56 “Many a genius, prob¬
ably there has been, which could neither write nor read
. . .”57 And in the Night Thoughts, again, — although
this particular passage has to do with religion — his respect
for the heritage of the past is representative :
“Wouldst thou on metaphysic’s pinions soar?
Or would thy patience amid logic’s thorns?
Or travel history’s enormous round?
Nature no such hard task enjoins: she gave
A make to man directive of his thought;
A make set upright, pointing to the stars,
As who should say, ‘Read thy chief lesson there’.”58
The reader wonders whether the “hard task” which learning
would enjoin was not influential in suggesting a method so
dear to that majestic indolence of Edward Young. Seldom
has there been a more irrational eulogist of reason. The
relation of his conception of reason to his nature cult is sig¬
nificant, and is a notable instance of a doctrine which was
in great vogue in later romanticism. It would require no
very dangerous stretch of the imagination to attribute the
following sentiment to Wordsworth himself :
“Nature, Thy daughter, ever-changing birth
Of thee the great Immutable, to man
Speaks wisdom; is his oracle supreme;
And he who most consults her is most wise.”59
Although there had been more or less controversy since
the time of Jonson and Shakespeare regarding the relative
merits of culture and native genius, the supporters of cul¬
ture and the classics, such as Dryden, Swift and Pope, gen¬
erally had the upper hand. Young’s Conjectures was a
powerful aid to the group who advocated reliance upon na¬
tive genius. Young insisted that an original “rises spon¬
taneously from the vital root of genius; it grows, it is not
made.”60 Nature, originality, genius, and spontaneity be-
06 N. T., VIII, 830.
57 Conjectures , p. 36.
58 N. T., IX, 866ff. Notice the consonance between the views expressed
in Nigjit Thoughts and in the Conjectures.
69 N. T., VI, 671-74.
60 Conjectures, p. 12.
Clark — Romanticism of Edward Young.
21
came the great watchwords. The best art was supposed to
be a sort of emanation of the folk, — those who lived closest
to the sacred heart of nature. Even as early as 1735
Blackwell's Homer had explained the author of the Odessey
as the result of a “concourse of natural causes." Allan
Ramsay became a shining example of “original genius."
Percy’s Reliques of Ancient Poetry (1765) were collected to
meet the growing demands for folk-songs ; and the tempta¬
tion to which Macpherson yielded had its roots in the same
enthusiasm for the naive and the spontaneous. Somewhat
later children came to be exalted as embodying these quali¬
ties, and we have Blake’s Songs of Innocence and Words¬
worth’s famous glorification of a “six years’ darling" as a
“Mighty prophet! seer blest!" Later, of course, the love
of the spontaneous became related to the primitivistic cult
and the praise of a dim Arcadian past. Following Black¬
well’s interpretation of Homer, it is interesting to find
Young among the first to proclaim the apotheosis of Shake¬
speare as an “original genius," a product of nature. “An
adult genius comes out of nature’s hand, as Pallas out of
Jove’s head, at full growth and mature: Shakespeare’s
genius was of this kind.’’61 Schiller also exalted Homer as
the supreme type of the naive poet : he “is nature.’’62 Of
the apotheosis of Shakespeare, however, Coleridge became
the high priest, assisted by DeQuincey ; the latter brings the
attitude proclaimed by Young to a supreme climax : Shake¬
speare is “like the phenomena of nature," inerrant, inscrut¬
able, transcendent.63 Although Young may be, as Professor
Steinke remarks, averse to “mystic" conceptions, it is evi¬
dent that his conception of the “genius" tends to belittle the
conscious adaptation of means to ends; the genius is like a
magician.
01 Conjectures, p. 31.
62 Essay on Naive and Sentimental Poetry (1795)
63 See the conclusion of “On Knocking- at the Gate in Macbeth” : “O
mighty poet ! Thy works are not as those of other men, simply and merely
great works of art, but are also like the phenomena of nature, like the sun
and the jea, the stars and the flowers, like frost and snow, rain and dew,
hail-storm and thunder, which are to be studied with entire submission of
our own faculties, and in the perfect faith that in them there can be no too
much or too little, nothing useless or inert, but that, the farther we press
our discoveries, the more we shall see proofs of design and self-supporting
arrangement where the careless eye had seen nothing but accident.”
22 Wisconsin Academy of Sciences , Arts, and Letters.
“For what, for the most part, mean we by genius, but the power of
accomplishing great things without the means generally reputed nec¬
essary to the end? A genius differs from a good understanding, as
a magician from a good architect; that raises his structure by means
invisible; this by a skillful use of common tools. Hence genius has
ever been supposed to partake of something divine.”64
VIII
Although Edward Young cannot be credited with having
inspired any detailed, minute, or picturesque description of
nature, his attitude toward nature in general is worthy of
examination. For his doctrine that the genius should rely
upon nature is not altogether unconnected with another doc¬
trine of signal importance, the doctrine that nature is a
manifestation of the deity. In one of his nocturnal rever¬
ies, he exclaims :
“This sacred shade, and solitude, what is it?
‘Tis the felt presence of the deity.”65
In a letter to the Duchess of Portland, 1742, he observes that
“the whole creation preaches ; I mean, that we can make no
just observation on any of the appearances in the material
world, but what will naturally have a good moral effect on
us. The Sacred Scriptures, therefore, are very justly re¬
garded as God Almighty's second volume, and creation as
his first ; which speaks to the same purpose, and if attended
to, is ever bettering the human heart."66 This, surely, is
somewhat radical for the orthodox, who of course regarded
supernatural revelation and the Scriptures as taking prece¬
dence over the volume Young ranks “first." But if Young
was unwittingly tainted by the doctrine of Shaftesbury
that nature is a manifestation of a benevolent deity, he
armed himself resolutely against the Shaftesburian assump¬
tion that the natural man is instinctively good. He refers
many times to “the depravity of our nature," and he once
defines religion as “little more than curbing the natural
84 Conjectures, p. 26. John Foster accused the romanticists of a “viola¬
tion of all the relations between ends and means.” (“On the Epithet Ro¬
mantic,”) Essays (1805-06), Vol. I, (3rd edition).
6B N.T., V, 17 Iff.
68 H. C. Shelley, op. cit ., quoted on p. 153.
Clark — Romanticism of Edward Young.
23
tendencies of our perverse hearts.”67 Perhaps, like Pope,
Young is a little muddled as to the ultimate implications of
his various statements. The nature which is '‘ever better¬
ing the human heart,” although this nature needs "curb¬
ing,”
. . . “is the Almighty’s oath,
In Reason’s court, to silence Unbelief.”68
Proof of a future life is found in the annual rebirth of
nature :
. . . “All, to re-flourish, fades;
As in a wheel, all sinks, to re-ascend.
Emblems of man, who passes, not expires.”69
We have already noticed his theory that nature "speaks
wisdom” ; he remarks elsewhere,
. . . “The stars call thee back;
And, if obeyed, their counsel set thee right.”70
It is scarcely necessary to mention the recognition of the
divine life in nature in romantic literature. Wordsworth
did homage to a god "whose dwelling is the light of setting
suns,” and Shelley proclaimed that "the universe is God”
and "mixed awful talk” with his "great parent.” Coleridge
became a pantheist in The Aeolian Harp and Frost at Mid¬
night. The rebellious Byron learned to "worship Nature
with a thought profound,” his altars being "the mountains
and the ocean, earth, air, stars,— -all that springs from the
great Whole.” Carlyle proclaimed that "The Universe is
but one vast symbol of God,” and his friend Emerson be¬
lieved that man and nature were but parts of that "Unity,
that Over-Soul, within which every man’s particular being is
contained and made one with all other.”
Probably no word in the eighteenth century was used
with a greater variety of meanings than the word "Nature.”
Pope, for example, had advocated that we "first follow Na¬
ture,” but we learn that this is a nature "methodized,” a na¬
ture "which is still the same,” that "Nature and Homer were
91 H. C. Shelley, op. cit., p. 58.
68 N. T., IX, 845ff.
69 N. T., VI, 687.
76 N. T., IX, 641.
24 Wisconsin Academy of Sciences , Arts, and Letters .
. . . the same.” In place of a natural universality we
find arising, along with the original genius theory, a natural
diversity, a glorification of natural idiosyncrasy. Young
opposes imitation because it forces us to “counteract na¬
ture/’ it “blots out nature’s mark of separation.”71 This
doctrine also differs, one should note, from Emerson’s self-
reliance which is, he tells us, reliance on “man’s share of
divinity,” on the universal Over-Soul which makes all men
brothers, the works of all mutually intelligible. The ori¬
ginal genius was supposed to be unique, singular, and indi¬
vidual, especially as regards his knowledge. “Genius,”
Young says, “is from heaven, learning from man. Learn¬
ing is borrowed, genius is knowledge innate, and quite our
our own.”72 Not only is genius a purely individual matter,
but nature itself, Young suggests, is partly a creation of the
individual mind.
“Objects are but th’ occasion; ours th’ exploit;
Ours is the cloth, the pencil, and the paint,
Which nature’s admirable picture draws.”73
This reminds us of Emerson’s Fichtean dictum that “the
Universe is the externalization of the soul,” or — as Profes¬
sor Myra Reynolds remarks — of Coleridge’s Dejection:
An Ode :
0 lady! we receive but what we give
And in our life alone does nature live;
Ours is her wedding garment; ours her shroud.”
IX
Many critics like to think of classicism and romanticism
as of alternate oscillations between collectivism and individ¬
ualism, as an “antithesis between the individual’s claims
and society’s, between synthetic law and individual liberty,
between obedience to restrictions imposed from without and
71 Conjectures , p. 42-43.
72 Conjectures, p. 36. The history of the “original genius” cult has been
brilliantly studied by Professor Paul Kaufman in “Heralds of Original Ge¬
nius,” an essay based on a Harvard doctoral dissertation, in Essays in Mem¬
ory of Barrett Wendell, Cambridge, 1924.
78 N. T., VI, 4 3 Iff.
Clark-— Romanticism of Edward Young.
25
the spiritual demand for inner freedom.”74 Whether or not
we accept this view as the whole truth, we cannot deny that
it represents an important element in the romantic move¬
ment. And it is quite fitting that individualism should be
remarkably salient in Young.75 Inasmuch as Young’s in¬
fluence was felt most strongly in Germany, it may be not
irrelevant to notice that Professor J. G. Robertson finds that
the romanticism— the modernness — of Germany lies in
“the over-weening, even one-sided, individualism of German art; it
deals more persistently and constantly with the individual human soul
than with the external world; it is essentially subjective.”76
It is to be expected from the rest of Young’s philosophy that
he should have gone to one extreme or the other — collectiv¬
ism or individualism — and to be consistent with his other
tenets he could not have chosen the former. As we shall
see, his individualism has an astonishing number of ram¬
ifications, only the more important of which it will be pos¬
sible to trace here.
In the first place the exaltation of the individual goes
hand in hand with Young’s refusal to imitate, for the imita¬
tion of a chosen model, whether literary or human, usually
results not in differences but in likenesses. In accordance
with the romantic edict “Follow Nature,” there was the feel¬
ing that imitation tended to efface the natural individuality
— one might almost say idiosyncrasy — which Young held to
be the inalienable right of every man.
“By a spirit of Imitation we counteract nature, and thwart her de¬
sign. She brings us into the world all Originals: No two faces, no
two minds, are just alike; but all bear nature’s evident mark of sep¬
aration on them. Born Originals, how comes it to pass that we die
copies? That meddling ape Imitation . . . blots out nature’s
mark of separation, cancels her kind intention, destroys all mental
individuality . . .”77
In other words, by making war upon imitation Young hoped
to nourish all those qualities which distinguished him from
74 J. G. Robertson, The Literature of Germany (Home University Library
Edition), p. 142.
75 In his attempt to account for the importance of Young’s Conjectures, M.
Thomas (Op. Cit. p. 484) concludes: “C’est surtout parce qu’il proclame les
droits de l’individualit6 et qu’il insiste sur la notion du ggnie.”
76 Op. Cit., p. 8.
77 Conjectures , p. 42-43.
26 Wisconsin Academy of Sciences, Arts, and Letters.
other men ; he cherished his points of difference, his individ¬
uality. Earlier theorists, advocating the imitation of a def¬
inite standard of excellence, imaginatively conceived, had
naturally stressed the development not of men’s differences
but their likenesses, their common humanity; this concep¬
tion was common to both the humanistic and the Christian
tradition. What, then, is Young’s incentive in refusing to
imitate? Very briefly, one may say it is his desire for su¬
periority ; for he firmly believed that to be different was to
be superior. In the Night Thoughts he imagines himself
as viewing the general race of men from some starry van¬
tage-point, and it is clear that he takes pride in the knowl¬
edge that he is unlike his fellows in all things; as did the
Pharisee, he thanks God that he is not as other men are.
“Earth’s genuine sons, the sceptered, and the slave,
A mingled mob, a wandering herd, he sees,
Bewilder’d in the vale; in all unlike!
His full reverse in all! What higher praise?
What stronger demonstration of the right?
The present all their care, the future his.”78
I am inclined to think that the following sentence from the
Conjectures compresses much of Young’s philosophy into a
nutshell :
“All eminence, and distinction, lies out of the beaten road; and ex¬
cursion, and deviation, are necessary to find it; and the more remote
your path from the highway, the more reputable.”79
78 Night Thoughts, VIII, 1089ff. Young's early insistence upon the merit
of his uniqueness reminds one of Rousseau’s famous words at the beginning
of his Confessions: “Je ne suis fait comme aucun de ceux que j’ai vus ; j’ose
croire n’etre fait comme aucun de ceux qui existent.” Obviously, for this
very reason — so far as it is true — Rousseau cannot serve as an example for
others ; universality, the “grandeur of generality,” is lost. Professor Arthur
Lovejoy ( P’MLA , Vol. XLII, p. 945) presents much scholarly evidence to
support his thesis that “the general transition from universalism to what
may be called diversitarianism” in the eighteenth century was promoted
chiefly “by the emphasis and reiteration given to the principle of plentitude.”
“There has, in the entire history of thought,” he concludes, “been hardly
any change of standards of value more profound and more momentous.”
Coming from such a discriminating scholar, this fundamental distinction
between neo-classicism and romanticism is of considerable interest ; I would
raise the question, however, whether what is called “diversitarianism” was
not promoted as much by the “original genius” cult as by “the principle of
plentitude.”
79 Conjectures, p. 23. It should be remembered that Young is simply the
central figure in a whole group of writers who expressed ideas similar to
his ; witness Sharpe, Duff, Gerard, Colman, Wood, Hurd, Pikerton, Blake
and others.
Clark — Romanticism of Edward Young.
27
Superiority is thus directly proportional to eccentricity.
And the motive for eccentricity— -for deviation from the
beaten road — is clearly the desire for eminence; his whole
program of distinctiveness therefore radiates from egotism,
and is contributory to egotism. Here we have the pivotal
idea in his doctrine of original genius : a deliberate choice
of extremes. This is the unifying centre— the nucleus —
around which his philosophy may be related. It accounts,
in part, at least, for his disdain of the world, “the beaten
road” ; for his love of solitude, “the remote” ; for his love of
“excursion” on the viewless wings of his imagination; and
it is at the core of several more ideas with which I have yet
to deal.
Accepting the desire for superiority as his determining
incentive, Young sees life in terms of individual rivalry.
Thus emulation becomes the key to his relations both with
the ancients and with the moderns. This attitude was
partly responsible for his hostility to the imitation of the
ancients, — ancients and moderns should no longer be con¬
sidered as “masters and pupils, but as hard-matched rivals
for renown . . .”80 “Imitation is inferiority confessed ;
emulation is superiority contested, or denied; imitation is
servile, emulation generous; that fetters, this fires; that
may give a name ; this a name immortal.”81
X
A man who is bent upon surpassing others is apt to con¬
centrate all his energy upon one chosen endeavor. This is
precisely what happened in the case of Young; it is entirely
consistent with his philosophy that he should have glorified
the master-passion, for obviously a man who centers all his
efforts upon one subject has more chance of “distinguish¬
ing” himself therein than the man who seeks a rounded,
symmetrical development. In the race for renown Young
is anxious to lay aside every weight that besets him, and to
focus all his attention on a single point.
“As for a general genius, there is no such thing in nature : A genius
implies the rays of the mind concentr’d and determined to some par-
80 Conjectures, p. 72.
81 Ibid., p. 66.
28 Wisconsin Academy of Sciences, Arts, and Letters.
ticular point; when they are scattered widely they act freely, and
strike not with sufficient force, to fire, or dissolve, the heart.”82
In contrast, we think of the neo-classic Dr. Johnson’s re¬
marks on the master-passion : "This doctrine is in itself
pernicious as well as false. . . True genius is a mind of
large general powers accidentally determined to some par¬
ticular direction.” Johnson was of course simply voicing
Aristotle’s attitude toward specialization and natural bias,
ah attitude motivated by the desire not for progress but
for individual happiness.
“We ought also,” said the Stagirite, “to take into consideration our
own natural bias, which varies in each man’s case, and will be ascer¬
tained from the pleasure and pain arising in us. Furthermore, we
should force ourselves off in a contrary direction, because we shall
find ourselves in the mean after we have removed ourselves for from
the wrong side, exactly as men do in straightening out a crooked
stick.”
Here we have the contrast between the theories rationaliz¬
ing modern specialization and the theories of the advocates
of the golden mean. It would be interesting to ponder upon
the relation between present industrial unrest in our large
cities and the warped, thwarted, and one-sided lives neces¬
sitated by the sub-division of labor of a machine civilization
which exalts material progress above the sacredness of the
individual human personality. And one of the factors be¬
hind this situation is certainly the glorification of the mas¬
ter-passion which evolved out of the theory of "humours” in
the eighteenth century. While the Renaissance, with its
classical heritage, had generally satirized the one-sidedness
of "every man in his humour,” near the end of the seven¬
teenth century we find Congreve tending to defend "hu¬
mours.” A "humour,” he writes Dennis, is "A singular and
unaviodable Manner of doing or saying anything, peculiar
and natural to one Man only ; by which his Speech and Ac¬
tions are distinguished from those of other Men.” He fur¬
ther declares that uHumour is from Nature, Habit from
Custom, and Affectation from Industry.”83 The relation
8a Conjectures, p. 85.
88 The Select Works of Mr. John Dennis, 1718, II, p. 514-525. The pas¬
sage quoted appears in a letter dated July 10, 1695, entitled “Concerning
Humour in Comedy."
Clark— -Romanticism of Edward Young.
29
between the “humour” and nature, and the slur upon In¬
dustry is significant. We cannot stop here to trace the
gradual transition from the theory of humours to that of
the ruling passion, but the relation is clear in Pope :
u . . . One master-passion in the breast,
Like Aaron’s serpent, swallows up the rest . . .
Each vital humor, which should feed the whole,
Soon flows to this, in body and in soul . . .
Nature its mother, habit is its nurse;
Wit, spirit, faculties but make it worse;
Reason itself but gives it edge and power.”
It remained for Young to incorporate the theory of the mas¬
ter-passion into the myriad-sided theory of original genius.
The theory of the master-passion, however, does not de¬
rive entirely from the “humours.” It is vastly significant
that Young confesses himself a disciple of Francis Bacon,
usually considered the father of our modern utilitarian
movement.84 Among the more important bequests of Bacon
to which Young fell heir was the faith in the possibility of
progress, or “perfectibility” as Godwin was to call it later.
He is ever haunted by the analogy between progress in the
world of science and the possibility of such progress in the
world of art. “While arts mechanic are in perpetual prog¬
ress and increase, the liberal are in retrogradation and de¬
cay.”85 And according to Young the source of this dis¬
parity lies in the prevalence of imitation; just as science
questions all authority and refuses to imitate, so ought art
to do. We were still intellectually moored to antiquity till
Young cut the cable and gave us a chance at the dangers
and glories of the blue water. His repudiation of authority
in the literary field soon gave rise to repudiation of author¬
ity in general ; and this break with the past and later with
the traditional standards of goods and evil came simul¬
taneously with the increased self-confidence which science
inspired by its conquest over matter.
The belief in perfectibility— that faith in a mechanical
and single-track advance of humanity — borrowed, as it is,
from modern science, is of signal significance in romanti-
84 “Two words,” said Macaulay, “are the key to the Baconian doctrine —
Utility and Progress.”
85 Conjectures, p. 41.
3
SO Wisconsin Academy of Sciences, Arts, and Letters.
cism.86 Indeed, Professor Barrett Wendell finds that the
only trait common to all the unruly members of the roman¬
tic school was their agreement “in looking forward to an
enfranchised future in which this world was to be far better
than in the tyrant-ridden past. This was the dominant sen¬
timent of English literature from the battle of the Nile to
that of Waterloo/'87 And it is interesting that Young
sounds the clarion of progress long before Godwin.
“Why should it seem altogether impossible that heaven’s latest edition
of the human mind may be the more correct and fair; that the day
may come when the moderns may proudly look back on the compara¬
tive darkness of former ages, on the children of antiquity?”88
It is clear, also, that his faith in perfectibility tempted him
to discard imitation. For imitation
“deprives the liberal and politer arts of an advantage which the me¬
chanic enjoy. In these, men are ever endeavoring to go beyond their
predecessors, in the former to follow them.”89
In his zeal to “go beyond” his predecessors Young re¬
minds us of the strange polarity which exists in the roman¬
tic movement : at one pole we have those who profess sym¬
pathy and brotherly love — Shaftesbury, Rousseau, Tolstoi;
at the other the votaries of the “will to power” — Hobbes,
Manderville, Napoleon, and Nietzsche.90 Here, once more,
we have the fatal weakness of the romanticists, the inability
to mediate between extremes; they are unable to steer a
middle course between Charybdis and Scylla, but must
splinter their keel on one or the other. The question pre¬
sents itself as to which of these extremes Young approaches.
George Eliot, with few exceptions, finds in him “hardly a
trace of human sympathy or of self-forgetfulness, in the
joy or sorrow of a fellow-being.”91 On the other hand, his
38 For Bacon’s belief in “progression” see Advancement of Learning , (Col¬
lected Works), Vol. I, p. 172.
87 A Literary History of America, N. Y., 1900.
88 Conjectures, p. 74.
™ Ibid., p. 41.
"Professor Kaufman (op. cit., p. 216) concludes: “Further developed in
more direct descent, the original genius becomes Carlyle’s hero and culmi¬
nates in Nietzsche’s superman. Applied to the general human level, the
faith in genius as one’s essential self, and hence the complete reliance upon
that self, is the pervasive teaching of Emerson and Whitman.”
81 Op. Git., p. 48.
Clark — Romanticism of Edward Young.
31
guiding motive — the desire for superiority — may be more
concretely described as the desire for fame ; he even went so
far as to call love of fame “The Universal Passion.”
“The love of praise howe’er concealed by art
Reigns more or less, and glows in every heart.”92
Manifestly, this is his principal incentive to literary compo¬
sition. He conceives of the press as “the Fountain of
Fame,” and he objects to imitation because “An imitator
shares his crown, if he has one, with the chosen object of
his imitation; an original enjoys an undivided applause.”93
But what is the ultimate consequence of this passion for
adulation? Toward which of the two poles does Young
tend ? His answer is unmistakable :
“For a man not to grasp at all which is laudable within his reach, is
a dishonor to human nature, and a disobedience to the divine.”94
Thus Young’s emulatory instincts, inflamed by his passion
for “distinction,” cause him to verge perilously near what
we have come to associate with the “will to power.” It is
perhaps worth pondering whether there is not an element of
similarity between Young’s hortatory eulogy on grasping
at all within one’s reach, and — -for example — Fichte’s en¬
deavours to arouse the martial spirit of the German people
by his proclamation of the omnipotence of the expansive
will: “What I will, I can; nothing is impossible.”95 One
need only mention the philosophy of Napoleon, or Nietzsche,
or even Bergson with his “elan vital.” Of course the im-
92 Satire I, 52.
93 Conjectures , p. 12.
94 Ibid p. 77.
95 Reden an die deutsche Nation. It should be recalled that Fichte was
largely responsible for developing the idea of original genius on a national
scale : nations are to cultivate those qualities which distinguish them from
other nations — a theory which leads to an inbreeding of idiosyncrasies. The
Germans were an UrvolJc, the elect not of Calvin’s God but of Rousseau’s na¬
ture ; the possession of character and German descent went together. “Char-
akter haben und deutsch sein ist ohne Zweifel gleichbedeutend . . .”
( Reden an die deutsche Nation , xii). Herder says “An original writer, with
few exceptions, is always a national author.” ( WerTce , Vol. I, p. 402.) J.
L. Kind, (op. cit.3 p. 44) traced Herder’s great “indebtedness to the Conjec¬
tures,” and says he “did some of his best work in this direction, applying
in a practical way to German literature the principles of Young, and thus
passing beyond the arena of speculative thought into the arena of a noble
patriotic cause.”
82 Wisconsin Academy of Sciences , Arts, and Letters.
plicit individualism in Young’s conception of original genius
which had been broadcast through France and Germany by
his Conjectures was powerfully supported by Rousseau’s
doctrine of individualism and by his exaltation of instinct
and emotion above reason. As regards Young, however, we
look for some means of limitation, some curb, for this ten¬
dency to reach out expansively. What is to restrain his de¬
sire to be individual and superior ? Inasmuch as he has os¬
tracised imitation, there is lacking the centripedal force, the
allegiance to a chosen standard, that might prevail against
the powers of self-assertion. It is scarcely necessary to re¬
mark that here we stand at the headwaters of a current
which was to sweep with devastating violence over the war-
torn fields of France in 1917. Once more we catch a
glimpse of the significance of this secluded spring high on
the watershed between the country of Pope and the country
of Rousseau.
XI
One implication of Young’s individualism, often unno¬
ticed, is the trend toward subjectivity, — a pronounced char¬
acteristic of later romanticism.96 By subjectivity here, I
do not mean so much the habit of introspection as the habit
of seeing the outer world in terms of his own mind ; every¬
thing, as we have seen, is painted on the walls of his own
consciousness and is derived from within.
“An inventive genius may safely stay at home. That, like the widow’s
curse, is divinely replenished from within, and affords us a miraculous
delight.”97 '
The view of the classicist, as expressed, for example, by Sir
Joshua Reynolds, is in striking contrast:
“The mind is but a barren soil; a soil which is soon exhausted and
will produce no crop or only one, unless it is enriched with foreign
matter.”98
96 According to James Russell Lowell, the “subjective tendency” . . .
“is one of the main distinctions between ancient and modern poetry.”
( Function of a Poet , Boston, 1920, p. 71.)
97 Conjectures, p. 44-45.
98 Fifteen Discourses on Art.
Clark — Romanticism of Edward Young.
33
In Young's satisfaction with the confines of his own mind
we have something that approaches the solipsism of later
romanticism. All outer reality came to be stained and cob
ored by the romanticist's own temperament ; he came to see
in nature only what he himself put there.
Of course, in the fresh morning of romance Young does
not go to this extreme; but the subjectivity of his literary
method, deriving from his all-embracing conception of orig¬
inal genius, is a straw in the current : “composition” opens
“a backdoor out of the bustle of this busy and idle world,
. . . the key of which is denied to the rest of mankind.”99
Thus the artist is distinctive and unique, and art becomes
the utterance of his own uniqueness. Young's influence
was here definitive: Doctor Johnson had insisted that art
carry a universal import, that it express what is common to
all men, the “grandeur of generality” ; we can perhaps un¬
derstand why the gruff old Londoner called but once on the
clerical recluse when we find Young proclaiming that genius
resides in one's idiosyncrasy, that ineffable something which
makes him unlike any of his fellows. Art thus became
uniquely subjective; the extreme romanticists could see
outer reality only through the tiny windows of their own
souls. And finally all that was left for them to universalize
was their own inner selves, — their dreams and their emo¬
tions. “Eternity,” declared Novalis, “is in us or no¬
where.” A method such as this led to constant introspec¬
tion ; and introspection may easily lead to self-pity and mel¬
ancholy. It is probably of this phase of romanticism that
Goethe was thinking when he made his famous dictum that
the romantic is diseased, the classic healthy. In the famous
first sentence of the Characteristics — “The Healthy know
not their health, but only the sick.”-— Carlyle echoes Goethe
in his judgment on the romantic subjectivity.
XII
Granting that we have now assayed the richest ore of
Young's philosophy of life, the question arises as to its ac¬
tual results. To leave this question unanswered would be
99 Conjectures, p. 5.
34 Wisconsin Academy of Sciences , Arts , and Letters.
to dodge the central current of his influence. For we must
beware of the deceitfulness of abstractions, “the shadows
of a shadow world” ; of these the supreme maxim must ever
be, “By their fruits ye shall know them.” We have no
guide to practical conduct here on earth save the oracles of
experience which direct us by the flickering signals of joy
and despair. If we admit that a man’s fitness for the world
in which he lives and his harmony with the laws of the uni¬
verse are attested by a state not of melancholy but of happi¬
ness, we can find few more serious indictments against a
view of life than that it ultimately leads to melancholy.
Yet this is precisely what happens in the case of Edward
Young; literary explorers have not been altogether wrong
in supposing the secluded spring on the watershed dark and
gloomy. However, his melancholy was not without prece¬
dent, and in order to understand it fully one should have a
general idea of what had gone before.
In glancing at the melancholy which preceded Young, it
is interesting to notice its close relation to solitude. Quite
appropriately, Francis Petrarch, modern in his tastes, is
noted for his melancholy, as one would expect who recalls
his love of solitude, his egoism, his extreme desire for
fame, and his hyper-sensibility. Along with other things
Petrarchan, melancholy was imported by the English Ren¬
aissance. The descriptions by Roger Ascham and Ben Jon-
son indicate that it had become fashionable by the time of
Elizabeth. Shakespeare’s Jacques, an early and striking
study of melancholy, is remarkable for his modern preten¬
sions: his melancholy is different from that of anyone
else, — it is his master-passion, his original genius ; and he is
proud of his excellent and affected differences.100 Richard
Burton, in his esoteric treatment of the subject, notes the
relation of melancholy to solitude, and the fact that it is
often cultivated for its atmosphere of distinction.101 Rep¬
resenting, as he does, the confluence of the Petrarchan and
Elizabethan streams, Burton is said to have suggested102 to
100 As You Like It, IV, Iff.
101 “Nullum solum infelici gratius solitudine, ubi nullus sit qui miseriam
exprobet.” ( Anatomy of Melancholy, Pt. I, Sec. 2, Mem.. 2, Subs. 6.)
102 By the prefatory stanzas to the Anatomy of Melancholy. These were
written about 1601, or twenty years before the completed work.
Clark — Romanticism of Edward Young.
35
Milton two poems of great influence in later romanticism —
U Allegro and II Penseroso. The latter poem, filled with the
spirit of Cambridge and sweet contemplation, represents
melancholy as being induced by solitude.103 After Milton
for nearly a hundred years melancholy was evidently
tabooed; the shadow of the grave and the mystery of the
future were shunned by the rational Augustans. In 1721,
however, Thomas Parnell104 restored melancholy to liter¬
ature, and, quite characteristically, a melancholy closely
related to solitude ; another element also appeared in Parnell
which was significant for later romanticists, — namely the
tendency to ruminate upon the futility of ambition and the
certainty of death. Thus the Petrarchan and the Eliza¬
bethan type of melancholy merge into nascent romanticism
in the early eighteenth century.
Young's Night Thoughts were published in 1742-45, 10 5
and, as we have seen, were greeted with immediate and
unusual popularity; in this love of the lugubrious we have
an index to the taste of an age which turned to questions of
‘‘Night, Death, and Immortality". Although continuing
the literary conventions of the melancholy which had gone
before and which was to become so distinct a note in later
romanticism, Young gives us a still deeper undertone in the
poetry of the graveyard than we find in the literature which
preceded or followed; it was thoroughly characteristic of
Young* that he should have carried his melancholy to ex¬
tremes. Considered in its historical relations, his type of
melancholy represents a transition to the sentimentalism
which resulted from the philosophy of Shaftesbury, of
which Richardson's Clarissa Harloive (1748), 106 Sterne's
Sentimental Journey (1768), and Mackenzie's Man of Feel¬
ing (1771), are examples. In the last book Mr. Henry
Morley find fifty outbursts of tears, and he did not stop to
count the sobs. And with this peculiar dripping senti-
103 The poem is made up of five solitary situations. Melancholy, in the in¬
vocation, is represented as the daughter of Solitude and Purity, or perhaps
of Solitude and Genius.
104 In A Hymn to Contentment and A Night Piece on Death.
105 In regard to the date see The Review of English Studies , Vol. IV, p. 300
and pps. 414-417.
106 A warm friendship existed between Richardson and Young, and the lat¬
ter was loud in his praise of his friend’s work.
36 Wisconsin Academy of Sciences, Arts, and Letters.
mentality Young was by no means unsympathetic; in his
eulogy upon that Christian paragon, Joseph Addison, he
finds but one fault: too much of a “philosophic reserve";
“he should have let loose all his fire, and have showed the
most tender sensibilities of his heart".107 In general, how¬
ever, Young's complaint is not so much what we recognize
today as sentimentality, as a sombre melancholy, or self-
pity. Of course tumid Edward Young is not exactly the
sort of man we should think of as indulging in endless
jollifications; making allowance, however, for his constitu¬
tional aversion to unseemly merriment, it is readily ap¬
parent that he lacks that deep spiritual health which, for
want of a better name, we call happiness. As Sir Leslie
Stephen observes, “Indeed, if Young is not capable of a
noble melancholy, he is in a thoroughly bilious condition."108
Inasmuch as external events frequently have a disagree¬
able way of altering the course of the finest philosophy,
one ought to glance at Young's life before laying the
undivided blame for his melancholy at the door of his
romantic philosophy ; his life was not without causes for
unhappiness. A study of Young's biography leads to the
conclusion that a natural predisposition to depression, the
complement of his subjective and egotistical nature, was
aggravated by sorrow and worldly disappointment. We are
told that in those formative and fruitful years of college
life he preferred to compose after midnight by the light of
a candle stuck in a skull. It was just after college that
Young’s thoughts were turned toward death and
“The grave, his subterranean road to bliss”,109
by the death of William Harrison with whom he enjoyed a
friendship not unlike that between West and Gray or that
between Hallam and Tennyson. In The Epistle to Lord
Lansdowne, (1713), Young laments for his friend, “the
partner of his soul".110 Notwithstanding repeated efforts to
win fame and preferment, “in his thirty-fifth year, — his
107 Conjectures, p. 83.
108 English Thought in the Eighteenth Century, London, Vol. II, p. 364.
189 N. T., VII, 10.
110 Line 509.
Clark — Romanticism of Edward Young.
37
prospects were no higher than in 1708”.111 Later, to be sure,
with the publication of Night Thoughts he enjoyed no incon¬
siderable fame, but according to Sir Leslie Stephen he “never
obtained the preferment to which he thought himself en¬
titled”, as we have seen. If we recall that his lode-star was
fame, we can perhaps imagine how this reception affected
him. To return to his early work, there is little of impor¬
tance to be noted, save that as M. Thomas has pointed out,
even in his youthful days he was “le poete de la pensee melan-
cholique”.112 Doubtless at first Young took delight in his
melancholy; real sorrows are decidedly uncomfortable, but
surely aesthetic ones are by no means unpleasant. Youthful
melancholy is as normal as dentition; Young, however,
seems to have been abnormally addicted to it, and as his
sense gradually deadened he increased the dose until he
ended in relatively genuine gloom. At first the debaucher,
he became the victim of his sensations. In 1727 he aban¬
doned the career of playwright, rewarded with meagre
success, for the church; in 1730, at the age of forty-seven
as we have seen, he was appointed Rector of Welwyn, where
he retired, bitter at his failure to win the attention he
thought he deserved, and henceforth almost as a theatrical
troglodyte he “peeped at the world through the loopholes
of retreat”. Doubtless his priestly duties of comforting
those who mourned, and his preoccupation with religious
and moral treatises, tended to make his life more sombre.
In 1741 his wife died, his step-daughter having died in
1736, and her husband in 1740. That he was not insensible
to these events we may judge from the words of Mr.
Shelley :
“Of all these losses, the one which affected him the most deeply was
that of his wife; that was the culminating burden of his sorrows,
having issue in sleepless nights and melancholy days.”113
Furthermore, there can be little doubt that his depression
was increased by his solitary habits and by the surround-
111 H. C. Shelley, op. cit., p. 40.
112 w. Thomas, op. cit., p. 316. In The Last Day (1714) Young says his
muse is a “Melancholy Maid” whom “dismal scenes delight.”
113 1 have corrected here a regrettable error made in my paper entitled “A
Study of Melancholy in Edward Young,” (MLN, XXXIX Nos. 3 and 4) — of
which the present study is an expansion — in citing from Mr. Shelley’s
valuable biography the date of Mrs. Young’s death.
38 Wisconsin Academy of Sciences, Arts, and Letters.
ings which he chose to remind him of the transitory char¬
acter of human life; he displays a marvelous ability to
extract the essence of melancholy from external objects.
It is unnecessary to repeat the story of the alcove in his
garden where he had a bench painted to give the illusion of
reality, bearing the words “Invisibilia non decipiunt”, or
the fate of the sundial inscribed “Eheu fugaces!” His last
years were lonely, — and melancholy.114 “While his health
permitted him to walk abroad, he preferred a solitary
ramble in his churchyard to exercise with a companion on
a more cheerful spot”.115 It was thus almost inevitable that
pondering upon graveyards should lead to reflections upon
the futility of ambition and universal mutability. His
“first contribution to theology” — addressed to Queen Caro¬
line — is supposed to be an argument against pessimism, but
the reader can judge for himself as to its cheering tone;
he has been considering all the passions of humanity as
“contributing to human misery”:
“If this account is just, as I think it is, what is human happiness?
A word, a notion, a day-dream, a wish, a sigh, a theme to be talked
of, a mark to be shot at, but never hit, a picture in the head, and a
pang in the heart of man! Wisdom recommends it gravely, learning
talks of it pompously, our understanding listens to it eagerly, our
affection pursues it warmly, and our experience despairs of it
irretrievably.”116
Nor is his final poem, Resignation (1762), much more
exhilarating :
“Of earth’s revenue would you state
A full account and fair?
We hope; and hope; and hope; then cast
The total up — Despair
It is true, of course, that he expressed at times the hope
of “endless joys” in the hereafter, but we have seen that
the expansive element in his religion led to atrabilious
reactions.
114 Sir Leslie Stephen, D. N. B., Vol. XXI, 1286, “Young’s last years were
melancholy.”
115 Rev. J. Mitford, “Life of Young,” lvil, in The Poetical Works of Young,
(Boston, 1854).
116 Quoted by Shelley, op. cit p. 98, from “A Vindication of Providence”
(1727).
Clark — Romanticism of Edward Young.
39
However, in exposing possible sources of melancholy in
the poet's life, as I have done, there is the danger of giving
a distorted impression. To outward appearance his life
was not as mournful as I may have suggested. Mr. Shelley
very properly stresses the pleasanter side of Young’s life,
especially the fact that he entertained guests occasionally
and could be witty in society. The truth of the matter is
probably given in the words of his own son :
“He was too well-bred a man not to be cheerful in company, but he
was gloomy when alone; he was never cheerful after my mother’s
death, and he had met with many disappointments.”117
We have already seen that he preferred solitude, — not so
much, perhaps, a physical as a psychic solitude due to a
certain intellectual isolation and uniqueness. However, not
to mention the good fortunes which were Young’s, countless
people — such as Milton, for example — have suffered far
greater sorrows than his and have lost neither their cheer¬
fulness nor their serenity of spirit. A more profound cause
for his melancholy must be sought, therefore, and for that
purpose we turn to his philosophy of life.
True happiness, it is probably safe to say, results chiefly
from a mediation between extremes. The great problem of
man is one of adjustment and compromise. At least this
was the teaching of Aristotle and, in part, of Plato. This
was the essence of the Horacian doctrine of the golden
mean — auream mediocritatem — and Cicero’s praise of me¬
dio critatem illam quae est inter nimium et parum. Mr.
G. K. Chesterton has recently advanced the thesis that
Christianity is peculiar in reconciling opposites, in its
balance. The classical and Christian traditions, at any
rate, unite in stressing the doctrines of imitation, restraint,
and a definitely focused aspiration. In the Middle Ages the
“law for measure” was allied to the theory of the four
humors, the balance of which meant health, the excess of
any one of which accounted for a man’s one-sidedness and
meant disease. It is a commonplace, of course, that in the
Renaissance— the rebirth of classicism — Ben Jonson satir¬
ized “Everyman in his Humor,” and the tragic heroes of
Marlowe and Shakespeare met their doom largely through
117 Rev. J. Mitford, “Life of Young,” lxi, in The Poetical Works of Young,
(Boston, 1854).
40 Wisconsin Academy of Sciences , Arts, and Letters.
some one weakness or one-sidedness. Hamlet, perhaps
Shakespeare's greatest creation, knows that those who, “by
the o'ergrowth of some complexion,” carry
. . . “the stamp of one defect, . . .
Shall in the general censure take corruption
From that particular fault.”
And of Horatio, his soul's choice, he exclaims,
. . . “blest are those
Whose blood and judgment are so well commingl’d.”
The Renaissance, however, gave birth to Bacon and mod¬
ern science with its faith in progress and specialization;
later the Royal Society lauded the virtuosi, men of sur¬
passing acquisition in some one field.
Now balance and poise are generally the result of the
disciplined and purposeful imitation of a balanced ideal en¬
visaged by the imagination. Edward Young, by opposing
imitation, by chartering of the imagination as a vehicle of
idyllic escape, by translating the theory of humors — with
the help of Bacon — into that of the master-passion, failed to
mediate between extremes. Indeed, it is worth consider¬
ing whether it was not precisely a failure of this sort
which underlies most of his much-discussed melancholy;118
for, if this study has served its purpose, it has indicated
that his whole philosophy may be grouped around a choice
of extremes as a nucleus. His scorn for the actual world
and his devotion to the “vast Unseen''; his love of a dis¬
tinctive solitude; his emancipation of the imagination; his
expansiveness posing as religious aspiration; his rejection
of the wisdom of the past and his preference for a unique
original genius ; his reliance upon the teaching of nature ; his
scorn for imitation and his insurgent individualism; his
praise of the master-passion and the faith in progress ; his
singular subjectivity — all these have a common denomi¬
nator: the choice of an extreme. I have endeavored else¬
where to demonstrate the relationship between Young’s
extremism and his melancholy.
118 See Amy Louise Reed, The Background of Gray’s Elegy, N. Y. 1924, p.
192ff. This is a scholarly study of the taste for melancholy poetry in the
first half of the eighteenth century.
Clark — Romanticism of Edward Young.
41
Perhaps the most important thing about Young’s mel¬
ancholy historically is its personal, subjective character.
Classical and neo-classical melancholy had been in general
objective and impersonal. Vergil’s “majestic sadness,” in
Tennyson’s phrase, resulted from the “doubtful doom of
human kind.” Gray brooded on the fact that “The paths of
glory lead but to the grave.” But Young, like Byron, is not
averse to bearing the pageant of his own bleeding heart
before the world.
“From short (as usual) and disturbed repose,
I wake: (how happy they, who wake no more!
Yet that were vain, if dreams infest the grave.
I wake, emerging from a sea of dreams
Tumultuous; where my wrecked desponding thought,
From wave to wave of fancied misery,
At random drove, her helm of reason lost.
Though now restored, ’tis only change of pain,
(A bitter change!) severer for severe.
The day too short for my distress; and night,
Even in the zenith of her dark domain,
Is sunshine to the colour of my fate.”119
This introspective tendency has many sources. In addi¬
tion to the expansiveness of Young’s religion which casts a
dark shadow on this world where, as Wordsworth said,
“we find our happiness or not at all,”120 his religion has a
strong taint of egoism, as apparent in his summary :
“In self-applause is Virtue’s golden prize.”121
“Virtue is true self-interest pursued.”122
We have already noted that his whole program of dis¬
tinctiveness is motivated by the passion for “eminence.”
Another consequence of his theory of original genius is the
impulse to introspection, to an escape into one’s self. He
loves night because
“It strikes thought inward; it drives back the soul
To settle on herself, our point supreme.”123
119 N. T., I, 5 ff.
120 Prelude, XI. Oliver Elton ( A Survey of English Literature 1780-1880,
London, 1912, Vol. II, p. 95) finds Wordsworth practically the only romantic
poet free from melancholy.
™N. T., VII, 148.
122 N. T.y VII, 143.
123 N. T.} V, 129ff.
42 Wisconsin Academy of Sciences, Arts, and Letters.
In the Conjectures “self-knowledge” and “self -reverence”
are exalted as the supreme canons.
“Therefore dive deep into thy bosom; learn the extent, bias, and full
fort of thy mind; contract full intimacy with the stranger within
thee. . . . Thyself so reverence, as to prefer the native growth
of thy own mind to the richest import from abroad.”124
Of course genuine sorrow — and much of Young's was gen¬
uine — commands respect. Do we respect, however, a sor¬
row which becomes a mark of superiority? “His grief,”
he tells us, “is but his grandeur in disguise.”125 Thus the
personal quality of his melancholy derives in part, at least,
from his myriad-sided theory of original genius. For
Young, believing that to be superior he must be unique,
becomes unique in feeling, and therefore in suffering, evi¬
dently following the generalization made by Walpole that
life is a comedy for those who think and a tragedy for
those who feel. On one hand, the distinctive quality of his
melancholy ministers to his egotism ;126 it distinguishes
him from the “mingled mob” and therefore makes him su¬
perior. On the other hand, the uniqueness of his melan¬
choly gives rise to what M. Thomas calls “un cri d’ an-
goisse tout personnel. Ce qui manque a ces effusions pathe-
tiques, ce n’est pas la sincerity mais Tuniversalite de 1’
emotion.”127 And there lies a deep significance in the fact
that, after the somewhat artificial objectivity of the neo¬
classic age, the return of “I” to literature — the subjective,
introspective, confessional note — should be inseparably
linked with melancholy.128
124 Conjectures, p. 24.
125 N. T., V, 558ff.
126 George Brandes {op. cit., p. 78) speaks of “the aesthetic stupor of self-
contemplation and self-absorption which was the final development of Ro¬
manticism” in Germany.
127 Op. cit., p. 458. One exception is noted, N. T., I, 238. It seems to me
that there are more exceptions. In regard to the charge' of hypocrisy in
connection with Young’s melancholy, M. Thomas says, “Cette supposition est
non moins injurieuse que peu vraisemblable.”
128 J. L. Kind (op. cit., p. 158) points out that “Herder discusses melancholy
poets of the day, and places Young in the first rank.” Goethe learned Eng¬
lish from Milton and Young, and in Aus Meinem Leben (Bk. XIII) he
“characterizes the effect that the gloomy English poets had upon Germany
at that time, and attributes to them, mentioning the Night Thoughts as the
work in which the pessimistic weariness of life is preeminently worked out,
the conditions that caused Werther to strike a responsive chord elsewhere.”
Clark — Romanticism of Edward Young.
43
Of course Young’s melancholy is but a foreshadowing of
the pervasive melancholy which one finds everywhere in
the romantic movement. Although “felicity is the great
fact of Wordsworth’s life,”129 even this serenest of the ro¬
manticists felt his anchor drag at times, as Ruskin noted:
“Great God,” the poet exclaims,
“I’d rather be
A Pagan suckled in a creed outworn;
So might I, standing on this pleasant lea,
Have glimpses that would make me less forlorn.”
Everyone remembers Coleridge’s painful confession, in
Dejection : An Ode , of
“A grief without a pang, void, dark, and drear,
A stifled, drowsy, unimpassioned grief,
Which finds no natural outlet, no relief,
In word, or sigh, or tear.”
John Keats forever lamented the transient loveliness of a
sensuous world
. . . “where men sit and hear each other groan;
Where palsy shakes a few, sad, last gray hairs,
Where youth grows pale, and spectre-thin, and dies;
Where but to think is to be full of sorrow
And leaden-eyed despairs.”
“I fall upon the thorns of life! I bleed!” Shelley confides.
In the Stanzas Written in Dejection his despair is laid
bare :
“Alas! I have nor hope nor health,
Nor peace within nor calm around,
Nor that content surpassing wealth
The sage in meditation found,
And walked with inward glory crowned —
Nor fame, nor power, nor love nor leisure.
“Yet now despair itself is mild,
Even as the winds and waters are;
I could lie down like a tired child,
And weep away the life of care. ...”
(Kind, op. cit., p. 109) In France', according- to E. Baldensperger, (“Young
et ses ‘Nuits’ en France,” in studies d’Histoire litteraire) Young was one of
the major influences behind the vogue of melancholy literature.
129 F. W. H. Meyers, Wordsworth, London, 1925, p. 72.
44 Wisconsin Academy of Sciences , Arts , and Letters.
Lord Byron tells us that, “grown aged in this world of
woe,” he writes Chiide Harold to “fling forgetfulness
around me,”130 and the Byronic egotist Manfred — with all
things offered him — prays only for “Self-oblivion.” And
at the end, in Don Juan , he confesses with infinite sadness,
“And if I laugh at any mortal thing,
'Tis that I may not weep; and if I weep,
Tis that our nature cannot always bring
Itself to apathy.”
We may safely conclude, then, that the shadows which
stole upon the author of Night Thoughts were but early
adumbrations of those which were to fall upon later ro¬
manticists.131
Summary
We have discussed the following major traits appearing
in Young's work:
(1) scorn for the commonplace and the actual world;
(2) praise of a solitude unique and distinctive;
(3) apotheosis of the lawless, creative, idyllic imagina¬
tion;
(4) indeterminate expansiveness in the guise of religious
aspiration ;
(5) praise of art as a means of play and of escape;
(6) contempt for rules and restrictions ;
(7) preference for native genius rather than culture and
the classics.
(8) recognition of nature as “the felt presence of the
deity;”
130 Young (N. T., I, 45) tells us, somewhat as Byron does, that his is “A
mind that fain would wander from its woe.”
131 A host of thinkers and scholars testifies to the pervasive melancholy of
the romantic movement and the nineteenth century. See, for example, the
following: J. R. Lowell, Prose Works, Boston, 1894, Vol. Ill, p. 94; John
Ruskin, Selections and Essays 1y Buskin, edited by F. W. Roe, N. Y., 1918,
p. 134 ; Walt Whitman, quoted by N. Foerster, American Criticism, Boston,
1928, p. 191; Thomas Hardy, Tess of the D'Uhervilles, N. Y. 1921, p. 132;
Arnold, Passim ; Walter Pater, Appreciations, London, 1889, p. 105; Bourget,
Essais de Psychologie contemporaine, in the essay on Flaubert ; H. A. Beers,
A History of English Romanticism in the Eighteenth Century, N. Y., 1916,
p. 115 (Victor Hugo is here said to have called melancholy the distinguish¬
ing badge of romantic art) ; Irving Babbitt, Rousseau and Romanticism,
Boston, 1917, p. 307; Oliver Elton, Op. cit., Vol. II, p. 95.
Clark — Romanticism of Edward Young .
45
(9) hostility to imitation and praise of militant individ¬
ualism ;
(10) glorification of the master-passion and the hope of
progress ;
(11) praise of a unique and idiosyncratic subjectivity ;
(12) the parading of a personal and singular melancholy.
While one may concede that these traits are not always the
major traits of the greater romanticists, I have tried to
show that all of these traits are manifested in one way or
another in the literature which by common consent we call
romantic.
Note: — In accordance with the terms of the award, statement is
here made of the fact that this study— in slightly different form —
was awarded the Bowdoin Prize in English in the Graduate School of
Harvard University for the year 1924. I wish to record my apprecia¬
tion of the kind suggestions of Professors Irving Babbitt and John L.
Lowes, under whose direction the study was originally begun, as well
as those of Professor Henry B. Lathrop of the University of Wiscon¬
sin. H. H. C.
The University of Wisconsin
January 10, 1929.
4
WISCONSIN INDIANS DURING THE AMERICAN
REVOLUTION
Louise Phelps Kellogg
When the Revolution began, Wisconsin had been for only
fourteen years under British sovereignty and the Indians
were not entirely reconciled to the officials sent to preserve
order in this distant part of the British Empire. They had
been restless from the moment a British force had advanced
from Mackinac and had taken possession of the old French
post at Green Bay. The Maryland officer who was sent to
pacify these more than thirty thousand savages, dependent
upon his post, was appalled at his task. However, he suc¬
ceeded better than most of his contemporaries, since when
Pontiac's Conspiracy broke out in 1763 the little post he had
grandiloquently named Fort Edward Augustus was the only
one in the West not attacked by hostile tribesmen and its
commandant was called upon to come with his Indian allies
to the aid of the captured officers and garrison at Mackinac.1
So faithful were Wisconsin Indians to British interests that
at the great conference called at Niagara in 1764 by Sir
William Johnson to settle differences the “Old King" of the
Menominee Indians was paid signal honors by Johnson him¬
self.2 Although Fort Edward Augustus was never regarri¬
soned, Wisconsin Indians were for the next decade, on the
whole, obedient to British orders. When Robert Rogers at
Mackinac in 1766-67 sent his subordinates, Captains James
Tute and Jonathan Carver to pacify the Wisconsin Indians
they induced the Sauk and Foxes as well as the Sioux to
abandon the French and Spanish traders, who were coming
up the Mississippi from St. Louis, and go to Mackinac to
renew their allegiance with the English king.3 Rogers gave
them large presents to bind their troth, so large that he was
rebuked by his superiors for his extravagance. Thus at the
'Fis. Hist. Colls. I, 24-48.
a Wis. Hist. Colls. XVIII, 268.
3 Wis. Magazine of History, XII, 139-140.
48 Wisconsin Academy of Sciences , Arts , and Letters.
beginning of the Revolutionary years, the British were in
control of the Indian situation. The trade centered at
Mackinac and passed through the Fox-Wisconsin route to
the Mississippi. Adventurous French Canadians had even
attempted to obtain the Spanish trade on the Missouri, but
were caught and punished by the watchful authorities at St.
Louis. Farther north along the Des Moines and the St. Pet¬
ers the British carried the fur trade into Spanish Louisiana
with impunity.
One reason the Wisconsin Indians were so well controlled
by the British authorities was because the French residents
of Wisconsin had taken an oath to the British sovereign and
their leaders had become officials of the Indian Department.
Charles de Langlade, a captain in the British service, re¬
sided at Green Bay and kept the eastern part of the present
state in order, while his nephew Charles Gautier lived at
Prairie du Chien and was influential with the Mississippi
tribesmen. When the American rebels threatened Canada
Sir Guy Carleton sent word to Langlade to come to his as¬
sistance and during the early years of the Revolution, Lang¬
lade led annually a contingent of Wisconsin Indians to op¬
pose Montgomery, Arnold, and the American officers who
were trying to add Canada as the fourteenth state to the
new American nation.4
Meanwhile, forces were gathering in the West which
were to make the British hold over Wisconsin Indians pre¬
carious. Early in 1778 a young colonel from Virginia
planned to protect the outlying settlements of that state
by carrying the offensive into the enemy territory. The
fourth of July of that year George Rogers Clark captured
the British post of Kaskaskia and within a few days had
taken Cahokia and all the outlying settlements for the Amer¬
icans. Clark also made friends with the Spanish command¬
ant across the river and thus controlled all the middle region
of the Mississippi. We speak of Kaskaskia as a British
post, and so it was technically ; its residents, however, were
all of French descent and when Clark informed them that
the French king had made an alliance with the American
4 Wis. Hist. Colls,. XVIII, 355-358.
Kellogg — Wisconsin Indians During American Devolution. 49
rebels, they all cheerfully took the oath of allegiance to the
new nation.
By this action the Wisconsin Indians were thrown into
a turmoil. Clark's French agents at once found their way
to the villages on Rock River and carried American peace
belts as far as Prairie du Chien and the portage. Overjoyed
at the prospect of the return of their old French “fathers,”
the Wisconsin chiefs hastened to the Illinois to meet the
conqueror. At Cahokia in August and September, 1778 an
immense concourse of northern Indians gathered and held
council with George Rogers Clark. Langlade was in Canada
with a large force of Wisconsin tribesmen supporting Carle-
ton ; no one in Wisconsin was able to restrain the curiosity
of the Indians left at home to see the “Big Knife” American.
This was not an individual name for Clark, but was the
Indian name for first, the governor of Virginia ; second, all
Virginians; third, all Americans.
Clark's account of the series of treaties he negotiated
with the Northwestern Indians at Cahokia on the Missis¬
sippi is interesting in the extreme.5 He states that there
were present Chippewa, Ottawa, Potawatomi, Mississaga,
Winnebago, Sauk, Fox, and Iowa Indians, who conducted
the ceremonies with the dignity and solemnity befitting the
occasion. “I suppose,” he wrote, these negotiations had “as
much Dignity and Importance in their eyes as the treaty
between France and America in ours.” Clark developed a
new technique in dealing with the natives ; he assured them
that he was absolutely indifferent whether they accepted
peace with the Americans or not. “I carry in my right
hand war and in my left peace,” he told them, and explained
that the Great Council fire at Philadelphia had sent him to
clear the roads for those who wished to be friends with the
Americans. They might choose which they pleased, peace
or war, but ifj the latter, then their women and children
would be subjected to grave danger.
The chiefs greatly admired the courage and indifference
of the “Big Knife”; they hastened to assure him of their
sympathy and that they would repudiate the British alliance
for one with him. Clark gravely responded to their wishes,
6 III. Hist Colls. VIII, 125-129, 243-248.
50 Wisconsin Academy of Sciences , Arts , and Letters.
and drew up treaties and certificates, clinching their new
allegiance. Two of these certificates have been preserved;
that with the Winnebago chief of Rock River, and one of
those given to the head Fox chief.6
A separate account of Clark’s dealings with the Milwau¬
kee Indians appears in his writings. The Indian village at
that place was of mixed composition — Ottawa, Chippewa,
and Potowatomi ; their chief Blackbird or Letourneau whose
Indian name was Siggenauk, was an Ottawa who had mar¬
ried a Potawatomi and had identified himself with the
latter tribe.7 Clark, however, called him a Chippewa.
Somewhere Siggenauk had acquired the manners of civiliza¬
tion and when visiting Clark, at the latter’s invitation,
“attempted to speak as much in the European manner as
possible.”8 Clark humored him and made fast friends with
him and so successful was he in attaching this chief and his
band to the American interest, that the British officer at
Mackinac spoke of them as “those runagates at Milwakie”,
and sent Gautier by ship to arrest him. Blackbird, how¬
ever, had flown and although a reward was offered for him
if fetched in either by fair or forced methods, he was not
captured and remained true to the “Big Knife” during all
the years of the Revolution.9
It would be tedious to relate in detail the duel that en¬
sued during the latter years of the Revolution between
Clark and Langlade for the alliance of the Wisconsin In¬
dians. After Clark’s capture of the British Governor Ham¬
ilton at Vincennes, his fame soared aloft among all the west¬
ern tribes, who fell away from the British to such a degree
that Mackinac trembled for its safety. It was at this time
that the garrison was removed from the south shore, and the
fort built on the island so as to be more secure from cap¬
ture by the enemy. The commandant reported that only
the Sioux Indians were faithful to the British.10 The next
year, 1780, the British expedition against St. Louis was
•Wis. Hist. Colls. XI, 113, 177.
7 Description of this chief and his son, the author of the Chicago massacre
of 1812, is in the Miners’ Journal, Galena, Ill., Oct. 30, 1830.
8 III Hist. Colls. VIII, 253-257.
•Wis. Hist. Colls. XI, 201; XVIII, 384.
10 Wis. Hist. Colls. XI, 144.
Kellogg — Wisconsin Indiums During American Revolution. 51
joined by a considerable Indian contingent, which deserted
its leaders on the rumor that Clark was at Cahokia prepar¬
ing to cross the river to aid the Spaniards in their defense.
Indeed, it was reported to British headquarters that the ex¬
pedition failed because of the treachery of the Sauk and
Foxes.11 The Winnebago, on the other hand, had a chief
and three men killed and four wounded, the only Indians
who suffered in the British cause.12
So the alliance wavered, now on one side, now on the
other, with the fortunes of the contestants. The close of
the Revolution, however, found the British largely in con¬
trol, due to their ability to bribe the tribesmen with pres¬
ents. The American Congress was too poor to send a large
amount of goods and ammunition to their Indian allies ; while
the British, with greater means and in control of the Great
Lakes, were able to forward to Detroit and Mackinac large
invoices of Indian presents. The Wisconsin Indians should
not be considered wholly mercenary, selling their friend¬
ship to the highest bidder. White men’s goods were neces¬
sary to their very existence : without powder and lead for
their guns they could not hunt; without blankets and cloth
they would be exposed to the elements, mayhap freeze to
death; without game and furs which they had to trap or
shoot with white men’s implements, they had no means of
livelihood. It should be remembered, however, especially
in this year of the George Rogers Clark Sesquicentennial,
that had not the Wisconsin and other Western Indians made
a temporary alliance with Clark at Cahokia in 1778, he
could not have maintained his position in Illinois, nor con¬
quered Vincennes, nor protected Kentucky. And had it not
been for Clark and his conquests, might not this great
Northwest, now so important a part of the United States,
have remained, as it then was, a part of Canada and become
a portion of the British Empire? These suggestions may
explain the importance of the role played by the Wisconsin
Indians in the American Revolution.
Wis. Hist. Colls. XI, 147-157 ; XVIII, 404-408.
13 Wis. Hist. Colls. XI, 156.
THE SETTLEMENT AND THE DISTRIBUTION OF
THE POPULATION IN WISCONSIN*
Guy-Harold Smith
Historical Introduction
The westward movement of settlers from the Atlantic
seaboard into the virgin lands beyond the Appalachians is
epochal in American history. The pioneers of early Amer¬
ican stock were joined by the foreign immigrants and to¬
gether they pushed westward into the new lands. After
1763 there began a steady stream of frontiersmen who cut
away the forests and established permanent settlements as
soon as, and even before, the land was acquired from the ab¬
origines. Everywhere they were confronted with differing
physical conditions to which they consciously or uncon¬
sciously adapted their activities.
About 1820 the frontier of settlement reached that por¬
tion of the Old Northwest that was later to be circumscribed
by the boundaries of the State of Wisconsin. The new lands
of the future state presented to the settlers a variety of
habitats each differing more or less from the others. These
differences in regional geography have produced a popula¬
tion sectionalism that has persisted for a hundred years.
Preliminary Exploration . The geographical conditions
of the State of Wisconsin, since the visits of the first white
men, have invited and guided exploration, immigration and
settlement. No doubt, the unwritten history of the ab¬
origines is filled with episode and incident reflecting the in¬
fluences of forests, prairies, lakes, and streams upon the
trails followed for game or for enemies, upon the selection
and division of hunting grounds, and the reservation of
ceremonial and burial sites.
Not only did the geographical conditions within the state
* A brief regional summary article on this subject entitled “The Populat¬
ing of Wisconsin” was published in the Geographical Review, Vol. XVIII, No.
3, July, 1928, pp. 402-421.
54 Wisconsin Academy of Sciences, Arts, and Letters.
influence the course of history, but the orientation of lakes
and streams beyond the present boundaries guided and di¬
rected explorers toward Wisconsin. Schafer writes that,
. . . The imposing geographic arch formed by the Mississippi
lands on the one hand and those of the Great Lakes and the St. Law¬
rence basin on the other has for its keystone the territory embraced
within the boundaries of Wisconsin. Resting lightly on Lake Su¬
perior but with a long shore line on both Lake Michigan and the
Father of Waters, that territory also holds the most convenient line
of communication between the two systems, the Fox and Wisconsin
Rivers, separated by a single short portage. This explains why so
much of the early history of the State not only connects but mingles
and blends with the French history of Canada and Louisiana . . /
Miss Kellogg, in one of a series of articles on the early
history of Wisconsin, has called attention to the importance
of geography in the discovery and exploration of the region.
She notes that,
. . . Wisconsin’s position at the headwaters of the two great
valleys of North America — the St. Lawrence and the Mississippi —
has been of supreme importance in the history of the State. To
these advantages of position is due its early discovery, its thorough
exploration, and its value as a link in the penetration of in¬
land America.1 2 * * *
Turner, the foremost student of American frontier his¬
tory, has made the following observation in his studies of
the fur trade.
. . . The importance of physical conditions is nowhere more mani¬
fest than in the exploration of the Northwest, and we cannot properly
appreciate Wisconsin’s relation to the history of the time without
first considering her situation as regards the lake and river systems
of North America.8
The French Regime, 1634-1763. The first white man to
set foot upon the territory, which was later to be included
within the boundaries erected to delimit the Territory, and
still later the State of Wisconsin, was Jean Nicholet. His
1 Joseph Schafer. A History of Agriculture in Wisconsin. (Wisconsin
Domesday Book, General Studies, Vol. 1), Madison, 1922, p. 1.
2 Louise Phelps Kellog. The Story of Wisconsin, 1634-1848. The Wis¬
consin Magazine of History, Vol. II, March 1919, p. 257.
8 Frederick J. Turner. The Character and Influence of the Indian Trade
in Wisconsin. Johns Hopkins University Studies, Vol. IX, 1891, p. 559.
Smith — Distribution of Population in Wisconsin.
55
journey under the direction of the indefatigable Champlain
illustrates not only that the latter . . wished to test
his theory that the route to the East lay through the Great
Lakes,”4 * * but how the lakes and tributaries from the begin¬
ning directed the course of exploration toward Wisconsin.
Nicholet ascended the Ottawa River and crossed by way
of Lake Nipissing into Georgian Bay which gave him easy
access by way of the Straits of Mackinac to Lake Michigan
and its important arm, Green Bay, near the head of which
he landed in 1634. (Fig. 1).
. . . We are now able to see how the river-courses of the North¬
west permitted a complete exploration of the country, and that in
these courses Wisconsin held a commanding situation. But these
rivers not only permitted exploration: they also furnished a motive
to exploration by the fact that their valleys teemed with fur-bearing
animals . . . The hope of a route to China was always influ¬
ential, as was also the search for mines . . .8
There were manifold motives that directed the pioneer
explorers into Wisconsin. To some the impelling motive
was a search for a passage across the North American con¬
tinent to the Pacific Ocean, and the discovery and explora¬
tion of the Great Lakes stimulated this vain hope. To
others exploration was secondary to the diplomatic services
they might render in securing the friendliness of the In¬
dians with whom the fur trade was to become the economic
basis of an important commerce. Still others hoped to find
mines and duplicate the wealth Spain was reputed to have
won from the Indians of Mexico and South America. What¬
ever motive was dominant the orientation of the lines of
communication brought Wisconsin in touch with the settle¬
ments along the lower St. Lawrence River at an early date.
After Nicholet' s visit of 1634 exploration progressed
rather slowly because of the hostility of the Iroquois In¬
dians, but the explorers, missionaries, and traders gradu¬
ally worked westward from lower Canada into the Upper
lakes country. Among the missionaries were the explorer
Louis Joliet and Father Jacques Marquette who in 1672
4 Louise Phelps Kellogg1. The French Regime in Wisconsin and the North¬
west, p. 78.
“Turner. Ibid. p. 562.
56 Wisconsin Academy of Sciences, Arts, and Letters.
ascended the Fox River, portaged to the Wisconsin and
descended it to its junction with the Mississippi.
Up to the end of the French and Indian War in 1763 Wis¬
consin’s bordering lakes and penetrating streams guided
Fig. 1. This identification map gives the names of the modern
counties of Wisconsin and the location of twenty important cities.
the hardy pioneers into the heart of the area. The many
straits, portages, river mouths and important lakes became
the strategic sites of the fur trading posts, and Green Bay
became the commercial capital of Wisconsin.
Smith — Distribution of Population in Wisconsin . 57
The traffic in furs was of such importance that the profit¬
able returns stimulated the traders to penetrate farther
into the interior of the country, and while very few of these
French, and French and Indian half-breeds became per¬
manent settlers there must have been a few hundred of
them who spent a major portion of their lives in Wisconsin.
They came, principally, from Quebec with their many iron
implements and interesting trinkets, guns, and ‘fire water'
with which they bargained for the furs. This systematic
trade with the Indians put into their hands a new weapon
and stimulated the Indian to hunt the fur-bearing animals
beyond their own requirements. “They passed also from
the economic stage in which their hunting was for food
and clothing simply, to that stage in which their hunting
was made systematic and stimulated by the European de¬
mand for furs."6
The traffic in furs led to an exploitation which was in
some ways detrimental to the Indians, but the traders ren¬
dered a valuable diplomatic service in preparing a way for
the people who were to come later. They became the liai¬
son agents between the Indians and the miners and farmers
who were soon to follow. Also they constituted the van¬
guard of the sedentary population.
The British Sovereignty , 1763-1816. The fall of Quebec
in 1760 virtually ended the French sovereignty in Wiscon¬
sin, but the change to the British control altered only
slightly the fur trade so far inland as Wisconsin. The
British began to displace the French in some of the import¬
ant posts and significant changes were made in the organi¬
zation of the industry. With the coming of the British —
particularly the Scotch, “. . . there began that long and
brilliant dynasty of Scotch fur barons which has endured
down to our own day.”7 The French-Canadians were not
displaced but merely utilized to carry out the details of the
traffic directed and financed by the British who established
posts and invited the Indians to bring in their furs. Under
this plan another step toward the permanent occupation of
the land had been taken.
6 Ibid., pp. 31-32.
7 Clarence W. Alvord. The Mississippi Valley in British Politics, I. p. 296.
58 Wisconsin Academy of Sciences , Arts, and Letters.
American Control after 1816. The Treaty of Paris in
1783, which marked the close of the Revolutionary War and
transferred Wisconsin from the British to the Americans,
was only a 'scrap of paper ’ so far as any change in the fur
trade was concerned. The traffic was maintained after the
war as before with little or no opposition except from a few
American traders. And concomitant with the commercial
activities there was maintained a virtual political sover¬
eignty of territory no longer British. John Jay was sent
to London to make a commercial treaty with the British in¬
volving among other things a settlement of the northwest
boundary troubles. He received a promise to withdraw in
1796, at which time Wayne’s army took over the post at
Detroit. From that date to the War of 1812 America ex¬
tended her sovereignty over Wisconsin, only to relinquish
it at the outbreak of hostilities when the British “. . .
with Indien assistance, seized the American posts up to a
line indicated by Toledo Bay, Fort Wayne, Peoria and St.
Louis.”8 After the war American sovereignty was again
established and Federal troops were sent to Green Bay and
Prairie du omen in 1816, when control by the United States
became effective for the first time.
The geographical remoteness of Wisconsin is an import¬
ant circumstance explaining this vacillation in political sov¬
ereignty and the delay in settlement of an area so long known
to white man. But after American dominance was estab¬
lished this section of the Old Northwest began to attract
settlers.
The First Settlements. By 1820 the white population had
begun to localize in one or two places, making possible an
estimate of the actual number of permanent settlers. Green
Bay and Prairie du Chien were the only important settle¬
ments in Wisconsin at this time. "The former, chiefly
limited to the stretch of five miles up the Fox River from
its mouth, consisted of about sixty houses, with a population
of perhaps 500 souls, besides which was a garrison.”9
Many of these people were settlers who utilized the land and
8 Frederic L. Paxson. History of the American Frontier. 1763-1893, p. 179.
9 Frederick J. Turner. The Character and Influence of the Fur Trade in
Wisconsin, Wisconsin Historical Proceedings, 1889, p. 92.
POPULATION MAP
WISCONSIN
19 0 0
Smith — Distribution of Population in Wisconsin.
59
the soil for agricultural purposes. They were tillers of the
soil for they raised . . potatoes, maize, oats, peas,
spring wheat, pumpkins, melons, cabbages, onions, and other
common vegetables/110 The ownership of the land was of
certain importance for they had divided the area, according
to the Canadian custom into narrow strips abutting upon
the river, the pioneers’ principal highway.
The farms were ribbon-like cotes common to Canadian settlements,
from one and one-half to eight arpents* wide and running back
eighty arpents from the river. As a rule, only about two or three
acres of this were cultivated.* 11
Prairie du Chien was the only other settlement of any
significance as measured by the number of people. In 1820
there were in the settlement approximately 600 people in¬
cluding the 100 soldiers stationed at Fort Crawford. Like
the Green Bay settlement, “The farms were the narrow
fields running back from the river and there had been a
common field where the inhabitants cut hay.”12 This com¬
mon meadow probably was one of the open prairie areas
found along the flood plain and on the terraces of the Wis¬
consin and Mississippi Rivers. The farm pattern shows
a French Canadian influence that has persisted down to the
present.
The Settlement of the Lead Region of Wisconsin,
1820-1850
The active settlement of southern Wisconsin was accom¬
plished in the three decades following 1820, This west¬
ward migration into Wisconsin may be considered as a part
of the great population readjustment that was taking place
in eastern United States. The sale of public lands — all of
which were west of the Appalachians — reflects the move¬
ment of people from the east to the west. Sales increased
rapidly after 1813, when only 140,000 acres were sold, to
1819, when 5,110,000 acres were purchased from the gov-
10 General Ellis’s Recollections, Wisconsin Historical Collections, Vol. VII,
1876, p. 216.
*'Arpent (linear measure) 192 feet and 6 inches.
11 Turner. Ibid., p. 92.
13 Ibid., p. 92.
60 Wisconsin Academy of Sciences , Arts , and Letters.
ernment, and then came a decrease to only 780,000 acres
in 1821.13 The westward expansion from the American
seaboard into the Mississippi Valley reached its most spec¬
tacular development in the 'great migration' which brought
into the Union Indiana in 1816, Mississippi in 1817, Illinois
in 1818, Alabama in 1819, Maine in 1820, and Missouri
in 1821. "With the admission of Missouri the great migra¬
tion came to an end so far as new states were concerned,
and the heavy shift of population subsided for another ten
years.14 Just as this historic movement began to abate
and readjust itself to the very large area contained within
the borders of the new states, Wisconsin was opened to
settlement. The first movement into the territory was
slow, echoing the decreasing vigor of the declining years
of the great migration.
Green Bay and Prairie du Chien were the only settle¬
ments of any importance before 1820. These were occu¬
pied because of their stragetic location for commerce in
furs and for the military advantages that their sites of¬
fered. These functions were not materially altered until
after 1835 when the agricultural frontier approached more
closely and the Indian menace had been removed.
The Lead Region. The lead deposits of the Upper Mis¬
sissippi Valley had been discovered more than a century
before the beginning of active settlement about 1820.
Even before white man came the Indians had their 'lead
diggings.' Later under the French influence, mining was
stimulated, particularly by Nicholas Perrot, who purchased
lead from the Indians in 1695. From that time until the
end of the French regime about 1760 " . . . the lead
mines were worked more or less constantly both by the In¬
dians and by whites, who used the product to supplement
the fur trade."15 A few traders found it profitable to bar¬
ter for lead, and some of the Indians willingly consented to
mine rather than hunt the fur-bearing animals. This new
adaptation of industry culminated in the settlement of this
section after 1820.
iaPaxson. Ibid., p. 221.
14 Ibid., p. 219.
15 Louise Phelps Kellogg-. The French R£geme in Wisconsin and the North¬
west, p. 361.
Smith — - Distribution of Population in Wisconsin. 61
This lead region lies . . . chiefly in Wisconsin, including,
however, a strip of about eight townships of land in Iowa, along the
western bank of the Mississippi, the greatest width of which strip
is on the Little Mequoketa, about twelve miles from east to west, and
including about ten townships in the northwestern corner of Illinois.
The portion of this lead region in Wisconsin includes about sixty-
two townships, or two thousand eight hundred and eighty square
miles; being about one-third larger than the state of Delaware. The
extreme length of this region, from west to east, is eighty-seven
miles; and its greatest width, from north to south, is fifty -four
miles.16
For more than 150 years, from the discovery of Wisconsin
in 1634 until after 1800, the traffic in furs was the domin¬
ant industry. Lead had made an ineffectual appeal to the
natives except when a few traders were able to stimulate
the mining industry. The continual exploration revealed
more and more the mineral wealth of the lead region, but
the distance from the markets and the state of scientific
knowledge precluded any intensive utilization of the re¬
source. Under such conditions, the early mining, viewed
from the perspective of over a hundred years, appears more
as ‘a promise than a performance/
In 1804 the Sauk and Fox Indians ceded to the United
States the larger portion of the lead bearing region of the
Upper Mississippi Valley. The treaty among other things
provided that “As long as the lands which are now ceded
remain their property, the Indians belonging to said tribes
shall enjoy the privilege of living and hunting upon them.”17
This stipulation was one of the indirect causes of the Black
Hawk War. The pioneers who passed beyond the agricul¬
tural frontier encroached upon land used and jealously
guarded by the Indians and any unlicensed intrusion was
regarded as trespass.
The active mining of lead began in Illinois about 1822
near the present city of Galena, and shortly thereafter pio¬
neer miners penetrated into Wisconsin. “In one sense the
settlements were an expansion of those of northern Illinois,
a widening circle of adventurers, who, not finding the
‘hoped for* wealth in the mines about Galena, pushed on
19 David D. Owen. Report of a Geological Exploration of Part of Iowa,
Wisconsin, and Illinois, in 1839. Revised Edition, 1844, p. 35.
17 Laws of the United States, Vol. 1, 1789-1815, p. 427.
5
62 Wisconsin Academy of Sciences, Arts, and Letters.
into Southwestern Wisconsin to seek for richer ‘leads’.”18
It was not until after 1827 when the Winnebago disturb¬
ances had quieted down that the miners became more dar¬
ing. “ . . . they ventured far beyond that protection
which numerical strength and the defensive organizations
near Galena secured.”19
The number of people in the lead region was very dif¬
ficult to determine but the report of the Superintendent of
the Mineral Lands stated that,
I am enabled to give with accuracy the number of persons at the
Public Mines of Illinois only, or rather at the mines near the north
boundary of that State, which are supposed to be within its limits,
commonly known as the Fever River Mines, viz. On the first day
of July, 1825, there were at those mines about one hundred men; on
the thirty-first of December, 1825, one hundred and fifty-one men;
on the thirty-first of March, 1826, one hundred and ninety-four; on
the thirtieth of June, 1826, four hundred and six; and on the thirty-
first of August 1826, (the date of last report,) there were four hundred
and fifty-three men. You will observe the increase has been grad¬
ual, and the numbers are still augmenting.20
From this Illinois nucleus the settlement spread northward
into southwestern Wisconsin.
As the frontier came nearer, the lead region tapped the
stream of migration and deflected a part of it northward
into the lands along the Upper Mississippi River.
The time had now come when this beautiful country was to be occu¬
pied by a hardy, resolute, adventurous and persevering population.
The laws which, as a rule, generally confine the migration of the
human race to isothermal zones and similarity of climate, were to be
set at defiance, and the emigrant from the mild climate of Tennessee,
Kentucky, Missouri and southern Illinois was to exchange, the balmy
and genial atmosphere to which he had been accustomed, for one in
which during nearly half the year all nature is bound with icy chains
and covered with its robe of snow.21
This explains how so many people of southern origin found
their way into southwestern Wisconsin.
18 W. O. Blanchard. The Geography of Southwestern Wisconsin. Wis.
Geol. and Nat. Hist. Survey. Bull. No. 65, p. 30.
19 Stephen Taylor Wisconsin — Its Rise and Progress, with Notices of
Mineral Point and Richland County. Wisconsin Historical Collections, Vol.
II, 1856, p. 485.
20 House Executive Documents, 19th Congress, 2nd Session, II No. 7, p. 8.
21 Moses M. Strong. History of the Territory of Wisconsin from 1836 to
1848, p. 118.
Smith — Distribution of Population in Wisconsin.
63
Influence of Geological Conditions . The mere presence
of lead in Wisconsin was not the only factor which induced
the migration of pioneer miners, for the mode of occurrence
of the lead-bearing minerals determined to a notable de¬
gree the distribution of the inhabitants. Within the min¬
eralized areas there is a zonal arrangement of the principal
minerals.22 Near the surface and some places at the sur¬
face, the lead ore, occupies the upper zone, occurring as
large cubical crystals of galena. The fortunate circum¬
stance that abundant ore occurred above the water table
made it possible for individuals to engage in the mining in¬
dustry. This created a population dispersion from the out¬
set. Some of the miners, who later were able to purchase
land for agricultural purposes were engaged in both indus¬
tries, depending upon season or price of lead to determine
to which they would give their services.
From Mining to Agriculture. Mining required less cap¬
ital than farming. “A miner could carry practically all of
the tools necessary for his work. Mining leases cost only
a share of the profits.”23 Gradually there came a shift
from mining to agriculture. “As agriculture developed
and mining became more difficult because the surface ores
had been removed, farmers turned their attention more
and more to the profits of agriculture to the neglect of min¬
ing.”24 The peak of lead production was reached about
1845 after which date the amount decreased to less than
half of the production of the early forties.
The Black Hawk War of 1832 removed the last menace to
white man’s occupancy of the lead region. At first the semi-
nomadic trader stimulated the Indians to search for furs.
Over this primitive occupation the miner overlaid a new in¬
dustry characterized by the intensive search for lead, and
localized effort replaced the extended activities of the
hunter and trader.
The coming of the farming element after 1832 added
32 U. S. Grant and E. F. Burchard. Description of the Lancaster and Min¬
eral Point Quadrangles (Wisconsin-Iowa-lllinois). U. S. Geol. Survey, Folio
No. 145, p. 12.
“Blanchard. Ibid., p. 66.
24 Selma L. Schubring. A Statistical Study of Lead and Zinc Mining in
Wisconsin. Trans, of the Wis. Acad. Sci., Arts and Let. Vol. 22, 1926.
p. 15.
64 Wisconsin Academy of Sciences , Arts, and Letters.
hundreds to the population, and the concentrated occupancy
of the land completely displaced the aborigines and erased
their culture from the land that had maintained them for
centuries. Cultural conditions of another order displaced
the old when white man replaced the red man.
The land with its fertile soil and the extensive prairie
areas was capable of yielding crops — particularly wheat —
that would equal those of other states. According to
Owen's report the land of this section was originally about
37 per cent prairie and 63 per cent woodland.25 This com¬
bination of woodland interspersed with prairie was a for¬
tunate circumstance for the early farmers, but many did
not realize it.
The Land Survey . It is significant that southern Wis¬
consin was surveyed from the west toward the east. The
land survey was begun in 1833 by the erection of the
Fourth Principal Meridian through the heart of the lead
region where a nucleus of permanent settlers required a
title to their land held previously by leases or by 'squatters'
rights.
The survey initiated a speculation in land, particularly
in sections adjacent to the mineral lands reserved from sale.
Probably two thirds of the land sales prior to 1837 were to
speculators.26 The 'specie circular' issued at the stipulation
of President Jackson ended speculation. The President de¬
sired to prevent public lands from falling into the hands of
capitalists who might do injury to settlers in search of new
homes.27 Speculation was curtailed, and the settlers con¬
tinued to press against the cutting edge of the frontier.
Statistical Measure of Settlement . The first reliable
quantitative data on population, other than estimates and
partial enumerations in Green Bay and Prairie du Chien, are
contained in the United States Census of 1830 which gives
a population of 3,245 for the three Wisconsin Counties, viz,
Brown County 964, Crawford County 692, and Iowa County
1,589. Almost half of the people were in Iowa County,
20 Owen. Ibid, p. 145.
28 Strong. Ibid., p. 217.
27 Senate Document No. 15 of December 14, 1836. 24th Congress Second
Session.
Smith — Distribution of Population in Wisconsin.
65
which included the lead region, and the remainder were in
the two long established posts of Green Bay and Prairie du
Chien.
Fig. 2. The population of Wisconsin in 1836. (Data from the
First Census of Wisconsin Territory, taken in July 1836.)
Wisconsin was organized as a Territory in 1836 and the
next census28 taken in July of that year gives the population
as 11,683. Brown county had 2,706 inhabitants, Crawford
38 The Territorial Census for 1836. Wis. Hist. Coll. Vol. XIII, 1895, pp.
247-270.
66 Wisconsin Academy of Sciences , Arts, and Letters.
850, Iowa 5,234, and Milwaukee 2,893. The lead region
continued to be the most densely settled section.
The two counties of Grant and Iowa included one fourth
of the population of the Territory. But with the rapid
Fig. 3. The population of Wisconsin in 1838, two years after the
formation of the Territory. (Data from Clark S. Matteson, An Illus¬
trated History of Wisconsin, pp. 259-260.)
growth of population in eastern and south central part of
the Territory the lead-producing counties had only 17 per
cent of the total in 1846, and by 1847 this was decreased to
Smith — Distribution of Population in Wisconsin.
67
13 per cent. Lead mining was declining but settlers con¬
tinued to come into the area. The gold rush to California
attracted many miners but the influx of agriculturalists pre¬
vented a decrease in population.
Fig. 4. By 1840 the frontier had crossed the southeastern quarter
of Wisconsin. (Data from the Journal of the Convention, January
3, 1848, pp. 156-163.)
With the exhaustion of the easily mined lead deposits,
the lands previously reserved from sale were placed upon
the market. Congress provided that the President be
68 Wisconsin Academy of Sciences , Arts , and Letters .
“ . . . authorized, as soon as practicable to cause the
reserved lead mines and contiguous lands in . . . Wis¬
consin . . . belonging to the United States, to be ex¬
posed to sale, in the same manner that other public lands
are authorized by law to be sold . . ,”29 By the time
Wisconsin was admitted to the Union in 1848 the popula¬
tion of the lead region was approximately 30,000. In 1825
there were only a few score in the region, and in 1836 when
Wisconsin became a Territory the number was only 5,234,
but rapid settlement in the middle forties changed the
lead region into an agricultural section.
The Settlement of Eastern Wisconsin
1832 to 1850
After 1832 the southeastern part of Wisconsin began to
tap the stream of western migration and by 1850 most of
the area from Lake Michigan westward to the lead region,
and northwestward toward the Fox River was completely
settled, and the frontier moved on. The United States Cen¬
sus for 1850 gives in detail the population of the state, and
the map prepared from these data pictures graphically the
distribution of the inhabitants. The map shows the rather
even film of rural population thinning out along the fron¬
tier, and the concentration of the people in urban commu¬
nities where the advantages of site or situation provided
the bases for the agglomerations.
The frontier of settlement, by 1850, was approaching,
and in a few places had crossed the Fox and Lower Wis¬
consin Rivers. That indefinite boundary between the set¬
tled and the unoccupied land was a progressively shifting
agricultural frontier. The westward and northward move¬
ment of this frontier up to 1850 had left behind it over 90
per cent of the total population, or 287,730 settlers, includ¬
ing 37,225 in the three lead mining counties of the south¬
west. The remaining 17,660 people were in the ten or
more northern wilderness counties.
The Beginning of Settlement . The settlement of south¬
eastern Wisconsin after 1832 is epochal. The lead region
29 TJ. S. Statues at Large. Vol. IX July 11, 1846, p. 37.
Smith — Distribution of Population in Wisconsin.
69
was first settled by a final wave of the great migration,
but southeastern Wisconsin was peopled by a current from
the Jacksonian migration which started about 1882, so far
as Wisconsin is concerned, and reached its crest about
Fig. 5. Between 1840 and 1842 the population of Wisconsin in¬
creased from, 30,945 to 44,478. (Data from the Journal of the
Convention, January 3, 1848, pp. 156-163.)
1837 ; but for many years thereafter the irresistable human
horde sought the newly opened lands. Most of the land
was entered either by speculators or by the actual settlers
in the years of 1836, 1837 and 1838. Even after 1840
70 Wisconsin Academy of Sciences , Arts , and Letters.
there was considerable land to be settled, but the major por¬
tion of the government lands had been sold. Resale by-
speculators permitted an intermingling of new settlers with
those who had preceded them by only a few years.
Fig. 6. By 1846 southern Wisconsin had a population of approxi¬
mately 150,000. (Data from the Journal of the Convention, January
3, 1848, pp. 156-163.)
Just as the explorers and traders sought the waterways
in their penetration of Wisconsin, the settlers were guided
in their great ‘trek’ to the new lands. They were ever
Smith — Distribution of Population in Wisconsin. 71
mindful that to depart from the waterways was to find as
obstacles, swamps and forests that were traversed only
with great effort. The first settlers into the lead region
came from the south and created a nuclear settlement that
expanded eastward. There were more people from Ken¬
tucky, Tennessee, and Missouri in the lead region in 1850
than in all the remainder of the state. Southeastern Wis¬
consin drew homeseekers from New York, New England
and other eastern states, thus the movement into Wisconsin
after 1832 was an along-the-parallels migration.
The new settlers came by way of the recently completed
Erie Canal and the Great Lakes to the western shore of
Lake Michigan. From the Ohio Valley many descended
the Ohio River to Cincinnati where they deserted nature's
highway and come across Indiana and Illinois to Wisconsin.
Enroute tLey were joined by natives of the states they were
traversing until the newcomers represented a homogeneous
mixture of native Americans and their immediate foreign
born ancestors. Among the immigrants from Indiana and
Illinois, came many people, who had participated in the
great migration.
The manuscript census30 for 1850 records many families
in which the father and mother were born in New York,
and two or three children in Indiana, and two or three more
in Wisconsin. Similarly many of the foreign born came
westward as far as Ohio and then moved on to Wisconsin
after a few years. The peopling of Wisconsin had become
a psychological movement, and the news accounts in the
eastern papers accelerated the migration.
Schafer, writing about the southeastern section of the
state, notes that,
The movement of settlers into the region was well started by the
spring of 1836. Most of the emigrants came by sloop or steamer
on the lakes, debarking at Milwaukee, Racine, or Kenosha, or else at
Chicago whence they made their way up the coast.31 (See figures
2, 3, 4, 5, 6, and 7)
The Land and Its Utilization. Southeastern Wisconsin
is almost entirely within the physiographic province called
80 The manuscript census reports for 1850, 1860, and 1870 have been de¬
posited in the Library of the State Historical Society of Wisconsin.
31 Joseph Schafer. Four Wisconsin Counties, Prairie and Forest, p. 64.
72 Wiscoytsin Academy of Sciences , Arts , and Letters .
by Martin ‘The Eastern Ridges and Lowlands.”32 (See
fig. 8). This region is underlain by eastward dipping sedi¬
mentary formations of the Paleozoic system. In pre-gla-
r
Fig. 7. In 1847, on the eve of the admission of Wisconsin into the
Union, the population was 210,546. (Data from the Journal of the
Convention, January 3, 1848, pp. 156-163.)
cial time the area had been maturely dissected into a relief
of approximately 300-500 feet. The major topographic
features are the cuestas developed upon these slightly dip-
33 Lawrence Martin. The Physical Geography of Wisconsin, Wis. GeoL
and Nat. Hist. Survey, Bull. 36, 1916, pp. 197-298.
Smith — Distribution of Population in Wisconsin .
73
ping sedimentary rocks. Superposed upon the maturely
dissected cuesta landscape is a veneer of glacial drift which
has reduced the relief and replaced the erosional topog¬
raphy with a depositional topography. However, along the
escarpment margin of the cuestas the drift is thin and the
bed-rock makes its influence felt through the mantle of
drift.
Fig. 8. The physiographic provinces of Wisconsin. (After Mar¬
tin.) (Courtesy of the Geographical Review published by the
American Geographical Society of New York.)
In the southern tier of counties from Lake Michigan to
the Mississippi there were originally extensive prairie
areas. Northward the prairies were smaller and not so
closely spaced. Around the prairies were the ‘oak open¬
ings’ and the more densely forested areas which took on a
distinctly primeval aspect farther north.
The distribution of improved land in 1850, and particu¬
larly the number of acres per capita indicate with fair ac¬
curacy the progress of settlement in Wisconsin. Rock
County ranked first with 143,235 acres; Walworth was sec¬
ond with 116, 750 acres; and Waukesha third with 105,269.
The counties having over 5 acres of improved land per cap¬
ita were Rock, Walworth, Dodge, and Green. In general
74 Wisconsin Academy of Sciences, Arts, and Letters.
the amount of improved land decreased rapidly toward the
frontier and particularly northward along the lake shore
where the heavy forests effectively retarded the improving
of the land.
Washington County had 2.2 acres of improved land per
person, Sheboygan 1.6 acres, and Manitowoc only 0.3 acres.
Similarly there was at that time a concomitant relation¬
ship in the number of people in these three counties.
Washington had a population of 19,485, Sheboygan 8,379,
and Manitowoc only 3,702.
Fig. 9. The prairie areas — in black — in southeastern Wisconsin.
(After Martin.) (Courtesy of the Geographical Review published by
the American Geographical Society of New York.)
This region to the east was heavily forested and the con¬
version of the unimproved to improved land required
years of strenuous labor; and then the stumps remained
for a generation, hindering the cultivation of the land.
The People. The value of any area for agricultural pur¬
poses is largely determined by the conditions of the natural
landscape, soil, climate, and proximity to other areas which
offer opportunities for exchange of products and ideas.
But under almost ideal environmental conditions a region
might remain undeveloped. “The lands of any country are
important for the human opportunity they represent. The
use which is made of them depends upon the people who
come into their possession.”83
33 Joseph Schafer. A History of Agriculture in Wisconsin, p. 23.
Smith — Distribution of Population in Wisconsin. 75
In southeastern Wisconsin there has developed a section¬
alism where the conditions of the physical landscape were
more or less uniform originally, except for a difference in
the natural vegetation. The southern part of this area was
characterized by the open oak forest with the extensive
prairies. (Fig. 9). Just to the north of the southern tier
of counties was a hardwood forest of maple, and some oak
and beech. Along the eastern coast a strip of conifers be¬
gan just north of Milwaukee and extended northward, be¬
coming wider until it embraced all of Door Peninsula.
In the settlement of southeastern Wisconsin, there was
enacted what may be called an experiment in historical
geography. The southern division was settled by the
Yankees and the British and the forested section by the
Teutons. When the Yankees settled the southern row of
counties they did not, for long at least, shun the prairie
lands as has often been stated. However, it is true that
certain prairie sections were not entered until a few years
after the wooded areas had been sold. Schafer has shown
. . . that the Yankee settlers in a prevailingly prairie township
of Racine County took up first every acre of forested land, together
with the prairie lands and marsh lands adjoining the woods, while
they shunned for some years the big, open, unsheltered prairie where
farms would be out of immediate touch with the woods.34 (Fig. 10).
These homeseekers were not destitute, and with the credit
extended to them by their friends and relatives, and because
of a tradition distinctly British in origin, the purchase of a
farm was looked upon as a method of creating an estate.
Large farms were the rule.
The number of Germans in Wisconsin before 1850
formed an important nucleus which determined the popula¬
tion conditions in the forest section of eastern Wisconsin
(Fig. 11) . The 38,064 Germans made a noteworthy contri¬
bution to the vital statistics.
About 30,000 old Lutheran subjects of Prussia . . . are shortly
to come over and settle in these United States. It is a religious
movement, these people preferring the good old orthodox doctrines to
the modern philosophy of Berlin. There are men of very large for-
84 Joseph Schafer. The Yankee and the Teuton in Wisconsin. Wisconsin
Magazine of History, Vol. VI, Dec. 1922, p. 9.
76 Wisconsin Academy of Sciences, Arts, and Letters.
Fig. 10. The number of German inhabitants in Racine, Kenosha,
and Walworth Counties in 1850 was small. The British and the
Yankees were the predominating population groups. Each symbol
represents 25 inhabitants; the dots, German and the circles, British.
(Data from a hand count made from the 1850 manuscript census
for the State Historical Society of Wisconsin by M. M. Quaife and
an assistant.)
Fig. 11. The distribution of German and British born settlers in
Dodge and Washington Counties in 1850. Each symbol represents 25
inhabitants ; the dots, German, and the circles British. Note the gen¬
eral decrease in the Germans westward from Lake Michigan. In
1850 Washington County included the present Washington and Okau-
kee Counties. (Data from a hand count made from the 1850 manu¬
script census for the State Historical Society of Wisconsin by M. M.
Quaife and an assistant) .
Smith — Distribution of Population in Wisconsin. 77
tunes among them; old German noblemen whose pedigrees date back
to the thirteenth century. They will make excellent western farm¬
ers, and are about to settle in Wisconsin — the coolest spot they can
select.85
These German immigrants coming to America took the
densely wooded areas. Why this selection? Schafer gives
an answer in the following statement.
It is at bottom a question of economic ability, not of personal or
racial tastes. The poor immigrants and the poorer natives also, with,
of course, many exceptions, settled in the woods because they could
not afford to encounter the risk of taking an ideal farm in the
‘Congress Land’ districts, nor could they afford to buy such land
from speculators or from farmers. They took what was at hand,
the heavily wooded lands avoided by persons who were in position to
pick and choose. In many cases they might have found lands on the
open prairies, which . . . were taken later than the other lands
even by Americans who had some means. But the person without
means would have been helpless in such a situation. He would need
money to buy lumber both for building and for fencing, while in the
timber his personal labor supplied these essentials, without cost, in
the process which at the same time cleared his land.36
The Germans who settled in the rural communities gener¬
ally purchased small farms, from 40 to 80 acres, and this
had the effect of creating a dense rural population in this
area which was originally a maple forest. And after three-
quarters of a century the small farms are common, in many
of the counties averaging less than a hundred acres. (Fig.
20). No other part of Wisconsin has so dense a rural pop¬
ulation as this German settled section.
In a generation or two the forest had been removed and
the two sections were essentially the same. Both the Ger¬
man settled section northwest of Milwaukee and the Yankee
settled strip along the southern tier of counties present a
cultural landscape of beauty and prosperity. This type of
controlled geographical experiment illustrates the import¬
ance of nationality, a social element, in influencing the rela¬
tion of man to his habitat.
88 Niles’ National Register. June 17, 1843, chronicle from the St. Louis
Republican June 7.
*8 Joseph Schafer. A History of Agriculture in Wisconsin, p. S9.
6
78 Wisconsin Academy of Sciences, Arts, and Letters.
The Northward Movement of the Frontier in Wiscon¬
sin 1850-1880
In the period from 1850 to 1880 the movement of popu¬
lation into Wisconsin completed the agricultural settlement
of that part of the state underlain by the Paleozoic sedi¬
ments. In central Wisconsin the earlier glaciated and
driftless portion of the older crystalline area also received
a large number of people, and across the heavily forested
Northern Highland thin threads of settlements followed the
Fig. 12. Land areas in farms, 1920. (Courtesy of the Geographi¬
cal Review published by the American Geographical Society of New
York).
rivers, roads and railways leaving isolated blocks of un¬
touched wilderness. By 1880 the frontier of settlement
had crossed Wisconsin, but within the state there remained
wilderness sections, around the periphery of which, the
frontier slowly encroached and penetrated.
The frontier seemed to be pivoted at Green Bay and
swept northward more rapidly across the Western Upland.
As it approached the densely forested upland to the north
the settlers departed from the deployed formation to one of
concentrated invasion along important salients. This sin¬
gle-file invasion has had the effect of leaving sections of vir-
Smith — Distribution of Population in Wisconsin.
79
gin forest and cut-over land unoccupied down to the pres¬
ent time.
With the extinction of the Indian titles, the lumbermen
with axe and saw overlaid a new culture on the site of the
fur-trading industry, which had declined after 1834 until it
was of little importance, and the roving hunters and trad¬
ers were replaced by more sedentary, yet transient,
pioneers. Lumbering became the dominant industry and
the removal of the forest prepared the way for the agricul¬
tural settlers who were soon to follow.
Wisconsin has been crossed by a succession of frontiers,
but due to the time of settlement the sequence has not been
the same in all parts of the state. Generally speaking the
earlier settled sedimentary Wisconsin was crossed by the
frontier of the hunter and trader in the period from 1634
down to 1820. This frontier prepared the way for the
miner in the southwestern part of the state. In less than
twenty years the pioneer agriculturists occupied the south¬
ern part of Wisconsin, and along the frontier the farmers
exploited the soil by raising wheat to the exclusion of soil
conserving crops. And finally a diversification of agricul¬
ture, — supplemented at first by dairying, and later made
subservient thereto-— slowly advanced a new kind of fron¬
tier across the land.
The westward movement involved the wasteful removal
of the forest to prepare the land for agriculture. This was
the guiding principle as the frontier marched westward
across the Old Northwest. But as the frontier advanced
northward into the highlands of the northern peninsula of
Michigan, Wisconsin, and Minnesota, the destruction of the
forest did not leave in its wake farm land of such high po¬
tential value as in the hard-wood forests to the south.
The settlement of the prairie areas of the Mississippi and
Missouri Valleys created a demand for lumber which the
Upper Lakes country was made to supply. Northern Wis¬
consin with Michigan and Minnesota became commercial
lumber areas where the removal of the forest for its lumber
was the principal motive in clearing the land and not a
preparation for agriculture. In fact the forest was gone
before much of the land was wanted for agricultural pur¬
poses.
80 Wisconsin Academy of Sciences , Arts, and Letters.
In spite of more invulnerable obstacles in the creation of
a livable habitat out of the new lands the settlements in
Wisconsin “ . . . moved at least one degree farther
north . . .”37 in the decade between 1850 and 1860.
The Settlement of Western Wisconsin . In 1850 there
were more than 9,000 people in western Wisconsin beyond
the lower Wisconsin River. Most of these were in Sauk,
Richland, and Crawford Counties and constituted an ad¬
vance guard of homeseekers who were pushing northwest¬
ward to settle the available lands of the Western Upland
and the adjacent Central Lowland. Settlement progressed
in a wave-like formation, the crest of which swept north¬
ward in the three decades from 1850 to 1880.
The 9,000 of 1850 were increased to over 100,000 in 1860.
The decade of the sixties added 98,645, bringing the total
to over 200,000. By 1880 the population of this part of
Wisconsin exceeded 300,000. Each of the three decades
added 100,000, and, in effect, accomplished the settlement of
the area, for in the decade of the eighties the increase had
dropped to about 75,000. The frontier of the farmer had
passed beyond the Western Upland into the crystalline
Northern Highland.
As previously stated western Wisconsin was settled by a
wave of homeseekers who pushed northward after 1850.
Sauk, Richland, and Crawford Counties were peopled in the
decade of the fifties; Vernon, Monroe, and La Crosse Coun¬
ties constitute the section that was settled in the sixties;
and in the decade of the seventies the wave of settlement
had advanced to the latitude of Pierce and St. Croix Coun¬
ties. (See population maps for 1850, 1860, 1870 and
1880).
The Settlement of Central Wisconsin. The frontier
passed very lightly across the sandy plain of central Wis¬
consin. By 1880 the stream of settlers had pushed beyond
the plain into the southern segment of the Northern High¬
land. The several counties that lie wholly or partially in
the sand country present an aspect of a flat plain with poor
sandy soil unsuited to intensive agricultural development.
87 u. S. Census, Population, 1880, p. XVIII.
Smith — Distribution of Population in Wisconsin.
81
In 1850 the population of this section was about 10,000.
In 1860 Adams County which lies entirely in the plain had
6,492 inhabitants; in 1870 the number had increased to
6,601, and by 1880 the number was 6,741. Settlement was
practically complete by 1870, and later enumerations
showed only slight increases, except in the strategically sit¬
uated cities and villages, and an increment in numbers was
due essentially to the excess of births over deaths. Emi¬
gration from the area had not yet become a population re¬
adjustment.
The Settlement of Northeastern Wisconsin . The north¬
eastern part of the state that was settled in the three de¬
cades from 1850 to 1880 included Brown, Door, Kewaunee,
and Outagamie Counties, and approximately half of Mari¬
nette, Oconto, and Shawano Counties. Much of this area
was either pine covered or there was an important mixture
of the conifers with the hardwoods. Door Peninsula was
more primeval in aspect than the section farther south, and
consequently, offered a more hostile environment to the in¬
coming settlers. Furthermore, the retarded settlement in
the peninsula was due to its location as a projection out into
the lake, and in effect it was passed by.
This section bordering Green Bay had about 35,000 in¬
habitants in 1860. By 1870 the number had doubled, and
the enumeration of 1880 gave these seven counties a total
of nearly 120,000. The following table shows the growth
of population in two lake shore counties.
I860 1865 1870 1875 1880
Sheboygan _ _ __26,875 27,671 31,749 34,021 34,206
Manitowoc „ _ — _ 22,416 26,762 33,369 38,456 37,505
These counties did not fill up so completely as some of the
southern sections largely because of the retarding influence
of the forests.
Brown and Outagamie Counties with the more valuable
agricultural lowland and the advantages of urban sites
along the Lower Fox River received the major portion of
the settlers in the northeastern part of Wisconsin. Brown
County increased from 11,795 in 1860 to 34,078 in 1880, and
in the same period the number in Outagamie County in-
82 Wisconsin Academy of Sciences , Arts, and Letters.
creased from 9,587 to 28,716, and because of the advantages
in this lowland, continued their growth after 1880.
Marinette and Oconto Counties were still in the pioneer
stage of their settlement in 1880. This is the only section
of sedimentary Wisconsin that was not completely settled
by agriculturalists by 1870. There is no doubt that the
pine forest operated as a deterrent to agricultural settle¬
ment. One writer describes the pine lands as follows :
There is comparatively very little farming done in the lumber re¬
gion. The country is rugged; there are few roads; and to man in
search of a home it looks like the work of a lifetime to acquire a foot¬
hold there. There are fairer fields; there are softer climates; there
are roads and school-houses, and more social advancement, farther
southward. So he turns away; and so we go on, year after year,
hewing down our wealth of timber and carrying it off to keep up the
prosperity of other states.38
The Settlement of Northern Wisconsin. That part of
Wisconsin, which lies north of the northern boundary of
Clark, Marathon, and Shawano Counties, was only sparsely
settled prior to 1880. In 1850 approximately 500 pioneers
had ventured into this region; in 1870 there were still less
than 2,000; but by 1880 the number was approaching the
10,000 mark. Before 1880, that part of Wisconsin under¬
lain by the ancient crystalline and associated rocks had not
participated in the active settlement of Wisconsin. Only a
scant 10,000 were in the region in 1880, but ten years later
the population was over 85,000, so the settlement of that
section of Wisconsin north of the latitude of the northern
boundary of Marathon County belongs to a later period
than the interval required for the agricultural occupation of
the crescent-shaped area north of the Lower Wisconsin and
Fox Rivers.
A small segment of the crystalline area embraced in
Clark, Marathon, Portage, Waupaca and Wood Counties
was settled before 1880, and from the standpoint of time of
settlement belongs with the regions to the south rather than
the more northern highland wilderness.
“ Charles D. Robinson. The Lumber Trade of Green Bay. Trans. Wis.
State Agr. Soc. 1858-59, pp. 404-405.
Smith — Distribution of Population in Wisconsin.
83
Population Changes in Older Wisconsin. Behind the
frontier there was an intensification that tended toward
filling the vacant spaces. The three counties in the lead
region increased very slowly from 1850 to 1870 when there
began a decrease in population. From 1870 to 1880 Grant
County decreased 127, Iowa 916, and La Fayette 1,380, an
aggregate decrease of 2,423 people. Lead, and after 1865
zinc, had attracted into the region more people than the
mineral and agricultural resources could support, and emi¬
gration became a necessary population readjustment.
In southeastern Wisconsin the decade of the fifties was a
period of active settlement. Several counties, particularly
those nearer the frontier, doubled in population. Dane
County increased from 16,639 in 1850 to 43,922 in 1860;
Dodge went from 19,138 to 42,818; Fond du Lac from 14,-
510 to 34,154; Manitowoc from 3,702 to 22,416; Winnebago
from 10,167 to 23,770; and Milwaukee from 31,077 to 62,-
518. In the decade of the fifties, older Wisconsin, or that
section south of the Lower Wisconsin arid Fox Rivers, re¬
ceived two-thirds of the people who contributed to the popu¬
lation increase. Not a single county showed a decrease.
Between 1860 and 1870 the increase continued, but the
section had less than half of the increase for the state. The
three counties of Kenosha, Ozaukee, and Walworth showed
a decrease in population. In the ten year interval from
1870 to 1880 the increase in population in the state was
260,827 and older Wisconsin accounted for less than a quar¬
ter of the total gain. It is obvious that the cutting edge of
the frontier was the site of the greatest increments to the
total population.
Population Growth and Readjustments 1880-1920
The history of the settlement of Wisconsin prior to 1890
is a narrative of human conquest of unoccupied lands. But
toward the end of the last century the new lands of Wiscon¬
sin had been so reduced in area that the frontier was prac¬
tically gone except in the more remote sections to the north.
In the United States Census for 1890 it is recorded that the
“Lumbering and mining interests have practically obliter¬
ated the wilderness of Michigan and have reduced that of
84 Wisconsin Academy of Sciences, Arts, and Letters.
Wisconsin to less than one-half of its former area.”39 This
decrease in the wilderness was accompanied by a reciprocal
increase in land in farms contingent upon the northward
Fig. 13. Cartogram showing the land in farms in 1920. Circles
are proportional to the size of the counties, and the black sectors
show the percentage of the land in farms.
movement of the agricultural population. The retarding in¬
fluence of the unfavorable geographical conditions was be¬
ing overcome, and the desire for new agricultural land was
3»U. S. Census. Population, 1890, p. XXVIIL
Smith — Distribution of Population in Wisconsin.
85
opening the north to settlement. “The census of 1890, which
notes the passing of the frontier, establishes a convenient
base from which to compute the pressure of that land short-
Fig. 14. Cartogram showing the general distribution of improved
land in 1920.
age which gradually brought the vast and fertile areas of
northern Wisconsin into requisition for general farming.”40
The distribution of land in farms is closely related to the
distribution of the rural population. (Figs. 12 and 13).
w Joseph Schafer. A History of Agriculture in Wisconsin, p. 139.
86 Wisconsin Academy of Sciences, Arts, and Letters.
The amount of improved land, and the proportion of the
land in farms that has been improved, are also closely re¬
lated to the number and distribution of the rural inhabi¬
tants. (Fig. 14). The maps showing the land in farms
and the improved land illustrate graphically the value of the
land for human occupancy.
Population Growth in Wisconsin. The settlement of
Wisconsin began about 1820, at a time when Pennsylvania,
one of the old states, already had a population of over a mil¬
lion. It required fifty years, from 1820 to 1870, for Wis¬
consin to reach the first million, and thirty years to add the
second, and probably the number will reach three million by
1930. 41
Population Growth in Wisconsin
1840 _ 30,945
1850 _ 305,391
1860 _ 775,881
1870 _ 1,054,670
1880 _ 1,315,497
1890 _ 1,693,330
1900 _ 2,069,042
1910 _ 2,333,860
1920 _ 2,632,067
1922 _ 2,708,85842
1923 _ 2,739,57442
1928 _ 2,953,00042
In the decade of the fifties the number added to the pop¬
ulation of Wisconsin was 470,490, a total greater than any
other ten year period in the history of the state. In the
decade of the sixties only 278,789 were added, and in the
seventies the declining rate of increase added but 260,827.
Between 1880 and 1890 the stream of immigration contrib¬
uted large numbers to Wisconsin’s population, for the in¬
crease was 377,838. This was maintained in the nineties
for an increase of 375,712. The two decades of the twen¬
tieth century added respectively 264,818 and 298,207 to the
population. (Fig. 15).
The continued growth of population has been main¬
tained, not only by an excess of births over deaths, but by
a continued immigration from foreign countries and from
the older states. In 1920 the native born Americans num¬
bered 2,171,582 out of the total 2,632,067 people.
41 Compare with W. O. Hotchkiss. Geography and Industries of Wiscon¬
sin. The Wisconsin Blue Book, 1925, p. 46.
42 Estimates by the U. S. Census Bureau for July 1 of the respective years.
Smith — Distribution of Population in Wisconsin.
87
The westward movement that brought so many people
from the northeastern quarter of the United States waned
somewhat with the passing of the new lands in Wisconsin,
but did not become extinct. The migration along the par¬
allels had gained a momentum that continued in spite of the
counter current that contributed so materially to the
growth of cities.
Fig. 15. Seventeen counties showed a decrease in population be¬
tween 1910 and 1920. (Courtesy of the Geographical Review pub¬
lished by the American Geographical Society of New York).
The movement of native Americans to Wisconsin has
been offset by emigration in recent years. There were liv¬
ing in the United States in 1920 a total of 2,460,101 Wis¬
consin-born people, of which 1,852,574 were still living in
the state. Over 600,000 Wisconsin natives had left the
state, but the loss was offset by a number, almost twice as
great, that had come to Wisconsin from other states. Emi¬
gration from Wisconsin has been to the adjacent states
where there is always a mutual exchange of peoples, to the
older northeastern states that contributed such a large num¬
ber to the early stock, and to the new western states. The
exodus to the western states began about 1850 when the
California gold rush stimulated the trans-Rocky Mountain
‘trek' to the Pacific states. Wisconsin then had much un¬
occupied good land, and emigration did not take many
88 Wisconsin Academy of Sciences , Arts, and Letters.
people from the state. The passing of the easily cleared
and prairie lands about 1880, and the availability of new
government lands in the west caused the emigration to the
trans-Mississippi states. This large number of emigrants
is indicative of the passing of the frontier in Wisconsin.
Population Decrease . Population growth in Wisconsin
has been attended by a readjustment in distribution that re¬
flects the influence of geographic and economic conditions.
Between 1880 and 1890 six counties showed a decrease.
These were Fond du Lac, Grant, Iowa, LaFayette, Ozaukee,
and Washington, all in the section that could be called older
Wisconsin where a shift in population was a normal read¬
justment, and in certain sections the decrease was only tem¬
porary. In the last decade of the century practically every
county showed an increase, only Fond du Lac continued to
decline.
Between 1900 and 1910 nineteen counties declined in pop¬
ulation, and most of these were located in western Wiscon¬
sin, which was settled between 1850 and 1890. Now a
wave of emigration was carrying away the people.
The oldest part of Wisconsin was the first to be affected
by emigration. Green County had its maximum popula¬
tion in 1870 when there were 23,611 people in the county.
The maximum population for Iowa County was recorded in
1870. Similar conditions existed in LaFayette County.
Emigration from the lead region set in about 1870, a few
years earlier than in most parts of older Wisconsin.
In the decade from 1910 to 1920 the population of Wis¬
consin increased 12.8 per cent, but seventeen counties re¬
corded decreases. This continued increase in population
with the shifts from the rural to the urban communities is
indicative of important readjustments in the social order.
(Fig. 16).
The two sections of Wisconsin that have suffered most in
the withdrawal of people are the Western Upland and the
Central Lowland. Under the economic conditions then
prevailing, the excess of births over deaths and continued
immigration produced a pressure of population upon a land
unsuited to continued economic development. As a result
an emigration movement set in about 1900 — locally even
Smith — Distribution of Population in Wisconsin.
89
earlier — and has removed many people from the farms and
smaller villages. The new farm lands in the west and the
greater opportunities in the larger urban centers have ab¬
sorbed the emigrants.
Counties such as Walworth, Ozaukee, and Kewaunee,
within the industrial shadow of Milwaukee and other large
cities, have experienced a decline in population. In Ozau¬
kee County even the city of Port Washington decreased
from 8,792 in 1910 to 3,010 in 1920.
Fig. 16. Twenty-eight counties showed a decrease in rural popu¬
lation between 1910 and 1920. (Courtesy of the Geographical Re¬
view published by the American Geographical Society of New York) .
The only county in northern Wisconsin to show a de¬
crease was Vilas, which declined from 6,019 in 1910 to 5,649
in 1920. The very small population was materially reduced
by the removal of the transient lumbermen with the pass¬
ing of the pinery. In general, northern Wisconsin has been
increasing in population since the first pioneers threaded
their way through the forests. The decline of the lumber
industry did not seriously affect the total population, for the
farmers followed close upon the lumbermen. However, in
Vilas County the high percentage of sand and swamp land
has retarded the agricultural occupancy of the land.
Urbanization in Wisconsin. In Wisconsin as in many
other states there has been a tendency toward urbanization
90 Wisconsin Academy of Sciences, Arts, and Letters.
of the population. In 1890 two thirds, or 68.8 per cent, of
the people were rural and 33.2 urban, a condition differing
only slightly from the average for the whole country; at
that time the people of the United States were classified as
64.6 per cent rural and 35.4 urban. Ten years later, at the
beginning of the twentieth century, the average for the
country was 60.0 per cent rural, and 40.0 per cent urban.
Similarly Wisconsin showed an increase in the proportion
of urban population ; the rural inhabitants made up 61.8 per
cent and the urban 38.2 per cent. Wisconsin has lagged a
little behind the United States in the movement of the peo¬
ple to the cities. In the nation 54.2 per cent were rural
and 45.8 per cent urban in 1910, but in Wisconsin 57.0 per
cent remained rural and 43.0 per cent urban. The census
of 1920 indicates that industrial development continued to
draw the people away from the rural communities to the
urban centers.
This change in the division of the population does not
mean that the rural population decreased. As a matter of
fact the rural population of Wisconsin has steadily in¬
creased.
Rural Urban Total
1890 _ 1,131,044 562,286 1,693,330
1900 _ 1,278,829 790,213 2,069,042
1910 _ 1,329,540 1,004,320 2,333,860
1920 _ 1,387,499 1,244,568 2,632,067
Because of the nature of the physical habitat, Wisconsin
combines within its borders an older settled section with a
frontier area in such a way that the population changes in
the state approximate those of the nation. Concomitant
with the expansion of agriculture in the north there has
been a decrease in the rural population in the older sections.
However, the net result of this change has been an increase
in the rural population. From 1900 to 1920 the number of
rural inhabitants increased at the rate of 5,000 per year.
In the same two decades the urban population increased
from 790,213 to 1,244,568 or a rate of over 21,000 per an¬
num for the first decade and over 24,000 per annum for the
latter.
The city of Milwaukee had, in 1920, 17.4 per cent of the
Smith — Distribution of Population in Wisconsin.
91
population of the state, and if the metropolitan district be¬
yond the city limits is included the proportion is increased
to 20 per cent. (Fig. 17).
The nine cities that had over 25,000 each contained al¬
most 29 per cent of the population. According to the clas¬
sification by the 1920 census there were a total of 82 urban
centers in the state, having an aggregate population of
1,244,568 or 47.3 per cent of the total. In addition there
were 271,900 people living in the 375 incorporated cities
and villages having less than 2,500. These are classed as
Fig. 17. The growth of population in Wisconsin has been steady
in spite of sectional decreases. (Courtesy of the Geographical Re¬
view published by the American Geographical Society of New York).
rural by the Census Bureau, but they constitute 10.3 per
cent of the population and when added to the 47.3 per cent
swell the total to 57.6 per cent. These may be considered
as strictly urban, reducing the rural element to only 42.2
per cent of the total population of the state.
The accompanying dot maps show graphically the dis¬
tribution of the principal cities and villages. Lake Michi¬
gan has been of major importance in concentrating the pop¬
ulation.
On the western boundary of Wisconsin is another waterway, once
regarded as an important traffic route; but the usefulness of the Mis¬
sissippi River to Wisconsin makes a poor showing when compared
with the Great Lakes. Ten manufacturing cities have grown up along
92 Wisconsin Academy of Sciences , Arts , and Letters.
the shore of Lake Michigan (including Green Bay) but only two on
the Mississippi River . . ,43
The five largest cities are Milwaukee, Racine, Kenosha,
Sheboygan and Manitowoc. From the time of the settle¬
ment of eastern Wisconsin down to the present the lake
shore cities have attracted a high percentage of the popu¬
lation. The advantages of water transportation caused the
selection of urban sites along the lake. Later when the
railroads came competing for the traffic, which had been al¬
most exclusively handled by lake carriers, the growth of
cities was in no way retarded. “These lake ports were the
termini from which the early railroads pushed their way
into the back country ; consequently, they became the gate¬
ways through which the products of farm and forests pro¬
ceeded to market . . .”44
The Fox-Winnebago Valley is second only to the Lake
Michigan shore as a contributing factor in the growth of
urban communities. From the time of Nicholet’s visit in
1634 down to the present the Fox River Valley has been an
important highway. The easy portage between the head¬
waters of the Fox and the Wisconsin made the Fox River
one of the principal routes followed into the interior of Wis¬
consin during the French and English regimes. Hardly
had the agricultural settlers entered the valley before there
came a demand for an improved waterway to give them the
full advantages of the river which nature had not made en¬
tirely navigable. The river was improved only to meet the
competition of the railroads, and its usefulness as an artery
of commerce declined. However, the nuclei of the present
cities had been formed and other advantages within the
valley contributed to urban development. The availability
of water power along the Lower Fox has been of major
importance in the growth of several cities. Green Bay
is — and always has been — a commercial city, and owes its
growth and importance to its strategic situation near the
head of the bay bearing the same name. Many other cities
as De Pere, Wrightstown, Kaukauna, Kimberly, Appleton,
48 R, H. Whitbeck. The Geography and Economic Development of South¬
eastern Wisconsin. Wis. Geol. and Nat. Hist. Survey, Bull. No. 68, 1921,
p. 31.
44 Ibid p. 32.
Smith — Distribution of Population in Wisconsin. 93
Menasha and Neenah, are located on water power sites.
Oshkosh on the western shore of Lake Winnebago is located
at the point where the Upper Fox River enters the lake.
This is a favorable location, for during the high tide of the
lumber industry Oshkosh became the center of the industry
that flourished along the Upper Fox and Wolfe Rivers.
Fond du Lac at the southern end of Lake Winnebago en¬
joys the advantage of a site favorable to commercial de¬
velopment.
In the lumbering days Fond du Lac derived a large advantage from
the water transportation afforded by the lake. While its recent
growth has but little connection with lake transportation, yet the lake
is the main natural factor, though an indirect one, in the city’s devel¬
opment. This comes about through the influence which the lake ex¬
erts upon the railway routes. The steep bluff of limestone along the
eastern shore of the lake renders that side less suited to the growth of
towns, and hence less attractive to railroads, and so the north-and-
south railways follow the west side of the lake. Lake Winnebago is
a barrier to east-and-west lines, and any such lines must bend north
or south around the lake. This causes Fond du Lac to be something
of a converging point for railway lines . . .45
In 1920 the eight cities along the Fox-Winnebago Low¬
land having more than 5,000 people, had a population of
132,668, or almost five and a half per cent of the total for
the state.
In southern Wisconsin along the Rock and Yahara River
Valley have grown up several important, though smaller,
cities of Wisconsin. Madison, the capital is the largest.
The others are Janesville, Beloit, Watertown, Fort Atkin¬
son, Jefferson, Edgerton, and Stoughton.
Along the Wisconsin River a chain of cities reflects the
usefulness of the river as a means of transportation and as
a source of power in the lumbering and wood using indus¬
tries. Northward from Portage, in order, are Wisconsin
Rapids, Stevens Point, Wausau, Merrill, Tomahawk and
Rhinelander.
The Chippewa with its two important cities of Eau Claire
and Chippewa Falls reflects on a smaller scale the same his-
torv of urbanization as occurred along the Wisconsin River.
48 R. H. Whitbeck. The Geography of the Fox-Winnebago Valley. Wls.
Geol. and Nat. Hist. Survey, Bull. No. 42, 1915, pp. 42-43.
7
94 Wisconsin Academy of Sciences, Arts, and Letters.
La Crosse is the only city of any size on the Mississsippi
River, although Prairie du Chien is an older, but smaller
city, that owes its early importance to the advantage of a
strategic position at the junction of the Wisconsin with the
Mississippi River. The Black River, like the Chippewa
and Wisconsin, drains the northern pinery, and La Crosse
at its mouth flourished during the lumbering period. Its
position on the Mississippi is at a point where the east-west
lines of transportation, after crossing the Western Upland,
reach the city by following the La Crosse River Valley.
Only two cities of any size have grown up on the Wiscon¬
sin shore of Lake Superior. These are Superior and Ash¬
land which had in 1920 a population of 39,671 and 11,334
respectively. Superior had 40,384 in 1910, a slight de¬
crease from the previous enumeration. Ashland decreased
from 13,074 in 1900 to 11,594 in 1910, and a further de¬
crease brought the total down to 11,334 in 1920. These
two cities not only serve a local community, but the more
extensive hinterland which has been the most important
factor in their growth. The passing of the lumber indus¬
try has produced a slight and temporary decrease in the
population, but continued agricultural expansion in north¬
ern Wisconsin will furnish a new basis for a revival in
growth.
In addition to the lake shore and river sites which have
been so important in the localization of the major urban
communities, there are numerous situations which are ideal
for smaller cities, which serve as collecting and distributing
centers each for a local community. Many of these smaller
cities as originally planned were expected to grow into
large cities. In pioneer days these sites were purchased by
speculators and held for prices above the government fig¬
ure. Many of these so called ‘paper towns' have become
the villages and the smaller cities, except in the cases where
the sites have proved to be favorable to continued popula¬
tion growth. The dot maps present a vivid picture of these
smaller urban communities.
One important result of the urbanization in Wisconsin
is shown by the movement of the center or oopulation
southward and eastward after 1900. (Fig. 18) . For each
of the last five census enumerations the center nf popula-
Smith — Distribution of Population in Wisconsin.
95
lion has been in Marquette County. From 1880 to 1900 it
moved almost directly northward, but only half as far in
the nineties as in the eighties. Settlement along the fron¬
tier had slowed down by 1890. From 1900 to 1910 the cen¬
ter moved eastward and slightly southward. This indi¬
cates clearly that the intensified urban growth more than
balanced the northward movement. In the next decade
from 1910 to 1920 the movement was slightly southeast¬
ward, illustrating Wisconsin's trend toward an industrial
state.
I WAUSHARA
i
• ?
! mooo'TlfeTo
j 1890® 7i920
• •
! uj
I 1880© r
i i GREEN
[marquetteJ. LAKE
Fig. 18. A map showing the movement of the center of popula¬
tion since 1880. Note the eastward and southward movement since
1900. (After the Statistical Atlas of the United States, 1924).
Regional Readjustments
Population distribution in Wisconsin can be effectively
studied by dividing the state into regions, each of which has
rather uniform environmental conditions over the entire
area, or because its history deserves especial examination.
(Fig. 19).
The federal census classifies all cities and villages under
2500 as rural, but in this regional study the people have
been separated on the basis of the incorporation of cities
and villages. All people living in incorporated districts
are classed as urban, and the remainder as rural. This di¬
vision, while still more or less arbitrary, separates more ex-
96 Wisconsin Academy of Sciences , Arts , and Letters .
actly the agricultural inhabitants from those who derive
their livelihood from other occupations. It is true that the
small incorporated villages have within their limits many
people who are strictly agricultural. Similarly, there are
Fig. 19. There is a population sectionalism in Wisconsin. The
provinces as delineated on this map permit a statistical examination
of the distribution of the rural inhabitants in particular.
unincorporated hamlets with many people engaged in occu¬
pations not directly concerned with farming. These two
cases mutually offset each other and probably balance any
apparent inaccuracy.
Smith — Distribution of Population in Wisconsin.
97
The dot maps were constructed from data as reported in
the United States Census, and all people living in cities and
villages having a population less than 200 have been shown
as rural. But there are many hamlets and larger towns
having a population of more than 200 — a few over 1000 —
which, if shown as rural, would give an inaccurate picture.
Other sources have been consulted, such as local histories,
Polk’s gazetteers, and particularly the many annotated
maps in the collections of the State Historical Society of
Wisconsin. In the newer parts of northern Wisconsin the
lumbering towns grew so rapidly that they attained a size
much larger than many of the incorporated villages of older
Wisconsin, yet they remained unincorporated, perhaps
awaiting the time when the lumbering industry would re¬
move many of the people to newer towns in the virgin for¬
est.
The Lead Mining Region. The early settled lead mining
region of Wisconsin experienced the population readjust¬
ments characteristic of mining areas. A general increase
continued up until 1870 when the maximum number of in¬
habitants was recorded. Between 1890 and 1900 there was
a slight increase from 79,033 to 82,954 for the three coun¬
ties of Grant, Iowa and La Fayette. After 1900 a decrease
brought the total down to 80,550 in 1920. If the number
living in cities and villages is deducted from the totals as
reported by the census the decrease in the rural people is
more marked. In 1890 there were in these three counties
18,534 who lived in incorporated cities and villages, and the
remaining 60,499 lived in the rural districts. The subse¬
quent decennial enumerations for 1900, 1910 and 1920 re¬
ported respectively 27,685, 28,022, and 30,705 living in
cities and villages. Corresponding to this increase there
was a reciprocal decrease in the rural peoples. In 1900 the
rural population was 55,269; by 1910 the number declined
to 53,557 and in 1920 to 49,845. Part of the increase in the
urban and the corresponding decrease in the rural popula¬
tion was due to the incorporation of six villages having a
total of 1928 people. The density of the rural population in
this section was only 19.2 persons per square mile in 1920
and the Agricultural Census of 1925, while not exactly com¬
parable, indicates a further decrease to 15.5 per square mile.
98 Wisconsin Academy of Sciences , Arts, and Letters.
The old lead mining section of Wisconsin was once the
goal of many pioneers, but a century has wrought changes
of far reaching importance. Like the rest of western Wis¬
consin the land is no longer attractive to homeseekers and
a decrease in rural people is the important change to be
noted. The land is only slightly less valuable than it was
a hundred years ago. General farming with dairying as
the specialized accompaniment has maintained the produc¬
tivity of the soil. In the century that lias elapsed since the
first settlers penetrated the region in search of lead, the
land has been continually improved. But the changing ec¬
onomic conditions of the country, and the use of improved
labor saving farm machinery, have made it possible for
fewer farmers to do the farming and at the same time in¬
crease production.
South Central Wisconsin. The counties drained by the
Rock River, and adjacent sections may be called South Cen¬
tral Wisconsin. It is the Rock River Valley that unifies the
region, though Green, Walworth and Columbia Counties
are on the periphery of the area. These may be considered
as transitional regions. Columbia County for example has
a sandy portion along the Wisconsin River that makes it
resemble very much the Central Plain, but the eastern part
is more densely populated and the section is capable of sup¬
porting a larger population than the western part. Simi¬
larly Green County reflects a condition more like the three
lead mining counties but Green County has never been a
producer of lead. It is not only midway between the old
lead mining country and the Rock River Valley but it is
transitional in population conditions. Walworth, between
the lake shore counties and Rock River Valley has suffered
an isolation that is comparable to the handicap imposed
upon Green and Columbia Counties. These three counties
have shown very little growth in population since 1890. In
fact Green County showed a steady decrease in total popu¬
lation. Dodge and Jefferson with a location similar to that
of Walworth showed increases of less than the average for
the state.
Dane and Rock Counties which include the three largest
urban centers in south central Wisconsin have shown the
greatest growth in the three decades under discussion.
Smith — Distribution of Population in Wisconsin.
99
Madison, the state capital, and Beloit and Janesville, indus¬
trial cities on the Rock River have absorbed the major por¬
tion of the increase in population of the two counties since
1900. Between 1900 and 1920 the total population of the
seven counties increased from 285,157 to 321, 709, a gain of
36,552. The aggregate population of Beloit, Janesville and
Madison increased from 42,785 to 77,955 in the same pe¬
riod, a gain of 35,170 or 1382 less than the total increase
for the entire area.
The result of this urbanization has been a gain in popu¬
lation greater than the average rate of increase for the en¬
tire state between 1910 and 1920. After 1900 there has
been an exodus from the farms that has reduced the strictly
rural population from 166,174 to 140,102 in 1920. The
land has ceased to be of fundamental importance in the
growth of population.
The Lake Shore Region. The lake shore counties ex¬
tend from the Illinois boundary to Door Peninsula, a dis¬
tance of more than 200 miles. Washington and Waukesha
Counties are included although they do not touch the lake.
Because of the latitudinal extent of the region it may be
subdivided into a northern and a southern section, but the
place of division is more or less arbitrary.
Door and Kewaunee Counties to the north are agricul¬
tural and contain no large urban centers. Sturgeon Bay,
in Door County, is the largest city and in 1920 had a popu¬
lation of 4553. The configuration of the peninsula limits
the commercial hinterland to the extent that this eastern
projection of Wisconsin has been passed~by. Farther
south Manitowoc and Sheboygan Counties, because of a
more favorable physical landscape and better commercial
opportunities, have a much larger population. Sheboygan
illustrates the growth in population, for the number in 1890
was 42,489, in 1900 it had increased to 50,345, in 1910 to
54,888 and in 1920 the total was 59,913. In contrast the
last four decennial enumerations have recorded for Kewau¬
nee successively 1890-— 16,153, 1900 — 17,212, 1910 — 16,784,
and 1920 — 16,091. Kewaunee is one of the fifteen counties
that showed a net decrease from 1900 to 1920, and one of
the seventeen recording a decrease from 1910 to 1920.
The southern lake shore counties showed a remarkable
100 Wisconsin Academy of Sciences , Arts , and Letters.
growth from 1890 to 1920 increasing from 358,914 to 754,-
354. The large total and the marked growth are due to the
urbanization that has concentrated so many people in the
three cities of Milwaukee, Racine and Kenosha. The
growth of these cities has caused the removal of rural peo¬
ple from some of the adjacent counties. Ozaukee increased
slowly from 1890 to 1910 but in the decade from 1910 to
1920 a decrease reduced the total below the number for
1900.
Washington and Waukesha Counties because of their in¬
land location did not grow so rapidly as the three southern
lake shore counties, but their proximity to Milwaukee is an
advantage and not a handicap. This German settled sec¬
tion with its small farms averaging less than 100 acres has
the densest rural population in the state. Urbanization
has produced a decrease in the rural inhabitants, but in
1920 Washington had 41.3 persons per square mile, and
Waukesha had 48.7.
Milwaukee, the smallest of the counties had the largest
population in 1920. The city of Milwaukee, the metropolis
of the state had in 1920 a population of 457,147, over 17 per
cent of the total population of the state. If the metropoli¬
tan district is included the proportion becomes 20 per cent.
Both Racine and Kenosha Counties, like Milwaukee, have
an important urban population. The city of Kenosha in¬
creased from 6,532 in 1890 to 11,606 in 1900, to 21,371 in
1910, and to 40,472 in 1920. Racine the second largest city
has had a similar growth, reaching a total of 58,593 in
1920.
All of this lake shore section except the northern penin¬
sula has maintained a large rural population. The geo¬
graphical conditions are favorable to a dense agricultural
population, but there can be no doubt that a social factor is
of some importance, for many of the German immigrants
purchased small farms from 40 io 80 acres, and from the
beginning of settlement this section has been very densely
peopled. (Fig. 20).
The Fox-Winnebago Basin . The six counties included
in the Fox-Winnebago Basin constitute a convenient unit.
Whitbeck included only Brown, Outagamie, Winnebago and
Fond du Lac in his Geography of the Fox-Winnebago Val-
Smith — Distribution of Population in Wisconsin . 101
ley,46 but Green Lake and Calumet are included in this area
because of their proximity and the difficulties involved by
including them respectively in the Central Plain or the
Lake Shore provinces.
These six counties showed a consistent gain in popula¬
tion from 1890 down to 1920, increasing from 203,841 to
269,121 a gain of 55,280, or over 1800 per annum. At the
Fig. 20. The small farms of less than 100 acres are found along
the German-settled Lake Shore counties and along the frontier in
northern Wisconsin. (Courtesy of the Geographical Review pub¬
lished by the American Geographical Society of New York).
beginning of the period the rural exceeded the urban pop¬
ulation, but by 1920 the urban inhabitants had been in¬
creased to almost two-thirds of the total. In 1920 the eight
principal cities had almost half of the population.
Accompanying the urbanization there has been a recipro¬
cal decrease not only in the proportion of rural population,
but in the aggregate. From 1890 to 1920 this section has
lost approximately 360 persons per annum from the farms.
Like all of older Wisconsin, the passing of the frontier has
brought in its wake a depopulating of the older agricultural
sections. Still the average density of the rural population
remains over 30 per square mile, less than in Washington
** Wis. Geol. and Nat. Hist. Survey. Bull. No. 42.
102 Wisconsin Academy of Sciences, Arts, and Letters.
County and greater than in the more rugged Western Up¬
land.
The Central Plain. The seven counties of Adams, Ju¬
neau, Marquette, Portage, Waupaca, Waushara and Wood
are not entirely within the sandy Central Plain; Adams,
Marquette and Waushara belong to the sand country
and may be chosen to illustrate the influence of the land
conditions upon density of population. Since 1900 the pop¬
ulation of the seven counties has increased slowly, reaching
a total of 158,143 in 1920 or about 32 persons per square
mile, or only 20 rural inhabitants per square mile. The to¬
tal area of these seven counties is almost 5000 square miles.
The reduction in rural population was at the rate of 400 per
year for the two decades after 1900, which means that each
year every 12 square miles of area lost one rural inhabitant.
A comparison between Adams and Waushara illustrates
the influence of glaciation on the habitat. Adams County
is in the Driftless Area and Waushara in the glaciated re¬
gion. In 1920 the population of Adams County was 9,287
or 13.6 persons per square mile. If the people living in the
three incorporated villages are deducted the density was
only 11.3 per square mile. WTaushara, — like Adams in all
essentials except that it was glaciated, — had in 1920 a pop¬
ulation of 16,712 or 25.9 persons per square mile, or if only
the strictly rural population is considered the density was
almost 20 per square mile. It is a significant conclusion
that the Central Plain was benefitted by glaciation. Ev¬
ery square mile of the glaciated plain supported at least one
more family than did the driftless portion of the plain.
Western Wisconsin. The fourteen counties of western
Wisconsin north of the Lower Wisconsin River showed an
almost static condition of the total population after 1900.
From 1890 to 1900 the population increased from 307,313
to 341,984, a gain of 11 per cent as compared with the 22.2
per cent for the state. Between 1900 and 1910 there was a
slight decrease, but by 1920 an increase brought the aggre¬
gate up to 347,274, the maximum attained in any enumera¬
tion.
An examination of the cities and villages reveals quite a
different condition. In 1890 the urban population numbered
93,867, almost a third of the total. The steady increase in
Smith — Distribution of Population in Wisconsin . 103
the urban population has materially reduced the rural ele¬
ment. In 1920 the urban element made up 41 per cent of
the total. This change is due not only to urbanization
characteristic of older Wisconsin, but emigration has
played an important part, particularly between 1900 and
1910.
Vernon County represents a fair cross section of the
rugged and unglaciated portion of the Western Upland.
The region is maturely dissected, but there are preserved
upon the upland broad areas that were originally prairie.
The slopes are too steep for utilization other than for pas¬
ture and woodland. From 1890 to 1920 the increase in the
total population was only 4,141. In the same period the in¬
corporated cities and villages grew from 2007 to 7614, a
gain of 5607 ; the increase in the urban group not only ab¬
sorbed the total gain of the county in the thirty years but
effected a decrease in the rural population. The emigra¬
tion from the farms averaged almost 49 people per year for
the 30 year period from 1890 to 1920.
In Vernon County as in most of the Western Upland the
momentum of settlement imposed upon the land too many
people who looked to it as a source of livelihood. The de¬
cline in the rural population is a normal readjustment, and
a continued decrease would help to relieve the agricultural
situation.
Northern Wisconsin. The newer northern part of Wis¬
consin contained after 1880 much land that was still to wit¬
ness the coming of the agricultural pioneers. The lumber¬
ing industry was at the height of its activity preparing the
way for the farmers who were cautiously penetrating the
wilderness and taking up the new lands.
The sixteen modern counties constitute a frontier section
of the state where the growth of population repeats the con¬
ditions of a half century ago when southern Wisconsin was
on the frontier. The growth in population is accounted for
by an increase in both the rural and urban elements, with
the rural increase exceeding the urban.
Adjacent to these sixteen counties of northern Wisconsin
are eight others that occupy a crescent shaped area near
the southern boundary of the Northern Highland. The to¬
tal area of these eight counties is somewhat smaller than
104 Wisconsin Academy of Sciences , Arts , and Letters.
the more northern counties but the population conditions
are strikingly different. In 1880 the northern counties had
only 11,710 people as compared with the 89,517 in the bor¬
der counties. At the subsequent decennial enumerations
the border counties failed to maintain this large margin in
population. By 1900 the border counties had a total of
222,790 and the northern counties 161,698, and two decades
later (1920) the border counties had 293,452 and the north¬
ern, 260,743. The two areas showed a growth of both ru¬
ral and urban population. Ashland and Superior, the two
largest cities in northern Wisconsin and serving a larger
hinterland than the adjacent tributary area, made up a
large proportion of the population. These two lake shore
cities contained half of the urban population in 1890.
The census enumerations for 1900, 1910 and 1920 show
that both the urban and the rural populations grew steadily
until there were 260,743 in the northern section in 1920.
Of this number 113,117 were urban, with the combined
population of Superior, 39,671, and Ashland, 11,334, mak¬
ing up almost half of the total, a condition that obtained 40
years previously. In 1900 the urban population in the bor¬
der and the northern sections was approximately equal, but
by 1920 the people living in incorporated cities and villages
numbered 113,117 in the northern counties and 102,511 in
the border counties. This leaves a larger rural population
in the smaller border section where agricultural conditions
are better.
The value of the cut-over land in northern Wisconsin has
been frequently over-stated. The land companies are
anxious to dispose of the land to settlers, and exaggeration
of the real value has caused much hardship. However,
there have been made more reliable statements that have
served to make some of the settlers more cautious in the se¬
lection of land. In 1896 Henry47 estimated that one to one
and a half million acres have swamp or humus soils and,
therefore, unsuited to agriculture. Interspersed between
the patches of wet and otherwise inferior land are exten¬
sive areas of valuable land which await the coming of far-
47 W. A. Henry. A Handbook for the Homeseeker, p. 6.
Smith — Distribution of Population in Wisconsin. 105
mers who are willing to devote a lifetime to the removal of
the forest.
In recent years a more conservative estimate of the
northern land deserves a wider circulation than it has re¬
ceived. Packer and Gunderson estimate that “forty per
cent of our unoccupied cut-over land is unsuited to farm¬
ing . . .”48 One of the tragedies of agricultural settle¬
ment on the poor land is a realization after a few years of
the futility of labor. A farmer and his family may have
come hundreds of miles to a new home which has been de¬
scribed to them as offering advantages and opportunities
approaching their greatest ambitions. When all of their
resources are invested in the land they cannot afford to
leave nor can they see a prosperous future. In a few years
it becomes home to them. Though they remain poor strug¬
gling farmers, their sentimental attachment to the soil is
too firm to be broken by changing economic conditions.
They find consolation in the hope that increased land values
will repay them in some measure for their years of toil.
In spite of the handicaps imposed by a more or less hos¬
tile environment the better farm lands of the northern part
of the state have been brought under cultivation. Steadily
the agricultural frontier encroaches upon the cut-over land,
and the increase in improved land is most marked along
the crescent margin of the northern highland, the area al¬
ready referred to as the border counties. The settlement
along the periphery of the northern section has been de¬
layed because the Central Plain acted as a deterrent.
It has been said that if all the settlers who in the past 50 years have
been sold land in the sandy central plain of Wisconsin and have failed
in consequence had been taken a few counties farther north and sold
some of the good silt loam lands, most of the good land in northern
Wisconsin would now be in farms. In Wisconsin in particular the
crescent-shaped . . . plain stretching all the way from Burnett
County in the northwest to Marinette County in the northeast, and
more than two counties wide in the center, has served as a very ef¬
fective barrier against development north of it.49
48 B. G. Packer and Oscar Gunderson. Preliminary Biennial Report of the
Wisconsin Department of Agriculture, Immigration Division, Madison, 1923-
1924, p. 45.
49 J. D. Black and L. C. Gray, Land Settlement and Colonisation in the
Great Lakes States, U. S. Department of Agriculture. Dept. Bull. No. 1295,
Washington, 1925, p. 12.
106 Wisconsin Academy of Sciences, Arts, and Letters .
The settlement of the more northern part of Wisconsin
has been in progress for half a century. The agricultural
expansion began about 1880 yet only about six per cent of
the area yields harvested crops. “At this rate it would
take over 750 years to complete the conquest of the north¬
ern woodland.”50 It is obvious that the settlement of north¬
ern Wisconsin in recent years resembles only slightly the
frenzied western migration that began the settlement of
older Wisconsin a century ago.
Conclusion
In the settlement of Wisconsin there has developed a sec¬
tionalism in the distribution of the popv lation which re¬
flects the physical conditions of the landscape, the soil, the
climate, as well as the social elements. The changes
wrought in almost three centuries of history present a se¬
quence of cultural stages. The type of people and the
stage of their civilization determine the use that they will
make of their regional habitat and the number that a given
area may support. “ . . . each mode of life has its
own space requirements which are larger for the hunter or
shepherd than for the agriculturist . . .”51
During the period of exploration, Wisconsin supported a
few hundred whites and a few thousand aborigines who
hunted, almost to extermination, the fur-bearing animals.
The effects of the glacial climate of a past geological period
persist into the present, creating an ideal habitat for the
fur-bearing animals. For more than a century a few thou¬
sand people in a primitive stage of civilization lived in Wis¬
consin and won a livelihood from the fur industry. Only
when the agriculturalists with their higher cultural aims
came into possession of the land was the intrinsic value of
the soil fully realized. Gradually the development of com¬
merce, manufacturing and service occupations has pro¬
duced readjustments in population distribution in keeping
with the cultural progress of mankind.
80 B. H. Hibbard, John SWenehart, W. A. Hartman and B. W. Allin, Tax
Delinquency in Northern Wisconsin. Agricultural Experiment Station.
University of Wisconsin, June 1928, p. 9.
81 Paul Vidal de la Blache. Principles of Human Geography, p. 155.
Smith — Distribution of Population in Wisconsin . 107
Beneath the film of population unevenly spread across
the regional landscape of Wisconsin the effects of soil,
drainage, relief, and climate persist as the eternal economic
bases of the relation of man to his habitat. But it must be
remembered that man, because of his intelligence, can exer¬
cise judgment in his relations to his physical environment.
He is not as specialized as the lower animals, “ . . .
for his mentality and communal co-operation are such that
he alone of the animal kingdom can produce artificial har¬
mony between himself and the environmental complex
where disharmonies actually occur.”52 Furthermore, social
inheritances may impel man to act in certain ways while
his neighbor, because of a different inheritance, behaves
differently. In spite of the influence of social conditions
“It is more than probable that soil and climate influence the
distribution of men on the surface of the globe, and facili¬
tate or hinder their concentration or dispersion. It is for
demography to undertake the study of both that concentra¬
tion and that dispersion.”53 In Wisconsin the settlement and
the distribution of the inhabitants reflect in a variety of
ways the significance of the frontier and the persistence of
the regional geography in the development of a population
sectionalism.
University of Illinois
Urbana, Illinois.
62 Raymond Swann Lull. The Ways of Life, p. 244.
68 Lucien Fe'bvre. A Geographical Introduction to History, p. 34.
GLACIAL GEOLOGY OF PART OF VILAS COUNTY,
WISCONSIN1
F. T. Thwaites
Introduction
This study of the glacial geology of part of Vilas County,
Wisconsin, was begun on September 23 and was terminated
on October 30, 1927. The area surveyed is shown on the
accompanying map (fig. 1). It comprises Townships 40 to
44, inclusive, and all of Ranges V, VI, VII and part of
Range VIII East. On account of the diagonal location of the
state boundary, this area includes only about fourteen and a
half townships, or approximately 522 square miles. Many
lakes and lakelets are situated within the area and this
study of the glacial geology of the region was carried out in
connection with an extended physical, chemical and biologi¬
cal investigation of these lakes. Only a few of the larger
lakes are shown in figure 1.
A study of glacial geology is primarily a study of topog¬
raphy and only secondarily a study of material. Exten¬
sive views of the surrounding territory are, therefore, of
chief importance, though exposures can by no means be
neglected. Since most of the region is covered with second
growth timber, this survey was made in the autumn after
most of the leaves had fallen from the trees ; even after the
leaves are almost gone, the density of the brush in many
places severely limits the field of observation. Work was
done preferably along roads and railroads, especially where
extensive views of the surrounding territory were available.
About 16 square miles per day were covered in the survey.
All of the maps of the region are extremely inaccurate
owing to inaccuracies in the original Government Survey.
Thus much more time must be spent at times in finding ex¬
act locations than in making the observations on the geol-
1 Published by permission of the State Geologist.
8
110 Wisconsin Academy of Sciences, Arts, and Letters .
ogy. Resurveys are being made in some parts of the area
and eventually more accurate maps will be available.
Acknowledgments. Aid was received from J. J. Mc¬
Donald, State Cruiser at Trout Lake ; Clarence Buck, Clerk
L_ Rv _ [ _ Rvj _ J _ R Vi i _ | Rvm _ |
HD Terminal Moraine (HjOutwash ^ Drumlin
Fig. 1.
of the town of Winchester; and William F. Kunschki, As¬
sessor of the town of Winegar. Profiles of the principal
railways were furnished by W. L. Towne, Chief Engineer of
the Chicago and Northwestern Railway, and C. F. Loweth,
Chief Engineer of the Chicago, Milwaukee, St. Paul and
Pacific Railroad.
Thwaites — Glacial Geology of Wisconsin.
Ill
Elevations. From the railroad profiles as a base, ane¬
roid readings were extended over the entire area, with par¬
ticular attention to the levels of the lakes. Great care was
used in making these readings and it is believed that the
great majority of the results are correct to the nearest 10
feet. Table 1 shows the results of these observations.
Table 1. Elevations of lakes in Vilas County , indicated in feet above
sea level.
112 Wisconsin Academy of Sciences , Arts, and Letters.
Table 1. Elevations of lakes in Vilas County, indicated in feet above
sea level. — (Continued)
only three are known in the region surveyed. Two ledges
in secs. 34 and 35, T. 43, R. 7 E., northwest of High Lake,
were visited by the writer; the rock is a coarse gray and
pink granite with pegmatite dikes. The exposures are in a
pitted outwash plain although erosion by glacial streams
Thwaites — Glacial Geology of Wisconsin.
113
doubtless had a part in uncovering the rock. Allen and
Barrett1 report a ledge of gneiss between Spider and Island
lakes which was not visited by the writer.
Drill holes. During the late panic over iron ore reserves
the area was explored by Allen and Barrett for the F. I.
Carpenter syndicate. The general results of this work
have been published but not the detailed logs of the numer¬
ous drill holes. No exploration is now going on and it is
reported that some of the lands which were purchased have
since been sold. The drill holes were for the most part on
magnetic lines and found granite, quartzite, slate, iron for¬
mation, and various types of schist. Few of the published
logs give the depth of drift, but these data were secured
from the files of C. K. Leith. In this area it varies from
129 to 234 feet. The relief of the bed rock surface is,
therefore, not great. The explorers named some of the
concealed ranges of much altered iron formation; that
which passes through the village of Winegar is the Turtle
Range and the much more irregular magnetic belt south of
it they called the Manitowish Range. The latter seems to
be underlain solely by schist and gneiss. The prevailing
strike of folds and schistosity is about N. 70° E.
Inferences from drift. Virtually no unassorted glacial
drift is found in Vilas County south of the latitude of Crab
Lake. This fact renders conclusions as to the character of the
bed rock rather difficult to arrive at since a large part of the
material of the drift may have been transported consider¬
able distances by water in addition to its journey by ice.
The transportation by water also removed most of the fine
material derived from slates, shales, and soft iron forma¬
tion. The pebble and bowlder counts show that pink and
gray granites and pegmatites, many of which are probably
local, predominate. Basalt, both dense and amygdaloidal,
diabase, rhyolite, red sandstone, and red shale, all obviously
derived from the Keweenawan rocks to the north, make up
a large part of the pebbles. There are very few fragments
of quartzite and iron formation. The fine material is in
large part quartz sand which in the till is mingled with a
1 Allen, R. C., and Barrett, L. P., Contributions to the pre-Cambrian geol¬
ogy of northern Michigan and Wisconsin : Michigan Geol. and Biol. Survey
Pub. 18, pp. 65-130, 1915.
114 Wisconsin Academy of Sciences, Arts, and Letters.
considerable amount of red clay probably derived from the
red Keweenawan and Huronian rocks to the north. It is
not at all probable from these data that any large areas of
Huronian rocks exist in Vilas County. The drilling showed
that such as are present are much altered by intrusive gran-
its as well as by regional metamorphism. It is highly doubt¬
ful that any areas of merchantable iron ore can exist in the
area surveyed. The writer is convinced that the bulk of
the bed rock is granite and gneiss. The immense amount of
sand came from the Keweenawan or Cambrian sandstones
to the north.
Topography
Elevation. The highest known point in the area surveyed
is the hill on which Muskellunge Fire Tower is situated
(sec. 34, T. 41, R. 7 E.) which reaches an elevation esti¬
mated at 1825 feet above sea level. The lowest measured
point is Shishebogama Lake in T. 40, R. 5 E. at 1565 feet.
The general surface of the country declines from about
1700 feet at the northeast to about 1600 feet in the south¬
west; 1650 feet is a general average for the country sur¬
veyed.
Relief. Vilas County is a region of relatively low relief
(fig. 2). Local differences of elevation of much over 50
feet are not at all common although on the other hand ex¬
tensive flats are rare. The roughest portion of the area is
at the north. Here the hills are very irregular both in out¬
line and in summit elevation and local differences of 75
feet are common. Interspersed among these hills are many
enclosed basins, a large number of which contain lakes and
ponds. This is by all odds the most picturesque portion of
the area. Farther south the landscape is a broken plain
which offers much more monotonous scenery. Locally
small hills and ridges rise above the general level. The
lakes and swamps are set in partially or wholly enclosed
depressions which range from a few feet to 50 feet in depth.
Most of the lakes are shallow and small ; Trout Lake is the
largest and deepest. The total depth of its depression is
about 115 feet.a
2 Juday, C., The inland lakes of Wisconsin : Wisconsin Geol. and Nat.
Hist Survey Bull. 27, p. 129, 1914.
Thwaites — Glacial Geology of Wisconsin.
115
Drift Deposits
Vilas County is remarkable for the monotony of the gla¬
cial geology, that is, for the large size of the individual areas
of the same origin (fig. 1) . It is also noted for the simplicity
of the geology and the lack of features with a complex
glacial history. In spite of this fact, the geologist is com¬
pelled by the lack of extended views to traverse the region
rather fully lest some relatively small feature escape him
and thus make his rendering of the story incomplete. An
effort was made to visit every section unless obviously all
swamp or all plain.
Types of deposits. The drift deposits of the area sur¬
veyed can be divided into (a) outwash, (b) terminal (re¬
cessional) moraines, (c) drumlins, (d) ground moraine,
and (e) eskers. Of these, the first covers by far the larg¬
est portion of the region and the second forms the most con¬
spicuous topographic features and the most striking coun¬
try. The other features cover only an inconsequential per¬
centage of the region.
Outwash. The most widespread and characteristic drift
deposit of the lake region of Vilas County is outwash which
contains numerous kettles, that is pitted outwash .3 The
material is nearly all horizontally bedded sand which for the
most part contains scattered pebbles and a few bowlders.
Some small cross bedding is generally present. Fairly well
sorted, locally very bowldery gravels are present in some
places. The topography varies from level as southwest of
Boulder Junction to so much pitted that no upland is left
between the kettles; this last type is well shown in the vi¬
cinity of Witches Lake west of Sayner. In many places
the uplands between the kettles are small but when the geolo¬
gist stands on one he can see at once that the other summits
form the remnants of a once continuous plain. Many of the
kettles extend below the water table and therefore contain
marshes or lakes. The majority of the lakes of Vilas
County are of this origin. They have low sandy and in
most cases uninteresting shores. In the very much pitted
areas the resemblance to terminal moraine is striking, es-
8 Thwaites, F. T.f The origin and significance of pitted outwash : Jour.
Geology vol. 34, pp. 308-319, 1926.
116 Wisconsin Academy of Sciences , Arts, and Letters.
pecially where bowlders are present. Discrimination is not
difficult, for in such cases neither the coarse gravel nor
clayey till of terminal moraines is present. In many places
the deposits are terraced into two or more distinct levels,
all pitted (fig. 2). It proved impracticable to map the dis¬
tribution of such terraces over any extended area because
of the lack of accurate topographic maps.
Terminal moraine. Terminal moraine topography con¬
sists of knobs with intervening sags; there is neither a
level upland nor an equality of summit levels. The ter¬
minal moraines form elevations above the adjacent outwash
areas. The material of the moraines consists of glacial till,
ill-assorted gravel, sand, and red clay. Bowlders are con¬
spicuous in most terminal moraine areas. Where the land
is still covered with virgin timber with its accompanying
vegetable mould and fallen leaves they are not easily seen.
Three distinct moraines, and traces of a fourth have been
discriminated as shown on the accompanying map (fig. 1).
Of these only the northernmost, the Winegar moraine, con¬
tains a large amount of till. This till is red in color and
contains pockets of bowldery sand and red clay. Locally
the surface is covered with a few feet of pebbly sand. The
red till is bleached to a yellowish gray to depths of several
feet from the surface. Lakes are abundant in the kettles
and some of the finest bodies of water in the area, such as
Crab Lake, are found in this moraine. The other moraines,
the Boulder and the Muskellunge, are, so far as could be dis¬
covered, composed wholly of assorted material. They can
be distinguished from the adjacent outwash by the great
abundance of bowlders, the coarseness and ill-assortment of
the gravels, and by their topographic form of ridges trans¬
verse to the direction of glacial movement. Both of these
moraines are discontinuous and are represented in some
places by isolated knolls of bowldery composition which
rise from the adjacent outwash plains.
Drumlins. Drumlins were not known in this portion of
Wisconsin previous to the present survey but had been found
in Iron and Gogebic counties, Michigan, by Leverett4 so
4 Leverett, Frank, Surface geology and agricultural conditions of Michi¬
gan : Michigan Geol. and Biol. Survey Pub. 25, Plate I, 1917.
Thwaites — Glacial Geology of Wisconsin.
117
that their discovery should excite no surprise. Mapping
of drumlins is exceedingly difficult in forested country and
it is possible that more drumlins might be recognized were
conditions more favorable for observation. It is probable
that many other drumlins lie buried beneath the outwash
plains from which only the highest project.
The mapped drumlins lie west and northwest of Trout
Lake with a single outlying specimen just south of Highway
70 in T. 40, R. 5 E. Of these, the latter may very well be a
portion of a group most of which lies south of the area map¬
ped, for drumlins rarely occur alone. It also seems possi¬
ble that the stony ridge in the so-called Game Farm east of
Trout Lake may be a drumlin. The recognized drumlins
range from less than a quarter of a mile in length to over
three quarters of a mile. The width varies from a third to
a quarter of the length. The maximum known height is
about 100 feet. Some of the most accessible and perfect
drumlins are situated west of Boulder Junction near the
Manitowish Fire Tower. All the observed drumlins have
a trend of 30° to 40° west of south.
Ground moraines . No true ground moraine, that is thin,
rolling drift through which the older rock topography
shows, is present in the area surveyed. An area southeast
of Big Papoose Lake in T. 43, R. 6 E. is bowldery, gently
rolling, and is apparently underlain by till. It was map¬
ped as ground moraine because of the low relief, but its
origin is doubtless associated with the drainage from the
ice front at this point which eroded and leveled some of the
border of the Winegar moraine.
EsJcers. Eskers are the beds of glacial streams which
were confined by walls of ice. They consist of relatively
low discontinuous ridges of coarse, ill-assorted gravel. The
discovery of eskers in a forested region is a matter of
chance. Many eskers are probably buried under the out¬
wash and in a few places the ice blocks which formed the
kettles served to protect a portion of an esker from such
cover. Discrimination of esker remnants from accidental
ridges b’etween pits is a matter of examining the material.
Unless cuts are present, such examination is very difficult.
One of the best eskers which was discovered is that crossed
118 Wisconsin Academy of Sciences, Arts, and Letters.
by the old railway grade in sec. 10, T. 42, R. 6 E. It is possi¬
ble that the ridge of coarse gravel west of Crawling Stone
Lake is really a gigantic esker and not a moraine as mapped.
Glacial History
The glacial history of the region surveyed is, so far as
the evidence there observed goes, relatively simple. It tells
only of the last, or Wisconsin, glaciation which in this re¬
gion ended in a relatively rapid retreat of the ice front in¬
terrupted by three or four halts. The times of relatively
stationary margin resulted in the formation of successive
moraines. The comparative durations of the halts may be
estimated from the size of the respective moraines; this
criterion shows that the formation of the most northerly or
Winegar moraine took longest. During each halt floods of
water from the melting ice buried the country just vacated
beneath their load of sediment.
Direction of ice movement. The direction in which the
glacier moved in Vilas County is shown by (a) the direction
of the long axes of the drumlins and eskers, (b) the trend
of the terminal moraines, (c) marks on bed rock, and (d)
the direction of the long axes of many of the lakes. All of
these indicate a motion toward the southwest (about S.
35° W.). The single observed groove on a ledge bears S.
50° W.
Formation of ground moraine and drumlins. When the
ice margin of the Wisconsin glacier stood at the outermost
moraine in Lincoln County, doubtless some drift, possibly
including some of the drumlins, was deposited. It is possi¬
ble, however, that these particular drumlins were not
formed until the ice edge had melted back some distance,
but they were undoubtedly in their present form before the
border reached the area surveyed, as drumlins are rarely
found within ten miles of the farthest extent of an ice sheet.
A considerable portion of the unassorted drift or ground'
moraine was undoubtedly formed during the last melting of
the ice.
First halt of ice margin. The first record of a halt of the
ice margin within the area surveyed consists in some scat-
Thwaites — Glacial Geology of Wisconsin.
119
tered moranic knolls in T. 40, R. 5 E. All these are small
and of such character that some might equally well be
simply large eskers or possibly in part imperfect drumlins.
If any definite moraine was formed, it is now almost wholly
buried in outwash.
Muskellunge Moraine. The halt of the ice margin which
allowed the deposition of the Muskellunge Moraine followed
upon such rapid melting of the glacier that retirement from
the area to the southwest was not complete. In hollows,
valleys, and depressions between drumlins masses of stag¬
nant ice from a few feet to two or three miles in width sur¬
vived just as isolated bodies of troops are left behind during
the retreat of a defeated army. Protected only by a mantle
of melted-out drift accumulated from their own burden,
these would soon have succumbed to the sun's rays; but
while the ice margin was at the Muskellunge Moraine, vast
quantities of water flowed from the glacier and buried these
isolated ice blocks in sand and gravel with scattered bowld¬
ers carried by ice bergs. This extra cover prolonged the
life of many of the glacial remnants. The material of the
moraine itself was also worked over by water. As originally
deposited the moraine rose above the outwash plain to the
south in only a few places, notably at the hill where the
Muskellunge Fire Tower now stands. Near the moraine
this plain had an elevation of about 1700 feet above sea
level. It sloped gently toward the south and southwest.
The outwash at Lac du Flambeau Station is unusually
bowldery and coarse. Such stony layers are doubtless
present elsewhere and await discovery by deep digging.
Boulder Moraine . What caused the alternating rapid re¬
treats and periods of relative stability of the ice margin is
not known. Certainly in Vilas County it was not due to
melting back to positions where the glacial front was pro¬
tected by hills although it is true that as a moraine accumu¬
lated, it tended to prolong the halt by protecting the ice
from the sun. More likely changes either in local climate
or in nourishment of the ice to the north were the cause.
The Boulder Moraine marks a retreat of about 8 miles and
the deposition of an outwash plain which buried blocks of ice
up to about four miles long and at least 150 feet thick. Such
120 Wisconsin Academy of Sciences , Arts, and Letters.
large blocks may have projected above the sand plain. The
deposits buried much of the formerly deposited terminal
moraine as well as all the intervening ground moraine and
many drumlins. The elongations of many of the ice blocks
in a northeast-southwest direction is doubtless explained
by their location in low tracts between drumlin uplands, for
drumlins occur in groups arranged parallel to the direction
of ice movement. Moreover, it is possible that preglacial or
interglacial valleys trended toward the southwest. The
streams from the new ice front found lower courses than
had prevailed when the plain south of the Muskellunge Mo¬
raine was completed. In part this was due to lower out¬
lets freed by the recession of the main body of the ice and
in part to melting of buried ice blocks in the outwash to
the south which opened new drainage lines. The result was
to cut away a large portion of the high level plain south of
the Muskellunge Moraine before all of the buried ice masses
had melted, for there are many kettles in the later drainage
lines. Large portions of the older moraine were also
eroded away or buried under outwash. When the ice blocks
melted, the bowlders they contained were deposited in the
resulting kettles or pits.
Winegar Moraine. The formation of the Boulder Mo¬
raine was followed by a retreat of about five miles after
which a prolonged halt of the border caused the deposition
of the big Winegar Moraine. That this moraine is one of
recession and not of readvance is demonstrated by the
gradation of the moraine into the pitted outwash south of it.
Had the ice front retired long enough to permit melting of
the buried ice blocks, unpitted outwash would have been de¬
posited along large portions of the border of the moraine.
No such deposits are present. The Winegar Moraine con¬
tains less water-sorted material than do the other moraines
of the area, but there are many kames and several large
patches of pitted outwash, probably not all mapped, within
the moraine proper. Some of these outwash plains had
drainage outlets over blocks of ice which have since melted
to form lake basins. Kettles formed when the isolated ice
masses which were buried in the till melted ; into those ket¬
tles which formed before the surrounding moraine was clad
S3 * Till and Coarse Gravel ESI “ Outwash E3 “ Granite E23 “ Schist and Gneiss
Thwaites — Glacial Geology of Wisconsin.
121
122 Wisconsin Academy of Sciences , Arts, and Letters.
with vegetation, red clay, fine sand, and some ice-rafted
bowlders were washed. In October 1927 a good example of
this could be seen just west of the station at Winegar. The
red color of the clay is probably due to its derivation from
red Keweenawan and Huronian rocks to the north rather
than to the plowing up of lake clays as in northeastern Wis¬
consin. The outwash streams from the Winegar Moraine
also formed a plain lower than the higher portions of the
Boulder outwash plain. The older moraines and outwash
plains were extensively eroded and buried. This was done
before the ice blocks had all melted. Little was left of the
Boulder Moraine. The lower plain may be seen cutting
across the higher plains along Highway 51 west of Trout
Lake and southwest of Sayner. The main level of outwash
from the Winegar Moraine is that seen at Boulder Junction
(fig. 1). It was itself extensively terraced along Manito-
wish River by flow coming through the outer part of the
moraine when the ice front had retired slightly farther
north (fig. 2).
Postglacial. The glacial history of the area closes with
the completion of the Winegar Moraine, for after that no
more glacial drainage seems to have reached this region.
This was due to the abrupt northward descent of the land
north of the moraine in Michigan which diverted the wa¬
ters to lower outlets than those across Vilas County. Since
the close of glaciation the surface of the land in Vilas
County has been altered by (a) erosion along some of the
principal streams forming valleys with a maximum depth
of 20 feet, (b) organic deposits in lakes and pools forming
marshes, and (c) weathering which has kaolinized the
feldspar of the sands to depths of one to three feet and has
oxidized the iron-bearing minerals to much greater depths.
In many places hydrous iron oxide has been redeposited in
veinlets to a depth of more than five feet from the surface.
These form irregular hard bands on the weathered surface
of an excavation. In the red till region the color has been
changed by hydration and solution to yellowish brown to a
depth of two to four feet from the surface.
Tkwaites — Glacial Geology of Wisconsin.
123
Economic Geology
Sand and gravel. Although the largest part of the area
surveyed is underlain by outwash and other forms of as¬
sorted drift, good gravel is not common. Most of the out¬
wash is fine sand. The best stony gravels are found in (a)
outwash close to the moraines, (b) kames within the ter¬
minal moraines, and (c) eskers. The following list of pits
is probably not complete, for small excavations near sum¬
mer resorts may have escaped observation.
Location Origin Remarks
T. 40, R. 4 E. Sec. 24. Kame (esker?) — Large pit in poorly sorted
gravels
— Several small pits in and
near Winchester
— On road to Little Long
Lake
— Several pits in rather fine
sandy gravel along
C. T. H. “W”
—-Roadside pit on new road
to Powell
— Largest pit in area,
mile long, 40 feet deep;
used for filling by C.
and N.W. R.R.
Playgrounds Association
T. 43, R. 5 E. Secs. 8.
and 9. Kames
Sec. 25. Kame
T. 42, R. 5 E. Sec. 4. Outwash
T. 41, R. 5 E. Sec. 30. Outwash
Sec. 34. Outwash
124 Wisconsin Academy of Sciences , Arts, and Letters.
Location
Origin
Remarks
T. 41, R. 7 E. Sec. 26. Kame
— On Sayner-Trout Lake
road
Sec. 36. Kame
— On Sayner-Trout Lake
road
T. 41, R. 8 E. Sec. 27. Kame buried
in outwash — On C. T. H. “S”
In addition to the above list there are many pits in the
weathered surface of the outwash or “top soil” and a num¬
ber in such sandy material that its use seems unwise. Many
showings of what seemed to be good stony gravel were ob¬
served both along roads and in the brush, but as these were
not confirmed by digging, they have been omitted. All the
gravel is composed of hard crystalline pebbles and a few
pebbles of sandstone. The gravels are inferior for both
surfacing and concrete pavement to those found in lime¬
stone regions.
Water. Underground water supplies have been devel¬
oped only to a very limited extent in the area surveyed. The
railroad tank at Boulder Junction is supplied from driven
wells in the outwash. Many summer resorts have shallow
dug or driven wells, but others depend upon lake water. It
is unlikely that large supplies could be developed at all points
as coarse gravel is so scarce. In many places till may be
found below the outwash and above the water table ; in such
situations little water could be obtained from wells. It is
likely that considerable iron will be found in the ground
water at most localities, for the forest mould and peat
swamps undoubtedly dissolve a considerable amount of that
substance.
Soils. The soils of the area here discussed have been
described by Whitson, Dunnewald, and others5 in connec¬
tion with the controversy over reforestation. The map
made for this report bears evidence of much careful and
painstaking work but of very limited knowledge of geology.
The following table represents the findings of the writer as
to the true origin of the several soil series described in the
report.
8 Whitson, A. R., and Dunnewald, T. J., and others, Soil survey of Vilas
and portions of adjoining- counties: Wisconsin Geol. and Nat. Hist. Survey
Bull. 43, 1916.
Thwaites — Glacial Geology of Wisconsin.
125
Soil Series Origin
Plainfield — Outwash, little pitted and only slightly weathered.
Vilas —Outwash with a few kames and some terminal moraine
where the till is covered with a few feet of sand; shows
more alteration than the Plainfield soils.
Antigo — Outwash, little pitted and considerably weathered.
Kennan — Sandy loams, mainly terminal moraine, especially the
rolling phase; level phase includes much deeply weathered
outwash; silt loams not yet investigated in this area.
Conclusion
General. Although the present survey covered only a
portion of the northern lake region, it is believed that it
showed the general type of geology which exists throughout
the area. Work over a much larger area will be necessary
to connect the moraines into the general history of the re¬
cession of the Wisconsin ice sheet, but the problems of the
origin of the lesser topographic features, such as the lakes,
have been solved.
9
CHEMICAL ANALYSES OF LAKE DEPOSITS
Charles Spurgeon Black
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. XXXI.
Introduction
For many years the Wisconsin Geological and Natural
History Survey has been studying the physics, chemistry
and biology of the inland lakes of the State. Up to the
present time the chemical work has been confined chiefly to
quantitative determinations of the gases, organic matter
and the various salts that are dissolved in the waters of
these lakes ; during the progress of these studies, the quan¬
tity and chemical composition of a number of aquatic or¬
ganisms have also been investigated. The analyses pre¬
sented here extend these chemical studies to the bottom de¬
posits of a number of Wisconsin lakes and to three Alas¬
kan lakes.
The collecting of bottom deposits for chemical and micro¬
scopical investigations was begun in 1918 and it has been
continued more or less regularly up to the present time.
During this interval a rather large collection of such mate¬
rial has accumulated; the chief aim, however, has been to
obtain representative samples from the various types of
lakes rather than to secure samples from as large a number
of lakes as possible. In fact samples have been taken in
only a relatively small percentage of the lakes visited dur¬
ing this period. The present analyses include only a small
part of the material now in hand.
Most of the samples were obtained in the deeper portions
of the various lakes; they were secured with an Ekman
dredge so that they represent only the upper 12 to 15 cm.
of the deposit. No attempt has been made to study the
differences between the top and bottom portions of these
samples. The bottom samples from the Alaskan lakes were
128 Wisconsin Academy of Sciences , Arts, and Letters.
collected by the late Dr. George I. Kemmerer during July
and August, 1927.
This investigation was carried out under the direction of
Dr. Kemmerer and the author wishes to express his indebt¬
edness to him for assistance and helpful suggestions.
Methods
The samples were spread out and air dried as soon as
possible after they were secured; this material was then
put into bottles and jars and kept until the analyses were
made. The air dried samples were ground in a disc mill
and the moisture was then determined by drying in an elec¬
trically heated vacuum desiccator; the material was spread
out on watch glasses in order to expose as much surface as
possible and it was dried for ten days at 70° C. The dry¬
ing was carried out at this temperature so as to avoid the
decomposition of the carbonaceous material. In general the
procedures given by Hillebrand1 were used for the various
chemical determinations.
Character of Water
The amount of salts held in solution by the waters of the
lakes under consideration differs very widely. The lakes
situated in southeastern Wisconsin possess relatively hard
waters; that is, they contain comparatively large amounts
of calcium and magnesium. On the other hand the lakes
of northeastern Wisconsin and of Alaska have waters that
range from medium hard to very soft, since they contain
only medium amounts to very little calcium and magnesium.
Table 1 shows the amount of residue, fixed carbon dioxide,
silica, calcium and magnesium in the surface waters of the
various lakes.
Determinations of silica, calcium and magnesium have
been made on six of the lakes, but the amount of fixed car¬
bon dioxide, or methyl orange alkalinity, serves to give a
general idea of the degree of hardness of the water. The
three lakes situated in southeastern Wisconsin, namely,
Mendota, Monona, and Okauchee, with 70 to 75 mgm. of
1 Hillebrand, W. F. Bui. 700, TJ. S. Geol. Survey. 1919.
Black — Chemical Analyses of Lake Deposits.
129
fixed carbon dioxide per liter, are classed as hard water
lakes, while Ike Walton, Long and Mary of the northeastern
group with less than 3 mgm. of fixed carbon dioxide per liter
belong to the soft water class; all of the others are inter¬
mediate, ranging up to medium hard.
Analysis of the Deposits
Table 2 shows the results obtained in the chemical analy¬
ses of the various deposits. The hard water lakes repre¬
sent one extreme and the soft water lakes the other extreme.
Si02. In the three hard water lakes (Mendota, Monona
and Okauchee) the silica constitutes from 28% to 36% of
the dry weight of the deposit. In the soft to medium lakes
there is a much wider variation; the range is from 22% in
Ike Walton in northeastern Wisconsin to more than 69% in
O'Malley, one of the Alaskan lakes, if the Forestry Bog is
excluded. This small bog represents a special type of de¬
posit and is not strictly comparable with deposits laid down
in lakes. The three Alaskan lakes have a much higher per¬
centage of silica than any of the Wisconsin lakes; this is
accounted for by the fact that a very large proportion of
the Alaskan deposits consists of diatom shells.
Fe203. There is a rather wide range in the amount of
iron in the various deposits. It ranges from a minimum
of 1.5% in Lake Laura to a maximum of approximately
9.5% in Trout Lake; both of these lakes are situated in
northeastern Wisconsin.
A1203. The percentage of aluminum ranges from a mini¬
mum of 1.4% in Long Lake to a maximum of 9.5% in Silver
Lake ; both of these lakes also belong to the group situated
in northeastern Wisconsin. The deposit from Thumb Lake,
Alaska is a close second to Silver Lake, however, since it
contains approximately 9.4% aluminum.
CaO. The analyses show very marked differences in the
calcium content of the various samples. In the three hard-
water lakes of southeastern Wisconsin calcium comprises
from 20% to almost 25% of the dry weight; in the medium
to soft water lakes of northeastern Wisconsin the percent¬
age varies from a minimum of 0.6% in Long Lake to a
maximum of 2.4% in Lost Canoe and Turtle Lakes. In the
130 Wisconsin Academy of Sciences , Arts , and Letters .
Alaskan lakes the calcium varies from 2.1% in Karluk to
3.6% in O’Malley. Table 1 shows that the water of Lake
Mendota contains two and a half times as much calcium as
that of Trout Lake, but the bottom deposit of the former
has fourteen times as much calcium as that of the latter.
Also the water of Lake Mendota contains approximately
twice as much calcium as the waters of the three Alaskan
lakes, but the bottom deposit of the former yields from five
to nine times as much calcium as the deposits of the latter.
These facts seem to indicate that a relatively small increase
in the calcium content of a lake water will produce a much
more marked increase in the calcium content of the bottom
deposit. The amount of calcium which gets into the bottom
deposit is the result of complex chemical and biological pro¬
cesses, but it is very greatly influenced by the amount in
the water.
MgO. Owing to the much greater solubility of magne¬
sium, there is a comparatively small amount of MgO in the
deposits of the hard water lakes as well as in those of the
soft to medium lakes. In the former group the maximum
is 3%, while in the latter it is 1.7%. In the three south¬
eastern lakes the percentage of CaO in the deposits is from
eight to fourteen times as large as that of the MgO ; in the
northeastern lakes the ratio of CaO to MgO ranges from a
little more than one to almost sixteen and in the Alaskan
lakes from about one and a half to three.
P205. There is about an eightfold range in the phos¬
phorus content of the various deposits, but there is no sig¬
nificant difference between hard and soft water lakes in the
relative amount of phosphorus.
S04. The smallest percentage of sulfate ion was found
in Lake Laura and the largest in Trout Lake. Both of these
lakes are situated in northeastern Wisconsin.
C02. As might be expected the three hard water lakes
of southeastern Wisconsin show a much higher percentage
of carbon dioxide than the medium and soft water lakes;
in the former group the percentage ranges from 9.3% to
16.3% and in the latter from about 0.2% to 2.4%.
Organic Carbon. The amount of organic carbon in the
various samples gives some idea of the relative quantities
Black — Chemical Analyses of Lake Deposits .
131
of organic matter in the different deposits. The samples
obtained from the lakes situated in northeastern Wisconsin
show the largest percentages; the organic carbon varies
from 10.5% in Turtle Lake to 82% in Long Lake, excluding
the Forestry Bog. In the samples from the lakes of south¬
eastern Wisconsin the range is from 4.7% to 7.2% and in
the Alaskan lakes from 4.4% to 8.2%. The Forestry Bog
yielded substantially 39%.
In five samples of the surface deposit from different lo¬
calities in Lake Balaton,2 Hungary, the silicia varied from
1.5% to 54%, the calcium from 12% to 52% and the mag¬
nesium from 0.7% to 4.6%.
Decomposition of Bottom Deposit
An experiment was undertaken to determine how rapidly
the organic matter in the bottom mud decomposes and also
to ascertain the quantity and chemical composition of the
gases given off during the decomposition. Two samples of
mud were secured from a depth of 24 m. in Lake Mendota
on October 21, 1927 ; they were transferred directly from
the dredge to bottles. One bottle had a capacity of 4 liters
and the other 5 liters ; in both bottles the space not occupied
by mud was filled with water brought up from the bottom
in the dredge. Both bottles were placed in the dark during
the experiment, but the 5 liter bottle was kept at room tem¬
perature and the 4 liter bottle in an electric refrigerator at
a temperature of 4°. At the conclusion of the experiment
the mud was dried and weighed ; the 5 liter bottle contained
600 grams of dry material and the 4 liter bottle 690 grams.
Between October 21, 1927 and April 9, 1928 the material
kept at room temperature in the 5 liter bottle (600 grams)
produced 2597 cc. of gas, of which 297 cc. consisted of car¬
bon dioxide and 1470 cc. of methane ; the remainder consis¬
ted of hydrogen and nitrogen. No carbon monoxide or
unsaturated compounds were found.
The sample kept in the refrigerator from October 21,
1927 to April 9, 1928 produced 593 cc. of gas, or only a little
more than a quarter as much as the one at room tempera-
2 Res. wissenschaft. Erforsch. des Balatonsees. Bd. I, T. 1, 17 pp. 1911.
132 Wisconsin Academy of Sciences , Arts , and Letters.
ture. Of this amount 39 cc. consisted of carbon dioxide.
320 cc. of methane and the remainder of hydrogen and ni¬
trogen.
Table 1. The amount of residue or total solids, fixed carbon dioxide ,
silica, calcium, and magnesium found in the surface waters of
the lakes for which analyses of the bottom deposits have been
obtained. The results are given in milligrams per liter of water.
Mendota, Monona, and Okauchee Lakes are situated in south¬
eastern Wisconsin, while the other Wisconsin lakes are situated
in the northeastern lake district. The Alaskan lakes are situated
on Kodiak Island.
Wisconsin Lakes.
Alaskan Lakes.
Table 2. Chemical analyses of bottom deposits. The results are stated in percentages of the dry weight. The depth
of the water at the points where the samples were taken is indicated in meters. Mendota, Monona and Okauchee
are situated in southeastern Wisconsin; Adelaide to Turtle inclusive , in northeastern Wisconsin; Karluk, O Malley
and Thumb on Kodiak Island , Alaska.
Black — Chemical Analyses of Lake Deposits,
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NOTES ON THE CHEMICAL COMPOSITION OF SOME
OF THE LARGER AQUATIC PLANTS OF LAKE
MENDOTA. III. CASTALIA ODORATA
AND NAJAS FLEXILIS
Henry A. Schuette and Hugo Alder
Contribution from the Department of Chemistry, University of
Wisconsin, and the Biological Laboratory of the Wisconsin Geological
and Natural History Survey. XXXII.
In earlier communications from these laboratories data
were presented on the composition of some of the attached
vegetation of Lake Mendota. Pertinent to these reports
were an alga, Cladophara,1 which attaches itself to the sub¬
merged rocks to be found along the shore-line in many
places, and representatives of the larger plants. Among
the latter were those which prefer a sandy soil, as the Pota-
mogetons2 those which flourish in muddy situations, as
Myriophyllum,1 and those which thrive equally well in sandy
soil as in mud, as Vallisneria.2 Herein we record the com¬
position of one of the species, Najas flexilis, which may be
found in the shallow bays and which from the standpoint of
abundance in this lake is exceeded only by Vallisneria spir¬
alis L. and the Potamogetons ; and of another which grows
in sandy or in gravelly soils, Castalia odorata. The former
has been reported as growing in water whose depth ranges
from 0.3 to approximately 5.5 meters; the latter in water
less than a meter in depth.
The species in question were part of the material collected
by Rickett3 in his determination of the quantities of the
larger aquatic vegetation which are found in the region of
plant growth in this lake, an area arbitrarily divided into
three zones. All three zones furnished specimens of Najas
flexilis , but only the first, or shallowest one, contributed the
Castalia odorata L. The latter in the amount of 1,500
grams, representing 146 grams of dried material, was gath¬
ered from five stations in this zone. Of the former there
136 Wisconsin Academy of Sciences , Arts , and Letters.
were obtained 3,455 grams of the green plant which on
drying yielded 384 grams.
Description of Plants
Castalia. The castalias, or more commonly the water-
lilies, though world-wide in distribution are restricted to the
northern hemisphere. Their grace and beauty have at¬
tracted man from the earliest time ; they have given him an
artistic inspiration in architectural design, and in India and
in Egypt have from time immemorial entered into his social
and religious life. There is evidence also that the seeds and
tuberous roots of various species have been used for food
by the natives of Australia, Madagascar, West Africa, and
Central America.4
The long rhizomes of the castalia group creep along in or
upon the muddy bottom of the lake in a tangle of vegetation.
They branch in every direction, dying off behind as they ad¬
vance by apical growth. Many of them have an erect stem.
This never acquires any considerable length, however, for
it stands with its apex about on a level with the mud in
which its lower portion is buried. Many stout fibrous roots
pass downward from the stem and anchor the plant in place
while the leaves and flowers rise upward. The leaves lie
flat upon the water surface, their upper sides being dry and
exposed to the air. Under some conditions the petioles
may be strong enough to hold up the leaves thus making
them totally aerial. All of the species produce at certain
periods of growth small thin leaves which lead a submerged
existence. The flowers of about half of the species open at
the surface of the water and seem to float upon it. Those
of the other half are raised on strong peduncles some dis¬
tance above the water-level. All have numerous petals ar¬
ranged in many rows, the innermost gradually passing into
the stamens. They are white, pink, yellow or blue and are
very showy. Only in one or two species is cleistogamy
known to occur. The flowers in such cases never rise to the
air, or only do so for a few hours, after self-pollination has
been effected. The fertile flowers, after anthesis, are
drawn down into the water by movements of the peduncle
and here the seeds ripen. The ripe seeds from the irregu-
Schuette* & Alder — Composition of Aquatic Plants. 137
larly bursting fruits rise to the surface of the water by
means of a buoyant aril and then float away far and wide.
Najas. The Najas are of no economic importance yet
they have aroused much academic interest because of the
simplicity and exceptional development of their floral parts.
This genus is commonly referred to as the water nymphs,
a name which appears to be inaptly applied to these humble
water weeds.
The members of the genus Najas5 are small, slender,
branching herbs growing under water, with opposite and
linear leaves which are somewhat crowded into whorls. The
former are spinulose-toothed, sessile and dilated at the base.
The flowers, which are dioecious or monoecious, are very
small. They are carried without a foot-stalk in the axils of
the leaves. Pollination is accomplished through the water
in which the plants grow. The fruit is a small seed-like
nutlet. It is enclosed in a loose and separable membrane.
Chemical Analysis
The material was made ready for chemical analysis by the
procedure which has been described previously.2 It was
analyzed by the methods of the Association of Official Agri¬
cultural Chemists8 which have been standardized and re¬
quire no description here. Manganese was determined col-
orimetrically and the alkali metals were separated by the
perchlorate7 rather than by the platinic chloride method.
Sand, which was considered to be extraneous matter, was
determined in each species and correction made for the
amount found. There proved to be 9.60 per cent in the
Najas flexilis sample and 1.73 per cent in Castalia odorata.
The analytical results (tables 1 and 2) , which represent
the averages of duplicate determinations, are expressed on
a sand-free and air-dry basis. The latter condition implies
the net results of desiccation in the air and oven-drying at a
temperature of 60°C. During the six-year interval which
elapsed between collection and chemical analysis, the ma¬
terial in question had been stored under conditions which
prevented the absorption of moisture.
138 Wisconsin Academy of Sciences , Arts, and Letters .
Table 1. Proximate composition of Castalia odorata and
Najas flexilis L.
proached from the standpoint of the relative amounts of
mineral matter which each species annually requires for
growth, and, in a sense, removes from the lake floor only to
return part, if not all, of it in the final stages of its life
cycle as soil constituent.
Rickett3 has estimated that in the zone of plant growth
in Lake Mendota — an area of approximately 1,040 hectares
— there are annually produced 40 metric tons of Najas flex¬
ilis. On reducing this weight to a dry basis and translat¬
ing it into terms of mineral matter, referable to the ash
content found, it follows that this species annually requires
some 7.6 metric tons of inorganic constituents for growth.
This is less than the drain made upon the soil and waters
of this lake by Vallisneria and Potamogeton2. A reinter¬
pretation of data1 previously reported brings out the fact
Schuette & Alder — Composition of Aquatic Plants. 139
that the annual requirements of Najas in Lake Mendota
exceed those of Cladophora; although the latter contains
more mineral matter, yet it is here less abundant. On the
other hand, Najas removes only about 45 per cent as much
inorganic material as does Myriophyllum 1 whose ash con¬
tent (20.72 per cent) is approximately the same. Quanti¬
tative data on the distribution of Castalia are incomplete in
the sources3 which furnished the above information, hence
a similar comparison can not be made for this form.
As for the species pertinent to this communication, it
may be said that the order of magnitude of the content of
alkali metals, chloride, sulfur, and phosphorus is substan¬
tially the same; that Najas flexilis annually requires more
iron, aluminum, calcium, and magnesium salts than does
Castalia odorata, but that the latter stores up more maga-
nese than does the former.
Literature Cited
1 Schuette, H. A. and Hoffman, Alice E. Trans. Wis. Acad. Sci. 20:529-531.
1921.
2 Schuette, H. A. and Alder, Hugo. Ibid., 23:249-254. 1927.
3 Rickett, H. W. Ibid., 20: 501-527. 1921.
4 Conrad, H. S. “The Waterlilies”. Carnegie Institute Publication 4. 1905.
6 Gray, “New Manual of Botany”, rearranged and revised by B. L. Rob¬
inson and M. L. Fernald. American Book Co., New York, 1908. 7th ed.,
p. 79.
6 Association of Official Agricultural Chemists. “Methods of Analysis.”
1925. Washington, D. C., p. 39 et seq.
7 Treadwell-Hall, “Analytical Chemistry”. Wiley & Sons, New York. 1911.
Vol. II. p. 50.
A NOTE ON THE CHEMICAL COMPOSITION OF
CHARA FROM GREEN LAKE, WISCONSIN
Henry A. Schuette and Hugo Alder
Contribution from the Department of Chemistry, University of Wis¬
consin, and the Biological Laboratory of the Wisconsin Geological
and Natural History Survey. XXXIII.
Little published information on the chemical composition
of Chara seems to be available except a statement from
botanical sources1 that a species of stone-wort, Chara foe -
tida, was found to contain potash, soda, lime, and silica in
the amounts of 0.2, 0.1, 54.8, and 0.3 per cent, respectively.
In view of the incomplete state of these data we have been
prompted to fill the existing gaps by reporting the results
of a chemical analysis which we made of a typical member
of this order.
An opportunity of securing representative samples of
Chara from Green Lake, Wisconsin, presented itself through
the activities of Rickett2 who made a survey of the annual
crop of higher plants of this body of water in 1921. He
observed that it is the dominant plant in the vegetation of
this lake and reported that it may be found growing almost
everywhere, sometimes mixed with other plants, and often
standing alone in such dense masses that no other form has
been able to obtain a foothold. This condition stands in
marked contrast to that which obtains in Lake Mendota, it
is stated, for here Chara plays but a minor role in the at¬
tached flora. Rickett advances the opinion that this situ¬
ation may be traced not only to differences in the charac¬
ter of the respective lake floors, but also to differences in
the transparency and temperatures of the waters in ques¬
tion. The waters of Lake Mendota are warmer and less
transparent than those of Green Lake. The floor of the
latter is covered with a fine marl in contrast to a mud
which is characteristic of Mendota’s bottom. Differences
in the chemical composition of the waters of these lakes are
10
142 Wisconsin Academy of Sciences, Arts, and Letters.
not such as to be conspicuous, hence this condition is prob¬
ably outside the scope of the discussion.
Description of the Plant
Members of the genus Chara are popularly known as the
stoneworts because they are incrusted with a calcareous
deposit which, it is believed, serves them as a defensive
armor. They are brittle and rough to the touch.
The most common species of this group is Chara fragilis 3.
It is wide spread in its distribution. The plant is about
twelve inches high and consists of an axis with whorled,
long, spiny, leaf-like structures at the nodes.
It reproduces vegetatively by outgrowths or sexually by
the formation of motile sperm cells. The sexual structures
are borne on the whorled branches at the nodes. Here are
found oogonia, each containing a large egg cell which is
visible to the eye because of its orange-red color. Beneath
the oogonia at the same nodes are found antheridia which
produce a large number of motile sperm cells. These enter
the oogonia through an opening at the top and after fertili¬
zation a resting spore (oospore) is produced.
It is probably not incorrect to state that these plants
have never been used for any practical purposes, yet very
good reasons suggest themselves for their use in correcting
the acidity of sour soils.
Others, however, of the so-called worts have had at one
time an economic importance. It may be pertinent to re¬
call, in this connection, that it was to the members of the
genus Salsola or saltworts that soap makers of early days
in Spain turned as a source of alkalies long before the de¬
velopment in France of the well-known Le Blanc process
for producing soda from common salt. The saltwort was
here burned for its ash because the latter is relatively high
in soda. This ash, which was known as “barilla”, was then
lixiviated with burnt lime whereby a solution of caustic
alkali was obtained.
Chemical Analysis
The material which was made available for analysis rep¬
resented the harvest from 38 collecting stations which were
Schuette & Adler — Chemical Composition of Chara. 143
variously located in the zones of plant growth — an area
comprising here some 8,570 square kilometers — over which
the depth of water ranged from zero to eight meters. It
had been air dried and then further dehydrated in an oven
maintained at a temperature of 60°C. During the five-
year interval which had elapsed between collection and
chemical analysis, it had been stored in closed containers.
Extraneous matter, such as shells, stones, sand, etc., was
removed by hand after which the dried plant was commin¬
uted in a drug mill to pass through a sixty-mesh sieve.
Iron introduced during the process of milling was removed
with an electro-magnet.
The analytical procedures of the Association of Official
Agricultural Chemists4 for the determination of organic
and inorganic plant constituents and that of carbon dioxide
in baking powder were followed with one exception, and
that was the method of separating the alkali metals. The
perchlorate method of Schlossing-Wense5 was substituted
for the official platinic chloride procedure. They are stand¬
ard methods which require no description here.
Sand was found to be present to the extent of 0.89 per
cent. In as much as this may very well be considered an
impurity, analytical data subsequently obtained were cor¬
rected to a sand-free basis. They are reported on “as re¬
ceived” or air dry basis under conditions which have been
outlined in preceding paragraphs. They are recorded in
the following tables.
Table 1. Proximate composition of Chara.
Percentage
Sand-free
Constituent air dry basis
Ash _ 41.22
Crude protein (N x 6.25) _ _ _ 4.50
Ether extract _ _ _ _ _ 0.76
Crude fiber _ 9.32
Pentosans _ 4.70
Nitrogen-free extract _ _ _ _ _ 39.50
144 Wisconsin Academy of Sciences, Arts, and Letters.
Table 2. Inorganic constituents of Chara ,
Percentage
Constituent
Silica (SiO,) _ - _
Ferric oxide (Fe203) _
Aluminum oxide (A1203) _
Manganomanganic oxide (Mn304)
Calcium oxide (CaO) _
Magnesium oxide (MgO) _
Sodium oxide (Na20) _ _ _
Potassium oxide (K20) _
Chloride (Cl) _ ___
Carbonate (C03) _
Total sulfur (S) _
Total phosphorus (P) _
Sand-free
air dry basis
_ _ 0.83
_ 0.06
_ 0.81
_ 0.08
_ 37.82
_ 1.19
_ 0.35
_ 0.58
_ 0.29
_ 39.00
_ 0.27
_ 0.06
It is generally accepted that the nature of the mineral
salts in a water, in a plant or its products is to a certain ex¬
tent a matter of conjecture since the salts found by analy¬
sis of the ash of the material in question are not exactly
the same as those present in the uncalcined sample. These
limitations make impractical the assignment to hypotheti¬
cal combination of the acidic and basic oxides which are re¬
corded in the foregoing tables. These data, however, ad¬
mit of certain gross generalizations of which the following
appear worthy of comment.
That the name stonewort which popular usage associates
with this plant is well taken becomes apparent upon analy¬
sis of the data in tables 1 and 2 and by comparison with
that obtained in the study of other forms of aquatic vege¬
tation 6’ 7 from Lake Mendota. Chara contains less ni¬
trogenous matter, less cellulosic material and a lower carbo¬
hydrate content, referable to crude fiber and pentosans, re¬
spectively, than do Vallisneria, Potamogeton, Myriophyllum
and Cladophora.
The fact that Chara contains, with some exceptions, a
lower content of silica, iron, aluminum, maganese, magne¬
sium, sodium, potassium, sulfur, phosphorus, and chloride
than do the other vegetative forms in question is not so
striking as that calcium oxide constitutes approximately 92
per cent of its ash. The value obtained, 37.82 per cent, is
not in agreement with that reported from other sources1,
yet this apparent discrepancy has no particular significance
Schuette & Adler — Chemical Composition of Char a. 145
of itself since the history of the material in question and
the conditions under which it was grown, gathered, and
analyzed are obscure.
Parallel with a high content of calcium oxide lies the
quantity of carbon dioxide which the dried plant yields on
treatment with acids. If one assumes that all of the car¬
bonate in this plant exists here in chemical union with cal¬
cium and translates the quantity of carbon dioxide found
into terms of calcium carbonate, there is obtained a value
for the latter of 65 per cent. That this assumption,
whereby is established the order of magnitude if not the
exact quantity of the calcium carbonate content of Cham,
is not seriously in error is established by the fact that the
actual amount of calcium oxide found by analysis is ap¬
proximately equal to that required by theory. On deduct¬
ing the calcium oxide content of the ash from that required
by the calcium carbonate of the incrusted deposit (36.40
per cent) there remains 1.42 per cent to be assigned to
other acidic oxides.
Birge and Juday8 state that Rickett’s survey2 indi¬
cates that about one half of the annual crop of higher
plants of Green Lake — some 1,528 metric tons — is contrib¬
uted by Cham. On correlating this observation with the
values reported herein it would seem that the annual
growth requirements of Cham for its major constituent
are 397 metric tons of calcium and 427 metric tons of car¬
bon in terms of carbon dioxide. There are returned every
year to this lake 993 metric tons of calcium carbonate.
Literature Cited
1 Kerner and Oliver, Natural History of Plants. Holt and Company, New
York. 1895. Vol. I, p. 69.
2 Rickett, H. W. A Quantitative study of the larger aquatic plants of
Green Lake, Wisconsin. Trans. Wis. Acad. Sci., Arts, and Let. 21: 381. 1924.
8 Kerner and Oliver, loc. cit., Vol. IV, p. 659.
4 Association of Official Agricultural Chemists, Methods of Analysis.
Washington, D .C. 1925, p. 39 et seq., 301.
5 Treadwell-Hall. Analytical Chemistry. New York, Wiley & Sons. Vol.
II, 1911, p. 50.
6 Schuette, H. A. and Hoffman, Alice' E. Notes on the chemical composi¬
tion of the larger aquatic plants of Lake Mendota. I. Cladophera and Myri-
ophyllum. Trans. Wis. Acad. Sci., Arts and Let. 20 : 529. 1921.
7 Schuette, H. A. and Alder, Hugo. Notes on the’ chemical composition
of some of the larger aquatic plants of Lake Mendta. II. Vallisneria and
Potamogeton. Trans. Wis. Acad. Sci., Arts, and Let. 23: 249. 1928.
8 Birge, E. A. and Juday, C. Organic content of lake water. Bull. Bur.
Fisheries, U. S. Dept. Commerce. 42: 191. 1926.
LIFE AND LETTERS OF EDWARD LEE GREENE
Angie Kumlien Main
Edward Lee Greene was born at Hopkinton, Rhode Island,
August 20, 1843. In his early boyhood his family moved
to Albion, Wisconsin and settled near the Seventh Day Bap¬
tist Church, of which they were devout members. He at¬
tended Albion Academy, but at the end of his junior
year in 1862 he, as well as his father and two brothers,
joined the Union army ; at the close of the Civil war he re¬
turned to Albion Academy and graduated with the class of
1866, receiving the Ph. B. degree.
Not far from Albion, near Lake Koshkonong, lived Thure
Kumlien, a naturalist of the old school, who became Ed¬
ward’s first teacher in the science of botany. A warm and
lifelong friendship developed between teacher and pupil,
which lasted until the death of the former in 1888.
From 1867 to 1870 Edward taught school at Monticello,
Illinois, where he kept up his botanical studies. In the lat¬
ter year he went to Colorado, where he studied for the min¬
istry and was ordained in 1871. He was instructor in bot¬
any at Jarvis Hall, Golden City, Colorado in 1871-72, after
which he was rector of churches at Greely, Pueblo, and
Georgetown, Colorado, and at Berkeley, California until
1882. He became connected with the department of bot¬
any of the University of California in the latter year and
remained there until 1895, serving as Professor of botany
during the last five years. In 1893 he was President of the
International Congress of botanists which was held at the
Chicago World’s Fair.
In 1895 he became professor of botany in the Catholic
University of America, Washington, D. C., where he re¬
mained until 1904; he then became associate in botany at
the Smithsonian Institution. During the twenty-five years
that he was in the west he devoted himself to an extensive
study of the western flora. His descriptions of his collect¬
ing trips through the Rockies, over the plains of Wyoming,
148 Wisconsin Academy of Sciences , Arts , and Letters.
through California, and away to the borders of New Mexico
teem with interest. He discovered and described many
new species of plants. He was the author of several books
and of numerous papers which were published in scientific
journals.
When Edward Lee Greene left home to enter the Civil
war, a correspondence was begun with Mr. Kumlien, which
continued for a period of twenty-six years. The following
letters have been selected from this correspondence and
they show his fine character, his love for the beautiful, his
appreciation of the truth, his love for his revered teacher,
and his gradual advancement as a high authority in botany.
In these letters, Mr. Greene used only the scientific names
of the plants mentioned, but the common names have been
added in parentheses in many instances.
Dover, Tennessee, Sept. 10th, 1862.
Dear friend Kumlien:
I am seated in my tent this afternoon to write you a few lines to
inform you that I am in the land of the living away down in Tennes¬
see. I have been on the march for the most of the time since I ar¬
rived in Dixie and have today returned from an expedition to Clarks¬
ville, a distance of about thirty-five miles. We went for the purpose
of expelling the rebel forces from that place and met them four miles
this side. Our artillery attacked them and routed them after firing
about one hundred shells and a few balls. We were so far from them
that none of our men were seriously injured. Neither our infantry
nor cavalry were allowed to fire at them at all but I think our mus¬
ket balls would have reached very easily if we had fired. Our forces
amounted to about twelve hundred and the rebels consisted of four
or five hundred soldiers and a great number of citizens armed mostly
with shot guns loaded with buck shot. We do not know exactly how
many of them were killed, but the negroes who were present tell us
that they estimated their loss at upwards of one hundred killed and
wounded. Our men in passing through the orchard (they were con¬
cealed in a peach orchard when we attacked them) after they sup¬
posed they had removed all their dead and wounded found two who
were most terribly mangled, one had his head severed from his body
and another was torn in two. They lay side by side and probably
were both killed by the same merciless bomb shell.
I have seen a great many new things in the vegetable world since I
left home but it has not been much more than an aggravation to me to
see when I could not have a chance to collect anything. But as we
shall probably remain here a few days I will try to send you a few
Filices (ferns) for I have found five or six new species here on the
rocky banks of the Cumberland river. Some of them are small and
Main — Life and Letters of Edward Lee Greene. 149
beautiful and one very interesting: one which is called the walking
fern. The frond is entire and the end bends over and after taking
root sends up another. I think the common name is very appropri¬
ate. I did not expect to find many flowers at this season of the year,
but I have seen quite a large number while on the march. There are
some pretty Labiates (mint family) and composites yet in flower and
many species of Desmodium (tick trefoil). These southern forests
must be delightful in early summer.
There are a great many handsome flowering trees here. The mag¬
nificent Liriodendron tulipifera (tulip tree) is quite common and also
Catalpa bignonoides (catalpa) and an Aesculus (horse chestnut) all
large trees.
Among the smaller trees the Cornus florida (flowering dogwood)
and Cercis canadensis (redbud) are very gay when in bloom, es¬
pecially the latter. The Castanea vesca (chestnut) is also common
and the beech and several species of Quercus (oak) are new to me.
One has leaves perfectly entire and two or three have leaves simply
dentate. Plantanus occidentalis (sycamore) and Laurus sassafras
(sassafras) and the gum tree are also large trees. Many of the forest
trees are entirely new to me. The Philadelphus (mock orange or
syringa) which we cultivate grows wild in abundance along the Cum¬
berland.
I have succeeded in collecting a few seeds and will try to send them
to you before long. I have also pressed two or three plants in a book.
I will send you one or two in this letter. They are not beautiful nor
even perfect specimens, but one of them is a climbing plant which I
took at Cairo, Illinois. I have not seen any more like it since, but I
examined the flower and the fruit which was not then mature and I
think it belongs near the genus Accerater (green milkweed). An¬
other is an Euphorbia (spurge). The piece which I am sending re¬
sembles E. maculata (milk purslane) but the plant is much larger
and nearly erect and lacks the spot on the leaf. The other is Lespe-
deza (bush clover) I suppose.
I have not seen many birds except turkey buzzards. I am not as
much of an ornithologist as I hope to be if I live. I will not attempt
to write anything about the birds for I know you will be tired when
you have read this letter, for I am at best a poor writer and am now
writing under very unfavorable circumstances, so I will draw to a
close. Give my best regards to Mrs. Kumlien and Sophia and re¬
member me as your friend,
Edward L. Greene.
Ft. Henry, Tennessee, (Written in the fall), 1862.
Dear friend Kumlien:
It is a lonesome, rainy day and I thought it would be very pleasant
to have a social chat with you for a few hours. But as that cannot
be I will try and write you a few lines to pass away the time.
We are at present encamped on the bank of the Tennessee river at
what is called Ft. Henry, but it is only an opening in the woods.
150 Wisconsin Academy of Sciences , Arts, and Letters.
There is no village here at all nor so much as a farm house within
two or three miles. We have been here about two weeks and I have
had a chance to search the woods a good deal.
I presume that the season of flowers is nearly passed in Wisconsin
by this time, but not so here. It is very warm yet in this section and
quite a large number of flowers are yet to be found and some very
beautiful ones.
I found a species of Chelone (turtlehead) while walking in the
woods. I consider it the most beautiful of anything I have seen. It
differs from the Wisconsin species in that it is not so tall, but more
branching and bears a profusion of rich purple flowers. It is a
pretty large plant, but I shall endeavor to press some small branches
and send them home. I have a book which I keep in my knapsack in
which I can press small plants quite well. I sent a few home about
a week ago and wrote mother to divide them with you. I shall send
some more in a few days. I suppose you have had a good time with
the Asters, Solidagos, Gentians and Gerardia this fall. There
are few of them here. I have not yet seen a Gentiana (gentian).
The Gerardias are very handsome here; G. purpurea and G. seta-
cea grow large here and flower very profusely. I saw a new species
while on the march from Fort Donelson and think it must have been
the G. pedicularia (fern leaved foxglove) if that is the correct name.
It was not so tall as G. quercifolia, more branching and the leaves,
which resemble those of the Pedicularia , clothe the branches very
thickly. It was pretty, but I did not dare to leave the ranks to notice
it closely.
We have taken a tramp of sixty miles south since we came here.
We went up the river on the steamboat to Cherokee landing which is
about forty five miles and then we marched the rest of the way. We
camped at noon near a creek in the woods and there spent the re¬
mainder of the day. And being in a new place I of course took a
tramp as soon as I was rested. I there found Mitchella (partridge
berry) in abundance growing on the bank of the creek. Here I also
found the beautiful Rhododendron nudiflorum (purple azalea) which
was an old Rhode Island favorite of mine. How I wished it had been
in flower, but I could not find even as much as a seed for they were
all gone. I also found a Vitis (grape) with small leaves and very
large fruit which was ripe and of an excellent flavor.
I will now tell you something about a coniferous tree which I have
found in the woods here at Ft. Henry. Whether it is an evergreen
or not I can not yet say, but I think it is not. It bears the fruit on
the upper part of the tree and the branches which bear the fruit have
a different foliage from the rest of the tree. This is one curiosity of
the tree, and another is that under the tree conical stumps arise from
the roots, some to the height of three feet, but of all sizes. They ap¬
pear like a group of small mountains. If I were an artist I would
draw a picture of the tree for it is very strange. I will send you
some twigs of this tree soon. Perhaps you can tell me what it is.
I want you to please write me a letter when you receive this and
Main — Life and Letters of Edward Lee Greene.
151
tell me how you prosper. If you will call on mother she will give you
some of the specimens which I have sent. I am going to send some
more in a day or two. I do not find much now for it is too late in the
season. But if we are so unfortunate as to be kept here until spring,
I expect to have some pleasant times, but I hope that I will be in Al¬
bion next spring in time to gather Arethusa and Pogonia with you
from that blessed little tamarack marsh.
We hear good news occasionally of victories on our side and may
the God of wars grant that the final victory may soon be won and
peace once more reign. For this war is ruining the best young men
of our land. I find that a great change has taken place in the young
men with whom I was acquainted. They are losing all taste for in¬
tellectual enjoyments and spend their hours in gambling and indulg¬
ing their vile appetites. Well I see I have written you a long and
tedious letter and I doubt your ability to read such a mess of scrib¬
bling for I have to write under very unfavorable circumstances. I
will enclose you a picture which you may remember when you see it
as your sincere friend and well wisher.
Edward L. Greene.
Co. H, 13th Reg.
Ft. Henry, Tennessee, Nov. 16th, (I think the year 1862)
My dear Friend:
It is Sunday morning and I have seated myself to try to pen a few
lines to you, although my stock of news is small and I fear it will not
prove very interesting. I suppose you have had some winter weather
in Wisconsin by this time. It begins to look like autumn here now,
although the weather is very warm and pleasant during the day, but
the nights are frosty. The leaves are falling from the trees and leav¬
ing them bare except now and then one whose branches are covered
with mistletoe which is yet green. Is it an evergreen or not? It is
indeed a very curious plant growing on the living branches of other
trees. Mother has received two small twigs which I sent to her by
mail and I presume that you have one of them by this time. When
you write again please give me its Latin name.
There is almost any quantity of nuts and acorns now in the woods
and a great many different species of both. I sent leaves of two spe¬
cies in my last package of flowers and was intending to send fruit of
both with the seeds which I sent you by father, but I found him just
ready to start when we returned from our last expedition and had no
time to get them before he started. I think I will take a tramp in
the woods before long and collect some nuts of all the new species and
put them in my knapsack for it would be quite a novelty in Wisconsin
to see some of these monstrous acorns and hickory nuts which grow
here.
Well, I guess I must lay aside my writing for the present and pre¬
pare to attend religious services which commence at ten o’clock.
We have been on the march for the most of the time since I last
wrote you and consequently I haven’t had much chance to collect
152 Wisconsin Academy of Sciences, Arts, and Letters.
seeds, and the bottle of insects which I attempted to fill would not
amount to much for it is now too late.
I sent all the seeds I had by father, but I had to do them up in such
a hurry that I could not mark them as I had intended to, for I knew
the names of some of them. One paper contains seeds of Plantago
lanceolata (rib grass), a common biennial in New England. Sow
some of the seeds in your garden for I think they will become natural¬
ized. It is very different in appearange from P. major (common
plantain). In another is Cassia marilandica (wild senna) and
another is an annual species of Cassia (senna). I think I sent speci¬
mens of it home. Another is a perennial Poly gala (milkwort). One
contains seeds of a shrub which I suppose to be a Hypericum (St.
John’s-wort) ; it resembles that genus in every particular except that
it is woody. I enclose some of the capsules in the paper so that you
will know it I think.
I will collect some more seeds if I have an opportunity, but it is
rumored that we are to start out on another expedition soon; but I
hope it will not prove true for I am quite tired of running about after
the little squads of guerrillas for they are all on horses and it is al¬
most impossible for us to catch them on foot and they always escape
us by flight. However, on our last trip we managed to overtake them
as they were secreted in the woods getting their supper. They were
about five hundred in number and our force happened to be superior
to theirs. They mounted their horses as soon as they were aware of
our approach and fired on us as they fled, but we returned the fire,
killing sixteen of them and wounding thirty or forty. None of our
regiment were seriously injured but several of the Iowa cavalry
were killed and wounded. We took I think about twenty prisoners
which we left at Fort Donelson as we returned.
During the short time which I have been in the service I have had
a chance to see a good deal of Kentucky and Tennessee and have seen
many places where I have thought it would be very pleasant to spend
a summer. The most romantic road which I have yet traveled is
situated between Ft. Donelson, Tennessee, and Hopkinsville, Ken¬
tucky. On the other side of the Cumberland as we leave Ft. Donel¬
son for about five miles the road runs through a deep ravine. On
one side of the road runs a stream of clear water which has its source
in a large spring at the upper end of the ravine. It increases in size
as it flows southward, other springs among the rocks contributing
each its share of clear, cold water to the beautiful stream. The rocks
on its edge are covered with mosses and higher up the ferns are still
green and beautiful, although in some places they are nearly buried
with the leaves which have fallen from the trees above them. I
thought as we marched along that it would be a beautiful place in
May or June and doubt not there would be many new things there
for you and me.
I am enclosing some specimens of a beautiful little grass which
grows along the river. It is all I have to send this time except my
Main — Life and Letters of Edward Lee Greene.
153
best wishes to yourself and family. Hoping that this rebellion may
soon be ended and peace restored, I remain,
Your sincere friend,
Edward L. Greene.
Write as soon as convenient and excuse my hastily written letter.
Ft. Donelson, Tennessee, March 12, 1863.
Dear friend Kumlien:
Yours of March 1st was received this morning with usual welcome
and read with pleasure. I excuse you of course for not answering
every letter of mine, knowing that you are otherwise engaged. I
am glad that you are ready now to begin the labors of another sea¬
son and wish you eminent success and regret that I cannot share the
pleasure of roaming about the prairies, woods, and marshes of old
Wisconsin with you. But I shall remain contented with gathering
what I can from Tennessee during the coming season and trust I
shall find many new things if we remain here. Since I last wrote to
you we have been removed from Ft. Henry and have taken up our
abode at this place and shall probably remain here for some time
to come. At least so long as there is any danger of another attack
being made, which there is at present. Father has been with us for
several weeks now. I thank you for the compliments you sent by
him, but I don’t think there is any great prospect of my becoming em¬
inent as a naturalist, situated as I am with nothing but my own
hands and nature before me. But as I love the study I shall give
all my leisure to it and the pleasure of the passing time will be
enough to repay me for my labors. Father brought me a new work,
one which mother procured for me, namely, “Principles of Zoology by
Agassiz and Gould,” a very interesting book to me and one which
has already given me many new ideas of nature beyond botany.
Should I ever be relieved from my present situation, I shall pursue
the study of the animal kingdom as eagerly as I do botany at present.
Spring is very evidently with us now. Hepatica triloba acuta is
in flower. I never saw any flower appear more romantically situated
than this little plant as it grows in crevices of the huge and almost
perpendicular masses of rock which rise from the south bank of the
river. High above the reach and among the verdant mosses this lit¬
tle herald of spring now shows its flowers, all of which are white. I
have seen no blue ones. Also Anemone thalictrordes (rue anemone)
is in flower on sunny hillsides in the woods, not so common as in Wis¬
consin and rather larger. A. nemorosa (wood anemone) I have not
seen and doubt whether it grows in this latitude. I find also with
Hepatica by the river a new plant, a specimen I will enclose. From
its resemblance to Saxifrage pennsylvanica (swamp saxifrage) of
Wisconsin I think it is a saxifrage, but don’t know for I have almost
forgotten the character of the genus.
Other plants (new to me) will soon be in flower, among them a
phlox which is almost ready. I also received a letter from Prof.
Wood this morning. It was in reply to one which I wrote him for
154 Wisconsin Academy of Sciences , Arts, and Letters.
information about getting a flora for this season’s use. The new
class book seems to be the one I want and he kindly offers to send it
to me postpaid for two dollars, although since the rise in the price of
paper the retail price of the book has been raised to two dollars and
a half. His chirography so closely resembles yours that before I
opened the letters I supposed them both from you. He writes a good,
sociable letter — says he has tramped about this forest and hill with
box, press, spade and compass and says it is a very rich field for bo¬
tanical researches.
I will send his letter to mother and if you wish to see it you can do
so by calling at mother’s. He gives the same name to that fern which
you do. Mother writes that one of her cacti is preparing to flower.
I will collect for you a bottle of insects as soon as they become plenti¬
ful. I undertook to do so last fall, but it was too late in the season.
About the reptiles of this section, I think they are more numerous
than at the north, especially snakes, which, according to what the
darkies say, are venomous. I must try to send you some specimens
of that flat little lacertian called the “swift.” It is quite common
among the rocks along the Cumberland. I killed two last fall but
could not preserve them, not knowing where to get the bottle of alco¬
hol. But I think I can procure some little things of that kind at the
hospital. I shall endeavor to make a collection of the shells of the
Mollusca of the Cumberland if we remain in its vicinity until the
river falls. I must close for this time, begging that you will excuse
bad penmanship. Give my respects to your family and think of me
ever as,
Your friend,
Edward L. Greene.
Nashville, Tennessee, April 18, 1863.
My dear Friend:
I am seated once again to pen you a few lines, not because I have
much to write, but I have a few specimens of an interesting little
plant now pretty nicely dried and I want to send them before they
get mouldy. I haven’t a book with me so cannot tell you what the
flower is, but you will readily find its name from the dried specimen.
I should at first sight pronounce it a Dicentra were it not for the fact
that only one of the sepals is hooded; it is quite a singular looking
flower and I think belongs near Dicentra. It grows in just such lo¬
calities as the dutchman’s breeches usually do, but the root is very
different from that plant.
Associated with it grows an Urtica (nettle). It is not more than
six inches high and was in flower when I took these specimens, and
today I am going to take specimens of it. I found a day or two since
plenty of Ranunculus pusillus (spearwort) but not yet in flower. I
have also, I think, found two new species of Dentaria, (toothwort or
pepper-root) ; one of them is diphylla (toothwort or crinkleroot) I
believe and the other is a very small Dentaria and is not yet in flower.
I have not yet been about in the wild lands much, but shall try to
Main — Life and Letters of Edward Lee Greene . 155
make a trip into the country soon. I may find a good many new
things even here in the vicinity of Nashville.
Our regiment has been assigned to duty at this place and we shall
no doubt remain here during the summer. About once a week I have
to go to Louisville, Kentucky as train guard. On this road we pass
within nine miles of the entrance to Mammoth cave. If it is possible
I will go there to see the great cave. Oh how gladly would I have
you with me if I am so fortunate as to go.
It will soon be with us here the pleasantest part of the year; al¬
ready “frondent silvae” and the birds and wild flowers are here in
plenty. But I suppose you have not collected many yet. May these
lines find you in good health and ready to commence your spring la¬
bors. Please answer when convenient and give me the name of the
enclosed plant. With my best wishes to your family and yourself, I
remain your friend,
Edward.
Ft. Donelson, Tennessee, May 6th, 1863.
Dear Mr. Kumlien:
I gladly seat myself this evening to reply to yours of April 29th;
glad to hear from you again so soon, and to know that you have re¬
sumed your labors for another season. I am still blessed with health
and remain in the hospital quite well contented with my situation.
Not many sick at present. I have my new book, thanks to Prof.
Wood! It is, I think, a great improvement on the old edition. Said
to describe all known plants, (lower Cryptogamma excepted) in the
U. S. except those of South Florida and west of the Mississippi river.
But I have taken specimens of an Allium (onion garlic) here in Ten
nessee that is far from being described in the book and also an Eri-
geron (daisy fleabane) that I think is not. The descriptions are, I
think, greatly improved in this edition and the number of genera is
reduced greatly. Pentalophus is again united to Lithospermum. Oto-
philla to Gerardia , and many others have returned to where they be¬
longed. It describes 186 Carices (sedges), 48 Solidagos (goldenrods)
and precisely the same number of asters, which is rather singular.
When the Carices come in fruit I shall find more species than I ex¬
pected to. There is a very beautiful one growing on the rocks along
the river with short leaves that are an inch broad, pale green, almost
white and translucent near the base. Many interesting flowers are
now appearing, Bignonia capreolata, (cross-vine) is just beginning to
flower, a fine evergreen climber with large dark green, eliptical
leaves and large yellow and red flowers. Houstonia caerulea (bluets,
innocence) grows on dry ground and is smaller with very dark blue
flowers. Houstonia purpurea (bluets) is handsome, with very deli¬
cate corollas, closely resembling those of Mitchella repens (partridge
berry). Cynthia dandelion (dwarf dandelion) is handsomer than
Cynthia virginica and bears a large tuber at the extremity of the root
about an inch below the surface. Silene virginica (fire pink, catch-
fly) is the richest thing that I have yet found, very common, with
156 Wisconsin Academy of Sciences , Arts , and Letters.
large scarlet flowers. Pyrus angustifolia (crabapple) is more deli¬
cate than the Wisconsin crabapple, but at a distance it appears the
same. I have at last found a little marsh of an acre or two, but it
contains no flower except a small delicate Scirpus (bulrush). But
it contains several plants that I do not find elsewhere; one of them I
suppose to be Geum rivale (water or purple avens) has very hand¬
some pinnate leaves, but no flowers as yet. You will find Trillium
grandiflorum (large flowering trillium) almost anywhere in the
woods between Clinton and Mr. Odell's. You will find it in the thick
young timber in moist shady places. It is not half as large as the
other species is here. Trillium sessile is the common one here. It is
small and not remarkable for beauty. I will give you the name of
most of the plants that I have taken on the opposite page and close.
With my best wishes I remain,
Ever truly yours,
Edward.
(There follows a list of fifty -two plants which I will not enumerate
here).
Ft. Donelson, Tennessee, May 12, 1863.
Friend Kumlien:
Yours of the 2nd inst. came to hand last evening and I am seated
this morning to pen you a few lines more. I am obliged to you for
your instructions and I shall do the best I can at collecting zoologi¬
cal specimens, but I do not know how much I shall be able to do in
that line, but shall probably send you some new things. Wish I were
able to stuff you some of our birds. I saw a red-headed woodpecker
the other day as large almost as two yellow-hammers, mostly black
except its red head. (The ivory-billed or pileated woodpecker). I
have seen several of them, besides many others that I never saw in
the north.
The Euphorbia (spurge) that I sent you is E. commutata I think
without doubt. It is distinguished from all others by its “floral
leaves, broader than long and so applied as to appear connate-perfol¬
iate.” I sent the whole plant except the root, but it is in fruit now,
also still in flower, but appears very different from what it did when
I took those. I shall preserve more when I go where it grows again.
It is by no means common here. Flowers are increasing in numbers
rapidly and they will get the start of me before long I fear. Yester¬
day I took a single specimen of Phlox maculate (wild sweet william).
It surpasses any phlox I ever saw in beauty, P. drummondii of the
gardens scarcely excepted. It is not very common here. Many very
rich flowering trees and shrubs are now in flower; among the most
beautiful are Wisteria frutescens, Bignonia capreolata (cross-vine),
Liriodendron tulipifera (tulip tree) etc. Pentstemon pubescens
(beard tongue) is in flower, quite handsome, and another species will
soon be. I found a small and elegant Desmodium (tick trefoil) in
flower yesterday, but could not determine the specific name with cer¬
tainty without the fruit.
Main — Life and Letters of Edward Lee Greene.
157
No orchids yet except Orchis spectabilis (showy orchis) and Cy-
pripedium pubescens (larger yellow ladyslipper) both out of flower
now. One plant which I had taken to be an orchid has flowered and
I am “up a tree” about it. I cannot find it described anywhere yet.
Certainly not an orchid but belongs to the Melanthaceae . It is not
described there nor in any of the adjacent orders. A two foot high
plant with a long nodding thick raceme of small, white fragrant
flowers, Hexandrous Monoggnous , stem nearly leafless, radical leaves
resembling those of Dodecatheon media (shooting star). Perhaps I
shall find its name when the fruit appears. It is very common on
high grounds and handsome. I have found a few specimens of Ap-
lectrum hyemale (putty root) the only new orchid, but it is not yet
in flower. The early flowering Carices (sedges) are now ready in
fruit. I have taken specimens of C. digitalis , C. steudelii, and
C. careyana, the latter a fine species with leaves an inch wide or
more, in which respect it does not agree with the description of which
says “6”, but it differs greatly from C. plantaginea in other respects,
but agrees in description of foliage.
I will send you a specimen of Silene virginica (fire pink) in this
if my specimens are not all too large. Its bright flowers have faded
in pressing.
I took a ride across the river last evening and found a new Ra¬
nunculus, R. parviflorus (creeping buttercup) a naturalized species
from Europe. I think of no more this time except to give you the
names of what I have taken since I last wrote to you. So good bye.
Edward.
P. S. Mother writes that several of her cacti are flowering this
spring. Go and see them. I wish I could. Remember my love to
your family all. We have just received glorious news from Virginia
and we begin to hope for an end of the war very soon.
(There follows a list of thirteen plants).
Ft. Donelson, Tennessee, May 25, 1863.
Dear Friend:
Although I have not received an answer to my last, I must chat a
little while with you this afternoon. I have just returned from a
tramp in the woods. The weather is very hot now and walking in the
middle of the day is very disagreeable. But still the flowers are in¬
creasing in numbers and I must try to keep up.
The composites are now beginning to flower. I saw two new ones
to-day that were just commencing, but did not examine them for I
have enough of others that are in full display and which will not wait
for me. I came very near losing Wisteria frutescens ; it is so un¬
common and I have had so many ways to go. But today I took four
specimens, but they are on the decline now. It is the most beautiful
native leguminous plant that I ever saw. Its thick raceme of large
blue flowers is often ten inches long.
11
158 Wisconsin Academy of Sciences, Arts, and Letters.
Of Bignonia capreolata (cross-vine) I got only two specimens, one
of them a very beautiful one and the other a poor one. Its flowers
are large and very evanescent, or like those of some species of Gerar-
dia, falling off when exposed to the sun for a short time. It is rather
more common than Wisteria , but grows only in certain peculiar situ¬
ations.
Tomorrow I must go after Aplectrum hyemale (putty root). It
must be out now, but I know of only one specimen which grows about
a mile and a half from here. I may find more. Philadelphus grandi-
florus (mock orange or syringa) on the river banks is now in flower
and how beautiful. I have found two new Vaccinia, V. stamineum
(squaw huckleberry) and V. arboreum (farkleberry) ; of the former
a small twig is enclosed, the latter is not yet in flower. I also enclose
a Verbena angustifolia (vervain). It is the most common species here,
growing in dry and open grounds everywhere. Wood says “corollas
deep blue” but they are invariably pale blue.
The Carices (sedges) are coming on finely. The most common one
I trace by the table to C. vestivalis very plainly, but it differs greatly
from the description and I cannot determine its name. The Aristo-
lochias (birthwort) are very interesting plants. A. serpentaria (Vir¬
ginia snakeroot) is quite common, but I have seen only one of A. to-
mentosa (similar to dutchman’s pipe). The little Ranunculus enclosed
I cannot name. It is not described in my book. Grows in a little
pond with Rumex verticillatus (swamp dock). I have not found more
than a dozen specimens.
Fraser a carolinensis (american columbo) is a large homely plant,
but would be handsome if the flowers were of some bright color. The
flowers are in a large terminal panicle and are quite curious looking.
I will give you the names of what new ones I have taken since I
last wrote you and close with my best wishes for your welfare.
Edward L. Greene.
(A list of twenty-five plants follows).
Ft. Donelson, Tennessee, July 15, 1863.
Mr. Kumlien, my Friend:
As I have a little spare time this morning, I will try to write you
a few words that you may know that I am well and still pursuing
my favorite employment. Within a few days I have taken specimens
of some very choice southern plants. Among them is the Platanthera
enclosed. It is, I believe, P. perzamaena (fringed orchis) and the
handsomest species that I have ever seen. It is quite common here
along the streams in shady places, and with the exception of a single
speciment of P. flava it is the only Platanthera that I have seen in
this country. The enclosed specimen is a very small one, the smallest
I could find, and notwithstanding I pressed with a hot iron the flow¬
ers have lost their color, which is a rich purple.
The little green Liliaceous flower, although it has no beauty to
recommend itself, is not unworthy of notice. It is a flower of Agave
Main — Life and Letters of Edward Lee Greene.
159
virginica (false aloe) and I do not hesitate to say that it is the sweet¬
est smelling flower that I ever saw, I have seen only one plant in
flower. The scape is more than six feet high and the raceme of flow¬
ers about three feet long. This specimen stands about half a mile
from where I am sitting and I go to it every evening to inhale its
delicious fragrance. But before many days I shall have to go out
and end it into specimens for the press. I hate to do it as long as
there are any flower buds to open, but I must. I wish I could find
more of it. It grows plentifully on the lime ridges around Ft. Henry.
I saw its leaves and the scapes of last year there last spring. I am
lucky in finding this one specimen here so I will make the best I can
of it and remain contented.
Summer is rapidly passing away and the fall flowers are preparing
to make their display. The Gerardia will soon be in flower.
I have taken specimens of Seymeria macrophylla (mullein foxglove)
which is closely related to Gerardia. It stands next to Dasystoma in
my book and you would take it to be a Dasystoma at first sight. It
is a large bushy herb six feet high, very diffusely branching.
Another common and very beautiful little herb, Sabbatia stellaris,
(sea pink) is just beginning to flower. The texture of the flower is
very delicate and I cannot press it so as to have it look natural. Since
I last wrote I have found another species of Passiflora (passion
flower) P. lutea, a small and slender species; have no specimens
pressed yet. Ozydendrum arbor eum (sour wood) is now in flower in
the woods and makes quite a show. It comes next after Andromeda
and scarcely differs from that genus, except in its size and manner
of flowering. It bears a large spreading panicle composed of numer¬
ous one sided racemes at the extremity of each branch. Tecoma ra¬
dio ans (trumpet flower), one of the most beautiful things that I have
seen in the south, is in flower. It belongs in the order Bignoniaceae
which is near to Scrophulariaceae. The flowers are large and beautiful
in clusters at the ends of the branches. The corolla is two and one-
half inches in length.
Mimulus alatus (monkey flower) takes the place of M. ringens
(northern species of monkey flower) here. Galium pilosum and G.
circaezans (bedstraws) are the most common species of this genus.
I have taken a few specimens of Aster oblongifolius. It is the earliest
species here. I don’t know that I have seen it in Wisconsin, but per¬
haps it grows there. I will enclose a specimen of Forsteronia diffor-
mis, a slender vine and rare here. I have seen only one plant. It
belongs to the Apocynaceae (dogbane family), next genus after Am-
sonia.
There are several species of Desmodium (tick trefoil) and Lespede-
za (bush clover) in flower but I have not examined any of them ex¬
cept Desmodium rotundi folium which is a large spreading species. The
Desmodium which bears leaves and flowers on different stems is pre¬
paring to flower. Yesterday I found Plantago aristata (plantain), a
very small and singular looking Plantago with linear leaves and bracts
attennated into long rigid awns.
160 Wisconsin Academy of Sciences, Arts, and Letters.
I would like to know just where you are today and what you are
doing. If our armies gain many more such glorious victories as we
have recently been doing I think we may hope for and expect peace
soon. I am making no other calculations than that I shall be at home
to commence botanizing in your company next spring. Perhaps I
shall be dissappointed but I think not. The news is cheering of late
and I think the rebels are about whipped. It will soon have been a
year since I enlisted, and how rapidly that year has passed away. I
have met with no serious misfortunes, my health has been preserved
and I have enjoyed my soldier life very well. I have seen many beau¬
tiful things in nature during my year’s travels through the south. I
am now ready to return to Wisconsin at any time when this war shall
be brought to an end. I thank God for all his mercies and blessings
which he has so kindly bestowed upon me and wait patiently until
the time comes when I and all the rest engaged in this cause shall be
permitted to return to our homes. I will close for this time with love
to you and to your family and hoping soon again to enjoy intercourse
with you all as in days past. I remain with all kind regards your
friend,
Edward.
P. S. Please write as soon as convenient.
Huntsville, Ala., Jan. 4, 1865.
Esteemed Friend:
Your kind letter of Nov. 13th is before me, and I will endeavor to
reply to it since railroad communications are once more, at least par¬
tially, opened between us. Your letter was a long time in finding its
way to me, and perhaps this may be as long in finding you. Doubt¬
less you are well aware that for a number of weeks past the position
of Hood near Nashville has prevented any correspondence between
us, and this is the reason why I have delayed writing to you for so
long.
In the first place I must give you a brief sketch of my military ex¬
perience since I last wrote to you, though I have not yet seen much
of what the ancients call the “gloria belli.” Amid all the recent fuss
and fighting in our department, our regiment has not been called to
take any active part. But we have indeed seen rather hard times,
suffering from sore feet and weary bodies; short provisions have been
experienced by many of us to an uncommonly oppressive degree.
I had well nigh lost all my botanical and other little collections on
my last march. We were ordered away from Claysville soon after I
wrote my last letter to you and were obliged to burn up a good many
valuable things to prevent them from falling into rebellious hands
after our departure. I managed to crowd most of my plants into my
knapsack by carrying some of my clothing in another manner and
have preserved now the most of them, and as soon as an express is
opened, which I am informed will be soon, I shall send them home.
We are perhaps permanently stationed here in Huntsville, the most
beautiful little city I have ever seen. We have fine, pleasant
Main — Life and Letters of Edward Lee Greene.
161
weather since the commencement of the month, though we had cold
rains and a few slight flurries of snow during the week intervening
between Christmas and New Years.
Three weeks more and we shall have spring flowers ! We had them
on the first of February in Nashville last year, crocus and such early
garden exotics.
This is probably my last winter in the U. S. Military service. I
have only seven more months to serve. I think, however, that it may
not be my last winter in the south if my life is prolonged. However,
I do not think of ever taking up my abode in this part. I should by
far prefer the opposite side of the Mississippi river somewhere. The
climate of this southern latitude is so much better than those hyper-
frozen regions, to my idea at least.
But I will cease from building these airy castles for the future
since it is so impossible as you say for us to know what it will realize
to us. I suppose we who acknowledge our dependence upon the mer¬
its of our Lord and Savior for our hope of eternal happiness should
be more willing to submit to do His will when, where and however He
may see fit, than to accomplish our own plans. Yes, certainly we
must and we cannot exercise proper faith in Him unless we are will¬
ing and ready to prefer what may seem to be His will to our own.
I am glad to perceive by your writing that your hopes of eternal fe¬
licity beyond this scene of strife and mortality are centered in the
merits of the crucified Son of God, which I too believe to be the only
hope for fallen and sinful humanity. And while it is a lamentable
fact that none of us who have come to years of understanding, have
not sinned and are not dependent upon the mercies of Christ; it is
also a fact, a blessed one too, that His salvation is able to save even
the most degraded sinners if they will but turn to Him and receive
Him as their Lord and Savior. Let us see then that with an unan¬
swering faith we rely upon Him and doubt not that He will save us
and that none other can. I find that it is not sufficient that I assent
to the fundamental articles of Christian faith, submit to the ordi¬
nances and sacraments of the church and abstain from outbreaking
sins. All these of themselves do not rob death of its gloomy aspect,
but these words, “God so loved the world that He gave His only be¬
gotten Son that whosoever believeth on Him should not perish but
have everlasting life,” are of themselves sufficient if I by faith apply
them to my own case.
There was a time in the life of the lovely and noble Martin Luther,
when these words of his credo “I believe in the forgiveness of sins”
gave him no consolation though he thought he believed them. But
when Staupits directed him to believe that not only were Paul's and
Peter’s sins but his own were forgiven, then it was that sorrow and
gloom left him, and death was stripped of its terrible aspect, and he
afterwards became what he was as a minister of the gospel, and re¬
former of the world. Pardon me if it may seem that I am writing
a tedious letter, but let me say to you as a brother in Christ, do not
only hope that Christ’s mercy will extend to you, but doubt not that
162 Wisconsin Academy of Sciences , Arts , and Letters.
it does , for I believe it to be the privilege of us all to have no fear of
death. All that is necessary to bring us into this blessed state is to
let Christ be indeed our own personal Savior and Author of eternal
life.
I must bring this to a close. The enclosed little lichens are some
which I took on the bank of the Tennessee river at Claysville on cedar
trees. I sought for more but they were evidently very uncommon.
It is a species entirely new to me. Let me hear from you again when
you find time to write. My best and kindest regards give to your
family. God bless you and them is the prayer of your unworthy
friend,
Edward.
Direct to Huntsville, Ala.
Huntsville, March 18, 1865.
Mr. Kumlien.
My dear Friend:
Before me lies your good letter of the 5th inst., which I will now
endeavor to answer immediately, for tomorrow we move to commence
our spring campaign. We are ordered to Knoxville, Tenn. and prob¬
ably from thence we shall go into Virginia. Until quite recently we
have been expecting to remain at this post during the season, but
being unexpectedly transferred to the 4th army corps, which is on
the move, our prospects are changed quite a good deal. It looks now
quite possible that we shall see more severe service than we have
done heretofore, and I shall probably have the pleasure of seeing
new territory before I get home.
Last night I dreamed of seeing lofty mountains whose sides were
clothed with the richest verdure of various kinds of trees and W'hose
summits rose above the floating clouds. I thought that with Chap¬
lain Foote I endeavored to ascend one, but it was so steep that I be¬
came extremely tired and then I awoke from my vision to find myself
on the same couch where I have slept for two or three months past.
But I am not sure but that I shall not at least partially realize my
dream for it is said to be a very mountainous country through which
we pass even on our way to Knoxville, and beyond that are several
ranges of mountains. It will soon be the most delightful part of the
year for traveling in this country. Trees are beginning to put forth
their leaves. Peach and plum trees are in bloom, elms and maples
in fruit. But it is rather strange that I find no new wild flowers yet.
Ranunculus recurvatus (hooked crowfoot) Taraxacum (dandelion)
and other flowers that are not the earliest in our country have been
in flower here for weeks past. Myosotis (scorpion grass, forget-me-
not) is the only plant I know of now in flower that does not grow in
Wisconsin.
In the gardens now are innumerable varieties of crocus, hyacinth,
narcissus, etc., etc., in bloom and the air is redolent with their
fragrance.
Among the feathered songsters are the mocking bird and pine gros-
Main — Life and Letters of Edward Lee Greene.
163
beak that I believe we do not have with us at home. Both are beau¬
tiful and accomplished musicians, taught not by man, but by their
creator.
I do hope that you may be permitted to prosecute your plan of a
trip into Kansas. Judging from what our boys say of that country
I conclude that you will find it a very rich place in both botany and
geology. But shall I not be deprived of your correspondence during
your absence from home? Probably this will be necessary for I shall
not know where you are. But if you spend three months there, Prov¬
idence permitting, I shall be at home again almost as soon as you
will and may see what you found and enjoy a narrative of your ex¬
pedition.
I say again I do hope you may go. By all means let me hear from
you once more before you start. I am glad to hear that although
your little ones have been unwell yet you by God’s blessings have
their health restored.
I thank you for your kind admonition with regard to abiding in
close intimacy with God our Saviour. Although in nature we see
many evidences of His love toward His creatures, but in the great
book of His we have a still more wonderful evidence of His goodness.
We have received the emblems of His broken body and shed blood and
I trust a portion of His spirit also. So let us cultivate it while this
life is given us so that death will be to us only the entrance to a true
life, a glorious one! I am glad for your sake and for her sake and
for the sake of God that you have a sister so able to give you counsel
and cheer in the comforts of our holy religion. I say our religion
for it is all the same in every fundamental principle and I with you
can see no reason why you need leave the communion of the church
into which you were baptized. With me it is all the same and I
would willingly for the sake of harmonizing the great body of
Christ’s divided church consent to unite with the one to which you be¬
long if it were expedient. I consider that every sect which does in¬
deed build on the true foundation, i. e. Christ, is a part of Christ’s
church. The superstructures are more or less imperfect in different
sects and which is more perfect I am unable to decide and so I con¬
clude that it is not material to salvation that we belong to this or
that denomination but are at liberty to unite our tastes and our judg¬
ment in this matter. But my Sabbatharian brethern say I am too
liberal in these views for a Sabbatharian and doubtless it is so. Nev¬
ertheless I am not obliged to sacrifice what I believe to be the truth
on that account. But I shall trespass on your patience, so good bye
and God bless and prosper you.
Ever your friend,
Edward.
My kind regards for Mrs. K. and your children. Ed.
164 Wisconsin Academy of Sciences , Arts , and Letters.
Camp of the 4th A. C. near Nashville, Tenn., May 2, 1865.
Mr. Kumlien.
My dear Friend:
Yesterday I was all day thinking about you and wondering whether
you had yet set out for Kansas. I was expecting a letter from you
and at night my expectations in that line were realized by the arrival
of yours, bearing date of April 23, 1865.
I am partly sorry that you cannot prosecute your plans for a wes¬
terly trip for it would be such a pleasant one if it could be made with
profit, and for the best part I am pleased with the hope that I shall
have the pleasure of a social interview with you sooner than I should
had you made the contemplated expedition.
We are all feeling quite confident that we shall be at home before
the first of July, and we hope even sooner than that. Evidently
the war is ended and every move seems now to indicate a speedy dis¬
missal of a large number of troops. Even today soldiers are being
mustered out here at Nashville.
Another year may afford a better sight for an expedition westward
and allow me the delight of accompanying you! That is if we can
arrange matters so that my presence with you would not be detri¬
mental to your success, etc. etc. I have almost made up my mind
that I would make my home somewhere in that trans-Mississippian
region though possibly farther to the southward than Kansas.
Well I have done some traveling and seen some new country since
I last wrote to you. Our Corps started from Huntsville about the
20th of March for Lynchburg, Va., for the purpose of assisting in
the capture of Lee should he vacate Richmond and endeavor to escape
to the west. We had proceeded but half way, however, when we re¬
ceived the intelligence that Lee had surrendered. Our Brigade went
as far as Jonesboro in the northeastern part of Tennessee, the re¬
mainder of the Corps stopping at other places in the rear of us. At
Jonesboro we remained two or three weeks, and then were ordered to
Nashville. We came back by railroad most of the way and here we
remain and probably we shall be kept here until mustered out. At
Jonesboro, Tennessee we were encamped in a full and delightful
view of the great Smoky Mountains of North Carolina, but the near¬
est point of the mountains was twenty miles distant and of course
a visit was impracticable. But you will easily imagine with what
longings of heart this child used to stand and gaze at those sublime
but rugged heights. I knew that there must be on those summits
many things which to me would be new and full of interest. No, I
seldom went out of my tent without casting a worshipful look into
the south. Indeed the picture in the distance was beautiful to look
upon in clear weather. We were informed that directly south of us
in that range was the highest mountain east of the Mississippi, and
geological reports etc., which Chaplain Foote picked up about town
increased our desire to make a trip, but it could not safely be done by
the few who were anxious to go, so it is all well.
Main — Life and Letters of Edward Lee Greene.
165
The immediate vicinity of Jonesboro was dry in botanical novel¬
ties to me. I only took two plants, Carices nigro-marginata, and um-
bellata (sedges), from its precincts. I will enclose a few of them in
this although they may not be new species.
On our advance from Huntsville we stopped half a day at Chatta¬
nooga, Tennessee, and during that time I with several others climbed
to the top of Lookout Mountain. Climbed , I say, and indeed climbing
it was, and the most severe half day’s journey I have recently made.
It seems to me almost incredible that an army stationed on its sum¬
mit should ever have been defeated by a force coming from below, yet
it was done successfully by General Hooker’s army. On the top of
this height I found a few flowers which I have never seen elsewhere,
and these are a sufficient compensation for my labors of that after¬
noon, severe as they were.
The lichen enclosed is from Lookout Mountain. There were large
ones from this species but dry weather had reduced them to such a
fragile condition that I could only preserve small pieces. They grew
on the sides of the rocks at the summit.
You express the same sentiment with regard to the death of Mr.
Lincoln which is felt by us here. We received the terrible tidings
while at Jonesboro, but were slow to believe such a horrible report-
But it is no great wonder that we who are but the merest children
in knowledge and foresight in comparison with the omnipotent, should
sometimes be shocked and almost disheartened by the manifestations
of His unerring hand, for His course sometimes seems to be directly
contrary to the one which our best wisdom and most perfect skill
would mark out.
He moves in a mysterious way
His wonders to perform
He plants his footsteps on the Sea
And rides upon the storm.
And dark and mysterious though they seem to our perceptions, yet
true faith leads us to submit all our ways to Him, and then we have
peace and assurance that we will be well in the end.
Even now we begin to believe that Mr. Johnson is going to be just
the man to decide the fate of the arch traitors, and we think that our
beloved Mr. Lincoln would have been too lenient towards them, and
by and by we may see that it was well that God called him who was
our choice anyway.
The memory of Lincoln is immortal in the minds of our people; he
has fulfilled his great mission with faithfulness and has laid aside
the mortal and put on the immortality. The great responsibilities that
were resting on him as President of the U. S. in the day of the na¬
tion’s peril, is removed and he rests, we hope, with God, for he con¬
fided in that name in which to trust is to gain the victory of death
and to secure to the soul of fallen man a life that is eternal.
166 Wisconsin Academy of Sciences , Arts , and Letters .
With best wishes to you and your family, I subscribe myself, with
love
Your friend,
Edward Lee Greene.
Write soon and address
3rd Brig., 3rd Div., 4th A. C.,
Nashville, Tenn.
Monticello, Ill., May 1, 1867.
Mr. T. Kumlien,
Albion, Wis.
My dear Sir:
It is now a little more than three weeks since I visited you, and this
morning I propose to have a brief chat upon familiar subjects, trust¬
ing the proposal will be met with pleasure on your part. And in the
first place I am well and, after having employed a couple of weeks in
visiting relatives, etc., I am at last settled down to labors and studies
for the season. I find myself very happily situated and have a fine
prospect of making many new acquaintances in Flora’s domain, and
some among the Mollusca for I believe there are a goodly number of
species of land snails in the low timber along the river and descrip¬
tions of them have commenced in the April No. of the Naturalist.
The weather here seems to continue rather cool and farmers com¬
plain of a late spring, neverless the woods are becoming green and
there are a good many flowers already.
On the 13th of last month I found in the woods growing and flow¬
ering with Hepatica, the little trillium which I enclose. What species
it is I am unable to determine. It is far too large for anything
which Mr. Wood describes except T. nivale (dwarf white or snow).
In nivale according to that author the peduncle is very short and
erect, whereas in this they are comparatively long and specimens
wdiich I took on the 15th had the flowers completely deflected beneath
the leaves. Besides, a damp northern forest hill side is not the place
for T. nivale whose habitat is “dry fields.”
Among the plants I have already taken are the following of inter¬
est perhaps to you namely, Isopyrum biternatum , Cardanine rhom-
boidea (bittert cress) Claytonia virginica (spring beauty) Trillium
recurvatum (wake robin) and Mertensia virginica (Virginia cow¬
slip, bluebells), the latter a plant of most exquisite beauty whose
equal in this respect I do not expect to find during the season. It is
going to be difficult to preserve, but with proper care doubtless I
shall save a few nice ones. Claytonia (spring beauty) is abundant
in all soil and extremely variable, yet I think there is but one species
here.
Five weeks hence I shall probably have my leisure hours all occu¬
pied if I attempt to preserve half a dozen representatives of each new
species that I shall find. Would that you could be here to “help your¬
self” and to add to my joy. Tell my dear friend Ludwig I wish he
were here to collect eggs and skins in Illinois and be my companion
Main — Life and Letters of Edward Lee Orreene.
167
in my rambles. I have seen not a few birds already which I have
not noticed in Wisconsin. A red-headed woodpecker is quite frequent
which is entirely different from our Wisconsin red-head, being rather
larger and I think much handsomer. The marking of its body are,
I should think, a kind of gray and white and the top of its head as
light as scarlet. A pair of cardinal grosbeaks are building a nest
near my boarding place. If I can keep the mischievous boys away
you shall have their eggs. Wild turkeys I have frequently met in
the woods. They breed here I am told, but they say it is time for
them to commence hatching. I have sought for their nests but thus
far in vain. Birds large and small are abundant here, and I am told
that the larger waders, swimmers, etc., have their breeding places in
those parts of the prairie that are yet unsettled and where the great
grasses and sedges still flourish in all their glory in a virgin soil.
The nearest locality of this description begins some twenty or thirty
miles east of us. I almost peruade myself that I shall go there in
spite of the difficulties and spend a day. This country is very differ¬
ent in its appearance from Wisconsin prairies; here are merely little
patches of a few hundred acres, but they spread out before the eye
just like the broad expanse of old ocean itself. Timber grows only
along the rivers. The land is generally low, rocks and hills scarce,
and the climate rather unhealthful.
If I have health, I remain here a year for teachers get very liberal
pay. I shall have no difficulty to get from fifty to eighty dollars a
month during next fall and winter after which I intend to go a little
way north to some good school. At Bloomington in this state and
only some thirty miles north, is located the state university. I am
going to see it sometime during the season if practicable. The state
society of Natural History has here its museum. According to the
published transactions this society bids fair to be something. Indeed
it already is. It is exclusively a society of Natural History and the
only one in the state. It is possible that I shall attend school there,
though hardly probable for I have no doubt the school is better at
Evanston near Chicago. But no more at present. With love to your
family and requesting the favor of a letter soon as convenient I am
yours in bonds of friendship.
Edward L. Greene.
Address —
Monticello, Pratt Co., Ill.
Monticello, Ill., Tuesday, Nov. 9, 1869.
Very dear Friend!
“Tis a time for memory and for tears.” One of the most genial and
refined spirits which it has been my happiness to know has passed
away from earth. The startling and painful news reached me a few
days in advance of your letter by one from my mother. And you
thought of me, so did I of you. The dear old man had few except
his kindred who appreciated him as we did, and how sadly it rests
upon my heart that neither my mother nor yourself knew of his
168 Wisconsin Academy of Sciences, Arts, and Letters.
sickness even until he was gone. Had it been possible I would have
been with him in his last moments. . It seems hard to think that they
who were so much attached to him as I know you and my mother
were should not have seen him after the spring of life began so rap¬
idly to fail, yet the fault, if there be any, is not yours. Yes, I shall
miss him. I have during my absence from home enjoyed an occa¬
sional letter from him, and now this time I sent him my first letter
on the very day he died.
When I shall return to Albion, be it sooner or later, I shall then
miss him more. Instead of a cordial welcome to his pleasant home,
I shall find his grave. And a gloomy place will be the spot where his
hand reared flowers. This sad event will admonish us anew that the
friendships of earth are only transient (or rather that their constant
enjoyment is a thing not to be expected) for “amicitia vera est sem¬
piternal and we cherish fondly the memory of friends departed and
hope to enjoy their society again “when the dream of life is fled.”
I now realize as I never did before, the fact that if I should live
to old age, I must see yet many dear friends pass away from my side
“into the silent land” and it seems that my old age will on that very
account be sadder and more devoted to sacred themes than was my
youth. No, it seems not strange that the aged are usually less mirth¬
ful than the young. I have never lost a parent nor a brother. I
have not known the pangs of parting for time with the very near and
dear, and the news of Mr. Clark’s death reveals to me that I loved
him as I have not loved many whose remains I have seen consigned
to the grave. I could scarcely control my feelings sufficiently to per¬
form my school duties on the day I heard of it. Seeing that these
dear ties of earth are so slenderly united, let us cherish them the
more fondly while they last, and when broken we shall await in
sweeter anticipation the day that shall in God’s infinite love and
mercy unite us again to the dear ones gone before.
Monticello, Ill., Sunday, March 27, 1870.
Prof. Thure Kumlien,
Bussey ville, Wis.
My very dear Friend:
You are kind indeed to give me two letters for one. I had thought
to wait until I should have obtained a view of the “everlasting hills”
and get some new things to tell you before I would answer yours
which I received some weeks since. But you write again in the
meantime and ask for a line before I start. Well here you shall have
it and very welcome you are to it. I had for a week or two been
looking for a letter from Ludwig. I am sorry to hear of his afflic¬
tion.
Well, I feel as if I were about done with this country for the pres¬
ent. One week more to teach and then I am off. I hope to start
from Decatur for St. Louis one week from tomorrow, which will be the
4th day of April. From St. Louis I go to Kansas City, thence by the
Main — Life cmd Letters of Edward Lee Greene.
169
Kansas Pacific railway directly to Denver, Colorado. The road is
now finished to within a few miles of Denver.
I am, as you suggest, going to take a good supply of paper. I
have reflected on the probable scarcity of that article at some points
I may reach. For Ludwig I have already packed some little vials
of alcohol for insects, and I will not fail to do my best for a few
skins for you.
We have spring now pretty well settled in this latitude. Spring
wheat and barley were sown last week quite extensively. But as it
rained a good deal yesterday, work of this kind will be delayed for a
day or two. Birds of spring are plenty. Three weeks ago or more,
I guess four, we had the bluebird and the robin. Blackbirds and I
cannot mention all I have heard within two weeks, and a week ago I
saw a pewee, and they have been singing every morning during the
past week. Ducks and geese in almost endless droves have been fly¬
ing, and some cranes. But the waders and swimmers are not so
plentiful as they will be in a couple of weeks. On Wednesday last
I saw a fine flock of sandhill cranes sailing high in the upper azure.
They were led by two beautiful specimens of the white plumaged spe¬
cies. I have seen several small flocks of the latter, but they seem
fond of the society of their brethern of the gray coats.
In a couple of weeks more I shall probably find flowers. Even now
the hazel bushes are in flower.
I think some of stopping on my way, for a few days, at the coal
mining region a few miles east of St. Louis. I have formed the ac¬
quaintance of several genial persons who live there, and they urge me
to call on them. It will be an interesting place especially for a geol¬
ogist, which I am not however. But I may nevertheless see interest¬
ing trees and possibly yet a few flowers, for the spring is two weeks
earlier there than here. In East St. Louis lives Mrs. Hinchclifl:
whom you have seen, but I have not, yet on account of the dear mem¬
ory of her late venerable father, I will try to find her.
Tell Ludwig I will perhaps send him a letter from some point on
my way and to you I will write as soon as I arrive in Colorado.
Now perhaps another thousand miles or more may separate us, ere
we hear from each other again, but we cannot join hands and say
farewell. Think of me sometimes. Go and see my mother’s flowers
for yourself and me when you pass that way. I would gladly have
seen my Wisconsin home and friends before I started on my long
journey, but it would take time and money, and not least, the sad¬
ness of the parting again, would almost neutralize the joy of meet¬
ing.
With much love to you all, I bid you for a time adieu.
Edward L. Greene.
Denver, Colorado, Probably about April 15, 1870.
My dear Friend:
Here I am at my writing table, but I almost believe I had better be
out getting flowers; but it is a dry, hot afternoon so I will wait until
170 Wisconsin Academy of Sciences , Arts , and Letters.
towards night. There are not yet many flowers. I have however, al¬
ready a collection of fifteen numbers and I know of several more I
might add today. Every tramp I make I find new and unexpected
ones. There seems to be no spring in this country. It is either winter
or summer. During the two weeks (nearly) that I have been here, we
have had many dry, hot days, and one severe snow storm! No
April showers, no bursting leaf buds, no springing green grass, but
yet flowers and seeds.
The country seems almost destitute of deciduous trees. There are
several species of poplars, but I have seen no others. Not an oak,
hickory, or anything of the kind, but oh what forests of pine and fir !
I have already been some twenty miles into the mountains, but I did
not stay. It is too soon. However I found an old Wisconsin favo¬
rite on the lower range. It was our Bulsatilla (pasque flower) with
flowers a trifle more deeply colored, but in no other way differing
from the same plant of Wisconsin. With it grew also Ranunculus,
but not R. glaberrimus as described in N. A. Flora. Last week I
sent my mother a plant which I wish you may see when it flowers.
I should never like this country for a home. I love too well the
pleasant May and June of the North. The season of green grass and
above all of blooming fruit trees which I suppose can never be ex¬
perienced here. The winters are so changeable from cold to extremely
hot days that fruit trees do not succeed at all and the summers are so
long and dry that I suppose the beautiful European grasses must also
fail. The grasses of the plains only grow in tufts and patches and
never carpet the hot soil, yet there are lots and lots of flowers among
them. I see them now springing up everywhere and different kinds on
every bluff and knoll. (A few lines torn off here) are too apt to bring
frosts even in midsummer for corn to be safe. Yet the few who have
during the past five years given themselves to farming are now im¬
mensely rich, so high have been the prices of farm products in the
extensive mining districts of the mountains. Garden vegetables are
plentiful along the streams where they can be irrigated, grow very
luxuriantly, and to sizes almost incredible.
Pork is not raised for want of corn, but beef is easily fattened
on the grasses of the plains where the dried pasturage of winter is
almost as good as it is in summer. I will not write more at present.
Write to me and by the time I get your letter I will have ready for
you a package of Rocky Mountain plants. Now this I know will
be an inducement. Direct to Denver, Colo. I am as ever with love
to you all,
Edward L. Greene.
Dudley’s Ranch, May 12, 1870.
My dear friend Kumlien!
Your letter is very , very welcome. It does me much good to hear
from home and friends. I am not altogether gone crazy over the
world of beauty and novelty that now surrounds me. I am among
strangers of the human kind and do really at times feel that I could
Main — Life and Letters of Edward Lee Greene. 171
appreciate as I never did before the society of those whom I have
long known. But this is only a natural sequence and is both pleas¬
ant and unpleasant. This afternoon I took five letters from the of¬
fice and it almost made me cry ! One from my mother and two from
brothers. What a blessed thing our mail system is!
Well, as you would suppose, I am having really a glorious time; all
the flowers I can possibly manage now. Most of the flowers are on
the plains, but I am in the mountains every week, and there by much
fatiguing effort climbing cliffs and descending precipices, by hanging
with my shoe toes in the crevices and my hands to pine and juniper
limbs, I get a good many fine things. It is almost too early for flow¬
ers to be plentiful in the mountains. The higher and more distant
ranges are yet white with snow to their very bases.
These mountains were very appropriately named Rocky. They
seem built of nothing else. On their summits and often on their ter¬
raced sides are large forests of Coniferae. It is in these forests
chiefly that I must look for flowers. I want to get up to the limit of
trees sometime in the summer if possible.
Now in a few days I am going to send you a small package of
flowers. You will find them chiefly leguminous plants and especially
Astragali (milk vetches). Of this genus I have already specimens
of half a dozen species, and I know of as many more that are hardly
yet in flower. They are, unlike our eastern ones, very handsome.
The little low silky one is a mountaineer. A. sincolencus of Gray, or
Phaca sericea of Nuttall grows in pretty spreading masses a foot
broad on bleak and barren summits of high hills or mountains. Prof.
Gray sent me a work of his entitled, “A Review of North American
Species of Astraglus and Oxytropis.” In this work Phaca is reduced
to Astragalus. It seems that although in the time of Linneaus the
genera were both well founded, yet the discovery of very many new
species through which the two run together, has broken down the
wall that separated them. I shall not be able to give you the names
of many of them until I have the fruits, for Gray has made almost
too exclusive use of the fruits in his descriptions and classifications.
One hundred and twelve species in N. A. of Astragulus, that are tol¬
erably well known either by flower or fruit or both. Oxytropis is
less known. On my last week’s tour in the mountains I found two
beautiful ones of which I will send you representations. You ask
me if I am going to stay all the season. I answer I intend to remain
at least two seasons. But it is uncertain. I can probably do better
in the winter at teaching here than I could in the states. It costs
considerable to come here I found and I want to get the good of the
trip by seeing and getting all I can. I want by all means to get as
far south as Santa Fe before I return, so that I can have a good deal
to talk about and a good deal to show you when I get back.
It is true as you perhaps know, that we have some prospects of
Indian difficulties here, but I hope not to be driven home by a war.
Indeed I think I will not. I have seen a great many of the Indians
of the Utah tribe already, and they are said to be at peace and
172 Wisconsin Academy of Sciences , Arts , and Letters .
friendly to the whites, but I do not like their faces, and by no means
would I like to commit myself to their mercies. We hear of miners
and others in remote and unsettled parts, being disturbed and driven
from their work by them. There is a difficulty starting about the
right of the whites to occupy North Park, a section of this territory.
This may end in something serious, but I hope not.
May 13, 1870.
The weather continues very warm and dry. I have just been out a
little way only upon the plains, and have recorded the collection of
specimens of a Nasturtium (water cress), a Senecio (groundsel, rag¬
wort, squaw-weed) and a Chenopodium (goosefoot, pigweed), the
latter in fruit. All new to me. But I saw more beautiful things
than those. I found a true Cereus in flower, C. veridiflorus with
greenish yellow, very fragrant flowers. It is just beginning to bloom.
Also a most beautiful Castilleia (painted cup) not described in any
of my books. It is low, like C. sessiliflora, which also grows here as
in Wisconsin, but is larger or rather stouter, with extremely large
and showy bracts, deeper red than any of C. coccinea I ever saw. It
would be an acquisition to flower gardens well worthy of a conspic¬
uous place. I will secure seeds without fail. I found also what
must be a night flowering Enothera (evening primrose) ; the very
large delicate pink flowers were withered somewhat and collapsed,
stem low and branching, leaves small, a lovely plant. Perhaps you
may raise it from seed if I get any, though it is evidently a biennial
and might not stand a Wisconsin winter.
Well, I do wish you were here to help me, or to get the birds and
mammals, reptiles and insects. If I stay during the winter I will see
what I can do in the line of bird skins for you, though I shall per¬
haps never become an expert in that art. I see a great many hawks
more than I ever noticed elsewhere. There is in the mountains a
very handsome bird called magpie, which I would like you to see. It
is black and white with a very long tail.
A mile from where I board is a republic of prairie dogs. I have
passed through it several times and seen and heard the little fellows,
but no rattlesnakes nor owls. A Pennsylvanian in the neighbor¬
hood, who has an Utah squaw for a wife, with a progeny of twelve
black haired, dark skinned hybrids, has near his house a colony of
forty tame prairie dogs. It consists of the progeny of a tame pair,
with perhaps the original ones also.
No buffalo here, but any quantity of their horns are to be found
on the plains, and also the prodigious large antlers of the elk, still
telling the passer-by what beasts once roamed these wide spreading
plains.
Well, if this letter reaches you some Saturday night and you stay
at home the following Sunday, I presume you will have time to read
most of this, but now I will close. Tell friend Ludwig I have a bottle
of alcohol into which I drop a beetle now and then and when it is full
Main — Life md Letters of Edward Lee Greene. 173
I have thought I would send it to him. But am going to stop and put
no more in until I hear from him.
So you still have a Scandinavian who is a parasite of Cornwall.
It is a pity. He may perhaps get satisfaction yet however as others
have done before him. I am sorry indeed to hear of Rasmus’ and
Karina’s misfortune. (Mr. and Mrs. Rasmus B. Anderson). They
have now an angel to invite them heavenward; one gone before who
stayed with them long enough to have become very dear no doubt.
I have for some time been expecting a letter from Rasmus. Now
adieu! With love to you all I remain as always,
Your sincere friend,
Edward Greene.
Denver, Col. Ter.
Please don’t fail to write again soon. Your letter was very inter¬
esting.
Saturday Evening, June 25, 1870.
My good Friend:
Many thanks for your letter, but it reached me rather late, only
yesterday. The trouble is that in the vicinity of Denver are two
Edward Greens and two E. L. Greens, three in all, who may get my
letters, two of whom have no business with them. Yours had evi¬
dently been opened and carried for a long time in some one’s pocket
and then returned to the P. 0. What shall I do to prevent this an¬
noyance? Surely I cannot invent.
Well, I am very glad to congratulate you on the pleasure you have
enjoyed in meeting with so excellent a gentleman from the old coun¬
try. Had I dreamed of such a thing I suppose I would have sent
three times as many flowers as I did. You of course did the very
best thing, to give him the little things I sent, and I shall almost be
beside myself at the sight of plants from the Dovrefjild.
If I thought I could come home within a year, I would have you
keep them for me. But you can send them in a package I suppose
with perfect safety for my papers and such things are never troubled
by others, and indeed I do not think I have lost any letters, but I
have repeatedly taken from the office letters to Ed. or Mrs. (?) E. L.
Green that evidently were not mine.
So you found Linnaea borealis (twin flower) in the little marsh!
So did I six or eight years ago, but could never find it the second time
though I often sought for it. I have found it abundant in several
localities in the mountains, always at a great altitude. Two or three
times since I wrote I have been away into the mountains and have
seen more than I can begin to tell you. I will get another package
which shall contain some of the fine things. On the 13th inst. I went
some twenty miles up the mountains by the road, but stopped when
I had arrived at the altitude where Populus tremuloides (American
aspen) was just putting forth tender leaves. I was yet twenty miles
from the snowy range. I am told that I need not go there until the
middle of July at the earliest. How strange it still seems to me to
12
174 Wisconsin Academy of Sciences , Arts, and Letters.
look over into the west on these hot midsummer days and see the dis¬
tant mountains streaked or capped with snow. Some of my finest
things now are plants too large to send in a mail package. I will
mention a few of them descriptively. First upon the list is Aquile-
gia (columbine) a mountain plant with very large flowers, blue and
white. It even leaves our cultived ones all in the shade. About four
species of Oenothera (evening primrose) are magnificent. A white
one (accidental) has flowers at least three inches broad with a frag¬
rance that resembles that of Nymphaea odorata (sweet-scented
water lily) but is more delicate. It grows in the mountains only and
flowers in the night. I almost went crazy over it when I found it
one morning before the sun was shining in the valley. One very like
it has golden yellow flowers as large but less fragrant. The corollas
of these beautiful things are very delicate in that one is obliged to
dry them without removing them from the paper until they are per¬
fectly dry. I have splendid specimens of some of them. None of
them is at all coarse looking like our 0. biennis, but are low
or stemless, with flowers purplish or white mostly. I have six
species of Pentstemon (beard tongue), but I don’t know whether you
are acquainted with the genus or not. There were two in central
Illinois, but I believe none in Wisconsin. They are splendid. You
will see from the specimens how Dodecatheon meadia (shooting star)
grows in the Rocky Mountains. I thought it another species when I
first found it growing in dense patches among wet moss on the rocks,
so small and so dark. I find that it varies in size in different locali¬
ties. The dark color of the flowers however is unvarying here.
None pink or white. Nor have I seen it more than half as largl as
the largest Wisconsin ones. I suppose the authorities are correct in
not giving it the specific rank. This little mountain species of Sene-
cio (groundsel) I know will interest you and the mountain Erigeron
(fleabane) too. One day this week I gathered Opuntia (prickly
pear, Indian fig) and a desperate time I had of it. Nuttall’s name,
Cactus ferox, is very appropriate for all the varieties of Opuntia
missouriensis. It is a detestable job for the hands for the articuli
have to be sliced with a knife or they would never dry and the spines
are all sizes and all over the plant. Three species of this genus are
all I find, though 0. rafinesquii and 0. missouriensis have each many
well marked varieties. The little O. fragilis is hateful. One cannot
come near it without the spines poking into one and the fragile joint
breaking off and clinging to one. What think you of Mr. Astragalus
pictus? Isn’t it queer? In fruit it is beautiful! Pads large, inflated
and richly spotted. Its foliage too is so strange for an Astragalus.
I will send you seeds of this and other Astragalus soon. I do not
know how many I have of them now, and I found more in the high
mountains not in flower.
Sunday, June 26, 1870.
I have just got Potentilla effusa (cinquefoil), a beautiful thing
with pinnate leaves white with soft tomentum and with diffuse leaf¬
less stems, very slender. The plains are now in places blue with
Main — Life and Letters of Edward Lee Greene.
175
lupines. I see only two species, L. pusillus and L. polyphyllus. The
latter is tall and very showy.
When I wrote you last I told you I had not seen the little owl that
associates with the prairie dog. I have since seen two pairs of them.
I will try to get a skin or two when I am done with the flowers, that
is in the fall or winter.
I have put in another specimen of Saxifraga nivalis that you may
have one to compare with your Dovrefjild Saxifraga. The other
plants I send are different from those I sent before.
With your letter I received one from Stephen T. Olney of Provi¬
dence, R. I. requesting specimens of Colorado Carices (sedges). That
gentleman is writing a monograph of the N. American species of
Carex. This is not a great country for Carices, however I am send¬
ing him a package of fourteen species that I have gathered up mostly
in the mountains. On the range in their season I shall probably get
a good many more.
Now if you could be here to go with me in two or three more weeks
up to the edges of eternal snowbanks! The distance is not less than
fifty miles and most of the way through the mountains. What a time
we would have! I shall probably go alone, but there is a good wagon
road as far as Georgetown, which is not more than eight miles from
the snow, or rather from the summit of the snowy range. Well, you
may perhaps hear in my next what I get if I am successful in making
the trip.
In looking over the package you will find several things not named,
because my botanical library is incomplete and very many of my
descriptions are by Nuttall, who is not very clear in his diagnoses.
Rubus nutkanus (salmon berry) is a beautiful bush with flowers as
large as roses. I am going to send you and mother seeds of it. Be¬
longing exclusively to the mountains I think it would thrive in Wis¬
consin. Write very soon. If you send my little Norwegian moun¬
taineers, direct just as you have done and I am confident I shall be
able to get them all right: otherwise I would wait until I should see
you, but that may be a long time.
With warm regards and best wishes to you all, I remain as ever,
Your friend,
Ed. L. Greene.
Golden City, Nov. 21, 1870.
My dear Friend:
For the last three or four months I have been thinking, while wait¬
ing to hear from you, that if you did not care enough about hearing
from me to answer my letters, I would not write. However as it has
now been half a year since I received the last word, I have thought it
possible that you may have written and that the letter may have been
lost. Though I am not certain that I have ever yet failed to receive
any letter which has been sent me. The person in Denver who bears
my name, and who has on one or two occasions taken letters which be-
176 Wisconsin Academy of Sciences , Arts, and Letters.
longed to me, I am now very well acquainted with, so that if he should
get them I am quite certain he would not retain them. And what in
the world is Ludwig doing that he does not write.
I have many interesting things to say to you, but I only write this
to let you know that for all these months, I have been awaiting in vain
for a reply to my last.
I remain and ever will, Your sincere friend,
Ed. L. Greene,
Golden City, C. T.
Jarvis Hall, July 22, 1871.
My dear friend Kumlien:
You are the same dilatory correspondent, are you not? Neverthe¬
less I overlook it again and take up my pen to talk with you for a few
moments.
A few days ago I returned from a ten days’ trip in the higher moun¬
tains where you can imagine what a time I had. Now it is to me a
surprising fact that the most beautiful wild flowers I ever saw in my
life, I have seen growing right in the vicinity of eternal snows ! I am
going to enclose a little Primula (primrose) (I know not what species),
of which a large patch was growing less than ten feet from the edge
of a deep snow drift which covered not less than ten acres of ground.
The flower has quite changed its color. It was most exquisite. The
little delicate Saxafraga grew also at the edge of the snow. I got
four other Saxifrages besides this one, all of which are new to me
and those of which I find described in the Flora Lapponica, I think.
I have all the spring been applying myself so closely to divinity stu¬
dies, that until within a few weeks I have quite neglected botany.
During the past week I have received a note from Prof. Gray, and one
from Dr. Engelman, both enquiring what has become of me. Dr. E.
tells me that he has two of my new species of Opuntia growing from
both seed and from joints I have sent. And, by the way, I am re¬
minded that I have not yet sent you O. greenei. I will do so now in a
day or two. There is a fine specimen growing on our college grounds.
I noticed it full of fruit this morning.
I expect a visit in a few days from Wm. M. Canby, a distinguished
botanist from Delaware. At least Mr. Olney tells me he has directed
him to call on me on his way to California. I hope he will come of
course.
On the day before yesterday someone knocked at my door, and as
my roommate opened the door, I instantly recognized the voice of
Charles Clausen asking for “Mr. Green.” I have not been so sur¬
prised in a long time. I was of course delighted. I learned from him
at once that his father was in Denver and that they were both to
start from Denver home again the next day. I returned with Charley
to Denver and tried to persuade the Reverend gentleman to remain
longer, but in vain. I like the Rev. Mr. C. very much though my ac¬
quaintance with him was very short. He tells me that it is quite
Main — Life and Letters of Edward Lee Greene.
177
possible that he will emigrate to this country in the spring. Now just
sell out that little place of yours, get a wagon and team and come
along, and let us enjoy each other’s society among these mountains for
a few years before we die.
Tell friend Ludwig that if he doesn’t write to me pretty soon I shall
give him over entirely. My love to all you and yours. Please write.
Edward L. Greene.
Denver, Colo., Sept. 25, 1871.
My dear Friend:
I never expect to hear from you again, but you shall know that I
think of you. I change my home this week and go to the town of Gree¬
ley in this territory to have charge of the congregation there during
the winter. If you can write, address me there.
As ever,
Love to you all.
Ed. L. G.
Greeley, Col., Easter Even., 1872.
Thure Kumlien Esq.
Busseyville, Wis.
My dear friend:
Certainly you have at last given me a treat indeed. I confess I
have, during the past week, indulged a faint hope that I should hear
from you this spring. I had actually been looking for this letter for
several days. Now it is all right and you may look out by and by for
some flowers. First of all I must tell you that two weeks ago I re¬
ceived a letter from Prof. Gray enclosing his photograph, and a
promise that he will be my guest for a few days next summer Des
volente! Don’t you think I was almost crazy with delight when that
letter came? You know that since I have been in Colorado I have
had many letters from him, and have sent him specimens of all my
collections, and he has named in my honor, at least two of my species.
I am sorry that my alpines are now all gone. Gray has himself
set several other botanists after me, who have begged me out of al¬
most everything, but they give me fine exchanges.
The little one flowered primrose is Primula integrifolia too. The
dear little beauty grew within three feet of the edge of an eternal
snowbank. But oh, if you could see the Primula parryi Gray, I
think you would at least throw up your hat and scream with delight.
It is the most magnificant wild flower I ever saw, and I believe I am
the third who has ever collected it. I preserved only three
specimens, for it is a large and rather succulent plant, and I
was on foot when I found it, and had to collect specimens as
large as I could each, of 125 species that I had never seen
before; consequently, save of the smallest, I only got two or
three specimens each. P. parryi is a foot high with leaves some¬
what like those of Dodecatheon with you, only twice as large and
178 Wisconsin Academy of Sciences, Arts, and Letters.
handsomer green. The stem bears a cluster of flowers each % of an
inch in diameter, of the most beautiful purple or perhaps mauve, and
are fragrant. And then where it grows! In the edges, among the
stones of those crystal mountain streams, in deep fir tree shades, about
a mile or a half mile from snow. The scenery of its locality is fit for
the home of such a wonderful flower. If I get to its locality another
year (and my good church wardens have already given me a permit
to take my vacation during Prof. Gray’s sojourn) I will not come back
without a specimen for you.
It seems to me now while I write that by all means, if possible, you
should come to Colorado and come to stay and collect here the rest of
your life. But you know better than I no doubt. If you could man¬
age to sell out and move, it would be a grand thing I believe. But I
will not suggest any more on this point. It is really singular that you
should have found those orchids as you relate. I have laughed heart¬
ily over your amusing account of it.
Goody era menziesii (rattlesnake plantain) and Corallorrhiza multi¬
flora (coral root) (both orchids) (a large one) I found last summer
growing together in the mountains near Golden City.
I should think your plant is C. odontorhiza Nutt, according to my
last edition of Gray. He says it is “rare northward” that is in Michi¬
gan, etc.
How nice a little affair that Primula scotica is. I am right glad to
see it.
Visiting a parishioner of mine the other day I obtained a very
pretty specimen of Erica vulgaris L., or Calluna vidgaris Salish.
(Neather). It had been sent from England in a letter. I shall be
right happy if you ever get around to send me a package of Norwe¬
gian plants. I guess I told you of my 59 species of Swedish Carices
received from Olney.
And so you fancy you see me with a wife — a mistaken fancy I as¬
sure you. I am not married, and to me it seems now quite probable
that I never shall be. I shall soon be twenty nine years of age, and
am not engaged nor in love. It seems with my preaching and botan¬
izing to have enough to keep me busy and happy. Happy as any mor¬
tal ought to expect to be. I am getting that old bachelor notion that
a wife would only be in the way. So I guess I am bound to know the
whole of single blessedness. I at least do not ask to be happier in the
world than I have been since I was ordained a minister. I rejoice
that we have spring. I saw the first wild flowers, Berberis aquifolium
(barberry) three weeks ago. Where is Ludwig that I never hear
from him? I think he is a remarkable fellow indeed, to let me go two
whole years without one word. Give my love to your dear ones, and
remember me ever as your friend.
Edward.
Main — Life and Letters of Edward Lee Greene.
179
Greeley, Col., Sept. 26, 1872.
My dear Friend:
Your letter of Aug. 30th was duly received, and I am much obliged
to you for being so prompt.
Yes, I had the honor of meeting the great Prof. Gray; and certainly
he is one of the most delightful men I ever saw. I went to the moun¬
tains ninety miles away to Empire City near Gray’s Peak, a week be¬
fore he came that I might not fail to meet him. However, he stopped
in Greeley, or rather just got off the train and enquired for me. He
thought he might meet me in Denver, where he found a letter from
me telling him where to meet me. I met him at Empire City on Sat¬
urday evening Aug. 10th, just at dark. Was presented to him by the
celebrated Dr. Parry, of fame as a collector in Mexico, California and
Colorado. Dr. Parry had been here some months and is yet in the
mountains collecting seeds. I expect a visit from him in about two
weeks. Well, on Monday the 12th Dr. Gray, Dr. Parry, and myself,
with some other unscientific gentlemen, made the trip to the top of
Parry’s Peak, and made our first tramp together, collecting no end of
fine things; were caught for some twenty minutes in a fearful snow¬
storm, accompanied with lightning that made our hair snap, and
stand on end with electricity at every flash. The snow covered up all
our botanizing for about half an hour, and then it disappeared al¬
most momentarily, when the sun came out. The storm was very un¬
pleasant, for we were far above the limit of trees, and had no protec¬
tion.
Wednesday and Thursday were occupied in making the ascent and
descent of Gray’s Peak, the highest mountain this side of the Parks.
That was a tiresome job, and yet well worth the trouble and fatigue.
For the last two or three miles toward the top no vegetation what¬
ever, but a few rock growing lichens met our eye, and it was
nothing but climb over sharp and often coarse rocks. And you may
understand that after twelve or thirteen thousand feet altitude is
reached, it requires some muscular exertion to get breath. Mt. Gray
is 14,245 feet high, and the view from the top is probably one of the
most magnificent which this world affords. I can never describe it
and will not try. Prof. Gray stayed in Empire and thereabouts just
one week only, and then we bade him farewell. His wife, a very
amiable lady, was with us, even to the top of the Peak. She is a
botanist too.
Dr. Parry is one of those quiet, diffident men who know a great deal
more than one would think. Gray regards him as a great botanist
though he has never published anything.
Well, I have seen more than Prof. Gray, for last week I was down
to Denver for a few days; at the depot one morning I noticed on the
platform waiting for the train a very aged gentleman, whom I took
to be a botanist when I saw among his hand baggage an unmistakable
bundle of specimens in press. I stepped up and begged his pardon for
asking his name. Judge of my delight when he replied, “I am called
180 Wisconsin Academy of Sciences , Arts , and Letters .
Dr. Torrey!” Yes, sure enough here I stood in the presence of and
shaking hands with the venerable teacher, and associate in labors of
Prof. Gray. He was on his way to the mountains to visit Gray’s Peak.
I was not able to go with him though he urged me. I had never cor¬
responded with him but he knew me and we were as familiar friends
from the moment I gave him my name. Car ex torrey i (sedge discov¬
ered by Dr. Torrey) had been one of the finest of my finds in Colo¬
rado, and I was the second who had found it in the U. S., Dr. Torrey
having been the first. He found it in Pennysyvania in a few speci¬
mens many, many years ago where nobody else has since been able
to detect it. I find and collect a hundred specimens in Colorado.
Well, I rode with Dr. Torrey to the end of the railway, seventeen
miles that morning, and have not seen him since, and it is not likely
that I ever shall again. He is some seventy-five years old, though in
apparent good health of body and vigor of mind. I missed him on
his way back, though I watched for him. But I am very glad to have
seen him and conversed with him for an hour. He of course will not
long be numbered with the living botanists, and in my old age, if I
live, I shall be glad to be able to say that I saw him.
Gray is sixty two, though he seems fifteen years younger. A hand¬
some man and one of the most delightfully pleasant men you ever saw.
Always smiling and always talking, and his speech affected by an
occasional slight stammering. It was very, very interesting to see
him pick up something which he himself had first described and
named, and ask, “Parry” or “Green,” “What is this” and then after
a closer examination, “Oh, it is _ _ _ _ isn't it? Oh, yes,
I never saw it growing before.” Yes, I have certainly had a rich de¬
light in seeing these three great botanists, and now I wish I could
see you and tell you more than I have time to write. Please let me
hear from you again. Have I sent you a specimen of Museniam
Greenei, an umbellifer discovered by me last year? My love to you
and your family,
Ever,
E. L. Greene.
Vallejo, Cal., April 30, 1874.
My dear Friend:
Your long and interesting letter is received. I am not going to
answer it just now, but I send you herewith the last four numbers
of the Naturalist, knowing that it may be a gratification to you to
read some of their contents. One of them contains the notice of Lud¬
wig’s bird. There are also a number of articles on botany — an un¬
usual number. Those from Prof. Farlow on the visits to European
historical fields will carry you back to your old home — at least In
fancy. The papers on Wyoming botany, by my dear friend Parry,
and those from my own pen, will also I trust interest you.
When you and Ludwig have read the magazines you can return
them. I have no recollection at all of having sent you any Apple-
Main — Life and Letters of Edward Lee Greene.
181
ton’s Journals, nor do I remember to have read what you refer to.
Some one else I think, has done you that favor. With love to all, I
am as ever,
Your affectionate friend,
Edward Lee Greene.
(This letter is to Ludwig Kumlien, son of his friend and teacher,
Thure Kumlien.)
Vallejo, Cal., Sept. 3, 1874.
My dear Ludwig:
It almost seems that I have an old friend risen from the dead, this
getting a letter from you after I guess almost five years of silence.
Well, I need not waste time and paper to assure you of how glad I
am to hear from you, and also to have a photograph of your face.
You have grown big and devised a little beard, and that is all the dif¬
ference I am able to detect in your looks, comparing of course the
photograph with the picture that is in my memory. I am greatly
obliged for so long a list of news. I have not heard so much about
Albion and vicinity I think, since I left there. Now that all our fam¬
ily are away I have absolutely no correspondence with any resident in
that section, except the very occasional letters I get from Monsieur,
your father.
I hope now you will not fail to do better by me, now that you have
succeeded so well in this lately renewed effort.
What a time I have had since I saw you, ranging over the won¬
derful plains of Wyoming and northern Colorado, and the cactus
deserts, even away down to the borders of New Mexico; and climb¬
ing about the perpetual snows of the Rocky Mountains, up to the alti¬
tude of 14,245 feet; and now at last have been six months on the
Pacific coast. I have naturally added somewhat to my knowledge of
botany and have a splendid collection of western plants; and have
been so fortunate as to discover a few new species, and have redis¬
covered a number of long lost ones, that had been found only by such
early botanists as Nuttall, James, etc. I have had a fine time, also
have worked my way into the pulpit so as to have no trouble about
the wherewithal to pay my expenses. But my new parish at Vallejo
is too much for me. I have a large congregation and good salary,
but with all that, so much pastorial work, that my scientific studies
are interfered with not a little. I have also the prospects of being
even called to a large San Francisco church on three thousand dol¬
lars a year, to which if I go, I fear I shall turn out either a poor
shepherd for so large a flock or else give up more still of my favorite
studies. However, I am thankful for so much happiness and success
as seems to attend me in several ways.
My people in Colorado are doing well indeed. I wish you could see
our Rocky Mountain home. It would delight you. I am heartily
glad to hear of your father’s prospects of becoming able to devote
himself to his favorite pursuits, with a fair remuneration. I hope
after a few years more, fewer than have elapsed since we met, that
182 Wisconsin Academy of Sciences , Arts, and Letters.
we shall all meet again. I long now to see old Wisconsin and bota¬
nize again along Lake Koshkonong and thereabouts. It would be as
good as a new field to rne, almost, not to mention the bliss of the com¬
panionship of a few old friends I could name. Write very soon.
Give my love to all your people and believe me as ever yours,
E. L. G.
I will send that stick before long.
(This letter also is for the son Ludwig.)
Georgetown, Colo., Nov. 22, 1875.
My dear Ludwig:
I shall no doubt surprise you again by writing still from George¬
town, Colorado. But you may learn after awhile never more to be
surprised at any of my unaccountable doings. On the very eve of
my departure for California, I came to a disagreement with the
Bishop of Northern California, to whose jurisdiction I was going, and
the result was, I said to him, “I will not go.” So here I am.
How did it come to pass that you heard I was to visit Wisconsin
this fall? Perhaps the wife of my brother Manzel who visited Al¬
bion, spoke of something of the kind for I did in the summer encour¬
age my brother who lives in Minnesota to think I would visit him be¬
fore ever I should return to the far Pacific coast. But there will, as
it appears, be time for that yet. However, I must confess since you
wrote me last I have been, for a few weeks, comparativly near you.
I was in central Illinois for four weeks and now have just returned.
When I determined not to go to California, I said to myself, “I will
take a trip somewhere anyway.” I wanted to see Dr. Engelman in
St. Louis, and my relatives in central Illinois, besides many other
friends that I made there while teaching in 1867, ’68 and ’69. You
will ask me I know, why I did not come on to Wisconsin. The rea¬
sons are several. For one important reason I give this, I did not feel
like seeing Albion, my old home, etc., so soon after that deplorable
event of my mother’s death. I should be sad all the time I knew if I
went. For another reason, I could not wish to go there without
several weeks to devote to calling on the different people whom I
should wish to see. Moreover if I should have gone to Wisconsin at
all there could have been no sort of an excuse for my not going on to
visit my brother in southern Minnesota. I had not at my disposal
the two or three months I should have needed for visiting all three
of the vicinities mentioned, so I determined to wait and visit Wis¬
consin and Minnesota perhaps another year.
I am very glad to hear of you at the University. Give my love to
Prof. Anderson and tell him the last book I read was his “Norse
Mythology” with which I was greatly pleased. I have no letter from
your father for now these many months. I trust he is well. How
much I would love to see you all.
I am wintering this year in a very interesting, if not pleasant
place. Georgetown is built in a little narrow valley at 8,400 ft. above
Main — Life and Letters of Edward Lee Greene. 183
the sea. The mountains on the east side of town rise very abruptly
2,300 ft. higher, and on the west 3,000 ft. higher, while to the south
Leavenworth Mountain summit is about 1,500 ft. above the town.
The sun rises at this season at half past nine, passes behind Leaven¬
worth Mountain at 12; reappears a little after one, and disappears
for the day shortly after two. We now have about a foot of snow
on the ground, but it is not extremely cold. In fact the thermometer
rarely falls below zero at Georgetown at any time during the winter.
It is what many would call a most dreary place in which to spend a
winter, but I doubt not I shall pass the days very pleasantly with my
books and my herbarium, and a great many specimens of things new
to me, to be studied and put away in their place. I have more than
three thousand specimens of foreign plants on hand now which I
have never looked at, and many from the new Northwest. I ought to
be very happy. I wish you could visit me in Colorado before long.
Can’t you do it? At least write to me often and believe me,
Your faithful friend,
Edward L. Greene.
Berkley, Cal., Jan. 9, 1885.
My dear Friend:
I did not wait for your letter. The printer was in a hurry, so I
gave in what you will see by the proof sheets enclosed. You will also
see that the plant has been handled by Asa Gray. But it was a
handling loosely and carelessly, just as he has handled many a score
of western genera and species which he knew naught about, saw in
poor specimens. You will perceive by my written character that,
by both flower and fruit, it is wide of Ranunculus. I am sorry to
say it is a rarity. I never saw it growing, but I hope I shall some
day. We have in the Herb. Cal. Acad, three specimens only! I
think I will give you one of these. For the rest you must wait until
I or some friend of mine shall collect more. Its nearest locality is
more than two hundred miles from where I am. I trust the bulletin
containing this and many more of my doings will be ready for dis¬
tribution soon and a complete one shall be sent you. I have not the
time to write more tonight.
Yours as always,
Edw. L. Greene.
(The following letter was written to O. A. Linder, Chicago, Ill., who
was preparing a paper on the “Life of Thure Kumlien.”)
National Museum, Washington, D. C., Nov. 7, 1908.
My dear Sir:
I take pleasure in enclosing to you, as a loan, the last letter I ever
had from Thure Kumlien. It is perfectly characteristic from begin¬
ning to end; shows the perfection of his beautiful handwriting at the
age of sixty-seven, and the autograph at the end, just as he always
wrote it.
184 Wisconsin Academy of Sciences, Arts, and Letters.
I think you will see from this letter that, with Kumlien, I was not
a companion in age, but rather, a boy of whom he was very fond and
helped many years before this letter. The fact is he and my father
were born in the autumn of the same year, 1819. Very curiously,
also, Kumlien’s oldest child (early deceased) was born on the same
day that I was. I am now sixty-five, but then if Kumlien and my
father had now been alive, each would have been in his ninetieth
year.
His opening of this fine letter in the use of an adage in Swedish
explains itself in this wise. At fifteen years of age I spoke Norweg¬
ian and English with equal readiness, on all matters appeartaining
to work and business. To Norwegians I was a marvel on this ac¬
count.
At Kumlien’s house, when once some of Mrs. Kumlien’s relatives
newly from Sweden were guests, and knew no English, I talked with
them in Norwegian, they answering in Swedish; and Mr. Kumlien
took a new pride in me from the fact that certain Swedish words, dif¬
fering considerably from their Norwegian equivalents, I had no diffi¬
culty in understanding, while they often failed to get the meaning of
certain Norwegian words of mine, though my pronunciation was al¬
ways said to be so faultless.
You must not locate Mr. K. as one of any Scandinavian colony.
His house was four or five miles away from Koshkonong Prairie,
and in the woods near, and in view of Koshkonong Lake. No Scan¬
dinavians were anywhere near him. When in my boyhood, I first
knew him, all his neighborhood was occupied by Yorkshire English,
small farmers. All his older children were baptized by Anglican,
that is, Protestant-Episcopalian ministers. Only the youngest
Frithiof, was christened by a Norwegian Lutheran minister.
I am sure you will be pleased with this piece of his handwriting,
and kindly be very careful not to lose it, and return it when it has
served your purpose.
Very truly yours,
Edward L. Greene.
In the letter from Berkeley, California, dated January 9, 1885, Mr.
Greene, who was then Professor of Botany at the University of Cali¬
fornia, wrote his old friend Mr. Kumlien that he had just named in
his honor Kumlienia hystricula. In his paper describing this flower
in the California Academy of Sciences, Professor Greene says, “I
gladly dedicate this very characteristic plant of our Sierras to Prof.
Thure Ludwig Kumlien, A. M., formerly Professor of Natural His¬
tory at Albion, Wisconsin, a learned and zealous naturalist, and my
first instructor in the science of botany.”
After the death of Mr. Kumlien in 1888, Mr. Greene wrote and
published in his Pittonia, a beautiful eulogy in honor of his friend.
In closing he says, “but these small tokens, like our worded tributes,
are all inadequate to speak the praise, or worthily perpetuate the
memory of a man so pure, so simple, so noble and so well beloved.”
Main — Life and Letters of Edward Lee Greene. 185
When Edward Lee Greene died on November 10, 1914, the world
lost not only a valued botanist, but also a man of sterling quaiities
of character, a man who set high value on friendship; he was a man
who loved the out of doors, not only the well trodden paths, but the
mountain peaks and the cactus deserts, and a man who loved truth
for its own sake.
Fort Atkinson, Wisconsin.
MICROCHEMICAL TESTS ON THE CELL WALLS OF
CERTAIN FUNGI. CELLULOSE AND CHITIN
E. W. Hopkins
The similarity of the life histories of the lower fungi and
many of the lower algae would seem to warrant the expec¬
tation that the composition of their cell walls would have
something in common. The analogies of structure and
function observed in the Phycomyeetes and the green algae
indicate what is apparently a direct relationship between
these forms. The Ascomycetes occupy a more doubtful
position, appearing to be related to the red algae, or to the
green algae through the intermediary of the Phycomyeetes.
The Basidiomycetes, however, show little or no relationship
to either the lower fungi or to the algae. It would be ex¬
pected that the Basidiomycetes would show a greater devia¬
tion from the algae in composition than would any other
class of fungi.
Earlier workers believed that the cell walls of fungi were
made up of a modification of cellulose which was called
“fungocellulose.” Richter (1881) made cellulose tests on
Agaricus campestris, Daedalea quercina, Polyporus f omen-
tar ius, and Mucors. Chlor-zinc iodide and iodine-sulfuric
acid tests were made on sections which had been previously
treated with alkali. Positive cellulose reactions were ob¬
tained with all except the Mucors, which gave no certain
result. The author concluded that fungocellulose was true
cellulose.
De Bary (1887) was of a different opinion. The so-
called “fungo-cellulose” differed from true cellulose in that
it was insoluble in Schweitzer’s reagent, and did not give
the color with iodine which is characteristic of cellulose.
True cellulose reactions, however, were given by the Sapro-
legnieae, Protomyces macrosporus, Peronosporeae, young
Mucors, and the cells of the resting perithecium of Penicil-
lium glaucum. Similar results are reported by Winter-
stein (1893) who removed fats and proteins from fungus
188 Wisconsin Academy of Sciences , Arts , and Letters .
material, and found that the residue was insoluble in
Schweitzer’s reagent. He concluded that it was a cellulose
differing from that in tissues of higher plants. Gilson
(1893) was unable to obtain crystalline cellulose from Mu-
cor vulgaris , Thamnidium vulgare, and Agaricus campes-
tris, while he succeeded easily with plant tissue. He be¬
lieved that fungus tissue did not contain cellulose.
Perhaps the greatest variety of materials wTas tested by
van Wisselingh (1898). This author reports the presence
of chitin in Myxomycetes, Peronosporales, Saprolegniales,
Chitridiales, Entomophtharales, Mucolares, and in almost
all of the higher fungi. Reactions for both chitin and cel¬
lulose were obtained with Myxomycetes and Phycomycetes,
although the reactions were given by different portions of
the hyphae. Chitin seemed to be confined to certain por¬
tions rather than to the entire cell wall. Wester (1909)
prepared chitosan salts from Mucor muceda , Xylaria hy-
poxylon, Peziza aurantia , and Xylaria polymorpha. Chitin
is reported by Vouk (1915) as being present in the follow¬
ing fungi : a Mucor, Helvella crispa, Peziza aurantia, Xy¬
laria polymorpha, Plicaria cervina, Agaricus fusipes, Ama-
nitopsis plumb ea, Boletus sanguinus, a Clitocybe, Cortinar-
ius obtusus, Hygrophorus conicus, Mutinus caninus , Psalli-
ota campestris, and Russula aeruginosa . Extensive stud¬
ies of chitosan salts were made by Brunswik (1921)
These salts were prepared from Lepiota procera, Pholiota
squarrosa, Lycoperdon caulatum, and an Aspergillus. These
preparations were identical with chitosan salts of animal
origin. Gwynne-Vaughan and Barnes (1927) made the
statement that the cell wTall of fungi is usually of cellulose,
or of a special variety known as fungal cellulose.
Thomas (1928) using several species of Fusaria, ex¬
tracted the proteins from the hyphae, and made an ammo-
niacal copper sulfate extraction of the residue. This re¬
agent dissolved a material which appeared to be cellulose,
giving cellulose color reactions, and hydrolyzing to reducing
sugars. The hyphae were still intact, and gave tests for
chitin.
The variations in results given by the above investigat¬
ors may be attributed in part to the different methods used.
One heated the material with glycerine to remove the cell
Hopkins — Microchemical Tests on Cell Walls of Fungi. 189
contents, while another let the sections stand in concen¬
trated alkali to remove the protein constituents of the cells.
Various adaptations of the tests for chitin and cellulose
were used. With the many variable factors concerned, it
is surprising that these workers did not report even more
conflicting results.
The methods used in our work were those recommended
by the most widely accepted authorities on micro-technique.
Three tests for cellulose were employed, and one test for
chitin.
Cellulose Tests .
1. Iodine-potassium iodide-sulfuric acid test. The sec¬
tions were saturated with the iodine-potassium iodide re¬
agent, the slide tilted, ana the excess solution removed by
placing a blotter at the lower edge of the drop. The blotter
did not come in contact with the section, and any cellulose
fibers adhering to the slide could not been confused with
material in the section because of their difference in appear¬
ance. Cellulose gives a very deep blue color when treated
with these reagents. Molisch (1923) .
2. Iodine-potassium iodide-phosphoric acid test. This
test was performed in the same way as the one above, ex¬
cept that it was necessary to heat the sections since the acid
used was weaker than that in the first test, and conse¬
quently did not penetrate the cell wall as readily. Cellu¬
lose gives a deep violet with this treatment. Zimmermann
(1901).
3. Chlor-zinc iodide test. This is one of the standard
tests for cellulose, giving a dark violet-blue color with that
compound. Molisch (1923).
Chitin Test.
Van Wisselingh (1898) sealed sections of material in
glass tubes with concentrated potassium hydroxide solu¬
tion. These sealed tubes were then heated in an oil bath to
180° C., cooled, the tubes broken, and the sections removed.
The alkali was removed by washing with 90 per cent alco¬
hol. After washing the section thoroughly with alcohol, it
was placed on a slide, saturated with an iodine-potassium
iodide reagent the excess of this solution removed as before
18
190 Wisconsin Academy of Sciences , Arts, and Letters.
by blotting, and dilute sulfuric acid added. Chitin gives a
violet color when treated in this way.
A much simpler method is given in detail by Vouk
(1915) and recommended by Eckerson. This method con¬
sists in boiling the material for 20-30 minutes in a satu¬
rated aqueous solution of potassium hydroxide in a beaker
and washing the material with 90 per cent alcohol. The
remaining steps are not explicitly stated, but since this
method was based on that of van Wisselingh, it is very
probable that the remainder of the procedure was left un¬
changed, i. e., the sections were treated with iodine-potas¬
sium iodide reagent and sulfuric acid. Vouk first states
that his preparations give the chitosan reaction “nach be-
kannten Weise mit Jodjodkalium.,, This statement does
not exclude the use of sulfuric acid from the test. Further,
this author states, “Nach dem Auswaschen in 90 p. ct. alko-
hol trat die Reaktion mit Jodjodkalium immer prompt ein.”
It seems doubtful that the use of sulfuric acid is indicated
by this statement. Miss Eckerson, in adapting the method,
does not state that sulfuric acid should be used. When the
test as described by this author is applied to flies’ bodies,
no violet color appears, but it does appear when sulfuric
acid is added. Other sources give tests for chitosan with
iodine-potassium iodide reagents, and no sulfuric acid, but
these tests all gave negative results when applied to parts
of flies’ bodies. The test for chitin was made by the pro¬
cedure recommended by Vouk and Eckerson except that
sulfuric acid was used to convert the chitosan to a material
which gave a violet color with iodine-potassium iodide.
The number of materials to be tested for chitin made it
impossible to boil them all together with alkali in a beaker.
In order to keep them separated, and to easily distinguish
the different materials tested, each was placed in an agglu¬
tination tube. These tubes were inserted for about half
f-heir length through the meshes ol a screen of the proper
size to hold them firmly. This screen, with the tubes held
firmly in place, was then suspended in an oil bath. The
samples in the tubes were covered with saturated KOH, the
oil bath heated to 130° C, and this temperature was main¬
tained for 25 minutes. The sections were then fished out
Hopkins— -Microchemical Tests on Cell Walls of Fungi. 191
of the tubes, washed with 90 per cent alcohol, placed on a
slide, and iodine-potassium iodide and sulfuric acid added.
Four cultures grown in the laboratory were tested daily,
and tests were also made on material collected at random.
The laboratory cultures were grown in Petri dishes as gi¬
ant colonies on malt agar (3 per cent malt extract, 1.5 per
cent agar.) Sections of the colony were removed daily,
and the duplicate tests made upon it. The agar did not
interfere materially with the tests, and in no case gave a
color which would be mistaken for a positive test. The cul¬
tures grown on agar, and tested systematically were: Mu-
cor rouxii , Aspergillus fumigatus, Thraustotheca clavata,
and Achlya imperfecta.
In table 1 are given the results of the tests on Mucor
rouxii . In the first 3 tests when the culture was 2, 4, and
5 days old respectively, all the tests were negative, although
a pale violet color was given with chlor-zinc iodide, and on
the last day a questionable chitin test was obtained. When
the culture was 7 days old, colors which were very similar
to these given by cellulose appeared with all of the cellulose
test reagents, and a positive chitin test was obtained. On
the 8th day, so much of the material had been removed
from the first Petri dish that it was necessary to take ma¬
terial from another plate. This gave negative cellulose
tests and a positive chitin test. Though both were started
at the same time, their development may be very different,
due to the vigor of the culture, the size of the inoculum, or
a lack of uniformity of favorable conditions. The lack of
concord between the results of different workers using the
same fungus may well be due to such factors. On the 9th
day, the chitin test was negative, while the cellulose tests
were questionable, a color being obtained which was similar
to that given by cellulose. On the 10th day, the cellulose
tests, except the chlor-zinc iodide test, were positive, and
the chitin test was negative. An old culture of Mucor
rouxii which happened to be available was also tested.
The cellulose tests except the chlor-zinc iodide test were
positive, and the chitin test positive.
Aspergillus fumigatus gave somewhat different results.
(Table 2.) At no time was a positive cellulose test ob¬
tained, although a pale violet color was observed with chlor-
192 Wisconsin Academy of Sciences, Arts, and Letters.
zinc iodide from the 7th to the 10th day. The chitin test
was questionable on the 4th and 5th day, but positive on
the 7th and 8th day, negative on the 9th, and positive again
on the 10th. On the 12th day, all tests were negative.
Thraustotheca clavata (table 3) and Achlya imperfecta
(table 4) at all times gave identical tests. The cellulose
tests were all strongly positive throughout, the color be¬
coming very intense as the cultures increased in age. The
chitin tests were all negative, although in the older cultures,
a blue coloration appeared which, however, was quite dif¬
ferent from the color given by chitin.
The results of the tests on the higher fungi are recorded
in table 5. No form gave a positive cellulose test. Chitin
tests were given by some, and not by others. In the cases
where the chitin test was most positive, the violet colora¬
tion seemed to be confined to the cuticle, never extending
far into the tissues.
Perhaps the greatest discrepancy occurs in table 1 in the
cellulose tests. It is probable that the second culture used
had not developed as far as the first one. In the chitin
tests given in tables 1 and 2, there is a most discouraging
lack of consistency. This may be due to several factors.
The test in itself is not easily made. The sections used,
though as small as possible, may nevertheless have been too
large to enable the reagents to thoroughly penetrate the
cell walls. The hyphae were always very much matted
after boiling in alkali, and it was not possible to separate
them. This made the test very difficult to observe, and
may account for the lack of consistency of the results. The
other two cultures used, Thraustotheca clavata and Achlya
imperfecta, had such large and rigid hyphae that the tests
were very easily observed.
In all of the tests made there is always a question re¬
garding interfering substances. The cellulose tests were
made in such a way that other substances giving a blue
color with iodine would not be mistaken for cellulose. This
was accomplished by saturating the section with iodine-
potassium iodide reagent, and examining under the micro¬
scope before sulfuric acid was added. No blue color was
observed in any case before addition of sulfuric acid.
Hopkins — Microchemical Tests on Cell Walls of Fungi. 193
Summary
Twenty-two species of fungi distributed through the
Oomycetes, Zygomycetes, and Basidiomycetes, were tested
for chitin and cellulose. The chitin test used was that of
Vouk. Three cellulose tests were made : iodine-potas¬
sium iodide-sulfuric acid test, iodine-potassium iodide-
phosphoric acid test, and the chlor-zinc iodide test.
Four fungi were grown on malt agar and tested daily.
These were Thraustotheca clavata , Achlya imperfecta , Mu-
cor rouxii, and Aspergillus fumigatus. The remaining
fungi were tested as they were collected.
The lower fungi, Saprolegniales, gave strong tests for
cellulose, one of the Mucorales gave tests for both cellulose
and chitin at different ages, and the higher fungi gave neg¬
ative tests for cellulose, though several gave chitin reac¬
tions.
It seems very likely that if these tests were made on a
greater variety of forms, or systematic tests of cultured
forms extended over a greater period of time, the results
would permit more definite conclusions than are possible
from this work.
The writer wishes to express his sincere gratitude to Dr.
E. M. Gilbert of the Department of Botany under whose
direction the work was done, and to Dr. E. B. Fred of the
Department of Agricultural Bacteriology for helpful criti¬
cisms.
University of Wisconsin
Madison, Wisconsin.
Bibliography
de Bary, A. 1887. Comparative morphology and biology
of the fungi, mycetozoa, and bacteria. London. 8.
Brunswik, H. 1921. Ueber die Mikrochemie der Chito-
sanverbindungen. Biochem. Zeit. 113: 111-124.
Eckerson, S. Microchemical methods. Unpublished notes.
Gilson, E. 1893. La cristallization de la cellulose et la
composition chimique de la membrane cellulaire veg-
etale. La Cellule 9: 397-441.
194 Wisconsin Academy of Sciences, Arts, and Letters.
G wynne- Vaughan, H. C. I., and Barnes, B. 1927. The
structure and development of the fungi. New York,
Cambridge.
Molisch, H. 1923. Mikrochemie der Pflanze. 3rd aufl.
Jena. 336.
Richter, C. 1881. Beitrage zur genaueren Kenntniss der
chemischen Beschaffenheit der Zellmembranen bei den
Pilzen. Sitzungber. d. Kais. akad. d. Wiss. in Wien.
83 : 494-510.
Thomas, R. C. 1928. Composition of fungus hyphae. I.
The fusaria. Am. J. of Bot. 15 : 537-547.
Vouk, V. 1915. Zur Kenntnis der mikrochemischen Chi-
tin-Reaktion. Ber. d. Deutsch. Bot. Ges. 33: 413-
415.
Wester, D. H. 1909. Studien liber das Chitin. Arch. d.
Pharm. 247 : 282-307.
Winterstein, E. 1893. Zur Kenntniss der Pilzcellulose.
Ber. d. Deutsch Bot. Ges. 11: 441-445 .
van Wisselingh, C. 1898. Mikrochemische Untersuchun-
gen liber die Zellwande der Fungi. Jahrb. f. wiss.
Bot. 31 : 619-687.
Zimmerman, A. 1901. Botanical microtechnique.
Translated by J. E. Humphrey. New York. 141.
Table 1. Cellulose and chitin tests on Mucor rouxii.
Hopkins — Microchemical Tests on Cell Walls of Fungi. 195
Table 2. Cellulose and chitin tests on Aspergillus fumigatus.
Table 3. Cellulose and chitin tests on Thraustotheca clavata.
Table 4. Cellulose and chitin tests on Achlya imperfecta.
196 Wisconsin Academy of Sciences , Arts, and Letters
Table 5. Cellulose and chitin tests on higher fungi .
THE HEPATICAE OF WISCONSIN
George Hall Conklin
Introduction
This report of the Hepaticae of Wisconsin is based upon
a re-examination of the specimens in the herbaria of the
University of Wisconsin and the New York Botanical Gar¬
den, and a study of the specimens collected in Wisconsin.
To this has been added the collections of the writer made
during the past 20 years in Douglas, Bayfield, Ashland, and
Iron counties. This report also includes an examination of
the collections of that indefatigable collector, L. S. Cheney
of Barron, Wis., who in the years from 1893 to 1897 made
a botanical survey along the upper Wisconsin River valley
from State Line, Vilas Co., to Kilbourn, Sauk Co., also
along the Montreal River, the White River, and the south
shore of Lake Superior from Montreal River, Iron Co., to
Superior, Douglas Co.
Only a few of the specimen numbers of the Cheney col¬
lections of 1893 were placed in the University Herbarium.
The work of identification was done by Dr. L. M. Under¬
wood. After Dr. Underwood's death his herbarium and
unidentified collections were purchased by the New York
Botanical Garden. In 1911 Miss C. C. Haynes named the
material, and a duplicate set was given to the University of
Wisconsin. An opportunity was given to verify this ma¬
terial. In 1926 Mr. Cheney gave to the botanical depart¬
ment of the University of Wisconsin the later collections
and remaining duplicates. These have been named and in¬
cluded in this report and returned to the University Herb¬
arium. Of more recent collections this report also includes
the following stations and collectors :
198 Wisconsin Academy of Sciences, Arts, and Letters .
Dr. C. E. Allen Devils Lake, Sauk Co. _ 1915, 1916, 1917
Solon Springs, Douglas Co. _ 1915.
Arena, Iowa Co. _ 1922,
Glen Haven, Grant Co. _ 1924.
Sawyer Co. _ 1926.
Green Lake, Green Lake Co. _ 1926.
Lake Wapogasset and Bear Trap
Lake, Polk Co. _ 1927.
Daisy S. Howe Osceola, Polk Co. _ 1901.
C. D. Baker St. Croix Falls, Polk Co. _ 1897.
N. C. Fassett and L. R. Wilson St. Croix Falls, Polk Co., and Mel-
len, Ashland Co., 1927.
J. M. Holzinger Along Mississippi and St. Croix Rivers, Polk, St.
Croix, Buffalo, and Trempealeau Counties, 1890 to 1906.
An examination has been made of all the known and avail¬
able material collected in the state, both named and un¬
named. It was thought that such a critical study of the old
herbarium material, together with the unreported collec¬
tions of more recent dates would be of value to future stu¬
dents in the study of plant distribution. Early virgin con¬
ditions of forests, waters, and surface soils are fast chang¬
ing in many counties of the state. Owing to the denuda¬
tion of forests, the drying up of lakes and swamps, and the
drying and erosion of waterways, the bryophytes have been
especially effected. In many places they are fast dying
out. The species here recorded of early dates have in
many instances wholly disappeared from the stations re¬
corded.
History
(1) The earliest known records of Hepaticae collected
in Wisconsin are found in the University of Wisconsin Her¬
barium. These are a few specimens from the herbarium of
I. A. Lapham, with the following data:
Reboulia hemisphaerica (L.) Raddi (labeled Fimbria -
ria tenella). Dells of Wisconsin River (Sauk Co.), Aug.,
1858.
Bazzania trilobata (L.) S. F. Gray (labeled Mastigobry-
um trilob atum) . Penokee (Ashland Co.), Sept., 1858.
Porella platyphylla (L.) Lindb. (labeled Madotheca
platyphylla) . Penokee Iron Range (Ashland Co.). No
date given.
Conklin — The Hepaticae of Wisconsin.
199
Porella platyphylla (L.) Lindb. Blue Mounds (Iowa
Co.), June 1, 1860.
Ricca fluitans L. (no. 43), ex. coll. I. J. Hale, 1861.
Several other specimens with dates unrecorded are from
the herbarium of I. A. Lapham and were probably collected
about this decade.
(2) There is one specimen of Ptilidium pulcherrimum
(listed as P. ciliare, no collector given), from Milwaukee,
Milwaukee Co., with the date of 1862.
(3) J. M. Holzinger collected Anthroceros laevis L,
August 18, 1890, at Marshland, Buffalo Co.
(4) In 1891 and 1892 L. S. Cheney and R. H. True col¬
lected in Dane Co., about Madison; in Fayette, Lafayette
Co., and Lake Mills, Jefferson Co. It was during these
years that Professor Cheney, at that time connected with
the University botanical department, made his first trip to
the north, collecting from Fond du Lac to Grand Portage,
Minn., along the north shore of Lake Superior. His trip
extended along the International Border from Grand Port¬
age to Ely, Minn., ending the trip at Barron, Wis. The
collected hepatics were reported in the Transactions of the
Wisconsin Academy (2). Most of the region covered in
this botanical survey was in Minnesota. A few collections
however, were made near Fond du Lac, Minn., on the Wis¬
consin side of the St. Louis River in Douglas Co., and when
the trip ended, some collecting was done about Barron, Wis.
This was in 1891, and 15 species from Wisconsin were re¬
corded and correctly named.
(5) In 1893 the American Association for the Advance¬
ment of Science met at Madison, and a collecting trip was
made to the Dells of the Wisconsin River. A few species
were found and bear the label of L. M. Underwood, collec¬
tor.
(6) Professor Cheney, beginning in 1893, began a sur¬
vey of the upper Wisconsin River from State Line, Vilas
Co., to Wausau, Marathon Co. He continued during the
succeeding years, 1894 to 1897, to explore the Wisconsin
River valley and the northern water shed. Collections
were made from Drummond, Bayfield Co., down the White
River ; along the Montreal River in Iron Co. to Lake Super¬
ior ; along the Lake Superior shore from Montreal River to
200 Wisconsin Academy of Sciences , Arts , cmcZ Letters.
Ashland, Bayfield, Oak and La Presque Islands, Cornuco¬
pia, Herbster and Superior.
What would one not give now and what hardships not en¬
dure for the sight of those virgin forests, moss-covered
ledges, forest-edged waterfalls, cascades, and rapids, which
he found along the northern Wisconsin rivers !
His collections were extensive, both of flowering plants
and of bryophytes, and are now the most valuable of any
made in the state. In 1895 the collection of the year 1893
was partially reported and published (3). He listed 33
species. The University herbarium contained only a few
of the published specimen numbers. The numbered pack¬
ets in the herbarium which contain additional species or in
which the name is changed upon re-examination are as fol¬
lows :
No. 998. Aneura latifrons is now known as Riccardia latifrons
Lindb., and contains also Sphenolobus exsectaeformis
(Breidl.) Steph.
No. 1284. Aneura species? is Riccardia palmata (Hedw.) Carruth.
No. 1548. is Pellia neesiana (Gottsche) Limpr.
No. 59. Jungermannia schraderi is Odontoschisma denudatum
(Mart.) Dumort., and Lophozia porphyroleuca (Nees)
Schiffn.
No. 2864. Jungermannia schraderi is J. lanceolata L.
Nos. 638, 1247. Jungermannia excisa is Lophozia incisa (Schrad.)
Dumort.
No. 628. Jungermannia exsecta is Sphenolobus exsectaeformis
(Breidl.) Steph.
Nos. 1032, 1980. Chiloscyphus polyanthos is C. fragilis (Roth)
Schiffn.
No. 1992. Chiloscyphus polyanthos is C. rivularis (Schrad.) Loeske.
Nos. 948, 951, 952. Kantia trichomanis is Calypogeia neesiana
(Massal. & Carrest.) K. Mull.
No. 463. Cephalozia multiflora is C. connivens (Dicks.) Lindb.
No. 1254. Cephalozia multiflora contains also C. pleniceps (Aust.)
Lindb.
No. 1263. Cephalozia multiflora contains also Lophozia porphyro¬
leuca (Nees) Schiffn. and Sphenolobus hellerianus
(Nees) Steph.
No. 1331. Cephalozia multiflora contains also C. connivens (Dicks.)
Lindb. and C. pleniceps (Aust.) Lindb.
Nos. 15, 20, 25, 27, 48, 89, 35, 561, 1560, 1757, 1871, 2116, 2192, 2194.
All Ptilidium pulcherrium. (Web.) Hampe.
No. 2127. Ptilidium ciliare as listed.
Conklin — The Hepaticae of Wisconsin.
201
No. 1325, 1327. P or ella platyphylla is P. platyphylloidea (Schwein.)
Lindb.
No. 1026. Radula complanata is R. obconica Sullivant.
No. 2365. Frullania asagrayana contains also F. brittoniae Evans.
(7) In the report of Cheney and True on the flora of
Madison and vicinity (2), fifteen liverworts were listed.
A re-examination of this material gives a few additions
and changes of nomenclature, as follows :
No. 188.
No. 883.
No. 884.
No. 888.
No. 890.
No. 899.
Frullania Asagrayana contains also F. brittoniae Evans.
Ptilidium ciliare contains also P. pulcherrimum (Web.)
Hampe.
Lophocolea bidentata is L. heterophylla (Schrad.) Du-
mort.
Preissia commutata is now known as P. quadrata (Scop.)
Nees.
Asterella hemisphaerica is now known as Reboulia hemis¬
phaerica (L.) Raddi.
Grimaldia barbifrons is now known as G. fragrans
(Balb.) Corda.
(8) In August, 1902, Prof. C. E. Allen collected exten¬
sively on two of the Apostle Islands, Madaline and La-
Presque, Ashland Co. This collection was reported in
August 1904 (4) . 21 species were listed. A re-examina¬
tion of this material added 9 species and the following cor¬
rections :
No. 4. Pellia endaevifolia is P. epiphylla (L.) Corda.
No. 10. Chiloscyphyus polyanthos is C. pallescens (Ehrh.) Dumort.
No. 17. Ptilidium ciliare is P. pulcherrimum (Web.) Hampe.
No. 19. Scapania undulata is S. subalpina (Nees) Dumort.
Additional species found in the material were :
Lophozia kaurini (Limpr.) Steph., L. porphyroleuca
(Nees) Schiffn. Sphenolobus exsectaeformis (Breidl.)
Steph., Plagiochila asplenioides (L.) Dumort, Cephalozia
catenulata (Huben.) Spruce, Jungermannia pumila With.,
J. sphaerocarpa Hook., Nardia hyalina (Lyell) Carringt.,
Odontoschisma denudatum (Mart.) Dumort.
(9) In 1914 G. H. Conklin published a preliminary re¬
port (6). The area covered in this report included Doug¬
las and Bayfield Counties in Wisconsin, and Carlton, St.
Louis, and Lake Counties in Minnesota. 83 species were
listed, 67 of which were found in Wisconsin. Since that
202 Wisconsin Academy of Sciences , Arts , and Letters.
publication 31 additional species have been found by the
writer in Douglas, Bayfield, Ashland, and Iron Counties,
namely :
Frullania asagrayana Mont., F. oakesiana Aust., Lejeu -
nea cavifolia (Ehrh.) Lindb., Lophozia bicrenata
(Schmid.) Dumort., L. excisa (Dicks.) Dumort., L. atten-
uata (Mart.) Dumort., L. alpestris (Schleich.) Evans, L.
badensis (Gottsche) SchifTn., Jungermannia sphaerocarpa
Hook., J. schiffneri (Loitles.) Evans, Scapania undulata
(L.) Dumort., S. paludicola Loeske & K. Mull., S. cuspidu-
ligera (Nees) K. Mull., Chiloscyphus fragilis (Roth)
Schiffn., Cephaloziella byssacea (Roth) Warnst., C. ela-
chista (Jack) Schiffn., C. bifida (Schreb.) Schiffn., C. ham -
peana (Nees) Schiffn., Odontoschisma denudatum (Mart.)
Dumort., O. macounii (Aust.) Underw., Diplophyllum api-
culatum (Evans) Steph., D. gymnostomophilum Kaal.,
Nardia hyalina (Lyell) Carringt., Jungermannia pumila
var. rivularis With., Pallavicinia fiotowiana (Nees) Lindb.
Ricciocarpus natans (L.) Corda, Sphenolobus scitulus
(Tayl.) Steph., Reboulia hemisphaerica (L.) Raddi, Porella
platyphylloidea (Schwein.) Lindb., L. ventricosa (Dicks.)
Dumort., Sphenolobus hellerianus (Nees) Steph.
The writer can now report 102 species from Wisconsin in
the counties bordering Lake Superior. The following spe¬
cies also have been collected by him in the Minnesota coun¬
ties of Carlton, St. Louis, Lake, and Cook which have not
yet been found in Wisconsin.
Lophozia longidens (Lindb.) Macoun, L. marchica
(Nees) Steph., L. grandiretis (Lindb.) Schiffn., L. ruthe-
ana (Limpr.) M. A. Howe, Calypogeia suecica (Arn. &
Perss.) K. Mull., Sphenolobus minutus (Crantz.) Steph.,
Marsupella emarginata (Ehrh.) Dumort., Aster ella Lud-
wigii (Schwaegr.) Underw., Scapania umbrosa (Schrad.)
Dumort.
The Hepaticae of North America are represented in Wis¬
consin by 44 genera and 116 species. This is a fairly good
number considering that only 33 counties in the state have
been explored. It will be seen that this review of the
known specimens found in the state adds 37 species which
have not before been correctly reported, namely :
Conklin — The Eepaticae of Wisconsin.
203
Metzgeria conjugata, Pallavicinia flotowiana, Chiloscy -
phus fragilis, Jungermannia cordifolia, J. schiffneri,
J. sphaerocarpa , J. rivularis, Lophozia alpestris, L. attenu-
ata, L. badensis, L. bicrenata, L. excisa, L. guttulata,
Diplophyllum apiculatum, D. gymnostomophilum, Nardia
hyalina, N. crenulata, Sphenolobus scitulus, Cephaloziella
bifida, C. byssaceu, C. elachista, C. hampeana, Odonto-
schisma denudatum , 0. macounii, Scapania cuspiduligera,
S. dentata, S. paludicola, S. undulata, Frullania inflata, F.
oakesiana, Anthoceros punctatus, A. macounii, Riccia soro-
carpa, R. beyrichiana, Grimaldia pilosa, G. rupestris, Frul¬
lania riparia.
It will be seen from the state-county map (fig. 1) that
only in a few counties has any systematic collecting been
done. The number of species collected in each county is
indicated by numerals. The most promising counties for
new species not as yet visited are these of Door, the coun¬
ties along the Mississippi in the unglaciated areas, and the
Red Cliff Indian Reservation in Bayfield Co. I know of no
regions more promising for new species or for the exten¬
sion of range of known species than these areas. This sur¬
vey of the Hepaticae of Wisconsin, which has been deferred
a long time, was made possible when L. S. Cheney in 1926
released to the University herbarium the bulk of the speci¬
mens collected in northern Wisconsin in the years 1893 to
1897. As stated, a large number of these specimens were
sent Dr. L. M. Underwood, some of which were reported in
the Transactions of the Academy in 1894. After Dr. Un¬
derwood’s death, the New York Botanical Garden pur¬
chased his herbarium and duplicates. After a lapse of
these many years, these three groups of collected specimens
have again come together and are deposited in the Univer¬
sity of Wisconsin Herbarium.
Grateful acknowledgement is here given to Dr. A. W. Ev¬
ans of Yale University, Miss Annie Lorenz of Hartford,
Conn., and Miss Caroline C. Haynes of Highland, New Jer¬
sey, for their assistance during the past fifteen years in
verifying and naming difficult species, and to Dr. C. E. Al¬
len and Dr. J. J. Davis of the University of Wisconsin for
their courtesy permitting a free examination of the mate¬
rials in the University Herbarium.
204 Wisconsin Academy of Sciences , Arts, and Letters .
The names of the collectors and herbaria which have
been abbreviated are:
L. S. Cheney of Barron, Wis., Cheney.
C. E. Allen of Madison, Wis., Allen.
G. H. Conklin of Superior, Wis., Conklin.
University of Wisconsin, U.
New York Botanical Garden, NY.
University of Minnesota, UM.
Sullivant Moss Society Hepatic Herbarium, SMS.
L. R. Wilson of Superior, Wis., Wilson.
N. C. Fassett of Madison, Wis., Fassett.
J. M. Holzinger of Winona, Minn., H.
In the catalog the specimens following a named collector
are of his collection and the stations following a named
county are from that county.
MARCHANTIALES
Riccieceae
1. Riccia fluitans L. Herb. I. A. Lapham, 1861, (no sta¬
tion recorded), ex. coll. F . J. Hale 43 (U) ; Dells Wisconsin
River, Sauk Co., July 7, 1883, E . G. Knight (NY) ; Barron,
Barron Co., Aug. 13, 1891, Cheney 62 (U) ; Newbold,
Oneida Co., July 10, 1893, 1584 (U) ; Stevens Point, Port¬
age Co., July 4, 1894, 3490 (U) ; Webster, Burnett Co.,
1894, 3441 (U) ; Drummond, Bayfield Co., June 22, 1896,
4040 (U) ; Superior, Douglas Co., July 26, 1897, 1713 (U) ;
same location, Aug. 3, 1907, Conklin 267, 2365 (SMS).
2. Riccia arvensis Aust. Madison, Dane Co., Oct. 10,
1891, Cheney ( U ) ; Webster, Burnett Co., 1894, 3441 p. p.
(U) ; Madison, Dane Co., Oct. 10, 1892, Cheney & True
(U) ; Arena, Iowa Co., Sept. 2, 1922, Allen (U). Near
Afton, Rock Co., Oct. 26, 1928, Cheney, 12871 (U), det.
M. A. Howe.
(Two Ricciae appear in the collections which can not be
named. Both specimens are without spores and imma¬
ture.)
3. Riccia beyrichiana Hampe. Bagley, Grant Co., June
5, 1927, Cheney, 12205 (U) , det. M. A. Howe.
Conklin — The Hepaticae of Wisconsin.
205
4. Riccia sorocarpa Bisch. Glen Haven, Grant Co., June
6, 1927, Cheney 12245 (U), det. M. A. Howe.
5. Ricciocarpus natans (L.) Corda. Black Earth, Dane
Co., (no date recorded), Hale (U) ; Madison, Dane Co.,
Feb. 20, 1892, Cheney “D” (U) ; Orienta, Bayfield Co., July
15, 1897, 7350 (U) ; Billings Park, Superior, Douglas Co.,
Oct. 1, 1924, Wilson, 2358 (SMS) ; Lodi, Columbia Co., Oct.
9, 1926, (SMS) ; Oconomowoc, Waukesha Co., June 27,
1925, (SMS) ; St. Croix Falls, Polk Co., Sept. 4, 1927, Fas -
sett & Wilson (U).
Rebouliaceae
1. Grimaldia fray vans (Balb.) Corda. Madison, Dane
Co., May 1891, Cheney & True (U) ; same location, June 1,
1891, Cheney (U) ; Potosi, Grant Co., May 19, 1928, 11295
(U) ; same location, June 1, 1926, 11407 (U) ; Glen Haven,
Grant Co., May 4, 1926, 12136 (U) ; Lake Wingra, Madi¬
son, Dane Co., Aug. 23, 1893, L. M. Undenvood (NY) ; St.
Croix Falls, Polk Co., C. D . Baker, 20 (NY) ; Glen Haven,
Grant Co., April 12, 1921, Allen 3 (U) ; Bradford Tp. Rock
Co., Aug. 7, 1928, Cheney 12760 (SMS.U), det. C. C.
Haynes; Hanover, Aug. 17, 1928, Cheney 12793 (SMS.U).
2. Grimaldia pilosa (Hornem.) Lindb. Glen Haven,
Grant Co., May 14, 1927, Cheney 12160, 12137, (U) ; Bag-
ley, May 30, 1927, 12206 (U) ; Osceola, Polk Co., May 6,
1901, Daisy S. Howe 24 (UM). Listed as Fimbriaria ten-
ella.
3. Grimaldia mtpestris ( Nees ) Lindenh. Burton, Grant
Co., June 4, 1926, Cheney 11423 (U) ; Potosi, May 7, 1926,
11210, 1140 p. p., (ver. A. Lorenz), (U).
4. Reboulia hemisphaerica (L.) Raddi. Racine, Racine
Co., I. A. Lapham, (U) ; Dells Wisconsin River, Sauk Co.,
(collector unrecorded), Aug. 1, 1858, (U) ; Madison, Dane
Co., May 26, 1891, Cheney 2 (U) ; Dells Wisconsin River,
Sauk Co., Aug. 9, 1893, L. M. Underwood (U & NY) ; Pet-
anwells Rock & Germantown, Adams & Juneau Cos., July
19, 1894, Cheney 3704 p. p., 3705 B. (U & NY) ; White
River Creek, July 23, 1894, (U & NY) ; Nakoosa, Wood Co.,
July 14, 1894, 3658 (U & NY) ; Bolin Valley, Buffalo Co.,
May 12, 1906, H. (SMS) ; Black River, Douglas Co., Oct.
ll
206 Wisconsin Academy of Sciences, Arts, omd Letters.
25, 1925, Conklin & Wilson 1828 (SMS) ; Potosi, Grant Co.,
May 19, 1925, Cheney 11297 (U) ; same location, May 25,
1926, 11340 (U) ; Trempealeau Mt., Trempealeau Co., May
17, 1890, H. (UM) ; Trevip Ridge, Trempealeau Co., Nov.
1, 1892, H. (UM) ; Bradford Tp. Rock Co., Aug. 7, 1928,
Cheney 12758 (U) ; Janesville, July 7, 1928, 12691, 12692
P.P. (U).
Marchantiaceae
1. Conocephalum conicum (L) Dumort. Herb. I. A.
Lapham, (U) ; Osceola, Polk Co., May 6, 1901, Daisy S.
Howe 25 (UM) ; Madison, Dane Co., May 26, 1891, Cheney,
3 (U & NY) ; Lac Vieux Desert, Yilas Co., June 20, 1893,
315 (NY) ; Noisy Creek, Oneida Co., July 19, 1893, 1968
(U) ; Whirlpool Rapids, Oneida & Lincoln Cos., July 20,
1893, 2005 (U) ; Petanwells Rock, Adams & Juneau Cos.,
July 19, 1894, 3700, 3705 (U & NY) ; Nakoosa, Wood Co.,
July 14, 1894, 3658 (U) ; Drummond, Bayfield Co., June 26,
1896, 4313 (U) ; Oak Island, Ashland Co., Aug. 7, 1896,
6041 (U) ; Maiden Rock, Pierce Co., Aug. 12, 1916, Allen
42 (U) ; Glen Haven, Grant Co., April 13, 1921, 67 (U) ;
Green Lake, Green Lake Co., July 16, 1922, (U) ; MitcheTs
Glen, Green Lake, July 20, 1922, (U) ; Wapogasset, Polk
Co., Sept. 4, 1927, (U) ; Winneboujou, Douglas Co., April
9, 1909, Conklin 1024 (SMS) ; Copper Creek, Aug. 5, 1909,
579, (SMS) ; Black River, Oct. 5, 1909, 1587, (SMS) ; Lake
Nebagamon, Sept. 3, 1911, 1208, (SMS) ; Brule River, July
22, 1923, 1898 (SMS) ; Squaw Bay, Bayfield Co., Aug. 10,
1917, 1330 (SMS) ; Orienta, Sept. 23, 1923, 1946 (SMS) ;
Siskiwit River, Oct. 5, 1924, Conklin & Wilson 2370
(SMS) ; Montreal River, Iron Co., Conklin 2004 Aug. 28,
1922, (SMS) ; Gurney Falls, Potato River, Sept. 7, 1925,
2542 (SMS) ; Mellen, Ashland Co., Aug. 29, 1922, 1739
(SMS) ; Osceola, Polk Co., Aug. 8, 1925, 2491 (SMS) ; Po¬
tosi, Grant Co., May 7, 1926, Cheney 11192 (U) ; Glen
Haven, Grant Co., May 28, 1927, 12202 (U).
2. Preissia quadrata (Scop.) Nees. Madison, Dane Co.,
May 26, 1891, Cheney 5, 4 (U & NY) ; Apostle Islands, Ash¬
land Co., Aug. 1902, Allen 28 B, 27 B, 27 C, (U) ; Petan¬
wells Rock & Germantown, Adams & Juneau Cos., July 19,
1894, Cheney 3700 (U & NY) ; Montreal River, Iron Co.,
Conklin — The Hepaticae of Wisconsin.
207
July 19, 1896, 5149 (U) ; Odanah Indian Reservation, July
21, 1896, 5190 (U) ; Oak Island, Ashland Co., Aug. 7,
1896, 5982 (U) ; Sand Bay, Bayfield Co., June 25, 1897,
6478 (U) ; Squaw Bay, June 30, 1897, 6633 (U) ; Herbster
and Port Wing, July 9, 1897, 7046 (U) ; Maiden Rock,
Pierce Co., Aug. 5, 1916, Allen 37 (U) ; same location, Aug.
17, 1916, 23 (U) ; Glen Haven, Grant Co., Aug. 30, 1921, 72
(U) ; Green Lake, Green Lake Co., July 25, 1922, (U) ;
Black River, Douglas Co., Oct. 5, 1909, Conklin 634 (SMS) ;
Squaw Bay, Bayfield Co., Aug. 10, 1917, 1287 (SMS) ; Sis-
kiwit Point, July 26, 1922, 1890 (SMS) ; Siskiwit River,
July 26, 1922, 1838 (SMS) ; Orienta Falls, Sept. 23, 1923,
1945 B. (SMS) ; Mellen, Ashland Co., Aug. 26, 1922, 1739
(SMS) ; Gurney Falls, Potato River, Iron Co., Sept. 7, 1925,
2679 (SMS) ; Osceola, Polk Co., Aug. 8, 1925, 2479 (SMS) ;
Potosi, Grant Co., May 25, 1926, Cheney 11341 (U) ; same
location, May 20, 1926, 11391 (U) ; June 9, 1926, 11442
(U) ; Bagley, Grant Co., May 30, 1927, 12207 (U) ; Glen
Haven, June 8, 1927, 12246 (U) ; same location, May 28,
1927, 12202 p. p. (U) ; Janesville, Rock Co., Sept. 3, 1928,
Cheney 12735 (U) ; same location, Sept. 22, 1928, 12692
(U) ; Beloit, July 7, 1928, 12639 (U).
3. Marchantia polymorpha L. Madison, Dane Co. I. A.
Lapham, (U) ; same location, Aug. 18, 1890, Cheney &
True, (U) ; same location, J. C. Carr (U) ; Osceola, Polk
Co., May 6, 1901, Daisy S. Howe 27 (UM) ; Blue Mounds,
Iowa Co., Aug. 1, 1903, (collector unrecorded), (U) ; Rug¬
by Junction Washington Co., Oct. 4, 1907, J. F. Brenckle
(U) ; Devil’s Lake, Sauk Co., July 11, 1903, Geol. & Nat.
Hist. Survey, 113 B. (U) ; Crystal Lake, Vilas Co., June 17,
1893, Cheney 219 (U) ; McNaughton, Oneida Co., July 6,
1893, 1439 (U) ; Granite Heights, Marathon Co., June 23,
1894, 2992 (U) ; Petanwells Rock, Adams & Juneau Cos.,
July 19, 1894, 3702, 3700 (U & NY) ; St. Louis River,
Douglas Co., July 31, 1897, 7918 (U) ; Apostle Islands, Ash¬
land Co., Aug. 1902, Allen 61 (U) ; Argyle, Lafayette Co.,
Sept. 17, 1925, (U) ; Sand Lake, Sawyer Co., Aug. 16, 1925,
(U) ; Glen Haven, Grant Co., April 14, 1921, 70 (U) ;
Maiden Rock, Pierce Co., Aug. 12, 1916, 45 (U) ; Gordon,
Douglas Co., Sept. 16, 1906, Conklin 82 (SMS) ; Superior,
Sept. 9, 1906, 918 (SMS) ; Copper Creek, Aug. 29, 1907,
208 Wisconsin Academy of Sciences , Arts, and Letters.
289 (SMS) ; Black River, Oct. 3, 1910, 1045 (SMS) ; Brule
River, July 22, 1923, 1904 (SMS) ; Montreal River, Iron
Co., Aug. 28, 1922, 1798 (SMS) ; Siskiwit River, Bayfield
Co., Aug. 26, 1923, 1839 (SMS) ; Orienta Falls, Sept. 23,
1923, 1909 (SMS) ; Osceola, Polk Co., Aug. 8, 1925, 2482
(SMS) ; Mellen, Ashland Co., Aug. 20, 1925, 2641 (SMS) ;
Potosi, Grant Co., May 15, 1926, Cheney 11262 (U) ; same
location, June 5, 1926, 11428 (U) ; Grant Co., 1927, 12207
(U) ; Bagley, May 30, 1927, 12207 p. p. (U) ; Bad River
Gorge, Mellen, Ashland Co., Aug. 9, 1927, Fas sett & Wilson
170 (U) ; Wallace Island, Lake Wapogasset, Polk Co., Sept.
3, 1927, Allen (U) ; Lake Wapogasset, Aug. 28, 1927, (U) ;
Beloit, Rock Co., July 5, 1928, Cheney 12637 (U).
JUNGERMANNIALES
Riccardiaceae
1. Riccardia pinguis (L.) S. F. Gray. Lac Vieux Des¬
ert, Vilas Co., June 1893, Cheney 510 (NY) ; same location,
June 22, 1893, 750 (U) ; Lake Nebagamon, Douglas Co.,
Oct. 1911, Conklin 1216 (SMS) ; Brule River, May 20, 1911,
1099 (SMS); Brule, (Whealdon’s), May 1, 1925, 2516
(SMS) ; Orienta Falls, Bayfield Co., Sept. 3, 1923, 1921 A.
(SMS) ; Cussen, July 25, 1913, 1341 (SMS) ; Mellen, Ash¬
land Co., Sept. 9, 1927, Fassett & Wilson (U).
2. Riccardia multi fida (L.) S. F. Gray. Tomahawk
Lake, Oneida Co., June 29, 1893, Cheney 998 (U) ; Rainbow
Rapids, July 1, 1893, 1145 (U & NY) ; Tomahawk Lake,
July 25, 1893, 2148 (U) ; Black River, Douglas Co., Oct. 3,
1911, Conklin 1036 p.p. (SMS) ; Brule River, Sept. 20,
1911, 1097 (SMS) ; Stone’s Bridge, June 12, 1916, 2156,
1438 (SMS) ; Mellen, Ashland Co., Sept. 7, 1927, Fassett &
Wilson (U).
3. Riccardia latifrons Lindb. State Line, Vilas Co.,
June 6, 1893, Cheney 470 (U & NY) : Apostle Islands, Ash¬
land Co., Aug. 1902, Allen, 92 C. (U) ; Sand Lake, Sawyer
Co., Aug. 28, 1926, (U) ; Solon Springs, Douglas Co., Aug.
1915, 24, 40 (U) ; Superior, Douglas Co., July 18, 1909,
Conklin 776 (SMS) ;Black River, Oct. 3, 1911, 1136 p.p.
(SMS) ; Lake Nebagamon, June 1, 1913, 1688 (SMS) ;
Conklin — The Hepaticae of Wisconsin.
209
Stone’s Bridge, Brule River, June 12, 1916, 1342 (SMS) ;
Balsam River, Sept. 15, 1925, 2570 (SMS) ; Cussen, Bay-
field Co., July 25, 1913, 1343 (SMS) ; Squaw Point, Aug.
10, 1917, 1325 (SMS).
4. Riccardia palmata (Hedw.) Carruth. Rainbow Rap¬
ids, Oneida Co., July 3, 1893, Cheney 1264 (U) ; Toma¬
hawk, Lincoln Co., July 25, 1893, 2184 (U) ; Drummond,
Bayfield Co., June 28, 1896, 4376 (U) ; St. Louis River,
Douglas Co., July 28, 1897, 7754 (U) ; Solon Springs, Doug¬
las Co., Aug. 14, 1915, Allen 62 (U) ; Solon Springs, Doug¬
las Co., Aug. 20, 1906, Conklin 107 (SMS) ; Black River,
Oct. 3, 1901, 1039 (SMS) ; St. Croix Lake, June 28, 1913,
1340 (SMS) ; Brule River, July 22, 1923, 1895 (SMS) ;
Cornucopia, Bayfield Co., Oct. 5, 1924, Conklin & Wilson
2387 (SMS).
5. Metzgeria conjugata Lindb. Dells Wisconsin River,
Sauk Co., July 7, 1884, E. G. Knight (NY) ; Coldwater
Canon, Adams Co., July 30, 1894, Cheney 3830 (NY & U).
6. Pallavicinia flotowiana (Nees) Lindb. Brule River,
(Whealdon’s), Douglas Co., July 22, 1923, Conklin 1850
(SMS).
Pelliaceae
1. Pellia epiphylla (L.) Cor da. Dells Wisconsin River,
Sauk Co, Aug. 19, 1898, L. M. Underwood (NY & U) ;
Grand Rapids, Wood Co, July 9, 1894, Cheney 3629 (NY &
U) ; Wisconsin River, Adams & Juneau Cos, July 27, 1894,
3788 (NY & U) ; Apostle Islands, Ashland Co, Aug. 1902,
Allen 29 A, 29 B, 27 A, 27 C, (U) ; Solon Springs, Douglas
Co, Aug. 22, 1915, 1, 18 (U) ; Lac Vieux Desert, Vilas Co,
Aug. 27, 1917, 57 (U) ; Eau Claire Lakes, Bayfield Co, Aug.
8, 1925, 71 (U) ; Lake Wapogasset, Polk Co, Aug. 28, 1927,
(U) ; Gordon, Douglas Co, Aug. 1907, Conklin 316 (SMS) ;
Copper Creek, May 5, 1909, 587 (SMS) ; Black River, Oct.
3, 1910, 1012, 1043 (SMS) ; Squaw Point, Bayfield Co,
Aug. 26, 1922, 1808 (SMS) ; Montreal River, Iron Co,
Sept. 6, 1925, 2551 (SMS) ; Mellen, Ashland Co, Sept. 7,
1927, Fassett & Wilson (U) ; Barron, Barron Co, April 26,
1928, Chaney 12565 (U & SMS).
2. Pellia fabroniana Raddi. Barron, Barron Co, Aug.
13, 1891, Cheney 64 (NY & U) ; Granite Heights, Mara-
210 Wisconsin Academy of Sciences, Arts, and Letters.
thon Co., June 22, 1894, 3018 (NY & U) ; Knowlton, Port¬
age Co., June 30, 1894, 3364 (NY & U) ; Mason, Bayfield
Co., July 8, 1896, 4686 (NY & U) ; Houghton Quarries,
July 25, 1896, 5429, 5457 (NY & U) ; Black River, Douglas
Co., April 9, and Oct. 7, 1911, Conklin 1066, 1026 (SMS) ;
Winneboujou, May 20, 1911, 1405 (SMS) ; Orienta Falls,
Bayfield Co., Sept. 23, 1923, 1933 (SMS).
3. Pellia neesiana (Gottsche) Limpr. Eagle River, Vi¬
las Co., June 27, 1893, Cheney 888 (NY & U) ; Rainbow
Rapids, Oneida Co., July 5, 1893, 1381 (NY & U) ; New-
bold, July 10, 1893, 1548 (NY & U) ; White River, Bayfield
Co., July 9, 1896, 4686 (U) ; Montreal River, Iron Co., July
19, 1896, 5029 (U) ; Houghton Quarries, Bayfield Co., July
25, 1896, 5429, 5457 (U) ; Devils Lake, Sauk Co., Aug. 16,
1917, Allen 25 (U) ; Lake Wapogasset, Polk Co., Sept. 4,
1927, (U) ; Gordon, Douglas Co., Sept. 16, 1906, Conklin
316 (SMS) ; Solon Springs, May 1908, 100 (SMS) ; Winne¬
boujou, April 9, 1910, 1027 (SMS) ; Black River, Oct. 3,
1910; 1045 (SMS); Stone’s Bridge, Brule River, June 12,
1916, 1436 (SMS) ; Mud Creek, June 7, 1925, 2473 (SMS) ;
Squaw Point, Bayfield Co., Aug. 10, 1817, 1298 (SMS) ;
Siskiwit Point, Oct. 5, 1924, 2375 (SMS) ; Cornucopia, Oct.
4, 1924, 2381 (SMS).
4. Blasia pusilla L. Newbold to Rhinelander, Oneida
Co., July 10, 1893, Cheney 1541 (NY & U) ; Drummond,
Bayfield Co., June 22, 1896, 4065, 4249 (NY & U) ; Mon¬
treal River, Iron Co., July 21, 1896, 5173 (U) ; Oak Island,
Ashland Co., Aug. 7, 1896, 6007 (U) ; Herbster, Bayfield
Co., Aug. 1, 1897, 6908 (U) ; Opposite Fond du Lac, Doug¬
las Co., Aug. 1, 1897, 7990 (U) ; Black River, Douglas Co.,
Oct. 3, 1910, Conklin 1040 (SMS) ; Squaw Bay, Bayfield
Co., Aug. 10, 1917, 1328 (SMS) ; Bark Point, July 27, 1923,
1929 (SMS) ; Siskiwit Point, July 29, 1923, 1888 (SMS) ;
Orienta, Sept. 23, 1923, 1849 (SMS) ; Montreal River, Iron
Co., Sept. 6, 1925, 2579 (SMS) ; Gurney Falls, Sept. 7,
1925, 2539 (SMS) ; Apostle Islands, Ashland Co., Aug
1902, Allen 6, 15 (U) ; Pine Creek, Maiden Rock, Pierce
Co., Aug. 15 and 16, 1916, 49, 50, 27 (U) ; St. Croix Falls,
Polk Co., July 12, 1890 H. (UM) ; Mellen, Ashland Co., Sect.
17, July 9, 1927, Fassett & Wilson, (U) ; Bad River Gorge,
Conklin — The Hepaticae of Wisconsin.
211
Aug. 9, 1927, (U) ; Barron, Barron Co., Nov. 17, 1929,
Cheney 12931 (U&SMS).
5. Fossomhroma foveolata Lindb. Witch’s Gulch, Adams
Co., July 31, 1891, Cheney 3883 (NY & U; Apostle Islands,
Ashland Co., Aug. 1902, Allen 28 A. (U) ; Siskiwit River,
Bayfield Co., July 29, 1923, Conklin 1847 (SMS) ; Arena,
Iowa Co., Sept 1, 1922, Allen (U) ; (Specimen collected is
immature, lacks spores and cannot be named with any cer¬
tainty. )
Lophoziaceae
1. Plagiochila asplenioides (L.) Dumort. Dells Wiscon¬
sin River, Sauk Co., July 7, 1883, E. G. Knight 12 (NY) ;
same location, July 8, 1893 (NY) ; same location Aug. 19,
1893, L. M . Underwood (NY) ; Lac Vieux Desert, Vilas Co.,
June 22, 1893, Cheney 547 (U) ; McNaughton to Newbold,
Oneida Co., July 8, 1893, 1526 (U) ; Mason, Bayfield
Co., July 8, 1896, 4641 (NY & U) ; Montreal River,
Iron Co., July 19, 1896, 5050 (U) ; same location. July
21, 1896, 5207 (U) ; Witch’s Gulch, Adams Co., July
28, 1896, 3892 (U) ; Presque Isle, Ashland Co., Aug. 6,
1896, 5951 (U) ; Herbster, Bayfield Co., July 7, 1897, 6906,
6944 p.p. 6947 (U) ; Port Wing, July 8, 1897, 7010 (U) ;
same location, July 12, 1897, 7214 (U) ; Brule River, Doug¬
las Co., July 19, 1897, 7525 (U) ; St. Louis River, July 31,
1897, 7963, (U) ; Apostle Islands, Ashland Co., Aug. 1902,
Allen 25 B, 25 (U) ; Devils Lake, Sauk Co., Aug. 6, 1915,
20, 39 (U) ; Solon Springs, Douglas Co., Sept. 5, 1915, 64
(U) ; Solon Springs, Douglas Co., Summer 1907, Conklin
294 (SMS) ; Copper Creek, Aug. 5, 1909, 542 (SMS) ;
Black River, Oct. 5, 1909, 663 (SMS) ; Brule River, May 7,
1911, 1220 (SMS) ; Lake Nebagamon, July 1, 1913, 1774
(SMS); Stone’s Bridge, June 12, 1916, 1414 (SMS);
Squaw Point, Bayfield Co., Aug. 10, 1917, 1332 (SMS) ;
same location, Aug. 26, 1922, 1976 (SMS) ; Bark Point,
Bayfield Co., July 27, 1923, 1880 (SMS) ; Siskiwit Point,
Sept. 1, 1923, 1891 (SMS) ; Cornucopia, Oct. 5, 1924, 2439
(SMS) ; Montreal River, Iron Co., Aug. 28, 1922, 2001
(SMS) ; same location, Sept. 6, 1925, 2561 (SMS) ; Bar¬
ron, Barron Co., March 31, 1927, Cheney 21080 (U) ; Bag-
212 Wisconsin Academy of Sciences, Arts, and Letters.
ley, Grant Co., June 4, 1927, 12227 (U) ; Mellen, Ashland
Co., Sept. 7, 1927, Fassett & Wilson (U).
2. Mylia anomala (Hook.) S. F. Gray. Black River,
Douglas Co., Oct. 3, 1910, Conklin 961 (SMS) ; Superior,
June 4, 1911, 1104, 2224 (SMS) ; Stone’s Bridge, Brule
River, June 26, 1916, 1437 (SMS).
3. Lophocolea heterophylla (Schrad.) Dumort. Milwau¬
kee, Milwaukee Co., July 1, 1862, I. A. Lapham (U) ; Mad¬
ison, Dane Co., April 19, 1890, Cheney & True (U) ; Fay¬
ette, Lafayette Co., Dec. 1891, Cheney 7, 17 (NY & U) ;
Lake Mills, Jefferson Co., March 30, 1891, G, H, I, K, M, N,
(NY & U) ; Dells Wisconsin River, Sauk Co., Aug. 19,
1893, L. M. Underwood (NY & U) ; Lac Vieux Desert, Yilas
Co., June 13, 1893, Cheney 52 (U) ; same location, June 20,
1893, 355 (U) ; State Line, June 21, 1893, 740 p.p. (U);
Thomas’ Place, Lac Vieux Desert, June 22, 1893, 578, 576
(U) ; Rhinelander, Oneida Co., July 18, 1893, 1824 (NY &
U) ; Tomahawk, Lincoln Co., July 27, 1893, 2297 p.p., 2359
(U) ; Mosinee, Marathon Co., July 27, 1894, 3231 (U) ;
same location, July 28, 1894, 3308 (U) ; Cold water Canon,
Adams Co., July 30, 1894, 3814 (U) ; Drummond, Bayfield
Co., June 29, 1896, 4376 (NY & U) ; Herbster, July 7, 1897,
6882, 6948 (U) ; Port Wing, July 12, 1897, 7185 p.p. (U) ;
White River, Ashland Co., July 13, 1896, 4781 (U) ; Wilson
Island, Aug. 6, 1896, 5901, 5939 (NY & U) ; Blue Mounds,
Iowa Co., Aug. 1, 1903, (U).
Two specimens Madison, Dane Co., April and May 1891,
Cheney; and Baraboo, Sauk Co., R . H. True (NY & U) ; are
referred here. Both are named Lophocolea bidentata (L.)
Dumort. The dioecious inflorescence of L. bidentata can¬
not be determined from condition of the specimens. They
are referred therefore to Lophocolea heterophylla (Schrad.)
Dumort. Barron, Barron Co., March 31, 1927, Cheney
12107 (U) ; Glen Haven, Grant Co., May 6, 1927, 12145
(U) ; same location, May 11, 1927, 12154 (U) ; May 31,
1927, 12213 (U) ; Bagley, Grant Co., May 20, 1927, 12187
(U) ; Bear Trap Lake, Polk Co., Aug. 30 and Aug. 31, Sept.
3, and Sept. 8, 1927, Allen (U) ; Lake Wapogasset, Aug.
28, Sept. 4, 1927, (U) ; Wallace Island, Lake Wapogasset,
Sept. 3, 1927 (U) ; Apostle Islands, Ashland Co., Aug. 1902,
78 C, 79 C, 22 A, 79 A, 19 B, 22 B, 64 B, 30 B, 51 A, 78 B,
Conklin — The ITepaticae of Wisconsin.
213
92 A, 23, 91 B, 79 B, 42 A, (U) ; Devil’s Lake, Sauk Co.,
July 11, 1903, ( collector unrecorded ), (U) ; Green Lake,
Green Lake Co., July 18, 1922, Allen (U) ; Argyle, Lafay¬
ette Co., Sept. 17, 1925, (U) ; Sand Lake, Sawyer Co., Aug.
12 to 28, 1925, (U) ; Solon Springs, Douglas Co., Aug. 5 to
14, 1915, 6, 14, 15, 29, 53, 69, 24, 47, 11 (U) ; Maiden Rock,
Pierce Co., Aug. 15, 1915, 27 (U) ; Eau Claire Lake, Bay-
field Co., Aug. 8, 1925, 66 (U) ; Oconomowoc, Waukesha
Co., June 27, 1925, Wilson (SMS) ; Solon Springs, Douglas
Co., May 29, 1907, Conklin 423 (SMS) ; Superior, July 18,
1909, 760 (SMS); Copper Creek, Aug. 29, 1909, 379
(SMS); Lake Nebagamon, Sept. 3, 1911, 1149 (SMS);
Black River, Oct. 2, 1912, 1064 (SMS) ; Brule River,
(Whealdon’s), July 22, 1923, 1977 (SMS); Squaw Point,
Bayfield Co., Aug. 10, 1917, 1333 (SMS) ; Lake Nameka-
gon, Ashland Co., Aug. 14, 1924, 2278 (SMS) : Marshland,
Buffalo Co., June 23, 1902, H. (UM) ; same location, June
24, 1902, (UM) ; Mellen, Ashland Co., Sect. 16, July 9, 1927,
Fassett & Wilson (U) ; St. Croix Falls, Polk Co., Sept. 4,
1927, (U) ; Barron, Barron Co., March 31, 1927, Cheney
12081, 12088, 12095, 12099 (U) ; Glen Haven, Grant Co.,
June 10, 1927, 12257 p.p. (U) ; Bradford Tp., Rock Co.,
Aug. 2, 1928, Cheney 12783 (U) ; Clinton, Aug. 14, 1928,
12791 (U) ; Neward Tp., Aug. 25, 1928, 12807 (U) ; same
location, Oct. 4, 1928, 12845 (U) ; Oct. 18, 1928, 12845 (U) .
4. Lophocolea minor Nees. Fayette, Lafayette Co., Dec.
1891, Cheney 11 (NY) ; Grandmother Bull Falls, Lincoln
Co., July 29, 1893, 2580 (NY & U) ; Mosinee, Marathon Co.,
June 27, 1894, 3308 (N Y& U) ; Stevens Point, Portage Co.,
July 4, 1894, 3486 (NY & U) ; Mason, Bayfield Co., July
8, 1898, 4645 (NY & U) ; Montreal River, Iron Co., July 21,
1896, 5128 (U) ; Montreal River, and Odanah Indian Res¬
ervation, July 21, 1898, 5191 (U) ; St. Louis River, Douglas
Co., July 30, 1897, 7882 (U) ; Dells Wisconsin River, Kil-
bourn, Sauk Co., Aug. 13, 1912, M. F. Somerville (SMS) ;
Black River, Douglas Co., Oct. 5, 1909, Conklin 673
(SMS) ; (Whealdon’s), Brule River, Aug. 24, 1924, 2279 B.
(SMS) ; Brule River, May 7, 1925, 2511 (SMS) ; Montreal
River, Iron Co., Aug. 27, 1922, 1799 (SMS) ; Orienta Falls,
Bayfield Co., Sept. 23, 1923, 1943 (SMS) ; Potosi, Grant
Co., May 7, 1926, Cheney 11200 (U) ; same location, May
214 Wisconsin Academy of Sciences , Arts, and Letters.
28, 1926, 11369 (U) ; Werley, Sept. 16, 1927, 12433 (U) ;
Glen Haven, May 16, 1927, 12168 (U) ; Bagley, June 8,
1927, 12253 (U) ; St. Croix Falls, Polk Co., Sept. 4, 1927,
Fassett & Wilson (U) ; Marshland, Buffalo Co., June 24,
1902, H. (UM & SMS) ; Winona Bluffs, Wisconsin side, Oct.
27, 1902, (UM) ; Glen Haven, Grant Co., June 10, 1927,
Cheney 12257 p.p. (U) ; Bradford Tp., Rock County, Aug.
14, 1928, Cheney 12792 (U) ; same location, Sept. 1, 1928,
12811 (U).
5. Chiloscyphus pallescens (Ehrh.) Dumort. Noisy
Creek, Oneida Co., July 19, 1893, Cheney 1964 (U) ; Wilson
Island, Ashland Co., Aug. 1896, 5901 (U) ; Herbster, Bay-
field Co., July 7, 1897, 6682 p. p. (U) ; Apostle Islands, Ash¬
land Co., Aug. 1902, Allen 62 (U) ; Maiden Rock, Pierce
Co., Aug. 17, 1915, 30 (U) ; Copper Creek, Douglas Co.,
Sept. 15, 1902, Conklin 250 (SMS) ; same location, Aug. 5,
1906, 591 (SMS) ; Gordon, Aug. 1907, 522 (SMS) ; Winne-
boujou, May 20, 1911, 1407, (SMS) ; Black River, Aug. 16,
1922, 1778 (SMS); (Whealdon’s) , Brule River, July 22,
1923, 1896 (SMS) ; Stone’s Bridge, Aug. 24, 1921, 2280
(SMS) ; Mellen, Ashland Co., Aug. 29, 1922, 1992 (SMS) ;
Montreal River, Iron Co., Aug. 26, 1923, 1974, 1982
(SMS) ; Siskiwit River, Bayfield Co., Sept. 1, 1923, 1910
(SMS) ; Bark Point, Oct. 4, 1924, 2434 (SMS) ; St. Croix
Lake, Douglas Co., Nov. 2, 1924, 2419 (SMS) ; Bridgeport,
Grant Co., July 19, 1927 Cheney 12346 (U) ; Bagley, Aug¬
ust 24, 1927, 12390 (U) ; Lake Wapogasset, Polk Co., Sept.
4, 1927, Allen (U).
6. Chiloscyphus fragilis (Roth) Schiffn. Tomahawk
Lake, Oneida Co., June 29, 1893, Cheney 1032 (U) ; Whirl¬
pool Rapids, Oneida & Lincoln Cos., July 20, 1893, 1980
(U) ; Argyle, Lafayette Co., Sept. 17, 1925, Allen (U) ;
Lake Nebagamon, Douglas Co., May 7, 1911, Merry man
1259 (SMS) ; Stone’s Bridge, June 12, 1916, Conklin 2195
(SMS) ; Squaw Bay, Bayfield Co., Aug. 10, 1917, 1326
(SMS) ; Stone’s Bridge, Douglas Co., July 4, 1924, 2203
(SMS) ; St. Croix Lake, Nov. 2, 1924, 2421 (SMS) ; Mud
Creek, July 7, 1925, 2474 (SMS).
7. Chiloscyphus rivularis (Schrad.) Loeske. Whirlpool
Rapids, Oneida & Lincoln Cos., June 20, 1893, Cheney 1992
(NY & U) ; Rainbow Falls, Oneida Co., July 1893, 1032
Conklin — The Ilepaticae of Wisconsin.
215
(NY & U) ; Knowlton, Marathon Co., June 30, 1894, 3349
(NY & U) ; Witch’s Gulch, Adams Co., July 31, 1894, 3898
(U) ; Houghton Quarries, Bayfield Co., July 25, 1896, 5483
(U) ; Winneboujou, Douglas Co., April 9, 1910, Conklin
1113 (SMS) ; same location, July 4, 1910, 1032 (SMS) ;
Stone’s Bridge, Brule River, June 12, 1916, 1411 (SMS) ;
Brule River, May 1, 1923, 2499 (SMS) ; Hatfield, Jackson
Co., Oct. 25, 1907, H. (UM & SMS).
8. Chiloscyphus polyanthos (L.) Corda. Copper Creek,
Douglas Co., Sept. 15, 1902, Conklin 458 (SMS) ; Superior,
Sept. 12, 1909, 718 (SMS) ; Stone’s Bridge, June 20, 1916,
1428 (SMS) ; Black River, July 26, 1922, 1767 (SMS) ; St.
Croix Lake, June 29, 1924, 2227 (SMS) ; Solon Springs,
Sept. 12, 1915, Allen 2, 4, 73 (U).
9. Harpanthus scutatus (Web. & Mohr) Spruce. Fay¬
ette, Lafayette Co., Dec. 1891, Cheney 12 (NY) ; Grand¬
mother Bull Falls, Lincoln Co., July 29, 1893, 2672 (NY &
U) ; Grand Rapids, Wood Co, July 9, 1894, 3631 (NY &
U) ; Coldwater Canon, Adams Co, July 30, 1894, 3814 p. p.
(U) ; Montreal River, Iron Co, July 22, 1896, 5244 p. p.
(U) ; Houghton Quarries, Bayfield Co, July 25, 1896, 5493
(U) ; Solon Springs, Douglas Co, July 1, 1909, Conklin
470 (SMS) ; Black River, Oct. 5, 1909, 1354 (SMS) ; Squaw
Bay, Bayfield Co, Aug. 10, 1917, 1291 (SMS) ; Brule River.
Douglas Co, May 1, 1925, 2518 (SMS) ; Sand Lake, Saw¬
yer Co, Sept. 28, 1925, Allen (U) ; Bass Lake, Sept. 15,
1925, (U) ; Solon Springs, Douglas Co, Aug. 1915, 7, 28,
(U) ; Glen Haven, Grant Co, Aug. 16, 1926, Cheney 11668
(U) ; same location, June 10, 1927, 12257 (U) ; Patch
Grove, Sept. 22, 1927, 12454 (U) ; same location, Sept. 24,
1927, 12465 (U) ; Bagley, Aug. 25, 1927, 12391 (U) ; Bear
Trap Lake, Polk Co, Sept. 8, 1927, Allen (U).
10. Geocalyx graveolens (Schrad.) Nees. Lac Vieux
Desert, Vilas Co, June 14, 1893, Cheney 82 p. p. (U) ; State
Line, June 14, 1893, 469 p. p. (U) ; Eagle River, Oneida Co,
June 28, 1893, 952 p. p. (U) ; Rainbow Rapids, July 3, 1893,
1264 p. p. (U) ; Coldwater Canon, Adams Co, July 30, 1894,
3814 p. p. (U) ; Mason, Bayfield Co, July 7, 1896, 4367
(NY) ; Oak Island, Ashland Co, Aug. 7, 1896, 5992 p. p.
(NY) ; Brule Swamp, Douglas Co, Aug. 9, 1915, Allen 17
(U) ; same location, Aug. 13, 1915, 56 (U) ; Solon Springs,
218 Wisconsin Academy of Sciences, Arts, and Letters .
Douglas Co., Oct. 1907, Conklin 387 (SMS) ; Superior,
Sept. 6, 1909, 489 (SMS) ; Winneboujou, April 9, 1910,
1030 (SMS) ; Hall’s Swamp, Winneboujou, May 20, 1911,
1460 (SMS) ; Lake Nebagamon, June 13, 1913, 1775
(SMS) ; Head St. Croix Lake, June 25, 1913, 1391 (SMS) ;
Cornucopia, Bayfield Co., Sept. 1, 1913, 1873 (SMS) ;
Stone’s Bridge, Douglas Co., June 12, 1916, 2198 (SMS) ;
Squaw Point, Bayfield Co., Aug. 11, 1917, 1280 (SMS) ;
Mellen, Ashland Co., Aug. 29, 1922, 1806 (SMS) ; Siski-
wit Point, Bayfield Co., July 29, 1923, 1889 (SMS) ; Mon¬
treal River, Iron Co., Sept. 6, 1925, 2559 (SMS) ; Barron,
Barron Co., March 31, 1927, Cheney 12098, 12104, 12105,
12115 (U) ; Marshland, Buffalo Co., June 1903, H. (UM).
11. Nardia crenulata (Smith) Lindb. Arena, Iowa Co.,
Sept. 1, 1922, Allen (U).
12. Nardia hyalina ( Lyell ) Carringt. Fayette, Lafay¬
ette Co., Dec. 1891, Cheney (NY) ; Dells Wisconsin River,
Sauk Co., Aug. 19, 1893, L. M. Underwood (NY) ; Nakoosa,
Wood Co., July 14, 1894, Cheney 3659 (NY & U) ; Petan-
wells Rock, Adams & Juneau Cos., July 16, 1894, 3680
(NY & U) ; same location, July 19, 1894, 3702, 3703, 3704
(NY & U) ; Witch’s Gulch, Adams Co., July 31, 1894, 3883
(U) ; Montreal River, Iron Co., July 20, 1896, 5115, 5128,
5152 (NY & U) ; Houghton Quarries, Bayfield Co., July 25,
1896, 5429 p. p., 5444 (U) ; Herbster, July 7, 1897, 6867,
6907, 6946 p. p. (U) ; Galesville, Trempealeau Co., 1919,
150 H. (Yale U) ; Apostle Island, Ashland Co., Aug. 1902,
Allen 27 A, (U) ; Squaw Point, Bayfield Co., Aug. 10, 1917,
Conklin 2322, 1276, 1282, 1289 (SMS) ; Siskiwit Point,
Aug. 27, 1922, 1813 (SMS) ; Bark Point, Sept. 23, 1923,
1980 (SMS) ; Orienta Falls, Sept. 23, 1923, 1924 (SMS) ;
Herbster, Sept. 23, 1923, 1920 (SMS) ; Black River, Doug¬
las Co., 1921, Conklin 1302 (SMS) ; Amnicon Falls, Oct. 5,
1924, 2019 (SMS) ; Marshland, Buffalo Co., June 23, 1902,
H. (UM) ; same location, June 24, 1902, (UM & SMS) ; Bag-
ley, Grant Co., June 4, 1927, Cheney 12230 (U) ; Patch
Grove, Sept. 24, 1927, 12390 p. p. (U).
13. Jungermannia lanceolata L. Fayette, Lafayette Co.,
Dec. 1891, Cheney (NY) ; Pine River, Lincoln Co., Aug. 3,
1893, 2864 (U) ; White River, Bayfield Co., July 8, 1896,
4634, 4637 (U) ; St. Louis River, Douglas Co., July 28,
Conklin — The Hepaticae of Wisconsin.
217
1897, 7778 (U) ; Brule Swamp, Solon Springs, Douglas Co.,
Aug. 9, 1915, Allen 17 A (U) ; Winneboujou, April 9, 1911,
Conklin 1026 (SMS) ; Black River, Oct. 5, 1912, 1653
(SMS) ; Stone's Bridge, June 12, 1916, 2193 (SMS) ;
(Whealdon's), Brule River, Oct. 7, 1923, 1900 (SMS);
Bark Point, Bayfield Co., July 27, 1923, 1851 (SMS) ; Sis-
kiwit River, Oct. 5, 1924, 2374 (SMS) ; Montreal River,
Iron Co., Sept. 6, 1925, 2559 (SMS).
14. Jungermannia pumila With. Petanwells Rock, Ad¬
ams & Juneau Cos., July 19, 1894, Cheney 3702 (U) ; Apos¬
tle Islands, Ashland Co., Aug. 1902, Allen 29 A (U) ; Cop¬
per Creek, Douglas Co., Aug. 5, 1909, Conklin 551 (SMS) ;
Black River, Oct. 3, 1909, 626 (SMS) ; same location, July
6, 1921, 1303 (SMS) ; (Whealdon’s), Brule River, Oct. 7,
1923, 2015 (SMS) ; Squaw Point, Bayfield Co., Aug. 10,
1917, 1310, 1290 (SMS) ; Siskiwit River, Aug. 26, 1922,
1781 (SMS) ; Orienta Falls, Sept. 23, 1923, 1923 (SMS) ;
Bark Point, Sept. 23, 1923, 1926 (SMS) ; Montreal River,
Iron Co., Aug. 28, 1922, 2005 (SMS) ; same location, Sept.
6, 1925, 2549 (SMS) ; Trevip Ridge, Trempealeau Co., Nov.
11, 1892, R. (UM).
15. Jungermannia pumila var. rivularis Schiffner. Sis¬
kiwit River, Bayfield Co., Aug. 26, 1922, Conklin 1780
(SMS) ; Montreal River, Iron Co., Aug. 28, 1922, 1954
(SMS).
16. Jungermannia cor difolia Hook. White River, Ash¬
land Co., Cheney July 13, 1896 4778 (NY & U).
17. Jungermannia schiffneri (Loitles.) Evans. Black
River, Douglas Co., Oct. 3, 1910, and April 15, 1915, Conk -
lin 1255, 1214, 1003 (SMS) ; Amnicon Falls, Aug. 12, 1923,
1867 (SMS) ; (Wheaidon’s), Brule River, Oct. 7, 1924, 2013
(SMS) ; Squaw Point, Bayfield Co., May 10, 1917, 1321
(SMS) ; same location, Aug. 26, 1922, 1785 (SMS) ; Siski¬
wit River, Sept. 27, 1923, 1832 (SMS) ; Montreal River,
Iron Co., Aug. 27, 1922, 1819 (SMS) ; same location, Sept.
7, 1925, 2550 (SMS).
18. Jungermannia sphaerocarpa Hook. Apostle Islands,
Ashland Co., Aug. 1902, Allen 29 A (U) ; Orienta Falls,
Bayfield Co, Sept. 23, 1923, Conklin 1944 (SMS).
19. Jamesoniella autumnalis (DC.) Steph. Fayette, La¬
fayette Co, Dec. 1891, Cheney (NY) ; Lac Vieux Desert,
218 Wisconsin Academy of Sciences, Arts, and Letters.
Vilas Co., June 13, 1893, 10, 55, 47 (U) ; State Line, June
21, 1893, 469, 636 p. p. (NY & U) ; Rainbow Rapids, Oneida
Co., July 3, 1893, 1264 p. p., 1262 A (U) ; Newbold, July 10,
1893, 1757 (U) ; Granite Heights, Marathon Co., June 22,
1894, 2994 p. p., 2954, 2999 (U) ; same location, June 23,
1894, 2981 (U) ; Mason, Bayfield Co., July 7, 1896,
4615, 4632 (NY & U) ; White River, July 8, 1896,
4633 p. p. 4635, 4646 (NY & U) ; Herbster, Cranberry
River, July 7, 1897, 6895, 6896 p. p. (U) ; Port Wing,
July 12, 1897, 7185 (U) ; La Chapelle, and Brule, Bay-
field & Douglas Cos., July 17, 1897, 7425 (U) ; Mon-
teral River, Iron Co., July 21, 1896, 5173 p. p. 5175
(NY & U) ; St. Louis River, Douglas Co., July 28,
7753, (U) ; same location, July 29, 1897, 7751 (U) ;
July 31, 1897, 7912 p. p. 7916 p. p. (U) ; Apostle Islands,
Ashland Co., Aug. 1902, Allen 13, 42 A, 92 B, 64, 12 B, 14
B, 42 B, 78 A, 14 A, 43 A, 22 A, 79 A, 19 B, 64 B, (U) ;
Devil’s Lake, Sauk Co., Aug. 6, 1915 ,20 (U) ; Sand Lake,
Sawyer Co., Aug. 13, 1925, (U) ; Bass Lake, Aug. 15,
1925, (U) ; Solon Springs, Douglas Co., Aug. 1915, 19, 21,
26, 31, 33, 36, 60, 63, 68, 47, (U) ; Solon Springs, Douglas
Co., Oct. 1907, Conklin 410 (SMS) ; Copper Creek, Aug.
5, 1909, 594 (SMS) ; Superior, Nov. 14, 1909, 607 (SMS) ;
Black River, Oct. 3, 1910, 1071 (SMS) ; Brule River, April
15, 1912, 1208 A (SMS) ; Stone’s Bridge, June 12, 1916,
2200 (SMS) ; Squaw Point, Bayfield Co., Aug. 10, 1917,
1329 (SMS) ; Siskiwit Point, Oct. 5, 1924, 2437 (SMS) ;
Potosi, Grant Co., May 19, 1926, Cheney 11309 (U) ; Glen
Haven, May 11, 1927, 12153 (U) ; Bagley, June 4, 1927,
12229, 12228 (U) ; Lancaster, Oct. 24, 1927, 12538 (U) ;
Mt. Hope, Oct. 15, 1927, 12518 (U) ; Mellen, Ashland Co.,
Sect. 16, Sept. 7, 1927, Fassett & Wilson, (U) ; Loon Lake,
Mellen, Sept. 8, 1927, (U) ; Bad River Gorge, Sept. 8, 1927,
(U) ; Bear Trap Lake, Polk Co., Sept. 8, 1927 Allen (U) ;
Lake Wapogasset, Aug. 31, and Sept. 4, 1927, (U) ; Newark
Tp., Rock Co., Aug. 25, 1925, Cheney 12808 (U) .
20. Lophozia alpestris (Schleich.) Evans. Goodrich,
Marathon Co., June 25, 1894, Cheney 3194 (NY & U) ; Oak
Island, Ashland Co., Aug. 7, 1896, 6023 (NY & U) ; Wilson
Island, Aug. 6, 1896, 5895 (U) ; Squaw Point, Bayfield Co.,
Conklin — The Hepaticae of Wisconsin .
219
Aug. 10, 1917, Conklin 1275 (SMS) ; Orienta Falls, Sept.
23, 1923, 1918, 1950 (SMS).
21. Lophozia attenuata (Mart.) Dumort. Black River,
Douglas Co., July 9, 1923, Conklin 1538 (SMS).
22. Lophazia badensis (Gottsche) Schiffn. Montreal
River, Iron Co., July 20, 1896, Cheney 5149 (U) ; Lake Su¬
perior, 3 miles from Montreal River, July 22, 1896, 5198
(U) ; Black River, Douglas Co., Oct. 5, 1912, 1360 (SMS)
same location, April 15, 1915, 1255 (SMS) ; Mellen, Ash¬
land Co., Aug. 28, 1922, 1854 (SMS) ; Siskiwit Point, Bay-
field Co., July 29, 1923, 1855 (SMS) ; Squaw Point, Aug.
26, 1923, 1970 (SMS) ; Orienta Fails, Sept. 23, 1923, 1948
(SMS) ; Montreal River, Iron Co., Aug. 28, 1923, 1993,
1995 (SMS) ; Gurney Falls, Sept. 7, 1925, 2538 (SMS) ;
Mellen, Ashland Co., Aug. 23, 1925, 2642 (SMS).
23. Lophozia barbata (Schreb.) Dumort. Lac Vieux
Desert, Vilas Co., June 14, 1893, Cheney 105 (U) ; Wiscon¬
sin River, Oneida Co., June 19, 1893, 1844 (NY & U) ;
Grandmother Bull Falls, Lincoln Co., July 29, 1893, 2559
2650, 2643 (NY & U) ; Between Herbster and Port Wing,
Bayfield Co., July 8, 1897, 7040 (U) ; St. Croix Falls, Polk
Co., Nov. 1897, C. F. Baker 94 (NY) ; Devil’s Lake, Sauk
Co., July 11, 1903, (U) ; Copper Creek, Douglas Co., Aug.
29, 1907, Conklin 232 (SMS) ; Black River, Oct. 3, 1910,
1086 (SMS) ; Montreal River, Iron Co., Aug. 28, 1922,
1787, 2022, 2003 (SMS) ; Gurney Falls, Potato River, Sept.
7, 1925, 2540 (SMS) ; Cornucopia, Bayfield Co., July 26,
1922, 1998 (SMS) ; Bark Point, July 27, 1923, 1878
(SMS) ; Orienta Falls, Oct. 4, 1924, 2388 (SMS) ; Mellen,
Ashland Co., Aug. 29, 1922, 1792 (SMS) ; St. Croix Falls,
Polk Co., Sept. 4, 1927, Fassett & Wilson (U) ; Mellen Ash¬
land Co., Sept. 8, 1927, (U) ; Bad River Gorge, Mellen,
Sept. 8, 1927, (U) f St. Croix Falls, Polk Co., June 19, 1927,
Mary E. Van Wert 37H (SMS).
24. Lophozia bicrenata (Schmid.) Dumort. Amnicon
Falls, Douglas Co., Oct. 1, 1922, Conklin 1740 (SMS).
25. Lophozia excisa (Dicks.) Dumort. Siskiwit River,
Bayfield Co., Sept. 1, 1923, Conklin 1853 (SMS).
26. Lophozia guttulata (Lindb. & Arnell) Evans. Lake
Nebagamon, Douglas Co., June 1, 1913, Conklin 2063
(SMS).
220 Wisconsin Academy of Sciences, Arts, and Letters.
27. Lophozia heterocolpa (Thed.) M. A. Howe. White
River, Ashland Co., July 13, 1896, Cheney 4781 (NY & U) ;
Herbster, Bayfield Co., July 7, 1897, 6944 (U) ; Port Wing,
July 14, 1897, 7274 (U) ; Copper Creek, Douglas Co., Oct.
5, 1909, Conklin 566 (SMS) ; Black River, Oct. 5, 1912,
1346, 1580 (SMS) ; Mellen, Ashland Co., Aug. 29, 1922,
1793 (SMS) ; Montreal River, Iron Co., Aug. 28, 1922,
1830, 1973 (SMS) ; same location Sept. 6, 1925, 2578
(SMS).
28. Lophozia incisa. (Schard.) Dumort. Lac Vieux
Desert, Vilas Co., June 14, 1893, Cheney 82 (NY & U) ;
Canover, June 26, 1893, 825 (NY & U) ; Rainbow Rapids,
Oneida Co., July 3, 1893, 1247, 1254, 1255, 1380 (U) ; Cold-
water Canon, Adams Co., July 10, 1894, 3823 p. p. (U) ;
Drummond, Bayfield Co., June 26, 1896, 4303 (U) ; St.
Louis River, Douglas Co., July 31, 1897, 7908, 7911 (U) ;
Solon Springs, Douglas Co., Aug. 9, 1915, Allen 16, 56, 59
(U) ; Superior, Douglas Co., July 18, 1909, Conklin 775
(SMS) ; Copper Creek, Aug. 5, 1909, 529 (SMS) ; Black
River, Oct. 2, 1910, 1065 (SMS) ; Solon Springs, June 28,
1913, 1345 (SMS) ; Brule River, June 12, 1916, 1347
(SMS) ; Montreal River, Iron Co., Aug. 28, 1922, 1975
(SMS) ; Mellen, Ashland Co., Aug. 29, 1922, 1905 (SMS) ;
Siskiwit River, Bayfield Co., Oct. 5, 1924, 2388, 2308
(SMS) ; Barron, Barron Co., March 31, 1927, Cheney 12,-
100, 12108 p. p., 12110, 12117 (U) ; Patch Grove, Grant
Co., Sept. 24, 1927, 12463 B. p. p. (U) ; Mellen, Ashland
Co., Sect. 16, July 9, 1927, Fassett & Wilson (U).
29. Lophozia kaurini (Limpr.) Steph. Houghton Quarries,
Bayfield Co., July 28, 1896, Cheney 5435 (U) ; Port Wing,
July 14, 1897, 7274 (U) ; Oak Island, Ashland Co., July 28,
1896, 5993 (U) ; Apostle Islands, Ashland Co., Aug. 1902,
Allen 26 B., 27 D., 27 A. p. p. 39 C., 27 C. (U) ; Black
River, Douglas Co, Oct. 3, 1910, Conklin 1138 (SMS) ;
Squaw Point, Bayfield Co., Aug. 10, 1917, 1324, 1285
(SMS) ; same location, Aug. 26, 1922, 1796, 1788 (SMS) ;
Siskiwit Point, July 29, 1923, 2027 (SMS) ; Mellen, Ash¬
land Co., Aug. 29, 1922, 1739 (SMS) ; Montreal River, Iron
Co., Aug. 28, 1922, 1989 (SMS).
30. Lophozia longi flora (Nees) Schiffn. Black River,
Douglas Co., Oct. 5, 1912, Conklin 1208 (SMS).
Conklin — The TIepaticae of Wisconsin.
221
31. Lophozia muelleri (Nees) Dumort. Black River,
Douglas Co., Oct. 3, 1910, Conklin 1002 (SMS) ; Squaw
Point, Bayfield Co., Aug. 10, 1917, 1323 (SMS).
(Recent studies of a large number of specimens of
Lophozia badensis from this region show a wide range of
variability in the size of the plants, margin of the bracts,
size of the leaf cell and cell structure. The above species
may be found upon further study, to be a robust form of
L. badensis. Lophozia badensis is rather common through¬
out Douglas, Bayfield and Iron counties, where similar
habitats are found; namely: The spray wet seams and
crevasses along the trap rock gorges of the streams cutting
the South Shore Range of Lake Superior.)
32. Lophozia porphyroleuca (Nees) Schiifn. Lac Vieux
Desert, VilasCo., June 1893, Cheney 59 p. p. (U) ; same
location, June 14, 1893, 82 (U) ; State Line, June 21, 1893,
469 p. p., 470 p. p. (U) ; Rainbow Rapids, Oneida Co., July
5, 1893, 1263, 1264 (U) ; Roy’s Points, Ashland Co., Aug.
5, 1896, 5850 (NY & U) ; Apostle Islands, Ashland Co.,
Aug. 1902, Allen 51 B (U) ; Superior, Douglas Co., Sept.
6, 1909, Conklin 724 (SMS) ; Black River, Oct. 3, 1910,
1006 (SMS) ; same location, Oct. 5, 1912, 1525 (SMS).
33. Lophozia quinquedentata (Huds.) Cogn. Cooper
Creek, Douglas Co., Aug. 5, 1907, Conklin 920 (SMS) ;
Black River, Oct. 3, 1910, 1046 (SMS) ; Montreal River,
Iron Co., Aug. 27, 1923, 1815 (SMS) ; Gurney Falls, Potato
River, Sept. 7, 1925, 2557 (SMS) ; Loon Lake, Mellen,
Ashland Co., Sept. 9, 1927, Fassett & Wilson (U).
34. Lophozia ventricosa (Dicks.) Dumort. Dells Wis¬
consin River, Sauk Co., Aug. 19, 1893, L. M. Under¬
wood (NY & U) ; Bass Island, Ashland Co., Aug. 5, 1896,
Cheney 5850 (U) ; Wilson Island, Aug. 6, 1896, 5895 (NY
& U) ; Oak Island, Aug. 7, 1897, 6025 (U) ; Apostle Islands,
Ashland Co., Aug. 1902, Allen, 31 B,, 31 C., 31 A., 25 B„
(U) ; Squaw Point, Bayfield Co., Aug. 10, 1917, Conklin 1293
p. p. (SMS) ; Orienta Falls, Sept. 23, 1923, 1931, 1917
(SMS) ; Herbster, July 27, 1923, 1961 A. (SMS).
35. Sphenolobus exsectaeformis (Breidl.) Steph. Lac
Vieux Desert, Vilas Co., June 14, 1893, Cheney 82
p. p. (U) ; State Line, June 23, 1893, 631 (U) ; Houghton
Quarries, Bayfield Co., July 28, 1896, 5552 (U) ; Apostle
16
222 Wisconsin Academy of Sciences, Arts, and Letters.
Islands, Ashland Co., Aug. 1902, 31 C (U) ; Copper Creek,
Douglas Co., May 5, 1909, Conklin 529 (SMS) ; Solon
Springs, June 28, 1913, 1351 (SMS) ; Black River,
April 18, 1915, 1349 (SMS) ; Stone's Bridge, June 12, 1916,
1347 (SMS) ; Squaw Point, Bayfield Co., Aug. 10, 1917,
1276, 1294 (SMS) ; Herbster, Bark Point, Aug. 27, 1923,
1851 p. p., 1851 (SMS).
36. Sphenolobus exsectus (Schmid.) Steph. Fayette, La¬
fayette Co., Dec. 1891, Cheney 30 (NY & U) ; Dells Wis¬
consin River, Sauk Co., Aug. 19, 1893, L. M. Underwood
(NY & U) ; Coldwater Canon, Adams Co., July 30, 1894,
Cheney 3823 (NY & U) ; Houghton Quarries, Bayfield Co.,
July 25, 1896, 5493 p. p. (U) ; Wilson Island, Ashland Co.,
Aug. 6, 1896, 5895 (U) ; Oak Island, Aug. 7, 1896, 6039
(U) ; Copper Creek, Douglas Co., Aug. 5, 1909, Conklin
599 (SMS) ; Black River, Oct. 2, 1910, 1000 (SMS) ; same
location, Oct. 5, 1912, 1368, 1651, 1560 (SMS) ; Brule
River, June 22, 1923, 1955 (SMS) ; Squaw Point, Bayfield
Co., Aug. 10, 1919, 1282 (SMS) ; Siskiwit Point, Aug. 16,
1922, 1997 (SMS) ; Orienta Falls, Sept. 23, 1923, 1949 C
(SMS) ; Mellen Ashland Co., Aug. 28, 1922, 2025 (SMS) ;
Millville, Grant Co., Sept. 13, 1927, Cheney 12419 (U) ;
Patch Grove, Sept. 24, 1927, 12464 p. p. (U).
37. Sphenolobus hellerianus (Nees) Steph. State Line,
Vilas Co., June 23, 1893, Cheney 631, 636 (U) ; Tomahawk
Lake, Oneida Co., June 29, 1893, 998 (U) ; Rainbow Rapids,
July 4, 1893, 1263 (U) ; Granite Heights, Marathon Co.,
June 22, 1894, 2981, 2994, 2999 (NY & U) ; Drummond,
Bayfield Co., June 29, 1896, 4376 p. p. (U) ; Mason, July 7,
1896, 4615 (NY & U) ; White River, July 8, 1896, 4632
p. p., 4635 (U) ; Frog Bay, Aug. 7, 1896, 6045 (U) ; Port
Wing, July 12, 1897, 7185 p. p. (U) ; St. Louis River,
Douglas Co., July 31, 1897, 7912 p. p. (U) ; Black River,
Douglas Co., Oct. 5, 1912, Conklin 1573 (SMS) ; Solon
Springs, June 28, 1913, 1348 (SMS) ; Mellen, Ashland Co.,
July 9, 1927, Fassett & Wilson (U).
38. Sphenolobus michauxii (Web.) Steph. State Line,
Vilas Co., June 23, 1893, Cheney 636 (U) ; White River,
Bayfield Co., July 8, 1896, 4633, 4635 (U) ; Frog Bay,
Aug. 7, 1896, 6045 p. p. (U) ; Solon Springs, Douglas Co.,
Conklin — The Hepaticae of Wisconsin .
223
July 30, 1911, Conklin 1109 (SMS) ; Mellen, Ashland Co.,
July 9, 1927, Fassett & Wilson (U).
39. Sphenolobus scitulus (Tayl.) Steph. Montreal River,
Iron Co., Aug. 28, 1922, Conklin 1988, 2009 (SMS).
Cephaloziellaceae
1. Cephaloziella bifida (Schreb.) Schiffn. Bark Point,
Bayfield Co., July 27, 1923, Conklin 1882 (SMS) ; Amnicon
Falls, Douglas Co., Aug. 12, 1923, 1884 (SMS) ; Glen
Haven, Grant Co., June 4, 1927 Cheney 12220 (U).
2. Cephaloziella byssacea (Roth) Warnst. Black River,
Douglas Co., Oct. 3, 1913, Conklin 1252 A., 1253 (SMS) ;
Solon Springs, Aug. 7, 1915, Allen 10 (U).
3. Cephaloziella elachisla (Jack) Schiffn. Superior,
Douglas Co., July 12, 1911, Conklin 1262 (SMS) ; Black
River, Oct. 5, 1910, 1048, 1364 (SMS) ; Lake Mebagamon,
June 1924, L. R. Wilson 426 (SMS) ; Amnicon Falls, July
13, 1924, Conklin 2239, 2238 (SMS).
4. Cephaloziella hampeana (Nees) Schiffn. Wilson Is¬
land, Ashland Co., Aug. 6, 1896, Cheney 5895 (U) ; Super¬
ior, Douglas Co., Sept. 12, 1909, Conklin 717, 716, 741, 739,
745 (SMS) ; Black River, Oct. 5, 1909, 621 (SMS) ; Ori-
enta Falls, Bayfield Co., Sept. 23, 1923, 1935 (SMS) ; Gur¬
ney Falls, Iron Co., Sept. 7, 1925, 2576 (SMS).
5. Cephaloziella myriantha (Lindb.) Schiffn. Superior,
Douglas Co., Sept. 12, 1909, Conklin 748 (SMS) ; Same lo¬
cation, Oct. 9, 1921, 1337 (SMS) ; Squaw Point, Bayfield
Co., Aug. 26, 1922, 1779 (SMS).
6. Cephaloziella sullivantii (Aust.) Evans. Superior,
Douglas Co., Sept. 12, 1909, Conklin 749 (SMS) ; Lake Neb-
agamon, Sept. 22, 1912, 1203 (SMS) ; St. Croix Lake, June
26, 1913, 1242 (SMS).
Cephaloziaceae
1. Cephalozia bicuspidata (L.) Dumort. Fayette La¬
fayette Co., Dec. 1891, Cheney (NY) ; Mosinee, Marathon
Co., June 28, 1894, 3308 (NY) ; Oak Island, Ashland Co.,
Aug. 7, 1896, 6023 p. p. (U) ; Apostle Islands, Ashland Co.,
Aug. 1902, Allen 32 B, 62 (U) ; Superior, Douglas Co., July
5, 1909, Conklin 783 (SMS) ; Squaw Bay, Bayfield Co., Aug.
224 Wisconsin Academy of Sciences , Arts, and Letters.
10, 1917, 1288 (SMS) ; same location, Oct. 8, 1927 3049
(SMS).
2. Cephalozia catenulata (Huben.) Spruce. Rainbow
Rapids, Oneida Co., July 3, 1893, Cheney 1262 (U) ; Granite
Heights, Marathon Co., June 28, 1894, 2954 (U) ; Coldwa-
ter Canon, Adams Co., July 30, 1894, 3814 p. p. (U) ; Oppo¬
site Fond du Lac, Douglas Co., July 28, 1897, 7753 (U) ;
Apostle Islands, Ashland Co., Aug. 1902, Allen 92 C. (U) ;
Solon Springs, Douglas Co., Aug. 7, 1916, 36 (U) ; Black
River, Douglas Co., Oct. 3, 1910, Conklin 1148 (SMS) ; Lake
Nebagamon, July 13, 1913, 1228 (SMS).
3. Cephalozia connivens (Dicks.) Lindb. State Line,
Vilas Co., June 21, 1893, Cheney 463 (U) ; Newbold, Oneida
Co., July 8, 1893, 1531 (U) ; Gordon, Douglas Co., Aug. 25,
1907, Conklin 260 (SMS) ; Black River, Oct. 3, 1910,
1037 (SMS) ; Solon Springs, July 1, 1911, 1017 (SMS) ;
Superior, July 1909, and Oct. 27, 1912, 767 (SMS) ; Lake
Nebagamon, July 18, 1924, 1607 (SMS) ; Solon Springs,
Summer 1925, Conklin & Wilson , 2418 (SMS) ; Solon
Springs, Douglas Co., Aug. 5, 1915, Allen 54 (U).
4. Cephalozia curvifolia (Dicks.) Dumort. Dells Wis¬
consin River, Sauk Co., Aug. 19, 1893, L. M. Underwood
(NY & U) ; Lac Vieux Desert, Vilas Co., June 20, 1893,
Cheney 355 p. p. (U) ; Canover, June 24, 1893, 698 p. p.
(U) ; Daugherty Lake, Oneida Co., July 3, 1893, 1281 (U) ;
Rainbow Rapids, July 3, 1893, 1262, 1263 (U) ; Grand¬
mother Bull Falls, Lincoln Co., July 27, 1893, 2288, 2297,
2359, 2356, 2357 (U) ; Granite Heights, Marathon Co., June
22, 1894, 2954, 2981, 2999 (U) ; Mosinee, June 28, 1894,
3308 (U) ; Coldwater Canon, Adams Co., July 30, 1894,
3814 (U) ; Drummond, Bayfield Co., June 29, 1896, 4376
p. p. (U) ; Mason, July 7 and 8, 1896, 4615, 4632 (U) ;
White River at Mason, July 8, 1896, 4635 p. p., 4633 (U) ;
Herbster, July 7, 1896, 6896 (U) ; Opposite Fond du Lac,
Douglas Co., July 31, 1897, 7916 (U) ; same location, July
29, 1897, 7751 p. p. (U) ; Montreal River, Iron Co., July 22,
1896, 5244, (U) ; same location, July 31, 1896, 5175 (U) ;
Apostle Islands, Ashland Co., Aug. 1902, Allen 28 C., 79
C., 79 A., 64 B., 78 B., 79 B., 78 A., (U) ; Solon Springs,
Douglas Co., Aug. 4 to 7, 1915, 12, 79, 49 (U) ; Sand Lake,
Conklin— The Hepaticae of Wisconsin .
225
Sawyer Co., Aug. 28, 1925, (U) ; Bass Lake, Aug. 15, 1925,
(U) ; Copper Creek, Douglas Co., Aug. 29, 1907, Conklin
270 (SMS) ; Solon Springs, May 1907, 408 (SMS) ; Black
River, Oct. 5, 1912, 1570 (SMS) ; Lake Nebagamon, July
1, 1913, 1769 (SMS) ; Brule River, June 12, 1916, 2205
(SMS) ; Brule River, (Whealdon’s), Sept. 28, 1925, 2489
(SMS) ; Loon Lake, Ashland Co., Sept. 8, 1927, Fassett &
Wilson (U).
5. Cephalozia macounii Aust. Lac Vieux Desert, Vilas
Co., June 13, 1893, Cheney 60 (NY & U) ; Newbold, Oneida
Co., July 10, 1893, 1581 (U) ; Black River, Douglas Co.,
Oct. 3, 1911, Conklin 1148, 1153 (SMS) ; Lake Nebagoman,
May 10, 1913, 1228, 1229 (SMS).
6. Cephalozia media Lindb. State Line, Vilas Co., June
21, 1893, Cheney 470 (NY & U) ; Tomahawk Lake, Oneida
Co., June 28, 1893, 959, 952 (NY & U) ; Clear Lake, July
2, 1893, 1196 (NY & U) ; Rainbow Rapids, July 3, 1893,
1254, 1262, 1262 A., 1263, 1264 (U) ; Newbold, July 10,
1893, 1581 (NY & U) ; Grandmother Bull Falls, Lincoln
Co., July 27, 1893, 2288 (U) ; Mosinee, Marathon Co., June
28, 1894, 3308 p. p. (U) ; Opposite Fond du Lac, Douglas
Co., July 31, 1897, 7912, 7916 p. p. (U) ; Apostle Islands,
Ashland Co., Aug. 1902, Allen 30 C. (U) ; Brule, Douglas
Co., Aug. 13, 1915, 56 (U) ; Superior, Douglas Co., Sept. 12,
1909, Conklin 743 (SMS) ; Solon Springs, July 1, 1909, 1016
(SMS) ; Black River, July 28, 1913, 1355 (SMS) ; Lake
Nebagamon, May 29, 1923, 1465 (SMS) ; Barron, Barron
Co., March 31, 1927, Cheney 12092, 12097, 12101 p. p. 12108
p. p. (U) ; Mellen, Ashland Co., Sect. 16, July 9, 1927 Fas¬
sett & Wilson (U) ; Bad River Gorge, July 9, 1927, (U).
7. Cephalozia pleniceps (Aust.) Lindb. Tomahawk Lake,
Oneida Co., June 23, 1893, Cheney 949 p. p., 933 p. p., 952
(U) ; Rainbow Rapids, July 3, 1893, 1253 (U) ; Newbold,
July 8, 1893, 1531 p. p. (U) ; Montreal River, Iron Co., July
19, 1896, 5149 (NY & U) ; Opposite Fond du Lac, Douglas
Co., July 31, 1897, 7911 (U) ; Solon Springs, Douglas Co.,
Aug. 5, 1915, Allen 19 (U) ; Brule River, Aug. 9, 1915, 34
(U) ; Superior, Douglas Co., July 15, 1909, Conklin 216
(SMS) ; Gordon, Oct. 25, 1909, 267 (SMS) ; Black River,
Oct. 3, 1910, 1058 (SMS) ; Lake Nebagamon, June 1, 1913,
1770 (SMS) ; Brule River, June 12, 1916, 1412 (SMS) ;
226 Wisconsin Academy of Sciences, Arts, and Letters.
Brule River, (Whealdon’s), May 30, 1925, 2490 (SMS);
Mud Creek, June 9, 1925, 2473 p. p. (SMS) ; Squaw Point,
Bayfield Co., Aug. 26, 1922, 1837 (SMS) ; Siskiwit River,
Oct. 5, 1924, (SMS) ; Barron, Barron Co., March 31, 1927,
Cheney 12103 p. p. (U).
8. Odontoschisma denudatum (Mart.) Dumort. Lac
Vieux Desert, Vilas Co., June 13, 1893, Cheney 59 (U) ;
State Line, June 21, 1893, 470 (NY & U) ; Canover, June
24, 1893, 698, 729 (U) ; Rainbow Rapids, Oneida Co., July
3, 1893, 1254, 1264 p. p. (U) ; Drummond, Bayfield Co.,
June 29, 1896, 4376 p. p. (U) ; St. Louis River, Douglas Co.,
July 28, to 29, 1896, 7751, 7753 p. p. (U) ; Apostle Islands,
Ashland Co., Aug. 1902, Allen 92 C (U) ; Solon Springs,
Douglas Co., Aug. 4, 1915, 21 (U) ; Solon Springs (Cope¬
land Farm) , Douglas Co., Oct. 7, 1907, Conklin 2162 (SMS) ;
same location, July 20, 1911, 1685 (SMS) ; Stone’s Bridge,
Brule River, May 14, 1916, 1264 (SMS) ; Lake Nebagamon,
Douglas Co., July 18, 1924, Wilson 2241 (SMS).
9. Odontoschisma macounii (Aust.) Underwood. Mon¬
treal River, Iron Co., Aug. 27, 1922, Conklin 1955 (SMS) ;
Squaw Point, Bayfield Co., Oct. 8, 1927, 3032 (SMS).
10. Calyyogeia neesiana (Massal. & Carest.) K. Mull.
Tomahawk Lake, Oneida Co., June 28, 1893, Cheney 933
(U) ; Eagle River, June 28, 1893, 948, 951, 952, 949 (U) ;
Winneboujou, Douglas Co., May 7, 1911, Conklin 1157
(SMS) ; St. Croix Lake, June 28, 1913, 1241 (SMS) ; Su¬
perior, Oct. 27, 1912, 1606 (SMS) ; Lake Nebagamon, July
11, 1913, 1773 (SMS) ; Stone’s Bridge, Brule River, June
12, 1916, 2207 (SMS) ; Brule Swamp, St. Croix Lake, Doug¬
las Co., Aug. 9, 1915, Allen 22, 34, 41, 55 (U) ; Barron,
Barron Co., March 31, 1927, Cheney 12108, 12114 p. p.,
12082, 12097 p. p., 12101, 12103 (U) ; Glen Haven, Grant
Co., June 24, 1927, 12463 A. p. p. (U) ; Cornucopia, Bay-
field Co., Oct. 8, 1927, Conklin 3051 (SMS).
11. Calyyogeia trichomanis (L.) Corda. Coldwater
Canon, Adams Co., July 30, 1894, Cheney 3814 (U) ; Solon
Springs, Douglas Co., Sept. 16, 1906, Conklin 289 (SMS) ;
Copper Creek, Aug. 5, 1909, 531 (SMS) ; Superior, Sept. 2,
1909, 489 (SMS) ; same location, Nov 14, 1909, 609 (SMS) ;
Black River, Oct. 2, 1910, 1062 (SMS) ; Mellen, Ashland
Co., July 9, 1927, Fassett & Wilson (U).
Conklin — The Hepaticae of Wisconsin.
221
12. Bazzania trilobata (L.) S. F. Gray. Penokee Iron
Range, Ashland Co., Sept. 1858, I. A. Lapham (U) ; Lac
Vieux Desert, Vilas Co., June 18, 1893, Cheney 54 (U) ;
same location, June 14, 1893, 86 (U) ; June 22, 1893, 575
(U) ; Canover and Eagle Rivers, June 26, 1893, 826 (U) ;
Rainbow Rapids, Oneida Co., July 4, 1893, 1318 (U) ; Mer¬
rill, Lincoln Co., July 31, 1893, 2715 (U) ; Granite Heights,
Marathon Co., June 23, 1894, 3150 (U) ; White River, Bay-
field Co., July 8, 1896, 4624, 4628 (U) ; Sand Bay, June 22,
1897, 6314 (U) ; Bark Point, July 3, 1897, 6737 (U) ; Wil¬
son Island, Ashland Co., Aug. 6, 1896, 5909 (U) ; Brule
River, Douglas Co., July 19, 1897, 7501 (U) ; Opposite Fond
du Lac, July 31, 1897, 7948 (U) ; Apostle Islands, Ashland
Co., Aug. 1902, Allen 18, 21, 52 (U) ; Bass Lake, Sawyer
Co., Aug. 15, 1925, (U) ; Solon Springs, Douglas Co., Aug.
26, 1906, Conklin 160 (SMS) ; Gordon, Sept. 15, 1906, 318
(SMS) ; Copper Creek, Aug. 5, 1909, 565 (SMS) ; Superior,
June 3, 1909, (SMS) ; Brule River, April 9, 1910, 1031
(SMS) ; Lake Nebagamon, June 1, 1913, 1771 (SMS) ; Win-
neboujou, May 1, 1925, 2497 (SMS) ; Cussen, Bayfield Co.,
June 25, 1913, 1356 (SMS) ; Drummond, Aug. 23, 1925,
2630 (SMS) ; Barron, Barron Co., March 31, 1927, Cheney
12109, 12112, 12116 (U) ; Mellen, Ashland Co., Sept. 16,
Fassett & Wilson, July 9, 1927, (U) ;
13. Lepidozia reptans (L.) Dumort. Fayette, Lafayette
Co., Dec. 1198, Cheney 37 (NY) ; Lac Vieux Desert, Vilas
Co., June 13, 1893, 10, 49 (U) ; same location, June 14, 1893,
82 p. p., 86 (U) ; State Line, June 23, 1893, 629 (NY & U) ;
Canover, June 26, 1893, 825 p. p. (NY & U) ; Tomahawk
Lake, Oneida Co., June 28, 1893, 952 (NY & U) ; Rainbow
Rapids, June 29, 1893, 1021 (NY & U) ; same location, July
3, 1893, 1264 p. p. (U) ; Newbold, July 8, 1893, 1520 (U) ;
same location, July 10, 1893, 1581 (U) ; Grandmother Bull
Falls, Lincoln Co., July 26, 1893, 2275, (U) ; Drummond,
Bayfield Co., June 26, 1896, 4392 (U) ; Mason, July 6, 1896,
4601 (U) ; White River, July 8, 1896, 4646 (U) ; Montreal
River, Iron Co., June 19, 1896, 5040 (U) ; same location,
July 20, 1896, 5148 (U) ; Wilson Island, Ashland Co., Aug.
6, 1896, 5910 (U) ; St. Louis River, Douglas Co., July 31,
1897, 7910 (U) ; Appostle Islands, Ashland Co., Aug. 1902,
228 Wisconsin Academy of Sciences , Arts, and Letters.
Allen 31 B., 43 A., 14 B., 42 B., 14 A., (U) ; Solon Springs,
Douglas Co., Aug. 4 to 9, 1915, 35, 60 B. (U) ; Copper Creek,
Douglas Co., Aug. 5, 1909, Conklin 544 (SMS) ; Superior,
July 5, 1909, 611 (SMS) ; Black River, Oct. 3, 1909, 1056
(SMS) ; Lake Nebagamon, Sept. 3, 1911, 1149 p. p. (SMS) ;
Stone's Bridge, Brule River, June 12, 1916, 1415 (SMS) ;
Brule River, (Whealdon’s) , May 1, 1925, 2495 (SMS);
Squaw Point, Bayfield Co., June 12, 1917, 1292 (SMS) ;
Orienta Falls, Sept. 23, 1923, 1951 (SMS) ; Herbster, June
23, 1923, 1961 (SMS) ; Siskiwit Point, Oct. 5, 1924, 2373
(SMS) ; Montreal River, Iron Co., Sept. 6, 1925, (SMS) ;
Barron, Barron Co., March 31, 1927, Cheney 12114 (U) ;
Patch Grove, Grant Co., Sept., 24, 1927, 12463 A., 12464
p. p. (U).
Ptilidiaceae
1. Ptilidium ciliare (L.) Nees. Milwaukee, Milwaukee
Co., July 1862 (U) ; Lac Vieux Desert, Vilas Co., June 13,
1893, Cheney 47 (U) ; Tomahawk, Lincoln Co., July 21,
1893, 2127 (U) ; Copper Creek, Douglas Co., Aug. 5, 1909,
Conklin 588 (SMS) ; Black River, Oct. 5, 1909, 613 (SMS).
Black River, Oct. 5, 1909, 613 (SMS).
2. Ptilidium pulcherrium (Web.) Hampe. Milwaukee,
Milwaukee Co., July 1862, I. A. Lapham (U) ; Fayette, La¬
fayette Co., Dec. 1891, Cheney 10, 19, 31, 38, (NY) ; Lake
Mills, Jefferson Co., March 30, 1891, “O” (U) ; Middleton,
Dane Co., Oct. 21, 1892, Cheney & True (U) ; Lac Vieux
Desert, Vilas Co., June 13, 1893, Cheney 15, 20, 25, 47, 48
(U) ; same location, June 14, 1893, 89 (U) ; June 1893, 561
(NY) ; Canover, June 24, 1893, 725 (NY & U) ; Newbold,
Oneida Co., July 10, 1893, 1560, 1757 p. p. (NY. & U) ;
Whirlpool Rapids, July 19, 1893, 6817 (U) ; Tomahawk
Lake, Oneida & Lincoln Cos., July 21, 1893, 2116 (U) ;
Tomahawk, Lincoln Co., July 25, 1893, 2192, 2194 (U) ;
Grandmother Bull Falls, July 26, 1893, 2320 (NY. & U) ;
Granite Heights, Marathon Co., June 23, 1894, 2981, 2994,
2998, 2999 (U) ; Mason, Bayfield Co., July 7, 1896, 4615
p. p. (U) ; White River, July 8, 1896, 4635, 4639 (U) ;
Drummond, July 22, 1896, 4157 (U) ; Siskiwit Bay, June
29, 1897, 6575 (U) ; Port Wing, July 12, 1897, 7185 (U) ;
Conklin — The Hepaticae of Wisconsin.
229
Orienta, July 15, 1897, 7358 (U) ; Herbster, July 7, 1897,
6928 A., 6932, (U) ; Apostle Islands, Ashland Co., Aug.
1902, Allen 1, 9, 12 A., 13, 19, 20, 31 C., 51, 79 A., 90, 91
B., 51 B. (U) ; Solon Springs, Douglas Co., Aug. 4 to 7,
1915, 8, 10, 13, 30, 44, 60 A., 65, 68 (U) ; Solen Springs,
Douglas Co., Aug. 26, 1906, Conklin 166 (SMS) ; Gordon,
Sept. 15, 1906, 917 (SMS) ; Superior, Sept. 12, 1909, 730
(SMS) ; Lake Nebagamon, Sept. 3, 1911, 1153 p. p. (SMS) ;
Brule River, April 15, 1912, 1007 (SMS) ; Head St. Croix
Lake, June 29, 1924, 2230 (SMS) ; Squaw Point, Bayfield
Co., Aug. 10, 1917, 1335 (SMS) ; Siskiwit Point, Aug. 26,
1922, 1995 (SMS) ; Bark Point, July 27, 1923, 1878 (SMS) ;
Mt. Hope, Grant Co., Oct. 15, 1927, Cheney 12519, 12520
(U) ; Barron, Barron Co., April 13, 1926, 11113 (U) ; same
location, April 2, 1927, 12130 (U) ; Mellen, Ashland Co.,
Sept. 16, Sept. 4, 1927, Fassett & Wilson (U) ; St. Croix
Falls, Polk Co., Sept, 4, 1927, p. p. (U) ; Loon Lake, Ash¬
land Co., Sept. 8, 1927, (U) ; Clinton, Rock Co., July 18,
1928, Cheney 12603 (U).
3. Blepharostoma trichophyllum (L.) Dumort. Fayette,
Lafayette Co., Dec. 1891, Cheney 35, 39 (NY) ; State Line,
Vilas Co., June 21, 1893, 469 p. p., 470 p. p. (NY & U) ; Lac
Vieux Desert, June 22, 1893, 576 (NY & U) ; same location,
June 23, 1893, 636 p. p. (NY & U) ; Rainbow Rapids,
Oneida Co., July 4, 1893, 1264 (NY & U) ; Newbold, July 8,
1893, 1522 (NY & U) ; Montreal River, Iron Co., July 19,
1898, 5040 p. p. (NY & U) ; Shore of Lake Superior, July
21, 1896, 5175 p. p. (NY & U) ; Oak Island, Ashland Co.,
Aug. 7, 1896, 5992, 6023 p. p. (NY & U) ; Drummond, Bay-
field Co., June 29, 1898, 4376 p. p. (NY & U) ; Herbster,
July 7, 1897, 6907 p. p., 6946 (U) ; Port Wing, July 8, 1897,
7017 p. p. (U) ; same location, July 12, 1897, 7197, (NY) ;
Orienta, July 14, 1897, 7278, 7274 (NY) ; Copper Creek,
Douglas Co., Conklin , Aug. 5, 1909, 595 (SMS) ; Superior
Sept. 12, 1912, 734 (SMS) ; Winneboujou, April 20, 1911,
1028 (SMS) ; Black River, Oct. 5, 1912, 1225 (SMS) ; Brule
River, June 12, 1916, 1419 (SMS) ; Squaw Bay, Bayfield
Co., Aug. 10, 1917, 1280 (SMS) ; Bark Point, July 27 1923,
1957 (SMS) ; Siskiwit River, Sept. 1, 1923, 1874 (SMS) ;
Montreal River, Iron Co., Aug. 27, 1923, 1832 (SMS) ; same
230 Wisconsin Academy of Sciences, Arts, and Letters.
location, Sept. 6, 1925, 2558 (SMS) ; Apostle Islands, Ash¬
land Co., Aug. 1902, Allen 29 B., 27 A., 30 A., 42 B., 30 B.,
30C., 22 A., 27 C., (U).
4. Trichocolea tomentella (Ehrh.) Dumort. Rainbow
Rapids, Oneida Co., July 1, 1893, Cheney 1165, 1168 (NY
& U) ; Coldwater Canon, Adams Co., July 30, 1894, 3817
(U) ; Montreal River, Iron Co., July 21, 1896, 5165 (U) ; St.
Louis River, Douglas Co., July 31, 1897, 7949 (U) ; Winne-
boujou, Douglas Co., May 20, 1911, Conklin 1198 (SMS) ;
Brule River, June 12, 1916, 2197 (SMS) ; Hall's Swamp,
Brule River, May 1, 1925, 2498 (SMS) ; Barron, Barron
Co., March 31, 1927, Cheney 12079 (U) ; Mellen, Ashland
Co., July 9, 1927, Fassett & Wilson (U).
SCAPANIACEAE
1. Diplophyllum apiculatum (Evans) Steph. Orienta
Falls, Bayfield Co., Sept. 23, 1923, Conklin 1941 (SMS).
2. Diplophyllum gymnostomophilum Kaal. Black River,
Douglas Co., Oct. 5, 1912, Conklin 959, 1361, 1527, 1591
(SMS) ; Gurney Falls, Potato River, Iron Co., Sept. 7, 1925,
2576 (SMS).
3. Scapania apiculata Spruce. Cranberry River, Bay-
field Co., July 7, 1897, Cheney 6882 p. p. (U) ; Herbster,
July 8, 1897, 7008 (U) ; St. Louis River, Douglas Co., July
31, 1897, 7955 (U) ; St. Louis Bay, Superior, Douglas Co.,
Aug. 1905, Conklin 382 (SMS) ; Copper Creek, Aug. 5, 1909,
530 (SMS) ; Wentworth, Oct. 19, 1910, 1004 (SMS) ; Black
River, Oct. 5, 1912, 1573, 1625 (SMS) ; Stone's Bridge,
Brule River, July 12, 1916, 2223 (SMS) ; Squaw Point,
Bayfield Co, Oct. 8, 1927, 3049, 3050 (SMS).
4. Scapania cuspiduligera (Nees) K. Mull. Squaw
Point, Bayfield Co, Aug. 10, 1917, Conklin 1301 (SMS) ;
same location, Aug. 26, 1922, 1788 (SMS) ; Bark Point,
Aug. 3, 1924, 2277 (SMS) ; Squaw Point, Oct. 4, 1924, 2361,
2429 (SMS).
5. Scapania curta (Mart.) Dumort. Montreal River,
Iron Co, July 10, 1896, Cheney 5115 (NY & U) ; Houghton
Quarries, Bayfield Co, July 25, 1896, 5444, 5493 (NY & U) ;
Oak Island, Ashland Co, Aug. 7, 1896, 6033 (NY & U) ;
Copper Creek, Douglas Co, Oct. 1907, Conklin 390 (SMS) ;
Conklin — The Hepaticae of Wisconsin.
231
Superior, Sept. 12, 1909, 738, 745 (SMS) ; Black River, Oct.
5, 1909, 635, 1074 (SMS) ; Squaw Point, Bayfield Co., Aug.
10, 1917, 1283, 1284, 1279, 1295, 1320, 1219, 1297 p. p., 1297
(SMS) ; same location, Aug. 25, 1924, 1783 (SMS) ; Oct.
4, 1924, 2385, 2435 (SMS) ; Siskiwit Point, July 29, 1923,
1886 (SMS) ; same location, Sept. 1, 1923, 1909 (SMS) ;
Orienta Falls, Sept. 23, 1923, 1934, 1847 (SMS) ; Herbster,
July 27, 1923, 1960 (SMS) ; Montreal River, Iron Co., Aug.
28, 1922, 1811, (SMS) ; Mellen, Ashland Co., Aug. 29, 1922,
1792 p. p. (SMS) ; Patch Grove, Grant Co., Sept. 24, 1927,
Cheney 12464 (U) ; Mellen, Ashland Co., Sept. 7, 1927, Fas -
sett & Wilson (U) .
(Dr. H. Buch, of Helsingfors, Finland has recently
studied S. curta and has split the composite species S. curta
into several species. Some of these specimens in Wiscon¬
sin he has referred to his new S. mucronata Buch. The
only specimen from Wisconsin sent him by Dr. A. W. Evans,
which he referred to S. curta was No. 1079, (duplicate of
1074 above.) Until Dr. Biich’s monograph is available for
study all specimens are referred under S. curta in this re¬
port (G. H. C.).
6. Scapania dentata Dumort. Squaw Point, Bayfield
Co., July 29, 1923, Conklin 1848 (SMS) ; Siskiwit Point,
Oct. 5, 1924, 2362, 2442 (SMS).
7. Scapania glaucocephala (Tayl.) Aust. Lake Su¬
perior, near Montreal River, Iron Co., July 21, 1896, Cheney
5191 (NY & U) ; Solon Springs, Douglas Co., Aug. 7, 1915,
Allen 11 (U) ; Black River, Douglas Co., Oct. 5, 1909, Conk¬
lin 620 (SMS) ; same location, Oct. 5, 1912, 1365 (SMS) ;
Solon Springs, Aug. 6, 1911, 1112 (SMS) ; Cussen, Bayfield
Co., July 25, 1915, 1244 (SMS) ; Brule River, (Whealdon’s)
Douglas Co., Aug. 24, 1924, Wilson 2282 (SMS).
8. Scapania irrigua (Nees) Dumort. Superior, Doug¬
las Co., Oct. 20, 1908, Conklin 1019 (SMS) ; same location,
Sept. 12, 1909, 735, 741 (SMS) ; Brule River, Stone’s
Bridge, June 12, 1916, 1415 (SMS) ; Mud Creek, June 8,
1925, 2472 (SMS) ; Lake Nebagamon, Douglas Co., June
1924, Nancy Bond (SMS) ; Mellen, Ashland Co., July 9,
1927, Fassett & Wilson (U) ; Siskiwit Point, Bayfield Co.,
Oct. 9, 1927, Conklin 3051 (SMS) ;
232 Wisconsin Academy of Sciences , Arts, and Letters.
9. Scapania nemorosa (L.) Dumort. Mosinee, Mara¬
thon Co., July 29, 1894, Cheney 3320, 3308 (NY & U) ; Cold-
water Canon, Adams Co., July 30, 1894, 3814 p. p. (NY &
U) ; Witch’s Gulch, July 30, 1894, 3897 (U) ; Oak Island,
Ashland Co., Aug. 7, 1928, 6033 p. p., 6039 (U) ; Devils
Lake, Sauk Co., Aug. 16, 1917, Allen 28 (U) ; Black River,
Douglas Co., Oct. 5, 1909, Conklin 643 (SMS) ; Amnicon
Falls, Oct. 1, 1922, 1741 (SMS) ; Squaw Point, Bayfield Co.,
Aug. 10, 1917, 1318 (SMS) ; same location, Aug. 27, 1923,
1836 (SMS) ; Siskiwit Point, July 29, 1923, 1885 (SMS) ;
Orienta Falls, Sept. 23, 1923, 1938 (SMS) ; Bagley, Grant
Co., June 24, 1927, Cheney 12310, 12311 (U) ; Mellen, Ash¬
land Co., Sept. 8, 1927, Fassett & Wilson (U) ;
10. Scapania paludicola Loeske & K. Mull. Drummond,
Bayfield Co., June 29, 1898, Cheney 4376 p. p., 4377 (NY &
U) ; Lac Vieux Desert, Vilas Co., Aug. 29, 1917, Allen 58
(U) ; Black River, Douglas Co., Oct. 3, 1910, Conklin 1009
(SMS) ; same location, Oct. 21, 1915, 1251, 1258 p. p
(SMS) ; Amnicon Lake, July 13, 1924, 2237 (SMS).
11. Scapania subalpina (Nees) Dumort. Montreal
River, Iron Co., July 20, 1896, Cheney 5115 p. p. (U) ; Oak
Island, Ashland Co., Aug. 7, 1896, 6033 p. p. (U) ; Apostle
Islands, Ashland Co., Aug. 1902, Allen 27 B, 29 B, 30 A,
32 A, 32 B, 32, 33, 34, 35 B, 72, 73 (U) ; Black River, Doug¬
las Co., Oct. 5, 1912, Conklin 1206 (SMS) ; Squaw Point,
Bayfield Co., Aug. 10, 1917, 1281, 1316, 1297 p. p. (SMS) ;
same location, July 29, 1923, 1986 (SMS) ; Squaw Bay,
Aug. 26, 1922, 1814, 1916 (SMS) ; Siskiwit Point, July 29,
1923, 1892, 1864 (SMS) ; Orienta Falls, Sept. 23, 1893, 1937
(SMS) ; Herbster, Aug. 27, 1923, 1851 (SMS) ; Montreal
River, Iron Co., Sept. 6, 1925, 2549 (U).
12. Scapania undulata (L.) Dumort. Squaw Point,
Bayfield Co., Aug. 10, 1919, Conklin 1317 (SMS).
Radulaceae
1. Radula complanata (L.) Dumort. Rainbow Rapids,
Oneida Co., June 29, 1893, Cheney 1026 (NY & U) ; Grand¬
mother Bull Falls, Lincoln Co., July 29, 1893, 2663 (NY &
U) ; White River, Bayfield Co., July 8, 1896, 4651 (U) ;
Devils Lake, Sauk Co., July 11, 1903, (collector unrecorded)
Conklin — The Hepaticae of Wisconsin,
233
(U) ; Gordon, Douglas Co., Sept. 10, 1906 Conklin 156
(SMS) ; Copper Creek, Aug. 5, 1909, 573 (SMS) ; Black
River, Oct. 3, 1910, 1226 (SMS) ; Brule River, July 22,
1923, 1902 (SMS) ; Lyman Lake, Aug. 20, 1923, 1870
(SMS) ; Siskiwit Point, Bayfield Co., July 29, 1923, 1887
(SMS) ; Montreal River, Iron Co., Sept. 6, 1925, 2552
(SMS) ; Bad River Gorge, Ashland Co., Sept. 8, 1927, Fas-
sett & Wilson (U) ; Wapogasset, Polk Co., Aug. 31, 1927,
Allen (U) ; St. Croix Falls, Polk Co., June 6, 1927, Mary
Van Wert, H. 37 p. p. (SMS) .
2. Radula obconica Sullivant. Goodrich to Granite
Heights, Marathon Co., June 25, 1894, Cheney 3192 (U) ;
White River, Bayfield Co., July 8, 1896, 4629, 4651, 4651
p. p. (U) ; Herbster, July 7, 1897, 6893 (U) ; Wentworth,
Douglas Co., Oct. 19, 1910, Conklin 963 (SMS) ; Stone's
Bridge, June 12, 1916, 2204 (SMS) ; Brule River, July 22,
1923, 1965 (SMS) ; Squaw Point, Bayfield Co., Aug. 10,
1917, 1334 (SMS) ; Cornucopia, April 29, 1923, 1996
(SMS) ; Barron, Barron Co., March 31, 1927, Cheney
12102, 12111 (U) ; Mellen, Ashland Co., Sect. 16, Sept. 7,
1927, Fassett & Wilson (U) ; Bad River Gorge, Sept. 8,
1927, (U).
PORELLACEAE
1. Porella pinnata L. Wisconsin River, Oneida Co.,
July 19, 1893, Cheney 1837, 1839 (U) ; Granite Heights,
Marathon Co., June 22, 1894, 2958 (NY & U) ; Witch's
Gulch, Adams Co., July 31, 1894, 3883 (NY & U) ; St. Croix
Falls, Polk Co., July 14, 1890, H. (UM).
2. Porella platyphylla (L.) Lindb. Penokee, Iron
Range, Ashland Co., I. A. Lapham (U) ; Madison, Dane Co.,
April 1886, A. B. Seymour (NY) ; Trempealeau Co., May
17, 1890, H (NY) ; Baraboo, Sauk Co., May 1, 1891, True
9, 12, (NY) ; Vilas Woods, Dane Co., May 3, 1902, (No col¬
lectors given), (U) ; Devils Lake, Sauk Co., July 11, 1903,
(U) ; Eagle Heights, Dane Co., April 22, 1902, (U) ; Blue
Mounds, Iowa Co., Aug. 1, 1903, (U) ; Northern Wiscon¬
sin, April 1907, /. F. Brenckle 1100, 1097 (SMS) ; Madison,
Dane Co., Aug. 25, 1890, Cheney (NY) ; Fayette, Lafayette
Co., Dec. 1891, 24, 26, 27, 43 (NY) ; Lac Vieux Desert, Vilas
234 Wisconsin Academy of Sciences , Arts, and Letters.
Co., June 20, 1893, 344 (NY & U) ; Drummond, Bayfield
Co., June 24, 1896, 4179 (U) ; Houghton Quarries, July 25,
1896, 5705 (U) ; Solon Springs, Douglas Co., Sept. 5, 1915,
Allen 61 (U) ; Apostle Islands, Ashland Co., Aug. 1902, 17
(U) ; Bass Lake, Sawyer Co., Aug. 15, 1925, (U) ; Parfrey’s
Glen, Sauk Co., May 23, 1926, (U) ; Gordon, Douglas Co.,
Sept. 15, 1906, Conklin 329 (SMS) ; Solon Springs, May
1907, 1023 (SMS) ; Superior, Dec. 27, 1908, 324 (SMS) ;
Copper Creek, Aug. 5, 1909, 534 (SMS) ; Black River, Oct.
3, 1910, 1068 (SMS) ; Lake Nebagamon, Sept. 3, 1911, 1149
p. p., 1146 (SMS); Amnicon Lake, May 6, 1923, 1754
(SMS) ; Lyman Lake, Aug. 20, 1923, 1872 (SMS) ; Cornu¬
copia, Bayfield Co., Aug. 26, 1922, 1996 (SMS) ; Solon
Springs, Douglas Co., June 22, 1924, Wilson 2360 (SMS) ;
Potosi, Grant Co., May 4, 1926, Cheney 11163 (U) ; same
location, May 19, 1926, 11296 (U) ; Barron, Barron Co.,
March 31, 1927, 12106 (U) ; Glen Haven, Grant Co., June
10, 1927, 12258 (U) ; same location, May 18, 1927, 12181
(U) ; Turvip Rock, Trempealeau Co., May 17, 1890, H
(UM) ; Insor Creek, May 17, 1890, (UM) ; Lake Wapogas-
set, Polk Co., Aug. 31 and Sept. 4, 1927, Allen (U) ; Turtle
Lake, Rock Co., July 7, 1928, Cheney 12755, 12757, 12756
(U) ; Bradford Tp., July 31, 1928, 12723 (U).
3. Porella platyphylloidea (Schwein.) Lindb. Madison,
Dane Co., Dec. 19, 1891, Cheney & True (U) ; Madison,
Dane Co., July 8, 1903, (No collector given), (U) ; Madison,
Dane Co., I. C. Carr (U) ; Rainbow Rapids, Oneida Co.,
July 4, 1893, Cheney 1327, 1325 (NY & U) ; Montreal River,
Iron Co., July 21, 1896, 5166, 5195 (U) ; La Pointe, Made¬
line Island, Ashland Co., July 30, 1896, 5705 (U) ; Drum¬
mond, Bayfield Co., June 24, 1896, 4164 (U) ; Mason, July
8, 1896, 4545 (U) ; Herbster, July 5, 1897, 6823, 6892 (U) ;
Port Wing, July 8, 1897, 7032 (U) ; same location, July 12,
1897, 7186 (U) ; Arena, Iowa Co., Sept. 3, 1922, Allen (U) ;
Green Lake, Green Co., July 16, 1922, (U) ; Gordon, Doug¬
las Co., Sept. 15, 1902, Conklin 157, 329 (SMS) ; St. Croix
Falls, Polk Co., Aug. 7, 1925, 2485 (SMS) ; Lodi, Columbia
Co., Oct. 4, 1926, Wilson 4 (SMS) ; Barron, Barron Co.,
Sept. 1, 1925, Cheney 9761 (U) ; same location, March 31,
1927, 12111 p. p. (U) ; Newark Tp., Rock Co., Aug. 30,
1928, Cheney 12810 (U).
Conklin — The Hepaticae of Wisconsin.
235
Lejeuneaceae
1. Lejeunea cavifolia (Ehrh.) Lindb. Grandmother
Bull Falls, Lincoln Co., July 29, 1893, Cheriey 2497, 2558
(NY & U) ; Montreal River, Iron Co., Aug. 27, 1922, Conk¬
lin 1801 (SMS).
2. Cololejeunea biddlecomiae (Aust.) Evans. Mason,
Bayfield Co., July 7, 1896, Cheney 4645 (NY & U) ; Copper
Creek, Douglas Co., Aug. 5, 1909, Conklin 384, 569 (SMS) ;
Black River, Oct. 3, 1910, 1055 (SMS) ; St. Croix Lake,
June 28, 1913, 1339 (SMS) ; Amnicon Falls, Aug. 12, 1922,
1868 (SMS) ; Cornucopia, Bayfield Co., Aug. 26, 1922, 1999
(SMS) ; Montreal River, Iron Co., Sept. 26, 1925, 2554
(SMS).
3. Frullania asagrayana Mont. Rainbow Rapids,
Oneida Co., July 4, 1893, Cheney 1314 (U) ; Grandmother
Bull Falls, Lincoln Co., July 27, 1893, 2353, 2365 (U) ;
Granite Heights, Marathon Co., June 22, 1894, 2964, 2995
(U) ; Montreal River, Iron Co., July 19, 1896, 5044 (U) ;
Odanah Indian Reservation, July 22, 1896, 5235, 5238 (U) ;
Fayette, Lafayette Co., Dec. 1899, 532 (NY & U) ; Potosi,
Grant Co., May 29, 1926, 11388 (U) ; Mt. Hope, Oct. 15,
1927, 12521 (U) ; Stone’s Bridge, Brule River, Douglas Co.,
June 27, 1926, Conklin 2646 (SMS) ; Mellen, Ashland Co.,
Sect. 16, July 9, 1927, Fassett & Wilson (U).
4. Frullania bolanderi Aust. Lake Nebagamon, Doug¬
las Co., Sept. 3, 1911, Conklin 1117 (SMS) ; Amnicon Lake,
May 6, 1923, 1753 (SMS).
5. Frullania brittoniae Evans. Madison, Dane Co., Dec.
19, 1891, Cheney & True (U) ; same location, Dec. 1899, 28
(NY & U) ; Grandmother Bull Falls, Lincoln Co., July 27,
1893, Cheney 2347 (U) ; Mason, Bayfield Co., July 6, 1896,
4558 (NY & U) ; Herbster, July 7, 1897, 6928 (U) ; Amni¬
con River, Douglas Co., July 20, 1897, 7558 (U) ; Gordon,
Douglas Co., Sept. 15, 1906, Conklin 351, 362 (SMS) ; Solon
Springs, Oct. 7, 1907, 362, 481 (SMS) ; Lake Nebagamon,
Sept. 3, 1911, 1148 (SMS) ; Barron, Barron Co., March 31,
1927, Cheney 12094, 12113 (U) ; Glen Haven, Grant Co.,
May 16, 1927, 12166 (U) ; same location, May 18, 1927,
12179 (U) ; Bagley, Grant Co., May 30, 1927, 12203 (U) ;
236 Wisconsin Academy of Sciences, Arts, and Letters.
Bradford Tp., Rock Co., July 2, 1928, Cheney 12728 (U) ;
same location, July 27, 1928, 12713 (U), Aug. 8, 1928, 12761
(U) ; Clinton, Sept. 11, 1928, 12784 (U) ; Afton, Oct. 26,
1928, 12870 (U).
6. Frullania eboracensis Gottsche. Madison, Dane Co.,
Dec. 9, 1891, Cheney & True (U) ; Madison, Dane Co., Dec.
1891, Cheney 20 A, 20 B, 22 (NY & U) ; Fayette, Lafayette
Co., Dec. 1891, 2, 4, 20, 21, 25, 33, 34, 41, (NY & U) ; Lac
Vieux Desert, Vilas Co., June 13, 1893, 1 (U) ; same loca¬
tion, June 20, 1893, 35, 320, 325, 342 (U) ; Rhinelander,
Oneida Co., July 18, 1893, 1819 (U) ; Rainbow Rapids, July
3, 1893, 1265 (U) ; Grandmother Bull Falls, Lincoln Co.,
July 26, 1893, 2276, 2268 (U) ; same location, July 27, 1893,
2437 (NY & U) ; Montreal River, Iron Co., July 20, 1896,
5127 (NY & U) ; Wilson Island, Ashland Co., Aug. 6, 1896,
5925 (U) ; Drummond, Bayfield Co., June 24, 1896, 4184
(U) ; Bark Point, July 5, 1897, 6732 (U) ; Herbster, July
7, 1897, 6902 (U) ; Cranberry River, July 7, 1897, 6936
(U) ; Amnicon River, Douglas Co., July 30, 1897, 7758 p. p.
(U) ; St. Louis River, July 28, 1897, 7786 (U) ; Devils Lake,
Sauk Co., July 11, 1903, (No collectors given), (U) ; Blue
Mounds, Iowa Co., Aug. 1, 1903, (U) ; Apostle Islands, Ash¬
land Co., Aug. 1902, Allen 89 (U) ; Arena, Iowa Co., Sept.
2, 1922, (U) ; Gordon, Douglas Co., Sept. 15, 1906, Conklin
366 (SMS) ; Copper Creek, Sept. 1, 1907, 506 (SMS) ;
Black River, Oct. 5, 1909, 629 (SMS) ; Superior, Aug. 1911,
348 (SMS) ; Lake Nebagamon, Sept. 3, 1911, 1153 (SMS) ;
Lyman Lake, Aug. 20, 1923, 1871 (SMS) ; Siskiwit Point,
Bayfield Co., Aug. 26, 1922, 1994 (SMS) ; Montreal River,
Iron Co., Sept. 26, 1925, 2555 (SMS) ; Solon Springs, Doug¬
las Co., June 29, 1924, Wilson 2228 (SMS) ; Barron, Bar¬
ron Co., March 31, 1927, Cheney 12087, 12089, 12090,
12091, 12118, 12083, 12084, 12085, 12086 (U) ; same loca¬
tion, April 1, 1925, 9720 (U) ; Glen Haven, Grant Co., May
16, 1927, 12165, 12167, 12169 (U) ; Bagley, June 3, 1927,
12224 (U) ; St. Croix Falls, Polk Co., July 12, 1890, H
(UM) ; Trempealeau Mt., Trempealeau Co., July 31, 1890,
(UM) ; Bad River Gorge, Ashland Co., Sept. 8, 1927, Fas-
sett & Wilson (U) ; Wapogasset, Polk Co., Sept. 3, 1927,
Allen (U) ; Bear Trap Lake, Sept. 8, 1927, (U) ; Midland,
Conklin — The Hepaticae of Wisconsin .
237
Buffalo Co., June 1903, H. (UM). Clinton, Rock Co., July
20, 1928, Cheney 12609 (U) ; same location, July 25, 1928,
1225 (U), July 26, 1928, 12626 (U) ; Bradford Tp., Aug.
8, 1928, 12762 (U) ; Avon, Aug. 19, 1928, 12795, 12797,
12796 (U) ; Newark Tp., Oct. 13, 1928, 12841 (U) ; same
location, Oct. 18, 1928, 12844, 12846, 12847 (U).
7. Frullania inflata Lehm. & Lindenb. Stevens Point,
Portage Co., June 30, 1894, Cheney 3358 (U) ; Herbster,
Bayfield Co., July 4, 1897, 6878 (U) ; Bridgeport, Grant Co.,
July 19, 1927, 12345 (U) ; Bridgeport, Grant Co., July 19,
1927, Cheney 12345 (U).
8. Frullania selwyniana Pears. St. Croix Lake, Solon
Springs, Douglas Co., Jan. 28, 1913, Conklin 1239 (SMS) ;
Stone's Bridge, Brule River, May 30, 1925, 1417 (SMS).
9. Frullania riparia Hampe. Potosi, Grant Co., May 2,
1926, Cheney 11155, 11342 (U) ; same location, May 4,
1926, 11162, 11164 (U) ; May 6, 1926, 11186 (U) ; May 10,
1926, 11222 (U) ; May 26, 11342 (U) ; Glen Haven, Grant
Co., Aug. 2, 1926, 11575, 11576 (U) ; same location, Aug.
21, 1926, 11690 (U) ; May 18, 1927, 12180, 12182 (U) ;
Bagley, June 3, 1927, 12225, 12226 (U) ; same location,
June 16, 1927, 12273 (IT) ; Millville, Sept. 13, 1927, 12420
(U) ; St. Croix Falls, Polk Co., July 12, 1890 H. (UM) ;
Turtle Lake, Rock Co., July 7, 1928, Cheney 12759 (U) ;
same location, July 24, 1928, 12703 (U) .
10. Frullania oakesiana Aust. Wentworth, Douglas
Co., Oct. 19, 1910, Conklin 2155 (SMS).
ANTHOCEROTALES
Anthrocerotaceae
1. Notothylas orbicularis (Schwein.) Suliivant. Madi¬
son, Dane Co., Aug. 10, 1892, Cheney & True.
2. Anthoceros laevis L. Marshland, Buffalo Co., Aug.
18, 1890, H. (NY & SMS) ; Madison, Dane Co., Oct. 8, 1892,
Cheney & True (U) ; Dells Wisconsin River, Sauk Co.,
American Association for the Advancement of Science
Meeting, Aug. 19, 1893, (U) ; Argyle, Lafayette Co., Aug. 6,
1892, Cheney (U) ; Merrill, Lincoln Co., July 31, 1893, 2725
16
238 Wisconsin Academy of Sciences , Arts , and Letters.
(U) ; same location, Aug. 1, 1893, 2822 (U) ; Petanwells
and Germantown, Adams & Juneau Cos., July 19, 1894,
3704 (U) ; Coldwater Canon, Aug. 1, 1894, 3916 (U) ; Su¬
perior, Douglas Co., Oct. 1904, Conklin 79 (SMS) ; Black
River, Manitou Falls, Oct. 1906, 459 (SMS) ; Brule River,
Oct. 7, 1923, 2011 (SMS) ; Montreal River, Iron Co., Aug.
27, 1922, 1803 (SMS) ; Cornucopia, Bayfield Co., Aug. 16,
1916, 2371 (SMS) ; Maiden Rock, Pierce Co., Aug. 16, 1916,
Allen 43 (U) ; Messinger Springs, Sauk Co., Aug. 16, 1917,
9 (U) ; Trempealeau Mt., Trempealeau Co., May 17, 1890,
H. (U) ; Cassville, Grant Co., July 7, 1927, Cheney 12337
(U) ; Bagley, May 30, 1927, 12204 (U) ; Wapogasset, Polk
Co., Aug. 28, 1927, Allen (U) ; Dodge, Trempealeau Co.,
June 21, 1902, H. (UM) ; Trempealeau Mt., Trempealeau
Co., Nov. 11, 1893 H. (det. L. M. Underwood). (Specimen
not seen. Reported XXVI. Bull. No. 9 Holzenger. Nat.
Hist. Survey of Minn.)
3. Anthoceros macounii M. A. Howe. Grandmother
Bull Falls, Lincoln Co., July 26, 1893, Cheney 2265 (U).
4. Anthoceros punctatus L. Arena, Iowa Co., Sept. 2,
1922, Allen 12 (U).
In the following pages the species found in each county
are briefly enumerated without authorities.
Adams County
Reboulia hemisphaerica, Conocephalum conicum, March-
antia polymorpha, Preissia quadrata, Metzgeria conju-
gata, Pellia epiphylla, Fossombronia foveolata, Chiloscy-
phus rivularis, Geocalyx graveolens, Harpanthus scutatus,
Jungermannia pumila, Lophocolea heterophylla, Lophozia
incisa, Nardia hyalina, Plagiochila asplenioides, Sphenolo-
bus exsectus, Calypogeia trichomanis, Cephalozia catenu-
lata, C. curvifolia, Trichocolea tomentella, Scapania nemo-
rosa, Porella pinnata, Anthoceros laevis.
Ashland County
Conocephalum conicum, Preissia quadrata, Marchantia
polymorpha, Riccardia latifrons, R. multifida, R. pinguis,
Blasia pusilla, Fossombronia foveolata, Pellia epiphylla,
Conklin — The Hepaticae of Wisconsin.
239
Chilosey pirns pallescens, Geocalyx graveolens, Jamesoniella
autumnalis, Jungermannia cordifolia, J. pumila, J. sphaero-
carpa, Lophocolea heterophylla, Lophozia alpestris, L. ba-
densis, L. barbata, L. heterocolpa, L. incisa, L. kaurini, L.
Fig. 1. County map of Wisconsin. Numerals designate the
number of species which have been found in that particular county,
porphyroleuca, L. quinquedentata, L. ventricosa, Nardia
hyalina, Plagiochila asplenioides, Sphenolobus exsectaefor-
mis, S. exsectus, S. hellerianus, S. michauxii, Cephaloziella
hampeana, Bazzania trilobata, Calypogeia trichomanis,
240 Wisconsin Academy of Sciences, Arts, and Letters.
Cephalozia bicuspidata, C. catenulata, C. curvifolia, C.
media, Lepidozia reptans, Odontoschisma denudatum, Ble-
pharostorna trichophyllum, Ptilidium pulcherrimum, Tri-
chocolea tomentella, Scapania curta, S. irrigua, S. nemo-
rosa, S. subalpina, Radula complanata, R. obconica, Porella
platyphylla, P. platyphylloidea, Frullania asagrayana, F.
eboracensis.
Barron County
Riccia fluitans, Pallia fabroniana, Geocalyx graveolens,
Lophocolea heterophylla, Lophozia incisa, Plagiochila as-
plenioides, Bazzania trilobata, Calypogeia neesiana, Cepha¬
lozia media, C. pleniceps, Lepidozia reptans, Ptilidium pul-
cherrimum, Trichocolea tomentella, Radula obconica, Por¬
ella platyphylla, P. platyphylloidea, Frullania brittoniae, F.
eboracensis, Blasia pusilla, Pellia epiphylla.
Bayfield County
Riccia fluitans, Ricciocarpus natans, Conocephalum coni-
cum, Preissia quadrata, Marchantia polymorpha, Riccardia
pinguis, R. latifrons, R. palmata, Pellia epiphylla, P. fa¬
broniana, P. neesiana, Blasia pusilla, Fossombronia foveo-
lata, Plagiochila asplenioides, Lophocolea heterophylla, L.
minor, Chiloscyphus pallescens, Ch. fragilis, Ch. rivularis,
Harpanthus scutatus, Geocalyx graveolens, Nardia hyalina,
Jungermannia lanceolata, J. pumila, J. pumila var. rivul¬
aris, J. schiffneri, J. sphaerocarpa, Jamesoniella autum-
nalis, Lophozia alpestris, L. badensis, L. barbata, L. excisa,
L. heterocolpa, L. incisa, L. kaurini, L. muelleri, L. quinque-
dentata, L. ventricosa, Sphenolobus exsectaeformis, S. ex-
sectus, S. heilerianus, S. michauxii, Cephaloziella bifida, C.
hampeana, C. myriantha, Cephalozia bicuspidata, C. curvi¬
folia, C. pleniceps, Odontoschisma denudatum, 0. macounii,
Bazzania trilobata, Calypogeia neesiana, Blepharostoma
trichophyllum, Ptilidium pulcherrimum, Lepidozia reptans,
Diplophyllum apiculatum, Scapania apiculata, S. undulata,
S. curta, S. cuspiduligera, S. dentata, S. glaucocephala, S.
irrigua, S. nemorosa, S. paludicola, S. subalpina, Radula
complanata, R. obconica, Proella platyphylla, P. platyphyl-
Conklin — The Hepaticae of Wisconsin.
241
loidea, Cololejeunea biddlecomiae, Frullania brittoniae, F.
eboracensis, F. inflata, Anthoceros laevis.
Buffalo County
Reboulia hemisphaerica, Lophocolea heterophylla, L.
minor, Nardia hyalina, Anthoceros laevis, Geocaylx graveo-
lens.
Burnett County
Riccia arvensis, R. fluitans.
Columbia County
Ricciocarpus natans, Porella platyphylloidea.
Dane County
Riccia arvensis, R. fluitans, Ricciocarpus natans, Grim-
aldia fragrans, Reboulia hemisphaerica, Conocephalum co-
nium, Marchantia polymorpha, Pressia quadrata, Lopho¬
colea heterophylla, Ptilidium pulcherrimum, Porella platy-
phylla, P. platyphylloidea, Frullania brittoniae, F. ebora¬
censis, Nothothylas orbicularis.
Douglas County
Riccia fluitans, Ricciocarpus natans, Rebouilia hemis¬
phaerica, Conocephalum conicum, Marchantia polymorpha,
Preissia quadrata, Pallavincinia flotowiana, Riccardia lati-
frons, R. multifida, R. palmata, R. pinguis, Blasia pusilla,
Pellia epiphylla, P. fabroniana, P. neesiana, Chiloscyphus
fragilis, Ch. pallescens, Ch. polyanthos, Ch. rivularis, Geo¬
calyx graveolens, Harpanthus scutatus, Jamesoniella au-
tumnalis, Jungermannia lanceolata, J. pumila, J. schiffneri,
Lophocolea heterophylla, L. minor, Lophozia attenuata, L.
badensis, L. barbata, L. bicrenata, L. guttulata, L. hetero-
colpa, L. incisa, L. kaurini, L. longiflora, L. muelleri, L.
porphyroleuca, L. quinquedentata, Mylia anomala, Nardia
hyalina, Plagiochila asplenioides, Sphenolobus exsectaefor-
mis, S. exsectus, S. Hellerianus, S. michauxii, Cephaloziella
bifida, C. byssacea, C. elachista, C. hampeana, C. myrian-
242 Wisconsin Academy of Sciences, Arts, and Letters.
tha, C. sullivantii, Bazzania trilobata, Calypogeia neesiana,
C. trichomanis, Cephalozia bicuspidata, C. catenulaa, C.
connivens, C. curvifolia, C. macounii, C. media, C. pleni-
ceps, Lepidozia reptans, Odontoschisma denudatum, Bleph-
arostoma trichophyllum, Ptilidium ciliare, Pt. pulcherri-
mum, Trichocolea tomentella, Diplophyllum gymnostomo-
philum, Scapania apiculata, S. curta, S. glaucocephala, S.
irrigua, S. nemorosa, S. paludiocola, S. subalpina, Radula
complanata, R. obconica, Porella platyphylla, P. platyphyl-
loidea, Cololejeunea biddlecomiae, Frullania asaprayana, F.
brittoniae, F. bolanderi, F. eboracensis, F. oakesiana, F.
selwyiana, Anthoceros laevis.
Grant County
Riccia beyrichiana, R. sorocarpa, Grimaldia fragrans, G.
pilosa, G. rupestris, Reboulia hemisphaerica, Conocephalum
conicum, Marchantia polymorpha, Preissia quadrata, Chilo-
scyphus pallescens, Harpanthus scutatus, Jamesoniella au-
tumnalis, Lophocolea heterophylla, L. minor, Lophozia in-
cisia, Nardia hyalina, Plagiochila asplenioides, Sphenolobus
exsectus, Cephaloziella bifida, Calypogeia neesiana, Lepi¬
dozia reptans, Ptilidium pulcherrimum, Scapania curta, S.
nemorosa, Porella platyphylla, Frullania asagrayana, F.
brittoniae, F. eboracensis, F. inflata, F. riparia, Anthoceros
laveis, Frullania inflata.
Green County
Conocephalum conicum, Preissia quadrata, Lophocolea
heterophylla, Porella platyphylloidea.
Iron County
Conocephalum conicum, Marchantia polymorpha, Preis¬
sia quadrata, Blasia pusilla Pellia epiphylla, P. neesiana,
Chiloscyphus pallescens, Geocalyx graveolens, Harpanthus
scutatus, Jamesoniella autumnalis, Jungermannia lanceo-
lata, J. pumila, J. rivularis, J. schiffneri, Lophocolea hetero¬
phylla, L. minor, Lophozia badensis, L. barbata, L. hetero-
colpa, L. incisa, L. kaurini, L. quinquedentata, Nardia hy¬
alina, Plagiochila asplenioides, Sphenolobus scitulus, Ceph-
Conklin — The Hepaticae of Wisconsin.
243
aloziella hampeana, Cephalozia curvifolia, C. pleniceps,
Lepidozia reptans, Odontoschisma macounii, Blepharos-
toma trichophyllum, Trichocolea tomentella, Diplophyllum
gymnostomophilum, Scapania curta, S. glaucocephala, S.
subalpina, Radula complanata, Porella platyphylloidea, Col-
olejeunea biddlecomiae, Lejeuna cavifolia, Frullania asa-
grayana, F. eboracensis, Anthoceros laevis.
Iowa County
Riccia arvensis, R. fluitans, Marchantia polymorpha,
Fossombronia (sterile), Lophocolea heterophylla, Nardia
crenulata, Porella platyphylla, P. platyphylloidea, Frullania
eboracensis, Anthoceros punctatus.
Jackson County
Chiloscyphus rivularis.
Jefferson County
Lophocolea heterophylla, Ptilidium pulcherrimum.
Juneau County
Reboulia hemisphaerica, Marchantia polymorpha, Preis-
sia quadrata, Pellia epiphylla, Jungermannia pumila, Nar¬
dia hyalina, Anthoceros laevis.
Lafayette County
Marchantia polymorpha, Chiloscyphus fragilis, Harpan-
thus scutatus, Jamesoniella autumnalis, Jungermannia
lanceolata, Lophocolea heterophylla, L. minor, Nardia hy¬
alina, Sphenolobus exsectus, Cephaloia bicuspidata, Lepid¬
ozia reptans, Blepharostoma trichophyllum, Ptilidium pul¬
cherrimum, Porella platyphylla, Frullania asagrayana, F.
eboracensis, Anthoceros laevis.
Lincoln County
Conocephalum conicum, Riccardia palmata, Chiloscyphus
fragilis, Ch. rivularis, Harpanthus scutatus, Jungermannia
244 Wisconsin Academy of Sciences , Arts, and Letters.
lanceolata, Lophocolea heterophylla, L. minor, Lophozia
barbata, Bazzania trilobata, Cephalozia curvifolia, C. me¬
dia, Lepidozia reptans, Ptilidium ciliare, Pt. pulcherrimum,
Radula complanata, Frullania asagrayana, F. eboracensis,
F. brittoniae, Lejeuna cavifolia, Anthoceros laevis, A.
macounii.
Marathon County
Marchantia polymorpha, Peliia fabroniana, Chiloscyphus
rivularis, Jamesoniella autumnalis, Lophocolea hetero¬
phylla, L. minor, Lophozia alpestris, Sphenolobus helleri-
anus, Bazzania trilobata, Cephalozia bicuspidata, C. cat-
enulata, C. curvifolia, C. media, Ptilidium pulcherrimum,
Scapania nemorosa, Radula obconica, Porella pinnata, frul¬
lania asagrayana.
Milwaukee County
Lophocolea heterophylla, Ptilidium ciliare.
Oneida County
Riccia fluitans, Conocephalum conicum, Marchantia poly¬
morpha, Riccardia multifida, R. palmata, Blasia pusilla,
Peilia neesiana, Chiloscyphus fragilis, Ch. pallescens, Ch.
rivularis, Geocalyx graveolens, Jamesoniella autumnalis,
Lophocolea heterophylla, Lophozia barbata, L. incisa, L.
porphyroleuca, Plagiochila asplenioides, Sphenolobus hel-
lerianus, Bazzania trilobata, Calypogeia neesiana, Cephal¬
ozia catenulata, C. connivens, C. curvifolia, C. macounii, C.
media, C. pleniceps, Lepidozia reptans, Odontoschisma de-
nudatum, Blepharostoma trichophyllum, Ptilidium pulcher¬
rimum, Trichocolea tomentella, Radula complanata, Porella
pinnata, P. piatyphylloidea, Frullania asagrayana, F. ebor¬
acensis.
Pierce County
Conocephalum conicum, Preissia quadrata, Blasia pusilla,
Chiloscyphus pallescens, Lophocolea heterophylla, Antho¬
ceros laevis.
Conklin — The Hepaticae of Wisconsin.
245
Polk County
Ricciocarpus natans, Asterella tenella, Grimaldia frag-
rans, Conocephalum conicum, Marchantia polymorpha,
Preissia quadrata, Blasia pusilla, Pellia epiphylla, P. neesi-
ana, Chilscyphus pallescens, Harpanthus scutatus, Jame-
soniella autumnalis, Lophocolea heterophylla, L. minor, Lo-
phozia barbata, L. quinquedentata, Ptilidium pulcherri-
mum, Radula complanata, Porella pinnata, P. platyphylla,
P. platyphylloidea, frullania eboracensis, F. riparia, An-
thoceros laevis, Grimaldia pilosa.
Portage County
Riccia fluitans, Pellia fabroniana, Lophocolea minor,
Frullania inflata.
Racine County
Reboulia hemisphaerica.
Riccia arvensis, Grimaldia fragrans, Conocephalum coni¬
cum, Preissia quadrata, Marchantia polymorpha, Reboulea
hemisphaerica, Lophocolea heterophylla, L. minor, Jame-
soniella autumnalis, Ptilidium pulcherrimum, Porella plat-
phylla, P. plalyphylloidea, Frullania brittoniae, F. ebora¬
censis, F. reparia.
Sauk County
Riccia fluitans, Reboulia hemisphaerica, Marchantia
polymorpha, Metzger ia conjugata, Pellia epiphylla, P. nee-
siana, Jamesoniella autumnalis, Lophocolea heterophylla, L.
minor, Lophozia barbata, L. ventricosa, Plagiochila asplen-
ioides, Nardia hyalina, Sphenolobus exsectus, Cephalozia
curvifolia, Scapania nemorosa, Radula complanata, Porella
platyphylla, Frullania eboracensis, Anthoceros laevis.
Sawyer County
Marchantia polymorpha, Riccardia latifrons, Jamesoni¬
ella autumnalis, Harpanthus scutatus, Lophocolea hetero¬
phylla, Bazzania trilobata, Cephalozia curvifolia, Porella
platyphylla.
246 Wisconsin Academy of Sciences , Arts, and Letters.
Trempealeau County
Reboulia hemisphaerica, Nardia hyalina Porella platv-
phylla, Frullania eboracensis, Anthoceros laevis, Junger-
mannia pumila.
Vilas County
Conocephalum conicum, Marchantia polymorpha, Ric-
cardia latifrons, R. pinguis, Pellia epiphylla, P. neesiana,
Geocalyx graveolens, Jamesoniella autumnalis, Lophocolea
heterophylla, Lophozia barbata, L. incisa, L. porphyroleuca,
Plagiochila aspleniodes, Splenolobus exsectaeformis, S. hel-
lerianus, S. michauxii, Bazzania trilobata, Cephalozia con-
nivens, C. curvifolia, C. macounii, C. media, Lepidozia rep-
tans, Odonto schisma denudatum, Blepharostoma tricho-
phyllum, Ptilidium ciliare, Pt. pulcherrimum, Scapania
paludicola, Porella platyphylla, Frullania eboracensis.
Washington County
Marchantia polymorpha.
Waukesha County
Riccia fluitans, Ricciocarpus natans, Lophozia hetero-
colpa.
Wood County
Rebouila hemisphaerica, Conocephalum conicum, Pellia
epiphylla, Harpanthus scutatus, Nardia hyalina.
Publications
1. Cheney, L. S., and True, R. H. On the flora of Madison
and vicinity, a preliminary paper on the flora of Dane
County, Wisconsin. Trans. Wis. Acad. Sci., Arts &
Let. 9 (1) : 45-185. 1893. (Hepaticae, pp. 133,
134.)
2. Cheney, L. S. A contribution to the flora of the Lake
Superior region. Trans. Wis. Acad. Sci., Arts & Let.
9 (1) : 233-234. 1893. (Hepaticae, pp. 233, 254.)
Conklin — The Hepaticae of Wisconsin.
247
3. - Hepaticae of the Wisconsin Valley. Trans.
Wis. Acad. Sci., Arts, & Let. 10 : 70-72. 1895.
4. Allen, C. E. Some Hepaticae of the Apostle Islands.
Trans. Wis., Acad. Sci., Arts & Let. 14 (2) : 485,
486. 1904.
5. Conklin, G. H. Brief notes on the distribution of Hepa¬
ticae. Bryologist 15 : 11,12. 1912.
6. — - — Preliminary report on a collection of Hepa¬
ticae from the Duluth-Superior district. States of
Minnesota and Wisconsin. Trans. Wis. Acad. Sci.,
Arts & Let. 17 (2) : 985-1010. 1914.
BUI
PRELIMINARY REPORTS ON THE FLORA OF
WISCONSIN. I. JUNCAGINACEAE,
ALISMACEAE
Norman C. Fassett
It is intended to issue from time to time, under this title,
accounts of the ranges, so far as they are known, of certain
groups of plants in Wisconsin. These reports are directed
largely to amateurs, without whose help the study of the
flora of any region is a tedious process. The professional
botanist may visit any given locality, and perhaps find a
fair proportion of the plants in flower or fruit at the par¬
ticular time, but only the local botanist, watching the same
region throughout the year and from year to year, can have
a true conception of the local flora. By presenting graphi¬
cally the known range of each plant it is hoped to show the
amateur how he can help fill the gaps in our knowledge of
these species. Plants familiar to him may prove to be im¬
portant extensions of range.
These maps are compiled entirely from herbarium speci¬
mens in the herbaria of the University of Wisconsin, of the
Milwaukee Public Museum and of Mr. S. C. Wadmond, Del-
avan, Wis. An important herbarium of Wisconsin plants,
that of the late J. H. Schuette, is unfortunately not easily
available. A number of Schuette’s collections, however,
are included in the herbarium of the late Professor L. M.
Umbach, recently acquired by the University of Wisconsin.
Specimens collected as hosts of fungi, particularly by Dr. J.
J. Davis, have been used In making these maps only when
their identity was absolutely certain.
JUNCAGINACEAE — Arrow Grass Family
1. Scheuchzeria
S. palustris L., var. Americana Fernald, Rhodora 25:
178. 1923. (Fig. 1). The American phase of Scheuch¬
zeria has been shown to differ from the European in the
250 Wisconsin Academy of Sciences , Arts , and Letters.
size and shape of the fruit. This plant occurs in Sphag¬
num bogs of northern and eastern Wisconsin. An un¬
named locality in Oconto County is not shown on the map.
2. Triglochin
T. maritima L. (Fig. 2) . Shore of Lake Superior, and
occasional on small lakes in the interior. Is this really ab¬
sent from the Lake Michigan shore ? The station in Racine
County is on Racine Prairie,1 five miles from the lake.
T. palustris L. (Fig. 3) . Shore of Lake Michigan, in
Door, Racine, and Kenosha Counties.
ALISMACEAE— Water-plantain Family
1. Sagittaria
S. latifolia Willd. (Fig. 4). The typical form of this
species seems to be found throughout Wisconsin. Speci¬
mens from Tripoli, Oneida County (August 17 & 19, 1925,
J. J. Davis) have the middle flowers perfect.
S. latifolia, f. hastata (Pursh) Robinson. (Fig. 5) .
Apparently absent from the lower Wisconsin and Missis¬
sippi Rivers.
S. latifolia, f. gracilis (Pursh) Robinson. (Fig. 6) .
Like the preceding, this form appears to be absent from
southwestern Wisconsin.
S. latifolia, var. obtusa (Muhl.) Wiegand, Rhodora
27 : 186. 1925. S. latifolia, f. obtusa Robinson. (Fig.
7) . Professor Wiegand’s statement that “the obtuse leaf
apex and the generally dioecious flowers as well as the
rather definite range suggest a more racial difference” ap¬
plies well to this plant as it grows in Wisconsin. It is the
most conspicuous Sagittaria on the Mississippi River bot¬
toms (in spite of the meagre collections from that region)
forming large patches, each often made up entirely of
plants of one sex.
S. CUNEATA Sheldon, Bull. Torr. Bot. Club 20: 283.
1893. S. arifolia Nutt, in herb. ; J. G. Smith, Rep. Mo. Bot.
Gard. 6: 32. 1895, and reprint, 6. 1894. (Fig. 8).
1 Wadmond, Trans. Wis. Acad. Sci., Arts & Let. 10:806. 1909.
Fassett — Preliminary Reports on Flora of Wisconsin. 1. 251
252 Wisconsin Academy of Sciences , Arts , and Letters,
Fassett — Preliminary Reports on Flora of Wisconsin. I. 253
17
254 Wisconsin Academy of Sciences, Arts, and Letters.
Sands of the Mississippi and Wisconsin Rivers, and south¬
ern Wisconsin northward to Shawano County.
S. heterophylla Pursh. Throughout the state, varying
as to leaf-outline. The typical form (fig. 9) seems to be
more common westward in the state, but is hardly dis¬
tinguishable from f. elliptica.
S. heterophylla, f. elliptica (Engelm.) Blake, Rho-
dora 15: 159. 1913. S. heterophylla Engelm. (Fig. 10).
Scattered.
S. HETEROPHYLLA, f. rigida (Pursh) Blake, 1. c. S. hetero¬
phylla, var. rigida Engelm. and var. angustifolia Engelm.
(Fig. 11). The commonest form.
S. HETEROPHYLLA, f. fluitans (Engelm.) Blake, 1. c.
(Fig. 12). This is a submerged form, with linear, usually
almost bladeless leaves. Occurs rarely northward and east¬
ward.
S. GRAMINEA Michx. (Fig. 13). Mostly northward.
This species and S. cuneata frequently form, when sub¬
merged, rosettes of narrowly lanceolate bladeless phyllodia.
These forms, usually sterile, may be distinguished in Wis¬
consin as follows:
a. Phyllodia with 12-20 longitudinal veins of almost equal
heaviness (fig. 14, a & b) _ S. graminea
a. Phyllodia with 3-5 heavy longitudinal veins, the inter¬
stices reticulate (fig. 14, e & f)__. - S. cuneata
However, S. graminea is a complex species, and this
character cannot be relied upon in other regions. It is a
good distinction in material from Maine, but a large series
of S. graminea from Illinois and Indiana (fig. 14, c & d) has
the veination ascribed to S. cuneata in Wisconsin.
Whether Michaux’s type of S. graminea was the plant
called by that name in Wisconsin or in Indiana, if either,
must remain a question at present.
The S . graminea of New England is seldom, if ever,
found with mature fruit. The plant of Wisconsin, appar¬
ently identical with it, at least in the veination of the phyl¬
lodia, occasionally sets fruit. A good series of this plant
collected at Hayward by Professor E. M. Gilbert and the
writer shows phyllodia and fruit. The achenes are like
those ascribed by J. G. Smith1 to S. cristata Engelm. It is
1 Rep. Mo. Bot. Gard. 6: 53. 1895, and reprint, 27: 1894.
Fassett — Preliminary Reports on Flora of Wisconsin. I. 255
possible, then, that S. cristata is really the northern repre¬
sentative of the complex group passing as S. graminea .
Michaux’s type of S. graminea may be the same plant.
Figure 14 represents portions of the phyllodia of six col¬
lections, as follows: a, S. graminea , Bowdoinham, Maine,
Fassett, no. 2847 ; b, S . graminea, Hayward, Wisconsin, Gil¬
bert & Fassett, no. 7436; c, S. graminea, Clarke, Indiana,
Umbach, June 4, 1898; d, S. graminea, Tolleston, Indiana,
Umbach, July 7, 1900; e, S. cuneata, Troutdale, Maine, Fas¬
sett, no. 3626; f, S. . cuneata , Bay City, Wisconsin, Fassett &
Wilson, no. 5288.
2. Lophotocarpus
L. calycinus (Engelm.) J. G. Smith. (Fig. 15). Rare,
but probably sometimes passed by as a Sagittaria. Sandy
shores of sloughs, etc., on the Mississippi and Wisconsin
River bottoms. This is a range not uncommon for plants
coming to Wisconsin from regions to the southward.
3. Alisma
A. Plantago-aquatica L. (Fig. 16). Dr. Gunnar Sam-
uelsson of Stockholm, Sweden, has identified the Wisconsin
Alisma of the University Herbarium. He separates our
A. Plantago-aquatica as a variety distinct from the Euro¬
pean plant.
The Wisconsin plant, as indicated by available specimens,
is absent from the great north-central shield of Archean
igneous rocks. Further field work must, however, check
this apparent fact.
A. subcordatum Raf. (Fig. 17). Rare in Wisconsin.
This is a segregate from the preceding species.
A. gramineum Gmel. (Fig. 18). This is the A. Geyeri
of Gray's Manual. It has been collected in this state only
once, at Alma. It was noted by the writer in the summer
of 1926, growing abundantly at several points on the Mis¬
sissippi River bottoms at Teepeota Point, below Wabasha,
Minnesota, and again directly across the river from Alma.
It seems to prefer damp places on the sand which silts in
between the wing dams. (The Mississippi River Commis-
256 Wisconsin Academy of Sciences , Arts, and Letters.
sion chart of 30 years ago2 shows the river at Alma as be¬
ing nearly a mile and a half wide, with a boom to direct
logs to the head of West Newton Chute, on the Minnesota
side. At the low water of August, 1926, the river was less
than a half mile wide at this point, and a series of wing
dams extended from Island 39 to Island 40, which had been
well toward the middle of the stream in 1897. A shallow
slough behind the new sand island formed about these wing
dams was only knee-deep. This part of the river was 4-8
feet deep in 1897. It was on this shore that A. gramineum
grew. This ground was flooded after the rains of Septem¬
ber, 1926, and the plant could not be found at the same
point in 1927.) The writer in 1926 spent an entire after¬
noon searching for this plant on the Wisconsin side of the
river, and finally collected only two individuals, a few miles
below Alma.
2 October 7, 1897.
PRELIMINARY REPORTS ON THE FLORA OF
WISCONSIN. II. ERICACEAE
Norman C. Fassett
1. General Discussion
The ranges of many ericaceous plants in Wisconsin illus¬
trate the control of plant distribution by geological and
physiographic features.
Bog plants are almost entirely absent from the unglaci¬
ated area of the southwestern part of the state. The to¬
pography of this area is of the type known as maturity,
with maximum slopes, almost perfect drainage, no natural
bodies of standing water (except, in this case, in the river-
bottoms) and (except as noted below) no bogs. The rest of
the state, recently glaciated, with many pockets where bogs
may form, shows such sphagnicolous plants as Cranberries
( Vaccinium macrocarpon, fig. 30, V. Oxycoccos, var. ovali -
folium , fig. 29), Creeping Snowberry ( Chiogenes hispidula,
fig. 22), Leather Leaf ( Chamaedaphne calyculata , fig. 18,
and Bog Rosemary ( Andromeda glaucophylla, fig. 17).
Parts of Jackson and Wood Counties, in the unglaciated
area, are flat and sandy, and the underground drainage is
so obstructed as to produce standing water; here we find
Vaccinium macrocarpon and the Chamaedaphne. The
former is represented by four collections, and the latter by
but two; future collections should show these species to be
more abundant in this part of the state.
Dry-soil types are rare in the unglaciated area, but for
another reason, concerned with the nature of the rocks
themselves. Trailing arbutus ( Epigaea repens, fig. 19),
Bearberry ( Arctostaphylos Uva~ursi, fig. 21), and the Blue¬
berries ( Vaccinium pennsylvanicum, fig. 24, and V. cana-
dense, fig. 26) avoid the limy rocks of southern Wisconsin.
On the maps of these species the distribution of the St.
Croixian Sandstone is indicated, and the plants may be seen
to be plentiful on this type of rock. They are equally plen-
258 Wisconsin Academy of Sciences, Arts, and Letters .
tiful on the granites and sandstones which lie north of this
formation, but south of this mass, where it is overlain by
Lower Magnesian and Galena Black River Domolites, they
are rare, except where patches of sandstone are not only
exposed by erosion but leached by weathering. Such
places may be seen on Picture Rock near Mt. Vernon (Dane
County), where Gaylussacia baccata and Vaccinium penn-
sylvanicum grow, and on Pine Bluff (named for its isolated
grove of Pinus Strobus) at Brodhead (Green County)
where Epigaea repens and Vaccinium canadense are found.
This type of plant also follows the shore of Lake Michigan.
Since the underlying rock here is Niagara Dolomite, the re¬
lation of these plants to sandy beaches, fossil or otherwise,
should be investigated.
Plants preferring woods are more general, but hardly
common, south of the granite and sandstone regions.
These include Princess Pine (Chimaphila umbellata, fig.
1), Wintergreen ( Gaultheria procumbens, fig. 20), and In¬
dian Pipe ( Monotropa uniflora, fig. 13) .
2. Ranges of Species and Varieties
The maps illustrating the following ranges are based on
herbarium specimens seen by the writer. These specimens
are in the Herbaria of the University of Wisconsin, of the
Milwaukee Public Museum, and of Mr. S. C. Wadmond.
Hosts of parasitic fungi in the Herbarium of the University
of Wisconsin were also used to determine localities for
these plants, except in the cases of Pyrola and Vaccinium.
Chimaphila umbellata (L.) Bart., var. cisatlantica
Blake, Rhodora 19: 241. 1917. C. corymbosa Pursh, in
part; Rydberg, N. Am. Flora 29, pt. 1: 31. 1914, in part.
(Fig. 1). Common northward in Wisconsin, rather rare
southward (see preceding paragraphs). The typical Chim¬
aphila umbellata is a plant of Europe, not found in the
United States.
MONESES UNIFLORA (L.) Gray. (Fig. 2). Common north¬
ward, very rare southward.
Pyrola minor L. (Fig. 3). Rare in Wisconsin, known
only from Cornucopia, where it was collected by Mr. L. S.
Cheney in 1897.
Fassett— Preliminary Reports on Flora of Wisconsin. 11. 259
260 Wisconsin Academy of Sciences, Arts, and Letters ,
Fassett — Preliminary Reports on Flora of Wisconsin. II. 261
P. SECUNDA L. (Fig. 4). Northern and eastern, with a
few isolated stations in central southern Wisconsin.
P. SECUNDA, var. OBUSATA Turcz. (Fig. 5) . Northward.
The characters of this variety as given in the following key
are mostly an abridgment of those listed by Fernald, Rho-
dora 28 : 223, 1926.
P. chlorantha Sw. (Fig. 6). Mostly northward, but
south to Racine on the Lake Michigan shore.
P. CHLORANTHA, var. paucifolia Fernald, Rhodora 22:
51, 1920. (Fig. 7). Range similar to the preceding.
P. CHLORANTHA, var. convoluta (Barton) Fernald, 1. c.,
page 52. (Fig. 8). Less common. The portion of the
key dealing with this species, on pages 262 & 264, is adapt¬
ed from that of Fernald, 1. c.
P. elliptxca Nutt. (Fig. 9). Our most abundant repre¬
sentative of the family.
P. rotundifolia L., var. Americana (Sweet) Fernald,
Rhodora 22 : 122, 1920. P. americana Sweet ; Fernald,
Rhodora 6: 201, 1904. (Fig. 10). Mostly northward.
In his earlier paper (1904) Fernald pointed out what he
then considered specific characters to separate this plant
from the European P. rotundifolia, but in 1920 he reported
the discovery of a plant of Newfoundland (later found also
in Novia Scotia) which showed intermediate characteris¬
tics. His description of this plant, var. arenaria Mert. &
Koch, in 1920, reads much like that of his earlier descrip¬
tion of typical P. rotundifolia. However, whatever may be
the status of this intermediate, a comparison of the com¬
mon American plant with that of Europe gives the impres¬
sion that it is better treated as a variety than maintained
as a species.
P. asarifolia Michx. (Fig. 11) . Mostly northern, south
rarely to Adams and Milwaukee Counties. (A station in
southeastern Dunn Co. was inadvertently omitted from the
map.)
P. asarfolia, var. INCARNATA (Fisch.) Fernald. (Fig.
12). Perhaps more common eastward and southeastward
than the preceding.
The following key to the species and varieties of Pyrola
in Wisconsin is intended not to supplant, but to supple-
262 Wisconsin Academy of Sciences , Arts , and Letters.
ment, the treatments in the current manuals, and to point
out some additional, mostly vegetative, characters.
a. Bracts, intermingled with the leaves at the base of
the stem, crowded, usually 1 cm. or more long,
obtuse, rounded or truncate at tip, often grad¬
ing into leaves b
b. Cauline bracts 1-5 (very rarely none) ovate-
lanceolate, their bases somewhat sheath¬
ing the stem; leaf -blades usually shorter
than the petiole ; sepals longer than broad
c
c. Sepals ovate-lanceolate, rather blunt, twice
as long as broad; petals white, rarely
pinkish-tinged, 6.5-10.5 mm. long___ P. rotundifolia,
var. americana.
c. Sepals triangular, acute or acuminate,
rarely more than 1.5 times as long as
broad; petals pink, about 5 mm. long
d
d. Leaf -blades cordate at base - P • asarifolia.
d. Leaf-blades subtruncate, rounded, or
tapering at base - P. asarif olia,
var. incarnata.
b. Cauline bracts none, or 1-3, narrowly lanceo¬
late, long-acuminate, not sheathing; leaf-
blades longer than the petiole ; sepals
about as broad as long e
e. Leaf -blades 3-7 cm. long, tapering at base ;
style declined; stigma not peltate — P .elliptica.
e. Leaf -blades 2-4 cm. long, rounded or trun¬
cate at base; style straight; stigma
peltate _ P- minor.
a. Basal bracts 2-4 mm. long, acute or acuminate, dis¬
tinct from the leaves, often absent /
/. Leaf -margins crenate-serrate, usually with min¬
ute teeth; scape with (1-) 2-4 ovate-
lanceolate bracts g
g. Basal bracts lanceolate, strongly involute,
firm; leaf -blades narrowed at tip,
1.5-6 cm. long _ _ _ _ _ P. secunda.
g. Basal bracts oblong to ovate, slightly in¬
volute, membranaceous ; leaf -blades
rounded at tip, 0.8-3 cm. long - P. secunda ,
var. obtusata.
/. Leaf -margins entire or somewhat undulate;
scape naked or rarely with 1 or 2 short
setaceous bracts h
Fassett — Preliminary Reports on Flora of Wisconsin . II,
263
284 Wisconsin Academy of Sciences, Arts, and Letters .
h. Leaf -blades 0.7-3. 3 cm. broad; calyx 3-4
(-5) mm. broad; anthers 1.6-3
(-3.3) mm. long i
i. Leaf -blades rounded to base and apex,
1. 5-3.3 cm. broad, 4-11 in a rosette P. chiorantka.
i. Leaf -blades mostly cuneate at base and
truncate or subtruncate at summit,
0.7-2.5 cm. broad, 0-7 in a rosette P. chlorantha,
var. paucifolia.
h . Leaf -blades 2-4.5 cm. broad; calyx 4.8-6
mm. broad; anthers 3-4 mm. long__ P. chlorantha,
var. convoluta.
Montropa UNIFLORA L, (Fig. 13). Throughout the
state, but rare in the unglaciated region, probably because
of the scarcity of damp woods there. Indian Pipes are
sometimes found, in this state, in deep Sphagnum.
M. Hypopitys L. (Fig. 14, dots). In woods, generally
distributed, but not common.
Pterospora andromedea Nutt. (Fig. 14, cross). Col¬
lected but once in Wisconsin, by Mr. Alvin Throne, at
Donges Bay, Ozaukee County, August 15, 1928.
Ledum groenlandicum Oeder. (Fig. 15). Mostly north¬
ward. Comes south to the Dells of the Wisconsin River,
where it grows on dry sandstone ledges along the river.
Rhododendron lapponicum (L.) Wahlenb. (Fig. 16,
cross) . This plant was collected but once in Wisconsin,
on May 30, 1898, by Mr. L. S. Cheney, on the high sand¬
stone walls of the gorge cut by the Wisconsin River at
Kilbourn, in the Dells. It has not been seen there since
1898.
Kalmia polifolia Wang. (Fig. 16, dots). Mostly north¬
ern, in Spahgnum bogs.
Andromeda glaucopiiylla Link. (Fig. 17) . The ungla¬
ciated area is indicated on the map ; see page 257.
Chamaedaphne calyculata (L.) Moench. (Fig. 18).
The unglaciated area is indicated on the map ; see page 257.
Epigaea repens L. (Fig. 19). The area of St. Croixian
Sandstone is indicated on the map ; see page 257.
Gaultheria procumbens L. (Fig. 20) . Rare southward.
The most abundant colonies I have seen in the southern
part of the state are in the pine woods at the Dells of the
Wisconsin River, in Adams County.
Fassett — Preliminary Reports on Flora of Wisconsin. II. 265
266 Wisconsin Academy of Sciences, Arts, and Letters
Fassett — Preliminary Reports on Flora of Wisconsin. II. 267
Arctostaphylos Uva-ursi (L.) Spreng., var. coactilis
Fernald & Macbride, Rhodora 16: 212, 1914. (Fig. 21).
The area of St. Croixian Sandstone is indicated on the map ;
see page 257. The Bearberry of this region differs slightly
from the typical form of the European plant in the char¬
acter of the tomentum of the twigs, and has been varietally
separated on that character.
Chiogenes hispidula (L.) T. & G. (Fig. 22). The un¬
glaciated area is indicated on the map ; see page 257.
Gaylussacia baccata (Wang.) K. Koch. (Fig. 23).
Although apparently more abundant southward, this
species is, south of the sandstone area, largely confined to
the sandstone outcrops discussed in connection with
Epigaea repens , etc., on pages 257-258.
Vaccinium pennsylvanicum Lam. (Fig. 24). The area
of St. Croixian Sandstone is indicated on the map ; see page
257.
V. PENNSYLVANICUM, var. myrtilloides (Michx.) Fer¬
nald, Rhodora 10: 148, 1908. This northern pubescent
extreme was collected by the writer in dry woods on the
Elephants Back, near the Dells of the Wisconsin River,
Adams County, on May 19, 1929.
V. pennsylvanicum, var. nigrum Wood. (Fig. 25).
Most of our collections of this variety are from central
Wisconsin, but it is probably more widespread than these
would indicate. Dr. J. J. Davis tell me that he has seen it
in Door County.
V. CANADENSE Kalm. (Fig. 26). The area of St. Croix¬
ian sandstone is indicated on the map ; see page 257.
V. cespitosum Michx. Fig. 27, crosses). The unglaci¬
ated area is indicated on the map. This is one of the
species accredited with preglacial age by Fernald, Mem.
Am. Acad. Sci. 15 : 282, 1925. Of the three known stations
in Wisconsin two are in the unglaciated area and one is to
the north of it. This is in line with the ranges of a number
of plants which seem to have survived the glacial period in
this region, and to have worked northward since the reces¬
sion of the ice, as, for example, Epipactis decipiens on the
south shore of Lake Superior, and Orplopanax horridum on
Isle Royale. Indeed, Vaccinium cespitosum is also found
268 Wisconsin Academy of Sciences, Arts, and Letters.
north of Wisconsin, on the Upper Peninsula of Michigan
and the north shore of Lake Superior.
V. Vxtis-Idaea L., var. minus Lodd. (Fig. 27, dot) . This
plant has long been known from Grand Marais, Minnesota,
near Duluth, and in 1926 it was collected in Superior,
Wisconsin, by Mr. L. R. Wilson, of that city.
V. Oxycoccos L. (Fig. 28). Northern.
V. Oxycoccos, var. ovalifolium Michx. ; Robinson & Fer-
nald, Rhodora 19: 54, 1909. Var. intermedium Gray.
(Fig. 29). The unglaciated area is indicated on the map;
see page 257.
V. macrocarpon Ait. (Fig. 30) . The unglaciated area
is indicated on the map ; see page 257. Cranberries are one
of the important crops of Wisconsin, the yield being sur¬
passed only by those of Massachusetts and New Jersey.
An old collection by T. J. Hale is marked “St. Croix’’ ; this
may be St. Croix Falls (where the plant has since been
collected), or Upper Lake St. Croix (Douglas County), or
somewhere along the St. Croix River between these points.
NOTES ON PARASITIC FUNGI IN WISCONSIN. XV
J. J. Davis
In “Notes” XIV, p. 185, Taphrina struthiopteridis Nishida
was recorded as occurring in Wisconsin. On referring to
the specimen in the herbarium it is found to be labeled
Taphrina hiratsukae Nishida.
In Annales Mycologici 23 : 63-4 is a note by Petrak on
Phyllosticta iridis Ell. & Mart, in which he refers it to
Phlyctaena. As he stated, the material that he examined
was over-mature. Had he examined younger material he
would have found that the pycnidia are perfect and hence
that there is no need of removing it from Phyllosticta be¬
cause of imperfect pycnidia.
Ascochyta wisconsina Davis (“Notes” II, 101) was re¬
ferred to that genus because a few septate sporules were
found. In a recent collection on Sambucus canadensis
nearly all of the sporules have a median septum. Some of
them are a little longer (12/*) than the description indi¬
cated.
In the original description of Septoria pachyspora Ell.
& Hoi. the pycnidia are said to be “on snow-white, thin,
small (1-2 mm.), subangular to nearly round spots sur¬
rounded by a broad purplish shaded border” (Journ. My col.
1:6). In the record of its occurrence in Wisconsin it was
stated that the spots lacked the colored border (Trans.
Wis. Acad. 9 : 177) . In a recent collection (Brodhead, Wis.,
Sept. 26, 1926) the leaves are somewhat faded and yellow¬
ish and the spots are merely small areas that have retained
the deep green color with a tendency to become blackish in
the central portion, the location of the pycnidia being indi¬
cated by whitish points due to exuded sporules. In the
original description, which was copied in Martin’s “Sep-
torias of North America” (Journ. My col. 3: 67) the width
of the sporules was given as 3/* which was perhaps a mis-
18
27 0 Wisconsin Academy of Sciences, Arts, and Letters.
print for 5/x inasmuch as they approach the phragmos-
porous form as indicated by the specific name that was
applied. In some collections the spots become arid and the
sporules are narrow, probably because of unfavorable con¬
ditions. In reporting upon the determination of specimens
a well known mycologist, who has described many species
of North American Fungi, stated that he had been influ¬
enced more by the character of the spots than by those of
the spores. It has become evident that spot characters
depend largely on environmental influences affecting the
host and should be used with caution in taxonomy.
A specimen on leaves of Helianthus from Lynxville
(July 17, 1916) shows angular spots 2-5 mm. in diameter
which are black-brown above, lighter on the lower surface.
The pycnidia are hypophyllous, small, scattered, the spor¬
ules mostly curved, 27-37 x l-Wfap. This is labeled Septo -
ria paupera Ellis. The host is a form with rather thin
triple-nerved leaves.
By an unfortunate error it was stated in “Notes” XIII,
p. 172, that Fungi Columbiana 2593 represents Gloeospor-
ium niveum Davis. The number should be 3491, 2593 being
as labeled.
What appears to be the parasite recorded in “Notes”
I, p. 89 under the name Colletotrichum helianthi n. sp. was
distributed in Ellis & Everhart North American Fungi,
second series 2778 as Vermicularia helianthi n. sp. The
specimen is from Kansas on Helianthus rigidus collected
by W. T. Swingle. Apparently no description was pub¬
lished.
In “Notes” XI, p. 291, the statement was made that I had
seen no description of the parasite on Fraxinus oregana
issued in Fungi Columbiani 4415, 4719, and 4816 under the
name Cylindrosporium fraxini (Ell. & Kell.) Ell. & Ev.
Ellis & Everhart North American Fungi, second series
1634, Septogloeum fraxini Hark, is the same fungus. As
the material was provided by Dr. Harkness it may be
assumed to be authentic, but the description of the spots
as given in Saccardo, Sylloge Fungorum 3: 802, “Maculis
minutis, albidis, angulosis” would not suggest it.
Davis — Notes on Parasitic Fungi in Wisconsin. XV. 271
In the first “supplementary list of parasitic fungi of
Wisconsin’" Phleospora oxyacanthae (Kze. & Schm.) was
recorded as occurring in the state. In carrying this over
into the “provisional list” the specific name was errone¬
ously printed “crataegi”. In “Notes” III, p. 254, a short
description of the parasite was given showing characters
quite different from those of Phleospora. In “Notes” X
it was stated that it had been compared with an authentic
specimen of Cercosporella mirabilis Pk. with which it
agreed. Since then I have examined Cylindrosporium
crataegi Ell. & Ev. as represented in North American Fungi
3182 and find that to be the same parasite. I do not find
the acervuli mentioned by Ellis & Everhart (Proc. Acad.
Nat. Sci. Phila. 189 4, p. 372) nor yet differentiated conidi-
ophores as stated by Peck and am labeling it Cercoseptoria
crataegi (E. & E.) n. comb. Peck stated that “This is a
remarkable, aberrant and variable species and possibly the
type of a new genus” (Report of the State Botanist, 1911,
p. 46). The genus to which it is here referred had not
at that time been proposed. Infected leaves may be un¬
spotted or spotted in various degrees, apparently accord¬
ing to age and amount of infection.
Saccardo referred to Cercospora nasturtii Pass, as sub¬
species barbareae, a form on Barbarea vulgaris , and fig¬
ured it in Fungi Ital. del. 648. A collection on Barbarea
stricta from Wauwatosa, however, bears more cylindrical
and multiseptate conidia like those of Cercospora armor -
aciae Sacc. as figured in F. Ital. del. 646. In this collection
on Barbarea stricta both conidiophores and conidia range
up to 125 /jl or more in length. It may be that such forms
on Cruciferae as have been named C. nasturtii Pass., C.
armoraciae Sacc., C. cheiranthi Sacc. etc., will be found
to be conspecific. Examination of a specimen labeled Cer¬
cospora cruciferarum E. & E. in Ellis & Everhart, North
American Fungi , 1995, showed Septoria sisymbrii Ellis.
In “Notes” VIII, p. 429, record was made of a Cercos¬
pora on Lespedeza capitata with the suggestion that it was
perhaps a northern form of the parasite which Atkinson
described, from material collected in Alabama, as Cercos¬
pora flag ellif era. The Wisconsin plant is doubtless Cer -
272 Wisconsin Academy of Sciences, Arts, and Letters.
cospora lespedezae Ell. & Dearn. which was described from
Canadian material. Length of conidiophores and of conidia
are variable characters in this genus. Cercospora latens
Ell. & Ev. is probably a form of the same species. In the
description of this (Journ. My col. 4: 3), the host was
given as Psoralea argophylla, but the error was later cor¬
rected.
In “Notes’’ XIV the statement was made that Amphi-
car pa monoica had been infected in the greenhouse with
teliospores from Andropogon as the infecting agent and
Aecidium falcatae Arth. as the result. In the spring of
1926 infection of Andropogon furcatus was brought about
in the greenhouse from exposure to aeciospores from Am-
phicarpa.
In June, 1926, Aecidium xanthoxyli Pk. was found in
abundance on Zanthoxylum americanum in the bottomlands
opposite Sauk City, but no clue had been secured as to the
alternate host. However, having potted plants of Andro¬
pogon furcatus in the greenhouse they were exposed to
infection from the Aecidium, with the result that uredinia
and at length telia developed. The uredospores from this
infection are of the Puccinia pustulata type referred to in
North American Flora as the Santalaceous race. The situ¬
ation with regard to Puccinia on Andropogon is evidently
somewhat complicated. As the plants of Andropogon that
were infected by aecia on Amphicarpa and on Zanthoxylum
had the same origin there is no question of racial differenti¬
ation of the telial hosts.
The first collection of aecia of Puccinia sorghi Schw. in
Wisconsin was made at Blue River, June 19, 1926 on Oxalis
corniculata. The aecidium was abundant in a station but a
few feet across on an old road running through brushland .
A collection on Oxalis “cymosa?” was made on the Univer¬
sity farm July 20, 1926, by J. G. Dickson.
A Phragmidium which occurs commonly in Wisconsin
on Potentilla canadensis was recorded under the name
Phragmidium potentillae-canadensis Diet, with the descrip-
tion of which all American specimens that I have examined
agree. This was separated from Ph. tormentillae Fckl.
{Ph. ohtusum (Strauss) Lk.) of Europe because of the
Davis — Notes on Parasitic Fungi in Wisconsin. XV. 273
smaller number of cells in the teliospores. October 2, 1925
a collection on P. canadensis was made at Madison having
teliospores extremely variable in size and septation from
which the following notes were made : Teliospores straight
or somewhat curved, 33-280/*, long, 2-16 celled, the cells
either uniform or varying in length and width in the same
spore, the septa usually transverse but exceptionally more
or less oblique. Size of individual cells measured 15-
36x14-33/1. Germ pores single. As but one collection of
this character has been made it is presumably an abnormal
development. Occasionally there is an appearance that
suggests proliferation of spores, but usually it seems more
like proliferation of cells.
Additional Hosts
Peronospora ficariae Tul. A very scanty development
on radical leaves of Ranunculus abortivus. Blue River.
Erysiphe cichoracearum DC. occurred at Brodhead on
Artemisia ludoviciana. Perithecia were very few but
pycnidia of Cicinnobolus were abundant but sterile.
Pseudopeziza medicagmis (Lib.) Sacc. On Melilotus
alba. Madison. (F. R. Jones) Sometimes abundant on
this host.
Phyllosticta decidua Ell. & Kell. On Eupatorium sessili-
folium . Prairie du Chien.
Septoria anemones Desm. On Anemone virginiana.
Potosi. I have seen no record of the occurrence of the
parasite on this host. The specimen corresponds with some
of those on A. nemorosa.
In the provisional list Aster Shortii was recorded as a
host of Septoria solidaginicola Pk. A recent collection on
this host made at Lynxville, July 17, 1926, however, is of
quite a different type as is indicated by the following notes.
Spots suborbicular, purple to brown above, lighter brown
below, often containing a small white arid spot, mostly 4-5
mm. in diameter; pycnidia few, scattered, globose to de¬
pressed-globose, extending through the leaf from epidermis
to epidermis but the ostiole epiphyllous where the wall is
274 Wisconsin Academy of Sciences, Arts, and Letters.
thickest, 100-165//, in diameter; sporules usually somewhat
curved, tapering toward the acute apex, 50-85*3^. This
is referred to Septoria atropurpurea Pk.
From a collection on Aster sagittifolius made at Madison
September 29, 1925 the following notes were made: spots
brown, suborbicular to angular, 5-10 mm. in diameter,
sometimes confluent ; conidiophores amphigenous, more
abundant below, fasciculate from a more or less stromatoid
base, hyaline, straight or somewhat curved, cylindrical to
subulate-cylindrical, usually simple, continuous, 12-30 x 2-
31/2/z.; conidia hyaline, cylindrical, mostly 1-3 septate, some¬
times catenulate, 15-50 X2-3/z. The longer conidia become
uniformly 3-septate. This has been labeled Ramularia vir-
gaureae Thuem. Perhaps it bears relation to Cercospora
viminei Tehon.
Cercosporella pyrina Ell. & Ev. On Pyrus ioensis. Brod-
head. In this collection the conidia are 25-50 X6-8//. What
appear to be immature perithecia occur beneath the upper
epidermis.
Cercospora caricina Ell. & Dearn. On Carex folliculata.
Millston.
Cercospora umbrata Ell. & Hoi. On Bidens connata.
Bridgeport and Camp Douglas. On Bidens vulgata. She¬
boygan Falls.
A collection on Ambrosia psilostachya (Brodhead, Sept.
13, 1926) bears Cercospora on spots which are amphi¬
genous, immarginate, at first yellowish green, then yellow
and finally brown with death of the included tissue, 2-5 mm.
in diameter, often confluent; conidiophores sometimes fas¬
ciculate, sometimes erect racemose branches from superficial
brown, repent hyphae or from scandent hypae that ascend
the trichomes, sub-hyaline, straight or somewhat tortuous
or geniculate, continuous, rarely branched, 20-40x4-6//;
conidia hyaline, obclavate-cylindrical, straight or somewhat
curved, 50-100x4//. The common Cercospora on Ambrosia
trifida has been referred to Cercospora racemosa Ell. &
Mart. (Fungi Columbiani 2117) and Cercospora ferru-
ginea Fckl. (F. Col. 3207). The former reference was
Davis — Notes on Parasitic Fungi in Wisconsin . XV. 275
based largely upon finding infected Teucrium and Ambrosia
growing together but field observation in Wisconsin sug¬
gests that the propinquity was casual. The specimen has
been labeled Cercospora ferruginea Fckl. It will probably
be some time before the relationships of the brown Cer-
cosporas on Compositae become known. In a growth of
the kind referred to here there is room for argument as to
where mycelium ends and condiophores begin. On Am¬
brosia trifida the conida are sometimes terete, constricted
at the septa, moniliform or finally catenulate. Apparently
it has never been found on the ubiquitous Ambrosia
artemisiae folia.
Uromyces plumbarius Pk. Uredinia on Gaura biennis.
Brodhead.
Puccinia vilfae Arth. & Hoi. Aecia ( Aecidium verbeni -
colum Ell. & Kell.) on Verbena urticae folia and V. stricta.
Prairie du Chien. This was abundant especially on the
latter host. The telial stage on Sporobolus asper was re¬
corded in the provisional list as Puccinia sydowiana Diet.,
a binomial that had been earlier proposed by Zopf for an¬
other rust.
Puccinia muhlenbergiae Arth. & Hoi. On Muhlenbergia
sylvatica. Blue River.
Additional Species
Not previously reported as occurring in Wisconsin.
Peronospora phlogina Diet. & Hoi. This was found in
very small quantity, with oospores, on Phlox divaricata at
Blue River.
Venturia gaultheriae Ell. & Ev. On Gaultheria procum -
bens. Camp Douglas.
Taphrina filicina Rostr. On Cystopteris fragilis. Brod¬
head.
A collection on leaves of Acer saccharum seedlings made
at Sheboygan was taken in the field for Gloeosporium sac-
charinum. On examination, however, it proved to be a
Septoria and one that does not agree with any species of
276 Wisconsin Academy of Sciences, Arts , and Letters.
which I have knowledge. The following notes were made :
On large dead leaf areas 2-5 cm. across which are reddish
brown, becoming pale with age; pycnidia epiphyllous, scat¬
tered, depressed-globose, 70-130/x in diameter, wall usually
thick and black about the ostiole ; sporules acicular,
straight, long acuminate, continuous, 20-30 x 1-2/x. On
leaves of seedling Acer saccharum. Sheboygan, Wiscon¬
sin, August 26, 1926. For the purpose of filing in the her¬
barium this has been provisionally referred to Septoria
seminalis Sacc. as forma orthospora . I have seen no speci¬
mens of this species which is said to occur on cotyledons of
Acer campestris and to have falcate sporules. The status
of this parasite awaits further information.
Septoria dodecatheonis n. sp. Spots dark olivaceous,
varying from circular or angular 2-4 mm. in diameter to
elongate 1 cm. in length, often confluent and with the death
of intervening and surrounding leaf tissue forming con¬
siderable areas; pycnidia small, numerous, scattered, glo¬
bose to ovoid or even flask-shaped, wall black, compact, os¬
tiole more or less prominent, 50-80/x in diameter; sporules
hyaline, straight, 20-40 x 1-1 On leaves of Dodeca-
theon Meadia. Blue River, Wisconsin, June 18, 1926. The
material is not quite mature and the mature sporules are
probably somewhat larger. The appearance under a hand
lens suggests stromata of a Hyphale rather than pycnidia.
[Collected in 1927 at Madison and New Glarus.]
Of a collection on Geum made at Sheboygan August 26,
1926, the following notes were made: Spots definite, cir¬
cular or subcircular, olivaceous with a narrow darker bor¬
der above, lighter below, 3-5 mm. in diameter, sometimes
confluent, conidiophores fasciculate, hyaline, straight,
curved or bent, sometimes denticulate near the apex, simple,
continuous, often congested, 10-30 x 1-2/*; conidia hyaline,
slender, straight, acute, continuous, 25-50 x lVia-2^4/*.
This has been referred to Ramularia gei (Fckl.) Lindau.
The species is evidently a variable one ranging from an Ov-
ularia to a Cercosporella type.
In the provisional list all specimens of Aecidium on He-
lianthus were referred to Puccinia helianthi Schw. In
North American Flora 7 : 754, a Wisconsin specimen on
Davis — Notes on Parasitic Fungi in Wisconsin. XV. 277
Helianthus was referred to Nigredo junci (Desm.) Arth.
( Uromyces junci (Desm.) Tul.). This form appears to be
not uncommon on Helianthus in Wisconsin but no uredinia
or telia of Uromyces junci (Desm.) Tul. are known to occur
in the state but U. junci-tenuis Syd. is common and U. jun -
ci-ejfusi Syd. is sometimes abundant.
Ramularia dispersa n. sp. Immaculate; conidiophores
fasciculate, hyaline, straight, simple, continuous, 20-35 x
3 fi; conidia hyaline, catenulate, fusoid to cylindrical,
straight, continuous, 10-23 x 2-3 fx. On Napaea dioica .
Browntown, Wisconsin, September 22, 1926: The white
tufts are widely scattered over the lower surface of the
unmodified leaves. In this collection the leaves contain
many holes apparently due to gnawing insects which sug¬
gests that the fungus may have been scattered by insects.
Aecidium trillii Burr. On Trillium grandiflorum. Bal¬
sam Lake.
Herbarium,
University of Wisconsin,
April, 1927.
NOTES ON PARASITIC FUNGI IN WISCONSIN. XVI
J. J. Davis
It is perhaps well to repeat that this series of notes is
supplemental to a list of parasitic fungi in Wisconsin pub¬
lished in the Transactions of the Wisconsin Academy of
Science, Arts and Letters 172: 846-984. Names of hosts
are according to Gray’s New Manual of Botany, seventh
edition.
Peronospora urticae (Lib.) DBy. was collected in 1883
at Kirkland (now Devils Lake) by Trelease and at La
Crosse by Pammel. These localities are 80 miles apart.
No further collections appear to have been made in Amer¬
ica except a record by Harkness & Moore of its occurrence
in California but there appears to be no specimen extant.
It has recently come to my attention that Geranium mac -
ulatum was recorded as a host of Erysiphe polygoni DC. in
the Provisional List while the parasite is really Sphaero-
theca humuli (DC.) Burr. While looking the matter up
in the herbarium I found that the specimen of Sphaero-
theca humuli (DC.) Burr, on Geranium in Ellis & Ever¬
hart Fungi columbiani continued 1438 was labeled Erysiphe
polygoni DC.
Because of the previous use of the name Keithia for a
genus of Labiatae by Bentham Maire replaces Keithia Sacc.
by Didymascella Sacc. & Maire and makes new combina¬
tions accordingly {Bull. Soc. Nat. Hist. Afrique du Nord.
18 : 120/*, Abstract in Review of Applied Mycology 7 :
59.) Keithia appears to have been used as a generic name
still earlier by Sprengel, applied to some phanerogamous
plant.
Pycnidia of Phyllosticta viticola (B. & C.) Thuem. ( Ph .
labruscae Thuem.) occur in which the sporules are mostly
4-7 x 2-3/*. This may be microconidial rather than im¬
mature.
280 Wisconsin Academy of Sciences, Arts, a7id Letters.
In the 34th Report of the State Museum of Natural His¬
tory (1881), p. 45, Peck described a leaf parasite of Apocy-
num androsaemi folium under the name Septogloeum apocyni
n. sp. and gave figures of a spotted leaf and of sporules on
Plate I, fig. 2 & 3. Because of the presence of a thin wall
surrounding the hymenium this was transferred to Stag-
onospora in Transactions of the Wisconsin Academy of Sci¬
ence, Arts and Letters 192: 699 (1919). In Hedivigia
58: 25 (1917) Bubak, apparently unaware of Peck’s pub¬
lication, described the parasite as Dearnessia apocyni n.
gen. & n. sp. The thinness of the pycnidial wTall and the
presence of superficial hyphae hardly seem to be generic
characters. Sydow, Fungi exotici exs. 749, represents the
Fungus.
In the provisional list a parasite of Gentiana andrewsii
was recorded under the name Leptothyrium gentianaeco-
lum (DC.?) Baeumler. Wisconsin specimens were distrib¬
uted in Ellis & Everhart North American Fungi 2766 la¬
beled Phyllosticta ( Depazea ) gentianaecola (DC.). In El¬
lis & Everhart’s North American Phyllostictas it was in¬
cluded under the name Phyllosticta gentianaecola with
Depazea gentianaecola given as a synonym. In North
American Flora 6: 30, it is Phyllosticta gentianaecola
(DC.) Ellis & Ev. In '‘Notes” X, pp. 272-3 it was sug¬
gested that it might be Asteromu gentianae Fckl. In an
attempt to clear the matter up material was sent to the
European mycologist, Dr. F. Petrak, who thinks it to be
distinct and suggests the new binomial Asteromella an¬
drewsii nom. nov. The ascogenous state that seems to be
connected with it being Mycosphaerella andrewsii Sacc.
As stated in “Notes” X there is one collection on Gentiana
puberula.
In July, 1927 a collection was made at Portage that ap¬
pears to be a better developed state of the parasite that
was described in Trans. Wis. Acad. 9: 99 under the name
Septoria brevispora Ell. & Davis (Syll. Fungorum 18:
396). In this collection the spots become sordid white, ex¬
cept the peripheral portion, the pycnidia dark brown with
a black ring around the pore and the sporules 15-30x2
%-4/a. On staining, a median division of the cytoplasm
Davis - — Notes on Parasitic Fungi in W isconsin. XVI. 281
appears. The host of the Portage collection was a coarse
grass without fructification which may well be Bromus cili -
atus or a cognate species. In the collections made at Ra¬
cine, of which there are 4, development may have been ar¬
rested by death of the host tissues.
In a collection of Septoria on leaves of what is perhaps
Populus nigra from Sauk City (Aug. 5, 1927) most of the
spots become finally round, white and arid, 1-2 mm. in
diameter. The sporules are biseptate only the short ones
having but one septum. This seems to be intermediate be¬
tween S . Populi Desm. and S. musiva Pk. and is of interest
in connection with the suggestion in “Notes” I, p. 83, of a
single variable species.
A specimen from this collection was sent to Dr. F. Pe-
trak who identified it as Septoria populi Desm. and stated
that S. musiva Pk. is a form of that species.
In a collection of Septoria negundinis E. & E. from Arena
the pycnidia are effused over areas that sometimes retain
the green color until full maturity of the parasite. Some
of the sporules exceed 50 /* in length. The hosts are door-
yard trees.
A collection made at Carmel, California, by Mrs. Effie S.
Spalding communicated by Dr. B. M. Duggar which I refer
to this species shows sporules 20-38 x 3-4/*, 3-sept ate.
In making comparison with specimens of Septoria davisii
Sacc. in the Davis herbarium the following notes as to size
of sporules were found on the packets: “33-40 x 2/*.
Type of Septoria canadensis Ell. & Davis,” “36-80/* long,”
“30-70 x 2-3/*,” “33-52 x li/2-2/* .” In the specimen that
was being compared they were 33-36 x 2/*. The host ap¬
pears to be what is now known as Solidago altissima L.
The parasite is probably not distinct from Septoria fumosa
Pk.
In addition to the typical form of Septoria rudbeckiae
Ell. & Hals. Rudbeckia laciniata bears a form in which the
spots are orbicular to angular, white and arid with a very
narrow dark margin, 1-2 mm. in diameter, sometimes con¬
fluent. The sporules are straight, 35-70 x 1/*.
282 Wisconsin Academy of Sciences , Arts , and Letters.
Gloeosporium apocryptum Ell. & Ev. was described as
having sporules 5-12 x 2*4-5/*. Wisconsin collections
show sporules up to 17 x Ip. In germination on a slide
dark “appressoria” were produced about 10 x 6p.
Dr. F. J. Seaver kindly compared Tuberculina argilJacea
Davis (“Notes” XI, p. 293) with Gloeosporium rubi Ell. &
Ev. ( Journ . Mycol. 4: 52) and found that they appear to
be the same. It is not typical of either genus. The refer¬
ence to Tuberculina was based largely on its apparent rela¬
tion to Caeoma.
Dr. Petrak states that a Wisconsin specimen labeled Ram-
ularia uredinis (Voss) Sacc. is Ramularia rosea (Fckl.)
Sacc. (Ann. Mycol. 25 : 222) . It was noted in the 1st and
3d supplementary lists of parasitic Fungi of Wisconsin that
it sometimes appeared on leaves of Salix on which no sori
of Melampsora were seen. What appears to be the same
organism has been collected on Caeoma-infected leaves of
Populus deltoides as was recorded in “Notes” IV, p. 679.
Typical Ramularia rosea (Voss) Sacc. is common in Wis¬
consin on leaves of various species of Salix not bearing Mel¬
ampsora.
In the transactions of the Wisconsin Academy of Science
Arts and Letters 16: 762-3 Ramularia paulula was pub¬
lished as a new species on Elodes (Hypericum) virginica.
It was afterwards found that the host was probably Lysi-
machia thyrsiflora and the parasite Ramularia lysimachiae
Thuem., hence it was omitted from the Provisional list.
In the description of Cladosporium humile (“Notes” V,
702) it was stated that the conidiophores are epiphyllous.
They occur on the lowTer surface of the spots also.
The record of Cercospora zebrina Pass, on Trifolium du-
bium in “Notes” XII, p. 160 was an error. It should be
Cercospora medicaginis Ell. & Ev. if that is distinct, on
Medicago lupulina. The collection was made hastily at a
junction point while changing trains.
While studying the germination of the spores of leaf
smut of Glyceria Bauch found that the sporidia of Usti-
lago longissima (Sow.) Tul. were of two classes as shown
Davis — Notes on Parasitic Fungi in Wisconsin. XVI. 283
by their behavior in conjugation while those of the var.
macrospora were of three kinds, (z eitschr. /. Botanik
15: 241 et seq.) On morphological grounds Liro consid¬
ered the variety specifically distinct, proposing for it the
binomial Ustilago davisi. ( Usilagineen Finnlands 1: 80).
This illustrates the different conceptions of the limits of
a group of organisms that shall be considered a species.
When the varietal distinction was first made the host was
known as Glyceria fluitans; later it was segregated under
the name G. septentrionalis.
A specimen of Enlyloma compositarum Farl. on Lepa-
chys pinnata collected at New Glarus June 20, 1927 shows
unusual development of conidia ranging up to 70 x 2/x.
In ‘‘Notes” XIV record was made of the development of
Puccinia on Andropogon furcatus in the greenhouse in the
spring of 1925 using aeciospores of Aecidium fcilcatae Arth.
on Amphicarpa monoica as the inoculum.
In 1926 similar infection was secured from Aecidium
xanthoxyli Pk. except that the uredospores were of the Puc¬
cinia pustulata type. Record of this is made in “Notes”
XV issued herewith.
In conversation with Dr. E. B. Mains he told me that he
had once infected Polygala Senega with Puccinia andropo -
gonis but that the return inoculation on Andropogon failed
and he did not publish for that reason. In the spring of
1927 two lots of rusted Andropogon furcatus that had been
overwintered out-doors were anchored in a locality where
Polygala Senega was growing but where no Aecidium had
been seen on it. One lot had been obtained at Blue River
and about it no aecia appeared. The other lot came from
Dill and around it Aecidium appeared on the Polygala.
With these aecia Andropogon furcatus was infected in the
greenhouse resulting in uredinia and a few telia. The
uredospores were of the typical P. andropogonis type.
Sheldon reported that he had brought about development
of Uromyces on Sisyrinchium by infection from aeciospores
of Aecidium houstoniatum Schw. on Houstonia caerulea.
He also reported that he had tried several times to inocu¬
late Sisyrinchium with aeciospores from Houstonia purpu -
284 Wisconsin Academy of Sciences , Arts, and Letters.
rea both in the field and in the greenhouse but without defi¬
nite success ( Torreya 9: 54-5). Aecidium houstoniatum
Schw. occurs in Wisconsin on Houstonia longifolia but no
rust has been found on Sisyrinchium although it has been
looked for in the vicinity of infected Houstonia. In June,
1927 an attempt was made to infect Sisyrinchium obtained
from two sources with Aecidium on Houstonia but although
the conditions appeared to be favorable no infection re¬
sulted.
In the Preliminary List of Parasite Fungi of Wisconsin
by William Trelease (Trans. Wisconsin Academy of Sci¬
ence Arts & Letters 6 (188), Uredo sp. (No. 216) on
Mimulus ringens and Aecidium pentstemonis Schw. (No.
240) on the same host were reported as having been col¬
lected at La Crosse by Pammel. I am informed that Dr.
Pammel’s specimens, which are in the herbarium of Iowa
State College, have been examined by Dr. Arthur who con¬
cluded that the host is Epilobium color atum, the Uredo
Pucciniastrum pustulatum (Pers.) Diet, and the Aecidium
that of Puccinia peckii (De Toni) Kell. The latter has not
been recorded as occurring on Epilobium in Wisconsin.
Additional Hosts for Wisconsin
It is customary to add each year another host for Syn-
chytrium in Wisconsin. For 1927 it is Steironema cilia-
tum and the parasite is referred to Synchytrium aureum
Schroet. In this collection which was made at Blue River,
the wall surrounding the sorus is often very irregularly
thickened. This may have been due to the character of the
season which was very dry.
Albugo Candida (Pers.) 0. Kuntze
On Cardamine rhomboidea. Madison.
Spotting of the leaves of water cress, Radicula nasturti-
um-aquaticum, growing in the waters of springs emptying
into lake Wingra at Madison due to Cercospora nasturtii
Pass, is not uncommon. In 1927 however abundant leaf
spotting occurred which was due to Peronospora parasitica
(Pers.) Tul.
Davis — Notes on Parasitic Fungi in Wisconsin. XVI. 285
Sphaerotheca humuli (DC.) Burr.
On Geum strictum. Sauk City.
Erysiphe cichoracearum DC.
On Lactuca spicata. Madison.
Microsphaera alni (Wallr.) Wint. ( M . ele'vata Burr.)
On Catalpa (cult.) Cudahy. A. C. Burrill.
Acanthostigma occidentale (E. & E.) Sacc.
Stylosporous state on Artemisia ludoviciana. Token
Creek. The stylospores are about 5 x 2p.
Stagonospora intermixta (Cke.) Sacc.
On Agrostis alba. La Valle. In this collection the pyc-
nidial wall is of uniform thickness, the 7-septate sporules
43-48 x 31/2
Septoria caricinella Sacc. & Roum.
On Carex straminea and C. Bebbii. New Glarus.
Septoria violae West.
On Viola blanda. Hollandale.
Septoria lysimachiae West.
On Lysimachia quadrifolia. La Valle. Scanty and
poorly developed in this collection.
Colletotrichum graminicolum (Ces.) Wilson.
On culms of Cinna arundinacea. Arena.
On Sorghastrum nutans. Browntown.
But few sporules in this collection. The spots are purple
bordered.
Marssonina coronaria (Ell. & Davis) Davis. ( Mar so -
nia coronariae Sacc. & Dearn.)
On Pyrus ioensis. Trempealeau and Highland.
Ramularia decipiens Ell. & Ev.
On Rumex obtusifolius. Browntown.
Fusicladium effusum Wint. var. carpineum Ell. & Ev.
On Garpinus caroliniana. Hollandale.
[Demaree finds that the conidia of Fusicladium effusum
Wint. on Carya are catenulate and refers the species to
Cladosporium with a new description ( Journ . Ag’l Re¬
is
286 Wisconsin Academy of Sciences , Arts , and Letters.
search 37: (86) The form on Carpinus appears to be of
the same character.]
Cercospora helianthi Ell. & Ev.
On Helianthus strumosus. Browntown. In this collec¬
tion the conidiophores are hypophyllous, tortuous, multi-
septate, ranging up to 200 /*, in length.
Isariopsis albo-rosella Sacc.
On Stellaria aquatica. South Wayne and Gratiot.
Abundant at the latter station but not well developed at the
time of collection. It was thought to be Graphiothecium
pusillum (Fckl.) Sacc. until septate conidia were found.
In 1911 Puccinia graminis Pers. was observed on a single
plant in a colony of Glyceria grandis near Butternut in
northern Wisconsin. In 1927 this experience was repeated
at New Glarus in southern Wisconsin. At Blue River a
plant of Leersia virginica was found bearing Puccinia im-
patienti-elymi Arth. (Klebahn) — P. impatientis Arth.
= P. ely mi-imp atientis Davis. This rust is abundant in
the locality on Elymus but in only the single instance has it
been found on Leersia although 3 species of the genus occur
there and it has been a favorite collecting ground in pre¬
vious years. There appears to be no record of its occur¬
rence on this host elsewhere. Such occurrences may lead
one to surmise that development of a species on a new host
may occasionally take place and that this may sometimes be
followed by adaptation, physiological changes due to the
new substratum resulting in isolation followed in time by
morphological differentiation and the development of a new
species.
Acerates floridana is recorded as an aecial host of Dicaeo-
ma jamesianum (Pk.) Arth. = Puccinia bartholomaei
Diet, in Wisconsin in North American Flora 7 : 320.
Puccinia seymouriana Arth.
Aecia on leaves and stems of Asclepias ovalifolia. New
Glarus.
Puccinia peckii (De Toni) Kell.
Aecia on Oenothera rhombipetala. Mazomanie.
Davis — ■ Notes on Parasitic Fungi in Wisconsin . XVI. 287
Solidago graminifolia should be included in the list of aecial
hosts of Puccinia extensicola Plowr. in Wisconsin.
Chrysomyxa pyrolae (Pers.) Diet.
Uredinia on Pyrola americana. Friendship.
Coleosporium solidaginis (Schw.) Thuem.
On Callistephus chinensis (cult.)
Oconto (A. C. Burrill)
Additional Species
Not hitherto recorded as occurring in Wisconsin.
Investigation of host relations of obligate fungous para¬
sites has shown that they are usually adapted to the kind of
host upon which they are found. That the adaptation may
be relative rather than absolute has also been shown, and
that it is not necessarily accompanied by morphological dif¬
ferentiation is well known. By some mycologists new spe¬
cies have been proposed based only upon adaptation to a
particular species of host. The practice of considering
these as physiological races seems the better one and hence
they are not recorded in these notes in which the hosts have
already been given.
Peronospora oxybaphi Ell. & Kell.
On Oxybaphus nyctagineus . Browntown and Brodhead
along the railroad. The conidia range up to SO/* in length
and 22/* in breadth and are ovate in outline.
At a station near Friendship a few plants of Melampy-
rum lineare were found, only 3 or 4 as I remember, and
none were found elsewhere in that locality. On the young
upper leaves of these plants was a parasite regarding which
the following notes were made: Forming a sordid gray
coating on the lower leaf surface ; conidiphores 300-400 /*
long, 3-4 times branched, usually dichotomously, but some¬
times the branches are at an obtuse or even a right angle,
ultimate branchlets short, straight, acute, divergent; con-
cidia fuscous, elliptical, acute at base, 24-38 x 20/*. No
oospores were found.
Plasmopam melampyri Buchoitz I know only from the de¬
scription in Saccardo : Sylloge Fungorum 21 : 861 but
28S Wisconsin Academy of Sciences , Arts , and Letters.
the Wisconsin plant seems to be referable to that species.
Bucholtz appears to have been in doubt as to whether it
should be placed in Peronospora or Plasmopara. To the
writer it furnishes additional evidence that they are con¬
generic and that the latter was given generic rank before
the evidence was all in. As this parasite has apparently
been found in the single locality in Russia and the very
restricted station in Wisconsin only, one might suspect that
it is well on the way toward extinction. However recrudes¬
cences have been known to occur in such cases.
[This was collected the following year at Radisson. As
in the previous collection it was confined to the lower sur¬
face of the upper leaves and would escape detection when
the hosts were observed from above as the infected leaves,
at least at the time of collection, were not changed in ap¬
pearance by the presence of the parasite. The germination
of the conidia has not been observed and oospores have not
been seen.]
Fig. 1. A conidiophore and three conidia of Plasmopara melam -
pyri Bucholtz. Drawn by E. M. Gilbert with the aid of camera lucida.
The lower surface of the older leaves of the same plants
of the Friendship collection bore Ramularia melampyri Ell.
& Dearn. I have not seen a specimen of Fusidium melam¬
pyri Rostr. but the description suggests that it is the same
species as the one described by Ellis & Dearness.
Davis — Notes on Parasitic Fungi in Wisconsin. XVI. 289
What has been known as Ascochyta pisi Lib. has been di¬
vided by Linford & Sprague into 2 species one retaining
the name and the other a pycnidial state of Mycosphaerella
pinodes (Berk. & Blox.) Stone with which the typical As¬
cochyta pisi Lib. is not connected. They also found a third
form which they designated temporarily Mycosphaerella
pinodes microform (?) ( Phytopath.il : 390-91.)
[For the “microform” the binomial Ascochyta pinodella
n. sp. has been proposed by Leon K. Jones (N. Y. State
Ag’l. Exp. Station, Geneva, Bull. 547, p. 10) and for the
pycnidial stage of Mycosphaerella pinodes , Ascochyta pino¬
des (Berk. & Blox.) n. comb. ( loc . cit. p. 4.) ]
While examining leaves of Anemone canadensis for the
presence of oospores of Plasmopara pygmaea it was found
that in addition to the oospores there were present pycni-
dia about 100//, in diameter containing hyaline oblong spor-
ules 18-30 x 3-7//, 1-2 septate. Whether or not this is
Stagonospora anemones Pat. I do not know.
Septoria polygalae Pk.
On Polygala Senega. New Glarus.
This name was given by Peck to a species occurring on
Polygala pauciflora. Subsequently he collected a form on
Polygala Senega bearing Aecidium also to which he gave
the name Septoria consocia. As described this differs
from the former in its shorter sporules which otherwise are
of the same character. On the Wisconsin collection there
is no Aecidium and the sporules are intermediate in length
(23-33 x 1/2-1//) as between the two descriptions.
Septoria calystegiae West.
On Convolvulus arvensis . Madison.
Septoria polemonii Thuem. ( S . polemoniicola Ell. & Mart.)
On Polemonium reptans. Gratiot and Token Creek.
5. polemoniicola Ell. & Mart, appears to have been sepa¬
rated mostly because of the small white spots but European
specimens bear similar spots. Ellis & Martin apparently
knew von Thuemen’s species from the description only.
While some kinds of plants are infected by various fungi
others are seldom found to be attacked. In Wisconsin no
290 Wisconsin Academy of Sciences, Arts, and Letters .
parasite had been seen on Lithospermum. In July 1927,
however, a few spotted leaves of L. canescens were ob¬
served near Portage. The spots are dark brown, more
conspicuous below, subangular, 1-3 mm. in diameter.
There are small spore bodies, usually near the periphery of
the spot on the upper surface and few in number bearing
tufts of hyaline conidia 40-60 x 2/3/a which are usually
curved, often strongly so. The better developed spore
bodies appear to be imperfectly formed pycnidia. It seems
possible that this was a chance infection by a Septoria that
normally develops on another host. I do not now recall
whether the infection was confined to a single plant but
the small number of infected leaves suggests that it may
have been. For the purpose of filing this has been labeled
Septoria lithospermi nom. herb.
Colletotrichum trifolii Bain is reported as occurring in
Wisconsin by John Monteith, Jr., in U. S. Department of
Agriculture Technical Bulletin No. 28.
Gloeosporium achaeniicola Rostr.
On epicarp of Heracleum lanatum. Mineral Point. The
type of this species was on fruit of Pastinaca sativa.
Cylindrosporium pimpinellae C. Massal. var. pastinacae
Sacc.
On Pastinaca sativa. Gratiot.
In this collection the conidia are mostly 60-70 x 3-4/a
curved, tapering both ways from the middle, becoming 1-3
septate.
Didymaria puncta n. sp.
On elongate light brown to sordid white areas; conidio-
phores borne on the outer aspect of globose brownish black
tubercles, congested, hyaline varying from 15 x 2-3/a to ob¬
solete; conidia apical, hyaline, straight, long fusoid, acute,
20-30 x 3/a developing a median septum. On stems and
bracts of Sisyrinchium campestre. New Glarus, Wiscon¬
sin, June 17, 1927. The tubercles are closely spaced, 40-
60/a in diameter. The conidia fall away readily and it was
not determined whether they are catenulate or not.
Davis — Notes on Parasitic Fungi in Wisconsin . XVI. 291
From collections on leaves of Saxifraga pennsylvanica
made at Portage, June 6 and 8, 1927 the following notes
were made : Spots circular, brown sometimes more or less
tinged with purple, becoming cinereous with the develop¬
ment of conidia and often zonate above, uniformly brown
below, 4-8 mm. in diameter; conidiophores arising from
dark tuberculoid stromata, congested, hyaline, straight or
somewhat flexuose above, often subulate, 10-30 x 2-4/*;
conidia hyaline, filiform, straight or lax, 30-100 x 1-2/*.
I do not doubt that this is the species collected at Dover,
Norway to which Rostrup gave the name Cercosporella sax -
ifragae which I know only from description. There is a
discrepancy in the length of the conidiophores which are
described as being long but that is a character that cannot
be stressed in this group.
Of a collection on leaves of Tanacetum vulgar e from a
garden in Highland the following notes were made : Spots
becoming definite and black; conidiophores amphigenous,
hyaline, fasciculate, simple, mostly straight and continuous,
20-40 x 3-4/* ; conidia hyaline, straight, cylindrical but us¬
ually with a slight taper, obtuse at each end, 24-43 x 3%-
5/*. On referring to the description of Ramularia tanaceti
J. Lind in the Sylloge Fungorum the statement was met
that the conidia are acute at each end. In Rabenhorst
Kryptog amen flora Lindau gave a description which he
stated was more complete than Lind’s and resulted from
examination of original material. This description is the
same as the one in the Sylloge including the statement that
the conidia are acute at each end. In the original descrip¬
tion by Lind {Ann. My col. 3: 431), however, the conidia
are described as being obtuse at both ends as they are in the
Wisconsin collection. There seems to be no question of the
identity of the Wisconsin parasite and Ramularia tanaceti
J. Lind. The similarity in the names of the three mycolo¬
gists Lind, Lindau and Lindroth sometimes leads to con¬
fusion. For instance in Oudemans’ Enumeratio Systema¬
tica Fungorum this species is attributed to Lindroth,
Cercospora thaspiicola n. sp.
Spots pale alutaceous becoming sordid white, angular,
limited by the veinlets, with a narrow brown border, 2-5 x
292 Wisconsin Academy of Sciences , Arts , Letters .
1-2 mm.; conidiophores amphigenous, fuligenous tinted,
caespitose, straight with an oblique apex and a shoulder,
simple, continuous, 35-50 x 4-5/a ; conidia hyaline, fusoid-
cylindrical, obtuse, straight or slightly curved, 1-3 septate,
37-73 x 4-6/a. On Thaspium aureum. South Wayne, Wis¬
consin, June 27, 1927. Ellis & Everhart, North American
Fungi 2585 is labeled Cercospora thaspii n. sp. On leaves
of Thaspium trifoliatum Wilmington, Delaware, November
1889. A. Commons. Apparently no description was ever
published. The specimen shows circular or sometimes an¬
gular brown spots which become white and arid with a
brown border 1-2 mm. in diameter. The conidiophores
are in small fascicles, erect or diverging, straight or but
little flexuose, deep brown, pluri-septate, 50-100 x 5-6/a.
As one might expect of a specimen collected so late in the
season and so long ago, no conidia were found. In Gray’s
Manual of Botany, edition 5, which was in use in 1889,
Zizia cor data DC. was referred to Thaspium trifoliatum as
var. apterum and it is a question as to whether N. A. F.
2585 is on Thaspium or Zizia and in the absence of conidia
what its relation may be to Cercospora ziziae Ell. & Ev. It
may easily be that collections of a species made in June and
in November would show differences but there does not
seem to be justification for stating that N. A. F. 2585 and
the Wisconsin collection are conspecific.
Although Puccinia muhlenbergiae Arth. & Hoi. has been
known in Wisconsin no Aecidium on Malvaceae had been
seen until 1927 when Aecidium napaeae Arth. & Hoi. was
collected on Napaea dioica at Gratiot, New Glarus, Brown-
town and South Wayne. These localities are in southern
Wisconsin. The host seems to be spreading along the rail¬
roads especially where they pass over low ground and are
ballasted with cinders.
Aecidium sparsum n. sp.
Pycnia hypophyllous, scattered, brown; aecia hypophyl-
lous, scattered, 14-1/3 mm. in diameter; peridium white,
lacerate, the lobes reflexed and caducous; spores yellow
globose, thin-walled, finely and sparsely verrucose, 18-22/a
in diameter. On Galium tinctorium on bottom lands of the
Davis — Notes on Parasitic Fungi in Wisconsin. XVI. 293
Wisconsin river opposite Sauk City, July 2, 1925 (Weber &
Davis,) July 11, 1927. The distribution of the aecia sug¬
gests systemic infection but there is no distortion or dis¬
coloration of the host.
Herbarium,
University of Wisconsin,
April, 1928.
NOTES ON PARASITIC FUNGI IN WISCONSIN. XVII
J. J. Davis
H. Sydow has proposed that Phacidium balsameae Davis
(“Notes” VIII, p. 424) be made the type of a new genus
Stegopezizella. (Ann. Mycol. 22 : 392)
To the description of Peziza (Mollisia) singularia Pk.,
afterwards referred to Pseudopeziza, was added the sug¬
gestion “Perhaps a form of Pseudopeziza ranunculi Fckl.”
(35th Report, p. 142) . This is doubtless the case and that
is a form of Fabraea ranunculi (Fr.) Karst. Septate
spores have been seen in Wisconsin material. The com¬
mon host in Wisconsin is Ranunculus pennsylvanicus but it
occurs also on R. septentrionalis.
Marsonia baptisiae E. & E. on Baptisia leucantha,
founded on a specimen from Iowa, was published in the
Bulletin of the Torrey Botanical Club 24: 291 [1897] with
the statement that the conidia become “faintly uniseptate”.
In “Notes” II, p. 103, this was recorded as occurring in
Wisconsin with the statement that “septation of the spor-
ules seems doubtful”. In “Notes” XIV p. 186 Ascochyta
baptisiae n. sp. was described with conidia uniseptate or
occasionally with 2 or 3 septa. In 1928 a collection was
made at Mazomanie in which the sporules are predomi¬
nantly 2-3 septate. Re-examination of the specimen re¬
ferred to Marssonina baptisiae (E. & E.) shows that the
sporules are borne in pycnidia with walls about two cells
thick. It is evident that the three collections represent
degrees of development of the same parasite. I have not
seen an authentic specimen of Marsonia baptisiae E. & E.,
but the description indicates that it is the same as the Wis¬
consin species. I am therefore labeling all of the speci¬
mens Stagonospora baptisiae (E. & E.) n. comb.
A parasite of Shepherdia canadensis was recorded in
“Notes” II, p. 105, under the name Cylindrosporium shep-
296 Wisconsin Academy of Sciences, Arts, and Letters .
herdiae Sacc. After examination of a collection from Idaho
Dearness changed the name to Septoria shepherdiae (Sacc.)
Dearn. ( Mycologia 20: 238). With the Wisconsin record
was a suggestion that this is close to Septoria argyraea
Sacc.
In the 29th Report of the State Botanist of New York a
fungus on living leaves of Trillium erythrocarpum was des¬
cribed under the name Vermicularia concentrica P. & C. n.
sp. The description was followed by the statement that
Judge Clinton had sent in a variety on Viola rotundifolia.
In the Sylloge Fungorum 3: 232, Saccardo changed the
name to Vermicularia peckii Sacc. and designated the form
on Viola var. violae-rotundifoliae. In the Report of the
State Botanist for 1919 the variety was raised to specific
rank as Vermicularia violae-rotundifoliae (Sacc.) House.
In the Botanical Gazette 26: 96-97 Miss Stoneman des¬
cribed and figured Volutella violae n. sp. on Viola cucullata.
In the Wisconsin “Notes” XI, p. 297 Colletotrichum vio-
larum n. sp. was recorded on Viola scabriuscula. This has
since been found on other species of Viola in Wisconsin.
Examination of authentic material from Dr. House and of
Miss Stoneman’s description and figures indicates that the
Wisconsin species is not distinct and that all of them should
be referred to Colletotrichum because of their acervulous
character. I am therefore labeling them Colletotrichum vi¬
olae-rotundifoliae (Sacc.) n. comb. This accords with the
results of Miss Duke's investigation in which she found that
Vermicularia is not distinct from Colletotrichum {Trans.
Brit. My col. Soc. 133). I quite agree with her that Colleto¬
trichum, which was properly described, should be retained
instead of Vermicularia as the name of the genus.
In “Notes” III, p. 263 a collection on Streptopus roseus
was referred to Vermicularia liliacearum West. There are
also Wisconsin specimens on Uvularia grandiflora (Nelson
Dewey State Park and Maiden Rock) , Oakesia sessili folia
(Plover and White Lake) and Smilacina stellata (Wood¬
man). All of these are on leaf spots and I am now desig¬
nating them Colletotrichum peckii (Sacc.) n. comb.
Of Ellis & Everhart North American Fungi 2778, Ver¬
micularia helianthi E. & K. n. sp. on Helianthus rigidus,
Davis — Notes on Parasitic Fungi in Wisconsin. XVII. 297
Manhattan, Kan. autumn 1887, W. T. Swingle I have seen
no description.
Collet otrichum helianthi Davis, “Notes” I, p. 88 is appar¬
ently the same parasite.
In a collection of Cladosporium humile Davis on Acer
saccharinum the development is on the peripheral portion
of Rhytisma spots sometimes extending on to the ascocarp.
The collection was made at Sauk City September 6, 1928.
In the description of Cercospora viciae Ell. & Hoi.
(Journ. My col. 1: 5 & 39) the conidiophores are said to be
“short, 25-30X3-4/*”. The Wisconsin specimens that have
been referred to this species bear conidiophores up to 80/*
long and the conidia sometimes attain 70/* in length. The
parasite is rather common on Lathyrws venosus, L. och-
roleucus and to a less extent on L. palustris in the northern
part of the state. There is but one specimen on Vicia
(V. caroliniana) which is from southeastern Wisconsin and
bears equally long conidiophores.
The smut of Polygonum sagittatum recorded in the pro¬
visional list under the name Sphacelotheca hydropiperis
(Schum.) DBy. is separated from that species by Liro and
given the name Sphacelotheca granosa Liro. He gives as
a distinguishing character the more prominent verrucosity
of the spore wall. (U stilagineen Finnlands I: 148-150)
In one of the “Notes” reference was made to an Aecidium
on Linaria canadensis with the suggestion that as the host
is a Scrophulariacea the Aecidium is probably connected
with Puccinia andropogonis Schw. Attempts to infect
Andropogon furcatus with the aeciospores in the green¬
house have failed, however, when the conditions appeared
to be favorable for infection.
In June, 1928 plants of Amphicarpa monoica and Poly¬
gala Senega were exposed to infection from Puccinia on
Andropogon furcatus in the greenhouse. Abundant aecia
were produced on the Polygala but none on the Amphi¬
carpa. The telia were obtained at New Glarus near Poly -
gala Senega plants that had borne aecia earlier in the sea¬
son. As far as the experiments have gone they suggest
298 Wisconsin Academy of Sciences, Arts, and Letters.
that Puccinia andropogonis Schw. is specialized as to aecial
hosts but perhaps not as to telial.
ADDITIONAL HOSTS
Peronospora Candida Fckl. (P. androsaces Niessl). On
Androsace occidentalis. On a hill opposite Prairie du Sac.
Plasmopara viticola (B. & C.) Berl. & De Toni. On Vitis
bicolor. Platteville.
Claviceps purpurea (Fr.) Tul. Sclerotia on Glyceria
borealis. Haugen.
Phyllachora graminis panici Shear. On Panicum ten -
nesseense. Sauk County.
Septoria bromi Sacc. On Bromus incanus. Couderay.
In this collection the more or less depressed pycnidia are
100-165 x 60-100/x, the sporules filiform, straight or but
little curved, 37-60X1-11^.
Septoria astragali Rob. On Vida americana. Radisson.
In this collection the pycnidia are very imperfect and the
sporules grow out to a length of 120-200/a resembling
Cylindrosporium. This appears to be but the second col¬
lection in the state, the first having been reported by Tre-
lease in 1884.
Of a collection on leaves of Solidago altissima made at
Mazomanie, Sept. 20, 1928 the following notes were made :
Spots subcircular, brown becoming cinereous above, 3-5
mm. in diameter, forming large areas through death of
the intervening tissue ; pycnidia epiphyllous, not prominent,
succineous, globose, wall thin, of thin flat polygonal cells,
80-100/a in diameter ; sporules hyaline, straight, cylindrical
with rounded ends, developing a median septum, 20-27X6-
7/a. Shorter continuous sporules are assumed to be im¬
mature. This is referred to Ascochyta compositarum Davis
(Trans. Wis. Acad. 192: 700).
A collection of the aecial stage of Puccinia bartholomaei
Diet, on Acerates longi folia made by Pammel at La Crosse
in 1883 appears to be the only one of that stage that has
been made in Wisconsin. [This was collected in 1929 on
Acerates lanuginosa near Prairie du Sac.]
Davis — Notes on Parasitic Fungi in Wisconsin. XVII. 299
Puccinia rubigo-vera (DC.) Wint. ( P . elymi West., P.
agropyri E. & E.). On Bromus incanns. Couderay. In
this collection the sori are epiphyllous and small and most
of the teliospores distorted.
Sclerotium deciduum Davis. On Bidens frondosa.
Radisson.
Additional Species
Dothichloe atramentaria (B. & C.) Atk.
On culms of Calamagrostis canadensis in a cranberry
marsh at Cranmoor (E. E. Honey).
Phyllosticta limitata Pk.
On Pyrus Malus. Lancaster. (V. H. Young & J. J.
Davis.)
Phlecspora mori (Lev.) Sacc.
On Morns alba . Madison. In all of the specimens of
this parasite that I have seen the conidia are borne extern¬
ally on a stroma of subcuticular origin and they might be
referred to Cylindrosporium.
In August 1928 small collections were made at Platteville
and Shullsburg of a parasite on leaves of Abutilon Theo-
phrasti from which the following notes were made : Spots
circular, alutaceous with a dark border, alike on both sur¬
faces of the leaf; when young 1-8 mm. in diameter, with
maturity becoming more irregular in outline, up to 6 mm.
in length and becoming lacerate; pycnidia epiphyllous,
depressed-globose, variable in size up to 200^ in diameter
with a black bordered pore up to 30,u across; sporules
hyaline, straight, continuous, 6-16 X 2-3!/2M. It may
be that this is Ascochyta abutilonis Hollos, in which the sep-
tation of the sporules is said to come late, of which I have
not seen an authentic specimen.
Leptostroma pinastri Desm.
On Finns Banksiana . Camp Douglas.
Sphaceloma symphoricarpi Barrus & Horsfall. ( Phyto-
path . 18:799.)
On Symphoricarpos racemosus (Cult.). Madison. The
spots are abundant on the fruit but acervuli are rare.
300 Wisconsin Academy of Sciences, Arts , and Letters.
The preservation of specimens of the more delicate Hy-
phales is often unsatisfactory because of the falling away
of the conidia and to a much less extent, the conidiophores.
When on flat leaves and carefully treated the pressure tends
to hold them in place and the detached ones do not wander
far. From cylindrical surfaces, however, they disappear.
This is by way of apology for calling attention to two species
of which the material in hand leaves something to be de¬
sired.
Cercospora eleocharidis n. sp.
Globose, black, subepidermal, stromatoid bodies which
sometimes extend into the epidermis from which spring fas¬
cicles of fuligenous fertile hyphae which reach the surface
and usually extend 3-15//, beyond; conidia apical, hyaline,
narrow cylindrical, straight or somewhat curved, 30-70 X
2 /a. On more or less extensive dying and dead areas on
culms of Eleocharis palustris. Brill, Wisconsin, July 23,
1928. This has been observed in Wisconsin for a number
of years but the collections have hitherto been discarded
when they came from the press because the conidia had
fallen away.
Cercospora jirnci n. sp.
Black, scattered, intraepidermal, stromatoid bases 20-30/a
wide; conidiophores fuligenous tinted, nodulose, 15-30 X
3-4/a ; conidia subolivaceous, obclavate-cylindrical, acute,
straight, 60-75 X 4-5/a. On dying and dead areas or entire
leaves of J uncus brevicaudatus. Brill, Wisconsin, July 23,
1928.
Cercospora setariae Atk.
On Setaria glauca. Browntown.
Cercospora parvimaculans n. sp.
Spots brown, angular, ^-l1/^ mm., often confluent and
sometimes with a white center on the upper surface; coni¬
diophores in small scattered fascicles or solitary, hypophyl-
lous, more or less fuligenous, becoming tortuous, denticu¬
late, and septate, 50-100 X 3-5/a; conidia subhyaline,
straight or curved, cylindric-obclavate to flagelliform, 100-
180 x 4-6/a. On leaves of Solidago serotina. Sauk City,
Davis — Notes on Parasitic Fungi in Wisconsin. XVII. 301
Wisconsin, September 6, 1928, type. Other collections are
from Lone Rock, Blue River and Wauzeka July and August
1921, but no description was published.
Cercospora silphii Ell. & Ev.
On Silphium laciniatum, Shullsburg. Silphium terebin -
thinaceum, Madison. In these collections the spots are pur¬
plish-brown to black and the tufts inconspicuous. The
conidiophores are often shorter than those of the type as
described. A form on the former host was designated
var. laciniatae by Tehon & Daniels ( Mycologia 19: 128).
Coleosporium terebinthinaceae (Schw.) Arth.
Uredinia and a few telia on Silphium perfoliatum . Lan¬
caster. In this collection the uredospores are 23-40/* in
length. That this species is a permanent member of the
Wisconsin flora is questionable.
Uromyces alopecuri Seym.
On Alopecurus geniculatus aristulatus . Haugen.
Puccinia physostegiae Pk. & Cl.
On Physostegia parviflora. Abundant at a station in the
bottom lands opposite Sauk City in 1928 attacking especially
the upper leaves and the inflorescence, destroying the
flowers.
While the fungous growths on “honey dew” on leaf sur¬
faces are not parasites their effects are probably ill. An in¬
teresting form that occurs in Wisconsin on leaves of various
plants is apparently the one to which Woronochin gave the
name Sclerotiomyces colchicus (Ann. My col. 24: 234). The
orbicular flattened sclerotia strongly resemble perithecia.
Herbarium,
University of Wisconsin,
April, 1929.
20
302
Wisconsin Academy of Sciences, Arts, and Letters.
INDEX TO PARASITES AND HOSTS REFERRED TO
IN NOTES XV, XVI, XVII
The names of Fungi are in italics.
Abutilon Theophrasti, 299
Acanthostigma occidentale (E. & E.)
Sacc., 285
Acer saccharinum, 297
Acer saccharum, 275
Acerates floridana, 286
Acerates lanuginosa, 298
Acerates longifolia, 298
Aecidium falcatae Arth., 272, 283
Aecidium houstoniatum Schw., 283,
284
Aecidium napaeae Arth. & Hoi., 292
Aecidium pentstemonis, 284
Aecidium sparsum n. sp., 292
Aecidium trillii Burr., 277
Aecidium verbenicolum E. & K., 275
Aecidium xanthoxyli Pk., 272, 283
Agrostis alba, 285
Albugo Candida (Pers.) 0. K., 284
Alopecurus geniculatus aristulatus,
301
Ambrosia psilostachya, 274
Ambrosia trifida, 274
Amphicarpa monoica, 272, 283, 297
Andropogon furcatus, 272, 283, 297
Androsace occidentalis, 298
Anemone canadensis, 289
Anemone virginiana, 273
Apocynum androsaemifolium, 280
Artemisia ludoviciana, 273, 285
Asclepias ovalifolia, 286
Ascochyta abutilonis Hollos, 299
Ascochyta baptisiae Davis, 295
Ascochyta compositarum Davis, 298
Ascochyta pinodella L. K. Jones, 289
Ascochyta pinodes (Berk. & Blox.)
L. K. Jones, 289
Ascochyta pisi Lib., 289
Ascochyta wisconsina Davis, 269
Aster sagittifolius, 274
Aster Shortii, 273
Asteroma gentianae Fckl., 280
Asteromella andrewsii Petr., 280
Baptisia leucantha, 295
Barbarea stricta, 271
Bidens connata, 274
Bidens frondosa, 299
Bidens vulgata, 274
Bromus ciliatus, 281
Bromus incanus, 298, 299
Calamagrostis canadensis, 299
Callistephus chinensis, 287
Cardamine rhomboidea, 284
Carex Bebbii, 285
Carex folliculata, 274
Carex straminea, 285
Carpinus caroliniana, 285
Catalpa, 285
Cercoseptoria crataegi n. comb., 271
Cercospora armoraciae Sacc., 271
Cercospora caricina Ell. & Dearn., 274
Cercospora cheiranthi, 271
Cercospora crucifer arum E. & E., 271
Cercospora eleocharidis n. sp., 300
Cercospora ferruginea Fckl., 274
Cercospora flagellifera Atk., 271
Cercospora helianthi E. & E., 286
Cercospora junci n. sp., 300
Cercospora latens E. & E., 272
Cercospora lespedezae Ell. & Dearn.,
272
Cercospora medicaginis E. & E., 282
Cercospora nasturtii Pass., 271, 284
Cercospora nasturtii barbareae Sacc.,
271
Cercospora parvimaculans n. sp., 300
Cercospora racemosa E. & M., 274
Cercospora setariae Atk., 300
Cercospora silphii E. & E., 301
Cercospora thaspii E. & E., 292
Cercospora thaspiicola n. sp., 291
Cercospora umbrata Ell. & Hoi., 274
Cercospora viciae Ell. & Hoi., 297
Cercospora viminei Tehon., 274
Cercospora ziziae E. & E., 292
Cercosporella mirabilis Pk., 271
Cercosporella pyrina E. & E., 274
Cercosporella saxifragae Rostr., 291
Chrysomyxa pyrolae (Pers.) Diet., 287
Cinna arundinaceae, 285
Cladosporium effusum (Wint.) Dem-
aree, 285
Cladosporium humile Davis, 282, 297
Claviceps purpurea (Fr.) Tul., 298
Coleosporium solidaginis (Schw.)
Thuem., 287
Coleosporium terebinthinaceae
(Schw.) Arth., 301
Collet otrichum graminicolum (Ces.)
Wilson, 285
Colletotrichum helianthi Davis, 270,
297
Colletotrichum peckii (Sacc.) n. comb.,
296
Colletotrichum trifolii Bain., 290
Colletotrichum viola e-rotundif oliae
(Sacc.) n. comb., 296
Colletotrichum violarum Davis, 296
Convolvulus arvensis, 289
Cylindrosporium crataegi E. & E.,
271
Davis — Notes on Parasitic Fungi in Wisconsin. XVII. 302a
Cylindrosporium f r ax ini ( Ell. &
Kell.) Ell. & Ev., 270
Cylindrosporium pimpinellae pastina-
ceae Sacc., 290
Cylindrosporium shepherdiae Sacc.,
295-6
Cystopteris fragilis, 275
Dearnessia apocyni Bubak, 280
Dicaeoma jamesiana (Pk.) Arth., 286
Didymaria puncta n. sp., 290
Didymascella Sacc. & Maire, 279
Dodecatheon Meadia, 276
Dothichloe atramentaria (B. & C.)
Atk., 299
Eleocharis palustris, 300
Entyloma compositarum Farl., 283
Epilobium coloratura, 284
Erysiphe cichoracearum DC., 273, 285
Erysiphe polygoni DC., 279
Eupatorium sessilifolium, 273
Fabraea ranunculi (Fr.) Karst., 295
Fraxinus oregana, 270
Fusicladium ejfusum carpineum E. &
E., 285
Fusidium melampyri Rostr., 288
Galium tinctorium, 292
Gaultheria procumbens, 275
Gaura biennis, 275
Gentiana Andrewsii, 280
Gentiana puberula 280
Geranium maculatum, 279
Geum, 276
Geum strictum, 285
Gloeosporium achaeniicola Rostr., 290
Gloeosporium apocruptum E. & E.,
282
Gloeosporium niveum Davis, 270
Gloeosporium rubi E. & E., 282
Glyceria borealis, 298
Glyceria fluitans, 283
Glyceria grandis, 286
Glyceria septentrionalis, 283
Helianthus, 270, 276
Helianthus rigidus, 270
Helianthus strumosus, 286
Heracleum lanatum, 290
Houstonia caerulea, 283
Houstonia longifolia, 284
Houstonia purpurea, 283
Isariopsis albo-rosella Sacc., 286
Juncus brevicaudatus, 300
Keithia, 279
Lactuca spicata, 285
Lathyrus ochroleucus, 297
Lathyrus palustris, 297
Lathyrus venosus, 297
Leersia virginica, 286
Lepachys pinnata, 283
Leptostroma pinastri Desm., 299
Leptothyrium gentianaecolum (DC.?)
Baeumler, 280
Lespedeza capitata, 271
Linaria canadensis, 297
Lithospermum canescens, 290
Lysimachia quadrifolia, 285
Mar sonia baptisiae E. & E., 295
Marsonia coronariae Sacc. & Dearn.,
285
Marssonina coronaria (Ell. & Davis)
Davis, 285
Medicago lupulina, 282
Melampyrum lineare, 287
Meliiotus alba, 273
Microsphaera alni (Wallr.) Wint., 285
Microsphaera elevata Burr., 285
Mimulus ringens, 284
Morus alba, 299
Muhlenbergia sylvatica, 275
Mycosphaerella andrewsii Sacc., 280
Mycosphaerella p in o d e s (Berk. &
Blox.) Stone, 289
Napaea dioica, 277, 292
Oakesia sessilifolia, 296
Oenothera rhombipetala, 286
Oxalis corniculata, 272
Oxalis cymosa ? 272
Oxybaphus nyctagineus, 287
Panicum tennesseense, 298
Pastinaca sativa, 290
Peronospora androsaces Niessl, 298
Peronospora Candida Fckl., 298
Peronospora ficariae Tul., 273
Peronospora oxybaphi E. & K., 287
Peronospora parasitica (Pers.) Tul.,
284
Peronospora phlogina Diet. & Hoi.,
275
Peronospora urticae (Lib.) DBy., 279
Peziza ( Mollisia ) singularia Pk., 295
Phacidium balsameae Davis, 295
Phleospora mori (Lev.) Sacc., 299
Phleospora oxyacanthae (Kze. &
Schm.), 271
Phlox divaricata, 275
Phragmidium potentillae-canadensis
Diet., 272
Phyllachora graminis panici Shear,
298
Phyllosticta decidua E. & K., 273
Phyllosticta gentianaecola (DC.) E.
& E., 280
Phyllosticta iridis E. & M., 269
Phyllosticta labruscae Thuem., 279
Phyllosticta Umitata Pk., 299
Phyllosticta viticola (B. & C.)
Thuem., 279
Physostegia parviflora, 301
Pinus Banksiana, 299
Plasmopara melampyri Bucholtz,
287-8
Plasmopara py g mae a (Ung.)
Schroet., 289
302b
Wisconsin Academy of Sciences, Arts, and Letters.
Plasmopara viticola (B. & C.) Berl. &
DeToni, 298
Polemonium reptans, 289
Polygala pauciflora, 289
Polygala Senega, 283, 289, 297
Polygonum sagittatum, 297
Populus deltoides, 282
Populus nigra, 281
Potentilla canadensis, 272
Pseudopeziza m e di c a g ini s (Lib.)
Sacc., 273
Pseudopeziza ranunculi Fckl., 295
Pseudopeziza singularia Pk., 295
Psoralea argophylla, 272
Puccinia agropyri, E. & E., 299
Puccinia andropogonis S c h w., 283,
297-8
Puccinia bartholomaei Diet., 286, 298
Puccinia elymi, West., 299
Puccinia extensicola Plowr., 287
Puccinia graminis Pers., 286
Puccinia helianthi Schw., 276
Puccinia impatienti-elymi Arth., 286
Puccinia muhlenbergiae Arth. & Hoi.,
275, 292
Puccinia peckii (De Toni) Kell., 284,
286
Puccinia physostegiae Pk. & Cl., 301
Puccinia pustulata Arth., 272, 283
Puccinia rubigo-vera (DC.) Wint.,
299
Puccinia seymouriana Arth., 286
Puccinia sorahi Schw., 272
Puccinia sydowiana Diet., 275
Puccinia vilfae Arth. & Hoi., 275
Pucciniastrum pustulatum (Pers.)
Diet., 284
Pyrola americana, 287
Pyrus ioensis, 274, 285
Pyrus Malus, 299
Radicula Nasturtium-aquaticum, 284
Ramularia decipiens E. & E., 285
Ramularia dispersa n. sp., 277
Ramularia gei (Fckl.) Lindau, 276
Ramularia melampyri Ell. & Dearn.,
288
Ramularia rosea (Fckl.) Sacc., 282
Ramularia tanaceti J. Lind, 291
Ramularia uredinis (Voss) Sacc., 282
Ramularia virgaureae Thuem., 274
Ranunculus abortivus, 273
Ranunculus pennsylvanicus, 295
Ranunculus septentrionalis, 295
Rudbeckia laciniata, 281
Rumex obtusifolius, 285
Sambucus canadensis, 269
Saxifraga pennsylvanica, 291
Sclerotiomyces colchicus Woronochin,
301
Sclerotium deciduum Davis, 299
Septogloeum apocyni Pk., 280
Septogloeum fraxini Hark., 270
Septoria anemones Desm., 273
Septoria argyraea Sacc., 296
Septoria astragali Rob., 298
Septoria atropurpurea Pk., 274
Septoria brevispora Ell. & Davis, 280
Septoria bromi Sacc., 298
Septoria calystegiae West., 289
Septoria canadensis Ell. & Davis, 281
Septoria caricinella Sacc. & Roum.,
285
Septoria consocia Pk., 289
Septoria davisii Sacc., 281
Septoria dodecatheonis n. sp., 276
Septoria fumosa, Pk., 281
Septoria lysimachiae West., 285
Septoria musiva Pk., 281
Septoria negundinis E. & E., 281
Septoria pachyspora Ell. & Hoi., 269
Septoria paupera Ellis, 270
Septoria polemonii Thuem., 289
Septoria polemoniicola E. & M., 289
Septoria polygalae Pk., 289
Septoria populi Desm., 281
Septoria rudbeckiae Ell. & Hals., 281
Septoria seminalis Sacc., 276
Septoria shepherdiae (Sacc.) Dearn.,
296
Septoria sisymbrii Ellis, 271
Septoria solidaginicola Pk., 273
Septoria violae West., 285
Setaria glauca, 300
Shepherdia canadensis, 295
Silphium laciniatum, 301
Silphium perfoliatum, 301
Sliphium terebinthinaceum, 301
Sisyrinchium, 283
Sisyrinchium campestre, 290
Smilacina stellata, 296
Solidago altissima, 281, 298
Solidago graminifolia, 287
Solidago serotina, 300
Sorghastrum nutans, 285
Sphaceloma symphoricarpi Barrus &
Horsfall, 299
Sphacelotheca granosa Liro, 297
Sphacelotheca hydropiperis (Schum.)
D By., 297
Sphaerotheca humuli (DC.) Burr.,
279, 285
Sporobolus asper, 275
Stagonospora anemones Pat., 289
Stagonospora apocyni (Pk.) Davis,
280
Stagonospora baptisiae (E. & E.) n.
comb., 295
Stagonospora intermixta (Cke.) Sacc.,
285
Stegopezizella , 295
Steironema ciliatum, 284
Stellaria aquatica, 286
Davis — Notes on Parasitic Fungi in Wisconsin. XVII. 302c
Streptopus roseus, 296
Symphoricarpos racemosus, 299
Synchytrium aureum Schroet., 284
Tanacetum vulgare, 291
Taphrina filicina Rostr., 275
Taphrina hiratsukae Nishida, 269
Taphrina struthiopteridis Nishida,
269
Thaspium aureum, 292
Thaspium trifoliatum, 292
Trillium erythrocarpum, 296
Trillium grandiflorum, 277
Tuberculina argillacea Davis, 282
Uromyces alopecuri Seym., 301
Uromyces junci (Desm.) Tub, 277
Uromyces junci-effusi Syd., 277
Uromyces junci-tenuis Syd., 277
Uromyces plumbarius Pk., 275
Ustilago davisi Liro, 283
Ustilago longissima (Sow.) TuL, 282
Uvularia grandiflora, 296
Venturia gaultheriae E. & E., 275
Verbena stricta, 275
Verbena urticaefolia, 275
Vermicularia concentrica P. & C., 296
Vermicularia helianthi E. & K,, 270,
296
Vermicularia liliacearum West., 296
Vermicularia peckii Sace., 296
Vermicularia peckii var. violae-rotun-
difoliae Sacc., 296
Vermicularia violae-r o tundi foliae
(Sace.) House, 296
Vicia americana, 298
Vicia caroliniana, 297
Viola blanda, 285
Viola cucullata, 296
Viola rotundifolia, 296
Viola scabriuscula, 296
Volutella violae Stoneman, 296
Vitis bicolor, 298
Zanthoxylum americanum, 272
,
A CYTOLOGICAL STUDY OF FERTILIZATION IN
ACHLYA HYPOGYNA COKER AND
PEMBERTON
George Olds Cooper
The question of fertilization in the Saprolegniaceae has
long aroused interest among botanists. Pringsheim
(1855), while he did not actually observe fertilization, be¬
lieved that it took place. Later workers, de Bary (1884),
Humphrey (1892), Hartog (1892-1895) studying various
genera, came to the conclusion that fertilization did not oc¬
cur. However, Trow (1895, 1904) disagreed with Hartog
and showed without doubt that fertilization does occur in
some species of the Saprolegniaceae.
Since Trow’s work further evidence that fertilization
does occur has been demonstrated by Claussen (1908),
Miicke (1900), Kasanowsky (1911), Carlson (1925), and
Couch (1925).
Materials and Methods
The fungus used as a basis for this study was secured
from dead minnows in Tomahawk Lake, Wisconsin. Cul¬
tures were grown on boiled corn endosperm in distilled wa¬
ter. As the early cultural studies were made in the field,
it was impossible to make single spore isolations ; therefore
growth from single sporanges was taken as a basis for clas¬
sification. Cultural studies were made from time to time,
and from Coker's “Classification of the Saprolegniaceae”
(1925) the species was identified as Adilya, hypogyna Coker
and Pemberton. This determination has been since verified
by Mr. J. V. Harvey.
The material was preserved in formol-acetic-alcohol, im¬
bedded by the paraffin method, and sections were cut to 5
microns in thickness. Overstaining with Haidenhain’s
iron-alum haematoxylin with light green as a counter stain
gave the best results.
304 Wisconsin Academy of Sciences , Arts, and Letters .
Observation and Discussion
In Achyla hypogyna Coker and Pemberton, the oogones
arise from hyphae on short lateral branches of the myce¬
lium. In the formation of the oogone there is a streaming
of the cytoplasm into the hyphal tip, forming in it a marked
swelling. The streaming continues until the swelling has
caused the hyphal tip to become almost spherical. A cross
wall is now laid down at the base of this spherical tip and
the young oogone is thus formed. The cytoplasm at this
stage is multinucleate ; conspicuous vacuoles are present,
surrounded by cytoplasm containing granules (mitochon¬
dria) of varying size and shape. A small dark body re¬
sembling a nucleolus was observed in each of the nuclei.
This body seems to be identical with that described by
Hartog as a chromatin body, by Trow as a combination of a
chromosome and a nucleolus, by Davis, Claussen, Mucke, and
Carlson as a nucleolus. Shortly, there appears a definite
vacuole near the center of the oogone. As this vacuole in¬
creases in size many of the nuclei migrate to the periphery
of the oogone and show signs of degenerating. The re¬
maining nuclei are found near the vacuole and vary as to
number, from a few to as high as sixteen in some oogones.
These nuclei undergo only one mitotic division (fig. 1).
Many of the nuclei formed as a result of this division de¬
generate, leaving seemingly only those that are to function
as nuclei of the eggs. These remaining nuclei enlarge
slightly and are readily recognized (fig. 2). That there is
only one division occurring within the oogone agrees with
the conclusions of Davis, Claussen, Mucke, Kasanowsky,
and Carlson in regard to closely related species. Trow
(1904) describes the occurrence of two divisions in the
oogone, the second of which he believed to be a reduction
division.
The central vacuole after mitosis enlarges greatly by fur¬
rowing the cytoplasm (fig. 2). Further furrowing extends
to the periphery of the oogone, leaving apparently small
rounded portions of the cytoplasm extending into the va¬
cuole. These partially rounded cytoplasmic masses are
known as egg initials and their number depends upon the
Cooper — Study of Fertilization in Achlya Hypogyna. 305
number of nuclei that remained following the divisions and
subsequent degeneration of some of the nuclei. This is
evidently the case, because in each of the initials only one of
these nuclei was found. Occasionally, however, an egg was
found containing two nuclei. In no instance were extra
nuclei found scattered about in the egg, unless they were
degenerating (fig. 3). Davis found in these egg initials a
body which he termed a coenocentrum and which he be¬
lieved was instrumental in the rounding up of the eggs. He
referred to this rounding up as the “balling” of the cyto¬
plasm. This body acted as a dynamic center which drew the
cytoplasm to it and rounded it up into a spheroid mass. Miss
Carlson and the writer found what might be termed a coen¬
ocentrum, but observed it only a few times. Claussen also
mentioned such a body. From continued observation of
the formation of the eggs and the infrequency of the find¬
ing of the “coenocentrum,” the writer is inclined to the be¬
lief that the coenocentrum is not the active agent. This
body was observed in eggs that had nearly rounded up. It
lay partially surrounding the nucleus and several fibrils
radiated from it. The agent that is instrumental in the
formation of the eggs is the central vacuole, according to
the present writer, and by its continual furrowing of the
cytoplasm the eggs are cut out. Later, when the vacuolation
has gone to completion and the egg initials have rounded
up, they come to lie close together so that their sides appear
to be in contact. Shortly afterwards they separate and re¬
main suspended in the liquid in the oogone.
During the formation of the eggs in the oogone, anthe-
rids have been approaching the oogone from the neighbor¬
ing hyphae. The name of this species was derived from the
fact that the antheridial branch often arises from the hy-
pha immediately beneath the oogone and attaches itself to
it. Antherids were observed to attach themselves to the
oogones after the formation of the eggs, but more fre¬
quently at the stage of the egg initial. The development
of the male gametes within the antherid is very similar to
that in the oogone. After the cytoplasm has streamed to
the tip of the antherid, a cross wall is laid down. The cyto¬
plasm is multinucleate and, as in the oogone, many of the
nuclei migrate to the periphery and degenerate. The re-
306 Wisconsin Academy of Sciences , Arts, and Letters .
maining nuclei undergo one mitotic division and many of
these nuclei also degenerate. Those nuclei remaining were
scattered in the cytoplasm and seem to enlarge slightly.
The antherid, upon coming in contact with the oogone,
may indent the oogone wall ; whether it finally ruptures the
wall and discharges its contents into the oogone was not ob¬
served. As is more often the case, the antherid adheres to
the oogone and sends one to several fertilization tubes
through the oogone wall. These fertilization tubes grow
until they come in contact with one or more eggs. How¬
ever, when a fertilization tube comes in contact with an
egg, other fertilization tubes from this same antherid either
grow to neighboring eggs or indicate signs of collapse.
The fertilization tubes vary as to length and width. Some
have walls that are fairly smooth, while others are quite
irregular. Several nuclei are often present within them
and the nucleus that is most terminal appears slightly lar¬
ger than the others and is generally the nucleus that func¬
tions as the male gamete nucleus. Upon the discharge of
the male gamete nucleus into the egg the other nuclei in the
fertilization tube soon showed indications of degeneration.
Previous workers observed that the wall of an oogone in
a number of species varied in thickness. The oogone wall
in surface view, appears to be made up of many circular
thickened plates at the intersection of which the areas were
thin-walled. In section view, this gave the appearance of
thickened areas separated by a thin membrane. It was be¬
lieved for a long time that the fertilization tube penetrated
the thinner areas only, due probably to some chemotactic
stimulus. Couch, in his studies with the water molds, has
observed the fertilization tubes penetrating the thickened
areas as well as the thinner ones, and so believes that there
is no ground for the assumption that the fertilization tubes
penetrate the thinner areas only.
Before the fertilization tube is in contact with the egg it
may contain several nuclei, the more terminal one of which
has enlarged. This tube continues to grow until it comes
in contact with the egg and the terminal nucleus enlarges
still more and in doing so probably ruptures the tip of the
fertilization tube (fig. 4). The male gamete nucleus, as it
is now recognized, can be seen within the egg (fig. 6) . The
Cooper — Study of Fertilization in Adilya Hypogyna. 307
female gamete nucleus is also present and is recognized by
the fact that it is slightly larger and denser. A nucleolus
was not observed in either of the gamete nuclei. It was not
uncommon to find the female gamete nucleus not in the cen¬
ter of the egg. In these preparations, while it may not be
characteristic of the species, the writer observed that the
male gamete nucleus was invariably discharged into the egg
at the side farthest away from the female gamete nucleus.
The male gamete nucleus, upon its discharge into the
egg, migrates to the female nucleus and evidently unites
with it (fig. 6). The nuclei were easily recognized, one
slightly over-lapping the other. The penetration path left
by the male gamete nucleus as described by Couch in Lep-
tolegnia caudata , was not observed. Upon the union of the
gamete nuclei the egg becomes a zygote (fig. 6) . A wall
now appears about the zygote, the cytoplasm becomes finely
vacuolate and often several oil bodies of varying size are
present. The zygote when mature is slightly contracted.
The results of this cytological investigation seem to defi¬
nitely establish the presence of fertilization in Achlya hy¬
pogyna and add, therefore, an eighth species of the Sapro-
legniaceae wherein fertilization has been observed. Up to
the present, fertilization has been observed in three Sapro-
legnias by Trow (1895) and Claussen (1908) ; in seven Adi¬
lyas by Trow (1899, 1904), Davis (1905), Mucke (1908),
Carlson (1929), and by the writer; in an Aphanomyces by
Kasanowsky (1911) ; and in Leptolegnia by Couch (1925).
Owing to the large number of genera and species in the
Saprolegniaceae, there is opportunity for further work to
be carried on in this field.
The writer wishes to express to Dr. E. M. Gilbert his ap¬
preciation for the advice and encouragement received dur¬
ing the course of the work.
Bibliography
Carlson, M. S. 1929. Gametogenesis and fertilization in
Achlya racemosa Hildebrand. Ann. Bot. 43:111-117.
Claussen, P. 1908. uber Eintwicklung und Befruchtung
bei Saprolegnia monoica. Ber. d. Deutsch. Bot. Gesell.
26:144. Pis. 6-7.
308 Wisconsin Academy of Sciences , Arts , Letters.
Coker, W. C. 1923. The Saprolegniaceae, with notes on
other water molds. Chapel Hill, N. C.
Davis, B. M. 1903. Oogenesis in Saprolegnia. Bot. Gaz.
35:233 and 320. Pis. 9-10.
- - 1905. Fertilization in the Saprolegniales. Bot.
Gaz. 39:61.
Kasanowsky, V. 1911. Aphanomyces laevis de Bary. I
Entwicklung der Sexualorgane und Befruchtung. Ber.
d. Deutsch. Bot. Gesell. 29 :210. PI. 10.
Miicke, M. 1908. Zur Kenntnis d. Eientwicklung und
Befruchtung von Achlya polyandra de Bary. Ber. d.
Deutsch. Bot. Gesell. 26a:367. PI. 6.
Trow, A. H. 1905. Fertilization in the Saprolegniales.
Bot. Gaz. 39 :300.
Explanation of Plate 2
All figures were drawn with a camera lucida and with a Leitz 4-
ocular and a 1/12 oil immersion. Magnification 1500.
Contents of the oogones were drawn with a Zeiss 12 compensating
ocular and a 3 mm. aproehromatic oil immersion.
Fig. 1. Young oogone with the nuclei in the metaphase; degener¬
ate nuclei at the periphery.
Fig. 2. Central vacuole prior to the formation of the egg initials.
Fig. 3. Oogone containing four egg initials; antherids present.
Fig. 4. The approach of the fertilization tube containing the male
gamete nucleus to the egg.
Fig. 5. The fertilization tube has discharged the male gamete nu¬
cleus into the egg.
Fig. 6. The male gamete nucleus about to be discharged from
the fertilization tube. Zygote; gamete nuclei in contact.
TRANS. WIS. ACAD. - VOL. 24
PLATE 2
5
CYTOLOGICAL STUDIES ON THE SPORANGE DE¬
VELOPMENT AND GAMETOGENESIS IN
BREVILEGNXA DICLXNA HARVEY
George Olds Cooper
Introduction
Primary interest has been directed to the sexuality of the
Saprolegniaceae and for many years there has been consid¬
erable discussion as to whether or not fertilization does ac¬
tually occur.
Pringsheim (1855, 1858, 1860), working with various
members of the Saprolegniaceae, while he did not actually
observe fertilization, believed that it took place. Zopf
(1882, 1883) declared that Pringsheim did not have fertili¬
zation, but that what he saw were “spermamoeben” zoos¬
pores of a parasitic fungus. Cornu (1872) believed fertil¬
ization was present in Achlya yolyandra Hild. and Achlya
racemosa Hild. He demonstrated that the structures which
Pringsheim considered antherids were the sporanges of
parasites, probably of a Chytrid.
De Bary (1884) did not share Pringsheim’s views and in
his “Comparative Morphology and Biology’' stated that
while the antherid may approach the oogone and penetrate
it and even continue on through and pass out the other side,
it does not discharge its protoplasmic content.
Humphrey (1892) was the first investigator to fix, im¬
bed, cut, and stain his material. He described the behavior
of the nuclei within the oogones of Achlya americana Hum¬
phrey and Achlya apiculata de Bary. He stated that the
nuclei did not divide, but repeatedly fused in pairs until
each egg contained only one nucleus. He agreed with de
Bary that the fertilization tube may enter the oogone and
approach the egg, but that it does not discharge the male
gamete-nucleus.
Hartog (1892, 1895), staining in toto, agreed with Hum¬
phrey that the nuclei in the oogone fused until each egg
310 Wisconsin Academy of Sciences, Arts, and Letters.
contained a single nucleus and that there were no mitotic
divisions within the oogone. He, too, insisted that fertili¬
zation did not occur; antheridia approached the oogones,
but there was no discharge of their protoplasmic contents.
Trow (1895) disagreed with Hartog as to the fusion of
nuclei and as to whether there was an actual fertilization.
He described nuclear divisions in the oogones and antherids
of Saprolegnia diocia and observed where the fertilization
tube had discharged the male gamete-nucleus into the egg.
From 1896 to 1899 there was a bitter controversy between
Hartog and Trow as to the behavior of the nuclei and as to
the occurrence of fertilization, but they came to no agree¬
ment.
Davis (1904) worked with Saprolegnia mixta which he
believed to be apogamous. He described one division
within the oogonium. The chromosome number was four
and there were no centrosomes present. He found in the
early formation of the egg a structure which he called a
coenocentrum, from which delicate fibrillae radiated. The
coenocentra were formed de novo, one for each spore ori¬
gin. Its function, he believed, was concerned with the
“balling” or rounding up of the egg and that it exerted a
chemotactic influence over nuclei that were near it. These
nuclei were nourished by the coenocentrum as they in¬
creased in size. Generally one nucleus persisted in the egg,
but occasionally two or three and in such instances the eggs
were bi- or tri-nucleate.
Trow (1904) worked on Achlya polyandra Hildebrand
and observed a fertilization tube in direct open communica¬
tion with a young egg. In Achlya deb ary ana Humphey
the nuclei undergo two divisions within the oogone, the sec¬
ond of which Trow believed to be in the nature of a reduc¬
tion division. Fertilization occurred in this species.
Claussen (1908) with Saprolegnia monoica (Prings.) ,
Mucke (1900) with Achlya polyandra de Bary and
Kasanowsky (1911), with Aphanomyces laevia de Bary
have observed fertilization to be present.
Carlson (1925) with Achyla racemosa Hildebrand de¬
scribed the divisions of the nuclei within the oogone. She
believed that the central vacuole in the young oogone was
the active agent in the cutting out of the egg initials and
Cooper — Cytological Studies on Sporange Development . 311
not the coenocentrum described by Davis (1905). She ob¬
served a body that might be termed a coenocentrum, but
observed it only a few times. As complete evidence for
fertilization she described the discharge of the male game¬
te-nucleus into the egg, the migration of the male gamete-
nucleus to the female gamete nucleus, and finally the union
of the two.
Couch (1925) with Leptolegnia caudata de Bary ob¬
served fertilization in this species. The penetration path
left in the cytoplasm by the male gamete nucleus is of in¬
terest as it has not been previously recorded.
Materials and Methods
The material was secured from Mr. J. V. Harvey who
isolated it from soil gathered in the vicinity of Madison,
Wisconsin. Cultures were grown on boiled hemp seed in
distilled water. Single sport cultures were made and
within three days to a week oogonia became abundant.
The material was identified as Brevilegnia diclina Harvey.
It was fixed in formol-acetic-alcohol. The material was
imbedded by the paraffin method and sectioned to a thick¬
ness of 5 microns. Haidenhain’s iron-alum haemotoxylin
stain with a counter stain of light green gave the best re¬
sults.
Development of the Sporanges
The mycelium of Brevilegnia declina Harvey is dense and
rather opaque, the hyphae are straight and sparingly
branched. The sporanges appear within a day in cultures
growing on boiled hemp seed, at nearly all the hyphal tips,
but also arise from the same point in sympodial groups.
The writer, while cooperating with Mr. Harvey in the ex¬
amination of material, observed two characteristic types
of primary sporanges, one long and slender, with the spores
arranged in a single row except for a small cluster of spores
at the tip of the sporange, the other ovate to long club-
shaped as in Thraustotheca clavala (de Bary) Humphrey.
Secondary sporanges appear in dense sympodial clusters
terminally or frequently below the primary sporanges. As
in Thraustotheca (Weston, 1918), typical sporanges are
312 Wisconsin Academy of Sciences, Arts , and Letters.
broadly cl a vat e and lack papillae of dehiscence. The spo-
ranges of Brevilegnia develop on the dimunition of the food
supply, agreeing in this respect with the general law estab¬
lished by Klebs (1899).
Sporange formation begins with the gradual streaming
of the protoplasm to the hyphal tip which, as a result, be¬
gins to swell slowly. In time the accumulation of proto¬
plasm fills the sporange initial and a cross wall is then
formed across the base of the sporange. The contents now
begin to differentiate into spores. In the sporange having
the spores arranged in a single row in section view, there
appears at the tip of the sporange a vacuole which by en¬
largement and furrowing cuts out the spore initials (fig. 2) .
The formation of the spore initials in the remaining part
of the sporange is caused probably by a furrowing of vacu¬
olar and plasma membranes.
In longitudinal sections of the larger club-shaped type of
sporange, the vacuole first appears near the tip and con¬
tinues on down through the center of the sporange (fig. 5) .
Here, as in the other type of sporange, the enlargement
and furrowing of the vacuole cuts out the polygonal spore
initials. In both types of sporanges the spore initials are
at first distinct and then for a space become slightly indis¬
tinct, due probably to the readjusting of the spore initials
and the movement of the liquid in the vacuole. The spore
initials, now granular in appearance, form a wall about
themselves and become more and more distinct. Weston,
in Thraustotheca, states that the sporangiospores imbibe
water and swell, but in Brevilegnia, on the contrary, the
spores throw off water and the cytoplasm appears to con¬
tract. This is now evidently an encysted state. The com¬
pleted spores occupy the same position as did the spore
initials.
In this form, as in Thraustotheca, it is to be noted that
there is no evidence of an intersporal substance, a condition
in agreement with the observations of Rothert (1890) and
Humphrey (1893) for other Saprolegniaceae.
It is interesting to note that in the formation of the spore
initials the vacuole does not cut out uninucleate masses of
protoplasm as has been observed in all other of the Sapro¬
legniaceae (fig. 2). In section view the sporangiospores
Cooper — Cytological Studies on Sporange Development. 313
(aplanospores) show a multinucleate condition. The nuclei
are quite distinct, each with a definite nuclear membrane
and a conspicuous irregularly shaped nucleole. A fine
granular network appears to run from the nucleole to chro¬
matin material at the periphery of the nucleus. The nuclei
vary slightly in size; the smaller ones, however, show no
indications of degeneration. Weston uses the term spo-
rangiospore to indicate a non-motile spore, thereby distin¬
guishing it from the term zoospore. Since the sporangios-
pores are monoplanetic the term sporangiospore is not
quite applicable in this case. In Brevilegnia the writer
suggests the term aplanospore as the spores are non-motile
at all times.
The method of the liberation of the spores agrees par¬
tially with that of Thraustotheca which Weston believes
differs from that of any other genus in the Saprolegniaceae.
As has been mentioned, the papillae of dehiscence which are
present in most members of the family are lacking in Brev¬
ilegnia, and the aplanospores escape by the disintegration
of the sporange walls. In a cross section of the larger
club-shaped type of sporange, the spores are arranged
about the central vacuole. There is no rupture of the wall
by the swollen sporangiospore as in Thraustotheca, but
rather a disintegration of the wall here and there due in all
probability to either chemical or enzyme action (fig. 6).
The aplanospores are shed either singly or more commonly
in small groups of two or three, often with portions of the
sporange wall adhering to them. Upon leaving the spo¬
range they round up, clinging together. This is in accord¬
ance with Weston, who observes a distinct adhesion and
mutual attraction among the spores. Aplanospores remain
sticking to any hyphae with which they may come in con¬
tact. They now have definite membranes or walls with
the protoplasm slightly contracted (fig. 7). The cytoplasm
is finely granular with small vacuoles, nuclei many, 7-15
having observed in some instances.
The aplanospores float about in the water, showing abso¬
lutely no signs of motility. Germinating spores or zo=
ospores were not observed. Harvey believes that the
aplanospores undergo a period of rest before germination,
omitting the zoospore stage, and produce a germ tube di-
21
314 Wisconsin Academy of Sciences, Arts, and Letters.
rectly. Couch, however, succeeded in producing zoospores
from aplanospores by placing some of them in distilled
water to a pH of about 4, or filtering them through animal
charcoal. Gemmae were not observed by Harvey or the
writer.
Development of the Oogones
The oogones arise singly and apically from hyphae on
short lateral branches of the mycelium. In the formation
of the oogone there is a streaming of the protoplasm into
the hyphal tip, forming in it a marked swelling. The
streaming continues until the swelling has caused the hy¬
phal tip to become almost spherical. A cross wall is now
laid down at the base of this swelling and the oogone is thus
formed (figs. 8-10). The cytoplasm at this stage is multi-
nucleate; conspicuous vacuoles are present surrounded by
cytoplasm containing granules (mitochondria) of varying
size and shape. A conspicuous, dark-staining body re¬
sembling a nucleolus is observed in each of the nuclei.
This body seems to be identical with that described by Har-
tog as a chromatin body, by Trow as a combination of a
chromosome and a nucleolus, by Davis, Claussen, Mucke,
and Carlson as a nucleolus.
A vacuole remains in the center of the oogone after the
formation of the cross wall at the base. Many of the nu¬
clei migrate to the periphery of the oogone, where they very
soon show indications of degeneration. The remaining
nuclei appear to enlarge slightly and to stand out clearly.
These nuclei appear to undergo one mitotic division only.
That there is only one division occurring within the oogone
(figs. 8, 9) agrees with the results found by Davis, Claus¬
sen, Mucke, Kasanowsky, and Carlson in regard to closely
related species. Trow (1904) describes the occurrence of
two divisions in the oogone, the second of which he believes
to be a reduction division. Contrary to other workers who
state that the nuclei divide simultaneously, the writer ob¬
served nuclei in resting stage and in various stages of divi¬
sion within a single oogone (figs. 8, 9). The chromatin-
linin network becomes very massed in the prophase and
with the nucleole still quite prominent it is very difficult to
make accurate determinations of chromosome behavior.
Cooper — Cytological Studies on Sporange Development. 315
The metaphase has been observed frequently (figs. 8-10).
The chromosomes are massed together at the equator of an
intranuclear spindle. Few spindle fibers are present. As¬
ters with only the suggestion of rays could be seen (fig. 9) .
In the anaphase the chromosome masses are to be noted.
The chromosomes can be seen at this stage but they are
so crowded it is impossible to make accurate counts. The
writer believes the number to be comparatively few (4-8)
and in no instance as many as 14-16, mentioned by Claus-
sen. The telophase has been observed in a few instances
and only a massing of cytoplasm at the poles could be made
out.
After the nuclear divisions, the oogone evidently throws
off water because there is a marked contraction of the cyto¬
plasm from the wall. The vacuole in the center disappears
and the cytoplasm becomes uniform throughout. This ag¬
gregated mass of cytoplasm becomes now the single egg
which is constant in this species.
Formation of the eggs
Davis, working with an oogone containing several eggs,
found a body which he terms a coenocentrum and which
he thinks is instrumental in the rounding up of the eggs.
The cytoplasm in the oogone remains adherent to the wall
and the central vacuole, by enlarging and furrowing, cuts
out masses of cytoplasm, each with its respective nucleus.
Beside each nucleus a body appears, the coenocentrum,
which he thinks aids in the rounding up or “balling” of the
cytoplasm. This body acts as a dynamic center which draws
the cytoplasm to it and rounds it up into a spheroid mass.
Miss Carlson and Claussen observed a body which they said
might be termed a coenocentrum but observed it only a few
times. In Brevilegnia, on the contrary, with the contrac¬
tion of the cytoplasm and the disappearance of the central
vacuole, there is no indication of a coenocentrum (fig. 12) .
The loss of water from the oogone causes the rounding up
of the cytoplasm.
The nuclei that have recently undergone division are
fairly evenly distributed throughout the cytoplasm, but
there is a tendency for them to arrange themselves at the
316 Wisconsin Academy of Sciences , Arts , and Letters.
periphery of the egg (fig. 12). All the nuclei except one
shortly begin to show signs of degeneration. The remain¬
ing nucleus enlarges slightly and is now the female gamete-
nucleus. The nucleus becomes slightly flattened and the
chromatin-linin network appears to radiate from the nu-
cleole to the nuclear membrane where the chromatin ap¬
pears to be more aggregated. The vacuoles in the egg are
fairly uniform in shape, those near the center being some¬
what larger than those at the periphery. The degenerated
nuclei persist until after fertilization. The mature egg is
found generally at one side of the oogone just prior to fer¬
tilization.
Behavior of the Antherids
During the formation of the egg in the oogone* antherids
have been approaching the oogone from the same hyphae
or from neighboring hyphae. Antherids have been ob¬
served to attach themselves to the oogones after the forma¬
tion of the egg, but more frequently they are present at the
stage of egg formation. The development of the male
gametes within the antherid was not observed in Brevileg-
nia, but on the basis of the observation of other workers it
is thought comparable with the changes taking place within
the oogone during the formation of the egg. The antherid
upon coming in contact with the oogone may indent the
oogone. The rupture of the wall and discharge of its
contents into the oogone was not observed. The pres¬
ence of a fertilization tube has not been observed in
this form. The antherid which adheres to the wall of
the oogone at the spot where the egg touches the wall ap¬
pears to be the functional antherid. There is evidently a
dissolution of the wall of the oogone at the point of contact
of the antherid and the egg. Apparently one of the gamete
nuclei migrates into the egg. Several nuclei are often
present within the antherid and the terminal nucleus ap¬
pears slightly larger than the others and is probably the
nucleus that functions as the male gamete-nucleus (fig.
15) . Upon the discharge of the male gamete-nucleus into
the egg, the other nuclei in the antherid show signs of de¬
generation.
After the formation of the egg, the wall of the oogone is
Cooper — Cytological Studies on Sporange Development. 317
quite irregular with several papillae Shortly before ferti¬
lization the oogone wall stretches slightly and the papillae
disappear. Previous workers have observed that the wall
of the oogone in a number of species varies in thickness.
The oogone wall, in section view, appears to be made up of
many circular thickened plates at the intersection of which
are thin-walled regions. In section view, this gave the ap¬
pearance of thickened areas separated by thin membranes.
It has been thought for a long time that the antherid
paused at a thin area or that the fertilization tube pene¬
trated the thinner areas only, due probably to some chemo-
tactic stimulus. Couch, in his studies with the water
molds, has observed that the fertilization tubes penetrated
the thickened areas as well as the thinner ones, and so be¬
lieves that there is no ground for the assumption that the
fertilization tubes penetrate the thinner areas only.
Fertilization
After the male gamete nucleus has been discharged into
the egg it migrates toward the female gamete nucleus.
The male gamete nucleus can be readily recognized within
the egg (fig. 17) . It is almost spherical and slightly smaller
than the female gamete nucleus. A nucleolus can be seen
within each of the nuclei. It was not uncommon to find
the female gamete nucleus at a distance from the center of
the egg. In many preparations, while it may not be char¬
acteristic for the species, the writer has observed that the
male gamete nucleus was invariably discharged into the
egg (fig. 17) at the side farthest away from the female
gamete nucleus.
Several stages have been observed of the male gamete
nucleus on its way to the female gamete nucleus. The male
nuclus leaves in its wake a path of dense cytoplasm, which
Couch calls the “penetration path” (figs. 17, 18, 19).
Couch describes such a path in Leptolegnia caudata de
Bary. The male gamete nucleus comes to lie next to the
female gamete nucleus, one slightly overlapping the other.
Upon the union of the gamete nuclei the egg becomes a zy¬
gote. A wall now appears about the zygote. The cytoplasm
becomes quite vacuolate. Several large vacuoles are found
318 Wisconsin Academy of Sciences , Arts, and Letters.
in the vicinity of the zygote nucleus, with many smaller
ones at the periphery. Shortly before fertilization a small
opaque body appears in the egg and after fertilization it en¬
larges rapidly (figs. 23, 24). Occasionally there may be
more than one. These bodies are easily recognized as oil
bodies because of their definite outline and because of their
staining reaction. Faint fibrillar lines are present over their
surfaces. An oil body may enlarge until in some instances
it is about two-thirds the size of the zygote.
The zygote evidently undergoes a period of rest before
germination. Germinating zygotes were not observed.
In several zygotes a multinucleate condition has been ob¬
served with resting nuclei (fig. 25), but a most careful
study of a very large number of preparations has given no
evidence as to when such division begins. No one has as
yet given a complete story of the processes which intervene
between fertilization and the germination of the zygote in
any of the Saprolegniaceae.
Summary
In Brevilegnia the development of the sporanges agrees
with other members of the family.
The sporangiospores are multinucleate, a fact which has
been observed in only one other genus of the Saprolegnia¬
ceae.
The method of spore liberation is by the dissolution of
the sporange wall and not by the rupture of the wall, by
the contraction and rupture of the wall by the enlarging
zoospores, nor by the assistance of an intersporal substance
as has been noted by earlier observers.
The sporangiospores are non-ciliate at all times. They
float about for a time, settle into substrate and germinate
directly into hyphae.
Prior to the liberation of the sporangiospores from the
sporange the spores appear to encyst as there is a marked
contraction of the cytoplasm from the wall of the spore.
A single division of the nuclei takes place in the young
oogone which is in agreement with the results of recent
workers in other members of the family.
Cooper — Cytological Studies on Sporange Development . 319
The oogone contains a single egg only, which is found
generally at one side of the oogone.
The antherids approach the oogone during the formation
of the egg. After attaching themselves to the oogone, the
antherid nearest the egg is the functional antherid. The
antherid possesses no fertilization tube. The separating
walls dissolve and the male gamete nucleus passes directly
from the antherid into the egg and migrates to the female
gamete nucleus leaving in its wake a penetration path.
The gamete nuclei fuse and the egg becomes the zygote.
No nuclear divisions were observed in the zygotes, but
several zygotes were observed in a multinucleate stage.
The writer wishes to express his appreciation to Prof.
E. M. Gilbert for his advice and encouragement during this
investigation.
Explanation of Plates
All drawings were made with the aid of a camera lucida at table
level with a Leitz No. 4 ocular and a 1/16 oil immersion, making a
magnification of 1750.
Plate 3
Fig. 1. Tip of young multinucleate sporange with the first ap¬
pearance of the central vacuole.
Fig. 2. Sporange with the spores in a row. Aplanospores being
cut out by the central vacuole and by the furrowing from the side oi
the sporange. Aplanospore initials present.
Figs. 3, 4. Cross section of the club-shaped type of sporange with
the aplanospores arranged about the central vacuole. Cytoplasm
has contracted slightly, evidently an encysted condition.
Fig. 5. Longitudinal section of the club-shaped type of sporange
showing the position of the central vacuole.
Fig. 6. Aplanospores of the sporange bearing the spores in a
single row, showing the irregular manner of the disintegration of the
sporange wall; the spores remaining clustered together.
Fig. 7. Club-shaped type of sporange at maturity, anterior por¬
tion of sporange disintegrated and the aplanospores about to be lib¬
erated.
Plate 4
Figs. 8, 9. Longitudinal sections of an oogone prior to its assum¬
ing typical spherical shape. Nuclei in various stages of division;
some nuclei in the resting stage.
320 Wisconsin Academy of Sciences, Arts, and Letters.
Figs. 10, 11. Longitudinal sections of young oogones which have
rounded up with the nuclei in various stages of division.
Fig. 12. Multinucleate egg after the division of the nuclei. The
cytoplasm contracted, the vacuole has disappeared.
Figs. 13, 14. Uninucleate eggs with the supernumerary nuclei
disintegrating.
Fig. 15. Antherids from adjacent hyphae attached to oogone.
Fig. 16. Anther id from same hypha in contact with oogone wall
opposite egg, showing position assumed at time of fertilization.
Plate 5
Fig. 17. Egg with the male gamete nucleus approaching the
female gamete nucleus, leaving in its wake a penetration path.
Fig. 18. Egg with the gamete nuclei lying in contact. The pene¬
tration path is still in evidence. Antherid attached to oogone wall.
Fig. 19. Egg with gamete nuclei about to fuse. Penetration path
still present.
Figs. 20, 21, 22. Zygotes with the gamete nuclei about to fuse.
Fig. 23. An abnormal zygote in which the gamete nuclei have not
fused. The presence of the oil bodies indicates that some time has
elapsed since the male gamete nucleus entered the egg.
Fig. 24. Fully matured zygote, with fusion nucleus and well de¬
veloped oil bodies.
Fig. 25. Zygote in a multinucleate stage prior to germination.
Large oil body present.
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3. Claussen, P. 1908. Uber Eientwicklung und Befructung bei
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4. Coker, W. C. 1923. The Saprolegniaceae, with notes on other
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11. - 1916. Observations sur le chondrioms des Saprolegnia, sa
nature, son origine, et ses proprietes. Bull. Soc. Myc. France
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12. Davis, B. M. 1903. Oogenesis in Saprolegnia. Bot. Gaz. 35:
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16. - 1884. Vergleichende Morphologie und Biologie d. Pilze.
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17. Guilliermond, A. 1920. Observations cytologiques sur la cyto-
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legniacee. Compt. rend. Acad, des Sci. Paris 170: 1320. 5
figs.
19. — - 1920b. Nouvelles observations cytologiques sur Saprolegnia.
Comptes Rendus Acad. Sci. Paris 171: 266. figs. 1-6.
20. Hartog, M. M. 1887. On the formation and liberation of the
zoospores of the Saprolegniaceae. Quart. Jour. Micr. Sci.,
N. S. 27: 427.
21. - 1889. Recherches sur la structure des Saprolegniees. Comp¬
tes Rendus, Paris 108: 687.
22. - — 1892. Some problems of reproduction. Quart. Jour. Micr.
Sci. 33: 1. (1891.)
23. - 1895. On the cytology of the vegetative and reproductive
organs of the Saprolegnieae. Trans. Roy. Irish Acad. 30 :
649.
24. - 1896. The cytology of Saprolegnia. Ann. Bot. 10: 98.
25. - 1899. The alleged fertilization in the Saprolegniese. Ann.
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26. Harvey, J. V. 1927. Brevilegnia diclina N. Sp. Jour. Elisha
Mitchell Sci. Soc. 42; 243-246.
27. Hildebrand, F. 1867-68. Mycologische Beitrage. I. Uber eini-
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16.
28. Humphrey, J. E. 1892 (1893). The Saprolegniaceae of the
United States, with notes on other species. Trans. Amer.
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29. Kasanowsky, V. 1911. Aphanomyces laevis de Bary. I. Ent-
wickling der Sexualorgane und Befruchting. Ber. d. Deutsch.
Bot. Gesell. 29: 210. PI. 10.
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30. Meyer, Arthur. 1904. Orientirende Untersuchungen fiber Ver-
breitung, Morphologie und Chemie des Volutins. Bot. Zeit.
62; 113. FI. 15.
31. Mottier, D. 1904. Fecundation in plants 6-7.
32. Miicke, M. 1908. Zur Kenntnis d. Eientwickling und Befrucht-
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sell. 26a: 367. PI. 6.
33. Pringsheim, N. 1855. Neber dei Befruchting d. Algen. Monats-
ber. d. K. P. Akad. d. Wissensch. zu Berlin, p. 133. Also in
Ann. Sci. Nat. Ser. 4, 3; 363. 1855 (Only a paragraph on A.
prolifera on p. 378.)
34. - 1858. Beitrage zur Morphologie und Systematik d. Algen,
II. Die Saprolegnieen. Jahrb. f. wiss. Bot. 1: 284. Pis. 19-
21. Also in Ann. Sci. Nat. Ser. 4, 11: 349. Pis. 6-7. 1859.
35. - 1860. Beitrage zur Morphologie und Systematik d. Algen,
IV. Nachtrage zur Morphologie d. Saprolegnieen. Jahrb. f.
wiss. Bot. 2: 205. Pis. 22-25.
36. - 1883. Ueber die vermeintlichen Amoeben in den Schlauchen
und Oogonium d. Saprolegnieen. Bot. Centralbl. 14: 378.
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ungsact von Achlya. Jahrb. f. wiss. Bot. 14: 111.
38. Rothert, W. 1888. Die Entwicklung d. Sporangien bei den Sa¬
prolegnieen. Cohn’s Beitrage zur Biologie d. Pflanzen 5: 291.
PI. 10; 1892 (1890). Distributed in 1888. Also published in
Polish in Sitzungsb. (Rospr.) Krakau. Akad. Math. Naturw.
Classe 17: 1. 1887.
39. Rudolph, K. 1912. Chondriosomen und Chromatophoren. Ber.
d. Deutsch. Bot. Gesell. 30: 605. PI. 18, 1 fig.
40. Schmitz, F. 1879. Untersuchungen fiber die Zellkerne d. Thal-
lophyten. Verh. des nat. Vereins d. Preuss. Rheinl. und West-
falens 36: 345.
41. Smith, F. E. V. 1923. On direct nuclear divisions in the vege*
tative mycelium of Saprolegnia. Ann. Bot. 37 : 63.
42. Trow, A. H. 1895. The karyology of Saprolegnia. Ann. Bot.
9: 609. Pis. 24, 25.
43. - 1899. Observations on the biology and cytology of a new
variety of Achlya Americana. Ann. Bot. 13: 131. Pis. 8-10.
44. - 1904. On fertilization in the Saprolegnieae. Ann. Bot. 18:
541. Pis. 34-36.
45. - 1905. Fertilization in the Saprolegniales. Bot. Gaz. 39:
300.
46. Weston, W. H. 1918. The development of Thraustotheca, a pe¬
culiar water-mold. Ann. Bot. 32: 155.
47. Zopf, W. 1882. Ueber Parasiten in den Antheridien, Oogonien
und Oosporen von Saprolegnieen. Bot. Centralblatt 12: 356.
43. - 1883. Erwiderung. Bot. Centralblatt 15: 156.
^77
TRANS. WIS. ACAD. - VOL. 24
PLATE 3
4
TRANS. WIS. ACAD. - VOL. 24
PLATE 4
15
TRANS. WIS. ACAD. — VOL. 24
PLATE 5
THE CYTOLOGY AND MORPHOLOGY OF SORDARIA
FIMICOLA CES. AND DE NOT
Addie Emma Piehl
INTRODUCTION
Recent cultural experiments dealing with the homothal-
lism and heterothallism of fungi relating to the Sordarias
has made it worth while to study the origin and nature of
the structures initiating the formation of fruiting bodies
in these forms.
Special attention has therefore been given to the growing
of Sordarias in pure cultures so that the presence or ab¬
sence of sex organs could be noted, as well as the nature
and function of such organs during their development.
Gross morphology was noted, but the nuclear phenomena,
especially as to the behavior of the nuclei during the early
stages in formation of the perithecium, has been given spe¬
cial consideration. As a preliminary to this study, a care¬
ful review has been made of the accumulated data, both cy-
toiogical and morphological, of previous workers on the As-
comycetes.
Historical Review
Pyronema confluens
The development and nature of the sex organs in Pyro¬
nema confluens have interested several investigators. The
first descriptions were supplied by De Bary (1863), by the
Tulasne brothers (1865, 1866), by Van Tieghem (1884),
and by Kihlman (1885). These early investigators ob¬
served the elongated antheridium, the globular oogonium,
the trichogyne, and the ascogenous hyphae characteristic
of Pyronema. More detailed work of a cytologicai nature
was necessary, however, before the history of Pyronema
confluens could be considered complete. This led to the in¬
vestigations of Harper (1900), followed by Dangeard
(1903, 1907), and by Claussen (1912).
324 Wisconsin Academy of Sciences, Arts, and Letters.
As a result of Harper's observations on Pyronema con -
fluens, there has been obtained a detailed life history, a re¬
view of which follows. The vegetative mycelium is multi-
nuclear, from six to twelve nuclei appearing in each cell.
Although the mycelium is sparse and loose, the reproduc¬
tive organs are abundant and produce ascocarps which,
when mature, are seen crowded together.
The first appearance of sex organs may be noted when
certain thick hyphae tend to stand at right angles to the
substratum. These hyphae, from the same or different
mycelia, may be seen grouped in pairs. One of the two
branches becomes swollen and differentiates into an upper
spherical cell, the oogonium, and a lower portion, the stalk,
composed of a few cells; the other branch has an upper
cell cut off which remains slender and is called the anthe-
ridium, with a lower portion forming the stalk cells. Very
early in the formation of these sex organs, the trichogyne
appears as a slight elevation at the “apex” of the oogonium.
This trichogyne elongates, becomes multinucleate, and is
separated from the oogonium by a wall.
The mature oogonium is a spherical or flask-shaped cell
filled with dense cytoplasm and containing many nuclei
larger than those of the vegetative cells. The nuclei of the
antheridium are almost as large as those of the oogonium
but the cytoplasm is less dense, due perhaps to the lack of
reserve materials.
Before fertilization, hyphae from the ascogonial and
stalk cells and from surrounding cells begin to grow up
and later envelop the sex organs.
The trichogyne and the antheridium grow towards each
other, the tip of the trichogyne finally coming in contact
with the apex or side of the male organ. When fertiliza¬
tion is about to take place, there is a single receptive spot
at the end of the trichogyne free from nuclei and made up
of dense, finely granular cytoplasm. At the point of con¬
tact of the antheridium and trichogyne, a pore is formed by
dissolution of the cell walls. During this process, the nu¬
clei of the trichogyne degenerate and when they are com¬
pletely disorganized, the nuclei of the antheridium migrate
through the pore and into the trichogyne, in which the con¬
tents have degenerated still further so that the cytoplasm
Piehl — Cytology and Morphology of Sordaria Fimicola Ces. 325
and nuclei together form a densely staining mass. The
male nuclei continue to pass into the trichogyne which
often appears filled and slightly swollen as a result. The
wall at the base of the trichogyne now breaks down. This
allows an opening for the male nuclei to enter the oogoni¬
um.
The greater part of the cytoplasm remains in the antheri-
dium and trichogyne and does not enter the oogonium. Af¬
ter migration of the male nuclei through the trichogyne
and into the oogonium, a new wall appears at the base of
the trichogyne so that the oogonium is again a single cell.
The male nuclei pair with the female nuclei and these
paired nuclei then fuse within the oogonium. During this
time slight protuberances make their appearance in the
walls of the oogonium which is called an ascogonium after
fertilization takes place. These projections elongate and
are young ascogenous hyphae into which the fusion nuclei
soon pass. These hyphae lengthen in a crooked fashion,
become septate, and grow among the vegetative branches
which have arisen from the stalk cells of the oogonium.
The vegetative branches grow faster than the ascogenous
hyphae and continue to grow upward, their extremities
becoming paraphyses. The ascogenous hyphae also grow
in a vertical direction and appear among the paraphyses,
but their upward growth soon stops and they then grow
horizontally, branching repeatedly, forming a network at
the base of the hymenium.
The tips of the ascogenous hyphae which extend upward
among the paraphyses, become curved. They contain, at
first, two nuclei which divide simultaneously so that four
nuclei are found in the curved portion, and these nuclei are
arranged in such a way that the two nuclei from separate
spindles lie together in the “crook” of the branch, one nu¬
cleus in the tip and one below the curve. Cell walls then
appear, cutting off the tip with one nucleus, the bent upper
part of the branch with two nuclei. It is from this binu-
cleate cell that the young ascus arises as an apical projec¬
tion into which the two nuclei pass and then fuse. This
fusion-nucleus or primary nucleus, as it is usually called,
divides to produce two, then four, and finally eight nuclei
326 Wisconsin Academy of Sciences, Arts, and Letters.
arranged throughout the entire length of the now fully
elongated ascus.
Each nucleus has a beak and to this are attached the as¬
ter rays which still remain. These fibers grow back
around the nucleus until they meet and a membrane is
formed around each nucleus. Thus the ascospores are cut
out by a process called a 'Tree cell-formation.” Later a
wall forms around each uninucleate spore.
Harper, therefore, finds that the nuclei in the antherid-
ium pass through the trichogyne and into the oogonium
where they pair with the female nuclei and fuse. These
fusion-nuclei then pass into the ascogenous hyphae. Har¬
per thus finds two fusions in Pyronema confluens, one in
the oogonium and one in the ascus when the two nuclei
from the penultimate cell unite to form the primary nu¬
cleus of the young ascus. Then chromosomes were counted
by Harper in the divisions of the nuclei in the ascogenous
hyphae and in the divisions of the primary nucleus in the
ascus but whether this number represents a double number
resulting from the fusion in the oogonium or whether re¬
duction division has already taken place, Harper was un¬
able to decide.
Dangeard (1903-1904) disagrees with these conclusions
given by Harper. The sex organs, he believes, are not
functional. The wall between the oogonium and the tri¬
chogyne does not disappear so that the migration of male
nuclei beyond the trichogyne can not take place. Asco¬
genous hyphae arise from the functionless sex organs and
the asci possess characteristics of a sex organ in which the
nuclei fuse as gametes. This fusion-nucleus then divides
by heterotypic division followed by two homoeotypic divi¬
sions. There is, therefore, according to Dangeard, only
one nuclear fusion and that in the young ascus.
The theory of a single nuclear fusion is also advanced by
Claussen (1912). His conclusions, concerning the develop¬
ment and function of sex organs and the migration of male
nuclei through the trichogyne and into the oogonium, agree
with those of Harper. But, according to Claussen’s inves¬
tigations, the male nuclei pair with the female nuclei but
there is no fusion in the oogonium. These paired nuclei
pass into the ascogenous hyphae and then fuse in the young
Piehl — Cytology and Morphology of Sordaria Fimicola Ces. 327
ascus as described by Harper and Dangeard. A single di¬
vision then follows with two homoeotypic divisions in the
ascus. Claussen has counted twelve chromosomes in these
divisions.
Brown (1915) finds no union of antheridium with tri-
chogyne, no fusion of nuclei in the ascogonium and none in
the ascogenous hyphae. He agrees with Claussen and Dan¬
geard in the single fusion thory, that fusion taking place
in the young ascus.
Pkyllactinia corylea
In Phyllactinia corylea , Harper (1905) finds a small coil
composed of the functional oogonium and antheridium
which appear at about the same time as lateral branches
from separate hyphae but from the same mycelium. The
oogonium is cut off from its hypha by a cross wall so that
a large upper cell and a small lower cell results. The upper
cell enlarges to form the oogonium and contains a single
nucleus, the lower cell divides to form the stalk cells of the
oogonium. The antheridium is formed in a similar man¬
ner but remains slender and erect and contains one nucleus.
The oogonium grows more rapidly and twists around the
short, erect antheridium. The walls separating the two
sex organs, at the point of contact, break down and a pore
is thereby produced for the passage of the male nucleus
into the oogonium. A part or all of the cytoplasm remains
in the antheridium. Fusion of the two nuclei then occurs
in the oogonium. The pore between the two sex organs
closes and the antheridial cell degenerates. While fertili¬
zation is going on, protective branches grow up from the
stalk cells of both sexual branches and begin the formation
of the perithecial envelope.
The oogonium, after fertilization, is called the ascogonium.
It enlarges and a division of the fusion-nucleus takes place.
This binucleate cell remains in this condiiton until the as¬
cogonium is completely enclosed by the enveloping hyphae.
Then the two nuclei divide and just how many divisions
occur before cell walls appear, Harper was unable to ob¬
serve. However, from three to five cells in a row appear
in the ascogonium, the end cell with one nucleus, the others
328 Wisconsin Academy of Sciences , Arts , and Letters.
with one or two nuclei. Ascogenous hyphae arise as lat¬
eral branches from the ascogonium cells and whether they
come from one cell or from several of the upper cells is un¬
known. It is certain that some arise from the penultimate
cell. The ascogenous hyphae develop considerably before
they become septate; then cells with two nuclei and with
one nucleus result. From the binucleate cells, the asci are
formed as lateral outgrowths probably in the same manner
as in Pyronema.
Each young ascus has two nuclei. The ascogenous cells
which formed the asci contained two nuclei, but these nu¬
clei are not necessarily daughter nuclei of the same nucleus.
As the asci elongate, the two nuclei in each ascus fuse,
forming the primary ascus-nucleus which moves to the
lower end of the ascus. By this time the asci have become
long, narrow structures with short stalks at the base. The
primary nucleus undergoes three nuclear divisions so that
eight nuclei are formed. By free cell-formation, eight
spores are cut out.
The perithecium at this time has a three-layered wall;
the innermost layer several cells in thickness, thin-walled,
and probably supplying food for the developing asci. Out¬
side of this layer is a strengthening and protecting zone of
cells with walls which appear lignified. The outermost
layer of the perithecium consists of thin-walled cells from
which characteristic appendages arise as stiff, pointed hairs
with swollen bases.
Ascobolus magnificus
Dodge (1920) found that cultures containing two strains
of mycelia were necessary for the production of ascocarps.
In such cultures four to six days old, he found paired
branches. These club-shaped branches elongate. One of
them usually grows faster than the other, coils about this
shorter branch, and is called the ascogonium; the shorter
structure remains a two or three-celled antheridium. Very
often both branches arise a short distance apart. In this
case, one remains an erect, short antheridium, the other
elongates into an ascogonium and forms a trichogyne which
grows to and becomes coiled about the antheridium.
PieJil — Cytology and Morphology of Sordaria Fimicola Ces. 329
Dodge was able to prove the presence of an antheridium in
any normal case by separating this same structure from
the trichogyne coil before fusion had occurred and if hyphal
branches had not grown out from the stalk of the oogonium
to envelop the male organ.
In normal development, the ascogonium enlarges and of¬
ten ascogenous hyphae grow out before sterile hyphae be¬
gin to form the fruiting body.
Since two strains of mycelia are necessary for the pro¬
duction of fruiting bodies, there is good indication that sex
has not been lost or reduced in at least this species of Asco-
bolus.
By germinating ascospores of Ascobolus magnificus,
Dodge obtained papulospores — defined by him as a spore
“in which one or two large storage cells are surrounded by
a covering of hyphae which develop from blister-like out¬
growths of the storage cell.” He transferred mycelia of
the single papulospore strains to different kinds of media
but obtained no ascospores of Ascobolus. Hotson, at
Dodge’s request, studied papulospores from cultures sent by
the latter and concluded that they were not the asexual
stage of Ascobolus magnificus. He was unable to obtain
ascospores from the cultures.
E. S. Schultz also made single spore cultures of this
papulaspora for Dodge, but was unable to obtain ascocarps.
Recently Dodge proved that papulospora is not a
case of parasitism on Ascobolus as he once thought, but is
an example of what has been described as self -penetration
or self-parasitism. Internal hyphae running in and out of
larger hyphae were found with papulospores arising from
branches growing out of these internal hyphae.
As already noted, ascocarps could not be obtained from
single spore cultures of papulospores and there is no record
of producing ascocarps from single ascospore cultures of A.
magnificus. Two strains are thus necessary for sexual re¬
production and the formation of ascocarps. Dodge planted
single ascospore cultures in separate petri dishes and ob¬
tained mycelium and papulospores but no sex organ
branches or ascocarps. Of the seven strains obtained he
planted five of them as follows:
22
330 Wisconsin Academy of Sciences , Arts , and Letters.
Strains - Results after 10 days
2 alone - Papulospores but no fruiting bodies
2 and 1 - - - Ascocarps
2 and 3 - Papulospores but no fruiting bodies
2 and 4 - Ascocarps
2 and 5 __ - Papulospores only
6 lost
7 not used
The results prove that strains 2, 3, and 5 when grown in
the combinations given in the table are sterile ; the same is
true for strains 1, 4, and 7. Fertile cultures, however, are
produced when any one of the first group is grown with
the different strains in the second group.
Strains 2 and 4 were further tested. It was found that
each is sterile when grown alone, but fertile when com¬
bined with the other. Strains 2 and 1, although sterile
when grown alone, produced ascocarps when grown to¬
gether. But strains 4 and 4 grown together in one petri
dish will show a zone between them free from hyphae.
When opposite strains 2 and 4 are planted, there is no such
zone between them and sex organs appear throughout the
culture. Therefore, according to Dodge, each strain is
self-sterile and in a single strain culture produces no sex
organs. Sexual reproduction occurs only in cultures con¬
taining two strains properly chosen.
Venturia inaequalis
Frey (1924) has reported the presence of an ascogonium,
a trichogyne, and antheridial branches which indicate sex¬
ual conditions in Venturia inaequalis. The coil usually
consists of two branches which arise from the same fila¬
ment. One branch functions as an ascogonium, the other
is not an antheridium but may function as a nutritive or¬
gan. Sometimes a single branch may be found producing
a coil. The coil increases in size, stains deeply, and sends
out an elongated projection called the trichogyne. Early
stages of the trichogyne and ascogonium are non-septate —
the former seldom has a nucleus, while the latter is multi-
nucleate. From the filaments near the ascogonium, the
antheridial hyphae arise. Their apical cells are enlarged
and some appear multinucleate, probably functioning as
PieJil — Cytology and Morphology of Sordaria Fimicola Ces. 331
antherids because they come in contact with the trichogyne
and seem to fuse with it. Just when, in the development
of the coil, fertilization occurs, is not known, but since later
stages show paired nuclei in cells of the ascogonium, it
probably takes place during the non-septate stage.
After fertilization, the ascogonium becomes septate and
contains seven or eight cells with paired nuclei, as already
mentioned. These nuclei do not fuse in the ascogonium.
Later there is an increase in number of cells in the ascogo¬
nium, some of which may contain as many as four nuclei.
These cells branch extensively, are septate, and certain of
these cells may become asci directly, other cells may again
branch, and these branches then become septate and form
asci. Frey does not consider these branches equivalent to
the ascogenous hyphae described by Harper and Claussen.
The ascogenous cells contain two or four nuclei which
enlarge and pair and these paired nuclei then fuse in the
young ascus. The nuclei resulting from the division of this
primary nucleus lie near the center of the ascus, but after
the second division when the ascus has elongated, the nuclei
pass to the periphery. After the third division the spores
are cut out by the astral fibers as described by Harper.
These spores are two-celled and uninucleate.
Killian (1915) reported a large trichogyne, a coiled asco¬
gonium, and also a branched antheridium in Venturia ina-
equalis. Frey found a few lobes of the apical cells of an-
theridial branches and perhaps these lobes correspond to
the branched antherids found by Killian. Killian suggests
the possibility of a septate archicarp at the time of fusion
of antherid and trichogyne because of pores found in cells
of the ascogonium.
Polystigma ruhrum
In Polystigma ruhrum, Nienburg (1914) found a coil
composed of an ascogonium, a trichogyne, and an antheri¬
dium. The ascogonium is an elongated cell with one nu¬
cleus ; the trichogyne is multinucleate, does not function as
a sex organ but perhaps has a nutritive value ; the antheri¬
dium is an elongated cell with many nuclei. The antherid¬
ium applies itself to the ascogonium and the walls at the
point of contact between the two cells break down. The
332 Wisconsin Academy of Sciences, Arts, and Letters .
male nuclei then pass from the antheridium into the asco-
gonium where one male nucleus enlarges to become the
functional male nucleus — the remaining nuclei degenerate.
No fusion of nuclei in the ascogonium was observed by
Nienburg. From the ascogonium, ascogenous hyphae, the
cells of which contain two nuclei, arise and whether this bi-
nucleate condition is due to a “constriction” division is an
unsettled question. Development of the ascus could not be
followed.
Nienburg did not observe a fusion of nuclei in the asco¬
gonium or in the ascogenous hyphae and believes there is
but a single fusion and that in the ascus.
Blackman and Welsford (1912) disagree with Nienburg.
They claim that the ascogenous hyphae develop from vege¬
tative cells, and that two nuclear fusions occur, one in the
ascogenous hyphae and one in the ascus.
Podospora anserina
An organism more closely related to Sordaria than the
forms described above, is Podospora anserina, a saprophyte
commonly found on waste material. It was investigated
for the purpose of determining spore formation because
the number of spores produced in a few species of Podos¬
pora varies from four to sixteen or more.
Wolf (1912) made a study of Podospora anserina and
found coils made up of two hyphae, but with no differentia¬
tion of sex organs. The nuclei are very small and for this
reason, fusion of nuclei was not observed. Vegetative hy¬
phae arise from the coil, envelop the coiled hyphae and de¬
velop into the pear-shaped perithecium.
Near the base of the perithecium, the asci develop in
groups, several asci branching from a single hypha. The
young ascus contains one large primary nucleus, much lar¬
ger than those found in the vegetative mycelium where
each cell is multinucleate. This primary nucleus under¬
goes divisions; only a few stages, however, were observed
by Wolf. After heterotypic and homoeotypic divisions
there is a short period of rest followed by an enlargement
of the nuclei containing knots of chromatin which may rep¬
resent chromosomes. After the third division a region of
Piehl — Cytology and Morphology of Sordaria Fimicola Ces. 333
less dense protoplasm appears between pairs of the result¬
ing eight nuclei. It was found that four spores are cut out
of the ascus in Podospora anserina , but whether only one
nucleus of each pair takes part in the process of spore de¬
limitation or astral rays from the two concerned are effec¬
tive was not determined. Since two nuclei are normally
included within each of the four spores, no disintegration
of the eight nuclei in the epiplasm takes place.
As the spores mature they increase in size, the nuclei mi¬
grate to the ends, the upper part enlarges and the lower
half becomes the hyaline appendage. The nuclei increase
in size and the colorless, young spores gradually change to
green, then to dark brown. The appendage remains hya¬
line, becomes equal in length to the body of the spores,
and often disappears in mature spores.
Sordaria fimicola Ces. and De Not
Sordaria fimicola Ces. and De Not. was first described by
Roberge in 1849 and given the name of Sphaeria fimicola
later in 1865. Cesalpini and De Notaris described the fun¬
gus more fully and gave it the name Sordaria fimicola ,
which name has been generally accepted. More recently,
Griffiths and Seaver, in listing the Sordariaceae for the
North American Flora, dropped the genus Sordaria as such
and substituted Fimetaria and gave the new combination
Fimetaria fimicola (Roberge) Griffiths and Seaver. As
Sordaria fimicola has been so long accepted and is found as
such in most literature, that name will be used in this
paper.
Materials and Methods
Pure cultures of Sordaria fimicola Ces and De not. avail¬
able for study, were easily isolated from refuse material
and from impure cultures of Sordaria, and grown at room
temperature on artificial media. It develops rapidly on
potato agar, forming a great number of black, flask-shaped
perithecia which stand upright above the medium. Spores
from the most characteristic perithecia were then chosen
and transferred to agar plates. Single spore cultures were
then made from these cultures and pure strains of Sordaria
334 Wisconsin Academy of Sciences , Arts , and Letters.
fimicoila were thus obtained. At different intervals, pieces
of agar bearing mycelium and perithecia in various stages
of development were cut out of the culture plates, fixed in
Flemming’s weak solution, imbedded in paraffin, and then
cut into sections varying from 5 to 20 microns in thickness.
A variety of stains were tried: Flemming’s triple, Dela-
field’s haematoxylin, Heidenhain’s iron alum haematoxylin,
and differential stains with the latter, such as licht gruen,
fuchsin, iodine green, orange-G, erythrosin and Congo red.
The best results were obtained with the 2 per cent iron
alum haematoxylin and without a counterstain.
Observations and Discussion
The cytoplasm in the mycelium of Sordaria fimicola is
highly vacuolate with strands of varying thickness extend¬
ing throughout the cells. Close to the walls can be seen
a thin, densely granular layer of cytoplasm. Imbedded in
the cytoplasm are numerous mitochondria, either scattered
singly, in aggregations, or often in pairs, probably as the
result of recent division. The cells are multinucleate.
With the exception of the nucleole which stains heavily, the
content of the nucleus is not readily distinguishable. Cross
walls with special thickenings as shown in figures 1, 4 and
5, are frequently present. A similar condition has been
reported in Pyronema by Harper, and in various Basidio-
mycetes by several workers. These thickenings appear as
darkly stained granules or as a mass of material centrally
located on either side of the cross wall.
In several instances, branches arising at right angles to
the mycelium and apparently from separate hyphae, grow
toward each other and come in contact (figures 1-6) . This
is not a condition of ordinary anastomosing, but rather one
of actual fusing of cell contents from the two closely ap¬
plied hyphae as the result of a dissolution of the separating
walls. This may occur at the tips or at various places in
the lateral walls of the two branches (figures 4, 5, and 6).
There is no differentiation of cytoplasmic content in the two
branches and the nuclei in each cell are variable, but in sev¬
eral cases a difference in size can be noted in the two hy¬
phae and in the mycelium from which the hyphae arise
PieJil — Cytology and Morphology of Sordaria Fimicola Ces. 335
(figures 2, 3, 6) . It is probable that the fusion of cell con¬
tents observed may be an initial step in coil formation, but
intermediate stages, which might give conclusive results,
are lacking. Miss Miller (1927), a research student in bot¬
any, made many single spore cultures of Sordaria and in
every instance secured normal perithecia containing nor¬
mal spores.
Repeated experiments with single spore cultures and
with mycelium from single spore cultures gave the same re¬
sults. She then made plantings of two spores and with
portions of mycelium from different cultures to see what
the effect would be, and in more than one hundred cultures
of this type the perithecia appeared throughout the cultures
indicating that all the strains with which she worked were
homothallic.
In the majority of cases a typical coil arises where two
hyphae come in contact (figures 7-20) . In plate cultures
four or five days old, upright branches which arise at right
angles to the mycelium make their appearance. These
branches originate from separate hyphae, grow toward
each other and come in contact. One of them usually
swells considerably, grows faster than the other, and ini¬
tiates the formation of the coil. The two branches elongate
and continue to coil about each other until a large coil is
produced. Dodge (1920) finds that the coil in Ascobolus
magniftcus is differentiated into a definite male organ, the
antheridium, which does not elongate but remains a short
erect branch, and into a female organ, the oogonium which
elongates, the end cell functioning as a trichogyne and coil¬
ing about the antheridium. Harper, in Pyronema, finds an
elongated antheridium, a large, spherical oogonium with a
trichogyne produced at the “apex” ; the trichogyne and an¬
theridium bring about fertilization. But in Sordaria fimi¬
cola no true oogonium, trichogyne or antheridium are dis¬
tinguishable. Sordaria resembles Podospora anserina in
this respect because both have little differentiation of sex
organs. Due to the swelling of one branch in the initiation
of the coil, there is a difference in size of the two sex organs
as seen in figures 16-20, suggesting the presence of male
and female elements.
The sex organs composing the coil are, therefore, elong-
336 Wisconsin Academy of Sciences, Arts, and Letters.
ated coiled branches with cross walls cutting off cells con¬
taining from one to seven nuclei. After fusion of certain
of these cells an increased number of nuclei can be found
in each cell of the coil proper. These fusion cells, probably
representing simple or reduced sex organs, give rise to a
variable number of hyphae which seemingly function as
ascogenous hyphae in that the ultimate branches will give
rise to asei. Whether they are true ascogenous hyphae such
as are found in Pyronema and other forms or whether they
more nearly resemble the ascogenous structures found in
Venturia, it is impossible to state. The nature and ar¬
rangement of nuclei in these structures suggest that fusion
has taken place, but the actual process of fusion has not
been noted.
These hyphae branch repeatedly and as they develop they
become more and more like the ascogenous hyphae de¬
scribed by others. The curved end of such a branch often
contains four nuclei arranged in a row. Cross walls are
then laid down in such a manner that the end cell contains
a single nucleus, the second cell two nuclei, and the third
cell again contains only a single nucleus (figures 22-25).
The growth of the second (penultimate) cell causes the tip
cell to bend backwards. In many instances it was found
that all the cells except that at the tip contained two nuclei.
In figures 23 and 24, the terminal cell can be seen to curve
downward and the whole to give the appearance of the
typical “shepherd’s crook” commonly found at the ends of
ascogenous hyphae in many Ascomycetes.
Most of these hyphae remain in large part within the coil,
but occasionally (figures 20-25) the hyphae project beyond
the coil and are then more characteristic of true “ascoge¬
nous hyphae.”
Since the above was written, a paper by Arnold has ap¬
peared in which he gives the development of the perithe-
cium and ascogenous hyphae in Sporomia leporina Niessl.
The perithecium, which originates as a single enlarged cell
of the vegetative mycelium, becomes a large rounded struc¬
ture composed of several layers of cells, the innermost of
which break down so that a cavity remains in the center.
Extending into this cavity are the long hyphae which arise
from the apex of the perithecium. Their enlarged tips,
Piehl — Cytology and Morphology of Sordaria Fimicola Ces. 337
found near the base of the perithecium, give rise to asco-
genous hyphae which bend upwards, become hooked, and
finally cells are formed in the usual way as described in
other forms.
Faull (1905) finds asci in Sordaria fimicola arising from
the terminal cells as well as from penultimate cells of the
ascogenous hyphae. In my material, the asci usually arise
as lateral outgrowths from any of the cells which contain
two nuclei. The young asci are short, swollen structures
(figures 24 and 25) with dense cytoplasm and a large pri¬
mary nucleus, the result of a fusion of the two nuclei which
were present in the ascogenous cells which develop into
young asci. The asci elongate and extend in a vertical di¬
rection beyond the coil which is located in the basal region
of the now enlarging perithecium (figures 26, 27).
The primary nucleus lies at the center or above the cen¬
ter of the ascus. It is a well defined and easily distinguish¬
able structure in contrast with the small indistinct nuclei
of the vegetative cells. Within the nuclear membrane is
one large nucleole and a network of chromatin material.
This chromatin is not in strands as Harper (1905) finds in
Phyllactinia, where the resting nucleus contains chromatin
threads attached to a central body located against the nu¬
clear membrane. These strands of Phyllactinia extend
into the nucleus and correspond in number to the number of
chromosomes counted later. In Sordaria, the primary nu¬
cleus in the resting condition has the chromatin in the form
of a network characteristic of nuclei in higher plants (fig¬
ures 28-30). The division of the primary nucleus has not
been observed in any of the material studied, but results
of division of the primary nucleus have been found. As
division goes on, the asci elongate and finally eight nuclei
are formed. Here again only the nucleole is visible in each
nucleus (figures 31-34). Further elongation takes place
as the asci mature. They become long, slender structures
with the eight nuclei in a single row, extending throughout
the entire length of the ascus. Eight spores are then cut
out of the cytoplasm surrounding the nuclei, probably by
‘Tree cell-formation” as Harper observed in several Asco-
mycetes, although no intervening stages were observed
from the time eight nuclei were visible until eight uninucle-
338 Wisconsin Academy of Sciences, Arts, and Letters.
ate spores were produced. No more than eight nuclei have
been found in a single ascus but spores have been observed
in which there are two nuclei, a condition which probably
results from division of the nucleus within the spore. The
spores are round when first formed, but become ellipsoid,
acutely rounded at one end with a vacuolated cytoplasm.
When mature, they have a thick wall surrounded by a gela¬
tinous sheath (figure 35).
In addition to these main studies, attention has been
given to the initiation of the perithecium. Vegetative hy-
phae arise in the neighborhood of the coil and envelop this
structure. They begin to enclose the sex organs when the
coil is still quite immature and by the time the “ascogenous
hyphae” are formed, the vegetative hyphae have formed
several layers of the perithecium. Sections of perithecia
were observed in which a coil had been retarded in its de¬
velopment although the perithecial walls were fully formed
and young asci were present. In such cases, the initiation
of the perithecium probably takes place with the develop¬
ment of another coil from which the young asci originate.
The inner layers of the fruiting body remain thin-walled,
but towards the exterior the walls become impregnated
with waxy substances, are thicker, and take the stain very
readily.
At the time of ascus formation, hyphae resembling para-
physes can be seen in the interior of the perithecium and
among the young asci. Later these hyphae disintegrate
and when the asci are mature, no paraphyses or hyphae re¬
sembling them are present.
I wish to express my sincere appreciation of the helpful
advice and criticism given by Dr. E. M. Gilbert during the
progress of this work.
Summary
Sordaria fimicola has recently been proved a homothallic
species. In single spore cultures, perithecia appear in
large numbers and in plate cultures inoculated with two
strains, perithecia are abundant throughout the material,
not limited to the region where the two different mycelia
meet.
Piehl — Cytology and Morphology of Sordaria Fimicola Ces. 339
The sex organs are elongated, multicellular branches
which coil about each other, forming large coils. They are
undifferentiated except for a slight difference in size; no
true antheridium, oogonium, or trichogyne are distinguish¬
able.
“Ascogenous hyphae” arise from fusion-cells within the
coil. Each hypha curves at the tip, forming a “crook” in
which are found a uninucleate cell at the end, a binucleate
penultimate cell and others with one or two nuclei.
Asci arise as lateral outgrowths from the binucleate cells,
usually the penultimate, and the two nuclei fuse to form a
primary nucleus. This nucleus undergoes three divisions
and eight nuclei result which finally are cut out with a por¬
tion of the surrounding cytoplasm to form spores.
The uninucleate spores become binucleate as a result of
the division of the nucleus present at the time the spores
were formed.
As soon as the coil is well organized, hyphae cells in¬
crease in number and very soon completely enclose the coil
and by a continued growth give rise to the various types of
cells which characterize the perithecium of a Sordaria.
Explanation of Plates
Plate 6
All figures were drawn with an Abbe camera lucida, using a Leitz
4-ocular and a 1/16 oil immersion objective; magnification of 1850.
Photomicrographs were made with a Leitz 4-ocular, 1/16 oil im¬
mersion, magnification of 1300, with the exception of fig. 29, which
was made with a 6 mm. objective, magnification of 500.
Figs. 1, 2. Two hyphae in contact. Cross walls with special
thickenings are present, cells are multinucleate with only nucleoles
visible. Fig. 2 shows hyphae of different size.
Figs. 3-6. As in Figs. 1 and 2. Dissolution of walls at place of
contact and fusion of cell contents.
Figs. 7-20. Coil formation. No differentation of sex organs,
merely a difference in size of the two hyphae can be noted.
Plate 7
Fig. 21. Tips of ascogenous hyphae showing multinucleate con¬
dition.
Figs. 22-25. Septate ascogenous hyphae. Fig. 23 shows the bi¬
nucleate penultimate cell which usually gives rise to asci. Figs. 24,
25 show young asci developing as lateral outgrowths.
Figs. 26, 27. Elongating asci with a primary nucleus visible in
some.
340 Wisconsin Academy of Sciences , Arts, and Letters.
Figs. 28-30. Portions of asci showing large primary nucleus with
distinct nucleole and a network of chromatin material.
Fig. 31. Portions of ascus after first division of primary nucleus.
Fig. 32. Ascus with two nuclei.
Figs. 33, 34. Asci with 8 nuclei, only 6 visible.
Fig. 35. At left, a spore from an ascus containing 8 spores. At
right, a spore from an ascus with 4 spores.
Bibliography
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245. 1928.
Ashby, S. F., and Nowell, W. The fungi of Stigmatomycosis. Ani-
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Claussen, P. Zur Entwicklungsgeschichte der Ascomyceten Pyro-
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Dodge, B. 0. Life history of Ascoholus magnificus and the origin of
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1920.
- Nuclear phenomena associated with heterothallism and homo-
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cytology of the ascus. Ann. Bot. XXIII: 537. 1909.
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Flora III. 1. Dec. 1910.
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PLATE 6
TRANS. WIS. ACAD. - VOL. 24
PLATE 7
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SOME NOTES ON ALLOMYCES ARBUSCULA
BUTLER
Joseph Henry Lugg
This fungus is one of the water molds whose exact rela¬
tionships have been in doubt, and which, as yet, have not
been the subject of much serious investigation. Its sapro¬
phytic habit, general mycelial structure, and apparent life
history indicate its connection with the Saprolegniales — if
this group can be raised to the rank of an order as Coker
suggests (2). In this order he places four families : the
Saprolegniaceae, the Leptomitaceae, the Rlastocladiaceae,
and the Monoblepharidaceae. Gaumann (4) has Oomy-
cetes as the order in which he includes five families : the
Monoblepharidaceae, the Blastocladiaceae, the Ancylista-
ceae, the Saprolegniaceae, and the Peronosporaceae. Kan-
house (7) finds distinctive characters in this form and its
relatives to suggest the order Blastocladiales in which she
would place the family Blastocladiaceae. The varying sys¬
tems are the result of the inability to accurately and defi¬
nitely define the structural characteristics which are to de¬
termine the place of a species in its relation to others.
This difficulty is enhanced by a lack of knowledge of the
true nature of the various features of structure. Hence a
clear description of the phylogenetic relationships and a
natural system of classification is, as yet, next to impossi¬
ble. Therefore, it seems best for convenience to adopt
Coker's classification in which we find the family Blasto¬
cladiaceae composed of two genera, Blasted adia and Allo-
myces. In the first of these we find four species men¬
tioned ; B. pringsheimii Reinsch, B, ramosa Thaxter, B. ros-
trata Minden, B. prolifera Minden. Barrett (1) had prev¬
iously associated these four species and included with them
B. strangulata Barrett, which, for reasons we later con¬
sider, Coker places in the genus Allomyces and gives the
name adopted throughout these notes. The same plant is
also described by Coker and Grant (3) under the name
344 Wisconsin Academy of Sciences, Arts, and Letters.
Septocladia dichotoma. Hence in the literature we find
this one form labeled any one of the following names:
Blastocladia strangulata Barrett, Allomyces arbuscula,
Butler, and Septocladia dichtoma.
Up to 1923 with the publication of Coker’s work on the
Saprolegniaceae, A. airbus cula had been reported only from
India by Butler, from Ithica by Barrett, and from the Phil¬
ippines by Weston. Coker considers it rare in occurrence
having found it only once at Chapel Hill. Harvey (5) re¬
ported it from Chapel Hill and also has it in his collection
of Mississippi soils from which he kindly supplied us with
material.
For reference throughout this paper we reproduce with
some omissions Coker’s description of the genus and spe¬
cies:
“Genus Allomyces : Plant small, slender, the short or long
stalk not conspicuously differentiated; branches usually
dichotomous, often verticellate in groups of 3-5, separated
from the nodes by distinct and complete septa, not con¬
stricted at intervals; in vigorous cultures repeating the
branching in the same way to form a complex plant. Spo¬
rangia oval, terminal, sympodially arranged, not rarely in
chains of several often clustered by the shortening of the
branches which continue the stem by one or more lateral
buds beneath. Spores bi-ciliate at times, but the two cilia
so closely approximated or fused as usually to appear as
one. Resting bodies borne in the same way as the sporan¬
gia and of the same size and shape, at maturity enclosed in
a thin hyaline sheath out of which they finally fall through
an apical slit ; the wall brown and conspicuously pitted ; the
whole representing a thin-walled oogonium completely filled
with a thick-walled parthenogenetic egg, or resting sporan¬
gium as thought by Barrett.
A sparophytic aquatic of anomalous structure and differ¬
ing from all the other Phycomycetes in the regular and nor¬
mal septation of the plant body.
Species A. arbuscula Butler: Characters of the genus,
threads extending about 4 or 5 mm. from the substratum of
hemp seed (Coker uses ants) about 10/x thick, growing
gradually more slender distally at each joint; joints of cen¬
tral region more elongated than those near the base, tips
Lugg — Borne Notes on Aliomyces Arbuscula Butler . 345
blunt, hyaline. Sporangia oval, spores escaping singly or
at times, according to Barrett, in a vesicle that soon bursts,
emerging through one or two usually apical holes or short
papillae, spores biciliate, oval when swimming, with cilia
apical, monoplanetic, amoeboid before encysting, 10/x thick
when at rest; sprouting by a slender thread. Resting
bodies appearing later than the sporangia but of the same
size and shape; the conspicuous pits apparently sunken
from the outside in regular fashion as in B. Pringsheimii,
at maturity slipping from the thin clasping sheath ; sprout¬
ing into zoospores after a rest (Barrett). The thick wall
is divided into two parts, an outerlayer pitted about 1.8 p
thick and a homogeneous inner one about lfx thick/1
The uncertainty as to the details of the life history of
A, arbuscula is due primarily to lack of knowledge of the
true nature of the reproductive bodies observed. Appar¬
ently there are two groups of these and in observing this
plant they constitute the most conspicuous features of its
structure. The mature plant bearing its crop of resting
bodies is easily recognized by its brick red color due to the
presence of these reddish organs. The other group of
bodies are the sporangia which have a lighter color. A
given plant may produce either one or both sporangia and
resting bodies. It is agreed the sporangia are asexual in
their origin while the bodies which Coker calls “resting
bodies'1 are oosporic in nature.
The exact time when the plant will develop sporangia
seems to depend almost entirely upon external conditions,
such as the presence or comparative lack of food, the
amount of moisture present, and perhaps on the temper¬
ature. Most investigators have grown the plant on such
media as sweet corn agar (Barrett, Kanhouse) while Coker
used termite ants also. Our cultures have been maintained
on boiled hemp seed and have appeared to thrive. Such a
medium is easily prepared ; the growing plants are removed
from it with no degree of difficulty and the amount of nu¬
trient supplied can be controlled with some measure of suc¬
cess. The observation of Kanhouse (6) that morphologi¬
cal development of such fungi is intimately related to ex¬
ternal stimuli i can be well demonstrated. However, we
23
346 Wisconsin Academy of Sciences, Arts, and Letters.
would regard our medium as superior to any of the agars in
nothing but the convenience and simplicity of its manipu¬
lation.
Our first cultures were started over a year ago from three
colonies. During these succeeding months we have varied
the growth conditions over as wide a range as possible. At
no time have we had difficulty with obtaining material in
almost any stage of growth. New cultures have been
started by touching the cut side of a bisected hemp seed
with fragments of an active colony. These pieces of seed
material are then placed cut side down in a Petri plate con¬
taining a quantity of distilled water sufficient to cover the
plate to the depth of a millimeter. The plate is then cov¬
ered and kept in a darkened place at a temperature not
above 20 degrees Centigrade. Three or four days will re¬
sult in well developed plants.
In the presence of comparatively large quantities of wa¬
ter and abundance of food with a temperature not below
15 degrees, immense numbers of sporangia were produced,
arranged in long chains often sympodically upon long fila¬
ments of mycelium. No resting bodies appeared in the cul¬
ture thus maintained for the period of a week. If, how¬
ever, the food supply was allowed to run low or the quan¬
tity of water was reduced to the point at which portions
of the plant were exposed to the air, resting bodies made
their appearance while the production of sporangia gradu¬
ally ceased. These resting bodies were produced in great
numbers as long as any food was available or moisture pres¬
ent. To investigate the potential vitality possessed by the
resting bodies the following procedure was tried; two sets
of cultures were chosen eleven months ago — one of these
sets was allowed to dry out slowly while the other was kept
moist with sterile water. During the next nine months oc¬
cassional inspection showed no change in the moist cultures
except for the gradual disintegration of the mycelial mass
indicating death of the plant. The dry cultures assumed
the form of brick colored incrustations on the surface of
the glass. The finger when touched to a colony came away
tinged with a rust colored spot caused by free resting bodies
adhering to the skin. At the end of the nine months’ inter-
Lugg — Some Notes on Allomyces Arbuscula Butler. 347
val these dried masses were scraped into a test tube of sterile
water and thoroughly broken up by means of a round ended
glass rod. The tube was set aside a few minutes until the
bottom appeared tinged with the reddish color of the rest¬
ing bodies which had settled. The water was then decanted
off and more sterile water added when the shaking and de¬
canting processes were repeated. The third addition of
water carrying the resting bodies was then poured into a
sterilized plate containing hemp seed which had been boiled
after the splitting process for fifteen minutes. The plate
was then covered and set aside in a temperature of 16 de¬
grees. Vigorous growth appeared in two weeks, the plants
subsequently passing through the sporange-producing stage
to the development of resting bodies during the succeeding
three days. The same treatment was accorded the set of
cultures which had been kept moist. These also germinated
successfully after one week in the new medium. Some
trouble was experienced with bacteria and protozoa of a
few types, but these were easily eliminated by washing the
cultures with distilled water at intervals after they were
fairly well established. Changing the water occasionally
prevented the foreign organisms from overrunning the cul¬
tures. Sterile water was used only with the idea of avoid¬
ing as much as possible the introduction of forms as yet ab¬
sent from these particular cultures.
The results thus obtained indicate the correctness of the
supposition that the resting bodies produced in the oogonia
function as their name implies. But the length of time re¬
quired for their germination is not necessarily a matter of
months as implied by Barrett. It is true, however, that
these bodies do not immediately germinate as none are ever
found in the germinating condition near the colony which
gave rise to them.
A close study of the mycelium reveals some complexity.
In living plants one readily observes a difference in the
contents of various hyphae. In some filaments the presence
of immense vacuoles is conspicuous (fig. 1). These com¬
pletely fill the axial region of the filament, giving it the
appearance of being empty. In other hyphae the vacuoles
are smaller, the cytoplasm is decidedly granular, and bodies
of several different forms are present (fig. 2). In these
348 Wisconsin Academy of Sciences , Arts, and Letters .
latter types of filaments streaming of the cytoplasm has
been observed when currents are seen in the outer region
setting back toward the proximal end of the cell and re¬
turning through the center. All hyphae with granular con¬
tents do not exhibit streaming, however.
The plastides found in the hyphal contents are
of more than one sort as to composition. There
are large, nearly spherical bodies as large as 2/x in diameter.
These bodies in the living hypha are slightly yellow and be¬
come brilliant red with triple stain. Reinsch (9) regarded
them as “endogenously produced cells”. Pringsheim (8)
holds them to the waste products of metabolic processes.
Both PringsheinTs and ReinsclTs views are mentioned by
Barrett (1). These bodies exhibit no internal structures
as might suggest nuclei, nor do they contain granules of
any sort. Hence PringsheinTs view seems the more rea¬
sonable. There are, however, bodies of nearly the same
size as these red-staining plastids whose staining reaction
is entirely different. These bodies are invariably light blue
or lilac in tint and close inspection shows their outline to be
irregular. They are usually centers from which strands
of protoplasm radiate. From their position and general ap¬
pearance we regard the mas nuclei. Occasionally larger
ovoid bodies are observable which take very little stain,
either triple or haematoxylin. The probability is strong
that they are accumulations of fatty material.
At intervals there occur very slight constrictions in the
hyphal wall at which points also are found the characteris¬
tic cross walls which Barrett (1) describes as bearing pores.
This finding is not corroborated by any other worker as
far as we can ascertain, and the present material exhibits
nothing of the sort, either living or sectioned (fig. 3) . In
our observations the cytoplasmic content of the hypha on
one side of the wall is entirely cut off from that on the other
side. Of course ultra-microscopic pores may, and probably
do, exist. But as far as available evidence goes, we have in
this peculiarly septated hypha a multicelled structure. In
Barrett’s paper of 1912 (1) we note that the micrographs
of his entire plant, figures 58 and 59, show the presence of
sterile hairs while figure 57 has no indication of them and
neither does his habit sketch, figure 12. Yet all these speci-
Lugg — Some Notes on Allomyces Arbuscula Butler. 349
mens are supposed to be typical of the species under discus¬
sion. Coker’s arbuscula (2) lacks these hair-like filaments.
We believe therefore, the question might be raised as to
whether the author of the incomplete septation observa-
sions was dealing with one species or had inadvertently used
a form like B. pringsheimii or B. rostrata where there are
sterile hairs and no septa tions. Coker (2) is quite clear in
his observation that we have in A. arbuscula a multicelled
mycelium.
At the points where cross walls are formed it will be ob¬
served that the hypheal diameter is increased forming a
slight bulge either side of the wall. These enlargements
could easily be due to a tendency to contraction of the cyto¬
plasm acting to shorten the filament. No bulges are ap¬
parent at young cross walls but seem to be a development
with age. Some change in cytoplasmic volume, therefore,
in time serves to cause this apparent constriction. The di¬
rection of least resistance and therefore of the greatest
change of shape is at right angles to the hyphal axis. But
the added reinforcement of the cross wall prevents a shrink¬
age at points where it is attached. Thus any reduction in
diameter is more apparent than real.
The basal cell from which a hyphal system is derived
is quite difficult to make out on material grown on hemp
seed. The cell is larger in diameter than the rest of the
hypha. It is also much shorter and Barrett’s description
mentions rhyzoids (1), but we have not seen them. The
hyphal cells vary in length ; when growth is slow they tend
to be short, while rapid growth results in long filamentous
cells bearing branches, each cell constituting a considerable
portion of the mycelium of one plant.
Under conditions which have not yet been worked out,
the tip of a growing hypha begins to enlarge taking on a
bulbous form. At the same time there is apparent a con¬
centration of nuclei near the tip of the filament (fig. 4).
The cytoplasm becomes lighter in appearance and lacks al¬
ways the large vacuoles which are often visible in some fila¬
ments. The nuclei that may number a score or more as¬
sume positions equidistant from each other and in a region
near the wall of the hypha. At this stage the bulbous de¬
velopment has resulted in an ovoid tip whose surface region
350 Wisconsin Academy of Sciences , Arts , and Letters.
is marked by the presence of these irregularly shaped nuclei
which now appear to be connected to each other by strands
of denser protoplasm (fig. 4). Much study might be ex¬
pended upon the method of cross wall formation at this
point in the development process. Apparently the first
evidence of a cross wall cutting off the bulbous terminal from
the filament is a line of granules extending from wall to
wall (fig. 5) . Whether or not a plasmal membrane has pre¬
ceded the appearance of these minute solid particles is un¬
certain. From the material we have observed, we believe
such to be the case as the cytoplasm in the sporange takes
the haematoxylin more readily, and the line of demarcation
between sporangial cytoplasm and that of the hypha is
quite distinct, although there is no separating medium ap¬
parent. Through accumulation of granular material a solid
partition is at last developed which takes on the character
of the hyphal wall. This process may be repeated in the
hyphal region immediately proximal to the first sporange.
Successive repetitions of this process produce a chain of
sporangia.
Thus far there has been no evidence of the peculiar fur¬
rowing process of spore development that is often seen in
Phycomycetes. But after the appearance of the cross wall
large vacuoles appear and the stranded structure disinte¬
grates into masses of varying density indicating the cleavage
furrows by which the cytoplasm is divided as shown by Bar¬
ret (1). This author also describes the subsequent behav¬
ior of the zoospores produced. They emerge from the spo¬
rangium through specially developed structures, papillae of
dehiscence, of which there may be one or several on the
walls of the sporange. Those we have seen are meniscus in
cross section and as Barrett observes, are made up of two
parts : an inner membrane surmounted by a convex hya¬
line elevation. The rupture of one or more of these papil¬
lae permits the meregence of the zoospores in single file.
These are uni-, bi, or tri-ciliate. The biciliate form is the
normal one and Coker (2) would admit the triciliate forms
as only bare possibilities.
Not all spores are discharged in some cases and those
which are retained in the sporangium become amoeboid and
remain in the sporange until they encyst (fig. 7). The
Lugg — Some Notes on Allomyces Arbuscula Butler. 351
empty sporangia are retained in position on the plant, often
forming long chains of six or more, some of which have
emptied sooner than others. No relation seems to exist as
to the order in which these members of a chain shall release
their spores.
It has not been our fortune to observe the zoospores in
the act of emerging but Thaxter (10) gives a short account
of the ciliated spore and Coker (2) includes diagrams of the
internal structure. It is not hard to find encysted spores in
a culture and also many such in the process of germinating
(fig. 8). The encysting wall is not marked nor of great
thickness and is hyaline. Before germination, stained
spores show a reticular cytoplasm. Two or three large va¬
cuoles are evident as are a number of large granules. Since
there is no great mass of undifferentiated protoplasm pres¬
ent, the entire contents of the spore may be a nucleus. There
is usually a single body more or less refractive that, by its
large size, may correspond to a nucleole. Spores are located
whose entire contents is granular and a smaller body within
this mass exhibits a reticulum. Spores sprout by hyphal
extensions. Hence the possibility that, as the spore enters
the germinating stages, the granular material is absorbed
as the nucleus enlarges. Germination results in the exten¬
sion of a slender thread of granular cytoplasm surrounded
by a hyaline sheath. Also we have often observed spores
whose contents have divided before the filamentous exten¬
sion is produced (fig. 9). If chromosome material exists
in distinct characteristic bodies anywhere in the structure
of the organism one would expect it apparent at this stage.
We do find several rather distinct granules in the spore con¬
tents especially at certain stages of germination. These are
too small to exhibit shapes which would make possible the
identification of any one chromosome. Yet these bodies
might well function as such. For cytologists are practic¬
ally united in the opinion that it is only the abnormal nucleus
that lacks chromosomes.
In the usual progress of growth in culture, the sporan-
gial period is followed by a later stage of growth character¬
ized by the appearance of large numbers of resting bodies.
Coker (2) suggests that they partake of the nature of par-
thenogenetic eggs. Since no other sex organs have been
352 Wisconsin Academy of Sciences, Arts, and Letters .
observed, if sex has dropped out of the life cycle, here is the
only remnant of that process. And we assume sex to have
disappeared to that point where antheridia have become
totally undeveloped. Of course one may regard A. arbus-
cuia as ascendant in the phylogenetic scale instead of degen¬
erate. If such a view be taken, it is hard to account for
the complex structure of these resting bodies. In early de¬
velopment they are practically identical with the sporangia
which have preceded them. They exhibit the reticular cy¬
toplasmic structure and regular arrangement of the nuclei
enclosed in the ovoid terminal of the hypha. This likeness
persists beyond the development of the cross wall. How¬
ever, groups of these bodies never occur in chains for the
reason that after the formation of a body at the hyphal ter¬
minal further growth occurs in the hypha by a lateral bud
at the base of the resting body. Until further work is done
on the cytology of the resting body we can go but little be¬
yond the facts we have mentioned. But this question might
be raised: from present knowledge of the nuclear material
of the hypha, what is to prevent the formation of these rest¬
ing bodies in chains similar to the sporangia? Their origin
is the same; their method of development is apparently
identical up to a very late stage of growth. Why then,
might not the contents of a considerable portion of the dis¬
tal end of a hypha break up into successive portions as hap¬
pens with the sporangia? We feel the answer will be found
in the behavior of the nuclear material of the resting body
immediately prior to the formation of the cross wall. By
some means these nuclei undergo an abnormal change.
This could take the form of a reduction process which would
be the natural thing to expect if this material were to form
oospores or eggs.
The next period of growth in the resting body is indicated
by the appearance of a clear refractive wall within the mem¬
branous sheath which originally functioned as the filament
wall and in which appear the papillae of dehiscence if the
body is a sporangium. This new inner coat appears to be
secreted from the outer cytoplasm. It attains some thick¬
ness before any indication of the characteristic pits appears.
However, soon the density increases, and the indentations
become evident. These pits are conical in longitudinal sec-
Lugg — Some Notes on Allomyces Arbuscula Butler. 353
tion with the wider diameter at the outside (fig. 11) . These
pits do not extend through the wall, but penetrate it to a
region which can be made out below the pitted stratum and
which is uniform in density. Coker regards these layers as
separate walls (2). His reason for doing so is not quite
apparent. In mature resting bodies the sporogenous mass
is surrounded by a second membranous wall within the pit¬
ted coat. This membrane is easily observed in material
that is slightly plasmolyzed.
As to the behavior of the mass of cytoplasm within this
case, little that is definite can be said. The numerous dark¬
ly stained nuclei arranged peripherally are not evident in
the mature resting body. The contents are uniformly gran¬
ular, or, at most, marked with a few masses of granular
structure a little more dense than the rest. However there
can be found a group of resting bodies whose walls are com¬
pletely formed, but whose contents consist of a couple of
dozen spherical bodies which stain red in safranin and in
living material are clear and slightly yellow. These may be
fat globules. If these fatty accumulations appear in the la¬
ter stages of development the question of what has become
of the nuclear bodies cannot be answered. There is a group
of bodies also whose contents resemble figure 12. These
bodies exhibit markings of some form of cell division. In
living material detached bodies are granular and fat-con¬
taining ones are never found free. Release of the resting
bodies takes place by the rupture of the external membran¬
ous wall when the body floats free, and even the part of the
heavy wall formed at the point of attachment to the hypha
shows pits (fig. 11). Coker mentions the cracking of the
pitted coat to release the material from which, he assumes,
a new mycelium is produced (2). From the nature of the
contents of the resting body and from the fact that the re¬
sult of the germination of these contents is a form no dif¬
ferent from that resulting from the germination of the zoo¬
spores, we may safely conclude that these resting spores
are not radically different in their make-up, unless some nu¬
clear modifications take place in the zoospores prior to ger¬
mination. This also strengthens the hypothesis of parthe¬
nogenesis.
354 Wisconsin Academy of Sciences, Arts, and Letters.
Summary
1. A convenient method of raising A. aribuscula has been
worked out.
2. Additional evidence of complete septation is provided.
3. The growth history of Coker's “resting bodies" has
been studied providing further indications of their zoogo-
nial nature.
Literature Cited
1. Barrett, J. T. The development of Blastocladia strangu -
lata . Botanical Gazette. 45: 353. 1912.
2. Coker, W. C. The Saprolegniaceae. Chapel Hill, N.
Car. 1923.
3. Coker, W. C., and Grant, F. A. A new genus of water
mold related to Blastocladia. Journ. Elisha Mitchell
Sci. Soc. 37; 180. PI. 32. 1922.
4. Gaumann, Ernst. Ein Beitrag zur Kenntnis der lapp-
landischen Saprolegnieen. Botaniska Notiser for
1918, p. 151.
5. Harvey, J. V. A study of the water molds and Pythi-
ums occurring in the soils of Chapel Hill. Jour. Eli¬
sha Mitchell sci. Soc. 41 : 151-164. Sept, 1925.
6. Kanhouse, B. B. On the distribution of the water
molds, with notes on the occurrance in Michigan of
members of the Leptomitaceae and Blastocladiaceae.
Papers of Mich. Acad. Sci, Arts and Letters.
5:105-114. 1925.
7. Kanhouse, B. B. A monographic study of special
groups of the water molds. Jour, of Botany,
14:287-357. 1927.
8. Pringsheim, N. Ueber die vermeintlichen Amoeben in
den Schlauchen und Oogonien d. Saprolegnieen. Bot.
Centralblatt. 14:378. 1883.
9. Reinsch, P. F. Beobachtungen iiber einige neue Sapro-
legnieae. Jahrb. f. wiss. Bot. 11 :283. 1878.
10. Thaxter, R. New and peculiar aquatic fungi. Bot.
Gaz. 21 :45. 1896.
TRANS. WIS. ACAD. - VOL. 24 PLATE 8
Lugg — Some Notes on Allomyces Arbuscula Butler. 355
Explanation of Plate 8
Fig. 1. Vacuolated cytoplasm in hypha. X 1900. Haematoxylin
stain.
Fig. 2. Types of hyphal plastids. X 1900. Triple stain.
Fig. 3. Typical cross-wall. X 1900. Triple stain.
Fig. 4. Developing sporange. X 1900. Safranin and licht grim
stain. The peripheral arrangement of the nucleii is often more pro¬
nounced than is shown here.
Fig. 5. Granular origin of the cross-wall of sporange. X 1900
Triple stain.
Fig. 6. A later stage in sporange development. X 950. Haema¬
toxylin stain. The arrangement of nucleii is very evident here as is
also the difference in staining reaction of contents of the sporange and
of the hypha.
Fig. 7. Amoeboid zoospores entrapped in sporange. X 950. Liv¬
ing material. The nature of the rupturing process of the papillae of
dehiscence may be easily made out as well as something of their
structure.
Fig. 7a. An empty sporange. X 440. Living material.
Fig. 8. Germinating spore. X 900. Living material.
Fig. 9. Spore which has divided without development of a process.
X 900. Living material.
Fig. 10. Young resting body with first indications of pitted wall.
X 1900. Living material. Contents not drawn.
Contents not completely shown.
FlG. 11. Mature detached resting body. X 1900. Triple stain.
Contents not completely shown.
Fig. 12. Spores found in mature resting body. X 2300. Triple
stain.
PRELIMINARY REPORTS ON THE FLORA OF WIS¬
CONSIN. III. LORELIACEAE, CAMPA-
NULACEAE, CUCURBITACEAE1
Kenneth L. Mahony
LOBELIACEAE
This family is represented in Wisconsin by the genus
Lobelia , with six species.
L. cardinal! s L. (fig. 1) This species is general over the
state with the exception of the extreme northern part.
L. SYPHILITICA L. (fig. 2) This is general over the state
except in the northern part. In common with many other
plants of southern range it is found in the valley of the
Wisconsin River north to Lincoln County and in the St.
Croix River valley north to Burnett County.
L. spicata Lam. (fig. 3) Is of southern range in Wiscon¬
sin, being found chiefly in the southwestern part of the state.
However, var. hirtella Gray (fig. 4) tends to run north¬
ward in the eastern part of the state to Marathon and Mari¬
nette Counties.
1 The ranges presented here are based on the collection in the Herbarium
of the University of Wisconsin and that of the Milwaukee Public Museum.
358 Wisconsin Academy of Sciences, Arts , and Letters .
Mahony — Preliminary Reports on Flora of Wisconsin. III. 359
L. Kalmii L. (fig. 5) This occurs principally in the south¬
ern part of the state, centering in Dane, Sauk and Colum¬
bia Counties along the Wisconsin River and also in Milwau¬
kee, Waukesha and Racine Counties in the eastern part of
the state. It has also been found in Door County and as far
north as the Apostle Islands and should be looked for in the
lake region of northern Wisconsin.
L. inflata L. (fig. 6) Is found to be quite common all
over Wisconsin.
L. Dortmanna L. (fig. 7) This species is entirely of
northern range, being found in the lake regions of Sawyer,
Oneida and Calumet Counties.
CAMPANULACEAE
This family is represented in Wisconsin by five species of
the genus Campanula and one species of the genus Specu -
laria.
Campanula rapunculoides L. A garden escape ; Racine
(J. J. Davis, 1879). Well established, according to Wad-
mond.2
C. Americana L. (fig. 8) Occurs in the southern part of
the state and also the western part along the Mississippi
and St. Croix River vallies as far north as Polk County.
This is a range that is characteristic of many plants of
southern Wisconsin.
C. rotundifolia L. (fig. 9) Is rather general through¬
out the state.
C. ULIGINOSA Rydb. (fig. 10) This is of northern range
in Wisconsin.
C. aparinoides Pursh. (fig. 11) Is found in the south¬
ern part of the state and north along the Wisconsin River
to Wisconsin Rapids. It has also been found in Polk
and Burnett Counties.
While there are well developed extreme specimens of C .
uliginosa and C. aparinoides which are quite distinct, there
are many intermediates which cause some confusion. Or¬
dinarily the little-branched C. uliginosa has large flowers
and linear leaves, while C. aparinoides is more branched,
•Trans. Wis. Acad. Sci., Arts and Let. 10: 869. 1909.
360 Wisconsin Academy of Sciences, Arts, and Letters,
Mahony — Preliminary Reports on Flora of Wisconsin. 111. 361
with small flowers and lanceolate leaves. Certain individ¬
uals which we term intermediates have small flowers and
linear leaves, or large flowers and lanceolate leaves, and are
very difficult to place. The characters that were principally
relied upon in this study were those given by Fernald and
Wiegand, Rhodora 25: 214. 1924.
C. aparinoides : naked portion of peduncle 0.3-3. 5 cm.
long; flowering calyx 1. 3-3.8 mm. long, its lobes 0.7-2 mm.
long; capsule 1.2-2 mm. long.
C. uliginosa : naked peduncle 1-6 cm. long ; flowering ca¬
lyx (3-) 4-6.7 mm. long, its lobes 2-4 mm. long; capsule
3.2-5 mm. long.
Specularia perfoliata (L.) A. DC. (fig. 12) This spe¬
cies is of southern range in Wisconsin, running north as far
as Jackson County in the western part of the state.
CUCURBITACEAE
This family is represented in this state by two genera,
each with one species.
SICYOS ANGULATUS L. (fig. 13) Comparatively few col¬
lections have been made of this species. These collections
have been made in the southern part of the state and along
the Mississippi River as far north as Pepin County.
Echinocystis lobata Torr. & Gray. (fig. 14) Is found to
be quite general over the state. It is extremely abundant in
the valley of the Mississippi River.
24
A STUDY OF TWO LIMESTONE QUARRY POOLS
Edward Joseph Wimmer
Introduction
Very little work has been done in the United States on
the chemical and plankton conditions in pools and ponds.
Scott (1910) made a study of the fauna of a solution pond.
Reed and Klugh (1924) studied the hydrogen ion concentra¬
tion of a granite and a limestone pool near Kingston in the
Province of Ontario. In Europe, more attention has been
given to this field. Griffiths (1916, 1922) and Atkins and
Harris (1924) have studied the heleoplankton and chemical
factors in quarry pools and ponds. Among the many other
workers are Alexander, Rylov, Nordqvist, Scahferna, Fric,
Diefenbach, and Sachse.
This paper is based upon a study of the physical, chemi¬
cal, and plankton conditions of two limestone quarry pools
near Milwaukee, Wisconsin. The larger of the two, which
shall be called the Wauwatosa pool, lies near the western
city limits of Milwaukee, in a recently annexed portion of
land, adjacent to the suburb of Wauwatosa. It is the site
of an old limestone quarry, in which operations ceased
about thirteen years ago, and which, since then, has been
slowly filling with water as there is no outlet, until it is now
23 meters (75 feet) deep. It is roughly rectangular in
shape with the long axis in an east-west direction, and cov¬
ers an area of approximately 1.36 hectares (3.36 acres).
The walls on all sides are precipitous, and rise on the south
side of the pool to a height of about 5 m. The height of the
walls decreases toward the north side where it is only about
1 m. above the surface of the water. On all sides the walls
descend abruptly to a depth of 5 m. except for a few re¬
cently submerged shallows, which are small in extent. The
limestone about the pool is overlaid by glacial gravel, which
forms a moraine on the south side, the crest of which is
about 200 m. from the quarry. The other sides are com-
364 Wisconsin Academy of Sciences, Arts , and Letters.
paratively low. The drainage into the pool is primarily
from the moraine to the south. Except for dwellings on
the crest of the moraine, there is little contamination from
civilization. Much carbon, however, settles on the pool,
from smoke of a railroad a few city blocks to the north, and
from factories nearby. The pool receives plenty of sun¬
shine due to its east-west position, and to the lack of vege¬
tation along the eastern and western shores.
Along the sides is a ledge about 5 m. below the surface
of the water. This ledge is very narrow along the eastern
three-fourths of the quarry, but forms the entire floor of
the quarry at its western portion. From this ledge the bot¬
tom descends abruptly again to a ledge at a depth of 15 m.,
which is the depth of the larger portion of the pool. In the
northeastern portion, at the site of the former “quarry
hole” is a large area, about one-fifth the area of the pool,
which is 23 m. deep. The bottom of the pool to within a
year ago was comparatively free from ooze. A sample of
the bottom brought up with a mud dredge showed the
greater part to consist of coal dust or carbon, and to be ap¬
proximately about two inches in thickness. In September
1926, clay was emptied into the pool. Within a compara¬
tively short time, it became distributed over the bottom of
the pool.
The rise of the water level of the pool during the time it
was studied is of interest. A photograph taken of the pool
in May 1921 shows the water level to be about 9 m. below
the level of August 1925 when this study was begun. Be¬
tween August 1925 and March 1928 the water level rose
about 2.1 m.
The study of the second pool was begun in September
1926. This also is an abandoned limestone quarry, about
five miles to the northeast of the one just described. This
pool is on the south limits of North Milwaukee, and will be
called the North Milwaukee pool. This body of water is
about one-third the size of the Wauwatosa pool, and is also
well isolated from contaminating sources. The walls rise
vertically from the bottom on all sides. The average depth
is about 4.5 m. The deepest place in the pool is 5.5 m. The
water level is less than 1 m. below the level of the solid rock.
This is covered with glacial gravel, overgrown on all sides
Wimmer — Study of Two Limestone Quarry Pools. 365
with grass. On the western side (the pool's long axis is in
a north-south direction) the gravel forms a ridge about 3 m.
high, with steep sides. There is little vegetation on the
shores except for a clump of trees in the southeast corner
and a few low wild crab trees near the center of the west
shore. The other shores are low. The eastern side has a
low dike to prevent a creek about 5 m. distant from over¬
flowing into the pool in times of flood. The level of the
creek is slightly lower than that of the pool and lies in solid
rock. The bottom of the pool is composed of soft black
ooze, but its depth is not known. About five years ago al¬
most all of the water was pumped from the pool after the
creek had overflowed into it. At this time the dike was
built. In winter ice is harvested from this pool, but not
from the Wauwatosa one. The smaller pool is about eight
to ten years old.
Methods
Practically all of the samples from the Wauwatosa pool
were taken in the open water at the spot where it was deep¬
est (23 m.) and about 10 m. from shore. This was made
possible by the use of a small flat-bottomed boat kindly
loaned by Mr. W. Manegold, who lives on the property, and
with whose kind permission and hearty cooperation this
work was made possible. At times it was impossible to use
the boat, and samples were then taken from the southwest¬
ern portion where the wall was vertical for about 5 m.
Samples were taken as far from shore as was possible by
pushing the line from the shore with the aid of a plank a
little more than a meter long. Samples from the North
Milwaukee pool were taken in the same manner from a
point near the middle of the west shore. During the win¬
ter, samples of the North Milwaukee pool were taken from
the center of the pool. So far as possible samples were
taken during the same time of the day. This was between
nine and twelve a. m.
All of the limnological apparatus was loaned by the Wis¬
consin Geological and Natural History Survey, through the
courtesy of Professor C. Juday, under whose supervision
and guidance this work was done.
The net plankton was collected by means of a closing net
366 Wisconsin Academy of Sciences , Arts , and Letters.
with No. 25 bolting cloth ( Tuday 1916). Hauls were made
from 0 to Y2 m., to 1 m., then 1 m. hauls to generally 5
m. and in the Wauwatosa pool further hauls of 5 m. each
to 20 m., and from 20 to 23 m. During 1927 horizontal
surface hauls were made. One was just below the surface
while the second haul was made about 10 m. below the sur¬
face. Each of these hauls of the net was for a distance of
2 to 3 m. The catch was killed and preserved immediately,
and before counting, was diluted to 20 cc.
A Sedgwick-Rafter counting cell was used in making the
plankton counts. The plankton in the entire cubic centi¬
meter contained within the cell was counted by the aid of a
mechanical stage, for such organisms as were not found in
great numbers. The abundant organisms were counted by
enumerating twenty squares with a Whipple micrometer
(Whipple, 1927). Entomostraca, when in small numbers,
were counted by looking at the total catch with a hand lens.
All of the counts were computed on the basis of the number
of organisms per liter.
Temperatures were taken with a Negretti and Zambra
deep sea reversing thermometer. It was generally found
necessary to take temperatures for every meter to 5 m., and
often this procedure had to be continued to 10 m. Then
readings were made at 5 m. intervals to the bottom, and
intermediate readings were taken whenever necessary.
The turbidity of the water was taken with a Secchi disc
10 cm. in diameter, and the depth at which it disappeared
was noted.
Dissolved oxygen was determined according to the modi¬
fication in Standard Methods of Water Analysis (1925) of
the Winkler Method (Winkler, 1888). Carbon dioxide was
determined by the Seyler method as given by Birge and Ju-
day (1911).
The hydrogen ion concentration was determined colori-
metrically using cresol red (pH 7.2 to 8,8) and thymol blue
(pH 8.2 to 9.8) as indicators and making a comparison with
the Clark Color Chart of Indicators (Williams and Wilkins
Co. 1922).
Nitrogen determinations were made as directed in Stan¬
dard Methods of Water Analysis. Free ammonia, organic
Wimmer — Study of Two Limestone Quarry Pools . 367
nitrogen, nitrates, and nitrites, were determined. Nitro¬
gen determinations were not begun until May 1927.
All of the chemical work other than oxygen, carbon diox¬
ide, and hydrogen ion concentration was done in the labora¬
tory of the Wisconsin State Laboratory of Hygiene, and
thanks are due to Dr. W. D. Stovall and to Dr. M. S. Nichols
for permission to use the laboratory. To Dr. B. P. Domo-
galla, Biochemist for the City of Madison, the author is
much indebted for guidance and assistance in all of the
chemical analyses in the laboratory, and also in assisting in
the transportation of the samples of water to Madison.
Soluble and total phosphorus were determined by the
modification by Domogalla of the method of Juday, Kem-
merer and Robinson (1927).
Chlorides were determined as directed in Standard Meth¬
ods of Water Analysis.
Soluble silica was determined by the Atkins modification
(1923) of the colorimetric method of Dienert and Wanden-
bulcke (1923).
The organic matter of the centrifuge plankton was deter¬
mined by centrifuging one liter of water in a Foerst electric
centrifuge (Juday 1924 and 1926). The organisms from a
liter of water (concentrated to 4.5 cc. of water left in the
bowl of the centrifuge) were transferred to weighed plati¬
num dishes of 8 cc. capacity. After drying at 60 °C. for 24
hours, the dishes were cooled in a desiccator and the dry
weight was ascertained. They were then ignited in an elec¬
tric furnace and after cooling were again weighed and the
loss on ignition was determined. Since the centrifuge bowl
contained 4.5 cc. of water, the amount of volatile matter in
solution in the water must be deducted from the loss on igni¬
tion. This factor was found by determining the loss on
ignition of 10 cc. of the centrifuged water, and the amount
of such “blank” volatile matter in 4.5 cc. was subtracted
from the loss on ignition of the centrifuged plankton.
In addition to the liter of water centrifuged for organic
matter, 500 cc., or sometimes 250 cc., were centrifuged and
diluted to 10 cc. for counts of the nannoplankton organisms
which escape through the meshes of the closing net.
368 Wisconsin Academy of Sciences , Arts , and Letters.
Temperatures
A comparison of the temperatures of the surface water
with the monthly meteorological tables issued by the Mil¬
waukee office of the United States Weather Bureau shows
that the water temperature increases more rapidly than
that of the air, and that it cools more slowly. The high
specific heat of water also prevents its showing as great
variations as are shown by the air. In the spring the heat
absorbed during warm periods is not dissipated to any great
extent during the cool spells so that the water continues to
increase in warmth at a comparatively uniform rate.
In the fall the water continued to cool until it reached a
temperature near 0°C. This temperature was not reached
in the Wauwatosa pool until early in December ; but in the
North Milwaukee pool, since there was a smaller volume of
water to be cooled, the temperature reached this point as
much as two weeks earlier. On the first quiet, cold night,
ice formed on the pool. This occurred for the Wauwatosa
pool on December 18, 1925, on December 4, 1926, and on
December 20, 1927. The smaller pool froze on November
13, 1926, and on December 8, 1927.
The ice left the smaller pool earlier than it did the large
one for there was less water to be warmed. Ice left the
North Milwaukee pool on March 13, 1927 and on March 20,
1928. The large pool was free of ice on April 17, 1926,
March 17, 1927, and March 28, 1928. The maximum tem¬
peratures at the times the pools were visited was 24.0° on
July 10, 1926 for the Wauwatosa pool, and 25.7° on July
16, 1927 for the North Milwaukee one. The minimum sur¬
face temperature for the large pool was 1.5° on January 3,
1926 while for the smaller pool it was 0.4° on January 2,
1927.
The Wauwatosa pool in 1925-1926 showed typical lake
conditions in regard to temperature. At the beginning of
August, 1925, the thermocline or layer of rapid change of
temperature began at about 5 m. On August 22 when the
boat was used for the first time, the top of the thermocline
was at 6.5 m. The thermocline continued to 12 m. with a
temperature change from 21.5° to 5.7°. On September 3
the temperature in this layer did not change to any extent.
Wimmer— Study of Two Limestone Quarry Pools.
Seasonal changes in temperature, expressed in degrees centigrade, at various depths in the Wauwatosa pool.
The depths are indicated in meters by the figures attached to the curves.
370 Wisconsin Academy of Sciences , Arts , and Letters.
The epilimnion, or layer above the thermocline, had cooled
a degree (21.0°). The hypolimnion or layer below the
thermocline began at 12 m. as before, with a temperature
of 5.7° which was uniform thence to the bottom (23 m).
By October 10 the surface had cooled to 16,5° bringing the
epilimnion down to 8.5 m. The thermocline continued as
before to 12 m. with the same temperature. On October 24
the surface layers had cooled uniformly to 10.9°, down to a
depth of 11 m. at which level the temperature was 10.4°.
This marked the lower limit of the epilimnion. Since the
temperature of the hypolimnion remained constant, this
narrowed the thermocline to only one meter with a fall in
temperature of 4.5°. On November 14 the temperature
was uniform from the surface to the bottom, namely 5.5°.
On November 26 it had further cooled one-fourth of a de¬
gree. Ice formed on December 18, to a depth of one inch
or 2.5 cm. By December 24 the ice was 2*4 inches or 6.3
cm. thick and on January 2, 1926 it was 6 inches or 15 cm.
thick. During the time observations were made, the ice
reached a maximum thickness of 15 inches.
During the winter the water at a depth of 23 m. main¬
tained a temperature between 4.0° and 5.0°. At 10 m. and
15 m. the temperature remained near 3.5°, while at 5 m.
the temperature slowly rose until in February, 1926 it was
almost the same as that for the 10 and 15 m. levels. The
surface in the meantime had also become warmer, so that
by the second week in April, when the ice left the pool, the
surface water was practically the same as that for the rest
of the levels. (Fig. 1).
With the increase in the length of the day in spring and
a consequent increase in the number of hours of sunshine,
the surface water warms very rapidly. At this time the
temperature of the water is slightly more than 5.0°, and,
being of a lower density than the water beneath it (which
is close to 4°), tends to float on the surface. This pre¬
vents winds and convection currents from mixing it with
the lower water so that the warm surface layer, or epilim¬
nion, is formed. (Birge 1908). This layer was not
clearly marked off until May. On May 9 it was only 1 m.
thick. Below it came the thermocline which extended to
5 m. with a fall in temperature from 7.7° to 3.7°. On May
A/eiers
Wimmer — Study of Two Limestone Quarry Pools. 371
29 the upper layer was still one meter in thickness but the
thermocline extended to 7 m. By June 22, 1926 the sur¬
face water had warmed to 20.0° and the epilimnion ex¬
tended to 4 m. The thermocline had its lower level at
10 m. with a temperature of 8.2°. The hypolimnion had
been warming up slightly. This condition continued for
the summer with the epilimnion increasing in depth and
warming slightly. The coldest water was near the 18 m.
level where the temperature was below 4.9° at all times.
(Fig. 2).
0°C 2. 4- 6 8 10 \Z 14 16 18 ZQZZ& 0° 10° £0°
Fig. 2. Diagrams showing the changes in the temperature of the
upper water between January 2 and August 19, 1926 in the Wauwa¬
tosa pool.
After September 16, 1926, clay was emptied into the pool
as already stated, which warmed the entire hypolimnion
and raised its upper level to 8 m. The entire water of this
lower layer had a temperature of 8.4°, and it steadily
372 Wisconsin Academy of Sciences , Arts, and Letters .
warmed so that the overturn, or period of uniform temper¬
ature, was brought on much earlier than usual. (October
30).
During the summer of 1927 the boat could not be used so
that bottom temperatures of the deep water were unavail¬
able. When it was used again in September 1927, the bot¬
tom temperature was the same as that for the 5 m. level.
This was probably due to the fact that clay was still being
emptied into the pool and continued to keep the lower water
at a temperature higher than usual.
The temperatures of the North Milwaukee pool were
substantially the same from surface to bottom, since the
pool showed no stratification. This condition is typical for
small, shallow lakes.
Turbidity
The turbidity of the water as measured by the Secchi
disc, corresponded in general with the plankton growths at
the time. In the case of the phytoplankton, an increase in
numbers will increase the turbidity and so cut off the pene¬
tration of the sun’s rays to the deeper waters and thus limit
the growth of these organisms to the upper water. Since
the North Milwaukee pool had at nearly all times a heavier
growth than the large pool, the turbidity was also found to
be higher than that for the Wauwatosa pool. The deepest
level at which the disc disappeared in the North Milwaukee
pool was 2 m. in July and September 1927, and in Febru¬
ary 1928. This was the minimum turbidity. The greatest
turbidity occurred in October 1927 and March 1928 when
the disc disappeared at 0.5 m. The Wauwatosa pool had a
much clearer water, in spite of the fact that clay was
emptied into it. The minimum turbidity was found in
August and September 1926, before the clay altered condi¬
tions. At this time the disc disappeared at 10 m. Im¬
mediately afterward, the disc was lost to sight at 6 and then
3 m. in October. Coincident with this, however, was an
increase in the growth of plankton. The greatest turbidity
in the large pool was found on November 30, 1926 when
the disc disappeared at 0.75 m. At this time there was a
heavy diatom growth and clay was still being dumped into
the pool.
Wimmer— Study of Two Limestone Quarry Pools.
Chemical Results
Oxygen. The story of dissolved oxygen for the Wauwa¬
tosa pool is very similar to that of the temperature, and is
very much in accord with the findings of Birge and Juday
and of other workers on Wisconsin Lakes. (Fig. 3).
OscOmS. /Liter 2.5
0^ Orr^. /Liter g
50
10
TO
14
2
£
u
T- o°
pH- 7.0
10
s.o
ao°
ao
28*
98
Fig. 3. Vertical distribution of the dissolved gases in the Wauwa¬
tosa pool on September 2, 1926.
Oxygen determinations were made in October 1925 and
were then resumed in March 1926. In October 1925 the
bottom water contained no free oxygen. The March de¬
termination showed no oxygen to be present in the bottom
water, so that whatever had been acquired by the overturn
of the fall had been used in the process of respiration and
oxidation. Since the covering of ice prevented the circu¬
lation of water, the lower water remained without oxygen
37 4 Wisconsin Academy of Sciences, Arts, and Letters.
until the spring overturn. By April 26, 1926, the bottom
water had acquired slightly more than 2 mg. per liter of
oxygen or 2 p.p.m., and this steadily rose until May 8 it
reached its maximum of 3.3 p.p.m. By this time the epi-
limnion had formed, effectively cutting off the lower water
from contact with the air, and in this manner depriving it
of further oxygen. The oxygen in the lower water was
gradually consumed until by July 10, it had again disap¬
peared in the water below 19 m.
With the dumping in of clay in the latter part of Sep¬
tember, oxygen was brought to the lower water, probably
partly by absorption to particles of clay and partly by
carrying down some of the surface water. The amount of
oxygen at the lower levels at once rose and soon was the
same as that at 10 m. With the overturn in the fall, all
layers contained oxygen, ranging from 80 to almost 100 per
cent saturation. The bottom water now continued to be
supplied with oxygen due to the dumping of clay, which
continued for a few weeks after the ice had formed. In
this way the amount of oxygen did not decrease in the bot¬
tom water as it had during the winter of 1925-1926. Dur¬
ing the spring of 1927 and during most of the summer the
bottom water could not be investigated due to the leaky
condition of the boat. When a test was made on October
10, 1927 before the fall overturn, the bottom water con¬
tained 5.5 p.p.m. of oxygen as compared to 1.0 p.p.m. the
year before. It must be remembered that clay was con¬
tinually being emptied into the pool about 150 m. from the
deepest portion.
The North Milwaukee pool presented no such conditions.
It was found that the temperatures varied little between
the surface and bottom. The same was true for oxygen.
The greatest difference between the surface and the bottom
was on March 12, 1927 when the surface held 14.7 p.p.m.
of oxygen in solution as compared with 6.0 p.p.m. for the
bottom water. Unlike the larger pool, the North Milwau¬
kee pool reached and even exceeded saturation with oxy¬
gen, several times during the year. The maximum surface
saturation was on November 26, 1926 with 138.1% satura¬
tion or 18.7 p.p.m. of oxygen. A similar high value was
reached on December 7, 1927 with 135.3% or 19.3 p.p.m.
Wimmer — Study of Two Limestone Quarry Pools. 375
The minimum saturation percentage was found in October
of both years-— 63.9% or 6.62 p.p.m. in 1927 and 44.9% or
4.5 p.p.m. in 1926. The maximum saturation is not un¬
usual, for Birge and Juday (1911) report a saturation for
Knight's Lake, Wisconsin of 364.5% and others have found
similar conditions to exist at times.
Carbon dioxide. With regard to the carbon dioxide, the
Wauwatosa pool is again found to be comparable to some
lakes. With the spring overturn the amount of fixed car¬
bon dioxide was practically the same from surface to bot¬
tom. As the stratification of the water progressed, the
amount of fixed C02 increased in the bottom water because
of the accumulation of free C02 which permits a greater
degree of solution of the carbonates. In August and Sep¬
tember, 1926 the bottom water contained 85 and 87 p.p.m.
of fixed C02, respectively, while the surface water at this
time had 57 and 52 p.p.m., respectively. (Fig. 3) With
the dumping of clay in the latter part of September the free
C02 disappeared from the bottom water and the amount of
fixed carbon dioxide decreased as a result. With the fall
overturn, the water again became uniform from surface to
bottom with respect to fixed C02. During the winter the
bottom water contained a larger amount of fixed C02, but
the increase was not more than 2 p.p.m.; before the ice
had left the pool a decrease in the amount of fixed C02 at
the bottom was found, so that there was no real gain dur¬
ing the winter. The difference between the surface and
bottom during the winter was very slight.
The free C02 followed in a general way the course of the
fixed C02. This was particularly true for the bottom
water, when in August 1926 there were 30 p.p.m. of free
C02, which was the maximum for the year. This cor¬
responded with the maximum of 85 p.p.m. of fixed C02.
The amount of free C02 in the surface water gradually di¬
minished until neutrality to phenolphthalein was reached
in September. After clay was emptied into the pool, the
surface water contained no free C02 except in December
1927 and March 1928 when 5.5 and 4 p.p.m., respectively,
were present.
The North Milwaukee pool as a whole contained more
fixed C02 than the Wauwatosa one. The maximum was
37 6 Wisconsin Academy of Sciences , Arts, and Letters.
reached on December 11, 1927 with 94 p.p.m. of fixed CO*.
The variations throughout the year were also much greater
in extent, reaching a minimum of 40 p.p.m. in October
1927. Determinations of the fixed C02 in the bottom water
of the North Milwaukee pool were not made during the
summer. In October 1927 when they were made, the
amount was the same for the surface and bottom, namely
85 p.p.m. During the summer of 1927 there was no free
CO*, in the surface water. This was probably due to its
utilization by algae during the process of photosynthesis.
Hydrogen ion. The study of the hydrogen ion concen¬
tration has played an important part in ecology. Shelford
(1923) and Jewell and Brown (1924) studied the reactions
of fish to varying concentrations of the hydrogen ion.
Juday, Wilson, and Fred (1924) studied the hydrogen ion
concentration in various lakes deep enough to permit strat¬
ification. The lower levels were found to have a lower pH
than the upper, due to decomposiiton in the former and
photosynthesis in the latter. Noland (1925) found that
ciliates exhibit a tolerance to rather a wide range of pH.
He also found decided diurnal variation due to photosynthe¬
sis during the daytime. Philip (1927) also found a wide
diurnal variation in a Minnesota lake and a horizontal vari¬
ation depending on the proximity to clumps of plants and on
the distance from the littoral region. Reed and Klugh
(1924) found a variation of pH from 7.6 to 9.2 in a lime¬
stone pool in October and a biota distinct from that of a
neighboring granite pool with a variation of pH from 6.2 to
6.8.
In the Wauwatosa pool the pH of the surface water var¬
ied from 7.8 to 8.8, being highest in September 1926, and
lowest in March 1927 under the ice. In March 1928 the
same value was reached with 4 p.p.m. of free CO*, present,
although for the previous year there was no free CO*, in
March. The bottom water varied between pH 7.2 and 8.6.
The low value was found in September 1926 when there
were 20 to 25 p.p.m. of free C02 and no free O* present.
(Fig. 3) . The high value came in 1926 just after the over¬
turn on November 30. After the ice was formed the pH of
the bottom water became lower, but did not vary more than
pH 0.2 from that of the surface water. During 1927 the
Wimmer — Study of Two Limestone Quarry Fools. 377
lower water did not reach as low a pH value as it had in
1926 because clay was dumped in as stated above.
The surface of the North Milwaukee pool varied between
pH 7.6 and 8.9; the greatest variations were found in the
spring and fall. The maximum was found on October 30,
1926 and the minimum on March 30, 1927. The lower
water, except during the spring and fall overturn, had a
lower pH value than the surface and varied between 7.4
and 7.8.
Chloride . The Wauwatosa pool had uniformly about
20 p.p.m. of chloride during the summer of 1927. The
determination of the chlorides was started in February
1927. The high values in the larger pool may be attributed
in part to the chlorides brought in with the clay. (This
may also be true for the soluble silica, next to be discussed.)
During the winter the chlorides rose to 21 p.p.m. Just
before the ice melted the chlorides in the surface water de¬
creased. In March 1927 and 1928 there was a decrease
of 7 to 8 p.p.m. This seems to be due to the melting of
the ice and a consequent dilution of the water under the
ice. A sample of the January ice on melting was found to
contain only 25 per cent of the amount of chloride found
in the water just beneath it. The bottom water had a
lower chloride content than the surface during the summer,
but with the fall overturn it approached the same concen¬
tration. Throughout the winter it remained lower than
the surface except in February 1927 when it was 4 p.p.m.
higher than the surface.
The North Milwaukee pool did not show such a high chlo¬
ride content, averaging about 12 p.p.m. However, dur¬
ing the summer, it exhibited a greater variation; in July
1927 it reached a maximum of 17 p.p.m. The two in¬
creases in the amount of chlorides in July and September
were perhaps due to additions as a result of drainage after
heavy rains ; or they may have been due to a concentration
as a result of evaporation, for the water level fell about
27.5 cm. during the summer, and only a small part of this
reduction in level was probably due to seepage through seams
in the rock. The minimum, as is the case for the large
pool, appeared in March, both in 1927 and 1928, just be¬
fore the ice melted. In March 1927 there was a decrease
25
378 Wisconsin Academy of Sciences, Arts, and Letters.
of 4 p.p.m. from that of the February value and in 1928
a decrease of 8.5 p.p.m. from the February reading with
the minimum value of 6.5 p.p.m. The bottom water
showed a slightly higher chloride content during the latter
part of the winter and during the summer; after the fall
overturn, both surface and bottom had the same concen¬
tration.
Silica. Birge and Juday (1911) found a periodicity in
the silica content of the water in Wisconsin lakes, corre¬
lated with the seasonal variations in diatom numbers.
They found that the silica content of the upper waters de¬
creased during the summer due to its utilization by dia¬
toms. The sinking of the dead diatom shells to the lower
water and their subsequent decomposition increased the
silica content of the lower water during the summer.
With the fall overturn the dissolved silica was uniformly
distributed from surface to bottom, thus furnishing favor¬
able conditions for the multiplication of diatoms. Pear¬
sall (1923) found that the increased diatom growth was
associated with heavy rains in spring and fall which
brought dissolved silica and nitrates into bodies of water
by drainage.
The amount of dissolved silica in the surface water of
the Wauwatosa pool varied from minima of 2.8 p.p.m.
in March 1928 and 3.0 p.p.m. in October 1927, to a maxi¬
mum of 7.5 p.p.m. in February 1928. The bottom water
was about 0.5 to 1.5 p.p.m. higher than the surface in sil¬
ica content before the overturn, but after the overturn in
November it had less than the surface. During the winter
the silica accumulated in the bottom water until it had
about 0.5 p.p.m. more than the surface water. The amount
of soluble silica fell in March in a manner similar to the
fall in chloride content described above, and may have been
due to the same cause.
The amount of silica in the North Milwaukee pool was
less than that in the large pool. The surface water was
low in soluble silica during the summer, with about 0.7
p.p.m. Both the surface and bottom increased in silica
content after the fall rains. The maximum for the surface
was reached in November 1927 with 5 p.p.m. During the
winter the silica increased in both the surface and bottom
X
Wimmer — Study of Two Limestone Quarry Pools . 379
waters. The maximum for the bottom was reached in
March 1928, with 5.5 p.p.m.
Nitrogen . No definite correlation could be found be¬
tween the various forms of nitrogen in the pools. In gen¬
eral the various forms of nitrogen tend to run parallel to
each other. In the Wauwatosa pool the maximum organic
nitrogen was found on October 31, 1927 with 3 p.p.m. A
minimum of 0.35 p.p.m. preceded this. The maximum is
correlated with a maximum crop of plankton, but the mini¬
mum is not so correlated with a minimum crop of plankton.
The higher organic nitrogen values seem to be coincident
with high counts of entomostraca. The high point of the
nitrate curve was the same for both pools and fell on Oc¬
tober 31, 1927. It is not due to nitrates brought in by
rains for there was no rain for the fortnight preceding this
date. The nitrates were low in the summer in the Wauwa¬
tosa pool, due to the demand made upon them by the phyto¬
plankton. The ammonia nitrogen was also low during the
summer. During the winter the bottom water of the Wau¬
watosa pool contained from 0.13 to 0.03 p.p.m. of ammonia
nitrogen less than the surface, although soon after the over¬
turn in November it had 0.5 p.p.m. more.
A similar condition was found for nitrogen in the North
Milwaukee pool. The variations between the surface and
bottom were not as great as they were in the Wauwatosa
pool. The nitrates were much lower than any of the other
forms of nitrogen, and agree with the findings of Domo-
galla and others for Wisconsin lakes. The nitrites in the
Wauwatosa pool varied between 0.0015 and 0.0095 p.p.m.
while those for the smaller pool varied between a trace and
0.0195 p.p.m.
Phosphorus. There is a discussion in regard to the role
of phosphorus in the economy of plankton growth. The
studies of Atkins (1923, ’24, ’26) and Atkins and Harris
(1925) introduced the theory that phosphorus is a limiting
factor in the growth of phytoplankton. The whole ques¬
tion is discussed by Juday et al (1927) in a study of lakes
of northeastern Wisconsin. They find no correlation be¬
tween the soluble phosphorus and the phytoplankton, nor
do they find any correlation between the amount of centri¬
fuge plankton and the organic phosphorus.
!$>£6 I9ST 196S
380 Wisconsin Academy of Sciences, Arts, and Letters ,
Fig. 4. Organic matter (curve marked OM) and ash (A) of centrifuge plankton of Wauwatosa pool, indicated in milli¬
grams per liter of water.
Wimmer — Study of Two Limestone Quarry Pools. 381
A similar condition appears to maintain for these pools.
The maxima of the soluble phosphorus came in September
and February in the large pool when the soluble phosphorus
reached 0.12 and 0.14 p.p.m. respectively. The average
was generally about 0.03 p.p.m. The bottom water held
less soluble phosphorus than the surface and there was no
accumulation of it at the bottom during the winter. The
organic phosphorus was not correlated with the net plank¬
ton or with the centrifuge plankton counts.
The soluble phosphorus in the North Milwaukee pool was
lower than that of the Wauwatosa pool. Its average was
about 0.003 p.p.m. and the highest points were found in
September and January when 0.08 and 0.06 p.p.m., respec¬
tively, were reached. Here, again, except for the high
peak in the curve in March 1928, there was no correlation
between the organic phosphorus and the plankton. In
March 1928 there was a coincident rise in the nannoplank-
ton.
Organic matter . A study of the volatile or organic mat¬
ter obtained by ashing the centrifuge plankton shows this
to correspond very well with the net plankton counts as
well as with the counts of the centrifuge plankton. (Fig.
4) The Wauwatosa pool had a maximum of organic mat¬
ter in February 1927 with 3.32 mg. per liter. February
1928 showed 2.43 mg. per liter of organic matter present.
The minima came in April and July, 1927 with slightly
more than 1.1 mg. of organic matter per liter. The influ¬
ence of diatoms as the dominant portion of the plankton
crop is seen by the increase in ash at such times.
The North Milwaukee pool, except for July 1927 with a
minimum of 0.75 mg. of organic matter per liter, showed
higher values than the large pool. This was due to the
greater abundance of plankton in the small pool. Here, too,
the organic matter followed the trend of the curve for the
plankton. The curve also shows the two annual maxima,
the one in spring, and the other in the fall. (Fig. 5).
Plankton
In addition to the physical and chemical factors, there
are the complex interrelations between the various inhabi¬
tants of the pools. The Wauwatosa pool had a deficiency
382 Wisconsin Academy of Sciences , Arts , and Letters .
and at times a complete lack of oxygen in the lower levels,
while the North Milwaukee pool was well supplied
throughout its depth at all times. Atkins and Harris
(1924) found that bodies of water with the bottom waters
low in oxygen, have diatoms and Peridinieae, while those
with a plentiful supply at all times support Chlorophyceae
and Protoccales, primarily. Pearsall (1921, 1922) found
that, in bodies of water where the value of the ratio Na
Fig. 5. Organic matter (curve marked OM) and ash (A) of cen¬
trifuge plankton of North Milwaukee pool, indicated in milligrams
per liter of water.
and K to Ca and Mg was high, a desmid flora predominated,
while a diatom flora predominated in bodies of water with
a low ratio and where the silica and nitrates were present
in sufficient quantities.
The Wauwatosa pool had low oxygen in the bottom water
and had a predominant diatom and dinoflagellate plankton.
Both pools had a large amount of Ca and Mg, consequently
a low value for the Na and K ratio, and the diatoms predom¬
inated at certain seasons of the year. In the North Mil¬
waukee pool, with oxygen at all levels at all times, the Chlo¬
rophyceae predominated during the summer.
384 Wisconsin Academy of Sciences , Arts, and Letters.
In the Wauwatosa pool the diatoms were abundant in
spring and fall, with the latter as the greater of the max¬
ima. (Fig. 6). The growth of diatoms begins in the late
summer and the maximum is reached in October or No¬
vember. Two genera are dominant, one succeeding the
other. Fragilaria appeared first and was followed by As-
terionella in greater numbers, which continued to be domi¬
nant for varying periods, often into March. The maxi¬
mum of diatoms was reached on October 31, 1927 with
1,000,000 colonies of Asterionella per liter in the net plank¬
ton. In the early months of 1926 Eudorina became the
dmoinant plankton organism, and continued so through
April, reaching a maximum of 250,000 colonies in March.
This was its only appearance in numbers in the pool during
the time it was studied. After the decrease in the number
of Eudorina, Sphaerocystis began to appear and increased
in numbers, overshadowing the diatoms in the spring.
During the latter part of summer Oocystis took the place
of Sphaerocystis and reached its maximum number in Sep¬
tember with 40,000 colonies and individuals per liter. In
October this organism was surpassed in numbers by dia¬
toms again. In 1926, Ceratium, which is dominant in
early fall, did not reach as great numbers as it did in 1927.
During the summer of 1927 there was a very meager plank¬
ton population. In the latter part of the summer the
dinoflagellates became the dominant organisms, with two
pronounced peaks in their numbers. The first peak came
in August with 70,000 cells per liter and the maximum came
in October, just before encystment, with 100,000 individu¬
als, chiefly Ceratium. Peridinium numbered a few hundred
cells per liter. After the cool weather of the second week of
October, Ceratium decreased in numbers and began to en¬
cyst. On October 31, 1927 the cysts numbered 1,000 per
liter as compared to 30,000 encysting individuals.
At this time, also, several cases of abnormalities in the
number of horns or spines of Ceratium were found, simi¬
lar to those described by Huber-Pestalozzi (1927). These
abnormalities consisted of unusual twistings of the spines,
especially one or more of the spines of the hypovalve. In
about ten cases, seen in the course of the plankton counts,
a doubling of one or more of the hypovalve spines was ob-
Wimmer — Study of Two Limestone Quarry Pools.
385
386 Wisconsin Academy of Sciences, Arts, and Letters.
served, similar to that shown in fig. II (No. 8) of Huber-
Pestalozzi’s paper. In one case each of the three hypovalve
spines was duplicated and in another, two extra spines
were present. In each case the spines were well developed
and not mere stumps.
Among the less abundant forms, the blue greens reached
their highest numbers in October, May and June. In the
fall, the common ones were Oscillatoria and Anabaena,
while in the spring and early summer Chroococcus was
most common.
The rotifers were abundant in the late summer. At
this time Polyarthra was the most common rotifer. In
1926 the maximum occurred in January, with 1,000 per
liter consisting of almost equal numbers of Polyarthra and
Anuraea. In 1927 the maximum came in August, with
1,600 per liter, the most common being Polyarthra. (Fig.
7).
The entomostraca were found in greatest numbers in the
fall, although there were a few exceptions to this; small
increases were found during the summer. Since the sea¬
sonal variations were studied primarily in the upper meter
of water, where the entomostraca are rarely found in great
numbers, the curve for them may be somewhat misleading.
The greater part of the entomostraca in the upper meter
of water consisted of nauplii. There seemed to be no defi¬
nite cycle in the numbers of the nauplii. This is also the
condition found by Birge and Juday (1922). Daphnia,
Cyclops, and Diaptomus were the most common of the en¬
tomostraca. These forms show a diurnal migration. (Ju¬
day 1904). In samples of water collected during the day,
Daphnia and Diaptomus were most abundant between two
and four meters, where at times 50 Daphnias and over 100
Diaptomi per liter were present. On July 15, 1927 samples
were taken at the surface at 9:00, 9:30, 10.00, and 11:00
p. m. An almost full moon rose at 10:00 p. m. At 9:30
the surface water yielded the greatest number of entomos¬
traca — about 200 Diaptomus and 75 Cyclops. Daphnia
were present in very small numbers — 1 adult and 11 young
individuals. The following noon another sample was taken
and no entomostraca, other than several nauplii were pres¬
ent. (Fig. 7).
Wimmer— Study of Two Limestone Quarry Pools. 387
388 Wisconsin Academy of Sciences, Arts , and Letters .
The North Milwaukee pool shows a much greater wealth
of organisms. The number varies between 40,000 and
over 29,000,000 per liter in the net plankton. The maxi¬
mum growth of plankton came in the spring instead of in
the fall as was the case for the Wauwatosa pool. The dom¬
inant forms were different from those in the large pool.
The diatoms were not the dominant plankton forms, ex¬
cept early in the spring, when Synedra and Diatoma were
most abundant for a short time. In the late summer and
fall the blue greens (Aphanizomenon and Anabaena) were
the most common organisms. In the fall of 1926, Synura
was next to the blue greens in numbers, reaching a total of
500,000 colonies per liter in October. Synura did not ap¬
pear in the fall of 1927, leaving the diatoms second in num¬
ber of individuals. (Fig. 8).
Toward the early part of the winter of 1926 Synura gave
way to a crop of Dinobryon of which, in January 1927,
there were 50,000 colonies of about 20 cells each. The
number of Dinobryon then decreased, but in May rose
again, and then were completely overshadowed by a heavy
crop of monads which in the net plankton numbered 29,-
000,000 per liter.
The dinoflagellates were not as conspicuous in the North
Milwaukee pool because other forms were more numerous.
The greatest number was found in July 1927 when 65,000
Ceratium and 2,000 Peridinium per liter of water were
present. The Ceratium in this pool was larger and
broader, with heavier spines than that in the Wauwatosa
pool. This form was followed in August by a slender and
smaller type, often with only two spines on the hypovalve.
The protozoa were more abundant in the smaller pool
than in the larger one. In October 1926 the protozoa
reached the highest point, with over 60,000 cells per liter.
The greatest number of them consisted of Codonella, al¬
though free swimming Vorticellas of several species num¬
bered over 2,000 per liter. In 1927 the crop of protozoa
was not as large, amounting to 2,000 per liter. The proto¬
zoa were most numerous in October and March with a
slight rise in numbers also in June.
The rotifers also were more numerous than in the Wau¬
watosa pool, reaching a maximum of over 50,000 per liter
19£6 _ mi _
Wimmer — Study of Two Limestone Quarry Pools .
389
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in the North Milwaukee pool.
390 Wisconsin Academy of Sciences , Arts, and Letters.
on March 27, 1927. This is about thirty times the maxi¬
mum number found in the large pool. This maximum con¬
sisted mainly of Polyarthra (22,000), Anuraea aculeaea
(20,000), A. cochlearis (6,000) and Triarthra (2,000). A
second peak in the rotifers was found in July, when they
numbered 25,000 per liter with A. cochlearis as the com¬
mon form. In October 1927 a third peak was found when
they reached 14,000 per liter with A. cochlearis var. tecta
most common. (Fig. 9).
The entomostraca were represented by Cyclops, primar¬
ily, and nauplii formed the greatest contribution to their
number. The maxima were found in late fall, late spring
and early summer. The following table will give some
idea of the numbers of entomostraca in the surface water
at the periods of greatest abundance.
Oct. 1926 May 1927 July 1927
Diaptomus _ 50 per liter 600 60
Cyclops _ 700 100 60
Nauplii _ 200 500 1200
Daphnia _ 25 — 35
Bosmina _ 80 100 700
This study of the plankton of the two pools shows that
they are very different with respect to the kinds and the
numbers of organisms they maintain. The increase in
phytoplankton does not seem at all times to be due to the
increase in the intensity of light in the spring, for at times
some phytoplankton organisms reach their maximum when
the light conditions are poor, (Eudorina in the Wauwa¬
tosa pool in March). In the North Milwaukee pool there
is an increase in the number of phytoplankton organisms
in the spring and this appears to be due in part to the in¬
crease in actinic power of the light at this time as sug¬
gested by Lloyd (1926). The greatest growth of diatoms
takes place when the water is between 10° and 15° C. and
at a time when the light is not too intense. (Whipple,
1927). In general, the protozoa occur in greatest numbers
when certain of the phytoplankton forms are most numer¬
ous, and similar relations seem to hold between the rotifers
and entomostraca. Of the Cyanophyceae, Aphanizomenon
is more tolerant of a wide range of temperature than most
Wimmer — Study of Two Limestone Quarry Pools. 391
algae, Whipple (1927). It occurred in greatest numbers
when the temperature of the water was near its highest
point (24°) in August, and persisted throughout the fall
and winter, in December reaching as many as 1,000,000 fila¬
ments per liter under the ice. In fact, the growth of
Aphanizomenon was so heavy that the ice became greenish
and was not harvested because of the discoloration. Whip¬
ple (1927) cites a similar case.
At various times, samples were taken at different sta¬
tions to study the horizontal distribution of the plankton.
Although some small variations were found at times, they
were not of sufficient magnitude to enable one to draw con¬
clusions. With very little wind, the plankton organisms
probably had a fairly uniform horizontal distribution.
Whipple (1927) believes that the horizontal distribution is
quite uniform.
The vertical distribution is, to a certain extent, depend¬
ent upon the wind. Some of the diatoms tend to sink in
quiet water, while a circulation of the water by the wind
will keep them in the upper stratum. Aphanizomenon
tends to float on the surface of the water, but a wind would
carry some down to the lower water. (Whipple 1927).
The vertical distribution in the North Milwaukee pool was
fairly uniform. The greatest growth was found within
the upper meter. The greatest number of organisms
may be found in either the first or the second half-meter of
water. From one meter downward, there was a gradual
diminution in numbers, with the smallest number at the
bottom. In the Wauwatosa pool uniform distribution was
noted only when the temperature was uniform and the
water was in full circulation. When stratification was
pronounced there was an abrupt change in the number of
plankton organisms at different levels. As in the North
Milwaukee pool, the greatest number of organisms was
found in the first meter of water, except for the entomos-
traca as noted above.
The nauplii were most abundant at the surface, while
Diaptomus and Cyclops were most common in the second
half-meter of water. Daphnia were most abundant be¬
tween two and four meters. Of the rest of the forms there
was a gradual decrease toward the bottom. When strati-
392 Wisconsin Academy of Sciences , Arts, and Letters.
fication is present there is no such gradual fall in numbers
from the surface to the bottom. At such times the ento-
mostraca were still found in greatest numbers between two
and five meters, the nauplii and Cyclops at higher levels
than the Daphnia.
Table 1 gives the numbers of the various organisms at
the different levels of the two pools during the four impor¬
tant seasonal periods of the year. The first column in May,
shows the vertical distribution when the temperature is al¬
most the same from surface to bottom, during the period of
the spring overturn in the Wauwatosa pool. The summer
period of stagnation and stratification is seen in the column
for August. The fall period of uniform temperature and
overturn comes in November, while the winter condition is
seen in February.
Summary
1. Physical, chemical and biological observations were
made on two limestone quarry pools in the vicinity of Mil¬
waukee, Wisconsin.
2. The temperature of the surface water of the two pools
followed that of the air in a general way, rising somewhat
more rapidly in the spring and falling more slowly in the
autumn. The Wauwatosa pool was thermally stratified
during the summer. With a maximum depth of only 5 m.,
the North Milwaukee pool was substantially uniform in
temperature from surface to bottom.
3. The North Milwaukee pool contained an abundance
of oxygen at all depths throughout the year ; dissolved oxy¬
gen entirely disappeared from the bottom water of the
other pool in summer and in winter. There was no defi¬
nite correlation between the various forms of nitrogen in
the two pools; in general there was a correlation between
the organic nitrogen and the plankton crop. Nitrates were
low in summer. No correlation was found between the
phosphorus and the plankton.
4. Diatoms and dinoflagellates were dominant in the
Wauwatosa pool, and Chlorophyceae and Cyanophyceae in
the North Milwaukee pool. The latter pool had a greater
number of plankton organisms per liter and also a greater
Wimmer — Study of Two Limestone Quarry Pools. 393
variety of species. The maximum crop of plankton was
found in the spring in the North Milwaukee pool, but it
came in the fall in the Wauwatosa pool. Protozoa and roti¬
fers were more abundant in the North Milwaukee pool than
in the Wauwatosa pool. With the exception of the ento-
mostraca, the largest number of plankton organisms was
found in the upper meter of water in both pools. The ver¬
tical distribution of the summer plankton in the Wauwa¬
tosa pool was similar to that of stratified lakes.
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- 1925. Seasonal changes in the phosphate content of
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26
394 Wisconsin Academy of Sciences, Arts, and Letters.
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Wimmer — Study of Two Limestone Quarry Pools. 395
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Table 1. Summary of the vertical distribution of the various plankton organisms in the two pools. The results
stated in the average number of individuals per liter between the depths given.
Wimmer — Study of Two Limestone Quarry Pools. 397
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398 Wisconsin Academy of Sciences, Arts, and Letters,
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Table 1. Summary of the vertical distribution of the various plankton organisms in the two pools. The results are
stated in the average number of individuals per liter between the depths given — Continued.
Wimmer — Study of Two Limestone Quarry Pools. 399
THE MORPHOLOGY AND DEVELOPMENTAL STAGES
OF A NEW SPECIES OF PIONA
Ruth Marshall
The material for this study was found by Dr. J. R. Hick¬
man while collecting bottom forms for the Wisconsin Geo¬
logical and Natural History Survey. The author is also in¬
debted to Dr. E. A. Rirge and Professor Chancey Juday
for permission to examine the water mites collected during
the summer of 1928 at the Trout Lake Laboratories.
On August 4, a large number of nymphs of a Piona were
found on branches of Nitella taken from Lake Kawague-
saga near Minocqua. Some of this material was kept un¬
der observation for the next ten days ; similar material was
later found in other lakes of the region. On the day they
were discovered, the nymphs appeared passive, with legs
outstretched and heads resting on the plant stems. The
body at this stage is rotund, with a yellow dorsal streak;
the epimera are small and well separated. The genital
plates are obliquely set and each bears two round acetabula
(fig. 7). In the water about these nymphs were found
great numbers of newly emerged males and they continued
to be abundant for several days, together with a few speci¬
mens of Hygrobates longipalpis. Very few females were
seen at first ; but three days later they, too, were abundant.
Copulating pairs then appeared and a few first larvae.
Pairing as observed here agrees with accounts given for
other species of this genus and illustrates the use of the
peculiarly modified legs of the male. The female remains
passive with legs outstretched. The male rests on the an¬
terior end of the ventral side of the female with the fourth
legs flexed so that the concavities of the fourth segments
are hooked over the first legs of the female. The club-
shaped distal segments of the third legs, which are usually
seen held together on the genital orifice, now transfer the
semen masses to the genital opening of the female. The
402 Wisconsin Academy of Sciences , Arts, and Letters.
entire process lasts but a few moments, when the pair sep¬
arate.
On August 8, egg masses were observed on the plant
stems (fig. 9). Each mass was about 2.5 mm. in length
and contained from twenty to thirty pale, oblong eggs
placed irregularly in a transparent jelly. Larvae were
emerging until August 14, at which time all of the second
larvae had apparently pupated and young females were
abundant. Material from Little Arbor Vitae Lake col¬
lected August 27 consisted chiefly of nymph cases and
young males. The first, or six-legged larva is a very deli¬
cate, active creature, measuring 0.475 mm. to the tip of
the capitulum (fig. 8). The body is broad, pale brown
with a pale blue area dorsally near the anterior end. The
epimera cover most of the ventral side, as in other recorded
larvae of the genus ; the surface is covered with fine oblong
hexagonal areas. The posterior end of the body bears two
papillae and numerous long hairs. The subsequent history
of these larvae is not known ; in other species they are be¬
lieved to attach themselves to the bodies of small water
beetles soon after hatching.
The newly emerged adults are very delicate and trans¬
parent, with a pale yellow to reddish Y-shaped dorsal mark.
The males are about 0.975 mm. in length; the females of
the same period are at first smaller, but soon exceed them
in size. The epimera now occupy much of the ventral sur¬
face; the second pair have well developed anchoral pro¬
cesses, the fourth pair are very large (fig. 1). The genital
areas are about half enclosed by the deep bay formed by the
concave margins of the last epimera, those of the male
barely joining them in the mid-line. The orifice in the
male is broadly trifoliate and large; in the female there is
a long slit. The genital plates are large, oval and similar
in the two sexes, with about thirty acetabula on each, the
latter variable in size and arrangement, with two on each
plate larger than the others (fig. 3, 4).
The maxillar shield is large and broad. The palpi ex¬
ceed the legs in width and bear few bristles ; the fourth seg¬
ment is slim, with one large and two smaller anteriorly di¬
rected papillae on the flexor surface with still another near
the distal end. The fifth segment is curved and ends in
Marshall — Morphology of New Species of Piom. 403
four finger-like processes (fig. 6). The legs are long and
well provided with bristles and swimming hairs. In the
male the third leg has a club-shaped distal segment with a
claw transformed into a long delicate hook (fig. 4) ; the
fourth segment of the last leg has a conspicuous concavity
bordered by rows of blade-like hairs (fig. 5) .
The species appears to be a new one, probably widely
distributed in this country and will be named Fiona ameri -
cana nov. spec. It bears some resemblance to P . turgida
(Wol.), a larger species with which it was at first confused.
It is closely related to P. coccinea (Koch), which, together
with its varieties, is widely distributed in Europe and re¬
ported also from parts of Asia and Africa. The two spe¬
cies are very similar in the form of the genital orifices and
the third and fourth legs of the male, but differ in details
of structure.
Biological Laboratory,
Rockford College,
Jan. 1, 1929.
Explanation of Plate 9
Piona americana
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Ventral view, male.
Genital area and epimera, female.
Genital area, male.
Sixth segment, third leg, male.
Fourth segment, fourth leg, male.
Left palpus.
Ventral surface, 2nd larva.
Ventral surface, 1st larva.
Egg mass on stem of Nitella.
404 Wisconsin Academy of Sciences , Arts , and Letters .
TRANS. WIS. ACAD., VOL. 24 PLATE 9
A PRELIMINARY LIST OF THE MOLLUSCA OF
DANE COUNTY, WISCONSIN
J. P. E. Morrison
Introduction
The purpose of the following paper is to tabulate the dis¬
tribution of the molluscs of this region, as far as known at
the present time. It is intended as a checklist, with local¬
ity records, of this particular portion of the fauna of the
area surrounding Madison. For convenience, the bounda¬
ries of the county have been taken as the limits of the area.
The Wisconsin River, the northwest boundary of the re¬
gion, has been included in the survey.
In the preparation of the list, material has been taken
from three sources, namely: (1) published lists, includ¬
ing scattered references, where found. (2) the collections
in the Museum of the University of Wisconsin. (3) collec¬
tions made by the writer in the area under consideration.
Acknowledgements are due the following people, who
have aided the work in various ways: Frank C. Baker,
University of Illinois, Mr. Chancey Juday, and Mr. Geo.
Wagner, University of Wisconsin. To Mr. Wm. J. Clench,
Harvard Museum of Comparative Zoology, for determina¬
tion of the Physidae; Dr. Victor Sterki, New Philadelphia,
Ohio, for determination of the Sphaeriidae; Dr. Bryant
Walker, Detroit, Michigan, are due many thanks for their
kind cooperation. Unfortunately, all of the Ancylidae in
the author's personal collections from the vicinity of Madi¬
son have recently been lost in the mails. As they had not
yet been examined by Dr. Walker, the records are unavail¬
able specifically, and not included.
I wish also to thank the many friends who have aided in
the field, by assistance in collecting, especially: Mr. Or¬
lando Park, of the University of Chicago; Mr. Ralph
Bailey, Mr. L. G. Gumbreck, Mr. J. H. Roberts, of the Uni¬
versity of Wisconsin ; Mr. Chatt Himley, of Madison.
406 Wisconsin Academy of Sciences, Arts, and Letters.
In 1823, Mr. D. H. Barnes, of New York, published an
article in the American Journal of Science, describing some
new species of Fresh Water Mussels. The specimens he
described were received by him from Capt. D. B. Douglass,
topographical engineer, and Mr. H. R. Schoolcraft, minera¬
logist, of the N. W. Expedition. The records are the first
from this region.
Mr. I. A. Lapham, of Milwaukee, was probably the first
Wisconsin man to make careful studies of the molluscan
fauna of the State. In 1852, he published a catalogue of
the fauna, both recent and fossil, and of the flora known
at that time from Yvisconsin. Included in the list were
some ninety forms of mollusks. Again in 1860, he pub¬
lished a list of the shells of the State. This second list,
published in an eastern journal, was an abridged copy of
the first one, and did not contribute more to our knowledge
of the fauna. However, Mr. LaphanTs original list is an
invaluable record of its kind.
Likewise among the early records from Dane County
are those of Prof. Spencer F. Baird, who was the first cura¬
tor of the Smithsonian Institution. In the summer of
1853, he visited Dr. P. R. Hoy, of Racine, doing quite a bit
of natural history work with him during the summer. On
July second, they set out, together with Dr. J. P. Kirtland,
on a longer excursion through southern Wisconsin. This
trip included a stop at Madison, where most of their collect¬
ing was done in the neighborhood of Gov. Farwell’s Mills,
located at the foot of Lake Mendota, between that lake and
Lake Monona. Baird’s records were published in Binney’s
monographs of 1865.
Between the years 1887 and 1899, Geo. Marston collected
extensively in eastern Wisconsin, including many collec¬
tions from the Wisconsin River.
In 1897, Mrs. E. C. Wiswall published a list of shells of
Southern Wisconsin. Some few of these records are refer¬
able to Dane County.
In the first decade of the present century, notably in
1904, Mr. Geo. Wagner collected from the lakes around
Madison, principally from Lake Mendota.
Incidental to reporting the occurrence of the element man¬
ganese in Fresh Water Mussels, Bradley (1910) mentions
Morrison — Preliminary List of the Mollusca of Dane County. 407
the two common mussels of the lakes near Madison. He
reports them as Anodonta — “the typical lake form,” and
Unio — (evidently Lampsilis, from the locality mentioned).
The writer has taken the liberty of referring these records
to the species to which they undoubtedly belong.
The records added by W. H. Dudley, who collected on the
Wisconsin River, etc., during the summer of 1918, have
been included wherever possible.
Muttkowski, working on Lake Mendota during the period
between 1913 and 1916, published on the fauna of the lake
(1918). He lists several species that were studied quanti¬
tatively, and mentions the fact that there were others in
the lake.
Perhaps the records based on the collections of D. S. Bul¬
lock, now in the Museum of the University, have added
most among previous ones to our knowledge of the kinds of
molluscs in the county.
Mr. Frank C. Baker, working in connection with the Ge¬
ological and Natural History Survey, made extensive col¬
lections in the State during the summers of 1920-22.
Some of these records are included, where they support re¬
cords from localities actually within Dane County. Not a
small number of records have been taken directly from
Baker's monograph (Published in December 1928) but
only those have been included that seem unquestionably
referable to the area under consideration.
Juday (1922) records Pisidium idahoense from the
deeper waters of Lake Mendota. It should perhaps be
mentioned here that in these studies he carried on, he was
dealing with not only the species mentioned, but also with
at least three others in addition (in all probability). In
his report, all of the pisidia mentioned as of the large class
(having a length of 7 mm. or more) were undoubtedly
idahoense, together with some young specimens listed in
the other classes. On the other hand, the great majority
of the pisidia mentioned (3,343 recorded as small; that is,
as having a length of from 1.5 to 3.5 mm.) were undoubt¬
edly not the above species, but of an undescribed species, in
large part. This statement is based on subsequent collec¬
tions by the writer. Since there is more than one species
in the deeper waters of Lake Mendota, Juday’s remarks as
408 Wisconsin Academy of Sciences , Arts , and Letters.
to the high rate of mortality of the pisidia have become un¬
tenable. In substitution, it may be stated that P. idaho-
ense is much less common in Lake Mendota than this unde¬
scribed species, which is usually no longer than 4 mm.
Finally, during the periods between September 1927 and
June 1928, and between September and December 1928, the
writer has made collections of molluscs from as many lo¬
calities and over as great a seasonal range as has been
possible in the limited time at his disposal. Besides adding
a few species to those previously known from the area,
these records have confirmed, in large part, those of the
other workers. There are, in the author's collections from
the region, some lots of species as yet undescribed. It has
been thought best to omit them entirely from the list, in
order to avoid any possibility of confusion.
For a full list of bibliographical references, the reader
is referred to Baker's Monograph (1928, b.). The follow¬
ing few references will serve as an introduction to the mol¬
luscs in the county, including, as they do, the published re¬
cords from this area.
Baker, F. C. 1902. The Mollusca of the Chicago area,
part 2, The Gastropoda. Bull. 3, part 2, Nat. Hist.
Sur., Chicago Acad. Sci.
- 1928.b. The fresh water Mollusca of Wisconsin.
part 1, Gastropoda. (Bull. 70. part 1. Wis. Geol. &
Nat. Hist. Sur.). Mon. Aquatic Gastropoda of Wis¬
consin. Wis. Acad. Sci. Art. Let.
— — 1928.b. The fresh water Mollusca of Wisconsin,
part 2, Pelecypoda. (Bull. 70. part 2. Wis. Geol. &
Nat. Hist. Sur.). Bull. U. of Wis. serial No. 1527,
general series No. 1301.
Binney, W. G., and Bland, T. 1869. Land and fresh
water shells of North America, part 1, Pulmonata Geo-
phila. Smith. Misc. Coll. No. 194.
Binney, W. G. lS65.a. Land and fresh water shells of
North America, part 2, Pulmonata, Limnophila and
Thalassophila. Smith Misc. Coll. No. 143.
— 1865.b. Land and fresh water shells of North
America, part 3, Ampullariidae, Vaivatidae, etc. Smith
Misc. Coll. No. 144.
Morrison — Preliminary List of the Mollusca of Dane County. 409
Bradley, H. C. 1910. Manganese of the tissues of lower
animals. Jour. Biol. Chem. 8:237-249.
Chadwick, G. H. 1906. Notes on Wisconsin Mollusca.
Bull. Wis. Nat. Hist. Soc. 4:67-99.
Juday, C. 1922. Quantitative studies of the bottom
fauna in the deeper waters of Lake Mendota. Trans.
Wis. Acad. Sci. Art. Let. 20 :461-493.
Lapham, I. A. 1852. Catalogue of the Mollusca of Wis¬
consin. Trans. Wis. State Agric. Soc. 2:367-370.
— — 1860. A list of the shells of the State of Wisconsin.
Proc. Phila. Acad. Sci. 1860:154-156.
Muttkowski, R. A. 1918. The fauna of Lake Mendota.
Trans. Wis. Acad. Sci. Art. Let. 19:174-482.
Pilsbry, H. A. and Johnson, C. W. 1898. A classified cat-
alouge, with localities, of the land Mollusca of North
America, north of Mexico, (reprinted from the Nau¬
tilus, August 1897— April 1898.).
Wiswall, (Mrs.) E. C. 1897. Shells of southern Wiscon¬
sin. Nat. Sci. Journ. 1 (2) :47-48.
The system of classification followed in this paper is that
used by Baker in his recent monograph, with additions.
That used for the land species is modified from the cata¬
logue of Pilsbry and Johnson cited above. It is not to be
presumed that the writer's arrangement of the groups of
land forms is anything more than a convenience.
In all, one hundred and fifty species and varieties are re¬
corded from Dane County. The following short list will
give an idea of the distribution of these species into the
major groups.
Fresh water univalves . . . . . „_53
Land univalves _ _ _ _ _ -25
Unionidae (bivalves) ... _ _ — _ _ .__42
Sphaeriidae (bivalves) _ _ _ _ -30
Total _„______-l 50 (species and varieties)
Of interest is the fact that four species are herein added
to those already known from the State. They are : Pisid -
turn concinnulum Sterki ; P. minutum Sterki ; JP. pusillum
Jen. ; Radix auricularia (Linn.) . The occurrence of Gy-
raulus crista (Linn.) ., in abundance, living, in the swamps
27
410 Wisconsin Academy of Sciences, Arts, and Letters .
near Lakes Mendota and Wingra is worthy of mention.
Also notable is the recent discovery of Acella haldemani
(Desh; Binney) as a fossil in the marl at the margin of
University Bay, Lake Mendota. The only other record of
this species' occurrence in the State is that of Lapham, who
recorded it from the Milwaukee River in 1852.
Undoubtedly further collecting in the area will add other
forms to those included in the present list. This is espe¬
cially true since so little of the county has been intensively
examined for Mollusca.
It is to be hoped that this list will in interest many ; those
interested in molluscs from a popular viewpoint, as well as
those interested from more technical viewpoints.
Any additions and corrections will be gratefully received.
The systematic list by species follows. Each name of a
species or variety is brought out to the left-hand margin
for ready reference; the localities where the species is
known to occur are stated; the names in parentheses fol¬
lowing the localities indicate the authority for the records
at the places cited.
SYSTEMATIC CATALOGUE OF SPECIES
Class GASTROPODA
Subclass STREPTONEURA Spengel.
Order CTENOBRANCHIATA Schweigger.
Suborder PLATYPODA Lamarck.
Superfamily Taeniglossa Bouvier.
Family VALVATIDAE Gray.
Genus Valvata Muller.
Valvata tricarinata (Say).
Four Lakes (Lapham) ; Lake Mendota (Muttkowski, Baker) ;
margin, University Bay, Lake Mendota; dredging, seven meters,
Lake Mendota; Merrill Spring stream; Pheasant Branch, near
mouth (Morrison).
Valvata sincera Say.
Four Lakes (Lapham) ; Madison (Lapham, Binney, Chadwick,
Baker).
Family VIVIPARIDAE (Gray) Gill.
Subfamily LIOPLACINAE (Gill) Baker,
Genus Campeloma Rafinesque.
Morrison — Preliminary List of the Mollusca of Dane County. 411
Campeloma integrum (Say).
Wisconsin River, “ascends as far as the dam at Kilbourn”
(Baker).
Campeloma rufum (Haldeman).
Yahara River, Madison (Pearse, Morrison) ; sandy shallows,
Lake Mendota; Black Earth Creek, Mazomanie; Koshkonong
River, Rockdale; outlet of Lake Ripley, Cambridge (Morrison).
Family AMNICOLIDAE (Tryon) Gill.
Subfamily AMNICOLINAE Gill.
Genus Amnicola Gould & Haldeman.
Amnicola limosa porata (Say).
Madison, (Lapham, Binney, Chadwick) ; Lake Mendota (Mutt-
kowski) ; University Bay, Lake Mendota; dredging off Picnic
Point, Lake Mendota (Morrison).
Amnicola limosa parva Lea.
Lake Mendota (Walker, Baker) ; Lake Mendota, dredging, seven
meters (Morrison).
Amnicola lustrica decepta Baker.
Lake Mendota (Hinkley, Baker) ; Lake Wingra (Bullock,
Baker) .
Amnicola walkeri Pilsbry.
Lake Wingra (Bullock, Baker).
Genus Cvncinnatia Pilsbry.
Cincinnatia cincinnatiensis (Anthony).
Yahara River, Madison; stream, four miles east of Belleville
(Morrison) .
Subfamily LITHOGLYPHINAE Fisher.
Genus Somatogyrus Gill.
Somato gyrus depressus (Tryon).
Prairie Du Sac (Hinkley, Walker, Baker) : Probably occurs on
the Dane Co. side of the river.
Somatogyrus tryoni Pilsbry & F. C. Baker.
Black Earth Creek, Mazomanie; Koshkonong River, Rockdale;
outlet of Lake Ripley, Cambridge (Morrison).
Family POMATIOPSIDAE Stimpson.
Genus Pomcrtiopsw Tryon.
Pomatiopsis lapidaria (Say).
Four Lakes, Madison (Lapham, Binney, Chadwick, Baker).
Family PLEUROCERIDAE Fisher.
Genus Pleurocera Rafinesque.
412 Wisconsin Academy of Sciences , Arts , and Letters .
Pleurocera acuta tracta (Anthony) .
Lake Mendota (Muttkowski) ; Yahara River, Madison (Juday,
Morrison) ; sandy shallows, Lake Mendota; Koshkonong River,
Rockdale; outlet of Lake Ripley, Cambridge; Black Earth Creek,
Mazomanie (Morrison).
Collections upstream at Kilbourn (Baker), and downstream at
Arena (Morrison), indicate its presence in the Wisconsin River
within the limits of Dane County.
Subclass EUTHYNEURA Spengel.
Order PULMONATA Cuvier.
Suborder BASOMMATOPHORA A. Schmidt.
Superfamily LIMNOPHILA.
Family LYMNAEIDAE (Broderip) Baker.
Genus Lymnaea Lamarck.
Lymnaea stagnalis jugidaris Say.
Fourth Lake (Lapham) ; Lake Mendota (Muttkowski, Baker).
Genus Stagnicola (Leach) Jeffreys.
Stagnicola palustris elodes (Say).
The Four Lakes (Lapham) ; Lake Mendota, Madison (Bullock,
Baker) ; ponds, streams, and swamp, south of Lake Wingra;
swamp, University Creek, University Bay, Lake Mendota (Mor¬
rison).
Stagnicola umbrosa (Say).
Lake Mendota (Bullock, Juday, Baker) ; near Madison (Wis-
wall, Baker); Windsor (Bullock, Baker).
Stagnicola umbrosa jolietensis (F. C. Baker).
Shore of Lake Mendota, Madison (Bullock, Wagner, Baker).
Stagnicola reflexa (Say).
Farwell’s Mills, Madison (Baird, Binney, Chadwick, Baker).
Stagnicola emarginata angulata (Sowerby).
Four Lakes (Lapham) ; Madison (Lapham, Binney, Marston,
Wiswall, Chadwick, Bullock, Baker) ; below old Chem. Lab.,
Lake Mendota (Wagner).
Stagnicola caperata (Say).
Pond, stream, south of Lake Wingra (Morrison).
Genus Acella Haldeman.
Acella haldemani (“Deshayes” Binney).
Fossil in Marl: Margin of University Bay, Lake Mendota (Mor¬
rison). Further search may reveal this species living in Lake
Mendota.
Genus Radix Montfort.
Morrison — Preliminary List of the Mollnsca of Dane County. 413
Radix auricularia (Linn.)*
A small but thriving colony of this European species has been
found in the aquarium of the Botany Dept. Greenhouse, at the
University of Wisconsin. The snails were accidentally intro¬
duced; probably on Elodea plants, from either Cincinnati
(Bryan), or Philadelphia (Denniston).
Genus Fossaria Westerlund.
Fossaria parva (Lea) .
Swamp, streams, south of Lake Wingra; Merrill Creek; swamp,
four miles east of Belleville (Morrison).
Fossaria modicella (Say).
Stream, pond, south of Lake Wingra; Pumping Station outlet,
University Bay, Lake Mendota (Morrison).
Fossaria obrussa (Say).
University Bay, Lake Mendota (Pearse) ; stream, swamp, ponds,
south of Lake Wingra; Pheasant Branch, near mouth (Morri¬
son).
Fossaria obrussa decampi (Streng).
Fossil: Lake Wingra near Madison (Bullock, Baker).
Fossaria exigua (Lea).
Streams, south of Lake Wingra (Morrison).
Family PLANORBIDAE H. & A. Adams.
Genus Helisoma Swainson.
Helisoma antrosa (Conrad).
Four Lakes (Lapham); University Bay, Tenney Park shores,
Lake Mendota, Madison (Morrison).
Helisoma antrosa unicarinata (Haldeman).
Lake Mendota (Muttkowski, Baker).
Helisoma trivolvis (Say).
FarwelPs Mills, Madison (Baird, Binnay) ; Madison (Baird,
Binney, Chadwick, Baker) ; Lake Monona, Lake Mendota, Madi¬
son (Bullock) ; swamp, University Creek; Pheasant Branch, near
mouth; ponds, south of Lake Wingra; Koshkonong River, Rock¬
dale (Morrison).
Helisoma pseudotrivolvis (F. C. Baker) .
Near Lake Monona (Bullock) ; near Murphy Creek, near Lake
Mendota, near Madison (Bullock, Baker).
Helisoma campanulata (Say).
Fourth Lake (Lapham); Lake Mendota (Muttkowski, Baker).
Genus Planorbula Haldeman.
414 Wisconsin Academy of Sciences, Arts, and Letters.
Planorbula armigera (Say) .
Ponds, stream, springs, swamp, south of Lake Wingra; swamp,
University Creek, University Bay, Lake Mendota (Morrison).
Genus Menetus H. & A. Adams.
Menetus exacuous (Say).
Swamp, south of Lake Wingra; swamp, University Creek, Uni¬
versity Bay, Lake Mendota (Morrison).
Menetus exacuous meg as (Dali).
The record of this variety from Lake Monona (Dane Co.) by
Hinkley is erroneous. The record should be Lake Monona, Min¬
nesota, not Wisconsin, as shown by other specimens in the Hink¬
ley collection (Baker).
Genus Gyraulus Charpentier.
Gyraulus hirsutus (Gould).
Canal, south of Lake Wingra; swamp, University Bay, Lake
Mendota (Morrison).
Gyraulus deflectus (Say).
Fourth Lake (Lapham) ; stream, south of Lake Wingra; Merrill
Spring; Pheasant Branch, near mouth; swamp, University Bay,
Lake Mendota (Morrison). The record from Lake Monona was
probably a lapsus pennae of Hinkley for Lake Monona, Minne¬
sota (Baker).
Gyraulus deflectus obliquus (DeKay).
Near Madison (Bullock, Baker).
Gyraulus parvus (Say).
Lake Mendota (Muttkowski, Bullock, Baker) ; dredging, seven
meters, Lake Mendota; Yahara River, Madison; Merrill Creek;
Merrill Spring stream; ponds, stream, swamp, south of Lake
Wingra; swamp, University Creek, University Bay, Lake Men¬
dota (Morrison).
Gyraulus altissimus (F. C. Baker).
Fossil in marl: near Lake Wingra (Bullock, Baker).
Gyraulus umbilicatellus (Cockerell).
Windsor (Bullock, Baker).
Gryaulus crista (Linn.).
Ponds, swamp, (on water plants) south of Lake Wingra; swamp,
University Creek, University Bay, Lake Mendota (Morrison).
Family ANCYLIDAE Menke.
Subfamily FERRISSINAE Walker.
Genus Ferrissia Walker.
Morrison — Preliminary List of the Mollusca of Dane County. 415
Ferrissia kirJclandi (Walker).
University Bay, Lake Mendota (Juday, Baker).
Family PHYSIDAE Dali.
Genus Phy sella (Haldeman) Baker.
Phy sella ancillaria (Say).
University shores, University Bay, Lake Mendota (Wagner).
Phy sella sayii (Tappan).
Along shore, Lake Mendota, Madison (Wagner) ; swamp, Uni¬
versity Bay, Lake Mendota; Yahara River, Madison; ponds,
south of Lake Wingra (Morrison).
Physella sayii crassa (Walker).
Lakes Monona and Mendota (Baird, Bullock, Muttkowski, Wag¬
ner, Baker).
Physella warreniana (Lea).
Pheasant Branch, near mouth; swamp, University Creek, Uni¬
versity Bay, Lake Mendota; Yahara River, Madison; canal,
streams, south of Lake Wingra (Morrison).
Physella gyrina (Say).
Farwell’s Mills, Madison (Baird, Binney, Chadwick, Baker) ;
stream, south of Lake Wingra (Bullock, Baker) ; ponds, stream,
canal, south of Lake Wingra; swamp, mouth of University
Creek, University Bay, Lake Mendota; Merrill Creek (Morri¬
son) .
Physella gyrina hildrethiana (Lea) .
Sun Prairie (Bullock).
Physella integra (Haldeman).
University shore, Lake Mendota, Madison (Wagner) ; rocky
shore, east of University Bay; shores in Tenney Park, Lake
Mendota; Yahara River, Madison; Koshkonong River, Rockdale;
outlet of Lake Ripley, Cambridge (Morrison).
Physella walkeri (Crandall).
Merrill Springs, Lake Mendota (Pearse, Baker) ; Lake Mendota
(Bullock, Muttkowski, Baker).
Physella michiganensis (Clench).
Merrill Spring (Morrison).
Genus Aplexa Fleming.
416 Wisconsin Academy of Sciences , Arts, and Letters .
Aplexa hypnorum (Linn.).
Windsor (Bullock, Baker) ; swamp, north of Picnic Point, Lake
Mendota, Madison; ponds, swamp, south of Lake Wingra;
swamp, near outlet, Lake Ripley, Cambridge (Morrison).
Superfamily AKTEOPHILA
Family AURICULIDAE
Genus Carychium Muller.
CarycJnium exiguum (Say).
Swampy ground, south of Lake Wingra; along Merrill Spring
stream; swamp, four miles east of Belleville (Morrison).
Suborder STYLOMMATOPHORA
MONOTREMATA
Vasopulmonata
ORTHURETHRA
Family VALLONIIDAE
Genus Vallonia Risso.
Vallonia costata (Muller) .
Sunset Point, three miles west of Madison; swamp, along Uni¬
versity Bay; along Merrill Spring stream; in willows, south of
Lake Wingra (Morrison).
Vallonia pulchella (Muller).
Sunset Point, three miles west of Madison; along Merrill Spring
stream; along Merrill Creek; swampy ground, south of Lake
Wingra; Picnic Point, near University Bay; swamp, four miles
east of Belleville (Morrison).
Family PUPILLIDAE
Genus Strobilops Pilsbry.
Strobilops affinis Pilsbry.
Swampy ground, south and west of Lake Wingra; Sunset Point,
three miles west of Madison; along Merrill Creek; along the
outlet of Lake Ripley, Cambridge (Morrison).
Genus Pupoides Pfeiffer.
Pupoides marginata (Say).
Sunset Point, three miles west of Madison (Morrison).
Genus Gastrocopta
Gastrocopta armifera (Say).
Sunset Point, three miles west of Madison; bluffs, northwest
margin of Lake Mendota (Morrison).
Gastrocopta contracta (Say).
Bluffs, northwest margin of Lake Mendota; along Merrill Creek;
in willows, south of Lake Wingra; Sunset Point, three miles
west of Madison (Morrison).
Morrison — Preliminary List of the Mollusca of Dane County. 417
Gastrocopta tappaniana (C. B. Adams).
Swampy ground, south of Lake Wingra; along Merrill Creek;
swamp, four miles east of Belleville (Morrison).
Gastrocopta pentodon (Say).
Bluffs, northwest margin of Lake Mendota; Sunset Point, three
miles west of Madison (Morrison).
Genus Vertigo Draparnaud.
Vertigo ovata (Say).
Swampy ground, south of Lake Wingra; swamp, four miles east
of Belleville (Morrison).
Vertigo milium (Gould).
Swampy ground, south of Lake Wingra (Morrison).
Family COCHLICOPIDAE
Genus Cochlicopa (Ferussac) Risso.
Cochlicopa lubrica (Muller).
Madison (Lapham) ; near springs, south of Lake Wingra; bluffs,
northwest margin of Lake Mendota (Morrison).
HETERURETHRA
Superfamily ELASMOGNATHA
Family SUCCINEIDAE
Genus Succinea Draparnaud.
Succinea retusa Lea.
Swampy ground, south of Lake Wingra (Morrison).
Succinea avara Say.
Swampy ground, south of Lake Wingra; bluffs, northwest mar¬
gin of Lake Mendota; swamp, four miles east of Belleville (Mor¬
rison).
SIGMURETHRA
Superfamily HOLOPODA
Family HELICIDAE
Subfamily POLYGYRINAE
Genus Polygyra (Say) Pilsbry.
Polygyra monodon (Rackett).
Along stream, south of Lake Wingra (Morrison).
Polygyra monodon fraterna (Say).
Swampy ground, south of Lake Wingra; bluffs, northwest mar¬
gin of Lake Mendota (Morrison).
418 Wisconsin Academy of Sciences, Arts, and Letters .
Polygyra multilineata (Say).
Madison (Lapham) ; swampy ground, south and west of Lake
Wingra (Morrison).
Superfamily AULACOPODA
Family ZONITIDAE
Subfamily ZONITINAE
Genus Vitrea Fitzinger.
Vitrea hammonis (Strom.).
Woods, south of Lake Wingra; Picnic Point, near University
Bay; along Merrill Creek (Morrison).
Vitrea indentata (Say).
Bluffs, northwest margin of Lake Mendota (Morrison).
Genus Euconulus Reinhardt.
Euconulus fulvus (Draparnaud) .
Swampy ground, south of Lake Wingra; Sunset Point, three
miles west of Madison (Morrison).
Subfamily ARIOPHANTINAE
Genus Zonitoides Lehmann.
Zonitoides arbor eus (Say).
Madison (Lapham) ; bluffs, northwest margin of Lake Mendota;
swampy ground, south of Lake Wingra; Picnic Point, near Uni¬
versity Bay; Sunset Point, three miles west of Madison; swamp,
four miles east of Belleville (Morrison).
Family LIMACIDAE
Genus Agriolimax Morch.
Agriolimax campestris (Binney).
Bluffs, northwest margin of Lake Mendota; Picnic Point, near
University Bay, Madison; swampy ground, south of Lake Win¬
gra (Morrison).
Family ENDODONTIDAE
Subfamily ENDODONTINAE
Genus Pyramidula Fitzinger.
Pyramidula alternata (Say).
Sunset Point, three miles west of Madison (Morrison).
Genus Gonyodiscus Fitzinger.
Gonyodiscus cronkhitei anthonyi Pilsbry.
Madison (Lapham); swampy ground, south of Lake Wingra;
along Merrill Creek; along outlet of Lake Ripley, Cambridge
(Morrison).
Genus Helicodiscus Morse.
Helicodiscus parallelus (Say).
Bluffs, northwest margin of Lake Mendota; in willows, south of
Lake Wingra; Sunset Point, three miles west of Madison (Mor¬
rison) .
Morrison — Preliminary List of the Mollusca of Dane County. 419
Class PELECYPODA Goldfuss.
Order PRIONODESMACEA Dali.
Superfamily NAIADACEA Menke.
Family UNIONIDAE (d’Orbigny) Ortmann.
Subfamily UNIONINAE (Swainson) Ortmann.
Genus Fusconaia Simpson.
Fusconaia flava (Rafinesque) .
Black Earth Creek, Mazomanie; Koshkonong River, Cambridge,
Rockdale (Morrison).
Fusconaia flava parvula (Grier)
Outlet of Lake Ripley, Cambridge (Morrison).
Fusconaia undata (Barnes).
Wisconsin River (Schoolcraft, Barnes, Marston, Baker), at
Prairie du Sac (Lapham), opposite Prairie du Sac (Morrison).
Fusconaia ebena (Lea).
Wisconsin River, “found as far as Kilbourn” (Baker).
Genus Amblema Rafinesque.
Amblema rariplicata (Lamarck).
Wisconsin River (Barnes, Baker), at Prairie du Sac (Lapham,
Wiswall). Probably occurs within the boundaries of the county.
Amblema costata Rafinesque.
Koshkonong River, Cambridge, Rockdale (Morrison).
Genus Quadrula Rafinesque.
Quadrula fragosa (Conrad).
Wisconsin River (Marston, Wiswall, Chadwick, Hinkley), at
Kilbourn (Baker). Possibly in the Wisconsin River in this area.
Quadrula pustulosa (Lea).
Wisconsin River (Marston, Baker) at Prairie du Sac (Lapham),
opposite Prairie du Sac (Morrison).
Quadrula metanevra Rafinesque.
Wisconsin River (Schoolcraft, Barnes, Marston, Baker), at
Prairie du Sac (Lapham), opposite Prairie du Sac (Morrison).
Genus Tritigonia Agassiz.
Tritig onia verrucosa (Rafinesque).
Wisconsin River (Douglass, Barnes, Bullock, Baker), at Prairie
du Sac (Lapham), opposite Prairie du Sac (Morrison).
Genus Plethobasus Simpson.
420 Wisconsin Academy of Sciences , Arts , and Letters.
Plethobasus cyphyus (Rafinesque).
Wisconsin River (Baker) ; Prairie du Sac (Dudley) ; Wiscon¬
sin River, opposite Prairie du Sac (Morrison).
Genus Pleurobema (Rafinesque) Agassiz.
Pleurobema coccineum (Conrad).
Black Earth Creek, Mazomanie (Morrison).
Pleurobema coccineum solida (Lea) .
Wisconsin River (Marston, Baker), at Prairie du Sac (Lap-
ham), opposite Prairie du Sac (Morrison.)
Genus Elliptio Rafinesque.
Elliptic dilatatus (Rafinesque).
“Inhabits the Wisconsin” (Douglass, Schoolcraft, Barnes) ; Wis¬
consin River (Lapham, Baker), opposite Prairie du Sac (Mor¬
rison).
Elliptio dilatatus delicatus (Simpson).
Outlet of Lake Ripley, Cambridge (Morrison).
Subfamily ANODONTINAE Ortmann.
Genus Lasmigona Rafinesque.
Lasmigona compressa (Lea) .
Black Earth Creek, Mazomanie; outlet of Lake Ripley, Cam¬
bridge ( Morrison ) .
Lasmigona costata (Rafinesque).
Wisconsin River (Douglass, Barnes, Lapham, Marston, Baker),
at Prairie du Sac (Wiswall) ; Black Earth Creek, Mazomanie;
Koshkonong River, Cambridge, Rockdale (Morrison).
Lasmigona complanata (Barnes).
Wisconsin River (Douglass, Barnes, Lapham, Baker), opposite
Prairie du Sac; Black Earth Creek, Mazomanie; Token Creek,
Koshkonong River, Rockdale (Morrison).
Genus Anodonta Lamarck.
Anodonta grandis Say.
University shore, Lake Mendota (Wagner) ; Lake Wingra
(Baker); south margin, Lake Wingra; canal, south of Lake
Wingra; Yahara River, Madison; sloughs, south end of Lake
Monona; Koshkonong River, Cambridge, Rockdale; Black Earth
Creek, Mazomanie; Wisconsin River, opposite Prairie du Sac
(Morrison).
Anodonta grandis footiana Lea.
Madison Lakes (Bradley) ; Lake Mendota (Dudley, Baker) ;
shallow water, University shore, Lake Mendota (Wagner) ; north
of Picnic Point, Lake Mendota; west margin of Lake Monona;
outlet of Lake Ripley, Cambridge (Morrison).
Morrison — Preliminary List of the Mollusca of Dane County. 421
Anodonta (grandis) gig ant ea Lea.
University Bay, Lake Mendota, Madison (Pearse) ; Yahara Riv¬
er, Madison (Pearse); Lake Wingra (Bullock, Twenhofel) .
These records are doubtfully referrable to gigantea.
Anodonta marginata Say.
Fourth Lake (Lapham) ; Lake Mendota (Dudley) ; Yahara
River, above Lake Kegonsa (Dudley) ; Token Creek, Token Creek
(Morrison) .
Genus Anodontoides Simpson.
Anodontoides ferussacianus (Lea).
Koshkonong River, Cambridge, Rockdale; creek, four miles east
of Belleville; Black Earth Creek, Mazomanie (Morrison).
Anodontoides ferussacianus subcylindraceus (Lea).
Token Creek, Token Creek; outlet of Lake Ripley, Cambridge
(Morrison) .
Genus Alasmidonta Say.
Alasmidonta calceola (Lea).
Black Earth Creek, Mazomanie; creek, four miles east of Belle¬
ville; Token Creek, Token Creek (Morrison).
Alasmidonta marginata (Say).
Black Earth Creek, Mazomanie (Morrison).
Genus Strophitus Rafinesque.
Strophitus rugosus (Swainson).
Wisconsin River (Marston, Baker) ; Wisconsin River, opposite
Prairie du Sac; Black Earth Creek. Mazomanie; Koshkonong
River, Rockdale (Morrison).
Strophitus rugosus lacustris F. C. Baker.
Outlet of Lake Ripley, Cambridge (Morrison).
Subfamily LAMPSILINAE Ortmann.
Genus Obliquaria Rafinesque.
Obliquaria reflexa Rafinesque.
Wisconsin River (Marston, Wiswall, Baker) ; Wisconsin River,
opposite Prairie du Sac (Morrison).
Genus Obovaria Rafinesque.
Obovaria olivaria (Rafinesque).
Wisconsin River (Marston, Baker), at Prairie du Sac (Lap-
ham); opposite Prairie du Sac (Morrison).
Genus Actinonaias Fischer & Crosse.
422 Wisconsin Academy of Sciences, Arts, and Letters ,
Actinonaias carinata (Barnes).
Wisconsin River (Barnes, Lapham, Marston, Wiswall, Chad¬
wick, Baker) ; Black Earth Creek, Mazomanie; Koskonong River,
Rockdale (Morrison).
Genus Truncilla Rafinesque.
Truncilla truncata Rafinesque.
Wisconsin River (Marston, Baker), at Prairie du Sac (Lap-
ham); opposite Prairie du Sac (Morrison).
Truncilla donaciformis (Lea).
Wisconsin River (Baker), at Prairie du Sac (Lapham); oppo¬
site Prairie du Sac (Morrison).
Genus Leptodea Rafinesque.
Leptodea fragilis Rafinesque.
“Inhabits the Wisconsin” (Schoolcraft, Barnes) ; Wisconsin River
(Baker), at Prairie du Sac (Lapham), opposite Prairie du Sac
(Morrison).
Genus Proptera Rafinesque.
Propter a alata meg apt era (Rafinesque).
Wisconsin River (Douglass, Barnes, Lapham, Baker). Probably
occurs, although no specific records are at hand.
Genus Carunculina Simpson.
Carunculina parva (Barnes) .
Black Earth Creek, two miles west of Mazomanie (Morrison).
Genus Ligumia Swainson.
Ligumia recta latissima (Rafinesque).
Wisconsin River (Douglass, Barnes, Marston, Baker), at Prairie
du Sac (Lapham), opposite Prairie du Sac (Morrison).
Ligumia ellipsiformis (Conrad).
Wisconsin River, opposite Prairie du Sac; Koshkonong River,
Rockdale; outlet of Lake Ripley, Cambridge (Morrison).
Genus Lampsilis Rafinesque.
Lampsilis siliquoidea (Barnes).
“Inhabits the Wisconsin” (Douglass, Barnes) ; Wisconsin River
(Lapham, Baker) ; Lake Mendota, Madison (Pearse) ; south
margin, Lake Wingra; Koshkonong River, Cambridge, Rockdale;
Wisconsin River, opposite Prairie du Sac (Morrison).
Lampsilis siliquoidea rosacea (DeKay).
Madison Lakes (Bradley) ; Lake Monona (Bullock, Baker) ; Uni¬
versity shore, Lake Mendota (Wagner) ; Tenney Park shores,
shores north of Picnic Point, Lake Mendota; west margin, Lake
Monona (Morrison).
Monisoyv~~Preliminary List of the Mollusca of Dane County. 423
Lampsilis ventricosa occidens (Lea).
“Inhabits the Wisconsin” (Schoolcraft, Barnes) ; Wisconsin
River (Lapham, Wiswall, Chadwick, Baker), at Prairie du Sac
(Wiswall), opposite Prairie du Sac; Black Earth Creek, Mazo-
manie; Koshkonong River, Rockdale (Morrison).
Lampsilis ventricosa lurida Simpson.
Fourth Lake (Lapham); west margin, Lake Monona; outlet of
Lake Ripley, Cambridge (Morrison).
Order TELEODESMACEA Dali.
Superfamily CYRENACEA Tryon.
Family SPHAERIIDAE Dali.
Subfamily SPHAERIINAE F. C. Baker.
Genus Sphaerium Scopoli.
Sphaerium sulcatum (Lamarck).
Stream, south of Lake Wingra; creek, four miles east of Belle¬
ville; Token Creek, Token Creek (Morrison).
Sphaerium crassum Sterki.
Yahara River, Madison (Juday, Morrison).
Sphaerium solidulum (Prime).
Yahara River, Madison; Black Earth Creek, Mazomanie (Morri¬
son).
Sphaerium bakeri Sterki.
Koshkonong River, Rockdale; Token Creek, Token Creek (Morri¬
son).
Sphaerium striatinum (Lamarck).
Outlet of Lake Ripley, Cambridge (Morrison).
Genus Musculium Link.
Musculium transversum (Say).
Black Earth Creek, Mazomanie; Koshkonong River, Rockdale
(Morrison).
Musculium truncatum (Linsley).
Stream, south of Lake Wingra (Morrison).
Musculium rosaceum (Prime).
Pheasant Branch, near mouth (Morrison).
Musculium securis (Prime).
Swamp, south of Lake Wingra; along University Bay, Lake Men-
dota (Morrison).
Subfamily PISIDIINAE F. C. Baker.
Genus Pisidium C. Pfeiffer.
424 Wisconsin Academy of Sciences , Arts , and Letters.
Pisidium idahoense Roper.
Lake Mendota (Juday, Baker) ; dredging, off Picnic Point, Lake
Mendota (Morrison).
Pisidium compressum Prime.
Wisconsin River, opposite Prairie du Sac; Pheasant Branch, near
mouth ; Koshkonong River, Rockdale ; outlet of Lake Ripley,
Cambridge; creek, four miles east of Belleville (Morrison).
Pisidium fallax Sterki.
Koshkonong River, Rockdale (Morrison).
Pisidium punctatum Sterki.
Yaliara River, Madison (Morrison).
Pisidium variable Prime.
Dredging, off Picnic Point, Lake Mendota (Morrison).
Pisidium glabellum Sterki.
Merrill Spring (four miles west of Madison) (Morrison).
Pisidium adamsi Prime.
Stream, south of Lake Wingra; Yahara River, Madison (Morri¬
son).
Pisidium sargenti Sterki.
Outlet of Lake Ripley, Cambridge (Morrison).
Pisidium neglectum Sterki.
Stream, south of Lake Wingra (Morrison).
Pisidium noveboracense Prime.
Merrill Spring; Merrill Spring stream; springs, along Merrill
Creek; spring fed streams, south of Lake Wingra; Yahara River,
Madison (Morrison).
Pisidium scutellatum Sterki.
Sandy shallows, Tenney Park shore, Lake Mendota (Morrison).
Pisidium walkeri Sterki.
Pheasant Branch, near mouth (Morrison).
Pisidium roperi Sterki.
Swamp, south of Lake Wingra; swamp, north of Picnic Point,
Lake Mendota (Morrison).
Pisidium politum Sterki.
Pheasant Branch, near mouth (Morrison).
Pisidium abditum xialdeman.
Stream, south of Lake Wingra (Morrison).
Morrison — Preliminary List of the Mollusca of Dane County. 425
Pisidium levissimum Sterki.
Springs, forest ponds, south of Lake Wingra (Morrison).
Pisidium rotundatum Prime.
Forest ponds, swamp, south of Lake Wingra; swamp, north of
Picnic Point; swamp, University Bay, Lake Mendota (Morri¬
son).
Pisidium minutum Sterki.
Spring fed stream, south of Lake Wingra (Morrison).
Pisidium subtruncatum Malmgren.
Dredging, off Picnic Point, Lake Mendota (Morrison).
Pisidium concinnulum Sterki.
Stream, south of Lake Wingra; outlet of pumping station, Uni¬
versity Bay, Lake Mendota (Morrison).
Pisidium pusillum Jenyus.
Forest ponds, swamp, south of Lake Wingra; swamp, University
Bay, Lake Mendota; swamp, four miles east of Belleville (Morri¬
son).
28
THE HYPODERMAL GLANDS OF THE BLACK
SCALE, SAISSETIA OLEAE (BERNARD)
I. The Dorsal Glands
Wm. S. Marshall
Not long ago there appeared an article (3) in which the
statement was made that very little is known of the hypo-
dermal glands of the Coccidae and that some of the work
which has been published on this subject is not always de¬
tailed or correct. Several months before the appearance
of this paper, work was started upon the hypodermal
glands of the black scale, Saissetia oleae; work not at first
so much about the structure of the glands and the functions
of their secretions as upon their origin. The work en¬
larged as it went on and a study of structure was added to
that of development; the other question, function, has been
omitted. It is generally the practice to call all hypodermal
glands of the Coccidae, wax glands, a classification which
may not be entirely true. To avoid any mistake it has been
deemed advantageous to classify these hypodermal glands
as to their position upon the insect’s body and they have
been divided into dorsal and ventral glands; only those of
the first category are described in this paper. Besides the
glands, if they are all such, there are the sense organs and
setae upon this surface of the scale insect’s body ; these, at
least at present, will not be described and this paper is re¬
stricted to the hypodermal glands upon the dorsal surface.
The methods used in preserving the scale insects were
four of the standard solutions, Carnoy, Kahle, Tower and
Bouin; the first two were found to be better than the oth¬
ers. A number of stains were tried; iron haematoxylin
was used for a number of slides but most of the sections
were stained in Delafield’s haematoxylin followed by an
aqueous solution of red eosin, Bordeaux red or acid fus-
chin; these colored the cuticula. An old method of stain¬
ing with pyroligneous acid and haematoxylin was found to
428 Wisconsin Academy of Sciences, Arts, and Letters .
be very good as it both stained the tissues and darkened the
cuticula. The ordinary formula was changed by decreas¬
ing the amount of haematoxylin. The slides remained in
this solution a few hours and then a large part of the hae¬
matoxylin was removed by acid alcohol; the preparations
were then placed in tap water for ten to twenty minutes
to regain their blue color and mounted. In several editions
of Bolles-Lee’s Microtomist’s Vade-mecum this method is
referred to as follows : “Burchardt’s pyroligneous acid hae¬
matoxylin would seem to be superfluous at least.” The pre¬
paration consists in dissolving 2 gr. potassium alum in 130
gr. pyroligneous acid and add to this 0.5 gr. haematoxylin
dissolved in 70% alcohol. As already mentioned a much
smaller amount of haematoxylin was found to give better
results. The solution of pyroligneous acid with potassium
can be used for coloring the cuticula and then followed by
some other stain for the tissues.
Sections of the embryo show that the hypodermis is
about the same over the entire body, but before emergence
a difference in the thickness of this layer is noticed between
that part covering the dorsal and the same layer on the ven¬
tral surface of the body. Dorsally the hypodermis is about
twice as thick as the cuticula ; the nuclei of its cells are el¬
liptical in outline with their longitudinal axis parallel to
the body's surface (fig. 1) ; there are no cell boundaries dis¬
cernible. Over the ventral surface the nuclei are more flat¬
tened, with only a strand of cytoplasm connecting them with
each other. The cuticula on the dorsal part of the body is
thicker than that covering the ventral surface.
The very young larvae, those which have been out of the
egg but a few hours, are small and active; before they
permanently settle they wander over the surface of the leaf
and, even after they have become fixed, they may change
their location as they are often found, of different sizes,
walking over the plant. Sections of these very young lar¬
vae show the layers of hypodermis and cuticula to be about
what they were in the old embryos; there are marked dif¬
ferences in both layers upon dorsal and ventral surfaces,
but transitions between the extremes of the two can readily
be found. Both layers are thicker on the dorsal (fig. 2)
than on the ventral surface but in these youngest larvae no
Marshall— Glands of the Black Beale, Saissetia Oleae. 429
traces could be found of any developing hypodermal
glands ; this was due either to the small size of the cells and
the thinness of the cuticula or they do not begin to develop
until the first instar is several days old. The nuclei of the
hypodermal cells were plainly visible but no cell boundaries
could be seen ; these cells are not numerous over the surface
of the body in this early stage ; in a longitudinal section of
the abdomen only four to six nuclei were seen in each seg¬
ment. These statements regarding the size of the hypo¬
dermal cells are very general, as many exceptions can be
found in different places on both dorsal and ventral sur¬
faces. In transverse sections one can see at the lateral
margins a gradual change in the size of the hypodermal
cells from the larger dorsal ones to the smaller and more
flattened cells on the ventral surface, also a errresponding
decrease in the thickness of the cuticula. The largest cells
in the hypodermal layer are those, dorsal on the head, above
the supraoesophageal ganglion. At different areas on the
dorsal surface some of the nuclei assume a spherical shape
(fig. 3), but elongated nuclei are more abundant; ventrally
all the nuclei are flattened. In the hypodermis a number
of mitotic figures can be seen, both nuclei and cells becom¬
ing more numerous as the young scale insect increases in
size.
The larvae during their first instar increase very much
in size and show great changes in the thickness of the cuti¬
cula and of the hypodermis. Quayle and Rust (13) state
that during the first instar the young larvae increase in size
from 0.35 mml 0.2 mm. to 0.7 mm.X 0.35 mm. and that
the first stadium lasts from a month to six weeks. This
long stadial period accounts for the changes which take
place in the hypodermis and the gland cells and why both
are so different towards the end of the first stadium from
what is found in the newly hatched larva. A few days
makes a marked difference in the size of the hypodermal
cells and in the thickness of the cuticula. Over the dorsal
and the ventral surfaces of the body the proportionate dif¬
ferences noticed in the embryo are still observed ; what this
is can be seen by comparing figures three and four with
figure one; these are all drawn to the same scale.
The earliest noticeable appearance of the glands (fig. 5)
430 Wisconsin Academy of Sciences, Arts, and Letters.
was found in small, first instar larvae after they had set¬
tled on the plant and increased slightly in size. No gland
was found without its pore through the cuticula; in fact
when hunting for the glands in the earliest stages it was
easier to find the pore in the cuticula and locate the gland
in this way than to attempt to first find the glands them¬
selves. A great many more pores were found than perfect
glands ; it was difficult to find both pore and gland well
shown in the same section and they were, at this early
stage, too small to follow from one section to the next. As
already mentioned the large cone shaped pore through the
cuticula was not difficult to find and it, the first trace of the
developing gland, was proportionally quite large; at the
upper surface of the cuticula it was narrow, but increased
in diameter towards the inner surface of the hypodermis.
About the first formation of the pore we know nothing and
can only suppose that the underlying cell failed to secrete
cuticula except the thin outer layer which is present and
through which is the small opening allowing the secretion
of the gland to pass out.
At first but a single cell of the hypodermis appears to
take part in the formation of the gland and this cell shows
but little difference from the others of this layer except that
its shape is changed by its protrusion into the pore. Hav¬
ing examined many of these early glands, similar to figure
five, I am unable to say whether the gland first appears as
a single cell or is composed of more than one ; none of them
showed any trace of a multicellular origin, although they
very early have more than a single nucleus. Examining
many of them and from different specimens one would
think that two or more nuclei within the boundary of the
cell could easily be distinguished ; this was not observed.
When the glands begin to increase in size, some of them
lie more or less parallel to the surface of the cuti¬
cula, others protrude into the body cavity away from
the hypodermis. The former of these are generally located
where some organ of the body lies adjacent to the hypoder¬
mis and the gland is thus unable to push as far away from
the neighboring hypodermal cells as if there was no tissue
at their base. Figure six shows such a developing gland
which in its growth has crowded against the cells of the hy-
Marshall — Glands of the Black Scale, Saissetia Oleae. 431
podermis. The gland now shows quite an increase in size,
much more so than do the regular hypodermal cells, and its
nucleus is much larger; this refers to the main nucleus,
probably the original one, as the others are no larger than
those of the regular hypodermal cells. The changes whicn
have so far taken place in the gland are its multinuclear
condition and its increase in size.
The next change in the gland is the appearance of a vacu¬
ole, small at first but increasing in size with the growth of
the gland of which it soon becomes the largest part. The po¬
sition of this vacuole (fig. 7) is constant; it is found in the
large part of the pyriform gland and between the nucleus
and the duct like portion. As the main part of the gland
has increased in size and pushed out into the haemocoele
there is always a narrow stem-like part which connects it
with the exit of the cuticular pore. This tubular part of
the gland often has the appearance of a regular duct and,
whatever its structure, it functions as an exit for the secre¬
tion of the gland which has collected within the large vacu¬
ole or bladder.
The further growth of the gland means principally an in¬
crease in its size and especially of the vacuole ; more nuclei
appear and two or three may be seen in a section. During
the growth of the glands there is some difference in their
general appearance; this is caused by the pressure against
them of parts within the body causing them to be more or
less intrahypodermal or they may not be subject to such
pressure and then protrude free into the cavity of the in¬
sect's body. Figures nine and ten show glands taken from
the dorsal surface of the same larva. Figure nine is of a
gland located about midway between the median axis of the
body and its margin. In this situation there are no large
organs in the haemocoele and the gland in its growth is
free to push down and acquire its normal shape. Figure
ten is taken from the median dorsal part of the same in¬
sect; it is in a position where the alimentary tract pushes
up against the dorsal wall of the body and the gland has in
part grown by pushing against the neighboring cells of the
hypodermis.
During the growth of the gland the regular hypodermis
has increased in thickness as has also the cuticular layer
432 Wisconsin Academy of Sciences , Arts, and Letters .
and the relative thickness of these layers varies in different
parts of the dorsal surface of the insect’s body. The pore
through the cuticula is necessarily longer, but it has the
same relative shape that it had in the younger larvae; it
gradually widens from the outer towards the inner surface
of the cuticula, but it is not so wide relative to its length
as in the earlier stages.
These hypodermal glands reach their maximum growth
(fig. 11) towards the end of the first instar and show no
difference except in size from those just described. The
appearance of the wall of the gland relative to the size of
the vacuole depends somewhat upon how near the center
the section is cut. No cell boundaries can be found but
there are a number of nuclei and of these there is always
one, probably the original of the earliest stage, consider¬
ably larger than any of the others. In this as in succeed¬
ing stages there is quite a difference in the thickness of the
hypodermal and cuticular layers, depending upon what part
of the dorsal wall is examined ; there is also a difference in
the shape of the cells of the former layer ; these are gener¬
ally longer in the median part of the body, but exceptions
to this can be found.
The problem of the comparative age of the specimens
which have been sectioned and studied in this work is one
about which little can be said and the criterion used was
the development of the glands in one specimen as compared
with their condition in the others. This has been followed
and no attempt made to rear a number of marked speci¬
mens which could be removed and sectioned at stated inter¬
vals. It was at one time thought that the width of the
specimens might be used as a comparison, but then a
greater or less convexity of the body might change this and
it soon became evident that sometimes a narrower insect
showed a further development of the glands and a thicker
cuticula than one which was wider.
A subject closely connected with this was the appearance
of the second type of gland, also of its origin. Judging
from two specimens sectioned, these were in the process of
ecdysis, it can be said that the first type of dorsal gland is
for the first instar and the second type appears just before
or during the change from first to second larval stage ; this
Marshall — Glands of the Black Scale, Saissetia Oleae. 433
last type of gland, with changes which will later be de¬
scribed, lasts until the insect becomes mature and probably
until its death. That these two specimens were in the pro¬
cess of ecdysis was shown by the outer layer of cuticula be¬
ing removed from an underlying one; in this outer layer,
the cuticula of the first instar, pores were found of the first
type of gland, none of the second type, while in the under¬
lying cuticula, that of the second instar, the pores all
showed the stopper characteristic of these glands (fig. 12).
One also noticed that in these two specimens all sections of
the tracheae enclosed a smaller tube, the intima of the tra¬
cheae of the first instar ; this had not been cast out because
the cuticula of the first instar had not yet been discarded.
The regular glands (fig. 11) found in the first instar lar¬
vae and which no doubt secrete the wax covering the dorsal
surface of the body during this period of the insect’s life,
apparently decrease in size towards the end of the first in¬
star, but whether the second type of glands becomes modi¬
fied from them or develops as a new set is uncertain; that
many of them do develop as new glands must be true as
there are very many more present in the second instar than
while the insect is in its first larval stage. The pore
through the cuticula leading to the surface is quite different
in the two and the presence of the stopper in the gland of
the second type and the increase in thickness of the cuti-
cular layer and later, the enlargement of the outlet, the
cavity in the cuticula, give the differences which readily
distinguish the one type from the other.
An early appearance of the second type of glands, those
developing after the first ecdysis, has been found in a num¬
ber of specimens and is well shown in that one where the
cuticula of the first instar is being cast. The stopper (fig.
13) of the pore is distinctly shown in the cuticula and pro¬
trudes a little below its inner surface. Its wall is thick and
it is easily seen in all sections cut through the center of the
pore ; in the older specimens, when the rest of the pore, the
cuticular cavity, widens, there are many sections of the
gland in which the stopper is not seen ; in these no outlet to
the exterior is shown, the sections being cut to one side of
the stopper. This peculiar, short, tube-like piece of the
cuticula, the stopper, is very similar in the black scale to a
434 Wisconsin Academy of Sciences, Arts, and Letters.
corresponding port found in other scale issects (8, 17).
The second type gland cell in this early stage is very similar
to a corresponding stage in the development of the first
type of gland but, being from a much older and much larger
larva the gland cell itself and the neighboring hypoder-
mal cells are very much larger. This is readily seen in a
comparison of figure twelve with figure five which is the
earliest stage in the development of the gland of the first
type.
The earliest gland of the second type (fig. 12) is very
similar to the other hypodermal cells ; it is longer, its nucleus
is larger, and, where it touches the cuticula, it is in connec¬
tion with a stopper. As in the first type of gland these also
appear of unicellular origin, but their connection with other
cells at a very early stage makes this uncertain.
When a slightly older gland is examined (fig. 14), one
in which the vacuole, not always circular in outline al¬
though this is the most common shape, has appeared, it
then becomes evident that more than one cell composes the
gland and, as we go on in its development, we will see that,
until the scale insect becomes mature, we are describing a
multicellular gland; there is an association of smaller cells
with the large one containing the vacuole. The number of
these small cells is doubtful but there are at least four to
each gland and probably more. The similarity of the vacu¬
ole in this gland to that in the early first type gland (fig. 7)
is also noticeable not only in appearance, but in position
within the gland. In the second instar, as in the first, one
finds that the largest and longest cells of the dorsal hypo-
dermis are near the median axis of the insect's body. Due
to pressure from other cells on its free end the gland cell
may be slightly curved.
So far in thei1" development the similarity, except in size,
between the first and second types of glands has been very
marked; from now on they begin to differ and the second
gland will not hereafter show any stage that can be com¬
pared with the corresponding development of the first type
of gland. Continuing with the development of the gland
of the second instar we find that it increases slightly in size,
but that the greatest change is in the vacuole ; this we will
now call the bladder. This for some time retains its spher-
Marshall — Glands of the Black Beale, Saissetia Oleae. 435
ical, or it may be flattened, shape to Anally become ovoid,
with its narrow end pointed towards the apex of the gland
or, put in another way, towards the outer surface of th£
cuticula. From the apex of the bladder the cell extends
through the cavity in the cuticula until it reaches the stop¬
per; this narrow part appears to form a tube through
which the contents of the bladder may pass to the outer
surface of the insect's body (fig, 14). With the increasing
age of the scale insect the bladder appears to slowly pass
into the cavity of the cuticula, but in reality the cuticula is
increasing in thickness to gradually envelop the bladder
and finally to surround it.
As the insect grows older the cells of the hypodermis con¬
tinue to secrete more and more cuticula, adding this of
course to the inner surface which becomes irregular.
Where each gland is situated there is, as already men¬
tioned, a conical cavity in the cuticular layer and this cav¬
ity situated as a cap over the gland becomes longer and
longer as the cuticular layer increases in thickness (fig.
15). At first the newly secreted cuticula surrounding each
cavity is only in part over the gland and its bladder is as yet
entirely surrounded by the layer of hypodermal cells.
With this growth it soon becomes apparent that each cuti¬
cular cavity contains a number of strands forming a reticu¬
lum; this is light colored at first, but becomes darker with
the age of the insect. After the appearance of this reticu¬
lum one easily notices that each cuticular cavity is becom¬
ing larger, not only by the inward growth of the cuticula,
but by lateral expansions on all sides ; an early stage of this
is seen in figure fifteen. This figure of only two of these
cavities shows no connection between them but this has, to
a slight extent, already taken place (fig. 16). This last
figure is taken from the same specimen as the preceding,
but at a position where no glands can be seen; it shows,
however, that such a communication between the cuticular
cavities has already begun.
Such a honeycombing of the cuticula is difficult to under¬
stand ; of what use it can be to a permanently fixed insect
is doubtful, but the figures from now on will all show that it
does occur and is found in all specimens. This dissolution
of a part of the cuticula must be due to some secretion and
436 Wisconsin Academy of Sciences, Arts, and Letters.
this probably comes from the small accessory cells of the
gland, those which are the dark staining ones. These
cells, at least some of them, become very dark in the stained
specimens and the cells or their secretion continue into the
cavity as a dark strand which passes along the margin of
the bladder to become lost in the reticulum (fig. 20).
There is also the possibility that these irregular cavities
might be formed by an unequal secretion from the cells of
the hypodermis. From the same specimen from which fig¬
ure eighteen was drawn, two transverse sections are shown
(fig. 19) . The smaller of these figures is a surface view
looking at the end of the stopper ; the clear space surround¬
ing this represents the apex of the cuticular cavity which
appears lighter than the surrounding cuticula. Figure b
shows a transverse section of the bladder and the cavity in
the cuticula in which it lies. Here one can notice the reti¬
culum which, closely surrounding the bladder, extends in
all directions to the margin of the cavity.
In the specimens of this and somewhat later stages there
is a difference in how far the bladder of the glands has be¬
come surrounded by the cuticula ; neither it nor the cuticu¬
lar layer have changed in appearance (fig. 17), but the lat¬
ter is thicker and its cavities have increased in size. In
all sections cut directly through the median part of the
gland, the original gland cell is prominent and it and its
large clear nucleus are easily seen, especially if the cell pro¬
trudes beyond the neighboring hypodermal cells. A view
of a section cut in some other plane may show some of the
other cells of the gland (fig. 18).
The shape of the cuticular cavities depends to a certain
extent upon the thickness of the cuticular layer; in the
long narrow ones, found especially at the posterior part of
the body, the bladder is also long and thin and its outer end
is, as in the shorter ones, more pointed than the end di¬
rected towards the center of the body. At this age the nu¬
cleus of the main gland cell has started to lose its regular
appearance and its chromatin contents is crowded to one
side. This is followed by a change in the cell near its nu¬
cleus (fig. 22) ; its regular structure has become modified
and in its place there appears a loose reticular arrangement
of the contents quite similar to what first appears in the
Marshall — Glands of the Black Scale , Saissetia Oleae. 437
cuticular cavities; this condition, with slight changes, re¬
mains until the cell disappears. The dark staining cells
are readily found in every gland ; each one has a long nar¬
row extension passing into the cuticular cavity and extend¬
ing to the side of the bladder (fig. 20). More than one of
these cells is usually found in each gland. In these last two
figures, twenty and twenty-two, the regular hypodermal
cells are still normal and have not begun to disintegrate as
occurs when the insect reaches maturity.
The duct passing from the bladder to the stopper is al¬
ways present and easily seen in sections cut through the
center of the gland and cuticular cavity. It develops from
the thinner part of the original gland cell and such a de¬
velopment can be seen in figures twelve, fourteen and fif¬
teen. After it is formed it remains unchanged for a con¬
siderable time during the growth of the gland, but, as the
scale insect reaches maturity, the wall of the duct becomes
darker and more distinctly seen; it becomes yellow or
brown in color. At this time and until the maturity of the
insect, the wall of the duct is thicker and darker near the
apex of the bladder (fig. 21).
The addition to the inner surface of the cuticular layer
which has been described, ceases after a time and in the
full grown scale insects this layer is, relative to the size of
the body, thinner than in the specimens that are half or
three-quarters grown. As the insect becomes mature the
increase in body size is due to egg formation and the re¬
tention in the body of the eggs until the embryo in each is
well along in its development. The cuticula shown in the
figures might be compared one stage with another if the
magnification was the same, were it not that variations in
the thickness of this layer occur on different parts of the
dorsal wall of the body.
The most noticeable change to now take place is in the
cells of the glands and of the hypodermis ; this change is so
great that in the oldest insects examined the hypodermis
has disappeared and its cells and their nuclei together with
those of the glands have all broken down and migrated, in
part or entirely, into the cuticular cavities. This seems to
be the most plausible explanation as these cavities now con¬
tain a dark and irregular mass in which are a number of
438 Wisconsin Academy of Sciences, Arts, and Letters.
small nuclear-like bodies ; these are no doubt from the regu¬
lar cells that have disappeared from their former and origi¬
nal position to finally pass into the cavities.
By a mature black scale we mean one within the body of
which a number of well developed embryos are found; all
the figures not yet mentioned, 23 to 27 inclusive, are from
these mature insects. In following these figures one can
notice what takes place until, in the last section shown (fig.
26) practically no hypodermis remains. This goes even
further than is shown in this figure as in some specimens,
evidently the oldest, no trace is left of any cytoplasm lying
along the inner wall of the cuticular layer. The bladder
appears to persist and the stopper is always present.
In the breaking up of the hypodermis and the glands the
cells of the gland, especially what appeared to be the main
one, lose the nucleus which becomes a dark staining, irregu¬
lar mass represented by what appears to be the chromatin
alone; later the contents of the cell changes to irregular
strands. This cell now becomes clearer than it was; it is
situated at the opening of the cuticular cavity into which
it appears to pass (fig. 24) . The beginning of this change
was noticed in an earlier stage (fig. 22) and it seems prob¬
able that these figures represent the disintegration of the
cell and its nucleus and their final disappearance into the
cavity.
The number of these glands in the dorsal wall of the
black scale is very large; they appear more numerous and
crowded together in the third instar before it reaches ma¬
turity. In some sections or parts of a section the glands
are crowded together as much as possible (fig. 15) and, as
the scale insect increases in size, the glands spread a little
further apart, probably due to a small amount of stretching
of the cuticula ; I know of no other way to explain it. Even
after the insect has become mature, sections show that the
cuticular cavities of the glands are not far apart (fig. 26).
A view of the dorsal surface of the mature insect (fig. 27)
gives one a better idea of their number and close proximity.
This last mentioned figure would have been about the same
as the one drawn if it had been taken from any other place
on the dorsal wall; the glands themselves are not shown,
Marshall — Glands of the Black Scale, Saissetia Oleae. 439
but the figure shows the cuticular cavities and each one of
these represents the position of a former gland.
In four specimens of very old insects that were sectioned,
another condition was noticed which has so far not been
described. In all of these insects the gland cells and those
of the hypodermis had in part or entirely disappeared and
in the cuticular cavity containing most of their contents
was noticed a dark yellow, irregular mass; this appeared
to partially fill some cavities and entirely fill others; this
same yellow substance was noticed in what remained of
the gland cells. This mass in one specimen was noticed as
a fine layer over the inner surface of the cuticula. The
meaning of this I do not know unless it is that the final
stage is the formation within the cuticular cavities of a
hard mass to completely block them ; whether this comes
to all insects of a certain age and shortly before their death
is unknown.
The old scale insects are difficult to section as the hard
cuticular covering makes it almost impossible to procure
anything like a series. The best method tried was to cut
the old insects into four or more pieces, after they were
preserved, and then section each piece. When these were
cut and mounted one could find, here and there, a section in
which the cuticular layer was in its normal position in con¬
tact with the body; many sections had to be discarded be¬
cause they were so badly broken that no piece could be
found where cuticula and body were in normal relation to
each other. Very often the cuticula was torn entirely away
from the body.
Summary
Two types of dorsal hypodermal glands occur in the black
scale. The first type is present only in the first instar and
these glands develop early in larval life from regular hypo-
dermal cells; a vacuole soon appears in each gland and this
soon grows to be its largest part. The gland communi¬
cates with the outside through a duct, part of the original
gland cell, and a small opening in the cuticula. These are
wax glands and secrete the thin layer of this substance cov¬
ering the dorsal surface of the body.
The second type of gland first appears in the early sec-
440 Wisconsin Academy of Sciences , Arts , and Letters.
ond instar and persists through the life of the insect, al¬
though it finally loses its normal shape and position. The
gland develops from the hypodermis and its early stages
are very similar to those of the first type gland. A vacuole
appears in the original gland cell; this later becomes the
bladder which connects with the outer surface through a
duct and a small cuticular stopper. The cuticular layer in¬
creases in thickness by secretion from the regular hypoder-
mal cells to finally surround the bladder, but leaving a cav¬
ity around it. This cavity increases in size by lateral en¬
largements; these are probably formed by some secretion
dissolving a part of the cuticula. These lateral enlarge¬
ments may increase until they communicate with each
other. The use of these glands is not known and they, with
the regular hypodermal cells, finally break down and pass
into the cavity of the cuticula.
Bibliography
1. Berlese, A. Gli Insetti. Vol. I, p. 505.
2. Bugnion, E. and N. Popoff. Les glandes cirieres de Flata
(Phromnia) marginella. Bull. Soc. Vaud. Sc. nat. XLIII, 1907.
p. 549.
3. Ferris, G. F. The wax-secreting organs of the Coccidae. Pan-
Pacific Entom. V, 1928. p, 67.
4. Fullaway, D. T. Description of a new Coccid species, Ceroputo
ambigua, with notes on its life-history and anatomy. Proc.
Davenport Acad. Sci. XII, 1910. p. 223.
5. Johnston, C. E. The internal anatomy of Icerya purchasi. Ann.
Entom. Soc. Amer. V, 1912. p. 383.
6. Kitao, Z. Notes on the anatomy of Warajicoccus corpulentus
Kuwana, a scale insect noxious to various oaks. Journ. Coll.
Agri. Imp. Univ. Tokyo X, 1928. p. 1.
7. Matheson, R. The wax secreting glands of Pseudococcus citri.
Ann. Entom. Soc. Amer. XVI, 1923. p. 50.
8. Mayer, P. Zur Kenntnis von Coccus cacti. Mitth. Zool. Stat.
Neapel X, 1891-93. p. 505.
9. Moulton, D. The Monterey pine scale, Physokermes nisignicola.
Proc. Davenport Acad. Sc. XII, 1907. p. 1.
10. Murdock, G. E. The wax-secreting mechanism in the adult fe¬
male of Icerya purchasi. Pan-Pacific Entom. V, 1928. p. 71.
11. Oguma, K. A new scale insect, Xylococcus alni, on alder with
special reference to its metamorphosis and anatomy. Journ.
Coll. Agric. Imp. Univ. Sapporo, Japan. VIII, 1918-20. p. 77.
Marshall — Glands of the Black Scale , Saissetia Oleae. 441
12. Putman, J. D. Biological and other notes of Coccidae. I. Pul-
vinaria innumerabilis. Proc. Davenport Acad. Sc. II, 1879.
p. 293.
13. Quayle, H. J. and E. W. Rust. The Black Scale. Univ. Calif.
Publ. Agric. Exper. Station Bull. No. 223, 1911. Also: Ma-
teriali per la storia di alcuni insetti dell ’olivo. Redia, IV,
1907. p. 48.
14. Sulk, K. Zur Anatomie der Cocciden. Zool. Anz. XXXIV, 1909.
p. 164.
15. Teodoro, G. Le glandule ceripare della femmina della Pulvinaria
camelicola. Redia, VII, 1911. p. 172.
16. Teodoro, G. La secrezione della cera nei mashi della Pulvinaria
camelicola. Redia, VII, 1911. p. 352.
17. Teodoro, G. Le glandule laccipare e ceripare del Lecanium
oleae. Redia, VIII, 1912. p. 312.
Explanations of Plates
Abbreviations used in plates
Bl., bladder. Gl., gland.
Ch., chorion. Hyp., hypodermis.
Cut., cuticula. Vac., vacuole.
Cut. Cav., cuticular cavity.
Plate 10
All the figures shown in this plate are of the first type of gland;
they are all of the same magnification, X 1400 diameters.
Fig. 1. A transverse section through the dorsal wall of an em¬
bryo. Due to contraction during preparation the chorion, Ch., is not
close to the cuticula, Cut., of the embryo.
Fig. 2. Section of a first instar larva shortly after its emergence
from the egg. This is from the dorsal wall of the body, anterior and
near the margin.
Fig. 3. From a transverse section of the body wall of the thorax
of a young first instar larva a few days older than the preceding.
Fig. 4. From the same specimen as figure three, but from the ven¬
tral body wall of the thorax.
Fig. 5. Section from the dorsal wall of a young first instar larva.
It shows the earliest noticeable appearance of the gland, Gl., which is
as yet of about the same size as the neighboring hypodermal cells,
but of a different shape.
Fig. 6. From a little older larva; the gland, Gl., is flattened
against the cuticula by the pressure of some adjacent organ. The
duct is cut a little to one side and it is not continuous to the outer
surface of the cuticula.
Fig. 7. Gland from a first instar larva showing the early forma¬
tion of the vacuole, Vac., within the gland, Gl.
29
442 Wisconsin Academy of Sciences , Arts, and Letters .
Fig. 8. Transverse section of a gland showing the vacuole and
two of the smaller nuclei.
Fig. 9. Gland from an older first instar larva.
Fig. 10. Two neighboring glands from the same larva as the pre¬
ceding one; the section is from the median part of the dorsal body
wall where the alimentary canal crowds up against it.
Fig. 11. Section of a gland from a nearly full grown first instar
larva.
Plate 11
Fig. 12. Section of a first instar larva in the process of ecdysis;
its loosened cuticula, Cut., shows a pore of the first type gland. This
figure is the earliest stage of the second type gland, Gl., at the end
of which in the cuticular layer, Cut. ’, is the stopper. X 1400.
Fig. 13. Diagram of a stopper in section, a, and surface view, b.
Fig. 14. Section from a young second instar larva showing the
gland, Gl., containing the vacuole, Vac., and with three small sec¬
ondary cells. X 1400.
Fig. 15. Two neighboring glands, Gl., each with bladder, Bl.,
from which a strand of cytoplasm or tube passes through the cuticu¬
lar cavity to the stopper. The extra cuticular secretion is forming
and has partially filled the cuticular cavities. X 940.
Fig. 16. Another section from the same specimen as preceding
figure and showing the hypodermal layer, Hyp., with the enlarged
cuticular cavities some of which are already connected with each
other. X 940.
Fig. 17. Section of a gland from an older larva than the preced¬
ing two figures. The cuticular layer, Cut., has grown in thickness
and now surrounds the bladder on all sides. Three of the smaller
gland cells are shown touching or adjacent to the main one. X 940.
Fig. 18. Gland showing some of the accessory cells, not the main
one shown in most figures. This is cut a little to one side; it shows
the enlarged cuticular cavity but not the stopper. X 940.
Fig. 19. From same specimen as last figure, 18. This shows at
a the surface view of a stopper with the apex of the cavity in the
cuticular layer shown as a clear surrounding space; b a transverse
section through the same cavity, but in the region of the bladder,
Bl., around which is seen the reticulum within the cuticular cavity.
X 940.
Plate 12
Fig. 20. Gland with a much elongated cuticular cavity. The nu¬
cleus of the main gland cell, Gl., has lost its normal appearance.
X 940.
Fig. 21. One of the ducts leading from the bladder to the stop¬
per. This becomes hardened in the old insects and in many speci¬
mens is yellow and very noticeable. X 1400.
TRANS. WIS. ACAD. — VOL. 24
PLATE JO
C-ni.
F iy 3.
01.
C u t.
H'JF
Cut
H,Jp
Cut.
Hyp.
fif 1
Cut.
r* typ
. \/ac.
Vac
■utr.
c-i.
CL
Fiq. !0
fF; h,
Cut.
Tifj. /
C-L.
■■m
Fi a. 8.
C-i.
0:V-
TRANS. WIS. ACAD. - VOL. 24
PLATE II
TRANS. WIS. ACAD. - VOL. 24
PLATE 12
.
T‘^-
Marshall — Glands of the Black Beale , Saissetia Oleae. 443
Fig. 22. Showing one regular cell of the hypodermis, Hyp., and
a few cells of the gland. The original large gland cell, GL, has
started to disintegrate; of the three other gland cells shown, two of
them are the darker cells which no doubt secrete the fluid forming
the reticulum. X 940.
Fig. 23. Showing the further disappearance of the hypodermis.
Two cuticular cavities, neither cut through the center, do not show
the stopper and the one to the right does not even show its communi¬
cation with the hypodermal layer. The bladder is present and one
is seen in the cuticular cavity to the right. This and the following
figures are all taken from insects containing well developed embryos
in their body. X 940.
Fig. 24. This specimen shows a quite large abnormal looking cell
at the opening of the cuticular cavity and, at its base, what appears
to be the remains of the nucleus. X 740.
Fig. 25. In this figure, as in all others of the old insects, the
bladder is still present in the cuticular cavity and would be con¬
nected to the stopper by its duct were the section through the mid¬
dle of the cavity. The hypodermis, Hyp., has lost its regular ap¬
pearance and has begun to disappear. X 740.
Fig. 26. In this figure two of the cuticular cavities are shown,
but neither cut near enough its center to show the stopper. The
hypodermis has nearly disappeared, no doubt having migrated into
the cuticular cavities. X 940.
Fig. 27. Surface view of an old black scale showing the spacing
of the glands over the dorsal surface; the clear spaces are the cu¬
ticular cavities. X 220.
PRELIMINARY STUDIES OF THE INTRACELLULAR
SYMBIONTS OF SAISSETIA OLEAE (BERNARD) *
A. A. Granovsky
In a course of study of mosaic diseases of economic
plants, many writers observed that sucking insects were
able to transmit and disseminate infective principles, which
cause the symptoms of these puzzling diseases. It has also
been shown that insects are far more efficient in inoculation
experiments of virous diseases than the modern methods
of pathological technique. In looking for a possible ex¬
planation of these facts, a number of sucking insects have
been examined for micro-organisms, hoping to learn some¬
thing of the cause and nature of mosaic diseases.
In this search for internal micro-organisms, a series of
insects, infesting numerous plants in greenhouses, were
taken for preliminary study during the winter of 1924-25
and the following winters up to the present date. Different
forms of organisms, varying from minute, highly refractive
spherical bodies to rather large oval or considerably elon¬
gated yeast-like bodies, have been found inhabiting several
species of aphids and scale insects. In their appearance
and location within the insects they resembled the sup¬
posedly symbiotic organisms reported by several workers.
These organisms were found to be especially numerous
within the scale insects. Because of the unusualness of
such association, attention has been given to one of the
species of scale insects, namely, the black scale of Cali¬
fornia, Saissetia oleae (Bernard), feeding on oleander
plants in the greenhouses. The reasons for selecting this
species of insect for study of intracellular symbionts were
as follows :
1. This scale insect was fairly abundant in number to in¬
sure sufficient material for study.
•Contribution from the Department of Economic Entomology, Wisconsin
Agricultural Experiment Station, University of Wisconsin.
446 Wisconsin Academy of Sciences, Arts, and Letters .
2. It was observed that the oleander leaves showed a con¬
siderable discoloration around the feeding punctures and
in severe cases the leaves were slightly distorted on account
of insect injury.
3. Every specimen of the insect showed the presence of
a large population of yeast-like organisms, which are rela¬
tively large in size and consequently are convenient to
study.
A Brief Historical Review
The symbiotic organisms of insects in general, and in
Homoptera especially, were first reported by F. Leydig in
1854. He described free living lanceolate bodies in the
lymph of Lecanium (Coccus) hesperidum Linn, and re¬
garded them as parasitic. In 1858 Huxley discovered the
symbionts in the embryos of parthenogenetic aphids. He
called the tissue with these organisms the “pseudovitellus”.
This name was used by several workers later, applying it
to certain cells in the developing ova. Metchnikov, 1866,
in his embryological studies of aphids, coccids and psyllids,
found the organisms infecting the ova, multiplying therein
and distributing themselves within the developing embryos.
He also used the term pseudovitellae for them.
J. D. Putnam in 1880 was the first in this country to
observe the yeast-like bodies in the ovaries of Pulvinaria
innumerabilis Rathvon. Besides giving an excellent de¬
scription of the organisms, he considered them after Leydig
to be parasitic. He also suggested that they may repre¬
sent the spermatophores and may be instrumental in the
fertilization of ova.
Until recently very little has been known concerning this
subject. Only the works of Berlese, 1893, and Pierantoni,
1910, in Italy; those of Buchner, 1910-1928, and Lindner,
1895-1907, in Germany; Conte and Faucheron, 1907, in
France; Sulc, 1910, in Czechoslovakia; Brain, 1923, in
South Africa; and those of Shinji, 1919, Brues and Glaser,
1921, and Schrader, 1923, in this country, contributed to
our knowledge of the intracellular symbionts of many in¬
sects, especially those belonging to Homoptera.
Several attempts have been made to classify the sym¬
bionts found in many insects. Moniez, 1887, was the first
Granovsky — Intracellular Symbionts of Saissetia Oleae. 447
who made this attempt, naming* the organism observed by
Leydig in Lecanium hesperidum Linn, as Lecaniascus poly-
morphus. He based his description on the characters ob¬
served in culture, believing that he had isolated the organ¬
ism. It was later shown, however, that he was dealing
with a contaminated culture. Lindner in 1895 renamed
the yeast-like bodies of this species of scale and placed it in
the genus of Saccharomyces. Sulc in 1906-1910, and
Buchner in 1912 erected several new genera with the de¬
scription of a few new species of symbionts found in differ¬
ent insects. These are accepted by workers on the subject
and have good possibilities of retaining their names in fu¬
ture literature on the subject. Brain in 1928, in his phe¬
nomenal work on classification “of the intracellular sym¬
bionts of insects with a short record of every known
species” revised the whole subject and erected several new
genera and described a number of new species of symbionts
found in coccids. He also gives a brief account of previous
work by other authors. His new genus Lecaniocola is of
interest in this paper. This genus is, no doubt, closely re¬
lated to Saccharomyces, but so far as is known no endos-
pores have been observed in these yeast-like bodies which
separate them from the latter genus. Another striking char¬
acteristic of the genus Lecaniocola is that the organisms
placed in it are found in scale insects, especially those be¬
longing to the sub-family Lecaniinae and are believed to be
in symbiotic relationship with the insects. The organisms
of this genus are found free in haemolymph or connective
fat tissue, as contrasted with the symbionts, belonging to
Kermincola Sulc, which are found in definite mycetom.
During the present study of symbionts in Saissetia oleae
(Bernard) it was found that Conte and Faucheron in 1907,
describing the organisms from Lecanium hemisphericum
Targ., made a brief statement is regard to the presence of
organisms in the black scale of California. This statement
in a free translation is as follows :
“ Lecanium hemisphericum is not the only coccid contain¬
ing yeast forms. We have found them always present in
very large numbers in Lecanium oleae, Lecanium hesperi¬
dum, Pulvinaria floccifera, etc. These present a different
aspect from those in Lecanium hemisphericum. The com-
448 Wisconsin Academy of Sciences , Arts, and Letters.
parative study of all forms only allows us to define their
affinities”.
This is the only original statement found in literature on
the subject in regard to symbionts met with in Saissetia
oleae (Bernard). It will be of interest to note, that Brain,
in describing all species of symbionts known to him, does
not list one from Saissetia oleae, although this insect oc¬
curs in South Africa on a number of plants such as olive,
Olea europaea Linn., Dovyalis caffra Harv. and Hibiscus
sp., but it is evidently not occurring on oleander, Nerium
oleander Linn, in South Africa, for Brain omits this plant
from his host list of the insect in question. A number of
species of scale insects are reported by Brain as feeding on
oleander, but there is no S. oleae among them. In Europe
this scale feeds primarily on olive, while in this country it
infests a number of plants such as orange, olive, oleander,
live oak and many others.
Methods of Study with Results of Observations*
As was stated above, all individuals of Saissetia oleae
(Bernard) are infested with the yeast-like organisms. If
any of the young nymphs, mature females, or even the eggs
are crushed on or between clean and sterile slides in a drop
of normal salt solution, the organisms may be seen quite
readily under low, or still better under the high power of
a microscope. They are easily separated by the normal
salt solution from the fat globules, and this enables one to
study them without much trouble from any extraneous mat¬
ter. Being heavier than the solution used, they always
drop next to the slide, while the fat globules remain next
to the cover glass. In such a simple preparation the sym¬
bionts are free in the normal salt medium and appear to be
colorless, with several large vacuoles and conspicuously
granular protoplasm, always having a few highly refrac¬
tive bodies. Their general form is oval, elongated, yeast¬
like, often with the buds on the pointed polar end. (Plate
13, fig. 1.).
*In this connection I wish to express my sincere gratitude to Dr. W. S.
Marshall and Dr. E. M. Gilbert for their constructive suggestions during
the progress of this work.
Granovsky — Intracellular Symbionts of Saissetia Oleae. 449
The half -grown nymphs and young females invariably
yielded a greater number of these organisms than the ex¬
tremely young nymphs or older females. This is probably
on account of the fact, that in the very young nymphs the
symbionts did not have enough time for multiplication to
reach their greatest number, while in the older, mature in¬
dividuals, the symbionts are reduced in number due to dis¬
integration or modification. Their development and in¬
crease in number, as well as subsequent disintegration is
apparently running parallel with the accumulation and as¬
similation of adipose tissue within the insect.
The leaves of oleander, infested with the scales, were
washed in sterile water and in normal salt solution and the
washings examined for the organisms, but, in spite of re¬
peated efforts, not a single yeast-like body, resembling the
organism in question, was detected. This seems to indi¬
cate that the surface of the infested leaves is free from the
organisms.
The scales were also washed in the normal salt solution
in a watch-glass and their washings examined, but they,
too, failed to yield the organisms. This, in its turn, indi¬
cates that the organisms are not present on the surface of
the insects.
A sufficient amount of honey dew given off by this species
of scale was collected and examined for the yeast-like
bodies, but their presence was not detected, in spite of the
fact that the honey dew was often rich in fungous and bac¬
terial flora. Evidently they are not found in the honey
dew of the insect, at least in their common forms ; although
very minute spherical and highly refractive bodies, not
more than granules, were often observed to be present in
the honey dew of these scale insects.
An attempt was made to study the multiplication of the
organisms in hanging drop cultures, using the normal salt
solution for the medium. To prevent contamination, every
insect was passed quickly through an alcohol lamp flame
several times before it was dissected on a sterile slide for
study. Mercuric bichloride 1 :500 was also used for sterilza-
tion with equal success. Only a small amount of fat tissue
of the insect containing the organisms was placed in hanging
drop cultures. A suitable field under a microscope was
450 Wisconsin Academy of Sciences , Arts , and Letters.
then located and the approximate positions of all the organ¬
isms in view were sketched, paying particular attention to
their forms, shapes, and budding. The position of the
stage was also noted and microscopes were left undisturbed
for the time of observation. It was repeatedly observed
that the organisms may multiply for a short duration of
time, although not rapidly. The buds as a rule develop at
the pointed end, and only rarely small spherical buds may
be produced at the rounded pole of the mother organism.
The buds at first may have the appearance of only small
spherical projections which gradually grow and increase in
size, and, upon reaching about half of the size of the mother
cell, they may separate. Often, however, the mature cell
produces a long neck, at the end of which a small swelling
begins to develop, and this swelling grows into a separate
individual, as the result of drawing the neck between the two
ceils to a very fine point. Buds often persist until they are
fully mature and only then separate. Most of the buds,
however, leave the parent cells while they are quite small,
and increase in size after their separation. It was found
that the organisms in the normal salt solution multiply and
increase in size quite rapidly only during the first 24 or 48
hours ; later there is no sign of multiplication. In old cul¬
tures many organisms evidently disintegrated and disap¬
peared. It was observed that greater growth was pro¬
duced near the insect tissue present in the cultures, than in
a pure normal salt solution. The organisms, evidently, in
hanging drop cultures drew on their own reserves as well
as on the insect tissue.
In the hanging drop cultures, and in the temporary nor¬
mal salt solution preparations, many minute spherical bodies
were also observed. They were often arranged in rosettes
or were floating free in the medium. What they are, is not
clearly known. They might be modified forms of the yeast¬
like organisms, or they may represent entirely distinct or¬
ganisms living within the scale insect.
The internal structure of the yeast-like symbionts has
been studied in smears, prepared in the ordinary method
and stained with Orange G, acid Fuchsin, Safranin, Eosin,
and Loeffler’s methylene blue alone, or in their combina¬
tions. The best smears, showing the internal structure,
Granovsky — Intracellular Symbionts of Saissetia Oleae. 451
were obtained with the combination of Orange G and aque¬
ous Safranin. It may not be amiss to mention that it is ex¬
tremely difficult to bring out the details of structure by va¬
rious combinations of differential staining. Triple Flem¬
ming stain also gave good results when carefully destained.
The best smears show the presence of several large vacu¬
oles, usually 3 or 4 in elongated and older symbionts, and
only one or two in smaller and younger cells. The proto¬
plasm is conspicuously granular and is arranged in streaks
between the vacuoles. It often stains deeply in two or
three places giving the impression of the presence of sev¬
eral nuclei, although it is doubtful if these deeply colored
spots are nuclei. The presence of a number of minute
clear spots is conspicuous. These are highly refractive
and are scattered in the protoplasm, especially toward the
pointed end of the organism, where budding takes place.
(Plate 15, fig. 5-9).
In preparing the microscope sections, ordinary histolog¬
ical methods were followed with special precaution to dehy¬
drate in the alcohols, as well as to clear in the xylol very
gradually. The same can be said of infiltration in paraffin.
A series of specimens were killed and fixed in Flemming's
medium, Tower’s No. 2, Kahle’s and Bouins’ fixing solu¬
tions. The best results were obtained with the Bouins’ and
Kahle’s solutions, although Flemming’s solution also gave
fairly good results. The staining is very difficult, but after
numerous trials as to combinations of differential stains,
and the time required in each of them, it was found that
the combination of Delafield Haematoxylin 20 minutes and
the saturated aqueous solution of safranin for one and one-
half minutes gave satisfying results. Two other com¬
binations gave good results. One of them was Orange G,
10 minutes and cyanin 5 minutes, when fixed in Bouins’ so¬
lution. The other was that of ordinary Flemming’s triple
stain.
After the cutting of serial sections of the nymphs and
mature females, as well as the eggs, collected from under
the shells of the insects, it was evident that the half-grown
individuals had a higher population of the organisms than
the mature females. The symbionts appeared to be intra¬
cellular in nature and were embedded in the connective fat
452 Wisconsin A cademy of Sciences , Arts, and Letters .
tissue in great numbers, and were usually present free in
the haemolymph. (Plate 13, fig. 2). The greatest num¬
ber of them was found toward the perifery of the younger
insects. In the mature females the symbionts were less
numerous and were distributed more evenly throughout
the whole insect, although most of them were restricted to
the connective fat tissue. At this time there is a noticeable
modification or disintegration of these micro-organisms,
for it is difficult to explain the reduction of their number
in any other way. They probably break down into minute
and hardly noticeable granules or gradually disappear en¬
tirely as the amount of adipose tissue is considerably di¬
gested and reduced by metabolic processes with the age of
the insect.
A number of serial sections reveal the entrance of these
micro-organisms into ova soon after their differentiation
from nurse cells and follicular epithelial cells, and before
the early stage of clearage. The manner in which the
symbionts enter the eggs is not clear; it is evident, how¬
ever, that they gain entrance in more than one way. They
are observed to enter the egg at the anterior end through
the nurse cells evidently with the stream of the nutriative
material, and are often lodged between the epithelial layer
and nurse cells, as well as the ovarian chamber. (Plate 14
fig. 3). Other sections seem to indicate that infection of
the ova takes place by mass action of the organisms at the
anterior pole of the egg. In this case a number of rather
small oval bodies penetrate the egg, instead of a few elon¬
gated forms of the organism passing from the nurse cells
to the egg with the nutriative material. (Plate 15, figures
10 and 11).
The organisms were occasionally found in the developing
embryos within the mother, and were always found in the
advanced embryos in the sectioned eggs, as well as in the
newly hatched insects. (Plate 14 fig. 4) . A critical study
of this phenomenon in all stages of the insect has not been
conducted, and consequently it must be passed without dis¬
cussion. However, such a study is in progress, and it can
be safely mentioned that the organisms in the young em¬
bryo multiply very rapidly until the maturity of the insect,
at which time their number is considerably reduced, and that
Granovsky — Intracellular Symbionts of Saissetia Oleae. 453
each individual is provided with its quota of these organ¬
isms upon hatching from the egg.
Systematic Position of the Organism
A careful comparative study of the organism found in
Saissetia oleae (Bernard) with those reported from other
scale insects shows that it closely approaches Lecaniocola
inglisiae Brain found in Inglisia geranii Brain. Yet, it has
some resemblance to Lecaniocola filippae Brain found in
Filippa chilianthi Brain. It resembles the first form by its
internal structure, while in the external morphology it ap¬
proaches the second form. So far as measurements are
concerned it is neither. Considering the above mentioned
facts it is believed to be a distinct form and is described
here as a new species.
Lecaniocola saissetiae sp. nov.
(Plate 15, fig. 5-9)
General form of the organism elongate, varying greatly
in size and shape, commonly spindle shaped, sometimes pear
or egg shaped, rarely spherical. Usually pointed at one
end, often with a long narrow neck, the other end being
rounded. Sometimes both ends are pointed. Protoplasm
coarse, conspicuously granular, highly vacuolated, having
from 2 to 4 or more large and a few smaller vacuoles, with
a few minute highly refractive bodies. Multiplication by
terminal budding at the pointed end; sometimes buds are
formed on both ends. Buds may be spherical at first and
later become oval or spindle shaped. More often the bud is
produced as an oval swelling at the end of the elongated
neck. Separation of buds in hanging drop cultures of nor¬
mal salt solution takes place between 24 and 48 hours.
More rarely the buds persist until fully grown. Only one
or two, rarely three, terminal buds observed in a chain.
Average size of typical mature organisms about 13 to 14 /*
long and 3.5 /* broad. Average size of the newly separated
individuals 6.5 to 7/* long and 3 to 3.5 /* broad. Average
size for small, egg shaped forms 3 to 5 /* long. Chains of
two or three buds often reach 26.5/* in length. The width
454 Wisconsin Academy of Sciences, Arts, and Letters .
varies from 2.5 to 4/*, the usual average being 3.5/x. The
width of the neck varies between 1 and 2/x.
Organism found in the connective fat tissue and free in
haemolymph of Saissetia oleae (Bernard).
Function of the Organisms
The exact function of the symbiotic fungi or mycetozoa
is not clearly understood. Their relation to the insects
within which they gain shelter is believed to be symbiotic
in nature. The benefit derived by the micro-organism is
apparent. It enjoys protection and abundance of food.
Several benefits may be suggested as being derived by the
insect from the presence of these organisms. The most log¬
ical one is that of assisting in breaking down the fat tissue
and assimilation of nutriative materials by some enzyme
action during the formation of ova and embryonic develop¬
ment of the insects. The fact that the number of organisms
is reduced within the mature insect after the formation of
ova, at which time most of the fat tissue is utilized, is
strongly in favor of that view.
The fact that every egg has symbiotic organisms within
its embryo indicates that they must play an important role
within the insect. It is quite possible that the organisms
play an essential and yet some unknown part in the rapid
development of the embryos within the eggs of this insect.
The males of this species of scale insect are usually rare,
and it is generally assumed that the insect reproduces par-
thenogenetically, at least in part. It is quite possible, in
the light of the demonstration by Loeb, 1912, of the pos¬
sibility of artificial fertilization of eggs of lower animals,
that these yeast-like organisms are instrumental in initiat¬
ing the embryonic development of parthenogenetic ova by
enzymic stimuli or by the combination of physical, chemical
and metabolic activities of organisms within the insect and
its eggs.
Granovsky-Intracellular Symbionts of Saissetia Oleae. 455
Bibliography
1. Brain , C. K. 1923. The intracellular symbionts of
some South African Coccidae. Ann. Univ. Stellen¬
bosch, 1, sec. A, no. 2 : 1-48. 12 plates.
2. - 1924. Host plant index of South African scale
insects (Coccidae) with a list of species found on each
plant recorded. Ibid. 2, Sec. A, no. 2:1-44.
3. Brues , C. T. and Glaser , R. W. 1921. A symbiotic
fungus occurring in the fat body of Pulvinaria innum-
erabilis Rath. Biol. Bui. 40: 299-324. 3 plates.
4. Buchner , P. 1921. Tier und Pflanze in intrazellularer
Symbiose. 462 p. 2 plates and figures. Berlin.
5. Conte , A. et Faucheron , F. 1907. Presence de levures
dans le corps adipeaux de divers Coccides. Comptes
Rendus. 145 : 1223-1225.
6. Putnam , J. D. 1880. Biological and other notes on
Coccidae. Proc. Davenport Acad. Sci 2, pt. 2 : 293-347.
1 plate.
7. Schrader , F. 1923. The origin of the mycetocytes in
Pseudococcus. Biol. Bui. 45: 279-302. 3 plates.
8. Shinji , G. 0. 1919. Embryology of Coccids with spe¬
cial reference to the ovary, origin and differentiation
of the germ cells, germ layers, rudiments of the mid¬
gut and the intracellular symbiotic organism Journ.
Morphol. 33: 73-167. 20 plates.
Explanation of Plates
Plate 13
Fig. 1. Photomicrograph taken from smear, showing yeast-like
bodies from fat tissue of Saissetia oleae (Bernard).
Fig. 2. Photomicrograph of a 6^ section, showing the distribution
of symbionts in situ of adipose tissue of a mature scale insect.
Plate 14
Fig. 3. Photomicrograph of a longitudinal section through the
egg within the insect, showing the yeast-like symbionts penetrating
the ovum.
Fig. 4. Photomicrograph of a 6^ section through the young em¬
bryo within the egg of the insect, showing the presence and multipli¬
cation of yeast-like symbionts therein.
456 Wisconsin Academy of Sciences, Arts, and Letters .
Plate 15
Fig. 5. Yeast-like body, Lecaniocola saissetiae n. sp. with elonga¬
tion at one pole, preparatory for budding.
Fig. 6. Formation of bud at the pointed pole.
Fig. 7. Formation of buds on both poles.
Fig. 8. The bud at the pointed pole is about to separate from the
mother organism.
Fig. 9. A newly separated yeast-like cell, showing tendency to
form a bud soon after separation.
Fig. 10. Penetration of ovum by means of mass attack of yeast¬
like organisms.
Fig. 11. A single organism penetrated ovum through the nurse
cell.
TRANS. WIS. ACAD., VOL. 24
PLATE 13
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X jj ' x
* A. ' ' * V. * 'A afg|||||| •> - „%k‘' . /
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TRAMS. WIS. ACAD., VOL. 24
PLATE 14
TRANS. WIS. ACAD. - VOL. 24
PLATE 15
10
11
THE BIRDS OF DANE COUNTY, WISCONSIN
A. W. SCHORGER
Most of Wisconsin remains virgin territory from the
standpoint of ornithology. Except for the Koshkonong re¬
gion east to Racine and along the Michigan shore to Green
Bay, very few comprehensive observations covering a period
of years have been made. The avifauna of a given area
usually changes greatly in the course of time. Some species
become extinct or rare, while others change the route of
their migration. It is a source of great regret that the
Madison region should have had so little of its bird history
recorded, especially since many of its earlier residents were
qualified to leave data that would be invaluable at this time.
The Golden Plover, though once common, no longer mi¬
grates through the state. There are no published records
for the county. Correspondence and conversations with
the oldest hunters failed to afford proof of their former
presence ; hence, that the Golden Plover visited Dane County
is only a reasonable assumption.
The advance or retreat of a species is always of interest.
Allan W. Carpenter1, in 1892, found the Vesper Sparrow
and Indigo Bunting to be rare summer residents at Madi¬
son. During the last 20 years, at least, they have been
common. Within this period also the Cardinal and Tufted
Titmouse have become permanent residents, and the West¬
ern Meadowlark a common summer bird.
It was decided to confine the present list of birds almost
entirely to Dane County. This may appear arbitrary in
view of the extensive work of the Kumliens on Lake Kosh¬
konong, a small area of which lies within our limits. The
fact that all the Kumlien skins examined were labelled Jef¬
ferson County had no bearing on this decision. Brewster
has stated rightly “that all arbitrary boundaries established
merely for convenience, should be regarded as more or less
1 Cited by Lynds Jones, Wilson Quart. 4, No. 1 (1892) 28 and 34.
30
458 Wisconsin Academy of Sciences , Arts, and Letters .
flexible.” The main purpose has been to give a picture of
the bird life as it has existed during the past fifteen years
and relatively little work has been done on Lake Koshko-
nong during that time. In addition, an attempt has been
made to rescue data on certain extinct species such as the
Passenger Pigeon before it is too late.
Physiography
Dane County is located centrally in the southern portion
of the state and is 24 miles from the Illinois line. Were it
not for the intersection of the northwest corner by the Wis¬
consin River, the county would be an almost perfect rect¬
angle. The distance from north to south is 30 miles, and
from east to west 42 miles. Of the total area of 1,238
square miles, only 120 square miles bordering the Wiscon¬
sin River fail to lie within the drainage area of the Rock
River basin.
Nature has endowed the county generously with those
features conducive to the presence of a great variety and
abundance of birds. Four of the largest lakes, Kegonsa,
Waubesa, Monona, and Mendota, have a combined area
of 28.7 square miles. The waters of these lakes run
through the Yahara River into the Rock River. This sys¬
tem forms a great highway for waterfowl. In spring the
line of flight passes over the chain of lakes north to the
Wisconsin River that contributes its quota of migrants
from the south and southwest. The Wisconsin River route
is favored particularly by geese on account of the extensive
sand flats in its wide and usually shallow bed, and Lake Wis¬
consin where the birds can rest in safety. Adjacent level
cornfields are used as feeding grounds.
The western portion, approximately one-third of the
county, lies in the driftless or non-glaciated area. The line
bounding this area enters the county at Sauk City, passes
between Black Earth and Cross Plains in a slightly south¬
eastern direction, and leaves the county near Belleville.
This portion of the county is a succession of high hills and
contains the most extensive tracts of woodland. The Sauk
County shore of the river has a high bluff, where the Duck
Hawk breeds and raids the adjacent lowlands of Dane
Schorger — Birds of Dane County , Wisconsin. 459
County. Here are extensive sandy wastes where the Lark
Sparrow occurs, extensive marshes, and large timber areas
that are unique in their number of the rarer birds. The
rolling country of the central and eastern portion of the
county contains most of the lakes and marshes.
Principal Trees
With the exception of the prairie that was limited to the
watershed, and naturally the lakes and marshes, the county
was formerly well timbered. Oaks predominate still, the
most abundant species being the white oak ( Quercus alba
Linn.), scarlet oak (Q. coccinea Muench.) , and red oak (Q.
rubra Linn.) Indigenous gymnosperms, with the ex¬
ception of the tamarack (Larix laricinia Koch) of the
swamps, are relatively scarce. A few black spruce (Picea
mariana B. S. P.) are to be found at Hook Lake, and scatter¬
ing white pines ( Firms strobus Linn.) at Pine Bluff, and
other places in the western portion of the county. The red
juniper ( Juniperus virginiana Linn.) is not uncommon, par¬
ticularly near the Wisconsin River. Boreal birds, however,
find such cultivated species as Norway spruce (Picea excelsa
Link.), and European larch ( Larix europea D. C.) to their
liking. The Redpolls, Crossbills, and Pine Grosbeaks pre¬
fer the cones of the European larch to all the others.
The following additional hardwoods are common:
Bur oak ( Quercus macrocarpa Michx.)
Glaucous willow (Salix discolor Muehl.)
Hoary willow (Salix Candida Fliigge)
Aspen (Populus tremuloides Michx.)
Large-toothed aspen (Populus grandidentata Michx.)
Basswood (Tilia americana Linn.)
Sugar maple (Acer saccharum Marsh.)
Silver maple (Acer saccharinum Linn.)
Box elder (Acer negundo Linn.)
Locust (Robinia pseudacacia Linn.)
Wild plum (Prunus americana Marsh.)
Choke cherry (Prunus virginiana Linn.)
Black cherry (Prunus serotina Ehrh.)
Serviceberry (Amelanchier canadensis Linn.)
White ash (Fraxinus americana Linn.)
460 Wisconsin Academy of Sciences, Arts, and Letters .
Black ash ( Fraxinus nigra Marsh.)
White elm ( Ulmus americana Linn.)
Butternut ( Juglans cinera Linn.)
Black walnut ( Juglans nigra Linn.)
Shag-bark hickory ( Hicoria ovata Britton.)
Bitternut ( Hicoria minima Britton.)
The following vines and shrubs are an important source
of fruits upon which birds feed:
Wild red raspberry ( Rubus strigosus Michx.)
Black raspberry ( Rubus occidentalis Linn.)
Dewberry ( Rubus sp.)
Blackberry ( Rubus sp.)
Prickly gooseberry ( Ribes cynosbati Linn.)
Missouri gooseberry ( Ribes gracile Michx.)
Wild black currant ( Ribes floridum L’Her.)
Common elder ( Sambucus canadensis Linn.)
Frost grape ( Vitis riparia Michx.)
Virginia creeper ( Psedera quinquefolia Greene)
Climbing bitter-sweet ( Celastrus scandens Linn.)
Poison ivy ( Rhus toxicodendron Linn.)
Life Zones
The county lies in the Alleghanian Area of the Transi¬
tion Zone with the exception of a narrow strip bordering the
Wisconsin River that it is unquestionably in the Carolinian
Area of the Upper Austral Zone. The division is based on
the avifauna which on account of its mobility always ex¬
tends beyond the plants peculiar to a given zone. Of the
trees characteristic of the Carolinian Area, sassafras, tulip
tree, hackberry, sycamore, sweet gum, rose magnolia, red
bud, and persimmon, all are absent from or excessively
rare in the county with the exception of the hackberry.
There is a single sycamore growing naturally at the edge of
the water at Colladay Point, Lake Kegonsa, but this may
have originated from the seed of introduced trees.
The following species of birds characteristic of the Caro¬
linian Area breed in the Mazomanie region: Tufted Tit¬
mouse, Blue-gray Gnatcatcher, Cardinal, Kentucky Warbler,
Louisiana Water Thrush, Blue-winged Warbler, Prothono-
Schorger — Birds of Dane County , Wisconsin.
461
tary Warbler, Yellow-breasted Chat, Red-bellied Wood¬
pecker, and Red-shouldered Hawk.
Some of the trees common to the bottom lands are : river
birch, white elm, silver maple, white ash, glaucous willow,
prickly ash, bur oak, red oak, black oak, white oak, large-
toothed aspen, basswood, red juniper, and hoary alder.
Seasonal Distribution of Birds
There are few birds that do not migrate to some extent
during the year. Certain species such as the Blue Jay and
Red-headed Woodpecker winter regularly, but the numbers
remaining represent but a fraction of the summer residents.
Permanent Residents .
Black Duck
Red-tailed Hawk
Ruffed Grouse
Prairie Chicken
Bob-white
Ring-necked Pheasant
Barn Owl
Screech Owl
Great-horned Owl
Barred Owl
Long-eared Owl
Short-eared Owl
Pileated Woodpecker
Irregular Winter Visitors
Goshawk
Snowy Owl
Saw-whet Owl
Horned Lark
Bohemian Waxwing
Northern Shrike
Evening Grosbeak
Red-bellied Woodpecker
Red-headed Woodpecker
Hairy Woodpecker
Downy Woodpecker
Blue Jay
Crow
Tufted Titmouse
Chickadee
White-breasted Nuthatch
Starling
Cardinal
English Sparrow
American Goldfinch
the Boreal Region.
Pine Grosbeak
Redpoll
Pine Siskin
American Crossbill
White-winged Crossbill
Snow Bunting
from
Summer Residents Occasionally Wintering.
Mallard Brown Thrasher
Cooper’s Hawk Robin
Marsh Hawk Bluebird
Sparrow Hawk Cedar Waxwing
462 Wisconsin Academy of Sciences , Arts , and Letters.
Coot
Mourning Dove
Flicker
Prairie Horned Lark
Prairie Marsh Wren
Meadowlark
Thick-billed Redwing
Cowbird
Song Sparrow
Birds Breeding to the Northward, but Migrating Regularly
Southward in Fall, Occasionally Wintering.
Canada Goose
Golden-eye
Merganser
Rough-legged Hawk
Wilson's Snipe
Herring Gull
Yellow-bellied Sapsucker
Red-breasted Nuthatch
Brown Creeper
Winter Wren
Golden-crowned Kinglet
Rusty Blackbird
Purple Finch
Slate-colored Junco *
Tree Sparrow *
White-throated Sparrow
Lapland Longspur
* Winters regularly
Acknowledgments
The data on migration are taken largely from the author's
notes that were begun in 1910, and kept regularly from 1913
to the present time. Use has been made of the notes of the
late Norman DeWitt Betts covering the years 1913 to 1916.
Special recognition is given to Mr. Warner Taylor who, ex¬
cept during late summer and early autumn, has done ex¬
tensive work in the field since 1913. Dr. Witmer Stone
kindly provided the revised nomenclature and arrangement
of species in advance of the appearance of the new check¬
list.
ANNOTATED LIST OF BIRDS
Part I
Family GAVIIDAE. Loons
1. Gavia immer (Briinnick) . Loon. This species is a
common migrant. Individuals in mature plumage are
found frequently in summer but there are no recent breed¬
ing records. It appears shortly after the lakes open, the
average date of arrival being April 3. In 1923, several
were seen on March 11. It occurs in largest numbers dur-
Schorger — Birds of Dane County , Wisconsin.
463
in g the first two weeks of April and only an occasional bird
is found after May 8.
In autumn, the Loon seldom arrives before the last week
in October. The earliest date is Sept. 11, 1914. It is most
common in November and has not been noted after De¬
cember 1.
Family COLYMBIDAE. Grebes
2. Colymbus holboelli (Reinhardt). Holboell’s Grebe.
An uncommon migrant. One was seen April 28, 1922 by
Mr. Warner Taylor1. Another was shot by a hunter on
Lake Mendota, Nov. 15, 1925, and identified by Mr. Tay¬
lor2 who gave the bird to me.
The skin is now in my collection.
3. Colymbus auritus Linnaeus. Horned Grebe. This
handsome grebe is a common migrant. It is found oc¬
casionally in marshes, but usually on the large lakes, in
small loose flocks. The average date of arrival is April 14 ;
the earliest date April 5, 1925. The latest date of departure
is May 14, 1917. The migration is usually completed before
May 7. : > ^M1J|
The fall migrants appear October 15 and depart before
December. The latest date is Nov. 28, 1920. This species
frequently swims into duck decoys. If a flock takes fright
and scatters, the birds utter a high-pitched collecting note,
“pee-your, pee-your”.
4. Podilymbus podiceps (Linnaeus). Pied-billed Grebe.
An abundant summer resident. Scarcely a marsh with
standing water or reedy pond lacks several breeding pairs.
The average date of arrival is April 6, the earliest date be¬
ing March 20 (Taylor). The fall migration is completed
by Nov. 27. The latest date is Dec. 20, 1925. There is
always the chance that an unusual date is due to a wounded
bird.
Nests with eggs have been found from June 9 to 19. It
is exceptional to find a nest the eggs of which have not been
carefully covered with a mass of decayed vegetable matter
by the departing bird.
1 Auk 40 (1923) 339.
2 Auk 43 (1926) 250.
464 Wisconsin Academy of Sciences, Arts, and Letters .
Family PELECANIDAE. Pelicans
5. Pelecanus erythrorhynchos Gmelin. White Pelican.
The white pelican had become a rare migrant in southern
Wisconsin by 1900. The route of migration is now mainly
west of the Mississippi River. Five were seen over Lake
Waubesa, April 22, 1925 by Warner Taylor3 ; and two were
seen on Mud Lake April 15, 1926 by Dr. W. V. Bryant4.
A white pelican was shot by Dr. Arthur G. Sullivan near
the mouth of the Yahara River, Lake Mendota, the latter
part of October in either 1917 or 1918. “This was a single
bird that sat out on the sandbar the greater part of the day
and flew north of Crescent Bog making a circle of the bay
every thirty or forty minutes, and returning to the sand¬
bar, which is west of Farwell’s Point. Each time the bird
passed over me my curiosity increased because I did not
know that it was a pelican. Finally, on about the fourth
or fifth excursion the bird made, I shot it.”
Dr. S. H. Chase informed me that he remembers the
pelican. Owing to destruction by fire of the record book of
the Crescent Club, the date cannot be fixed more closely.
Family PHALACROCORACIDE. Cormorants
6. Phalacrocorax auritus auritus (Lesson). Double-
crested Cormorant. There has been a marked increase in
this species since 1920. Flocks numbering several hun¬
dred birds are now not uncommon. The habit of flying
abreast in only slight V-formation produces an impressive
black line against the sky. The birds arrive as early as
April 2, and the height of the migration is reached the last
week in the month. The earliest date of arrival is March
31, 1928 (Taylor). Final departure from the Madison
lakes takes place May 15 to 18.
This species nested with Great Blue Herons at Okee in
1921, in the trees killed by the formation of Lake Wiscon¬
sin5. I found the herons nesting here in 1919, but there
8 Auk 43 (1926) 250.
4 Schorger, Auk 43 (1926) 556.
5 Stoddard, Yearbook Public Museum Milwaukee for 1921 (1922) 36 ;
Shrosbree, Ibid. 28.
S charger — Birds of Dane County, Wisconsin.
465
were no Cormorants present. A flock of 12 birds was seen
on a pond at Springfield Corners, July 17, 1927.
The southward movement in fall takes place largely be¬
yond our limits since relatively few Cormorants are to be
seen. The extreme dates of arrival and departure are Sep¬
tember 23, 1925 and November 15, 1922.
Family ARDEIDAE. Herons and Bitterns
7. Ardea herodias herodias Linnaeus. Great Blue Heron.
Common summer resident, arriving on the average April 6.
The earliest date of arrival is March 22, 1925. The latest
fall record is October 25, 1919.
A heronry at Okee, Lake Wisconsin, contained 14 nests
in 19196, and was occupied through 19217. The nesting
trees, standing in the water, have since been felled.
Most of the eggs are deposited the last two weeks in
April. The period of incubation is about 28 days. Since
the eggs at Okee began to hatch May 11, some of them must
have been deposited before April 15.
8. Casmerodius egretta (Gmelin). Egret. Though for¬
merly nesting in the state, the Egret is now only a rare
straggler from the south. When found in southern Wis¬
consin, it is usually in August or September. One was
seen May 30, 1925 at Mud Lake, near Waterloo, Jefferson
County, by Warner Taylor8. The locality is only a few
miles from the county line.
9. Butorides virescens virescens (Linnaeus). Green
Heron. Common summer resident, arriving the first week
in May. The earliest date is April 25 (Taylor). It has
not been noted after September 30.
Nests with eggs have been found from May 19 to June
16. The birds usually nest apart but colonies are some¬
times found. A colonial nesting site may be abandoned for
no apparent reason and then reoccupied after an interval
of one or more years. The condition of a group of six
nests found in an oak thicket on June 5 was: one nest
with 5 eggs ; one with 6 eggs ; two with 3 eggs and 2 young ;
8 Schorger, Auk 37 (1920) 143.
7 Stoddard, vide supra.
8 Auk 43 (1926) 250.
466 Wisconsin Academy of Sciences , Arts , cmd Letters .
one with one egg1 and 3 young; and one with 4 eggs and 1
young. Trees near creeks and ponds are preferred as nest¬
ing sites.
10. Nycticorax nycticorax naevius (Boddaert). Black-
crowned Night Heron. The “Quak” of this heron flying
to its feeding grounds is a familiar night sound. Owing
to its crepuscular and nocturnal habits, it may be over¬
looked in regions where it is common. Except during the
breeding season, the day is usually spent in trees or in the
midst of rushes. The first migrants arrive March 20 (Tay¬
lor) to April 15. It has not been noted in fall after Sep¬
tember 30. The birds separate following the nesting sea¬
son and are widely distributed from July 15 to September
1. A thriving colony has existed in a woods near Cooks-
ville for several years. The eggs are deposited mainly be¬
tween May 15 and 30, and the period of incubation is 24
to 26 days.
11. Botaurus lentiginosus (Montagu). Bittern. Com¬
mon summer resident. The generic name of this bird is
obscurely descriptive of its notes, and is perpetuated in the
vernacular by “Bog-Bull”. Certainly for the American
species the notes resemble far more the sound of the old
fashioned wooden pump than the bellowing of a bull. The
average date of arrival is April 16. My earliest record is
April 9, though Bent9 gives March 30. The latest date of
departure is October 28, 1923.
A nest found June 27, contained one addled egg and 2
small young. As soon as I approached, the young “hissed”
and crept slowly through the vegetation, where in a frozen
position they were very difficult to distinguish. Another
sound produced by clicking the mandibles together was ac¬
companied by pronounced distention of the chin.
12. Ixobrychus exilis (Gmelin) . Least Bittern. On the
basis of “intelligence” this species should be placed at the
foot of the bird class. One allowed my dog to take it alive
under inexcusable circumstances. The average date of ar¬
rival is May 18; the earliest May 10, 1914 (Betts). The
latest date of departure is September 23.
The nests are almost invariably placed on a platform of
8 North American Marsh Birds, Bull. 135, U. S. Nat. Museum, p, 83.
Schorger — Birds of Dane County , Wisconsin . 467
cat-tails. I have not found full sets of eggs before June 1.
On June 13, 1914, I examined sixteen nests in an area of
two acres. Seven contained 6 eggs, seven 5 eggs, and two
4 eggs each.
Family ANATIDAE. Swans, Geese, Ducks
13. Cygnus columbianus (Ord.) Whistling Swan. This
handsome bird is an uncommon migrant and has been ac¬
corded full protection by law to save it from extermina¬
tion. There are the following spring records: 6 seen on
Lake Mendota, April 5, 1925 by Taylor10; 4 on Lake Ke-
gonsa, April 10, 1926, and 40 on the following day by Dr.
W. V. Bryant; 7 on April 23, 1926 by Dr. S. H. Chase* 11;
and 3 on Lake Mendota March 27, 1927 by the writer.
The fall records consist of 7 seen on Lake Mendota Oc¬
tober 23, 1925 ; and an immature bird shot for a goose by a
Madison hunter Nov. 18, 1926. The latter bird was iden¬
tified by John Main.
14. Branta canadensis canadensis (Linnaeus). Canada
Goose. Common migrant, exceeding greatly in number all
other species of geese. This hardy bird is a not uncom¬
mon winter resident, especially on the Sauk prairie. The
earliest date of arrival in spring is March 6, the average
date being March 16. All have departed by April 18. The
Wisconsin River is the great resting place during the
spring migration, and on Lake Wisconsin “literally thou¬
sands^ of Canada Geese can be seen.12. The fall migration
takes place with great regularity. During eight of sixteen
years' observations, the first migrants appeared between
October 19 and 24. The southward movement is practi¬
cally completed by November 15.
15. Chen hyperboreus hyperborews (Pallas). Snow
Goose. The records for the Snow Goose are meager, and
probably misleading as to its numbers. The indications
are that the great majority of these geese pass through the
county without stopping. Gromme13 observed a heavy
10 Auk 43 (1926) 250,
11 Schorger, Auk 43 (1926) 556.
12 Stoddard, Yearbook Public Museum, Milwaukee for 1921 (1922) 36.
18 Auk 44 (1927) 96.
468 Wisconsin Academy of Sciences , Arts , and Letters.
flight over Lake Winnebago November 1, 1926. Strag¬
glers are occasionally reported by hunters as occurring in
flocks of Canada Geese along the Wisconsin River. I found
a lone Snow Goose on the shore of Lake Kegonsa, May 5,
1928. This bird, in sound condition, was shot the follow¬
ing day by John Main. The skin is in my collection.
There is only a remote chance for the occurrence of the
Greater Snow Goose ( Chen hyperboreus nivalis ), there be¬
ing but one good record for the interior.14
16. Chen caerulescens (Linnaeus). Blue Goose. Rare
migrant. The only record is a flock of 40 observed over
Lake Mendota by John Gundlach, December 10, 1927.
17. Anas platyrhynchos platyrhynchos Linnaeus. Mal¬
lard. Abundant migrant and common summer resident.
This hardy bird occasionally winters. It pushes north¬
ward in spring as rapidly as water becomes available. The
earliest record is March 5, 1921; average date of arrival
March 19. The fall flight is at its height the last two
weeks of October.
During August the young and old birds congregate in
favorite marshes, as many as one hundred having been seen
together. The wariness of the Mallard permits it to sur¬
vive in breeding grounds from which other species have
long since vanished.
18. Anas rubripes tristis Brewster. Black Duck.
When Brewster in 1902 described the Red-legged Black
Duck ( Anas rubripes rubripes) , he started a discussion on
the validity of a species upon which experts have continued
to disagree. Rubripes has bright coral red legs and toes,
in contrast with the yellow or orange-yellow of tristis , and
presumably occupies a boreal region north of the range of
tristis. Bent15 considers rubripes as the more hardy, ma¬
ture Black Duck.
The species breeding locally is unquestionably tristis
while rubripes may be sought among the late fall migrants
and wintering birds. Local hunters recognize the follow¬
ing: up to the middle of October the ducks shot have
greenish brown to yellow legs; there follows a hiatus of
14 Bent, Bull. 130 U. S. Nat. Mus. (1925) 173.
“Bull. 126 U. S. Nat. Mus. (1923) 64.
ScJiorger — Birds of Dane County , Wisconsin. 469
about a week, then from October 25 to November 20, or as
long as the marshes are open, the ducks shot are larger and
have orange legs. Personally, I have never seen a speci¬
men in the flesh that could be referred to rubripes. The
latest date on which I have shot a Black Duck is November
10, and the legs of this bird were bright yellow.
The Black Duck arrives in spring simultaneously with
the Mallard and does not depart in fall until the marshes
are frozen. A few winter regularly where the Yahara
River remains open. Flocks of 25 to 30 birds are common
in August. Mallards and Black Ducks intermingle while
feeding, but when flushed there is segregation according to
species.
19. Chaulelasmus streperus (Linnaeus). Gadwall.
Though formerly common, the Gadwall has become one of
the rarest of the native marsh ducks. It has been observed
in spring from March 27 to May 10, and in fall from Sep¬
tember 3 to October 18. Taylor16 saw four October 19,
1922. The average date of arrival in spring is April 13.
20. Mareca penelope (Linnaeus). European Widgeon.
While a rare species, the American records are too numer¬
ous to consider the European Widgeon a straggler. It has
probably become established as a breeding species. It
has been noted on three occasions at Madison in spring
only as follows : April 22, 1917, and April 3, 192717 ; April
14, 1918. 18 This species associates in migration with the
Baldpate to which it is closely related.
21. Mareca americana (Gmelin). Baldpate. The Bald¬
pate is an early migrant, the average date of arrival being
March 19. The earliest record is March 12, 1922. The
fall migration lasts from Sept. 30 to Nov. 27. The Bald¬
pate formerly bred in southern Wisconsin, but is now
known only as a transient.
22. Dafila acuta tzitzihoa (Vieillot). Pintail. The
sight of a flock of Pintails flying low over a marsh on a
March morning renders the observer oblivious to chilling
winds. The long neck, long tail, and white underparts of
the male produce the mirage of a frigate under full sail.
18 Auk 40 (1923) 339.
17 Schorger, Auk 35 (1918) 74; ibid. 45 (1928) 106.
18 Taylor, Auk 36 (1919) 277.
470 Wisconsin Academy of Sciences , Arts , and Letters .
This species arrives March 12 and is common by the last
week of the month. Most of the birds have departed by
April 15, but the occurrence of mated pairs is not uncom¬
mon up to the last week in May. This fact coupled with
the presence of mature birds in August is the only evidence
that the Pintail may breed, as formerly, in this region. In¬
clusive dates for the fall migration are Sept. 24 and Nov.
13.
23. Nettion carolinense (Gmelin). Green-winged Teal.
This handsome teal arrives as early as March 18 and is most
common during the first week in April. It has not been
observed after April 23. The fall migration lasts from
Sept. 16 to Oct. 22. One spent the winter of 1928-9 with
the Mallards kept by Prof. J. G. Dickson at the Nakoma
spring.
The flight of the Green-winged Teal is singularly swift
and erratic. When in good form, the evolutions performed
will take the conceit from any except the best shots. I
have had them appear suddenly over the decoys, do a figure
eight, and be gone so swiftly as to leave me with a cold gun.
24. Querquedula discors (Linnaeus). Blue- winged Teal.
This species is a later migrant than the Green-winged Teal.
Though it has been noted by March 25, the average date of
arrival is April 9. The period of greatest abundance is
April 23 to May 1. It has not been noted after October 27.
The Blue-winged Teal shares with the Mallard and Black
Duck the distinction of nesting regularly in the county.
Records: May 17, 6 eggs; May 28, 7 eggs; and July 2, a
nest with 10 eggs, 9 of which were hatched on July 12. The
three nests examined were on high ground 50 to 600 feet
from the water in the marsh. Cattle, aside from natural
enemies, are destructive to nests by trampling.
25. Spatula clypeata (Linnaeus). Shoveller. In 1921
and 1922 this species arrived March 12. Normally, the
first migrants appear March 27 to April 8, and the species
is common until May 26. The latest fall record is Oct. 16.
A female with six half-grown young was seen July 18,
1925. The Shoveller probably breeds more frequently than
this single record indicates.
26. Aix sponsa (Linnaeus). Wood Duck. This duck is
the loveliest of all our waterfowl. The accompanying at-
TRANS. WIS. ACAD., VOL. 24
PLATE 16
Fig. 1. Nest of Bluewinged Teal.
Fig. 2. Nest of Virginia Rail.
TRANS. WIS. ACAD., VOL. 24
PLATE 17
Fig. 3. Nest of Coot.
Fig. 4. Nest of Least Bittern.
Schorger — Birds of Dane County , Wisconsin.
471
tributes of gentleness and confidence in man led it far on the
road to extermination. It is gradually increasing in num¬
bers, though still one of our rarer species. The migration
dates are so few as to be subject to considerable revision as
further data accumulate. It has not been noted in the
Madison region before April 29, though it may be looked
for the last of March. The average date of arrival at
Heron Lake, Minnesota is April 4. The species is occasion¬
ally found in the marshes around Madison in August, and
occurs in greatest numbers during September and the first
week in October.
The Wood Duck has received the protection of the law
but many are shot unwittingly by hunters in the dim light
of early morning. This is especially true of the first day
of the hunting season when these ducks are entirely unsus¬
picious. On the morning of September 16, 1922, I was at
Mud Lake, Columbia County, a few miles from the Dane
County line. Twenty Wood Ducks were seen, six of which
were dead birds left on the water where they fell. This
useless destruction is a difficult problem with which to cope.
On July 11, 1926, 13 old Wood Ducks and 6 small young
were seen on the same pond. It undoubtedly nests spar¬
ingly in Dane County.
27. Nyroca marila (Linnaeus). Scaup Duck. This
species is a regular migrant on the large lakes, but is de¬
cidedly less numerous than its smaller relative. Judging
from sight records and the bags of hunters, there is one
marila to fifty affinis. Hollister19 has recorded the number
of ducks killed by his party, at Delavan Lake, from the fall
of 1892 until the fall of 1899. During this period there
were 20 Scaup and 182 Lesser Scaup obtained.
I was in a blind on Lake Kegonsa, Nov. 13, 1927, when
one Scaup was shot out of a flock of twenty. It arrives in
spring from March 12 to 25.
28. Nyroca affinis (Eyton). Lesser Scaup Duck. This
is the most abundant species of ducks. Rafts of 100 to 600
birds are not uncommon. It decoys readily and is killed
in large numbers. The average date of spring arrival is
March 20; the earliest March 5. It is common until the
19 Auk 37 (1920) 367.
472 Wisconsin Academy of Sciences , Arts , and Letters.
middle of May and the last stragglers, probably non-breed¬
ing birds, do not depart until May 22.
Fall migrants appear Oct. 20, and are abundant through¬
out November. The final departure is regulated by the
freezing of the lakes. During the winter of 1913-1914,
they remained until Jan. 10.
29. Nyroca collaris (Donovan). Ring-necked Duck.
This is a common migrant. It associates with the Lesser
Scaup, and their migrations are coeval. In the spring,
both species are found commonly on small ponds and
flooded marshes; but in fall their activities are confined
mostly to the larger lakes where like the other sea ducks
they dive to considerable depths for aquatic plants.
30. Nyroca americana (Eyton). Redhead. Though for¬
merly abundant, the Redhead is no longer common. Flocks
greater than twenty to thirty birds are seldom seen. This
species occurs usually in small numbers intermingled with
the Lesser Scaup. It arrives as soon as the lakes show open
water, the earliest date being March 11. Nearly all have de¬
parted by the last week of April, the latest date being May
14. The fall migration begins Oct. 5 and ends on the clos¬
ing of the lakes.
The decrease of this species is due to its being an excel¬
lent table bird and to the fact that on certain days it will
decoy with ridiculous ease.
31. Nyroca valisineria (Wilson). Canvasback. The
Canvasback with its Hanoverian profile is the aristocrat of
the duck family. While praised by epicures, it is in fact
no better in taste than the other members of the genus
Nyroca. This species attains large size. A male weighing
1648 grams was shot at Fox Lake, Nov. 8, 1928.
The average date of arrival in spring is March 18; the
earliest March 10, 1927. Most of the birds have departed
for the north by April 20, but stragglers have been noted
up to May 11. The fall migrants appear the middle of Oc¬
tober and remain in large numbers until the lakes freeze.
On January 10, 1914 there was a raft of 2000 Canvasback
on Lake Mendota between Picnic and Second Points.
It is gratifying to state that this fine bird appears to be
maintaining its numerical strength. Rafts of several hun¬
dred birds are common. This species loves to take to the
Schorger — Birds of Dane County, Wisconsin. 473
air on bright days. About 10 o’clock in the morning and
again at 8 o’clock in the afternoon all the Canvasback leave
the water and go “on parade”. There is nothing more in¬
spiring than the sight of flock after flock of these powerful
birds sweeping along the shore of the lake. The first ar¬
rivals decoy quite readily, but they soon become so wary
that flocks numbering more than a few birds will seldom
come in.
The Canvasback secures its food from the bottom of the
lake, hence is an expert diver. It is practically useless to
pursue a winged bird if the water is at all rough. After
swimming beneath the surface for a considerable distance,
the bird rises for air but only the bill appears above water.
The Canvasback is reputed to feed largely on eel grass or
“wild celery” ( Vallisneria spiralis Linn.). Stomachs of
birds shot at Fox Lake, Lake Mendota, and Lake Kegonsa
were filled largely with the propagating buds of this plant.
At Lake Koshkonong, however, Mr. H. L. Skavlem20 found
that 60 to 80 per cent of the food consumed consisted of the
buds of the pond-weed Potamogeton pectinatus Linn. He
further states that the Canvasback is comparatively indif¬
ferent to wild celery.
The above plants form a large portion of the aquatic
vegetation of Lake Mendota.21 Canvasback use Lake Men¬
dota year after year. Lakes Kegonsa, Waubesa, and Mo¬
nona follow in their order. In some years this species is
common on Lake Kegonsa, and again as in the fall of 1928,
very few birds are present. The effect on the food sup¬
ply of seining for carp has never been satisfactorily deter¬
mined. The Canvasback nests rarely in this county. Prof.
J. G. Dickson informs us that in June, 1927, a pair had a
nest with nine eggs near the Nakoma spring. Eight of the
eggs hatched and seven of the birds survived. In 1929, a
pair was seen frequently in Hammersley Marsh up to May
24.
32. Glaucionetta clangula americana Bonaparte. Golden¬
eye. Common migrant. A few always winter where the
creeks and springs provide open water. Hunters com-
20 Bull. Wis. Nat. Hist. Soc. 3 (1905) 168.
21Rickett, “A quantitative study of the larger aquatic plants of Lake Men¬
dota”. Wis. Acad. Sci. 20 (1922) 501-527.
31
47 4 Wisconsin Academy of Sciences , Arts, and Letters.
monly call it Winter Duck or Whistler. The Golden-eye
arrives early in March, the average date being the 11th.
The earliest date is March 2, 1924 when ten were seen.
The birds depart the second week in April and only a few
are found after the 18th. The southward movement in fall
seems to be controlled by the freezing of the northern lakes.
It arrives in the Madison region from Nov. 4 to 16, and re¬
mains until the lakes close.
The Golden-eye has a restless disposition and seems to
spend as much time in the air as on the water. It is not
gregarious, except when forced to become so by limited
open water. I have never seen more than 40 of this species
in one flock when the lakes were open, and this number is
exceptional. Ordinarily less than a dozen birds are seen
together. It does not decoy easily.
83. Charitonetta albeola (Linnaeus) . Buffle-head. This
handsome little duck is a regular migrant. It has de¬
creased so greatly that in point of numbers it is scarcely
more common than the Wood Duck. Protection from
shooting should be provided for a period of years. It is
ordinarily found in pairs, flocks of 8 to 12 birds being the
exception. It decoys easily for the hunter by whom it is
known as “Butter-ball”.
The Buffle-head is found in spring from March 22 to
April 25 and in fall from October 9 until the lakes close.
34. Clangula hyemalis (Linnaeus). Old-squaw. This
hardy northern duck is decidedly uncommon away from
Lake Michigan. It is difficult now to accept the statement
of Kumlien and Hollister22 that this species is “anything
but rare on most of the larger lakes”. Of the 1167 ducks
killed at Delavan Lake from 1892 to 1899, only 8 were Old-
squaw.23
One was flushed from a spring at Lake Wingra, Jan. 19,
1913. 24 Mr. John Gundlach informs me that one was ob¬
served daily on Lake Mendota from April 5 to 13, 1925.
On March 2, 1929, Prof. J. G. Dickson found a male
Old-squaw in the lagoon of the Nakoma Golf Club. The
22 Birds of Wisconsin, Wis. Nat. Hist. Soc. 3 (1903) 25.
23 Hollister, 1. c.
24 Schorger, Auk 43 (1926) 556.
Schorger— Birds of Dane County , Wisconsin. 475
bird was very unsuspicious and was photographed at close
range.
35. Melanitta deglandi Bonaparte. White-winged Scoter.
In view of the breeding of this species in North Dakota and
the Canadian prairie provinces, the White-winged Scoter
might be expected to occur regularly in Dane County. The
records as a matter of fact are few, for we seem to be on
the edge of the path of migration. The movement of these
ducks from the above breeding grounds is by way of the
Great Lakes, where it winters commonly, to the New Eng¬
land coast, and vice versa.
There are no spring records. I shot a pair at Picnic
Point, Lake Mendota, Oct. 30, 1910.25 Decoys appear to
be irresistible. On Nov. 20, 1927, I happened to stop at a
blind on Lake Mendota when three White-winged Scoters
came in to the decoys and escaped a fusillade unscathed.
After flying half a mile, one bird turned back, alighted
among the decoys and was killed. Taylor26 saw one Dec.
3, 1922.
The White-winged Scoter feeds preferably on shellfish
and its flesh is not a delicacy. The feathers are so firmly
attached that it is preferable for the mildly religious to be¬
gin by skinning rather than plucking.
36. Erismatura jamaicensis rubida (Wilson). Ruddy
Duck. Migrant in small numbers. It usually arrives
from April 1 to 15, and has stayed until May 10. April is
the accepted month for its arrival in the northern states
though Eaton 27 gives March 10 to 20 for New York. My
earliest record is March 15, 1914. The notes of Norman
De Witt Betts give three occurrences for this month:
March 30, 1913; March 14, 1914; March 31, 1916. The
southward movement takes place in October. I shot one
from a flock of four flying low over the Widespread, Oct.
16, 1910. It has not been noted after Nov. 13.
We are within the breeding range of the Ruddy Duck
and it may nest occasionally. An adult male was seen on
a pond near Verona July 16, 1927.
25 Auk 43 (1926) 556.
2a Auk, 43 (1926) 250.
27 "Birds of New York” 1 (1910) 226.
47 6 Wisconsin Academy of Sciences , Arts, and Letters .
37. Lophodytes cucullatus (Linnaeus). Hooded Mer¬
ganser. This Merganser is still a common migrant, but
its numbers have decreased greatly. Of the ducks killed
at Delavan Lake from 1892 to 1899, 13.4 per cent were
Hooded Mergansers.28 This percentage would not obtain
today. The male is a handsome bird and is seen at his best
during the spring courtship. One moment the head with
depressed crest rests on the breast; the next, the neck is
outstretched and the white crest outlined in black flashes
forth. It prefers swamps, streams, and ponds to the large
lakes. Its flesh is barely palatable, but since it decoys
readily, few hunters resist the temptation to burn their
powder.
This species arrives the middle of March (March 13,
1927) , and is most common the first week in April. It has
remained until May 6. It is a late fall migrant. It ar¬
rives in November and remains until the lakes close. The
earliest record is a female shot November 6.
There are two summer occcurrences. Two were seen on
one of the Verona ponds July 16 and 17, 1927. They were
feeding on small bull-heads (Amiurus) which were swal¬
lowed without difficulty. Betts saw one June 28, 1914.
38. Mergus americanus Cassin. Merganser. The sight
of the handsome male Mergansers uttering their hoarse
“kwerr” as they fly from one patch of open water to an¬
other renders one an easy victim to the belief that spring
is not “far behind”. This species is a powerful flyer and
expert diver. Its flight is so loon-like as to be deceptive
at a distance. It is sometimes shot over decoys placed on
a point, but this is due I believe to the directness of its
flight. I have never seen one show indications of having
been influenced by them. The hardy Merganser occasion¬
ally winters and pushes northward in spring ahead of all
the other wild fowl; hence it is difficult to state when mi¬
gration begins. The males arrive first. Mud Lake and
the Yahara River between Lakes Monona and Waubesa are
favored localities. At the latter place I counted 125 on
February 15, 1925. Rafts of 100 to 300 are to be found
from March 11 to 21. They leave correspondingly early
28 Hollister, Auk 37 (1920) 369.
Schorger — Birds of Dane County , Wisconsin.
477
and are seldom seen after the middle of April. In fall they
appear by October 21 and remain until the lakes freeze.
At this season they do not collect in large flocks.
39. Mergus serrator Linnaeus. Red-breasted Mergan¬
ser. Common migrant, more numerous than L. cuculiatus
but less so than M. americanus. Occurs in spring from
March 12 to April 25. It is most abundant from April 8
to 20. It arrives in the fall Nov. 13 and remains until the
lakes close.
This species nests regularly on the lakes in the northern
counties and especially about Green Bay.
Family ACCIPITRIDAE. Hawks and Eagles
40. Astur atricapillus atricopillus (Wilson) . Goshawk.
This freebooter from the north is an irregular winter
visitor. Its presence is due presumably to the scarcity of
grouse and rabbits within its natural range. An adult
Goshawk29 remained in the Lake Wingra woods from Jan.
1 to March 5, 1927. On one occasion it was flushed from
the skeleton of a rabbit, but its main diet during the above
period was Bob-white. The kills were invariably devoured
on a stump or log. Another was seen Jan. 30, 1928 near
Fitchburg. One was seen by Taylor30 on Jan. 24, 1923.
41. Accipiter velox (Wilson). Sharp-shinned Hawk.
This hawk is a migrant in small numbers. According to
my notes, it is present in spring from April 8 to May 14
and in fall from September 5 to October 7. Betts saw one
April 6, 1913.
42. Accipiter cooperi (Bonaparte). Cooper’s Hawk.
The Cooper’s Hawk is a common summer resident and ex¬
ceeds in number all other members of the family. The av¬
erage date of arrival is April 5 ; the earliest March 28. It
remains until October 24. This species winters rarely.
One was seen December 19, 1920.
Nests with eggs have been found from May 1 to June 6.
Mr. George French found eggs April 27, 1926. Most of
the nests contain full sets of eggs by May 15.
29 Schorger, Auk 44 (1927) 271.
30 Auk 43 (1926) 380.
478 Wisconsin Academy of Sciences , Arts, and Letters .
48. Buteo borealis borealis (Gmelin). Red-tailed Hawk.
Permanent resident. There is a well defined southward mi¬
gration of northern birds in October. This species ranks
third in number. Full sets of eggs have been found
from April 10 to 22.
44. Buteo borealis krideri Hoopes. Krider’s Hawk.
This is a western form of the Red-tailed Hawk, presumably
of regular occurrence in western Wisconsin in autumn.31
Taylor32 reports one seen in the spring of 1921 (Feb. 6).
He has the following unpublished sight records: Jan. 23,
1923; and Oct. 14, 1925.
I am not aware of a specimen having been taken in the
county. There are so great variations in the several races
of Red-tails, B. borealis borealis, B. borealis calurus, and B.
borealis krideri, all of which may occur in Wisconsin, that
their identification is frequently difficult even with the skins
in hand.
45. Buteo lineatus lineatus (Gmelin). Red-shouldered
Hawk. Known only as an uncommon spring migrant at
Madison from March 14 (Betts) to April 13. One was seen
by Betts Jan. 11, 1914. This species is of regular occurence
in the western part of the county and breeds in the Wiscon¬
sin River valley.
Mr. Alvin Cahn seems to have been more fortunate with
this species for he states33: “Not an uncommon visitor - .
A male collected had eaten two toads, a garter snake, and a
number of May beetles.”
46. Buteo platypterus (Vieillot). Broad-winged Hawk.
Common migrant and fairly common summer resident.
Arrives April 3 (Taylor) to May 1. The southward mi¬
gration is completed for the most part by October 1. This
is the least wary of all our hawks and it has been seen in
well wooded, populous sections of Madison.
A set of three eggs was obtained May 27, 192234. Mr.
George French took a set of eggs May 24, and found a nest
with young July 4, 1927.
31Kumlien and Hollister, “Birds of Wisconsin” p. 63.
32 Auk 39 (1922) 273.
33 “An Ecological Survey of theWingra Springs Region, near Madison, Wis¬
consin.” Bull. Wis. Nat. Hist. Soc. 13 (1915) 159.
34 Schorger, Auk 39 (1922) 574.
Schorger — Birds of Bane County , Wisconsin.
479
47. Triorchis lag opus sancti-johannis (Gmelin). Rough¬
legged Hawk. This hawk appears in November and, un¬
less driven south by heavy snows that render the procuring
of food difficult, remains in diminished numbers throughout
the winter. It is most common in spring during the last
two weeks in March. It has been seen as late as April 20.
Kumlien and Hollister35 record the finding of a nest in
the eastern part of Dane County in May, 1872. The female
had received a wound in the wing, that presumably pre¬
vented her from migrating.
48. Aquila chrysaetos (Linnaeus). Golden Eagle.
Rare migrant. Information on this species is extremely
meager. Captured eagles, reported as belonging to this
species, have proved on examination to be immature Bald
Eagles. Taylor reports one seen near Mazomanie October
26, 1928.
49. Haliaeetus leucocephalus leucocephalus (Linnaeus) .
Bald Eagle. The Bald Eagle is a fairly common visitor to
the Madison region. Most of the records are for the winter
months, just before the lakes freeze. I have seen it on the
following occasions: September 10, 1922; December 12,
1925 ; December 11, 1927 ; and January 10, 1928. Betts
saw one January 8, 1914. The following records are from
Taylor's notes : October 14, 1921 ; September 14, 1922 ; May
6, 1923; May 2, 1924 ; February 11, 1927; and May 27, 1928.
It is a not uncommon winter resident near Mazomanie
where it is able to obtain from the Wisconsin River its usual
diet of fish.
50. Circus hudsonius (Linnaeus). Marsh Hawk. . Com¬
mon summer resident ranking second to the Cooper’s Hawk
in abundance. A few winter regularly unless the snow be¬
comes too deep on the marshes. The average date of arrival
in spring is March 16. There appears to be no well defined
period to the fall migration, the decrease being gradual.
The Marsh Hawk is beneficial since it feeds largely on
small rodents. I have, however, seen it in the act of strik¬
ing down or devouring the following species of birds. Flo¬
rida Gallinule ( Gallinula chloropus cachinnans) ; Bobolink
( Dolichonyx oryzivorus) ; Swamp Sparrow ( Melospiza
35 “Birds of Wisconsin,” p. 64.
480 Wisconsin Academy of Sciences , Arts, and Letters .
georgiana) ; and a chicken. The latter was from the Uni¬
versity farm. From the band that it carried, it was found
to be 58 days old.
A full set of eggs has been found by May 8. Mr. George
French obtained a set of 5 eggs May 21, 1927. A second
clutch of the same pair containing the exceptional number
of 7 eggs was found June 12.
51. Pandion haliaetus carolinensis (Gmelin) . Osprey.
The Osprey is a fairly common migrant. It has been ob¬
served in spring from April 15 to May 24 ; and in fall from
September 6 to 25.
52. Falco peregrinus anatum Bonaparte. Duck Hawk.
This is one of our rarer hawks. The dates on which it
has been noted are insufficiently early or late to cover the
migrating period. It should arrive the middle of March
and remain until November, if not a permanent resident.
The Duck Hawk nests preferably on cliffs and there are
no suitable sites actually within the county. Three adjacent
places are used with more or less regularity: Ferry Bluff,
Sauk County, just across the river from Dane County; Dev¬
il's Lake, Sauk County; and Gibraltar Rock, Columbia
County.
53. Falco columbarius columbarius Linnaeus. Pig¬
eon Hawk. In my experience this is one of the rarest of the
hawks, appearing regularly in the state. I saw one April
14, 1928, and this is the only occasion on which identification
was positive. Taylor has seen it on the following occasions :
May 7, 1923; May 22 and October 9,1924; and April 27,
1925.
54. Falco sparverius sparverius Linnaeus. Sparrow
Hawk. This dainty falcon is a fairly common migrant, but
an uncommon summer resident. A pair bred in the Wingra
woods in 1914, and another pair on Picnic Point in 1928.
In July the young birds are quite noisy and readily betray
their presence. It is a rare winter resident, one being seen
February 9, 1913.
The Sparrow Hawk sometimes appears during the latter
half of March, but usually not until April. The average
date of arrival is April 9 ; the earliest March 9, 1913. The
fall migration begins in September and is completed by Oc¬
tober 18.
S charger— Birds of Dane County, Wisconsin.
481
Family TETRAONIDAE. Grouse
55. Bonasa umbellus umbellus (Linnaeus). Ruffed
Grouse. Permanent resident, most numerous in the west¬
ern half of the county. The actions of the Ruffed Grouse
of the wilderness border on stupidity, but no cleverer bird
exists than the grouse of civilized regions. In 1914 a brood
was reared where the city reservoir now stands, near Sun¬
set Point. This species is most widely distributed during
winter. One or two birds usually winter in the Eagle
Heights region, but disappear in late spring. A nest found
at Cross Plains May 30, 1925 contained 11 eggs.
56. Tympanuchus americanus americanus (Reichen-
bach). Prairie Chicken. Permanent resident. There
are indications of an influx of northern birds during the
winter months when flocks containing 40 to 50 individuals
are occasionally seen. The persistent draining of marshes
renders it difficult for the species to maintain its ground.
The large marsh at the north end of Lake Kegonsa is occu¬
pied permanently, and will probably remain so as it is un¬
likely to be changed by further drainage. While the va¬
rious marshes seem equally attractive, on some the Prairie
Chicken will not be found. No satisfactory explanation for
this situation can be advanced.
A nest found in one of the Lake Wingra marshes on April
26, 1914, contained two eggs.
Family PERDICIDAE. Bob-whites, Quails
57. Colinus virginianus virginianus (Linnaeus). Bob-
white. Common permanent resident. There has been no
apparent change in the number of Bob-whites during the
past fifteen years, though they have been protected by law.
This situation is due largely to lack of suitable cover as a
protection against natural enemies and the rigors of winter.
While normally roosting upon the ground, I have found
covies roosting in the dense foliage of spruce trees. Good
indigenous cover is a thick stand of blackberry (Rubus).
During seasons of deep snow the best protection is afforded
by the canopies formed by the wild grape overgrowing
small trees and shrubs. In a winter of heavy snow the Bob-
482 Wisconsin Academy of Sciences , Arts , and Letters .
white will remain in the vicinity of locust (Robinia
pseudacacia) thickets and feed largely on the seeds. Many
of the seed pods of this species remain on the tree until
spring.
58. Perdix perdix perdix (Linnaeus). Hungarian Part¬
ridge. This species has been introduced from Europe
where it is known as the Common or Grey Partridge. The
name accepted in this country has its origin in the circum¬
stance that the first birds were obtained from Hungar¬
ian dealers. The male is easily recognized by a horseshoe¬
shaped patch of chestnut on the lower breast. This par¬
tridge is partial to open and cultivated fields, and in other
habits resembles the Bob-white. The first liberation of the
species in Wisconsin was made in Waukesha County by Mr.
Gustave Pabst, who on February 6, 1929 wrote as follows:
“I started about nineteen years ago, both importing and
buying stock birds from abroad and from dealers in this
country. I have planted between five and six thousand
birds, covering a territory of twelve to fifteen miles, replant¬
ing same every second year after the first planting and cov¬
ering a total period of about six or eight years.”
A flock of about twenty was seen ten miles east of Madi¬
son in May, 1927, by the late Professor George Kemmerer.
Family PHASIANIDAE. Pheasants
59. Phasianus colchicus torquatus (Gmelin). Eastern
Chinese Ring-necked Pheasant. This bird was introduced
very recently. Mr. William Aberg liberated 35 birds at
Shorewood Hills in 1926. In April, 1927, the local chapter
of the Izaak Walton League liberated near Lake Wingra 22
birds that had been trapped in South Dakota. In the fall
of the same year, 160 birds were liberated in various parts
of the county. The League was instrumental in having 600
birds hatched from eggs liberated in 1928.
Family RALLIDAE. Rails, Coots, and Gallinules
60. Rallus elegans Audubon. King Rail. Fairly com¬
mon summer resident. The skill that so large a bird as the
King Rail displays in hiding in sparse cover is astonishing.
Schorger — Birds of Dane County , Wisconsin.
483
The earliest spring arrival for Madison is April 21, 191138
(by Roland E. Kremers). Betts observed it at Madison
April 23, 1916. It has been found at Delavan as late as
October 22, 1894, but in the Madison region not after Sep¬
tember 24.
61. Rallus virginiana Linnaeus. Virginia Rail. This
miniature of the King Rail is a common summer resident,
arriving the first week in May and becoming common by the
10th. The earliest arrival37 is April 28 (Taylor). The
latest fall record is the capture of an exhausted bird at Mad¬
ison on the night of October 21, 1913, during a premature
snow storm. A very interesting description of the food
habits of the captive has been given by Cahn38.
Sets of eggs have been found from May 30 to June 5.
The nest is usually placed in a tuft of marsh grass on a
hillock.
62. Porzana Carolina (Linnaeus). Sora. The jaunty
Sora is an abundant summer resident arriving usually the
last week in April. It is sometimes common by the middle
of the month as 10 were seen April 17, 1916. The earliest
date is April 10 (Taylor). The latest fall record is Octo¬
ber 23, 1921. The Sora is a hardy bird. Of two seen in a
frozen marsh on December 28, 1913, one was caught and
found to be minus the right wing.39 At the report of a
gun, it has the disconcerting habit of dropping to the
ground whether hit or not.
George French found two nests on May 30 containing 9
and 11 eggs respectively.
63. Coturnicops noveboracensis (Gmelin). Yellow Rail.
A specimen was taken May 2, 1920 by Mr. Warner Taylor
and this is the only known capture for the county. Bent40
gives a record for Madison for May 13, 1911. The Yellow
Rail is so secretive and difficult to flush that it is probably
much more common than the records indicate. It has been
found in Wisconsin from April 23 to October 13.
86 Bent, U. S. Nat. Mus. Bull. 135 (1926) 266.
87 Bent, vide supra , p. 300, gives April 19, 1917. This is an error and ap¬
plies to the Sora.
88 Auk 32 (1915) 91.
39 Schorger, Auk 31 (1914) 256.
40 U. S. Nat. Mus. Bull. 135 (1926) 325; record taken from Wisconsin
Arbor and Bird Day Annual for 1912.
484 Wisconsin Academy of Sciences, Arts, and Letters .
64. Gallinula chloropus cachinnans Bangs. Florida Gal-
linule. Summer resident, arriving the end of April, and
becoming common by May 10. The earliest record is April
16, 1927. It has not been noted in fall after October 8.
Nests with apparently full sets of eggs have been found
from May 30 to June 13. A nest found by G. French on
May 30 contained 13 eggs. Deposition of eggs begins the
middle of May. The young hatch irregularly and leave
the nest soon afterwards, at least on the slightest disturb¬
ance, as I have found only young recently hatched. Very
small young have been noted as late as August 11.
65. Fulica americana Gmelin. Coot. The “Mud-hen”
is a common summer resident. The wintering birds are
mostly crippled. It arrives usually the last week in March,
and is common to abundant by April 8. In 1922 it arrived
March 12 and was common on the 26th. In fall the Coots
begin to mass by Sept. 20, and rafts of one to five thousand
birds are not uncommon in October. It remains in num¬
bers, especially on Lake Mendota, until the ice forms. One
hundred were seen December 16, 1928.
Nests with eggs have been found from June 4 to 13.
Shore Birds
The county in general is not attractive to shore birds.
They prefer mud flats as feeding grounds. The beaches
that exist on the lake shores consist largely of sand; as a
result the shore birds are found mainly about the muddy
margins of flooded marshes and small ponds. The Wil¬
son’s Snipe prefers marsh land with standing water and
with vegetation sufficiently short that the view is unob¬
structed. There remain still areas of grazed marsh land
that attract this species in large numbers.
There have been two occasions on which shore birds have
been present in unusual numbers. In May 1915 a dredge
working above the Widespread created a level area of sev¬
eral acres of black mud and marl. Indian Lake near
Springfield Corners was drained the summer of 1927. On
July 17, on an area of about 20 acres of mud, there were
500 shore birds consisting mainly of Lesser Yellowlegs,
Solitary Sandpipers, Semipalmated Sandpipers, Least
Schorger — Birds of Dane County , Wisconsin. 485
Sandpipers, and Pectoral Sandpipers. Previous to being
drained this lake drew very few birds, so that it was not a
customary stop during migration. It is apparent that
those that chanced to pass within sight of the lake bottom
recognized it as a rich feeding ground and stopped accord¬
ingly. It is doubtful if birds beyond an air path two miles
wide were drawn down. If the same attractions had ex¬
isted elsewhere, a line running east and west through the
county would have contained over 10,000 shore birds.
Family CHARADRIIDAE. Plovers and Turnstones
66. Charadrius semipalmatus (Bonaparte). Semipal-
mated Plover. Common migrant. The average date of
arrival is May 9, and it becomes common by May 15. The
earliest date is May 3, 1914 (Betts), It has not been noted
after May 31.
This plover is less common in fall than in spring. The
extreme dates are July 30, 1922 and September 19, 1926.
Most of the birds pass south the last week in August.
67. Oxyechus vociferus vociferus (Linnaeus). Killdeer.
This noisy shore bird is an abundant summer resident, and
is the first of the family to arrive from the south. The
average date of arrival is March 17 ; the earliest March 5,
1921. Large flocks numbering as high as 150 birds are to
be found in September. They disappear gradually, the
average date of departure being October 25. Frequently
they do not depart until November, the latest date being
Nov. 10, 1923.
The eggs are usually deposited in May, but occasionally
they must be laid the latter part of April since tiny young
have been found by May 9. They have also been found as
late as July 18. The late dates undoubtedly represent a
second attempt as a result of the destruction of the first
nest. This species relies on concealing coloration and not
natural cover to protect the nest. The latter is usually
placed in a pasture, a recently ploughed field, or one about
to be ploughed. As a result the first set of eggs is fre¬
quently destroyed.
68. Squatarola squatarola (Linnaeus). Black-bellied
Plover. This plover is now an uncommon migrant. It
486 Wisconsin Academy of Sciences , Arts , and Letters .
was formerly abundant in migration on the virgin prai¬
ries of the state. Mr. H. L. Skavlem informed me that
this species and the Golden Plover were very tame and fol¬
lowed the plow like chickens, especially when prairie land
was being broken. Among hunters it had the euphonius
name of “Bull-neck”.
There are no more than a score of records for the Black-
bellied Plover during the past 15 years. Nearly all of
these fall between May 20 and 80. The earliest date of ar¬
rival is May 17, 1914 (Betts). There is but one fall record.
This bird, seen near De Forest on September 18, 1921, had
the black breast and belly characteristic of the full breed¬
ing plumage.
69. Arenaria interpres morinella (Linnaeus). Ruddy
Turnstone. The bizarrely colored Turnstone is an uncom¬
mon spring migrant but cannot be called rare. It has been
found from May 19 (Betts) to May 27. Mr. John Gund-
lach found 18 on the bar in University Bay May 27, 1925.
There are no fall records.
Family SCOLOPACIDAE. Woodcocks, Snipes,
Sandpipers, etc.
70. Philohela minor (Gmelin). Woodcock. Summer
resident. The Woodcock is an early migrant, reaching us
the latter part of March. The sight of this bird standing
in snow so deep that it can scarcely walk is incongruous.
The extreme dates are March 18, 1918 to October 21, 1921.
This species is most numerous in the western part of the
county, particularly along the Wisconsin River. It is sel¬
dom, however, that more than two or three birds are
flushed in a day. Removal from the list of game birds is
advisable.
The Woodcock nests frequently in the Wingra woods.
Here French found one with young May 22, 1926. Taylor
found a nest with 2 eggs near Mazomanie, May 24, 1925.
On June 5, 1927 I found a nest with 2 eggs at Cross
Plains. It was situated in the edge of a woods on a high
hill and within five feet of a well used cow-path.
71. Capella delicata (Ord). Wilson’s Snipe. This fine
game bird is an abundant migrant. It is not uncommon
Schorger — Birds of Dane County , Wisconsin.
487
as a winter resident about certain spring holes. Though
formerly a common summer resident, there is no satis¬
factory breeding record within recent years. It frequently
arrives the end of March, the earliest date being March 19,
1927; average date of arrival April 6. Nearly all have de¬
parted by the end of April, but individuals are frequently
found up to May 16. The average date of fall arrival is
Aug. 2; the earliest July 24, 1920. The birds do not be¬
come common until the last week in September. From this
it might be concluded that the earliest birds are local, were
it not that during the period of two months covering the
nesting season no snipe have been noted. The height of
the migration is reached in October when as many as 200
birds have been seen in one day. A gradual decrease in
numbers has taken place during the past 15 years. Strag¬
glers remain until the marshes freeze. The latest date is
Nov. 25, 1920.
The stomachs of birds taken in Dane County in autumn
were forwarded to the Biological Survey for examination.
The principal food is the white larva of a fly of the family
Dolichopodidae. Frequently a large portion of the food
consists of seeds of: sedge (Carex; Cyperus) ; bur-reed
(Sparganium) ; mare's tail (Hippuris vulgaris) ; buck bean
( Menyanthes trifoliata) ; and crowfoot (Ranunculus).
On Oct. 31, 1925, I shot a Wilson's Snipe along the shore of
Lake Mendota. As soon as the bird drifted in it was placed
on a sheet of paper, whereupon leeches ( Dina parva) be¬
gan crawling from its mouth. In all 39 entire leeches,
ranging from three-fourths to one inch in length, were ac¬
counted for subsequently. The size and nature of the meal
is surprising.
72. Numenius hudsonicus Latham. Hudsonian Curlew.
Rare migrant. Taylor saw one May 24, 1926. Early
in the morning of October 1, 1928, Mr. John Gundlach saw
three curlews alight on a mud bar in the Widespread. The
birds left before he could approach near enough for a shot.
They were very probably of this species. The Eskimo
Curlew is practically extinct. The Long-billed Curlew
was never common in Wisconsin and disappeared even from
Minnesota twenty-five years ago.
488 Wisconsin Academy of Sciences, Arts, and Letters .
73. Bartramia longicauda (Bechstein). Upland Plover.
This species is a summer resident in very small numbers.41
There are seasons when none have been observed. The
average date of arrival for southern Wisconsin is April
18.42 Taylor observed it April 11. The latest fall record
is Aug. 8, 1926.
74. Actitis macularia (Linnaeus). Spotted Sandpiper.
Common summer resident. The average date of arrival is
April 29; the earliest April 20, 1914 (Betts). The fall mi¬
gration is usually completed by September 20. The latest
date is October 10, 1926.
75. Tringa solitaria solitaria (Wilson). Solitary Sand¬
piper. This sandpiper is a common migrant, the average
date of arrival in spring being May 2 ; the earliest April 25
(Taylor). The average date of departure for the north is
May 18; the latest May 27, 1917. Its sojourn in the north
is short. It was seen by Betts July 4, 1913, and by the
writer July 7, 1929. The average date of arrival is July
18. The migration is completed on the average by Sep¬
tember 30, the latest date being October 8, 1926.
76. Catoptrophorus semipalmatus inornatus (Brewster).
Western Willet. Very rare migrant. There is but one
record, that of a bird found on the bar in University Bay
by Mr. John Gundlach, July 2, 1925. Taylor43 attempted its
collection.
77. Totanus melanoleucus (Gmelin). Greater Yellow-
legs. A regular migrant in small numbers. It is much less
numerous than T, fiavipes. The proportion one of the for¬
mer to seventy-five of the latter is not far wrong. The
average date of arrival is April 20; the earliest April 3,
1921. In 1914 it was observed as late as May 17 by Betts.
The early date (by Roland E. Kremers) of July 29 for ar¬
rival at Madison in fall is given by Bent.44 Nearly all the
records fall between Sept. 25 and Oct. 11. It is exceptional
to find a flock at any season numbering more than 6 to 8
41 Taylor, Auk. 39 (1922) 273; 40 (1923) 339; 43 (1926) 251; Schorger,
Ibid. 44 (1927) 235.
42 Cooke, Biological Survey, Bull. 35 (Revised), (1912), p. 66.
43 Auk 43 (1926) 251.
44 U. S. Nat. Mus. Bull. 142 (1927) 335.
Schorger— Birds of Dam County , Wisconsin .
489
birds. One was shot out of a flock of 8 Oct. 11, 1919, the
latest date on which it has been found.
78. Totanus flavipes (Gmelin). Lesser Yellow-legs,
Abundant migrant arriving on the average April 27. The
earliest date is April 2, 1916 (Betts). Ordinarily it is not
common before May 5, though 38 were seen April 14, 1928.
The average date of departure is May 22; the latest May
80, 1928.
Scarcely six weeks elapse before the note of the Lesser
Yellow-legs is heard again. The average date of arrival is
July 19, the earliest July 7, 1929. It is common from July
30 to Sept. 24. The average date of departure is Sept. 28 ;
the latest Oct. 15, 1927.
79. Pisobia maculata (Vieillot) . Pectoral Sandpiper.
This sandpiper is a common migrant. It is found fre¬
quently in cultivated fields and meadows as well as near the
water. On May 3, 1925, a flock of 200 was found in a par¬
tially ploughed clover field near Lake Kegonsa. The spring
arrivals are erratic, dating from the last of March to the
first of May. The earliest date is March 25, 1928. On
March 31, 1917 a flock of 20 was found on ice cakes in Uni¬
versity Bay. The average date of departure is May 19;
the latest May 26, 1928.
The average date of arrival in fall is July 23; the earliest
July 9, 1927. The average date of departure is Oct. 12;
the latest Oct. 18, 1916.
80. Pisobia fuscicollis (Vieillot). White-rumped Sand¬
piper. The “White-rump” is one of the rarer shore birds.
Taylor45 states: “On May 21, 1923, I collected one speci¬
men; on May 28, 1924, under favorable conditions I identi¬
fied six; on May 3, 1925, two; on May 30, I collected one,
and on June 5, three among a flock of a dozen or so.”
I have found it on the following occasions: one was
taken May 10, 46 and another seen May 20, 1914; two seen
May 22, 1915; three seen May 20, 1928; and one seen Aug.
12 and 14, 1926. Betts had the following records : May 9,
17, and 20, 1914; and May 22, 1915.
45 Auk 43 (1926) 250.
46 Auk 43 (1926) 556.
32
490 Wisconsin Academy of Sciences , Arts , and Letters .
81. Pisobia bairdi (Coues). Baird’s Sandpiper. This
sandpiper also is not common. It occurs much more fre¬
quently in fall than in spring. The only spring record is
one taken by Taylor47 June 5, 1925 from a flock of White-
rumped Sandpipers.
This species was found on nine occasions during the fall
of 1926 and 1928, between Aug. 8 and Sept. 15. All but
one of the records were for August. In 1926, I took a lone
female Aug. 15, and another female from a flock of four
Aug. 27.48
82. Pisobia minutilla (Vieillot). Least Sandpiper.
Common migrant whose average date of arrival is May 18.
The earliest date is May 3 (Taylor) . It is most numerous
May 15 to 20. The latest date of departure is May 27,
1928.
The average date of the return is July 16. The earliest
date is July 5, 1913 (Betts) . The migration is usually com¬
pleted by Sept. 15, the latest date being Oct. 7, 1928.
83. Pelidna alpina sakhalina (Vieillot). Red-backed
Sandpiper. This shore bird is common in spring, but rare
in autumn. Most of the birds appear to return from the
Hudson Bay breeding grounds by way of the Atlantic
coast. The average date of arrival in spring is May 13;
the earliest May 9, 1926. The height of the migration is
May 17 to 22. A flock of 75 birds was seen May 22, 1915.
The latest date of departure is May 31, 1914.
The fall records are scanty. The following dates are
given by Betts: Nov. 1, 1914; and Nov. 15, 1915. I have
found it on only one occasion, Nov. 13, 1927, when two were
collected at Lake Kegonsa.
84. Limnodromus griseus scolopaceus (Say). Long¬
billed Dowitcher. A specimen was referred to this form
by Dr. A. Wetmore. It is unlikely that the eastern bird ( L .
griseus griseus) will be found. Formerly an abundant mi¬
grant, now uncommon. The extreme dates of its occur
rence in spring are May 6, 1928, and May 22, 1915. The
average date of arrival is May 15. I have never found
more than 4 birds in one flock except on May 17, 1925 when
47 Auk 43 (1926) 251.
48 Auk 44 (1927) 262.
Schorger — Birds of Dane County, Wisconsin. 491
26 were seen.49 The only fall record is a male taken Aug.
12, 1926.50
85. Micropalama himantopus (Bonaparte). Stilt Sand¬
piper. The Stilt Sandpiper is a rare migrant. John Main
took one May 18, 1927, and this is the only spring record.
In the summer of 1926, I found this species on several
occasions as follows:51 8 were seen and 3 taken Aug. 7; 2
were seen and one taken Aug. 8 ; 2 were seen and one taken
Aug. 14; and a lone bird was taken Aug. 25. John Main
took one Sept. 12. All of the above records were obtained
on the same pond.
86. Ereunetes pusillus (Linnaeus). Semipalmated
Sandpiper. Common migrant. The average date of ar¬
rival is May 15; the earliest May 3 (Taylor). It is found
in largest numbers from May 28 to 31. Strangely, it has
not been noted after the latter date.
The extreme dates for the return migration are July 16,
1927 and Sept. 15, 1928. The height of the movement is
Aug. 14 to 28.
87. Limosa fedoa (Linnaeus). Marbled Godwit. The
Marbled Godwit was never common in Wisconsin and is
now merely a straggler from its principal range, which is
west of the Mississippi. Taylor52 has reported one seen
on May 18, 1922.
88. Crocethia leuchophaea (Pallas). Sanderling. The
Sanderling is an uncommon migrant. The data are too
few to furnish reliable limits to the migration periods. It
has been found in spring from May 11 (Taylor) to May 23
(Betts). My fall records extend from Aug. 26 to Sept. 15.
The latest record is Nov. 5, 1926, when Mr. John Gundlach
collected a Sanderling on Picnic Point, Lake Mendota.
The statement of Kumlien and Hollister53 that this spe¬
cies “frequents the sandy shores exclusively” cannot be con¬
firmed. I have seen it feeding on mud flats with other
shore birds, both at Lake Koshkonong and in the Madison
region.
49 Auk 43 (1926) 557.
00 Auk 44 (1927) 261.
01 Auk 44 (1927) 261.
62 Auk 40 (1923) 339.
53 Bull. Wls. Nat. Hist. Soc. 3 (1903) 48.
492 Wisconsin Academy of Sciences , Arts, and Letters .
Family, PHALAROPODIDAE. Phalaropes
89. Steganopus tricolor Vieillot. Wilson’s Phalarope.
The exquisite Wilson’s Phalarope is found in small numbers
during the migration period and probably breeds within
the county. Mr. Louis Sumner collected a set of eggs June
5, 1887 in the marsh at the fair grounds at Madison. Tay¬
lor54 saw two June 19, 1925 at Mud Lake (Jefferson Co.).
The earliest date of arrival is May 3 (Taylor). This
species leaves its breeding grounds early, the southward
movement being completed usually in August. It has been
taken on the extreme dates July 12 and Aug. 13.
The spinning motion which the Phalaropes undergo while
sitting on the water, is reputed to result in bringing to the
surface minute forms of animal life upon which the birds
feed. I watched a pair of Wilson’s Phalaropes May 14,
1928, feeding upon small insects hovering thickly over the
water. The birds went through precisely the same spin¬
ning motion in capturing the insects from the air. They
were merely observing a simple mechanical principle. By
pivoting, food can be seized most rapidly over a large area
with a minimum of effort.
90. Lobipes lobatus (Linnaeus). Northern Phalarope.
Rare migrant. Warner Taylor reports one seen May 20,
1917. I collected one Sept. 3, 1923 in a marsh near Madi¬
son, and another Sept. 24, 1927 at a pond near Springfield
Corners.55 The date Oct. 14, 1927 given in the latter ref¬
erence is an error.
Family LARIDAE. Gulls and Terns
91. Larus argentatus Pontoppidan. Herring Gull.
This species, though present throughout most of the year,
does not breed. It frequently winters where the Yahara
River remains open, and immature and non-breeding birds
are common throughout the summer. This gull arrives
early in March, the average date being the 13th. It is num¬
erous until the middle of April, occasionally until the middle
of May.
84 Auk 43 (1926) 250.
85 Auk 43 (1926) 556; 45 (1928) 106.
Schorger — Birds of Dane County , Wisconsin.
493
In fall it remains for the most part on the lakes, but in
spring hundreds may be found in cultivated fields and
around temporary pools. It is less common in fall than
in spring, arriving on the average Oct. 7 and remaining
until the lakes close.
92. Larus delawarensis Ord. Ring-billed Gull. Com¬
mon migrant, but less numerous than the preceding species
with which it associates. The average date of arrival is
March 30; the earliest March 15, 1913 (Betts). It usually
departs by May 20, but on May 29, 1927, a mixed flock of
immature birds consisting of 15 Herring Gulls and 35 Ring¬
billed Gulls wTas seen in a meadow near Lake Wingra. It
is occasionally found in July and August. It appears in
fall the middle of October, the earliest date being Septem¬
ber 20, 1914.
93. Larus pipixcan Wagler. Franklin’s Gull. Kum-
lien and Hollister56 state that in May, 1870 considerable
numbers of this species were seen following the plow near
Rockdale, Dane County. There is only one subsequent re¬
cord. Fragments of a bird of this species were found Ap¬
ril 23, 1911, on the shore of Lake Mendota. The wing by
which the species was identified at Washington is preserved
in the Biological Department of the University.57
94. Larus Philadelphia (Ord). Bonaparte’s Gull. This
gull is a fairly common spring migrant. The average date
of arrival is April 26; the earliest April 9, 1916 (Betts.)
It has not been noted after May 20. The fall migration
takes a southeasterly direction over the Great Lakes, and
records for the interior are scarce. I saw an immature
Bonaparte’s Gull at Lake Koshkonong August 26, 1928, but
this bird may have spent the summer there.
95. Sterna hirundo Linnaeus. Common Tern. The
information on the status of this species and Forster’s Tern
(S. forsteri) is very unsatisfactory. Identification by the
eye is unreliable. Each species has characteristic notes,
but too frequently they cannot be heard. S. hirundo utters
a harsh “tee-ar-r-r”, and S. forsteri a hoarse “zru-r-up.”
The Common Tern is the more abundant bird, but the mi-
56 Bull. Wis. Nat. Hist. Soc. 3 (1903) 10.
#7 Conover, Auk 29 (1912) 388.
494 Wisconsin Academy of Sciences , Arts , and Letters.
gration dates given below cannot be referred to this species
alone with certainty.
The Common Tern arrives on the average May 5, the
earliest date being April 22, 1928. It is common the last
two weeks in May, and remains until June 15. Individuals
are occasionally seen in summer. It has been observed in
fall from September 10 to October 14, but it is far less
common at this season than in spring.
96. Sterna forsteri Nuttall. Forsters Tern. There
is only one record backed by a specimen. Mr. Warner Tay¬
lor took one May 14, 1923.58 Based on identification by
note, Forster's Tern occurs mainly between May 17 and 27.
97. Sterna caspia Pallas. Caspian Tern. This large
tern is a fairly common migrant. There are the following
spring records: one seen May 8, 1913 (Betts) ; two seen
May 17,192259; one May 18, 1919; and two May 27, 1924
(Taylor).
The fall records are as follows : two seen September 18,
1915 (Schorger) ; seven seen September 12, 1925 (John
Gundlach) ; small flocks September 16, 1926 and September
10, 1927 ; and one bird August 30, 1928 (Harry Anderson) ;
two seen September 16, 1923 (Taylor).
98. Chlidonias nigra surinamensis (Gmelin). Black
Tern. Abundant summer resident. The average date of
arrival is May 6 ; the earliest April 24 (Taylor) . The aver¬
age date of departure in fall is September 12 ; the latest Sep¬
tember 20, 1914.
Nests with full sets of eggs have been found from May
30 to June 19.
Family COLUMBIDAE. Pigeons and Doves
99. Zenaidura macroura carolinensis (Linnaeus).
Mourning Dove. Common summer resident, occasionally
wintering. The average date of arrival is April 2 ; the ear¬
liest March 20, 1927. The average date of departure is Oc¬
tober 13.
The breeding season is extraordinarily long. I have
found nests with eggs from April 18 to September 12.
88 Auk 43 (1926) 250.
89 Taylor, Auk 40 (1923) 339.
Sciiorger — Birds of Dane County , Wisconsin.
495
Family CUCULIDAE. Cuckoos
100. Coccyzus americanus americanus (Linnaeus). Yel¬
low-billed Cuckoo. This species is a fairly common sum¬
mer resident that is heard more often than seen. As a rule
it is one of the latest of the migratory birds to arrive. It
is irregular in its movements, arriving one year the middle
of May, and in another not until the first week in June.
The earliest date of arrival is May 12 (Taylor). The av¬
erage date of arrival is about May 26. The average date
of departure in autumn is September 19 ; the latest Septem¬
ber 27, 1914 (Betts).
Nests with eggs have been found from June 5 to Aug. 5,
the majority the second week in June.
101. Coccyzus erythrophthalmus (Wilson). Black-billed
Cuckoo. This species is more numerous than the preceding
and arrives earlier, the average date being May 18. The
earliest date is May 10, 1927. The average date of de¬
parture is September 20 ; the latest September 26, 1926.
Nests with eggs have been found from May 31 to June 6.
Family TYTONIDAE. Barn Owls
102. Tyto alba pratincola (Bonaparte). Barn Owl.
Uncommon permanent resident. One was seen August 25,
1917. On October 1, 1924, five fully grown young were
taken from a gable at the State Hospital on Lake Men-
dota60. These were placed in the zoological collection at
Vilas Park and at the present date four are living. War¬
ner Taylor saw one near Oregon April 3, 1920.
Family STRIGIDAE. Horned Owls, etc.
103. Otus asio asio (Linnaeus). Screech Owl. Com¬
mon permanent resident. Eggs have been found April 11
(French) and young able to fly have been seen by June 11.
104. Bubo virginianus virginianus (Gmelin). Great
Horned Owl. Common permanent resident. Eggs have
been found from February 11 to March 16.
60 Schorger, Auk 42 (1925) 131.
496 Wisconsin Academy of Sciences , Arts , and Letters.
105. Nyctea nyctea - (Linnaeus) . Snowy Owl. Oc¬
casional winter visitor. One killed near Morrisonville was
brought to Madison to be mounted Nov. 30, 1921. In
February, 1922, a Snowy Owl spent ten days on the farm of
George Williamson on Lake Waubesa 61.
106. Strix varia varia Barton. Barred Owl. Perma¬
nent resident, more common in the woods of the Wisconsin
River bottoms than elsewhere.
107. Asio wilsonianus (Lesson). Long-eared Owl.
This owl is a fairly common permanent resident. It uses
the same roost continuously, if undisturbed, this being lo¬
cated in conifers when available.
On May 5, 1917 three eggs and two callow young were
found in an old crow’s nest. The sitting bird when flushed
uttered a “meuw” that brought its mate instantly. The
parents would drop to the ground and flutter along as
though crippled.
108. Asio flammeus (Pontoppidian) . Short-eared Owl.
Permanent resident. During the winter months concen¬
tration of local birds or migrants frequently takes place in
a marsh where field mice are numerous. On March 8, 1913
eleven were flushed from the marsh at the west end of Lake
Wingra. When the marshes are buried in snow, this owl
commonly spends the day in those oak trees near the
marshes that retain a large portion of their leaves.
109. Crypto glaux acadica acadica (Gmelin). Saw-whet
Owl. This tiny owl is a rare migrant. It may be resident
throughout the year, but there are no summer or breeding
records. One was captured at Madison, October 15, 1915
by Professor E. R. Maurer. The skin, No. 1169, was pre¬
served by Prof. George Wagner and is now in the Univer¬
sity collection. Mr. Warner Taylor on April 22, 1926
found a wing and foot of this species in the Wingra woods.
I took a male near College Hills, November 19, 1927.61a
Family CAPRIMULGIDAE. Whip-poor-wills,
Nighthawks
110. Antrostomus vociferus vociferus (Wilson) . Whip-
poor-will. Rather common summer resident, especially in
« Schorger, Auk 39 (1922) 574.
#ln Auk 46 (1929) 250.
Schorger — Birds of Dane County , Wisconsin . 497
the western portion of the county. The average date of
arrival is May 10; the earliest date April 21, 1923. The
average date of departure in fall is September 21 ; the latest
date October 1, 1918. The Whip-poor-will is vocal through¬
out the entire period of its presence.
A nest with one egg was found June 18. Another nest
contained a half-grown young June 17.
111. Chordeiles minor minor (Forster). Nighthawk.
Common summer resident migrating in large flocks. The
average date of arrival is May 14 ; the earliest date May 6,
1913 (Betts). The fall migration begins in August and
is completed on the average by September 10. The latest
date of departure is September 25, 1915.
A nest with 2 eggs was found June 12. The young are
able to fly by July 17.
Family MICROPODIDAE. Swifts
112. Chaetura pelagica (Linnaeus). Chimney Swift.
Common summer resident, arriving on the average April
24. The earliest date is April 9, 1917. The birds are num¬
erous on the date of arrival or shortly thereafter. The
final date of departure varies greatly. Some years none
are seen after the middle of September, in others they may
be found through the third week in October. The average
date of departure is October 6, the latest date being Novem¬
ber 4, 1922.
I have seen a nest built on the interior side of a barn.
Family TROCHILIDAE. Hummingbirds
113. Archilochus colubris (Linnaeus). Ruby-throated
Hummingbird. The Ruby-throat is a common summer res¬
ident. The average date of arrival is May 17, the earliest
date May 10, 1914. The average date of departure in the
fall is September 18 ; the latest date September 23, 1916.
Eggs have been found from July 13 to August 5.
Family ALCEDINIDAE. Kingfishers
114. Ceryle alcyon (Linnaeus). Belted Kingfisher.
The familiar rattle of the Kingfisher is to be heard the end
498 Wisconsin Academy of Sciences , Arts, and Letters.
of March or the first of April. The average date of arrival
is March 31 ; the earliest date March 16, 1919. The aver¬
age date of departure is October 27. It is occasionally
found near creeks long after the lakes are closed. The lat¬
est date is December 20, 1925.
Family PICIDAE. Woodpeckers
115. Colaptes auratus luteus Bangs. Northern Flicker.
Common summer resident, occasionally wintering. The
Flicker appears the end of March or the first of April, the
average date being March 28. The earliest date is March
16, 1919. The arriving Flicker invariably calls while the
wintering birds are usually silent. The average date of de¬
parture is October 19. November and December birds may
be final departures or winter residents.
Nests with eggs have been found from May 10 to 80.
116. Ceophloeus pileatus ahieticola (Bangs). Northern
Pileated Woodpecker. This magnificant woodpecker is a
permanent resident in the woods near the Wisconsin Riv¬
er62.
117. Centurus carolinus (Linnaeus). Red-bellied Wood¬
pecker. Fairly common permanent resident near the Wis¬
consin River. During the winter the range is extended.
It is rare in the Madison region during the summer, but not
uncommon at other seasons.
118. Melanerpes erythrocephalus (Linnaeus). Red¬
headed Woodpecker. Permanent resident whose numbers
are greatly reduced in winter. There appears to be no rela¬
tion between the abundance of mast and the number of win¬
tering birds. The migrants appear from April 25 to May
2, and the majority have departed by the first week in Sep¬
tember. Approximately as many birds of this species are
killed by automobiles as of all others combined.
Nests with eggs are found usually from May 30 to June
5.
119. Sphyrapicus varius varius (Linnaeus). Yellow-
bellied Sapsucker. Common migrant, wintering rarely.
One was collected Dec. 25, 192663. The average date of
«2 Taylor, Auk 39 (1922) 273; Schorger, Ibid. 44 (1927) 236.
68 Schorger, Auk 44 (1927) 262.
Schorger — Birds of Bane County , Wisconsin.
499
arrival is April 4 ; the earliest March 20, 1920. The migra¬
tion is at its height April 15, and is completed largely by
May 1, though stragglers have been noted as late as May 19.
The main movement in autumn takes place the last week
in September and the first week in October. The extreme
dates of arrival and departure are September 15, 1918 and
October 25, 1920.
120. Dryobates villoms villosus (Linnaeus). Hairy
Woodpecker. Common permanent resident. The opening
to the nest of this species is usually drilled through a shell
of live wood to reach a decayed interior. A nest found
May 13 contained 3 eggs.
121. Dryobates pubescens medianus (Swainson).
Downy Woodpecker. Common permanent resident.
Nests with eggs have been found from May 13 to June 5.
(To be concluded).
RAINFALL MAPS OF WISCONSIN AND ADJOINING
STATES
Eric R. Miller
The maps accompanying this paper are based on a new
computation of the average rainfall. Earlier maps and
discussions of the rainfall of Wisconsin are listed at the
end.
The data are derived from observations made by and un¬
der the supervision of the U. S. Weather Bureau, and with
standard rain gages. The observing stations are of two
classes, (1) regular stations, manned by paid observers,
and equipped with automatic recording gages, (2) cooper¬
ative stations, operated by public spirited citizens who per¬
form the duty voluntarily, equipped with non-registering
gages. The relative number and density of stations from
which data were used are given in the following table :
$ Including Omaha, Neb.
The period of 31 years from 1897 to 1927 inclusive was
taken as standard, and the records for only those years
were used for stations with longer records. Missing obser¬
vations within that period were estimated, and shorter
periods of observation at 46 places in Wisconsin, and 16 in
upper Michigan were extrapolated by comparison with
surrounding observation points.
No corrections have been applied, beyond the extrapola¬
tion just mentioned, and the elimination of typographical
errors.
502 Wisconsin Academy of Sciences , Arts , and Letters .
It is well-known that the gaging of rain is not independ¬
ent of wind, the catch diminishing the stronger the wind.
The diminution is still greater in the case of snow. The
regular stations of the Weather Bureau are generally lo¬
cated on high buildings in cities, and in plotting these
charts it was found that their catch in the warmer months
was less than at cooperative stations, which usually have a
ground exposure. In winter the professional observer ad¬
justs record of snow to correspond with the accretion of
fresh snow on the ground, while the cooperative observer
continues gage measurements. The cooperative observers
are found in general to have less winter precipitation than
the regular observers. As an example, the regular station
at Duluth has a 31-year average precipitation for January
of 1.01 inches, while the cooperative observer in the contig¬
uous city of Superior, has only 0.58 inches adjusted aver¬
age, based on 19 years of observations. On the other hand,
the Duluth average for the 6 months, April-September, is
18.70 inches, against 19.76 for Superior.
Although precipitation is neither increasing nor decreas¬
ing permanently, its yearly deviations from the average are
highly erratic. It is for this reason that care was taken to
average the same period of years at all stations. Beyond
this, the variations are local, especially in the warmer
months, whence it is easily conceivable that some of the fea¬
tures of these maps are accidental. It is intended to study
the dispersion of the data employed in these maps at some
later time, but in the meantime it is of interest to remark
that previous studies have shown that the average deviation
of the annual rainfalls rises from 10% of the average in
the Upper Peninsula of Michigan to 20% in southwestern
Iowa.
That the main features of the distribution of rainfall
shown by these maps are persistent can be verified by com¬
parison with figures 2 and 8 of Kincer, (4).
Most of the precipitation in Wisconsin and adjacent
states falls in connection with cyclonic storms. These
move from west to east, but on account of their centripetal
circulation and the western mountain barrier, their supply
of moisture comes from the Gulf of Mexico and the Atlantic
ocean. Decrease of rainfall with distance from source is
Miller — Rainfall Maps of Wisconsin and Adjoining States. 503
plainly seen on these maps, especially from Illinois and
southern Iowa to northwestern Minnesota. The import¬
ance of the Great Lakes depends upon the season. In win¬
ter these lakes remain open, so that they are relatively much
warmer than the land. Their vapor is then the source of
heavy snowfalls on lee shores, e. g. the northern shore of
the Upper Peninsula in map 3. A reach of wind across
more than 30 miles of water appears to be essential, as the
heavy snowfalls do not appear along the western end of
Lake Superior, until the prevailing northwest winds have
passed Bayfield Peninsula. In summer the heavy rains occur
mostly in summer thunderstorms, which must be regarded
as local convectional overturnings, although they mostly
occur during the passage cf large cyclonic whirls. In sum¬
mer, however, the Great Lakes are relatively cool, condi¬
tions adverse to local convection. Map 2 shows that the
summer rainfall is deficient around the lake shores.
The rising of air is considered by meteorologists to be the
cause of rainfall. This is paradoxical to the lay mind
which has become attached to the erroneous explanation
that the rain is due to contact with the cold upper atmos¬
phere. The correct explanation is that rising air expands
to equalize its internal pressure with the external pressure,
which diminishes with height. This expansion does work
at the expense of the temperature. Consequently the cool¬
ing goes on continuously at a rate proportional to the rate
of rising. We expect to find heavier rainfall where the as¬
cent is fastest. Ascent in cyclonic whirls and thunder¬
storms is not localized, but highlands cause localized ascents
of air, and it is this factor that explains the heavier rain¬
fall in western and northern Wisconsin, and the smaller
rainfall of the level lands extending from Green Bay up the
Fox River valley, down the lower Wisconsin valley, and up
and down the Mississippi valley. Additional discussion of
the rainfall distribution of this region will be found in the
papers listed below, of which Kincer must be mentioned as
containing an extensive bibliography.
504 Wisconsin Academy of Sciences , Arts , and Letters.
References
1. Bormann, W. R. Development of water power in Wis¬
consin and the relation of precipitation to stream flow.
(Map, average annual rainfall of Wisconsin) Monthly
Weather Review, 1913, p. 1020.
2. Climatological reports, Wisconsin, 1907, monthly and
annual summary. (Average monthly and annual rain¬
fall of Wisconsin.) Milwaukee, 1907.
3. Day, P. C. Precipitation of the drainage area of the
Great Lakes. (Maps). Monthly Weather Review.
1926, pp. 85-106.
4. Kincer, J. B. Precipitation and humidity. U. S. Dept.
of Agriculture, Office of Farm Management, Atlas of
American Agriculture, Part 2, Section A, Washing¬
ton, D. C. 1922.
5. Lapham, I. A. Rainfall of Wisconsin. Geology of Wis¬
consin, Vol. 2, 1877, pp. 35-44 and maps.
6. Seeley, D. A. The climate of Michigan and its relation
to agriculture. (Map, average annual rainfall, Michi¬
gan, p. 16) Rep’t State Board of Agriculture, 1917,
pp. 683-715.
7. Whitson and Baker. The climate of Wisconsin and its
relation to agriculture. Wisconsin Agricultural Sta¬
tion, Bulletin 223, 1912. 2d Rev. Ed. 1928.
Miller — Rainfall Maps of Wisconsin and Adjoining States .
505
83
Map. 1. Average annual precipitation, 1897-1927 inc.
506 Wisconsin Academy of Sciences, Arts, and Letters.
Map 2. Average warm season precipitation, April-September, 1897-1927 inc.
MiNN
Miller — Rainfall Maps of Wisconsin and Adjoining States.
507
Map 3. Average cold season precipitation, October-March, 1897-1927 inc.
508 Wisconsin Academy of Sciences, Arts , and Letters.
Map. 4. Percentage of the annual rainfall that falls in the six
warmer months, April-September.
TRANSMISSION OF SOLAR RADIATION BY THE
WATERS OF INLAND LAKES
E. A. Birge and C. Juday
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. XXXIV
Contents
1. Introduction _ 510-520
Acknowledgments _ 511
Instruments described _ 512
Calibration _ 514
Value in calories _ 514
Ratio, water-air — . — - - 514
With plate glass cover _ 514
With hemispherical cover _ 516
Adjustment for sun’s altitude _ 517
Zenith sun _ 518
2. Observations on lakes _ 520-564
General results _ 520-540
Data _ 520
Conclusions _ 525
Factors determining transmission _ _ _ 526
Illustrations of effect _ 527
Transmission in upper meters _ 534
Transparency and transmission _ 536
Lake Mendota _ 541-549
Data _ __ _ 541
Results _ _ 542
Table 5 _ 543
Transmission during warming period, mean sun_ _ 545
Calories delivered, Apl. 15 — Aug. 15 _ 546
Per cent of light transmitted _ 548
The Southeastern Lakes _ 549-554
General results _ 549
Groups of lakes _ _ 551
The Four Lake group _ 551
The Oconomowoc district _ 551
The Waupaca lakes _ _ _ 552
Elkhart Lake __ _ 553
Geneva Lake _ 553
Green Lake _ 554
Devils Lake _ 554
510 Wisconsin Academy of Sciences, Arts, and Letters.
The Northeastern Lakes _ _ _ _ _ 555-558
Data — - - 555
General results _ 556
Other Lakes _ 558
Finger Lakes, New York _ 558
Okoboji Lake, Iowa _ 558
General Tables _ 559-563
Table 10, transparent lakes _ 559
Table 11, all lakes _ 560
3. Comparison with work of other observers _ 564-574
Total radiation compared with light _ 564
The selenium cell, Regnard _ _ 566
Photographic methods - 566
Linsbauer, Traunsee _ 566
Klugh, Bay of Fundy, Chamcook Lake _ 567
Oberdorfer, Bodensee _ 568
Photoelectric cell _ 569
Shelford and Gail, Puget Sound _ 569
Compared with Crystal Lake _ 570
Poole and Atkins, ocean near Plymouth _ 570
Shelford and Gail, Lake Mendota _ 572
General results _ 572
4. Literature cited _ 574
5. Appendix, Computation of mean path of radiation _ 576-580
1. INTRODUCTION
This paper deals with the amount of solar radiation de¬
livered to a unit of horizontal surface at various depths in
the upper water of lakes. The present paper states the re¬
sults obtained and discusses their relation with similar ob¬
servations made by other methods. The discussion of other
conclusions which naturally come from the data is deferred
to a later paper.
The study began in 1900 in a tentative way, employing a
black-bulb thermometer in vacuo as the instrument. This
instrument and its use have been described and illustrated
(Birge 9 22 , pp. 539-542). The rate of transmission was
determined to a maximum depth of five meters and numer¬
ous observations were made in Lake Mendota during 1900
and 1901 and visits were made to eleven other lakes. None
of these readings are used in the present paper, since they
do not add to the knowledge obtained later. The results
Birge & Juday — Transmission of Radiation in Inland Lakes. 511
are in general agreement with those here reported and are
from lakes also reported here.
After 1901 the study was discontinued until 1912, when
the first pyrlimnometer was constructed. This instrument
has been described and illustrated and, with certain im¬
provements and modifications, it is still in full use. Ail ob¬
servations reported in this paper have been made with it.
Its essential part is a thermopile, whose electric current,
when exposed to the sun's radiation, is measured by a gal¬
vanometer or millivoltmeter.
In the tables of this paper the amount of radiation re¬
ceived below the surface of the lake is reported as a per cent
of the total radiation delivered to the thermopile when held
horizontally just above the surface of the water at the time
of observation. Radiation is stated in calories per square
centimeter per minute.
The unit of depth usually employed is the meter. The
coefficient of transmission is the per cent of radiation de¬
livered at the upper surface of the meter stratum in ques¬
tion which is found at the lower surface of the stratum.
This is called “the transmission," as a short title ; a “trans¬
mission of 63 in the 1-2 m. stratum" means that the swing
of the galvanometer at 2 m. is 63 per cent of the swing at
1 m. The coefficient of absorption of the stratum is the dif¬
ference between the transmission and 100 per cent. In the
example given the coefficient of absorption is 37 per cent.
In this paper values are given in terms of transmission.
The transmission as stated in the tables is computed for
zenith sun (see p. 517), and is stated to the nearest whole
per cent. The transmission of the upper meter is given
to the nearest tenth of one per cent, although there is no
reason to think that much value attaches to the last figure.
In computing results the fractional per cent was used and
the final results were rounded off.
Acknowledgments
We have received valuable aid from many persons, with¬
out whose assistance our work could not have been done.
Especial thanks are due to the members of the department
of physics, University of Wisconsin. From Professor C. E.
512 Wisconsin Academy of Sciences , Arts , cmd Letters.
Mendenhall we have received much counsel ; Professor J. R.
Roebuck has determined the transmission of various glasses
and ray filters; Professor L. R. Ingersoll has given un¬
wearied aid and counsel in our innumerable difficulties,
Mr. E. R. Miller, of the United States Weather Bureau, has
constantly helped us in all the many matters relating to his
department, and has also taken part in observations on
lakes. In all mathematical questions constant recourse has
been had to Professor H. W. March, of the department of
mathematics, University of Wisconsin. We are indebted to
Mr. F. E. Fowle of the Smithsonian Institution and Dr. H.
H. Kimball of the U. S. Weather Bureau, Washington, D. C.
for help and counsel.
The thermopiles with which we have worked and the gal¬
vanometer which we used until 1926 were constructed by
Mr. J. P. Foerst of the University department of physics.
INSTRUMENTS
The pyrlimnometer has already been described and illus¬
trated and the description need not be repeated. See Birge,
’22, pp. 543-547. The modifications and improvements since
1922 are as follows:
1. In the original instrument the thermopile was made
from iron-constantan couples. In 1926 a new thermopile
was made from bismuth-silver couples. This gives for equal
radiation about twice the swing of the iron-constantan ap¬
paratus. It was feared that the bismuth wire would prove
too brittle for safe transportation, but no accident has hap¬
pened, although the instrument has been carried several
thousand miles over roads by no means exceptionally good.
For carrying about the country it is placed on an inflated
air-cushion in a basket.
2. The electrical current from the thermopile was meas¬
ured by a D’Arsonval galvanometer (Birge, ’22, p. 544).
At first this was taken out in the boat, but while the instru¬
ment is not disturbed by motion parallel to the axis of the
galvanometer, it is sensitive to motion at right angles to
this axis. While, therefore, it was not difficult in a small
lake to find places for observation which were sheltered
from the wind, it was not possible to escape the effect of
Birge & Juday — Transmission of Radiation in Inland Lakes. 513
swells caused by the movement of launches, etc. It was
found better, therefore, to keep the galvanometer on shore
and connect it with the boat by a long cable (l.c., p. 545).
In 1926 the galvanometer was replaced by a Rawson milli-
voltmeter. This instrument has a scale of 100 divisions and
four ranges, respectively 20 mv., 10 mv., 5 mv., and 2 mv.
The swing of the needle on the 2 mv. range is about 2.5
times as great as on the 10 mv. These are the two ranges
most used. Full sunlight at or near noon gives a swing on
the 10 mv. range of about 80 divisions of the scale, equiva¬
lent to about 200 divisions on the 2 mv. range. The 2 mv.
range is used for all readings in water at or below one meter,
except in the case of the most transparent lakes, where if
may be necessary to employ the 5 mv. range for the upper
meters. This instrument is rapid in action and steady when
used in a boat. Radiation can be followed with it as far
down as one per cent of the amount in the air. In 1928 a
more sensitive millivoltmeter was also used. This has a
scale of 100 divisions and two ranges, 2 mv. and 0.333 mv.
The last range gives six times the swing of the 2 mv. range
and radiation can be followed correspondingly further. It
is steady when used in the boat but the current is so feeble
at considerable depths that the movement of the needle is
slow. The instrument has been chiefly used to secure read¬
ings at moderate depths in highly colored lakes, in which
radiation is very rapidly absorbed, and also for readings
designed to follow the course of light of various colors, ray-
filters being placed over the thermopile.
3. In the observations from 1913 to 1925 inclusive the
thermopile had a hemispherical glass cover, not unlike that
of the Callendar sunshine recorder or the Kipp and Zonen
solarimeter. The bismuth-silver thermopile, constructed in
1926, was provided with a flat plate glass cover, so that
ray -filters could be used with it. It appeared that the ratio
between the reading in air and that at any given depth
differed with the form of the cover, the hemispherical cover
giving a decidedly lower ratio than the flat one. This led
to a study of this ratio, the results of which are reported
later.
4. In general it may be said that the instrument as now
constructed serves well the purpose for which it was de-
514 Wisconsin Academy of Sciences, Arts, and Letters.
signed. The radiation can readily be followed to a depth
where the result is sufficient for general biological purposes
in an inland lake. The apparatus is simple and easily
handled; it can be easily transported; it is not heavy and
can be used from a rowboat. Observations are made rapidly
and with sufficient accuracy.
CALIBRATION OF INSTRUMENTS
There are two calibrations to be made on these ther¬
mopiles: (1) the value in calories of the scale readings, and
(2) the ratio between the readings in air and in the water.
1. The first calibration is easily made. The station of the
United States Weather Bureau at the University of Wis¬
consin keeps continuous records of radiation from sun and
sky by means of a Callendar recorder. Very frequent com¬
parisons of readings of the thermopile with those of the
Callendar have been made. When the thermopile has a
hemispherical glass cover no further comparisons are neces¬
sary. When a fiat glass cover is used the effect of the
angle between radiation and glass must also be determined.
2. The correct ratio between the reading in air and in
the water is more difficult to determine. The thermopile is
sensitive to radiation of every wave length, but water is
highly selective in its power of transmitting radiation. The
glass cover of the thermopile is also selective toward radia¬
tion and it behaves differently in air and in water. The
solar radiation is mainly direct, from the sun, but a large
fraction is diffuse, coming from the sky, and the two kinds
behave differently both in regard to reflection and to trans¬
mission. Hence arises a complex condition, to which may
be added the fact that the flat glass cover behaves differ¬
ently from the hemispherical one both in air and in water.
The calibration was done with the flat cover, since experi¬
ment was more easy with that than with the curved surface.
The standard depth of water used for comparison with the
air is one meter, since at that depth the radiation coming
from wave lengths beyond 0.762/a has been almost com¬
pletely absorbed and no further consideration need be given
to it. The following facts were determined: (a) the plate
Birge & Juday — Transmission of Radiation in Inland Lakes. 515
glass cover cuts off 18-19 per cent of the direct radiation
at normal incidence in air. This was first determined with
a Marvin pyrheliometer by Mr. Eric Miller, who is in
charge of the United States Weather Bureau Station at the
University. His results ranged from 17.8 to 18,9 per cent
and averaged about 18 per cent. This result was used in
the paper given at the meeting of the Geophysical Union in
Washington, April 26, 1928 (Birge and Juday, ’29, p. 64).
This work was done not long before going into the field for
the summer of 1928 and under this condition we did not
wish to dismount any of the thermopiles for the study.
After returning from the field one of the iron-constantan
thermopiles was uncovered and mounted as a pyrheliome¬
ter. It appeared from numerous experiments that the
plate glass cut off an average of about 19 per cent of the
direct radiation rather than 18 per cent as found in the
Marvin instrument. This value has been used in all results
except those already reported at Washington. Since the
difference is small and quite within the ordinary variation
of sunshine from minute to minute, it did not seem neces¬
sary to correct the figures already given.
The behavior of the cover below the surface of the water
could not be directly observed since it is not possible to use
an uncovered thermopile in the water. Determination of
the transparency of the glass showed that there is no ap¬
preciable absorption of radiation at wTave lengths which are
left in the water at a depth of one meter. At any rate, no
more would be absorbed than by clear water of the same
thickness as the glass, so that no special allowance need be
made for it. The refractive index of the glass is 1.52, cor¬
responding to a reflection of 4.26 per cent of radiation at
normal incidence. This amount would not exceed 5 per
cent until the angle of incidence exceeds 40°.
The result for the plate glass used in water at one meter
or deeper is as follows : little or no allowance need be made
for reflection from the upper or glass-water surface of the
cover; there is a reflection of 4.26 per cent or somewhat
more from the lower or glass-air surface of the cover; lit¬
tle or no allowance need be made for absorption of radia¬
tion in passing through the glass. It was therefore de¬
termined to allow a total loss of 5 per cent for the effect of
516 Wisconsin Academy of Sciences f Arts , and Letters .
the glass cover in water at depths of one meter or more.
At smaller depths there would be a greater loss correspond¬
ing to the depth and to the amount of long wave radiation
left in the total ; but in no case would the absorption of such
radiation by the glass exceed that of a corresponding thick¬
ness of water, so that the allowance of 5 per cent will still
be reasonable. Thus the reading of the thermopile in air
is determined as 81 per cent of its true value and that in
water as 95 per cent of its value; and the observed swings
of the millivoltmeter are corrected accordingly.
It will be noted that this calibration deals with the direct
radiation only, diffuse radiation being neglected. No way
was found of dealing with radiation from the sky separ¬
ately from that direct from the sun. The amount of dif¬
fuse radiation reflected from the glass cover and from the
surface of the water is different from the amount of di¬
rect radiation reflected. The mean path of diffuse radia¬
tion in water is also greater than that of direct, so that a
greater per cent will be absorbed in reaching the depth of
one meter below the surface. Thus the percentage stated
as present at one meter can not be exact and is probably
somewhat too great. The error can not be very large,
however ; and even a considerable error in determining the
amount at one meter makes very little difference in the gen¬
eral result.
The hemispherical cover was calibrated by comparison
with the results reached by experiment with the plate glass
cover. The curved surface does not lend itself to direct
experiment as well as the flat one. One of the iron-con-
stantan thermopiles was repeatedly compared with the
other, when both were provided with plate glass covers.
Then one was furnished with a hemispherical cover and in
this condition was compared with the other. Its readings
were also compared with those taken with the plate glass
cover by alternately placing the two covers on the same
thermopile. In this way the difference between the two
covers in air was determined. For comparison in the
water the two thermopiles with different covers were
mounted in the same frame and were read alternately at
different depths in the water of Lake Mendota. The re¬
sult was to show that the hemispherical cover gave higher
Birge & Juday — Transmission of Radiation in Inland Lakes . 517
readings in air and lower in water than the flat cover. This
would be expected since the form of the dome and the thin¬
ner glass of which it is made would allow a greater pene¬
tration of radiation in air. In the water the concave lens
caused by the water-glass surface would lower the reading.
It was found that the readings of the thermopile with the
hemispherical cover could be brought into agreement with
the corrected readings of the other instrument, by dividing
the reading in the air by 1.5. All observations made be¬
tween 1913 and 1925, inclusive, have been thus adjusted.
This adjustment of readings concerns only the relation
of the reading in the air with that in the water. Either in¬
strument gives the same ratio between readings both of
which are made in the water. Thus the transmission in
water is the same with either form of thermopile; adjust¬
ment is needed to carry these readings in water over into
the air so as to determine in calories the value of the trans¬
mitted radiation.
Adjustment for Altitude of Sun
The amount of radiation in the water of the lake is meas¬
ured at certain definite distances below the surface, one
meter, two meters, etc. The distance through which radi¬
ation will have traveled in order to reach this depth varies
with the altitude of the sun, and since the distance varies,
the amount of absorption varies correspondingly. If the
sun is only 5° above the horizon its direct rays pass
through 1.505 m. of water in reaching the depth of one
meter below the surface. They have passed through one
meter of water at the depth of about 66.4 cm. This is an
extreme case and is a condition under which observations
on the sun would not be made. But many observations will
be made when the sun is at altitudes between 40° and 50°
and the radiation has passed through one meter at depths
between 82 cm. and 88 cm.
In all cases therefore, it is advisable to reduce observa¬
tions to a standard altitude of the sun and the best altitude
is that of a zenith sun, since from this the situation at any
other elevation can be readily derived. The method em¬
ployed is to plat the observations, reduced to percentages
518 Wisconsin Academy of Sciences , Arts , and Letters.
of radiation in air, on coordinate paper, to join the points
thus ascertained by a line, and to mark on this curve the
depths corresponding to the successive cosines of the angle
of refraction. Mathematical methods may also be used.
(See Shelf ord and Gail, '22. )
In practice the altitude of the sun is taken from a large
diagram on which is drawn a series of curves giving the al¬
titude for each half hour between 8 :30 a. m. and 8 :30 p. m.
and from April 1 to October 15. The altitude for inter¬
mediate times, such as 9:48 a. m. is estimated from the dis¬
tance between the curves. No attempt is made to get the
altitude closer than the nearest degree. The corresponding
distance to be employed is taken from a diagram in which
the cosine of the angle of refraction in water is platted
against the altitude of the sun. This is easily read to 0.1
cm., but no attempt is made to use distances smaller than
one centimeter.
As coordinate paper for platting transmission curves
there can be used either standard paper or semi-logarithmic
paper. The latter is the more convenient since the angle
made by the transmission curve with the lines represent¬
ing depth or per cent varies only as the transmission per
cent varies, while in standard cross section paper this angle
depends also on the number of units gained or lost. (See
figs. 2 and 8.) Thus the curve comes to run nearly parallel
to the depth lines near the surface when many units are
lost by absorption ; and when the radiation is reduced to a
small amount the curve runs nearly parallel to the per cent
lines, although in each case the transmission may be prac¬
tically the same. It is impossible to mark the curve ac¬
curately if drawn on ordinary coordinate paper.
An illustration of the effect of reducing observations to
zenith sun may be taken from two series made on Lake
Mendota, July 10, 1916, as given in table 1 and figure 1.
The first series was made at 12:20-12:40 p. m. and the
second at 4:20-4:35 p. m. In the first case the sun’s alti¬
tude was 68.5° and the corresponding cosine of the angle
of refraction is 96; at 4:30 the altitude was 31.8°, whose
cosine is 77. No readings were taken above the depth of
one meter and the transmission was fairly uniform between
one meter and three meters. The percentages found by
Birge & Juday — Transmission of Radiation in Inland Lakes. 519
observation have been platted and the transmission curves
have been marked for the corresponding cosines (fig. 1).
The two curves are quite different but when adjusted for
zenith sun they show that the transmission was practically
identical for the two series.
Fig. 1. Observations on Lake Mendota, July 10, 1916. For de¬
tails see text. Small circles mark the percentages observed. An in¬
verted V marks percentages as corrected for zenith sun. In the noon
series (A) the radiation passes through one meter of water in the
vertical distance of 96 cm.; at 4:30 (B) the corresponding distance
is 77 cm. The curves are marked accordingly and the dots show that
substantially equal percentages of radiation are transmitted by each
meter of water, if zenith sun is assumed; while the direct observa¬
tions seem to show a much lower transmission in the afternoon.
Table 1. Lake Mendota , July 10, 1916
Depth, Per cent, Noon Per cent, 4:30 p. m.
meters, Observed Zenith sun Observed Zenith sun
Surface _ _ _ 100 100 100 100
1.0 16.2 16.7 14.1 17.7
2.0 7.5 8.2 5.0 8.2
2.5 4.9 6.5 2.9 5.4
3.0 3.3 3.7 1.7 3.5
The transmission in the 1~2 m. stratum was 47 at noon
and 35 at 4 :30 ; in the 2-3 m. stratum it was 44 and 34, re¬
spectively. When adjusted to zenith sun the transmission
became 48 and 47, and 44 and 43, or substantially the same
in both cases. At noon the incident radiation would have
been reduced to one per cent at the depth of 4.5 m., at 4 :30
520 Wisconsin Academy of Sciences , Arts, and Letters.
the same per cent would have been found at 3.5 m. When
adjusted for zenith sun both series gave 4.8 m. as the depth
for one per cent.
In this case, as in others, all radiation is treated as com¬
ing direct from the sun. No account is made of diffuse
radiation because it was impossible to determine the amount
of this under the conditions of observation. It is even less
possible to determine how the origin of the diffuse radia¬
tion is distributed over the sky and thus determine its av¬
erage angle of incidence on the water. The omission of
this factor makes the cosine somewhat larger than the true
value, and correspondingly decreases the correction applied
to the percentages observed at any given depth.
Thus in the general tables the results of observation are
corrected for zenith sun and on the assumption that all
radiation is direct from the sun.
2. OBSERVATIONS ON LAKES
There are reported in this paper 36 series of observations
in the upper water of Lake Mendota and 96 series on 55
other lakes. Of these lakes 26 are in South eastern Wis¬
consin, 25 in Northeastern Wisconsin, 3 in New York, and
1 in Iowa. The results are given in table 11 at the end of
this section; the general results are summarized and illus¬
trated in the following pages.
GENERAL RESULTS
Data
Table 2 and figures 2 and 3 show the transmission of
radiation through the water of a series of lakes which il¬
lustrate all types found in Wisconsin. Table 2 gives four
series of data from each lake, in order to illustrate the kind
of data obtained and the methods of handling them. The
first line under each lake gives the swing of the galvanome¬
ter or millivoltmeter, stated in divisions of the scale and ad¬
justed as described on p. 515; the second line states the per
cent of radiation found at each depth, taking the incident
Birge & J uday— Transmission of Radiation in Inland Lakes. 521
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522 Wisconsin Academy of Sciences , Arts, and Letters ,
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Meters
Birge & J uday — Transmission of Radiation in Inland Lakes. 523
radiation in air as 100 ; the third line gives the per cent as
corrected for zenith sun. These last are the values used in
the regular tables and diagrams. The transmission for
each meter stratum is derived from them by dividing the
Per Cent Incident Radiation
Fig. 2. Transmission curves for various lakes. In general the
curves are started at one meter depth, but in a few cases begin at
50 cm. in order to show the course of the curve in the upper water.
They are carried down in the intersection with the line indicating
one per cent of incident radiation. Where the extension is long the
accuracy is correspondingly uncertain.
In a diagram platted on regular coordinate paper the length of the
abscissa to the intersection with any transmission curve indicates
the amount of radiation left at the depth of the abscissa.
per cent at the bottom of any stratum by that at its top.
This transmission for these lakes is stated in tables 10 and
11. The fourth line gives the values for “mean sun” as de¬
fined on p. 545. These may be used in computing the num-
524 Wisconsin Academy of Sciences, Arts, and Letters.
Fig. 3. The same transmission curves as those in figure 2, but
platted on semi-logarithmic paper. This platting shows the rate of
transmission and its changes in different strata much more accurately
than the regular coordinate paper; but it does not show to the eye
the amount of radiation left at various depths. Note that in the
most opaque lakes the transmission curve is practically a straight
line from 50 cm. down, indicating a constant transmission. In Crys¬
tal Lake the curve is constantly deflected toward the right, indicating
a constantly increasing transmission. Blue Lake and Weber Lake
show variations either accidental or systematic. Weber Lake, be¬
tween 4 m. and 5 m. shows a marked increase of transmission, prob¬
ably accidental. The case is inserted to illustrate this very common
occcurrence. Blue Lake seems to show an increasing transmission
between 5 m. and 9 m. with an accidental drop between 9 m. and 10 m.
Birge & Juday — Transmission of Radiation in Inland Lakes. 525
ber of calories delivered by direct insolation at any given
depth during the warming period, April-August, assum¬
ing that conditions are not dissimilar to those at Madison.
Figures 4 and 5 contain similar transmission curves for
a number of other lakes reported in the general tables.
Figure 6 shows maximum, minimum, and mean curves for
a moderately transparent lake and for an extremely trans¬
parent one; similar curves for Lake Mendota, a relatively
opaque lake, are given in fig. 3. Figure 7 plats the trans¬
mission for a series of lakes, in order to show its system¬
atic and accidental variations. Figure 8 shows transmis¬
sion curves in the upper two meters of several lakes.
In figures 2 and 3 the transmission curves are platted on
two types of co-ordinate paper, fig. 2 on standard paper and
fig. 3 on semi-logarithmic paper. The same facts are
shown on each diagram so as to bring out the relation of
the two types of platting. The standard paper gives to the
eye the better idea of the amount of radiation delivered at
different depths, and the semi-logarithmic paper gives a
more definite notion of the rate of transmission.
Results
Transmission is determined by three main factors: (1)
the selective action of water, which is transparent to short
wave radiation and opaque to long waves; (2) the selective
effect of stain which acts more strongly on the short wave
radiation and is effective in proportion to the amount and
kind present; (3) the action of suspended matter — organic
and inorganic — which offers more obstruction to short wave
radiation, but is not definitely selective.
Thus the selective action of water is a constant factor in
all lakes ; that of stain is very variable as between different
lakes, and somewhat variable in the same lake at different
times. It is usually, but not always, the same in the same
lake at different depths of the same series. The influence
of suspended matter may differ widely in different lakes, in
the same lake at different times, and at different depths in
the same lake at the same time. In eutrophic lakes its in¬
fluence is very great and often dominant; in lakes with lit¬
tle plankton it is less and stain or water itself may be more
526 Wisconsin Academy of Sciences , Arts , and Letters .
important factors in reducing radiation. In the surface
meters there is more plankton, and to this fact is due part
of the relatively low transmission in the 1-2 m. stratum.
The presence of an actual or potential scum of algae in the
surface water may add greatly to the absorption in the
upper meter (Lake Kegonsa, table 11). If a crop of algae
or other suspended matter is settling, the surface water
may be much more transparent than that below (Marl
Lake).
Where high stain and much suspended matter combine
to obstruct radiation, the transmission is low and practi¬
cally constant at all depths. When the transmission curve
is platted on semi-logarithmic paper it is indistinguishable
from a straight line, perhaps from the depth of 0.5 m. on.
In most lakes, however, the transmission of the 1-2 m.
stratum is decidedly lower than that of the strata below;
but in the highly eutrophic lakes of Southeastern Wiscon¬
sin there is but little rise in transmission after passing the
depth of 3 m. This situation holds in general for lakes
whose transmission does not exceed 60. Where transmis¬
sion reaches 70, as in Green Lake, it is apt to increase up
to 4 or 5 m. (table 10) and to remain steady thereafter,
showing only accidental variations in the deeper water
down to 10 m. When transmission reaches 80 or more, as
in Crystal Lake, the increase may continue in some cases as
far as observation has gone, i. e. up to 10 or 11 m. (see table
10) and at that depth transmission may exceed 90. For il¬
lustrations both of the uniformity of transmission and of
its increase with depth, see figs. 3-6 and fig. 8.
Table 2 and figures 2 and 3 illustrate the principles of
transmission in lakes. For illustrations we will confine
our attention to the third line, zenith sun. This is derived
from the observed per cent, as shown in fig. 1, and the
fourth line, mean sun, is derived from zenith sun as shown
in fig. 9.
The great loss of radiation in the upper meter is obvious.
No lake has at the depth of one meter as much as 40 per
cent of incident radiation. Some of the loss is due to re¬
flection, but most comes from absorption in passing through
the water. Pure water transmits through the first meter
less than 50 per cent of incident radiation, the amount
f!
Birge & Juday — Transmission of Radiation in Inland Lakes. 527
varying with the altitude of the sun and the quantity of
precipitable water in the atmosphere. Thus a large part
of the loss of radiation in the first meter comes from water
itself. In Crystal Lake this is the main source of loss, and
most of the remainder is due to suspended matter. In per¬
haps all of the other lakes stain plays a larger or smaller
part in reducing the radiation. No other lake much ex-
Fig. 4. Transmission curves, chiefly from southeastern lakes.
Turtle Lake from the Northeastern District has a lower transmis¬
sion than any southern lake visited. The curve for Pope Lake in
the Southeastern District is practically identical with that of Alle-
quash Lake. These lakes are slightly below the minimum for Lake
Mendota, and Nagawicka Lake has a transmission practically iden¬
tical with the maximum of Mendota. Columbian Lake lies between
the maximum of Mendota and the regular curve of Green Lake.
The curve of Marl Lake approaches that of the most transparent
lakes in its upper part and shows a great decline of transmission
at 8 m. and below.
ceeds 80 per cent at the depth of one meter, Weber Lake
alone going as high as 84 per cent. Both Blue and Weber
lakes are recorded with zero color in the tables, and no
doubt suspended matter has more influence than stain in
them.
Meters
528 Wisconsin Academy of Sciences , Arts , and Letters .
At the other end of the series are Turtle and Mary lakes,
in which both stain and suspended matter combine to re¬
duce the amount of radiation at one meter to 10 per cent or
less. In Turtle Lake stain alone may bring down the
transmission of the first meter as low as 11 per cent. In
Devils Lake and Green Lake stain plays a minor part and
the variations shown in Green Lake in fig. 7 depend mainly
Per Cent
CO 1.5 2 3 4 5 6 7 8 9 10 15 20 30 40 50 60 70 80 100
Fig. 5. Transmission curves for lakes from the Northeastern Dis¬
trict. Wolf, High, and Wild Cat lakes have a fairly high stain and
much plankton; they come within, or (Wolf) just outside the limits
of Lake Mendota. White Sand and Lost Canoe lakes are near the
mean for Mendota. Trout Lake lies between Green Lake and the
maximum for Mendota. Clear Lake is close to Blue Lake (fig. 3)
Two curves for Trout Lake are given, one a regular one and the other
with accidental irregularities. Lost Canoe Lake shows a similar de¬
cline of transmission between 4 m. and 5 m.
on variations in plankton. The same is, of course, true of
the variations in Crystal Lake in fig. 7.
The transmission curves for Lake Mendota (figs. 3 and
10) give an instructive picture of conditions in a lake with
low stain (color 6-10 on the platinum-cobalt scale) and with
large and varying amounts of plankton. The per cent at
Meters
Birye & Juday — Transmission of Radiation in Inland Lakes. 529
one meter may be as high as 30, thus closely approaching
Green Lake. It may be as low as about 14, not far above
Lake Adelaide (table 11) with its color of 25 or more. The
effect of plankton is even more strongly shown in Lake Ke-
gonsa (table 11), which, with a color close to that of Men-
dota had at one meter only 7.2 per cent of the radiation
present at the surface. Within the range shown by Men-
dota come the great majority of the southeastern lakes and
Per Cent
1.0 1.5 2 3 4 5 6 8 10 15 20 30 40 50 60 80 100
Fig. 6. Transmission curves for Green and Crystal lakes. These
show the mean, minimum, and maximum curves for a lake with moder¬
ately high transmission and one whose transmission is exceptionally
high. Corresponding curves for Lake Mendota are given in fig. 3.
The curves for Green Lake are from Aug. 22, 1922 and Aug. 4, 1919.
Those for Crystal Lake are from Aug. 17, 1926 and July 6, 1926.
See table 10.
many of the northeastern. One such lake — Beasley— is
platted on figs. 2 and 3, in order to illustrate this fact.
Table 2 and the accompanying figures show that in the
less transparent lakes the radiation is followed to the depth
where one or two per cent of the observed incident radia¬
tion is left. No lake is studied below 10 m. and at this
depth as much as 4 to 8 per cent remains in Crystal Lake,
but not much more than one-half that maximum in any
Meters
530 Wisconsin Academy of Sciences , Arts, and Letters.
other. In all diagrams the transmission curve has been
prolonged until it intersects the line indicating one per cent
of incident radiation. This prolongation has a very dif¬
ferent length in the several cases. Often the lowest ob¬
served point is close to one per cent and in other cases the
line must be extended through several meters. Where the
distance is short and the curve on semi-logarithmic paper
is practically a straight line, the extension may be confi¬
dently made. Its precise course is uncertain in proportion
to its length and to the nature of the curve. Where a note¬
worthy change in transmission occurs with the observation
at the greatest depth, it is doubtful whether the line should
follow the general trend of the curve or the altered direc¬
tion. For this note the curves for Blue and Weber lakes
in fig. 3. So, also, if transmission is still increasing at the
deepest observation— as in Crystal Lake — it is not certain
how long increase will continue or how far it will go. Ap¬
proximate accuracy only can be assured in such cases. In
most of the lakes in the list the general situation is clear
enough before the last reading is passed.
The depth at which one per cent of incident radiation
can be delivered by the sun varies from about 2 m. to more
than 20 m. The latter distance is reached only by Crystal
Lake among those which have been studied. In one case
one per cent would not have been reached at a depth less
than 30 m., if conditions were the same below 10 m. as they
were above that depth. Since the depth of the lake is 21
m., more than one per cent of the radiation would have been
present in the bottom water, assuming a sun in the zenith.
This was a condition of exceptional transparency; in other
cases in the same lake radiation would have been reduced
to one per cent at a depth of 16 m. or 18 m., the difference
depending on the amount of plankton present.
From the diagram it is also possible to discover the thick¬
ness of the stratum of water below the surface meter or
two, which will reduce radiation to one-tenth of the quan¬
tity delivered to its upper surface. The thickness of this
stratum varies from a minimum of 1.4 m. in Lake Mary to
more than 12 m. in Crystal Lake ; it perhaps reaches nearly
20 m. in exceptional cases in the latter lake. In Lake Men-
dota it ranges from 2.3 to 5.6 m. with a mean of 3.3 m. ;
Birge & Juday— Transmission of Radiation in Inland \ Lakes. 531
and this range would include the cases of the ordinary eu-
trophic lakes, especially those of the southeastern district.
Green Lake, with about 7 m., and Blue Lake with 8.7 m.
illustrate the cases of the more transparent lakes, having
relatively little plankton and little stain.
In the relatively opaque and shallow lakes there would be
small chance of serious error if these figures were applied
to the deeper water of the lakes. Beasley lake, for in¬
stance, had one per cent of incident radiation left at 7.1 m.
and radiation is reduced to one-tenth of its surface value
by passing through a stratum 4.6 m. thick. We are fairly
justified in applying this figure to the remainder of the
lake, which is 15 m. deep. At this rate the value of the
radiation at the bottom would be 0.0002 of the incident ra¬
diation or 2X10-4. In Lake Mendota the radiation might
be followed down in the same way. The curve for mean
transmission shows one per cent just below 5 m. This
would bring one-tenth of one per cent about 8.5 m. and re¬
duction would continue at the rate of 90 per cent in 3.5 m.
to the bottom at about 24 m. In the case of the maximum
observation the depth at which one per cent is reached is
8.9 m. and an additional stratum 5.6 m. thick is needed to
reduce this value to one-tenth of one per cent. Figure 9
shows the situation in Lake Mendota, giving the data for
both zenith sun and mean sun. The curve for filtered
water in this lake is also platted.
These diagrams well illustrate the fact to which Poole
and Atkins call attention (’28, p. 475) ; viz. that the de¬
crease in radiation with depth goes up very rapidly as the
per cent of transmission decreases, since this decrease
makes the radiation fall off in a geometrical ratio. Thus
at one meter the amount of radiation in the lake ranges
from 0.14 to 0.28 — a two-fold ratio. At 10 m. the maxi¬
mum shows that there is left 6X10'3, while the minimum is
only 2xl0~5. At 5 m. the maximum shows nearly 0.05,
while the minimum has only 0.0026, or practically one-six¬
teenth as much.
The following table summarizes the most important facts
regarding the lakes shown in figures 2 and 3. They will
also be found listed in table 11, where the transmission in
532 Wisconsin Academy of Sciences, Arts, and Letters.
the several meter strata is given ; and the more transparent
lakes are also found in table 10.
Table 3. Summary of data regarding lakes shown in
figures 2 and 3
Stratum
transmit-
Figure 7 shows for a few lakes the coefficients of trans¬
mission platted on a large scale, so as to illustrate various
details. Among these are: the marked rise in transmis¬
sion in the 1-2 m. stratum as compared with the 0-1 m.
stratum, and the smaller rise when the 2-3 m. stratum is
reached; the tendency toward a uniform rate of transmis¬
sion in almost all cases, together with a gradual systematic
rise in the upper strata of the more transparent lakes; the
kind and amount of accidental variation which may be ex¬
pected in such observations.
In the extreme case — Crystal Lake — transmission must
nearly reach the maximum for inland lakes, since at 10 m.
it is above 90. The transmission of Blue Lake seems to be
steady at about 80. Marl Lake, very transparent near the
surface, shows a marked and progressive loss of transmis¬
sion in the deeper water, which is probably due to sus¬
pended matter which had settled out of the surface water.
Green Lake offers a typical case of a moderately high trans¬
mission — going above 70 — together with the accidental
variations which are often found in such a series. Some¬
thing of the same sort is shown by Beasley Lake, on a lower
level of transmission. Spider Lake has a very low trans-
Birge & Juday — Transmission of Radiation in Inland Lakes . 533
mission in the upper meter, due to abundant plankton. The
same fact causes uniformly low transmission below two
meters. The color of the lake water is low (8) and would
not of itself cause a low transmission. Compare in this
01 23456789 10
Meters
Fig. 7. Transmission curves platted to show the variations in the
several one-meter strata. The per cent of transmission is platted
in the middle of each stratum and the points thus indicated are con¬
nected by straight lines.
respect the record of Beasley Lake or Nagawicka Lake
(table 11) with the same color; or Kawaguesaga and Ocon-
omowoc lakes (table 11), both of which had a color of 14
with a transmission rising above 60.
534 Wisconsin Academy of Sciences , Arts , cmd Letters.
Transmission in the Upper Meters
Figure 8 shows on a large scale the transmission from
the surface to the depth of 2 m. The curves are from lakes
which represent all types from the most transparent to the
most opaque in the list. The cases platted are given in the
general tables and are as follows :
Adelaide, July 11, 1928
Crystal, August 10, 1928
Mary, July 11, 1928
Okauchee, August 9, 1919
Pauto, August 18, 1928
Seneca, August 1, 1918
Trout, August 24, 1928
Turtle, August 30, 1918
All but two of these are from the northeastern group of
lakes. Very few readings have been taken at the depth of
10 cm. in the southern lakes. A series from Lake Mendota
was thus taken on May 29, 1928, which is recorded as the
maximum series. It would almost exactly coincide with
the curve from Lake Pauto. All series were taken on oc¬
casions when the surface of the lake was free from waves
or even larger ripples. However, the readings at 10 cm.
cannot claim the accuracy that belongs to those at 100 cm.
The rate of transmission is changing rapidly at this depth
since the long wave radiation is being rapidly absorbed. A
difference of even a half centimeter in depth would make a
decided difference in the per cent recorded ; and in working
over the side of a rowboat small errors in depth are sure
to occur. However, the general correctness of the results
is clear.
The curves platted in figure 8 are not corrected for zen¬
ith sun, but are taken from the results directly observed.
The rate of transmission is changing so rapidly in the sur¬
face centimeters of the lake that it is not advisable to make
such corrections without having a much larger amount of
data, and indeed without data obtained with greater ac¬
curacy than is ordinarily practicable in observations made
from a boat.
The rapid loss of long wave radiation in the upper 10 cm.
appears in the fact that, at this depth, there remains only
Birge & Juday — Transmission of Radiation in Inland Lakes. 535
from 35 to 60 per cent of the radiation received by the sur¬
face. The percentage of loss falls off rapidly as depth in¬
creases, since at 25 cm. there is left from 21 to 51 per cent
Per Cent
1.0 1.5 2 3 4 5 6 8 10 15 20 30 40 50 60 80- 100
Fig. 8. Transmission in the upper two meters. In this diagram
the scale of depth is ten times that of the other diagrams. The re¬
sult is to bring out the small variations as well as the major changes
as depth increases. Turtle Lake has an unusually irregular curve,
and was selected for that reason. The other curves show the or¬
dinary and normal variations in lakes covering the scale of trans¬
parency. The curves are platted as observed, and are not corrected
for zenith sun.
of the surface radiation, or from 60 to 80 per cent of that
present at 10 cm. At 50 cm. only 10 per cent of surface ra¬
diation is left in the case of Lake Mary, while in Crystal
Centimeters
536 Wisconsin Academy of Sciences , Arts, and Letters.
Lake nearly 45 per cent remains. At 50 cm. in Lake Mary
there has been lost about 50 per cent of the radiation pres¬
ent at 25 cm.; the loss between 25 cm. and 50 cm. may be
less than 15 per cent in Crystal Lake.
At 100 cm. there remains in Lake Mary about 4 per cent
of the radiation delivered to the surface, while in Crystal
Lake about 38 per cent is present. Thus, while there is
about a two-fold range in the amount of radiation present
at 10 cm., the range is nearly ten-fold at one meter. In
Lake Mary the 50-100 cm. stratum transmits about 40 per
cent of the radiation received by its upper surface; while
in Crystal Lake more than 85 per cent is thus transmitted.
In all of the lakes transmission necessarily increases with
depth. In Crystal Lake the transmission of the several
strata 10-25 cm., 25-50 cm., 50-100 cm. is of the same or¬
der of magnitude, i. e., about 85 per cent, in spite of the dif¬
ference in thickness of the stratum: and this transmission
is not far different from that of 100 cm. at depths somewhat
greater. In this and in similar cases this increase is due
mainly to the rapid absorption of long wave radiation by
the colorless water, with a corresponding reduction of the
base for comparison of the several strata. At the other
extreme, the transmission in Lake Mary is practically uni¬
form from the depth of 50 cm. and in lakes Turtle and Ade¬
laide from the depth of 25 cm. only accidental variations of
transmission appear. This situation is due to the high
color of the water and the large amount of plankton found
in it. The increase of transmission in Lake Mary at and
below 50 cm. was due to the fact that the surface centime¬
ters were more highly charged with dark gummy extractives
from peat than were those just below. In all these cases
color cut off so much of the short wave radiation that the
energy spectrum quickly assumed what may be called a
symmetrical form and its content thereafter fell off at a
nearly uniform rate.
Transparency and Transmission
It is commonly said that there is no observable correla¬
tion between the depth at which Secchi’s disc is visible and
the transmission of radiation (Aufsess ’03, p. 46, Oberdor-
Birge & Juday — Transmission of Radiation in Inland Lakes. 537
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538 Wisconsin Academy of Sciences, Arts, and Letters.
fer ’28a, p. 478) . This statement is true in general and is
indeed the only conclusion which one would reach as the
result of a few observations. Stain affects transmission
much more than it does transparency and where this is a
dominant factor in the lake, correlation is absent or diffi¬
cult to determine. Since the amount of suspended matter
is one main factor in determining both transparency and
transmission, there should be at least a rough correlation
between the results where this factor has a large part to
play.
Where numerous observations are available such a rough
correlation between disc and transmission is easily found.
Figure 9 shows the result for Lake Mendota of platting
transmission against transparency or the depth to which
Secchi’s disc was visible. For every unit of depth there is
a large range of transmission, provided there are enough ob¬
servations. Thus, in the 1-2 m. stratum there is a range of
transmission from 37 to 50 when the transparency is from
2 m. to 3 m. ; the range is from 42 to 58 when the trans¬
parency is between 3 m. and 4 m. Where the transparency
was 3.1 m. the transmission in three cases was 49, 51, and
58.
On the other hand, if the mean transparency and mean
transmission are determined it will be found that 18 cases
come below the mean of transparency and 18 at or above it.
In the 1-2 m. stratum (marked with a circle in the diagram)
there are 6 cases below the mean transparency and at or
above the mean transmission ; but of these only 2 are more
than 1 per cent above the mean. Of the 18 cases above the
mean transparency, 16 are also above the mean transmis¬
sion. Much the same result is seen also in the cases from
the 2-3 m. stratum, which are marked with a cross in the
diagram.
It would be possible to draw a mean curve, representing
the relation between transparency and transmission; but
the number of cases is so small and the range in each rubric
is so great that it is not worth while to attempt it. It is
evident that transmission rises faster than transparency at
first and more slowly afterwards ; so that when transparency
is at or near mean almost all cases of transmission are
Birge d; Juday — Transmission of Radiation in Inland Lakes. 539
•3
<o
S
e
CO
£
540 Wisconsin Academy of Sciences , Arts, and Letters .
above the mean. This situation means that in the higher
rubrics of transparency the transmission comes to depend
more and more on the effect of stain and water, while in the
lower rubrics the suspended matter is a larger factor in de¬
termining the result, both for transparency and transmis¬
sion.
A similar result is reached when the relation between disc
and transmission is studied in the list of table 11. The re¬
sults are shown in table 4.
This table, in which all lakes except Mendota are repre¬
sented, shows relations of much the same order as does the
large series of observations from Mendota. Low trans¬
parencies are accompanied with low transmission and high
transparencies with high transmission. There is also a
wide range of transmission under each rubric of trans¬
parency and a wide range of transparency under each rubric
of transmission. The two cases which carry the range of
transmission beyond 50 when the disc was visible between
1 and 2 m. both come from the marly lakes of Waupaca;
and these lakes are exceptional in this relation (see p. 000).
However, the transmission in those lakes whose visibility
was 2-3 m. has also a great range and the cases are scattered
along the whole distance. There is no case of a lake with
transparency more than 4 m. and transmission below 50;
similarly, all lakes with visibility beyond 5 m. have trans¬
mission greater than 55. No lake goes much beyond 70
unless its transparency is greater than 6 m.
In these 91 cases the transparency lies between 2 m. and
6 m. in 65 cases or 71 per cent, and the transmission lies be¬
tween 50 and 69 in 58 cases or 64 per cent. The cases of
relatively high transmission are dominated by the 27 series
from three lakes, Trout, Green, and Crystal ; so that no de¬
tailed discussion would be profitable.
It should be noted that this table deals with the 1-2 m.
stratum only; if deeper strata were included the transmis¬
sion would be higher, as may be seen from the tables from
which these data are taken.
Birge & Juday — Transmission of Radiation in Inland Lakes. 541
Lake Mendota
Data
Lake Mendota is a lake typical of southeastern Wiscon¬
sin. It lies in lat. 43.1° N. ; long. 89.4° W. It is relatively
shallow (25 m.), has hard water (about 40 to 55 p.p.m. of
CaO and MgO), has very abundant plankton, and water
which is not highly colored (6 to 10 on U. S. G. S. scale).
Under these conditions transmission of radiation is low and
very variable, the variations depending primarily on the
amount of suspended matter in the water.
Many observations have been made in the lake at all times
during the warmer season and wide extremes of transmis¬
sion have been recorded ; so that practically all of the other
similar southeastern lakes come within their range, as do
also many lakes of the same type in northeastern Wiscon¬
sin.
In table 5 are recorded 36 series of observations on Lake
Mendota, taken from 1912 to 1928, mostly during the sum¬
mer. Many other series are not included because they do
not add to the knowledge derived from those in the table.
In some of the earlier series the readings extended only to
two meters. All of these are omitted, although the trans¬
mission below two meters is ordinarily much the same as
that in the 1-2 m. stratum. In other cases the observations
were interrupted by cloud or made irregular by streaks of
haze or high cirrus. The recorded observations give both
maximum and minimum cases and yield a mean which is
sensibly the same as that which would result from the inclu¬
sion of all.
In many cases the observations ended at the depth of three
meters. In 16 cases out of the 36 in the table they went to
four meters and in 7 to five meters. The swing of the gal¬
vanometer in the deeper water is small ; at three meters it
was ordinarily four or five divisions of the scale; at four
meters it would be about half that amount and correspond¬
ingly smaller at five meters. Under such conditions the
computed transmission comes to depend on the estimate of
the fraction of a division in reading the scale and is corre¬
spondingly uncertain. In general the conditions of trans-
542 Wisconsin Academy of Sciences, Arts, and Letters.
mission below the surface meter are nearly uniform down
to the depth which we have set as the limit of our study —
that at which one per cent of the incident radiation is found.
This depth is well within the limits of a uniform epilimnion
during the summer.
Results
The per cent of incident radiation present at the depth
of one meter below the surface has been computed on the
assumptions that the sun is in the zenith and that all radia¬
tion comes directly from the sun. In that case the radia¬
tion passes through one meter of water in reaching the same
depth below the surface. The result is as follows :
Mean per cent, 20.7 ; maximum, 27.8 ; minimum, 18.9.
Table 5 gives no ground for inferring that the altitude of
the sun affects this result, if observations are reduced to
zenith sun. The two series on July 10, 1916, give sub¬
stantially identical results (see p. 518 and fig. 1) , and in gen¬
eral there is no correlation between the hour of the day and
the percentage at one meter.
The minimum per cent happens to be just one-half of the
maximum. It is not probable that this relation would be
found if a larger series of observations were taken with
special reference to it. It is not likely that much more than
28 per cent would be found at one meter, for no lake of
similar type in Wisconsin has yielded a higher per cent, as
may be seen from table 11. On the other hand, the water
of Lake Mendota has often been more densely filled with al¬
gae than on any occasion when observations were made on
radiation; and under such conditions the transmission in
the surface water would be lower, as may be seen from the
case of Lake Kegonsa in table 11.
Readings were taken in all cases at the depth of 50 cm.,
with the following result, when computed for zenith sun :
Mean per cent, 33.1; maximum, 41.0; minimum, 28.0.
In 18 cases readings were taken at the depth of 25 cm. be¬
low the surface. The mean result was to give at that depth
1.33 times the per cent at 50 cm. This would give a mean
of 44 per cent of incident radiation present at 25 cm. depth.
These results have been used in platting the mean transmis¬
sion curve of radiation (fig. 10).
Birge & Juday — Transmission of Radiation in Inland Lakes. 543
Transmission below the depth of one meter
The transmission of radiation through the successive one
meter strata of water below the surface is as variable as is
the per cent present of the depth of one meter. The results
are :
1- 2 m. stratum
Mean per cent, 47.3 ; maximum, 65 ; minimum, 29.
2- 3 m. stratum
Mean per cent, 48.7 ; maximum, 66 ; minimum, 34.
Thus the mean transmission increases slightly with depth
in the upper meters, as would be expected. If the trans¬
mission curve is platted on a large vertical scale, this in¬
crease is apparent. But when platted on the scale used in
figures 3-6, the fact hardly appears. It is also true, as table
Table 5. Transmission of solar radiation in the surface water of
Lake Mendota.
544 Wisconsin Academy of Sciences , Arts , and Letters .
5 shows, that an increase below 2 m. is by no means always
present, and figure 10 shows the reason. The transmission
curve of the filtered water is shown in comparison with
those of the lake. At 3 m., for instance, there would be
about 20 per cent of incident radiation present in filtered
water. In the lake the maximum found at 3 m. is 11 per
cent, or about half as much, and from this it ranges down¬
ward to 1.8 per cent, or about 9 per cent of that present in
the filtered water. At 5 m. the filtered water would have
reduced radiation by about 8 per cent, leaving 12 per cent;
while in the lake there remain from 5 to 0.25 per cent. Thus
suspended matter decreases transmission by a very large and
a very variable amount and may quite obscure the more
modest effects of water and stain at depths below on 3 or
two meters. The occluding effect of suspended matter de¬
pends mainly on its relative abundance, which is very vari¬
able. Hence the transmission in deeper strata may be
greater or smaller or remain about the same, according to
the balance of these variable factors. In general, there is
a slight increase.
In 16 cases the radiation was followed through the 3-4 m.
stratum and the mean result was a transmission of exactly
50 per cent. The few cases where the transmission through
the 4-5 m. stratum was measured gave no reason for infer¬
ring any further increase. In computing the average prog¬
ress of radiation through the lake a transmission of 50 per
cent per meter has been assumed for all depths below three
meters. At this rate the radiation would be reduced to one
per cent of its surface value at the depth of 5.3 m. below
the surface.
The depth at which one per cent of incident radiation is
found is computed for the single observations on the as¬
sumption that the transmission found in the lowest meter
observed is continued in the underlying meters. The maxi¬
mum depth thus found is 8.8 m., May 29, 1928. The mini¬
mum depth is 3.5 m., May 20, 1926. This difference was
due to a marked difference in the plankton, which also
showed itself in the transparency as measured by Secchi’s
disc. In the first case readings extended to 6 m. and in the
second to 4 m. The series of 1928 was made by the aid of
the new and sensitive millivoltmeter.
Birge & Juday — Transmission of Radiation in Inland Lakes. 545
Table 6. Mean effect of insolation , computed for zenith sun.
Depth, Per cent Transmission
meters present Depth Per cent
1 _ 20.70 1-2 m. 47
2 _ _ 9.73 2-3 m. 49
3 — _ _ 4.77 3-4 m. 50
4 _ _ 2.38 4-5 m. 50
5 _ 1.19 5-6 m. 50
6 _ _ _ _ 0.60
No considerable error would be made in asserting as a
general statement, that with zenith sun 20 per cent of inci¬
dent radiation is present at one meter depth in Lake Men-
dota; and that the average transmission per meter below
that depth is 50 per cent.
The observations recorded in table 5 are reduced to
zenith sun and are therefore maximum. They do not rep¬
resent the average per cent of radiation which is found at
one meter during a day or during the season. The average
path of radiation direct from the sun in reaching the depth
of one meter is longer than that assumed, and the presence
of much diffuse radiation from sky and cloud adds to the
length of the average path. The amount of incident radia¬
tion which is reflected increases as the angle of incidence
decreases and thus the amount that enters the water is di¬
minished.
Mean path of radiation ; “mean sun”
The mean path of total radiation, including both direct
and indirect, which enters the lake has been computed with
allowance for reflection, as stated on p. 576. The lake
reaches the temperature of 4° C. about April 15 and both
warming of the water and the growth of plants from the
bottom are substantially completed by August 15. This
may be called the “warming period” of the lake. The
mean path of radiation in reaching the depth of 100 cm.
below the surface during this period is computed as 116 cm.
The radiation at 1 m. below the surface has therefore suf¬
fered an average loss equivalent to that from 1.16 m. with
zenith sun. The transmission curve for zenith sun is
marked at depths of 1.16 m., 2.32 m., etc. The figures thus
546 Wisconsin Academy of Sciences, Arts, and Letters .
obtained show the average situation during the warming
season, of course on the assumption that the transmission
remains the same.
Applying this method to the curve of mean transmission
with zenith sun gives the following result :
Table 7. Mean results during warming period.
Depth, Per cent Transmission
meters present Depth Per cent
1 _ 18.30 1-2 m. 43
2 _ 7.80 2-3 m. 45
3 _ 3.45 3-4 m. 45
4 _ 1.55 4-5 m. 45
5 _ 0.70
The base for computing per cent in column 2 is the total
radiation in air, not that which enters the water.
During the period April 15-August 15 the lake receives
about 60,000 cal. cm.2, and of this amount about 4,620 cal.
are reflected from the surface. The following table shows
the quantity of radiation delivered to different depths in
the lake, on the above assumptions.
Table 8. Calories per square centimeter delivered during warming
period. Computed for umean sun,” April 15-August 15.
Depth, Cal. Depth, Cal.
meters meters
1 _ 10,980 0-1 m. 44,400
2 _ 4,650 1-2 m. 6,300
3 _ 2,070 2-3 m. 2,610
4 _ 930 3-4 m. 1,140
5 _ 420 4-5 m. 510
Below 5 m. 420
Most of the enormous quantity of heat which enters the
surface meter is used in evaporation, is rapidly returned to
the air, or is distributed by wind to deeper strata. The
longer wave lengths penetrate only a very short distance.
The 1-2 m. stratum gets from the sun nearly three times
as much heat as is retained; its mean maximum temper¬
ature may be placed at about 24°, requiring 2,000 cal. cm.2
to warm it from 4° to this point. The 2-3 m. stratum,
Per Cent
Birge & Juday — Transmission of Radiation in Inland Lakes. 547
S«313J/\[
Fig. 10. Transmission curves for Lake Mendota, showing the curves for filtered water, and mean, maximum, and
minimum for the lake, extended to 0.1 per cent. The last three are given in two forms, that for zenith sun and that
for mean sun. See text p. 548.
548 Wisconsin Academy of Sciences, Arts, and Letters.
which has the same average temperature, gets from the sun
directly only a little more than it retains ; the meters below
get from insolation much less than they receive altogether,
and below 5 m. the effect of direct insolation is almost neg¬
ligible so far as the warming of the lake is concerned.
The region of the solar spectrum between 0.360 ^
and 0.762/a contains about one-half of the total energy pres¬
ent in the whole spectrum, and practically all of the energy
delivered by insolation to depths of one meter or more be¬
longs to this region. If, therefore, the percentages of radi¬
ation given in tables 5 and 11 are doubled they will yield a
fair estimate of the radiation present as light at these
depths. This is energy measured in calories per square
centimeter of area.
Rickett (’22, p. 500) found that higher plants grow from
the bottom in Lake Mendota to a maximum depth of about
6 m. At this depth there should be present an average
amount of about 0.6 per cent of the energy received by the
surface in the form of radiation within the visible spectrum
as defined above. When the sun is high in the sky this per
cent might be doubled or trebled, and at maximum trans¬
parency and high sun more than 4 per cent might be pres¬
ent at this depth. The region of the spectrum to which the
water of Lake Mendota is most transparent has its center
about 0.550/a and a much larger percentage of a spectral
band in this region would reach the depth named. This
band lies in the region of green and yellow light.
Figure 10 shows the transmission curves for Lake Men¬
dota, the minimum case, the mean of 36 series, and the max¬
imum ; they are given for zenith sun and also for mean sun.
There is also platted on the same diagram the transmission
curve of the filtered water of the lake ; this curve is a mean
from five series. The minimum curve was read to the
depth of four meters ; the maximum to six meters ; all are
adjusted for zenith sun.
All curves are extended until they intersect the line repre¬
senting 0.1 per cent of the radiation delivered to the surface ;
or until they intersect the line representing 15 m. The
depth at which one per cent is reached is: minimum, 3.6
m. ; mean, 5.2 m. ; maximum, 7.9 m. ; filtered water, 16.0 m.
Since the color of the water does not vary greatly at differ-
Birge <& Juday — Transmission of Radiation in Inland Lakes. 549
ent times we may conclude that the difference in transmis¬
sion is largely due to the varying amount of suspended mat¬
ter in the water.
The difference between the several curves increases rap¬
idly as they are carried down. The following table shows
the situation.
Table 9. Lake Mendota. Per cent of radiation present at various
depths.
Thus at 5 m. the maximum amount of radiation is about
20 times as great as the minimum; at 10 m. the mean is
nearly 20 times the minimum and the maximum about 20
times the mean; so that the maximum is nearly 400 times
the minimum, or the difference is 20 times as great as at
5 m. Similar relations can be traced if the changes in each
curve are followed from 5 m. to 10 m.
The Southeastern Lakes
Data
Tables 10 and 11 contain the data from the lakes both of
southeastern and northeastern Wisconsin. They also con¬
tain data from four lakes outside of Wisconsin.
In table 10 are given records from the lakes in which ob¬
servation extended to the depth of 9 or 10 m. These in¬
clude 9 of the more transparent lakes from all regions.
The per cent present at each depth is recorded and also the
transmission through the several one-meter strata. Table
11 gives in summary form the records from all lakes to the
depth of 5 m., if observation went so far. The transmis¬
sion curves from many of these lakes are shown in figures
3-6.
Results
The list includes 26 lakes from the southeastern district,
besides Lake Mendota, with 45 series of observations. In
550 Wisconsin Academy of Sciences, Arts, and. Letters.
general, their records agree with those from Lake Mendota.
This would be expected, since the general character of the
lakes is similar. They are glacial lakes situated in a region
of drift that contains much calcium and magnesium. The
water of the lakes, therefore, contains much of the salts of
these elements and the amount and nature of the plankton
are partly determined by this fact. The amount of stain in
the several waters does not differ greatly. It is not sur¬
prising that the figures given in table 11 should show that
in many of the lakes the amount of radiation found at a
depth of one meter and the transmission through successive
meters lie within the limits observed in Lake Mendota.
In each case a long series of observations would probably
disclose a different mean result from that found in Lake
Mendota. Shallow and active lakes like Kegonsa will be
found to have much more plankton than Mendota and the
transmission of radiation will suffer accordingly. Deep
lakes like Green and Geneva have much less plankton and
the transmission is correspondingly higher. Lakes like
Okauchee, Nagawicka, and North are affected by the large
amount of material brought in by affluents from a wide
area. Pine, Elkhart, and Mouse are lakes fed by springs
and are without affluents or adjacent marsh; they are cor¬
respondingly transparent.
In spite of these differences of detail the resemblances be¬
tween these lakes are far more noteworthy than the differ¬
ences. But there are two obvious exceptions to these state¬
ments — the marl lakes of the Waupaca group and Devils
Lake. In the marl lakes the presence of this substance is
decisive in determining the character of the lake. The dis¬
solved salts of the water are not greatly different from
those in other lakes (Birge and Juday ’ll, pp. 170-171),
but the presence of numerous particles of marl in the water
affects its color and the plankton which it carries. These
lakes have a low transparency as measured by Secchi’s disc,
owing to the lack of contrast between disc and back¬
ground; but they have a high transmission of radiation
and a relatively large per cent is present at the depth of one
meter. The location and surroundings of Devils Lake give
it a character widely different from any other southeastern
lake. It is really an outlying member of the northeastern
Birge & Juday — Transmission of Radiation in Inland Lakes. 551
group of soft water lakes— a group which lies 150 miles
(250 km.) to the north.
The lakes of southeastern Wisconsin are members of
three groups besides five single lakes. They may be ar¬
ranged as follows:
1. Three of the lakes of the Four Lake group — Men-
dota, Monona, and Kegonsa. These are all con¬
nected by the Yahara River.
2. Eight lakes from the Oconomowoc group — Garvin,
Mouse, Nagawicka, North, Pine, Oconomowoc,
Okauchee, Upper Nemahbin. This group lies
close to lat. 43.1° N., long. 88.2°-88.5° W.
3. Eleven lakes from the Waupaca group — Beasley, Co¬
lumbian, Knights, Long, Marl, McCrossen, Otter,
Pope, Rainbow, Round, Taylor.
4. Five single lakes — Crystal and Elkhart, Devils, Ge¬
neva, Green.
All of these lakes are fully described in Juday, ’14 and
only those matters will be mentioned here which are essen¬
tial to the understanding of the table.
1. Monona and Kegonsa lakes are less deep than Men-
dota, and they have a larger amount of plankton for each
unit of volume. The amount beneath each unit of surface
in these lakes is much the same and since the depth is less
the number of organisms in a cubic meter is greater. The
result is to cause a greater obstruction to the passage of
light. This does not appear in the series from Monona
which happened to come on a day when the plankton was
unusually low; but it is very conspicuous in that from Lake
Kegonsa. The color of the water is much the same as that
of Mendota and has a similar effect on the light; and the
transmission will be greater or less according to the abun¬
dance of the plankton.
2. The several lakes of the Oconomowoc district fall into
two groups. North, Okauchee, and Oconomowoc lakes are
strung along on the Oconomowoc River, and Nagawicka
and Nemahbin lakes are on the Bark River. In both cases
the situation is like that of the Four Lakes. Pine Lake and
Mouse Lake are spring-fed, with little or no marsh adjacent
552 Wisconsin Academy of Sciences, Arts, and Letters .
and without affluent or effluent. They show a correspond¬
ing transparency and transmission, but both are within the
limits of Lake Mendota. Garvin Lake has a similar situa¬
tion; but it is so small that its character is determined by
the wash from the adjacent slopes.
It will be noted that the transmission in the 1-2 m. stra¬
tum of North Lake on August 9 differs greatly from that of
the 2-3 m. stratum. The visit to the lake on August 19 was
was made to ascertain whether the peculiar situation on the
9th was due to temporary or permanent causes. It ap¬
peared that the former was the case and probably the low
transmission on the 9th was due to an unusual concentra¬
tion of plankton.
3. The Waupaca group of small, deep lakelets offers very
interesting differences. The Waupaca Chain o’ Lakes
consists of 12 lakes or lakelets and extends about 5 km.
from Otter Lake on the northeast to Marl Lake on the
southwest. The lakes are therefore small, ranging from
Beasley with an area of 5.4 ha. to Rainbow with 55.7 ha.
Their depth is considerable, ranging from 12 m. to 29 m.
There are substantially no affluents and all lakes discharge
through Long Lake. Six lakelets lie to the east and five to
the west of the outlet. They are in lat. 44.4° N. ; long.
89.2° W.
There are two widely different types of lake — the ordi¬
nary type and marl lakes. Among the lakes examined,
Rainbow, McCrossen, Round and Columbian (naming them
in order) are marl lakes. At the extreme end of the series
Marl Lake has a like character. They have large marl de¬
posits on the bottom and many fine particles of marl are
suspended in the water. The last matter is especially evi¬
dent in summer when navigation on the lakes is active.
The water has a bluish-green color when viewed in the
lake and is nearly colorless when filtered. The transpar¬
ency of the lakes, as measured by Secchi’s disc, is low and
the transmission of radiation is high. The lake bottoms
in shallow water support only a scanty plant life.
The lakes of the other type are Beasley, Knights, Long,
Otter, Pope. In these there are no marl springs ; the water
is obviously stained ; there is an abundant growth of plank¬
ton and of plants on the bottom; the transparency is us-
Birge & Juday — Transmission of Radiation in Inland Lakes. 553
ually greater than that of the marl lakes, but the trans¬
mission of radiation is less. Taylor Lake seems to be some¬
what intermediate in character, though nearer to the marl
lakes.
In this group was found the smallest transmission among
the southeastern lakes — Pope Lake, 18 per cent; also the
highest transmission — Marl Lake, 80 per cent. The marl
lakes had ordinarily a transmission above 60 per cent,
though with exceptions; lakes of the other type rarely ex¬
ceeded 60 and often fell below that number.
The three series recorded for Marl Lake offer an instruc¬
tive case of the effect of suspended matter on transmission.
There is no reason to suppose that the absorption of radia¬
tion by the water of the lake differed notably on the three
occasions. There is no adjacent marsh and there is very
little opportunity for drainage water to enter from the sur¬
rounding country. The difference in transmission was due
to suspended matter, chiefly particles of marl. There is a
rough correlation between transparency and transmission
in these cases. One of them furnishes the only case from
the southeastern lakes where the transmission exceeds 80.
4. Among the single lakes, Crystal Lake must not be con¬
fused with its namesake in the north. It is a very differ¬
ent lake in all respects. It is situated not far from Elk¬
hart Lake and it presents the ordinary type of a rather
shallow southeastern lake. Each of the other four lakes
in the class has points of special interest.
Elkhart Lake (lat. 43.8° N.; long. 88.0° W.) is a small
deep lake (121 ha., 34.5 m.) lying in a hollow of the Kettle
Moraine. It is exceeded in depth only by Geneva and
Green lakes in the southeastern group; and Trout Lake in
the north is perhaps a meter or two deeper. It has no af¬
fluent and little or no adjacent marsh so that its water is
clear. In accordance with this situation and with the depth
of the lake, it has a high transmission, showing over 30 per
cent of incident radiation at one meter, and a transmis¬
sion reaching 75 per cent below that depth.
Geneva Lake (lat. 42.6° N. ; long. 88.5° W. — 2210 ha.,
43.3 m.) in the extreme southeast of the state, lies in an
old drainage valley which has been blocked by glacial ac¬
tion. As would be expected from its depth the transmis-
36
554 Wisconsin Academy of Sciences, Arts, and Letters.
sion is relatively large. On both occasions when the lake
was visited the transparency was greater than usual and it
is probable that the transmission was also higher than a
larger number of series would show.
Green Lake (lat. 43.8° N.; long. 88.6° W. — 2972 ha., 68
m.) also lies in a preglacial valley. It is the deepest in¬
land lake in Wisconsin and apparently the deepest inland
lake between the Finger lakes of New York and the lakes
of the Rocky Mountain region. It is also the largest lake
of Wisconsin except the very shallow Lake Winnebago (558
sq. km., 6.4 m.) . It is the only lake in Wisconsin that fully
complies with the character of an oligotrophic lake as de¬
fined by European limnologists. Much study has been
given to it by the Survey, since it presents characteristics
widely different from Lake Mendota and the other south¬
eastern lakes. The relation to radiation is recorded in ten
series in six different years from 1912 to 1923. There are
other series which are omitted as either too short or other¬
wise imperfect. Their results are in general agreement
with those which are in the list. All but one of these series
extended to the depth of 8-10 m. The shore of the lake is
so steep in places that a depth considerably exceeding 10 m.
can be found within less than 100 m. of the shore. In gen¬
eral the readings were limited to the epilimnion, which ends
at eight or ten meters in July and August. It is often im¬
possible to get satisfactory readings of the pyrlimnometer
at the level of most rapid transition to the thermocline, as
the small and rapid oscillations of temperature in the water
obscure the effect of the sun.
The average amount found at one meter was 27.3 per
cent of the incident radiation, with a maximum of 30.6 and
a minimum of 23.3 per cent. The average transmission in
the 1-2 m. stratum was 64, rising to 69 in the 3-4 m. stra¬
tum and remaining close to that figure below, though rising
beyond 70 at 9 or 10 m. In all of these particulars there
is a close resemblance between Green Lake and Trout Lake
in the northeastern group of lakes.
Devils Lake (lat. 43.3° N.; long. 89.7C W., 146 ha., 12 m.)
differs widely from all the other southeastern lakes. It
lies in the gorge where the preglacial valley of the Wiscon¬
sin River passed through the quartzites of the Baraboo
Birge & Juday — Transmission of Radiation in Inland Lakes. 555
Range. This valley was deeply filled with drift and on top
of this filling the lake lies between two sandy ridges. On
the east and west sides the lake is bounded by the talus of
the quartzite cliffs. The water of the lake is soft, contain¬
ing 5-7 p.p.m. of fixed carbon dioxid, or less than one-tenth
as much as the average southeastern lake. The plankton
is small in quantity and the lake is the only one in southern
Wisconsin in which Holopedium is regularly found. There
are no affluents and very little adjacent marsh. The water
has little color and the lake is unusually transparent.
The average transmission of the lake exceeds that of any
other southeastern lake and approaches that of the clearest
lakes in the northeastern district. Over 80 per cent of in¬
cident radiation may be found at one meter and the trans¬
mission may rise to 80. This was also reached by Marl
Lake, but only as an exceptional case, while there is no rea¬
son to doubt that it may often be attained in Devils Lake.
Green, Geneva, and Elkhart lakes belong to a group of
moderately transparent lakes, which also includes Trout
Lake from the northern district. These have little plank¬
ton, low color, and a transmission which regularly exceeds
the maximum for Lake Mendota. So far as known, the
variation in transmission (see fig. 6) is small, as would be
.expected from the small amount of plankton. Devils Lake
forms a transition to the extremely transparent lakes of
the northeastern district.
The Northeastern Lakes
There are 25 lakes listed from the district in Northeast¬
ern Wisconsin, with 47 series of observations ; as compared
with 26 lakes (besides Lake Mendota) with 45 series from
the Southeastern groups of lakes. This group of lakes lies
close to lat. 46° N. and between 89° and 90° W. long.
The lakes of the northern group are much more varied
and interesting than those from the south. The difference
between the transmission of radiation in the southern lakes
is largely a matter of the quantity of plankton; stain is in
general a subordinate factor. The marl lakes, indeed, offer
an interesting problem; having very clear water, made
more or less milky by mechanical mixture of marl particles.
556 Wisconsin Academy of Sciences, Arts, and Letters .
But these are only a few lakelets and the problem is largely
a mechanical one.
No lake in the southern group has a color that exceeds 14
and none would be wholly devoid of color. In general, the
stain ranges about 6 or 8. Among the northern lakes there
are six whose stain exceeds 20, one of them going to 118,
and four others have a color between 14 and 20. There
are four lakes whose stain is regularly too small to measure
and is recorded as zero. One of these, Crystal Lake, is
definitely a “blue” lake, since Secchi’s disc or other white
objects appear distinctly blue when viewed through a few
meters of its water. No other lake has been found which
is so definitely blue, though several others, not in the pres¬
ent list, approach it. Thus both the presence and the ab¬
sence of stain produce effects in the northern lakes on a
scale which is quite absent from those in the south.
The northern lakes have soft water; the fixed carbon
dioxid does not exceed 35 p.p.m., while in the southern lakes
it ranges from 65 to 100 p.p.m., and may go beyond the
larger figure.
Among the southern lakes only one had less than 15 per
cent of incident radiation present at one meter; and this
case (Kegonsa) was obviously due to growths of algae near
the surface. Among the northern lakes there were 11
cases of a very low per cent of radiation at one meter, from
8 different lakes. Only two southern lakes (Marl and Elk¬
hart) exceeded 31 per cent at this level, and two others had
percentages between 30 and 31. One northern lake
(Crystal) exceeded 35 per cent, one other (Weber) nearly
reached that number, three others (Clear, Pauto, Trout)
were above 31, and one (Clear Crooked) was above 30. All
grades may be found connecting these extremes.
Thus the northeastern lake district contains lakes which
may furnish typical examples of every type of factor af¬
fecting transmission of radiation. These are present in
considerable numbers for every type and they show every
grade within the type. The few lakes reported probably
offer cases close to the limits of transparency for inland
lakes. There are lakes much more deeply stained than any
in the list; and further study will yield a much more com-
Birge & Juday — Transmission of Radiation in Inland Lakes . 557
plete series and one in which the various factors may be
assigned more definite values.
The most deeply stained waters are those of Mary and
Turtle lakes. Lake Mary (1.1 ha., 22 m.) is a small, deep,
moraine lakelet; it normally overflows into Lake Adelaide,
but in 1928 there was no overflow and the upper meter or
two of water was not only deeply stained but also contained
much gummy extractive from peat. Turtle Lake lies in
the course of a small stream issuing from marshes; it has
much plankton as well as stain. Dead Pike Lake has much
stain, but little organic plankton. Lake Adelaide is high in
both; and both lakes are relatively deep. Allequash and
Wolf lakes are shallow and weedy and have much sus¬
pended vegetable debris. In all of these lakes the trans¬
mission is in general below 40.
There follows a group of lakes with colors ranging from
12 to 18, including Fishtrap, High, Kawaguesaga, and Lost
Canoe lakes. These have also much plankton, which has
much influence on transmission. This is usually below 50
and often below 40. When the plankton is low it may ex¬
ceed 60, as in Kawaguesaga Lake.
The lakes with low stain — 5 to 8 — have a transmission
which is rarely below 50, ordinarily exceeds 60 and may
touch 70 (Trout Lake), or even decidedly exceed 70 (Blue
Lake). It should be said that readings of color below 8
have to be estimated and may easily be lower than stated.
In the lakes which are regularly recorded as having no
stain the transmission depends on the amount of plankton
and the selective effect of water. The quantity of plank¬
ton is usually small ; if it were large the color of the water
would be affected. In these lakes the transmission does
not fall below 60 in any recorded case ; it ordinarily exceeds
70; it may rise above 80 and even touch 90 (Crystal Lake).
The observations on the Northeastern lakes for 1926 and
1927 are also given in the table found in Birge and Juday,
*29, p. 69 (p. 8 of separate) . In a few cases the figures there
given differ slightly from those reported in the present
paper. These variations are due to revision and recompu¬
tation and are without general significance. The series
given in table 10 for Crystal Lake, July 6, 1926, is not men¬
tioned in the earlier paper, either in the table or the discus-
558 Wisconsin Academy of Sciences , Arts, and Letters .
sion. This was the only case in which transmissions ap¬
proaching or exceeding 90 had been found and we were un¬
willing to accept them. In 1928 another series was taken
with transmissions essentially similar, and consequently
both are now included.
When the transmission thus increases in the deeper water
the question naturally arises whether this may not be due
to the removal of the shadow of the boat. Oberdorfer at¬
tributes a very considerable effect to this shadow (’28, p.
150). We are not prepared to discuss the question at pres¬
ent, though we have given it some attention. Examination
of table 10 will show cases where such an effect would be
suspected and other similar cases where it is obviously ab¬
sent. The effect would not be great if, as in our work, all
observations are made in bright sunhine. The matter is
also complicated by the fact that transmission must tend
to rise in the deeper strata, as the longer wave-lengths of
the spectrum are absorbed. The effect of possible reflec¬
tion from the side of the boat on the reading at one meter
must also be considered. This effect must be present to
some extent in readings taken as ours were, and it will vary
with the color of the boat.
Other Lakes
The transmission of radiation has been measured in four
lakes outside of Wisconsin; in Canandaigua, Cayuga, and
Seneca lakes in New York; and in West Okoboji Lake, Iowa.
The last named is a lake much like those of southeastern
Wisconsin ; having hard water, abundant plankton, and cor¬
respondingly low transmission. The New York lakes are
large, deep lakes, whose water has little stain and also little
plankton. They are lakes of the same type as Green Lake,
Wisconsin, but though they are much larger and deeper,
their transmission is not higher than that of Green Lake.
These three lakes may be taken as showing the usual situa¬
tion in lakes belonging to the type called oligotrophic. Trout
Lake in northeastern Wisconsin belongs to the same type,
so far as plankton, stain, and transmission are concerned.
In all there may be expected as much as 80 per cent of in=
cident radiation at one meter, or perhaps somewhat more,
and a transmission rising to 70 or above, but not reaching 80.
Birge & Juday — Transmission of Radiation in Inland Lakes . 559
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564 Wisconsin Academy of Sciences , Arts , and Letters .
It should be noted that the new calibration of the thermo¬
pile with the hemispherical cover (p. 516) adds 50 per cent
to the percentages reported for these lakes at the depth of
one meter. See Birge and Juday, ’20, ’21. It has a similar
effect on the calories reported for various depths. The
transmission remains the same as reported. The general
result is that one per cent of incident radiation is found in
Seneca Lake at the depth of 11.4 m. instead of at 10 m., as
reported in the earlier paper; in Cayuga Lake the shift is
from 8.6 m. to 9.4 m. ; in Canandaigua Lake from 7.5 m. to
8.6 m. ; in Okoboji Lake from 6.4 m. to 6.8 m. It is of
course understood that in stating these depths no great ac¬
curacy is claimed for the last figure in the number indicat¬
ing the depth since it is reached by prolonging the curve of
transmission, in most cases through three or four meters.
3. COMPARISON WITH WORK OF OTHER
OBSERVERS
The Wisconsin Survey seems to have been alone in using
the thermopile for measuring the penetration of solar ra¬
diation. Other observers have used photometers of various
kinds. In all cases the amount of radiation received in air
is the basis on which is computed the per cent left at dif¬
ferent depths below the surface.
Since these authors deal with light and we with total ra¬
diation, some ratio between these two bases must be found,
at least provisionally. However “light” may be defined,
that part of the solar energy spectrum included in it varies
with the altitude of the sun, with the transparency of the
air, with the quantity of precipitable atmospheric water,
and with other factors. We have obtained a partial notion
of this variation from tables kindly furnished us by Mr.
F. E. Fowle of the Smithsonian Institution. These enable
us to compare this value for air masses 1 to 5 and for 0.5
cm. to 2.5 cm. of precipitable water. The per cent of radia¬
tion found between 0.300/* and 0.764/x (practically the A
line) with air mass 1 and precipitable water 0.5 cm. is about
57.5; with air mass 5 and water 2.5 cm. it is 32.6 per cent;
and all intermediate results may be found as these two va¬
riables are given different values.
Birge & Juday — Transmission of Radiation in Inland Lakes. 565
With air mass 1.5 and water 2.0 cm. the value of the ra¬
diation in the region named is about 55.7 per cent; and this,
rounded off to 56, we have used in comparing our results
with those of others. The amount of precipitable water as¬
sumed is lower than our summer conditions ; and perhaps it
would have been as well to assume the rough value of about
50 per cent.
Solar radiation which has passed through one meter of
pure water has lost practically all radiation of greater wave
length than 0.764/*. If the absorption coefficients of Asch-
kimass are accepted, between one and two per cent of the
radiation left at one meter may come from the region be¬
tween wave lengths 0.764/* and 0.850/*. If the coefficient
of Nichols at 0.779/* is accepted, as in the Smithsonian
Physical Tables (p. 307), no appreciable part of the radia¬
tion comes from this region. Hence we have repeatedly
stated that the radiation measured by the pyrlimnometer at
the depth of one meter or more is “practically” all in the
form of light, as defined above.
Light may be more narrowly defined so as to exclude ra¬
diation from wave lengths shorter than 0.360/* or 0.400/*. In
that case the per cent of the total energy spectrum repre¬
sented by light is correspondingly reduced and a larger per
cent of radiation present at the depth of one meter lies in
the region thus excluded from light. But it is not neces¬
sary to discuss this aspect of the matter further.
From this statement it follows that observations with
thermopile and photometer will have certain points of gen¬
eral agreement and difference.
1. The percentage of light radiation shown to be present
at any depth by a photometer is greater than the percentage
of total radiation shown by the thermopile. The ratio will
be variable and dependent on the nature of the photometer
and the character of the water; but in clear water the
amount of radiation at any depth should be roughly twice
as large a per cent of light as of total radiation.
2. The transmission of radiation to the depth of one me¬
ter, as measured by the photometer should be correspond¬
ingly greater than the transmission measured by the ther¬
mopile.
566 Wisconsin Academy of Sciences , Arts, and, Letters.
3. The transmission below one meter should be about the
same when measured by either type of instrument; unless
the selective action of the photometer and the conditions
of stain or suspended matter in the water combine to pro¬
duce a different result. The observations with which ours
are to be compared were made in water free from stain and
in general with little plankton. A general agreement would
be expected in transmission as measured with photometer
and thermopile ; and in fact such an agreement is present.
Three types of photometer have been used for measuring
transmission of radiation in wafer; the selenium cell, the
photographic plate, and the photo-electric cell.
Selenium Cell
The earliest photometer to be employed was the selenium
cell, which was used by Regnard (’91, p. 216) to measure
radiation to the depth of 10 m. in the Mediterranean Sea.
One series only is recorded and this shows the defects of the
cell as a receiver. As pointed out by Shelford and Gail
(’22, p. 144) the results near the surface are too low and
at greater depths too high ; and they do not require further
discussion here.
Photographic Methods
Photographic plates or paper have been used by numer¬
ous observers and this method is, no doubt, the best for se¬
curing records of the feeble light which penetrates to great
depths. The method has not so often been applied to de¬
termining the transmission near the surface, meter by me¬
ter, and thus obtaining records which can be compared di¬
rectly with ours. There are three observers whose work
may be mentioned, Linsbauer, Klugh and Oberdorfer.
Linsbauer (’05) constructed an apparatus for employing
under water the photographic methods of Wiesner. He ap¬
plied it for measuring radiation in the Traunsee, and he,
therefore, furnishes one of the few records from fresh wa¬
ter. His mean results follow :
Table 12. Radiation in the Traunsee.
Depth in meters _ 0 0.5 1 2 3 5 10
Radiation, per cent _ 100 29 19 4,9 3.0 1.4 1.4
Birge & Juday — Transmission of Radiation in Inland Lakes. 567
These results, like those of Regnard are evidently too low
near the surface and too high at greater depths. It is possi¬
ble to imagine conditions under which there might be so
rapid and so great an increase of transmission with depth;
but such conditions, if present at all, must be very rare, and
the series does not show results comparable with ours.
The work of Klugh (’25, ’27) is quite different. He de¬
vised methods of using photographic plates in an instru¬
ment called the ecological photometer; and in certain cases
he employed this in the same way that we have used the
pyrlimnometer. Klugh’s first paper gives a full descrip¬
tion of the instrument and its use. It is sensitive to radia¬
tion from wave length 0.400^ to the end of the visible spec¬
trum (’27, p. 91).
Two of Klugh’s observations may profitably be compared
with ours. In his first paper (’25, p. 230) is a series of
readings to the depth of 10 m. in the Bay of Fundy near
St. Andrews, N. B. His standard of illumination is the
radiation at 2 cm. below the surface and the recorded per¬
centages are accordingly higher than would have been the
case if the original reading had been taken in air. It is
also true that the transmission in the first meter (or 98 cm.)
is much greater than would be expected from the transmis¬
sion below one meter. The per cent at one meter is record¬
ed as 65; while the transmission in the 1-2 m. stratum is
42 and in the meter below it is 44. Below 3 m. the trans¬
mission rises, but as there is an evident irregularity in the
reading at 4 m. the transmission cannot be calculated in de¬
tail. The average transmission between 3 m. and 10 m. is
about 53. The situation thus disclosed is quite like that in
our less transparent lakes, so far as transmission below 1
m. is concerned. The series might well have come from
Lake Mendota at rather less than average transparency
down to 3 m. and a little above the average below that depth,
down to 10 m.
In the paper of 1927 (p. 91) a similar series is given for
Chamcook Lake. This is a lake near St. Andrews, N. B.,
about 3 miles long and 35 meters deep. It lies “in a gran¬
itic region” and is a soft water lake with very clear water
and little plankton. Readings were taken at 0.5, 1, 3, 5,
and 10 m. The transmission between 1 m. and 5 m. is about
568 Wisconsin Academy of Sciences f Arts , and Letters .
uniform at 85; the interval between 5 m. and 10 m. seems
to have been slightly less transparent, averaging about 80.
If the reading at 10 m. had been 13 per cent instead of 10,
the transmission from 1 m. to that depth would have been
uniform; and a slight change in the solar radiation might
have caused this difference. But, accepting the record, the
upper water of Chamcook Lake was nearly as transparent
as the clearest strata of our Crystal Lake. Several of our
lakes have a transparency as great as that in the 5-10 m.
stratum, and the general record is one which we might ex¬
pect to duplicate in the clearest lakes of northeastern Wis¬
consin.
Klugh notes that the five readings in this lake required
the time from 11:00 a. m. to 3:15 p. m. We have never
used photographic methods and will not compare their ad¬
vantages and disadvantages with those of the thermopile.
But the pyrlimnometer has at least the advantage of greater
speed. Such a series of 5 readings would be completed in
about 10 minutes, if only one reading were taken at each
depth.
Oberdorfer (’28, ’28a) devised an apparatus for expos¬
ing a sensitive paper or film under a neutral-tint wedge, the
“Eder-Hechtsche Graukeil.” The apparatus is relatively
simple and permits rapid work, since a series of 12-14 ex¬
posures to the depth of 25 m. can be made in 5-10 minutes.
Oberdorfer not only determined the transmission of light,
but he also discusses the effect of the shadow of the boat,
a matter which we have not investigated. He also made an
ecological use of his photometric results by determining the
depths within which various species of algae are found and
the amount of light which they demand.
Oberdorfer’s work was done in the Bodensee (Lake Con¬
stance) during two years, January, 1926 to December, 1927
and extended to the depth of 25 m. The highest transmis¬
sion was found in March, when the average transmission
to the depth of 10 m. was 82.1 per meter; to 20 m. it was
82.7. The lowest was in August when the average trans¬
mission to 10 m. was 72.2 per meter, and to 20 m. it was
71.7.
Thus the water of the Bodensee has substantially the
same transmission as that of our more transparent deep
Birge & Juday — Transmission of Radiation in Inland Lakes. 569
lakes. Our observation in Seneca Lake (table 9) shows
that it belongs to the same general class as Lake Constance.
The average transmission found in Seneca Lake to 10 m. on
August 1 was about 71.3 per meter, practically the same as
that found in August by Oberdorfer. There is no reason
to doubt that there would be a similarly close correspond¬
ence at other seasons.
It may be noted that Oberdorfer found no exceptional
loss of light in the surface centimeters of the lake. In this
respect his results agree with ours and differ from the ob¬
servations made in the ocean.
The Photo-Electric Cell
Recent studies with the photo-electric cell have been made
by Shelford and Gail 022, p. 14), Poole and Atkins 027,
p. 177 ; '28, p. 455), and by Shelford and Kunz 026, p. 284) .
Each supplies data which can be compared directly with
ours. The instrument which they employed is far more
sensitive than the thermopile and their observations extend
to much greater depths than ours. Shelford and Gail took
readings as deep as 40 m. in Puget Sound; and Poole and
Atkins went to depths of 65 m. in the sea near Plymouth,
England. Their studies furnish numerous series of obser¬
vations with whose readings in the upper 10 m. we may
compare ours. It will be seen that there is a close general
agreement so far as our readings extend and that the deeper
readings in the sea show that the general conditions of the
upper water continue in the deeper strata.
Shelford and Gail (table 5, p. 155) give the transmission
in Puget Sound to the depth of 10 m. as determined from
16 series of observations. Our observations on Crystal
Lake, the mean of 6 series, may be compared with theirs in
the following table.
570 Wisconsin Academy of Sciences , Arts, and Letters.
Table 13. Per cent of radiation present in Puget Sound (P) and
Crystal Lake (C).
Per cent of Transmission
In our observations the percentages present are given in
terms of total radiation ; those of Shelf ord and Gail in terms
of light. We may recompute our results as light, since only
short wave radiation remains at depths of one meter and
below. We have assigned 56 per cent of the solar energy
spectrum to the region between 0.300/x and 0.762/x. This is
certainly quite large enough. The results are given in the
third line of the above table and they show that Crystal
Lake and Puget Sound may yield almost identical results.
The transmission meter by meter is also practically the
same. Two series from the same water would not be likely
to show a closer agreement than do these, of which one is
from salt water and the other from fresh. Figure 11 shows
the transmission curves corresponding to table 13 and their
parallel course is obvious.
Poole and Atkins also worked with the photo-electric cell
and their observations extended to 65 m. Both of their
papers report series which can be compared with ours. The
paper of 1928 reports a larger number of series which they
tabulate in detail and show in fig. 3, p. 476. The curves
show on a large scale for depth exactly the same phe¬
nomena as do ours to the depth of 10 m. They show also
that the curves may be essentially straight lines when plat¬
ted on semi-logarithmic paper, and that in general they con¬
tinue at greater depths the same course that they have
taken in the upper water. They also show the same kind
of changes and irregularities of transmission that we have
found at our lesser depths. The series reported in 1928 fall
Birge & J uday — Transmission of Radiation in Inland Lakes. 571
into two groups, one with low transmission and one with
high.
We may compare the average transmission of some of our
lakes with that in the upper 10 m. of the curves in the group
with low transmission. The average transmission in these
cases, when measured over a straight run of 6 m. to 9 m. is
from 68 to 77. The lowest is found in series 12 and 18,
where transmission is from 68 to 71; that of Nagawicka,
Per Cent
t 2 3 4 5 6 7 8 iO 15 20 30 40 50 60 80 106
c p c
Fig. 11. Transmission curves of radiation in the water of Puget
Sound and of Crystal Lake, Wisconsin. The curve from Puget
Sound (P), is the mean of 16 series of observations by Shelford and
Gail; that from Crystal Lake (C) is the mean of 6 series, reported
in table 9. The curves are parallel, but the percentages determined
by the pyrlimnometer are lower than those by the photo-electric cell,
since one set is based on total solar radiation and the other on light.
In the curve marked C' the percentages from Crystal Lake are re¬
computed as light, assigning 56 per cent of solar radiation to light.
Pine, and Cayuga lakes is just below 70 and readings of
about 70 are found in Green, Trout, and Seneca lakes. Se¬
ries 11 and 14 show a transmission of 74-75, about like
that reached by Devils, Elkhart, and Geneva lakes. Series
9 has about 77 which compares with the upper part of Marl
Lake.
In the other group of series (16-22) in the report of
Poole and Atkins — that with high transmission— -the per
Meters
572 Wisconsin Academy of Sciences , Arts, and Letters .
cent ranges from 85 to 90. This is a condition reached only
by Crystal Lake in our list, and is reached by that lake only
under conditions of exceptional clearness. (See table 9).
The lake belongs to the same type, but only just reaches it.
In nearly all of our lakes the transmission in the 2-3 m.
stratum is smaller than that below. (See fig. 7). This is
notably true also of certain series reported by Poole and
Atkins, such as series 13 and 14. In other cases this con¬
dition is not present. Less uniform conditions of trans¬
mission would be expected in our lakes, small and without
currents, than in the sea with its large waves and vigorous
currents due to wind and tide.
In 1925 observations with various types of photo-elec¬
tric cells were made by Shelford and Kunz in Lake Men-
dota to the depth of 10 m. Their paper (’26, p. 290) shows
the transmission meter by meter as measured by different
cells and with various ray filters. The results are also
shown in fig. 9, p. 298. The transmission is much the same
as is shown by the pyrlimnometer. The readings of the
potassium cell without a ray filter show transmission from
33 to 66; a result quite comparable to ours, although the
curve is unusually irregular.
It thus appears that the photometer and the thermopile
give similar results when employed for the measurement of
the transmission of radiation through the water of lakes.
Transmission is of the same order of value and platted
transmission curves run parallel whenever observations by
the two types of receiver are compared. This is true for
waters devoid of stain and with little plankton. No read¬
ings have been made with the photometer in lakes with
deeply stained water, and only one series in Lake Mendota,
the lake which may represent the larger part of those which
we have studied.
There remains to be compared the transmission in the
upper meter of the water as measured by photometer and
thermopile. On this transmission, combined with the
losses at the surface, depends the per cent recorded as pres¬
ent at one meter. The two types of instrument yield dif¬
ferent results and different forms of photometer should also
give different results, according as their sensitivity extends
Birge & Juday — Transmission of Radiation in Inland Lakes. 573
more or less into the red end of the visible spectrum. In
any case the thermopile must give a smaller per cent, since
in air it registers the total solar radiation, which is about
twice as great as the radiation in the form of light, and of
which almost all except the light is lost in the first meter.
In table 13 and figure 11 the transmission curve of Crys¬
tal Lake is transposed from the basis of total solar radia¬
tion to that of light radiation. The part of the spectrum
between 0.300 /*, and 0.762/*, is reckoned as light and com¬
puted as 56 per cent of the total radiation. This change
brings the curve for Crystal Lake very close to that for
Puget Sound, but the apparent correspondence is closer
than is warranted by the probable facts. The red part of
the spectrum, included by us in the 56 per cent designated
as light, is rapidly absorbed by water, while the photo-elec¬
tric cell responds only to those wave lengths to which water
is most transparent. It would seem, therefore, that the
per cent reported by Shelford and Gail is too low or else
ours is too high. Poole and Atkins ('25, p. 195) suggest
that low readings may be caused by the concave lens formed
by the water-glass surface of the photo-electric bulb used
by Shelford and Gail. We have already said that our read¬
ings are probably too high as a result of leaving the diffuse
radiation out of account.
The transmission curves of the series reported by Poole
and Atkins may be extended upward to one meter from the
depth of 1.1 m. to 1.5 m. where the upper reading was
made. In that case the curve for series 14 shows that
about 37 per cent of total illumination was present at one
meter. Series 16 gives about 70 per cent at that depth
and series 6 from the paper of 1927 about 75 per cent. The
others would come between these extremes. These figures
would correspond roughly to 20-35 per cent of total radia¬
tion as shown by the pyrlimnometer and such a rough cor¬
respondence is indicated in the several cases whose trans¬
mission has just been discussed. It is not worth while to
say more on this subject at present, especially as both of
the papers reporting the results of using the photo-electric
cell record also large losses of light at the surface. This
part of the subject has not been especially investigated in
our work. The readings which we have taken in several
574 Wisconsin Academy of Sciences , Arts , and Letters.
cases at the depth of 10 cm. have not disclosed any such
large and irregular losses at the surface as those reported
by Shelf ord and Gail ; but our observations are by no means
numerous enough to constitute a study of the subject.
4. Literature Cited
For an historical account of earlier work and a good bib¬
liography, see especially Klugh 1925; for a discussion of
optical and physical principles, see Shelford and Gail 1922,
and Poole and Atkins 1926.
Aufsess, Otto von, 1903. Die Farbe der Seen. Inaugural
Dissertation. 64 pp. Miinchen.
Birge, E. A., 1922. A second report on limnological ap¬
paratus. Trans. Wis. Acad. Sci., Arts and Let. 20:
532-551.
Birge, E. A. and Juday, C., 1911. The inland lakes of Wis¬
consin. 1. The dissolved gases of the water and their
biological significance. Bui. No. 22, Wis. Geol. and
Nat. Hist. Survey. 259 pp. Madison.
Birge, E. A. and Juday, C., 1920. A limnological recon¬
naissance of West Okoboji. Univ. of Iowa Studies,
Nat. Hist. 9:1-56. Iowa City.
Birge, E. A. and Juday, C., 1921. Further limnological
observations on the Finger Lakes of New York. Bull.
Bureau of Fisheries. 37 :211-252. Washington.
Birge, E. A. and Juday, C., 1929. Penetration of solar
radiation into lakes, as measured by the thermopile.
Bull. Nat. Research Council. No. 68:61-76. Wash¬
ington. This paper is essentially a partial summary
of the present paper, based on results from northeast¬
ern Wisconsin.
Juday, C., 1914. The inland lakes of Wisconsin. II. The
hydrography and morphometry of the lakes. Bui. No.
27, Wis. Geol. and Nat. Hist. Survey. 137 pp. Madi¬
son.
Kimball, H. H., 1920. Variations in total and luminous
solar radiation with geographical position in the
United States. Monthly Weather Review. 47:769-
793. Washington.
Birge & Juda/y — Transmission of Radiation in Inland Lakes . 575
Klugh, A, B„ 1925. Ecological photometry and a new in¬
strument for measuring light. Ecology 6 :203-237,
Klugh, A. B., 1927. Light penetration into the Bay of
Fundy and into Chamcook Lake, New Brunswick.
Ecology 8:90-93.
Linsbauer, L., 1905. Photometrische Untersuchungen ii.
d. Beleuchtungsverhaltnisse im Wasser. Sitzber. d.
Akad. d. Wissens. Wien 114 :51-76.
Oberdorfer, E., 1928. Ein neuer Apparat zur Lichtmes-
sung unter Wasser. Archiv fur Hydrobiol. 20:134-
162.
Oberdorfer, E., 1928a. Lichtverhaltnisse und Algenbesied-
lung im Bodensee. Zeitschrift fur Botanik. 20 : 465-
568.
Poole, H. H. and Atkins, W. R. G., 1926. On the penetra¬
tion of light into sea-water. Jour. Mar. Biol. Assoc.
14: 177-198.
Poole, H. H. and Atkins, W. R. G., 1928. Further photo¬
electric measurements of the penetration of light into
sea-water. Jour. Mar. Biol. Assoc. 15: 455-483.
Regnard, P., 1891. Recherches experimentales sur les
conditions physiques de la vie dans les eaux. 500 pp.
Paris.
Rickett, H. W., 1922. A quantitative study of the larger
aquatic plants of Lake Mendota. Trans. Wis. Acad.
Sci., Arts and Let. 20 : 501-527.
Schmidt, W., 1915. Strahlung u. Verdunstung an freien
Wasserflachen, u. s. w. Ann. d. Hyd. u. Marit. Meteo-
rologie 42 : 111-124, 169-178.
Shelford, V. E. and Gail, F. W., 1922. A study of light
penetration into sea water made with the Kunz photo¬
electric cell with particular reference to the distribu¬
tion of plants. Pub. Puget Sound Biol. Sta. 3: 141-
176.
Shelford, V. E. and Kunz, J., 1926. The use of photo-elec¬
tric cells of different alkali metals and color screens in
the measurement of light penetration into water.
Trans. Wis. Acad. Sci., Arts and Let. 22 : 283-298.
576 Wisconsin Academy of Sciences, Arts , and Letters.
Appendix
Determination of the mean path of radiation in Lake Mendota.
By the “mean path of radiation” we understand the average dis¬
tance through which solar radiation travels in passing through a
stratum of lake water one meter in vertical thickness. This mean
path may be computed for radiation delivered to the lake during any
desired period of time. If all radiation came direct from the sun
and if the sun were fixed in the sky, this path would be the secant
of the angle of refraction for parallel rays reaching the surface of
the lake from the sun. But since radiation is both direct and dif¬
fuse and since the sun constantly travels over the sky, the problem
of the mean path is more complicated.
Since one main purpose of the study of radiation was to ascertain
the effect of direct insolation on the warming of lakes, it was ad¬
visable to determine this mean path for the warming period of Lake
Mendota. This incidentally involves the solution for 5 months of
the year, April to August inclusive.
The general result is to assign to the radiation reaching the lake
during this period a mean path in the water of about 116 cm. in
passing through a stratum 100 cm. in vertical thickness. This cor¬
responds to the path of direct radiation from the sun at the altitude
of about 47.5°. In tables 2, 7, and 8, this result is employed and is
referred to as “mean sun”. This expression agrees in form with
“zenith sun” in the same tables and means that all radiation is re¬
garded as direct, and that during the warming period such radiation
has an average path corresponding to that from the sun at the alti¬
tude stated above. The application of this result to Lake Mendota
is explained on p. 546.
The following statement gives an account of the method of com¬
putation.
A. The warming period of Lake Mendota is taken as extending
from April 15 to August 15. The earlier day marks the mean date
when the lake reaches the temperature of 4.0° C.; there is usually no
increase of temperature after the middle of August.
B. The records of the United States Weather Bureau at Madison
show (1) the total radiation received from sun and sky on a hori¬
zontal surface. This is recorded automatically by a Callendar
sunshine recorder and is stated in calories per square centimeter
per hour. (2) The duration of sunshine, recorded automatically by
a Marvin sunshine recorder and stated in the records as per cent of
sunshine for each hour of the day.
C. In the record sheets from April to August, inclusive, and from
1911 to 1920 inclusive, all hours were marked in which sunshine was
recorded for the entire hour. From these data was computed the
value of a mean sunshine hour for each hour of each day and each
month. The records of the Weather Bureau show the mean per
cent of sunshine for each hour of each month. Hence could be com-
Birge <& Juday— Transmission of Radiation in Inland Lakes . 577
puted the amount of radiation received from sun and sky during
each hour. The difference between this amount and the total radia¬
tion recorded is that received from sky during cloudy hours. It was
assumed that the radiation from cloud came in equal amount from
each unit area of sky. See Fig. 12 and table 15.
D. The mean altitude of the sun for each hour of each month was
determined from tables giving the altitude for each hour on the 1st,
10th, and 20th of each month.
E. The mean hourly radiation from sun and sky was platted as a
curve on coordinate paper and the curve of the sun’s altitude was
platted on the same sheet. The amount of radiation received be¬
tween successive intervals of solar altitude was thus determined.
See Fig. 12 and table 16.
F. The percentage of sky radiation was computed from data given
by Kimball (’19, p. 777) as follows:
Altitude of sun _ 60.0° 41.7° 30.0 ' 23.5° 19.3° 14.3° 10.2°
Per cent from sky 16 16 19 21 24 27 37
The per cent of radiation reflected from the surface of the lake
was computed from data given by Schmidt (’15, p. 171) as follows:
Altitude of sun 90° 80° 70° 60° 50° 40° 30° 20° 10° 0°
Per cent reflected 2.04 2.04 2.06 2.14 2.45 3.36 5.97 13.35 34.79 100.0
The loss of diffuse radiation by reflection, assuming that unit areas
of sky send out equal quantities of radiation, is 17.3 per cent
(Schmidt, 15, p. 176).
G. The length of the path of direct radiation in reaching the depth
of 100 cm. below the surface was computed for sun’s altitude 5°, 15°,
etc.
H. From these data the mean path of direct radiation was deter¬
mined for each month and for the whole period. (Tables 17 and 18.)
I. The diffuse radiation was computed on the assumption that equal
areas of sky send out equal amounts of radiation. The hemisphere
was divided into 10° zones whose area was determined; the percent¬
age reflected and the path in the water was computed for the center
of each zone. The mean path of diffuse radiation was thus deter¬
mined as 119 cm. in reaching the depth of 100 cm. below the surface.
See Poole and Atkins ’27, p. 184, table 1.
J. For direct radiation and for direct and diffuse combined the re¬
sults are shown in the following table.
37
578 Wisconsin Academy of Sciences, Arts, and Letters.
Table 14. Mean path of radiation, April-August.
The mean path for the period is based on the mean path and the
mean number of calories received during each month. It may be
more exactly stated as 115.9 cm. If the period is limited to April 15-
August 15 the mean path is 115.6 cm.
K. This computation is evidently approximate. The division be¬
tween sun and cloud is not very accurately given by the Marvin re¬
corder, which records “Cloud” when the sun does not cast a shadow,
and the amount of illumination at which the sun ceases to cast a
shadow is determined by the judgment of the observer. It is ap¬
proximately true that equal areas of sky send out equal amounts of
radiation when the sky is covered by a uniform layer of clouds; but
this is by no means true when the sky is partly covered by clouds,
whether these obscure the sun or not. The estimated percentages of
sky radiation assume a sky free from clouds. There seem to be no
measurements of sky radiation under average conditions of mixed
sky and cloud, or with haze and smoke. Kimball’s estimates of sky
radiation are, therefore, lower than average figures would be. In¬
crease of the per cent of sky radiation would reduce correspondingly
the amount of direct radiation, and would increase the amount re¬
flected and the mean path in the lake.
On the other hand, if all radiation were diffuse the mean path
would be 119 cm.; if all were direct the mean path for any month
in the period would not be less than 113 cm. (table 14) and for the
whole period it would be about 115 cm. Thus, the possible error in the
accepted result of 116 cm. cannot be so great as seriously to affect
the purposes for which it is used. It will be found that any reasonable
assumption as to the ratio between direct and diffuse radiation will
leave the result close to 116 cm. The same may be said of the re¬
sults given for the single months in table 14.
L. We have not studied the change of direction of radiation and
consequent change of length of path, due to dispersion by particles
suspended in the water. We have not considered the matter of radia¬
tion directed upward or laterally in the water.
M. Figure 12 and the following tables are added in order to show
the methods of computation in certain cases. Figure 12 shows the
diagram from which is derived the distribution of calories in June
Birge & Juday — Transmission of Radiation in Inland Lakes. 579
by altitude of sun. The computation is made for a mean day and the
total calories given in table 16 are derived from this result. Table
15 shows the computation for June.
In this table the data given under “per cent of sunshine” and “total
calories” are taken from the records of the United States Weather
Bureau. “Calories, full sunshine” are computed as stated in this
note. “Calories, cloud” are derived by difference.
Fig. 12. Distribution of calories by hour and by altitude of sun,
mean day, June, 1911-1920. The upper curve gives the course of the
sun; T represents total calories; S represents calories from sun;
and SS represents those from sun and sky. Calories for the cor¬
responding hours of morning and afternoon are combined; the day
is divided by sun time. The time is given at the bottom of the dia¬
gram; the numbers 1, 2, etc. on the side represent hundreds of cal¬
ories or successive increases of 10° in the altitude of the sun. Ver¬
ticals drawn from the solar altitude line intercept on the calorie lines
the number received during the passage of the sun through the 10
degree intervals.
580 Wisconsin Academy of Sciences, Arts, and Letters ,
Table 15. Computation of mean daily calories for June 1911-1920.
Table 16. Calories received by Lake Mendota from sun and sky and
from cloud.
Table 17. Computation of direct radiation and its path in the lake.
Notes — “Total calories” are taken from table 16.
“Per cent sun ’ ’ comes from Kimball’s data p. 000.
“Per cent to lake” equals 100 per cent less the percent reflected.
Table 18. Final computation of mean path.
Calories from cloud _ 8,740
Calories from sky _ 10,043
Total diffuse radiation _ 18,783
Reflected at 17.3 per cent _ _ _ _ _ 3, 249
Diffuse radiation to lake . 15,534, or 2S.2 per cent
Direct radiation to lake _ _ _ _ , _ 51,579, or 76.8 per cent
Path, diffuse radiation 119 cm.
Path, direct radiation 115 cm.
Mean path 115.9 cm.
A PRELIMINARY REPORT ON THE GROWTH OF
THE ROCK BASS, AMBLOPLITES RUPESTRIS
(RAFINESQUE), IN TWO LAKES OF
NORTHERN WISCONSIN.1
Stillman Weight
Assistant Aquatic Biologist
U. S. Bureau of Fisheries
I. Introduction
For several years the Wisconsin Geological and Natural
History Survey has been making a limnological study of
lakes in the northern part of the State. Because of the fact
that so much has been learned of the physical, chemical and
biological conditions in these lakes, the region seems par¬
ticularly favorable for a study of the growth rates of fishes
in relation to environmental factors.
The U. S. Bureau of Fisheries undertook such a study in
the summer of 1927 and continued the collection of material
in the summer of 1928. Other duties have prevented the
writer from spending all of his time on the problem so that
much of the material has not been examined. However,
work on the rock bass collected in Trout and Muskellunge
Lakes during both summers has been completed, and since
the Bureau has found it necessary to discontinue the in¬
vestigation for the time being, it seems advisable to publish
a preliminary report of the results.
The data are admittedly inadequate and any conclusions
reached must be tentative. The number of individuals
and the number of age groups represented must be greatly
increased to form the basis of final conclusions. The Wis¬
consin Geological and Natural History Survey plans to con¬
tinue the work and we may confidently expect further in¬
formation on the growth of the rock bass.
No attempt has been made to correlate growth of the rock
bass with environmental factors. The data of this paper
1 Published by permission of the U. S. Commissioner of Fisheries.
582 Wisconsin Academy of Sciences, Arts, and Letters .
indicate that, in all probability, local races exist among the
rock bass of Trout Lake, and that each race has its own
characteristic growth rate. If this is true, then it is ap¬
parent that in order to correlate the growth of the fish with
factors in the environment, it is necessary to study these
factors at each collecting station. In the lakes considered
here, limnological observations were made only in the
deeper areas and not in the protected, shallower areas
where the rock bass were captured.
The writer takes this opportunity of thanking Mr. Chan-
cey Juday, who suggested the problem and made available
equipment and space in the field laboratory of the Wiscon¬
sin Geological and Natural History Survey at Trout Lake.
Dr. John Van Oosten, U. S. Bureau of Fisheries, by his sug¬
gestions and critical examination of the data, has made an
invaluable contribution to the work. Thanks are also due
Dr. Carl L. Hubbs, Curator of Fishes, University Museums,
University of Michigan, who has offered many helpful sug¬
gestions.
II. The Lakes
Trout and Muskellunge Lakes are situated in morainal
basins in the western part of Vilas County, Wisconsin.
Trout Lake is much the larger and deeper, but in many re¬
spects the two lakes are similar — both are deep enough for
thermal stratification and both have rather extensive areas
covered with large aquatic plants.
The following data on Trout Lake have been taken from
Juday (1914) :
Location: Towns 41 and 42 North, Range VI and VII East.
Length: 7.24 km. (4.5 miles) in N-S direction.
Width: 3.86 km. (2.4 miles).
Area: 16.83 sq. km. (6.5 sq. miles).
Depth: 35 meters (115 feet).
No data have been published on Muskellunge Lake: the
information given below was taken from the unpublished
records of the Wisconsin Geological and Natural History
Survey :
Wright — Growth of Rock Bass in Two Lakes of Wisconsin. 583
Location: Town 41 North, Range VII East.
Length: 3.3 km. (2.05 miles) in NE-SW direction.
Width: 1.18 km. (0.73 miles).
Area: 3.74 sq. km. (1.45 sq. miles).
Depth: 19.3 meters (63.3 feet).
III. Materials and Methods
Most of the fish were taken in gill nets of various sizes
of mesh. It was found that seines could not be used effect¬
ively because of snags along the shores, although a few fish
were taken by this method. A small number of fish were
also captured by hook and line.
With few exceptions, the fish captured in Trout Lake
during 1927 were taken in Blaisdell’s Bay, which is located
at the extreme southeast corner of the lake, between the
point of land known as Rocky Reef and the mainland on
the east. In 1928 the collections were made on Camp
Franklin Shoal, which is located between Crescent Island
and the mainland at the point on which Camp Franklin is
situated. The shoal is approximately one mile west of
BlaisdelFs Bay, from which it is separated by Rocky Reef
and an area overlain by relatively deep water.
With the exception of a few individuals, the fish in Mus-
kellunge Lake were taken along the north shore between
the Musky Cottages on the east and Camp Osoha on the
west. So far as is known there are no barriers which
would prevent migration of the fish along this shore.
Standard length measurements in millimeters were made
on a measuring board similar to the one described by
Thompson (1916), and a sample of the scales of each fish
was taken for age determination. No weights were re¬
corded for the 1927 fish, but in 1928 weights were deter¬
mined in grams by means of a Chatillon balance. These
data are not included in this report.
The method known as the scale method was used to de¬
termine age and growth rates. The reader may refer to
Van Oosten (1923 and 1929) and to Creaser (1926) for a
discussion of the method. Up to the present, no one has
tested the soundness of the scale theory on the rock bass,
but since Creaser (1926) has found it usable for the closely
related pond sunfish, Eupomotis gibbosus, and since other
584 Wisconsin Academy of Sciences, Arts, and Letters.
authors have found it valid for many species of fish, the
writer has accepted it as valid here.
Scales were mounted in a glycerine-gelatine jelly on mi¬
croscopic slides and the image projected, at a magnification
of about 40, on the ground-glass plate of the apparatus de¬
scribed by Van Oosten (1923). The rock bass has a typi¬
cal ctenoid scale and the annuli are readily located on most
of the scales. Of the 418 scale samples examined, 22 were
discarded because the age could not be determined with con¬
fidence. Although most of the discarded scales were taken
from fish which belonged in age groups discussed in this
paper, the number discarded (5.2%) is considered too small
to affect the results markedly.
In order to calculate the length of a fish at the end of
each year of its life, measurements were made of the dis¬
tance of each annulus and the border of the scale from the
focus. The measurements were made to the nearest milli¬
meter along the anterior radius. The border of a scale was
taken to be the outermost circulus, thus excluding the clear
marginal area. In making calculations, the following for¬
mula was employed :
L x Sj
Li = - >
S
where LA = length of the fish at the end of year N,
L = length of the fish at the time of capture, Sx = length
of anterior radius of scale at end of year N, and S — length
of anterior radius of scale at the time of capture. This
formula is believed to give only approximate results since
it is based on the assumption that the anterior radius of
the scale of the rock bass increases in length proportion¬
ately with the length of the fish, which assumption is prob¬
ably only approximately true. No correction was made
for Lee’s phenomenon or for the fact that the fish attains
a certain length before forming scales. Since only rela¬
tively old fish are employed, Lee’s phenomenon may not be
an important factor in the calculated length data.
Ages are designated by Roman numerals ; a fish taken in
the first summer of its life and which shows no annulus on
Wright — Growth of Bock Bass in Two Lakes of Wisconsin. 585
its scales, belongs to age group I; a fish taken in the sec¬
ond summer of its life and which shows one annulus be¬
longs to age group II.
IV. Discussion
a. Selection of data
Tables 1 and 2 give the number of fish in each age group
taken from Trout and Muskellunge Lakes in 1927 and 1928,
with the exception of 22 fish which were discarded be¬
cause their scales could not be read with confidence.
Table 1. Trout Lake. Number of fish in each age group taken in
1927 and 1928.
♦Taken in seine.
Table 2. Muskellunge Lake. Number of fish in each age group
taken in 1927 and 1928.
It is evident that most of the fish fall into a few age
groups in each collection, a fact which is explained by the
selective action of the gill nets. The obvious difference in
age composition of collections from the two lakes will be
commented on later. Not only does a gill net select fish of
certain lengths, but it seems certain that it selects the
smaller individuals of the oldest and the larger individuals
of the youngest age group taken. Such selected individ¬
uals are not representative of their year classes and must
be discarded in a study of the average growth rate of a pop-
586 Wisconsin Academy of Sciences , Arts, and Letters.
ulation. Not only must representative age groups be
chosen, but for a comparative study, only those representa¬
tive age groups which were taken in adequate numbers in
both lakes. On this basis, fish of age groups VI, VII and
VIII seem to meet the requirements for a comparative
study.
There may be some question whether the fish of age
group VI in the four collections are really representative
of their respective year classes. In the 1927 collection
from Trout Lake, fish of age group V are discarded and it
follows that the fish of the sixth age group are in all prob¬
ability representative. No fish of the fifth age group were
taken in the 1928 collection, but the sixth age group is con¬
sidered representative because the average length of the
Vl-year fish at the time of capture is greater than that of
the Vl-year fish of the 1928 collection from Muskellunge
(tables 4 and 6). Since gill nets of the same size mesh
were used in both lakes, it seems probable that V-year fish,
with an average length less than that of the Vl-year fish,
would have been taken in Trout Lake, had they been pres¬
ent. The fact that only two V-year fish were taken in the
1927 collection from Muskellunge Lake suggests that the
Vl-year fish are not representative. If this were true,
then the assumed fact that the Vl-year fish were the larger
individuals of their year class, combined with the influence
of Lee’s phenomenon, should make their calculated lengths
greater than those of the same year class (age group VII)
of the 1928 collection. Comparison of the calculated
lengths of age group VI (table 5) and age group VII (table
6) does not support this contention. It might also be
claimed that the Vl-year fish of the 1928 collection from
MuskelJunge Lake were the larger individuals of their year
class, but the calculated length data in table 6 give no evi¬
dence to support the claim.
b. Typical growth curve
Tables 8, 4, 5, and 6 show the average actual and calcu¬
lated lengths, in millimeters for each year of life, for fish
of age groups VI, VII and VIII taken from Trout and Mus¬
kellunge Lakes in 1927 and 1928 respectively. They also
Wright — Growth of Rock Bass in Two Lakes of Wisconsin. 587
show, for each year of life, the averages resulting from
combining the calculated lengths of all three age groups,
and the average annual increments derived from these cal¬
culated lengths.
Table 3. Trout Lake , 1927. Number of fish, average actual and cal¬
culated lengths in millimeters, combined average calculated
lengths and annual increments for fish of age groups VI, VII and
VIII.
Table 4. Trout Lake, 1928. Number of fish, average actual and cal¬
culated lengths in millimeters, combined average calculated
lengths and annual increments for fish of age groups VI, VII and
VIII.
Table 5. Muskellunge Lake, 1927. Number of fish, average ac¬
tual and calculated lengths in millimeters, combined average cal¬
culated lengths and annual increments for fish of age groups VI,
VII and VIII.
588 Wisconsin Academy of Sciences , Arts, and Letters.
Table 6. Muskellunge Lake, 1928. Number of fish, average ac¬
tual and calculated lengths in millimeters, combined average cal¬
culated lengths and annual increments for fish of age groups VI,
VII and VIII.
Examination of the grand averages of annual growth in¬
crements for the four collections brings out certain general
features of the growth history of the rock bass. It will be
seen that the second year invariably shows a decline in rate
of growth over the first year; that in general the incre¬
ments later increase progressively to the highest point in
the fifth year of life (in the sixth year in table 6) , and then,
with one exception (table 4), decline to the lowest point in
the seventh year. Thus, the growth curve typically has
the form of a double sigmoid (fig. 1). The curves shown
by Creaser (1926, fig. 11) indicate the same type of growth
for the pond sunfish, Eupomotis gibbosus . Virtually all
age groups (table 7, this paper), though not all individual
fish, show this type of growth, so it may be stated that, in
general, the double sigmoid growth curve is characteristic
of the rock bass. The exception noted in table 4, where the
grand average increments reached the highest rather than
the lowest point in the seventh year, is explained by the un¬
usually slow growth of the VH-year fish throughout life.
This slow growth resulted in a lowering of the grand aver¬
ages of calculated lengths, and consequently of increments,
for each year of life except the seventh. The average an¬
nual increments of this VH-year group (table 7), however,
exhibit the typical rock bass growth curve.
The grand average increments show further that the
second hump of the double sigmoid curve, which usually
reaches its highest point in the fifth year of life, is greater
than the first, which occurs in the first year of life. That
is, the grand average increments of the fifth year of life
Wright — Growth of Bock Bass in Two Lakes of Wisconsin. 589
are consistently greater than those of the first year; and
with two exceptions (tables 4 and 6) are greater than those
of any other year of life. Table 7 shows that in six of the
twelve age groups the greatest growTth occurred in the fifth
year, and that of the remaining six age groups, in five the
greatest growth occurred in the fourth or sixth years, or
was equally great in the fifth and sixth, or in the fourth,
fifth and sixth.
This fact is more strikingly demonstrated from a dif¬
ferent point of view in table 8, which shows the distribu¬
tion of individual fish of age groups VII and VIII accord¬
ing to the year of life in which the greatest growth oc¬
curred. The fish of age group VI were not included in the
table because individuals of that age had no opportunity to
demonstrate their rate of growth in the sixth year of life.
If included, they would only contribute to the number hav¬
ing the greatest growth in the first five years of life. It
appears proper to include the VH-year fish for the reason
that not a single V Ill-year fish grew most rapidly in its
seventh year of life. It will be seen that, of the 200 fish,
78 (87%) grew most rapidly in the fifth year of life, 54
(27%) grew most rapidly in the sixth year, and 84 (17%)
in the fourth year. Only 39 (20%) of the 200 fish showed
the most rapid growth in years other than the fourth, fifth
and sixth.
Table 7. Trout and Mushellunge lakes , 1927 and 1928. Average
growth increments, by age groups, of fish in age groups VI, VII
and VIII. (Data derived from average calculated lengths in
Tables 3, 4, 5 and 6).
590 Wisconsin Academy of Sciences , Arts, and Letters.
Table 8. Trout and Muskellunge lakes, 1927 and 1928. Distribution
of indioidual fish of age groups VII and VIII according to the
year of life in which greatest growth occurred.
In most fresh-water fish the most rapid growth takes
place in the early years of life and there is a progressive
decrease with increase in age. Two explanations have
been offered for the occurrence of rapid growth at a rela¬
tively late period in the life of the rock bass. The growth
curves shown by Greaser (1928, fig. 11) seem to indicate
that the most rapid growth of the pond sunfish occurs in
the two years just preceding that in which sexual maturity
is reached. There is a possibility that an internal adjust¬
ment associated with the approach of sexual maturity
would result in an increase of the rate of growth, but there
are no data on the rock bass which bear on that question.
Another suggestion offered is that when the rock bass at¬
tains a certain length, there is a change in the type of food
eaten, which results in an increase in the growth rate.
Whatever the true explanation for rapid growth rela¬
tively late in life may be, there is some evidence that the
influence of conditions in the environment may determine
the particular year in which the most rapid growth takes
place. More individual fish from Muskellunge Lake grew
most rapidly in 1925 than in any other calendar year cov¬
ered in the collections. Nineteen of 25 Vl-year fish, 19 of
45 VH-year fish, and 11 of 21 VUI-year fish of the 1927
collection; 4 of 38 Vl-year fish, 10 of 29 VH-year fish, and
14 of 36 VUI-year fish of the 1928 collection grew most
rapidly in the calendar year 1925. In this respect the re¬
maining fish are well distributed through the other calen¬
dar years. Trout Lake fish give no evidence of having
grown most rapidly in 1925 or in any other calendar year,
Wright — Growth of Rock Bass in Two Lakes of Wisconsin. 591
and since a favorable year for Muskellunge Lake rock bass
would, in all probability, be equally favorable for those in
Trout Lake, the evidence cited is not regarded as highly
significant.
c. Comparison of growth rates of populations in
in Trout and Muskellunge Lakes
A comparison of the average calculated lengths of the
age groups taken from Trout Lake in 1927 (table 3), with
corresponding age groups taken in the same year from
Muskellunge Lake (table 5), shows that, in every year of
life and in each of the three age groups, the Trout Lake
rock bass attained the greater length. The grand aver¬
ages of calculated lengths in these tables (see also curves A
and B, fig. 1) point to the same conclusion: that the rock
bass of Trout Lake grew more rapidly than those of Mus¬
kellunge Lake.
A comparison of the average calculated lengths of the
age groups taken from Trout Lake in 1928 (table 4), with
the corresponding age groups taken from Muskellunge
Lake in the same year (table 8) shows that, on the whole,
the rock bass of Muskellunge Lake grew more rapidly
than those of Trout Lake. An examination of the grand
averages in these tables (see also curves C and D, fig. 1)
corroborate this conclusion. Thus it appears that the data
of 1927 and 1928 conflict.
If we compare now, for each lake, the growth data of
corresponding age groups and the grand averages of these
growth data, for fish taken in 1927 with those for fish taken
in 1928, we find that while all age groups taken in the two
years grew at approximately the same rate in Muskellunge
Lake (see also curves B and C, fig. 1), those taken from
Trout Lake in 1927 grew much more rapidly than those se¬
cured in 1928 (curves A and D, fig. 1). The conflict in the
data is thus obviously due to the marked difference in
growth rates of the fish of the two collections from Trout
Lake.
That this difference in the growth rates of the Trout
Lake fish of the two collections is real, and not due to the
inadequacy of the size of samples or of the method of sam-
592 Wisconsin Academy of Sciences, Arts, and Letters .
pling, seems probable from a comparison of the growth
rates of age groups which belong to the same year class.
Thus we may compare the growth rates of V-, VI- and
VHX-year fish of 1927 with those of the VI-, VII- and
VUI-year fish of 1928 (tables 8 and 4) . The average cal¬
culated lengths in millimeters, at the end of each year of
life, of the thirty V-year fish of 1927 are not shown in
table 3 ; they are 23, 43, 66 and 99. The average measured
length of the fish was 114 mm. The data show that in no
instance did the growth rates of any two age groups of a
year class agree; those of age groups taken in 1927 were
consistently much greater than those of age groups taken
in 1928.
The evidence seems to indicate that the two collections
from Trout Lake represent separate populations or races,
each with its own rate of growth, while both collections
from Muskellunge Lake represent the same population.
This conclusion agrees with the fact that in both years the
Muskellunge Lake fish were taken in the same general lo¬
cality (p. 583), whereas the two collections from Trout Lake
were taken in two different localities (Blaisdell’s Bay and
Camp Franklin Shoal), separated by what appears to be a
barrier to migration of the rock bass. For the reasons
given in the introduction (p. 581), it is impossible, with the
available information, to correlate this difference in growth
rates with environmental factors.
The difference in age composition of collections from the
two lakes (tables 1 and 2), as noted on p. 585, is partially
explained on the basis of growth rates and selective action
of gill nets. Most of the fish from the Trout Lake collec¬
tion of 1927 fell in age groups V, VI, VII and VIII, while
those from Muskellunge Lake of both years fell in age
groups VI, VII, VIII and IX. This difference is explained
by the more rapid growth of the 1927 Trout Lake fish; on
the average, they would attain a greater length at any
given age than would those from Muskellunge Lake and
consequently more of the younger Trout Lake fish would be
taken by the gill nets. The bulk of the collection of 1928
from Trout Lake fell in age groups VI, VII and VIII. The
absence of V-year fish is to be expected from the fact that
the fish of this collection grew more slowly than the 1927
Wright — Growth of Rock Bass in Two Lakes of Wisconsin. 593
fish and would thus have to grow for a longer period be¬
fore reaching a size great enough to be selected by the gill
nets. However, it may be due to the absence of V-year
fish from the lake as suggested on p. 586. No explanation
can be offered for the failure of the nets to take more IX-
and X-year fish in 1928.
V. Summary
1. This paper is a preliminary report on the growth of
the rock bass in Trout and Muskellunge Lakes,
Vilas County, Wisconsin.
2. The growth curve of the rock bass typically has the
form of a double sigmoid; that is, there are two
periods of rapid growth, each followed by a decline.
3. The rate of growth in the second period of rapid
growth is greater than that in the first. The first
period usually occurs in the first year and the sec¬
ond in the fifth year. More individuals grow most
rapidly in the fifth year of life than in any other;
the second largest number grow most rapidly in
the sixth year, and the third largest number in the
fourth year.
4. Rock bass of collections from Muskellunge Lake made
in 1927 and 1928 grew at approximately the same
rate. The collections were made in the same gen¬
eral locality and are regarded as representing a
single population.
5. Rock bass taken from Trout Lake in 1927 grew much
more rapidly than those taken in 1928. The two
collections were made in different localities and are
regarded as representing two distinct populations,
each with a characteristic rate of growth.
6. Trout Lake rock bass taken in 1927 grew more rap¬
idly than those taken from Muskellunge Lake in the
same year. Muskellunge Lake rock bass captured
in 1928 grew more rapidly than those taken from
Trout Lake in the same year.
38
594 Wisconsin Academy of Sciences, Arts, and Letters.
Literature
Birge E. A. and C. Juday. 1911. The Inland Lakes of
Wisconsin : The dissolved gases of the water and
their biological significance. Wis. Geol. and Nat. Hist.
Sur. Bull. 22: i-xx and 1-259. Madison, 1911.
Creaser, C. W. 1926. The structure and growth of the
scales of fishes in relation to the interpretation of their
life-history, with special reference to the sunfish, Eupo-
motis gibbosus. Univ. Mich. Mus. Zool., Misc. Publ.,
17 : 1-82. Ann Arbor, 1926.
Juday, C. 1914. The Inland Lakes of Wisconsin: The
hydrography and morphometry of the lakes. Wis.
Geol. and Nat. Hist. Sur. Bull. 27 : i-xv and 1-137.
Madison, 1914.
Thompson, W. F. 1916. A contribution to the life-history
of the Pacific herring: Its bearing on the condition
and future of the fishery. Rept. Comm. Fish., British
Columbia. 1916: 39-87.
Van Oosten, John. 1923. The whitefishes (Coregomts clu-
peaformis) : A study of the scales of whitefishes of
known ages. Zoologica 2 (17) : 381-412. 1923.
— — 1928. The life-history of the lake herring (1) of Lake
Huron as revealed by its scales, with a critique of the
scale method. Bull. U. S. Bur. Fish. 44:265-428.
1928 (Issued 1929) Doc. No. 1053.
Wright — Growth of Bock Bass in Two Lakes of Wisconsin. 595
n
AGE IN YEARS
Fig. 1. Trout and Muskellunge lakes, 1927 and 1928. Average calculated length, at the end of each year of life, of rock
ss of age groups VI, VII and VIII: A, Trout Lake 1927: B, Muskellunge Lake 1927; C, Muskellunge Lake 1928; D, Trout
ike 1928. E, annual growth increments taken from Curve A.
PROCEEDINGS OF THE ACADEMY
Fifty-seventh Annual Meeting, 1927
The fifty-seventh annual meeting of the Wisconsin Aca¬
demy of Sciences, Arts and Letters, in joint session with
the Wisconsin Archeological Society and the Midwest Mu¬
seums Conference, was held at the University of Wisconsin,
Madison, on Thursday, Friday and Saturday, April 7, 8 and
9, 1927.
The following program was presented:
THURSDAY, APRIL 7
Afternoon Session, 2:00 O’clock
Auditorium of State Historical Museum
Midwest Museums Conference.
1. Model Museum at the State Fair.
a. What the Historical Department should contain. Charles
E. Brown.
b. What the Natural History Department should contain.
Owen Grumme, Huron H. Smith, Ira Edwards and
T. E. B. Pope.
c. Methods of cataloguing, Nile Behncke.
2. A course in museum work for midwest members. S. A. Barrett.
3. Museum news exchange.
4. The museum specializing in some particular field. R. N. Buck-
staff.
5. A report on the recently formed Art Classes at the Oshkosh
Public Museum. Nile Behncke.
Business Meeting.
FRIDAY, APRIL 8
Morning Session, 9:00 O’clock
Auditorium of Biology Building.
General Meeting.
6. History of Costume. Hazel Manning. Illustrated.
7. The natural history department of a small museum. R. N. Buck-
staff.
598 Wisconsin Academy of Sciences, Arts, and Letters.
8. Life and letters of Edward Lee Green. Angie K. Main.
9. The scope of botany. Huron H. Smith. Illustrated.
10. A forty year old hobby. S. C. Wadmond.
11. Catalogue of the species of Hepaticae found in Wisconsin. George
H. Conklin. By Title.
12. Notes on parasitic fungi in Wisconsin. XV. J. J. Davis. By
Title.
13. The chemical composition of Chara from Green Lake, Wisconsin.
H. A. SCHUETTE. By Title.
14. Structure and behavior of century-old cells. J. B. Overton.
Afternoon Session, 2:00 O’clock
15. The measurement of educational processes and products. M. V.
O’Shea.
16. The intellectual resemblance of twins. Curtis Merriman.
17. The life and customs of the Tarahumari Indians in the Sierra
Madre Mountains. R. M. Bagg. Illustrated.
18. Colonial debtors and English creditors. Curtis Nettels.
19. Recent photographs of Mars. Frank E. Ross. Illustrated.
SATURDAY, APRIL 9
Morning Session, 9:00 O’clock
20. Commemorative postage stamps of the United States. F. M. K.
Foster. Illustrated.
21. Fort Winnebago. W. C. English.
22. The relation between Erasmus and Luther as shown in their cor¬
respondence. Ernst Voss.
23. Organic content of the waters of small lakes. E. A. Birge.
24. Studies of the development and prevention of apple scab epi¬
demics. G. W. Keitt.
25. Reconstruction in Japan. L. S. Smith. Illustrated.
Report of Secretary, April 1, 1926 to March 31, 1927.
Membership
Honorary Members _ 6
Life Members - 15
Corresponding Members - 20
Active Members _ 316
Total _ _ _ _ _ _ 357
Resigned - - - — - - - - 1
Dropped for nonpayment of dues - - - , - 2
Deaths _ 2
Total
5
Proceedings of Annual Meetings.
599
The following members died during the year:
Frederick Belding Power, March 26 _ 1927
Addison Emery Verrill _ 1926
John Barber Parkinson, April 3 - 1927
The Secretary presented the following applications for membership.
On motion he was unanimously instructed to cast the ballot in their
favor :
H. R. Aldrich, Madison
Anselm M. Keefe, West Depere
Curtis Putnam Nettels, Madison
Frank E. Ross, Williams Bay
Otto Struve, Williams Bay
George Van Biesbroeck, Williams Bay
Ruth I. Walker, Madison
Stillman Wright, Madison
J. Sidney Hooton, Williams Bay
Officers elected April 9, 1927 for three years:
President, Samuel A. Barrett, Milwaukee
Vice-President, Sciences, Storrs B. Barrett, Williams Bay
Vice-President, Arts, Arnold Dresden, Madison
Vice-President, Letters, E. K. J. H. Voss, Madison
Secretary-Treasurer, Chancey Juday, Madison
Curator, C. E. Brown, Madison
Librarian, Walter M. Smith, Madison
The annual dinner was held at the University Club on
Friday evening, April 8, 1927, attended by 47 members and
guests. Following the dinner Dr. Joseph Schafer delivered
an address entitled “On the Gold Trail, 1849”.
Chancey Juday,
Secretary .
Report of Treasurer from April 1, 1926 to March 31, 1927.
Receipts
Balance in State Treasury April 1, 1926 _ $4,353.17
Receipts from dues _ 228.10
Received from sales of Transactions _ 7.85
Received for reprints _ 45.97
Received from Mrs. E. J. B. Schubring _ _ _ _ _ 50.00
Interest _ 1.00
Annual appropriation, July 1, 1926 _ 1,500.00
$6,186.09
600 Wisconsin Academy of Sciences , Arts, and Letters .
Disbursements
Drafting for Mrs. Schubring’s paper _ $22.00
Telephone message _ _ .40
Envelopes (University) _ 1.00
Postage _ 61.50
Annual allowance of Secretary _ 200.00
Printing _ 2,202.13
$2,487.03
Balance in State Treasury April 1, 1927 _ $3,699.06
Securities and cash on hand April 1, 1927
City of Madison bonds _ $2,900.00
Mortgage bonds on Chapman Block, Madison _ 400.00
Certificate of deposit _ 34.72
Cash _ 8.80
$3,343.52
Chancey Juday,
Treasurer.
Audited and found correct.
George Wagner,
R. N. Buckstaff,
Auditors.
FIFTY-EIGHTH ANNUAL MEETING, 1928
The fifty-eighth annual meeting of the Wisconsin
Academy of Sciences, Arts and Letters, in joint session with
the Wisconsin Archeological Society and the Midwest Mu¬
seums Conference, was held at Lawrence College, Appleton,
on Friday and Saturday, April 6 and 7, 1928.
The following program was presented :
FRIDAY, APRIL 6
Morning Session, 9:30 O’clock
General Business.
Presentation of Papers.
1. Wisconsin’s oldest families. N. C. Fassett. Illustrated.
2. A call for amateur botanists. N. C. Fassett.
3. Notes on the flora of Lake Wingra. Jeanett Kendall.
4. Forestry in Wisconsin. Huron H. Smith. Illustrated.
Proceedings of Annual Meetings.
601
5. Flora of Wisconsin sand dunes. A. M. Fuller. Illustrated.
6. Botanical groups. Huron H. Smith. Illustrated.
7. Visual methods and devices for museums and schools. John B.
MacHarg.
8. Discussion of Lawrence Vale Coleman’s “Manual for the small
museum.” Leaders of the discussion, R. N. Buckstaff, and
S. A. Barrett.
9. Notes on the chemical composition of some of the larger aquatic
plants of Lake Mendota. III. Castalia and Najas. Henry
A. Schuette and Hugo Alder. By title.
10. A note on the chemical composition of Chara from Green Lake,
Wisconsin. Henry A. Schuette and Hugo Alder. By title.
Afternoon Session, 1:30 O’clock
Presentation of Papers.
11. Eskimo picture writing. George A. West. Illustrated.
12. Wisconsin Indians after the British conquest, 1761-1775. Louise
P. Kellogg. By title.
13. Family names of civilized Indians. Vetal Winn.
14. The pottery repository at Ann Arbor. Charles E. Brown.
15. Sacred springs of the Poygan region. George Overton.
16. The Reedsburg cache of flint implements. Milton K. Hulbert.
17. Two fluted stone implements. Vetal Winn.
18. Indian trade clasp knives from Lake Koshgonong. Theodore T.
Brown.
19. Balance and psychology of museum work. George R. Fox.
20. Airplane photograph of a Kenosha County efligy mound. C. W.
Beemer.
21. Indian fishing camps of the Wolf River. Francis S. Dayton.
By title.
22. An early Chicago immigration society. Albert O. Barton.
23. Museum classification and storage methods. T. E. B. Pope. Illu¬
strated.
24. Trail-side and outdoor museums. A discussion led by S. A. Bar¬
rett and R. N. Buckstaff.
25. Hawaii, the Paradise of the Pacific. S. A. Barrett. Illustrated
with moving pictures.
SATURDAY, APRIL 7
Morning Session, 9:00 O’clock
General Business.
Presentation of Papers.
26. The sexes of hybrid pigeons and doves. L. J. Cole.
27. The University bird banding station. George Wagner.
28. The transmission of solar radiation by the waters of Wisconsin
lakes. E. A. Birge.
39
602 Wisconsin Academy of Sciences, Arts, and Letters.
29. Mineral content of the waters of the Great Lakes. L. A. Youtz.
30. Redetermination of the atomic weight of arsenic. L. A. Youtz.
31. Recent studies of the Wisconsin Cambrian. Ira Edwards.
32. Merestomes of the Wisconsin Cambrian. Gilbert 0. Raasch.
Illustrated.
33. The origin of Crater Lake. Ira Edwards. Illustrated.
34. The Devonian section of Little Traverse Bay, Michigan. Gilbert
0. Raasch.
35. The red salmon of Kariuk. George I. Kem merer. Illustrated
with moving pictures.
36. William Langland’s “Piers the Plowman” and Johannes von
Saaz’s “Der Ackerman aus Bohmen.” Ernst Voss. By title.
Report of Secretary, April 1, 1927 to March 31, 1928.
Membership
Honorary Members _ 6
Life Members _ 15
Corresponding Members _ 20
Active Members _ 288
Total _ 329
New Members not included above _ 5
Grand total _ 334
Membership losses during year
Deceased _ 4
Resigned _ 1
Dropped for nonpayment of dues _ 28
Total _ 33
The following Members died during the year:
Frank Gaylord Hubbard, March 15, 1928
Victor Lenher, June 12, 1927
Maurice McKenna, August 23, 1927
Harry E. Cole, April 13, 1928.
The Secretary presented the following* applications for
membership. On motion he was unanimously instructed
to cast the ballot in their favor.
Demetrio di Demetrio, Trieste, Italy
Marcel Elias, Liege, Belgium
Paul S. Henshaw, Madison
Proceedings of Annual Meetings.
603
Stephen John Martin, Madison
Samuel B. Sklar, Minneapolis, Minnesota
Vladimir N. Rimsky-Korsakoff, Madison
Willis L. Tressler, Madison
Charles H. Skinner, Marquette University, Milwaukee
Charles D. Higgs, Fontana
E. E. Honey, Madison.
The Annual dinner was held at Brokaw Hall on Friday
evening, April 6, and was attended by 45 individuals. Af¬
ter the dinner President Wriston delivered an address on
the subject “Opportunities for scholarly work in the De¬
partment of State”.
The following motion was carried unanimously:
That we extend a hearty vote of thinks to Lawrence Col¬
lege for its kindly hospitality, and to Professor Bagg for
the complete and efficient manner in which he looked after
the local arrangements for our meeting.
Chancey Juday,
Secretary .
Report of Treasurer, April 1, 1927 to March 31, 1928.
Receipts
Balance in State Treasury April 1, 1927 _ _ $3,699.06
Receipts from dues _ _ _ _ _ _ 243.00
Received from sales of Transactions _ _ _ 26.62
Received for reprints _ 32.31
Annual appropriation, July 1, 1927 — _ . _ 1,500.00
Total _ $5,500.99
Disbursements
Annual allowance of Secretary _ _ $200.00
Postage _ . _ _ 109.00
Printing — - - 2,902.74
Shipping labels - 12.35
Total- - - - $3,224.09
Balance in State Treasury April 1, 1928 - $2,276.90
604 Wisconsin Academy of Sciences , Arts, and Letters.
Securities and cash on hand April 1, 1928
City of Madison bonds _ _ _ _ _ $2,000.00
Mortgage bonds on Chapman block, Madison _ 400.00
Trust agreement (Cent. Wis. Trust Co.) _ 1,000.00
Certificate of deposit _ _ _ 58.50
Cash _ 40.47
Total _ $3,458.97
Chancey Juday,
Treasurer.
Audited and found correct.
George Wagner,
Huron H. Smith,
Auditors.
FIFTY-NINTH ANNUAL MEETING, 1929
The fifty-ninth annual meeting of the Wisconsin Academy
of Sciences, Arts and Letters, in joint session with the Wis¬
consin Archeological Society and the Midwest Museums
Conference, was held at Yerkes Observatory, Williams Bay,
on Friday and Saturday, April 12 and 13, 1929.
The following program was presented :
FRIDAY, APRIL 12
Morning Session, 10:00 O’clock
General Business
Presentation of Papers
1. The Potawatomi Indians of Lake Geneva. Paul B. Jenkins.
Illustrated.
2. Wisconsin Indians during the American Revolution. Louise P.
Kellogg.
3. An Abraham Lincoln Indian Medal. Theodore T. Brown.
4. An ancient village site in Winnebago County. George Overton.
5. Native plants used by Quileute Indians of Washington. Albert
B. Reagan.
6. The Indian copper mines of Isle Royal. George A. West. Illu¬
strated.
7. Swamp dwellers. George R. Fox.
Proceedings of Annual Meetings.
605
Afternoon Session, 1:30 O’clock
Presentation of Papers
8. Large telescopes. Frank E. Ross. Illustrated.
9. Founding of the Republican party. S. M. Pedrick.
10. Some general observations on immigration. J. Schafer. Twen¬
ty minutes.
11. Excavation of Paleolithic deposits of Algeria. George L. Collie.
Illustrated.
12. The Lions of Cochiti. John B. McHarg. Illustrated.
13. Diary of an immigrant gold seeker in California. Albert 0.
Barton.
14. Pioneer mills of Rock County. May L. Bauchle.
15. Indian village sites of the Lower Rock River in Wisconsin. C. E.
Brown.
16. The conservation of our Wisconsin lakes and streams. 0. W.
Smith.
17. Problems encountered in pathological investigations of prehis¬
toric bone material. Herbert W. Kuhm, George Adami and
Alton K. Fisher. Illustrated.
18. Phases of museum extension work. The Ohio Valley. Roy S.
Corwin. Illustrated.
19. Fostering zoological collecting among younger people. W. E.
Dickinson.
20. Wisconsin herpetological notes. T. E. B. Pope. Illustrated.
21. New restoration of Stegosaurus and Allosaurus. Ira Edwards.
Illustrated.
SATURDAY, APRIL 13
Morning Session, 9:00 O’Clock
General Business
Presentation of Papers
22. Extremes of heat and cold in Wisconsin. Eric R. Miller. Illu¬
strated.
23. New facts regarding the Wisconsin Cambrian. Ira E wards.
24. New facts regarding the Wisconsin Devonian. Gilbert O.
Raasch.
25. A remarkable abnormality in cats. George Wagner.
26. The status of the European starling in Wisconsin. George Wag¬
ner.
27. Color inheritance in pigeon hybrids. Leon J. Cole. Illustrated.
28. Resting-spores of Entomophthora species found on Aphis spirae-
cola. E. M. Gilbert.
29. Observations on Protoachyla. James A. Lounsbury. Illustrated.
30. Delayed germination of Nelumbo lutea. J. A. Jones.
606 Wisconsin Academy of Sciences , Arts, and Letters.
31. Occurrence of ferns in Wisconsin. W. N. Stiles. Illustrated.
32. Some ecological studies of the Missouri River bluffs. David F.
Costello. Illustrated.
33. Newer biochemical aspects of cold resistance in wheat. W. E.
Tottingham.
34. Botanizing Wisconsin’s lost lake. N. C. Fassett. Illustrated.
35. Phases of museum extension work. The Ozark region of Missouri
and Arkansas. W. D. Kline. Illustrated.
36. Plant research among the Winnebago Indians. Huron H. Smith.
Illustrated.
37. Notes on parasitic fungi of Wisconsin. XVI and XVII. J. J.
Davis. By title.
38. Wordsworth, science, and poetry. Arthur Beatty.
39. Plan for a small astronomical museum. R. N. Buck staff. Illu¬
strated.
Report of Secretary, April 1, 1928 to March 31, 1929.
Membership
Honorary Members _ 5
Life Members - 14
Corresponding Members _ 17
Active Members _ _ _ 345
Total _ 381
Membership losses during year
Deceased _ 5
Resigned _ 3
Dropped for nonpayment of dues _ 3
Total _ 11
The following Members died during the year:
Thomas Chrowder Chamberlin, November 15, 1928
John Merle Coulter, December 23, 1928
Linnaeus Wayland Dowling, September 16, 1928
George I. Kemmerer, November 18, 1928
Benjamin W. Snow, September 21, 1928
William H. Wright, May 3, 1929
The Secretary presented the following applications for member¬
ship.
On motion he was unanimously instructed to cast the ballot in their
favor.
Walter M. Banfield, Madison
H. W. Beams, Madison
Ethelwynn R. Beckwith, Milwaukee
George C. Blakeslee, Williams Bay
Proceedings of Annual Meetings.
607
Lester M. Blank, Madison
Newton Bobb, Ashland
H. C. Bradley, Madison
H. N. Calderwood, Madison
Ralph M. Caldwell, Madison
Mary R. Calvert, Williams Bay
Thomas C. Carter, Madison
John O’Neill Gloss, Madison
Grace L. Clapp, Milwaukee
William Wheeler Coleman, South Milwaukee
Delmer C. Cooper, Madison
David F. Costello, Milwaukee
Hobart W. Cromwell, Madison
L. A. V. DeCleene, West Depere
John E. Dudley, Jr., Madison
B. M. Duggar, Madison
C. T. Elvey, Williams Bay
Henry Ericson, Milwaukee
R. I. Evans, Madison
J. R. Fanselow, Appleton
Edward A. Fitzpatrick, Milwaukee
J. B. Goldsmith, Madison
L. F. Graber, Madison
William H. Gunther, Sheboygan
J. M. Hamilton, Madison
J. M. Hansell, Madison
Edmund Heller, Milwaukee
C. A. Herrick, Madison
George Hilton, Oshkosh
Wm. Horlick, Jr., Racine
Mark H. Ingraham, Madison
Charles A. Jahr, Elkhorn
James Albert Jones, Fond du Lac
H. H. Ketcham, Madison
Ruth H. Lindsay, Nashotah
Elizabeth F. McCoy, Madison
S. M. McElvain, Madison
J. D. Mac Lean, Madison
Carol Y. Mason, Milwaukee
V. W. Meloche, Madison
Albertine E. Metzner, Plymouth
Edward L. Miloslavich, Milwaukee
George W. Moffitt, Williams Bay
Clarence W. Muehlberger, Madison
Beatrice I. Nevins, Madison
Helen T. Parsons, Madison
Ernest A. Petzke, Hixton
A. H. Poetker, Milwaukee
608 Wisconsin Academy of Sciences , Arts, and Letters .
Louise Pollitz, Oshkosh
Maximilian Rakette, Frankfurt, Germany
Allan F. Reith, Milwaukee
W. D. Richtmann, Madison
Caryl Ripley, Williams Bay
Walter E. Rogers, Appleton
H. W. Rohde, Milwaukee
Fred. George Russell, Milwaukee
Elbert B. Ruth, Madison
David A. Schaefer, Lake Geneva
Erwin R. Schmidt, Madison
B. L. Schuster, Waupun
Joel Stebbins, Madison
Walter E. Sullivan, Madison
James H. Taylor, Madison
Robert Triebel, Beuthen, Germany
K. H. Tutunjian, Ripon
J. H. Van Vleck, Madison
Ralph M. Walters, Madison
Frederick L. Wellmann, Tela, Honduras
A. H. Wiebe, Fairport, Iowa
Edgar J. Witzeman, Madison
The annual dinner was held on Friday evening, April 12,
at the Rose Lane Resort; it was attended by 90 members
and guests. Following the dinner Professor Edwin B.
Frost gave a very interesting address on “The Architecture
of the Heavens’'. After the address, members and guests
were given an opportunity to look through the 12 inch and
the 40 inch telescopes.
At the business meeting on Saturday the following mo¬
tion was unanimously adopted :
We extend to the Director and the Staff of the Yerkes
Observatory our thanks and our deep appreciation of their
kindly hospitality and thoughtful solicitude in making this
meeting of the Academy so enjoyable a success.
Chancey Juday,
Secretary.
Proceedings of Annual Meetings.
609
Report of Treasurer, April 1, 1928 to March 31, 1929.
Receipts
Balance in State Treasury April 1, 1928 _ $2,276.90
Receipts from dues _ _ _ 312.00
Received for publication of paper (Prof. Owen) _ 711.41
Received from sales of Transactions _ 107.18
Received from sales of reprints _ 15.65
Annual appropriation July 1, 1928 _ 1,500.00
Total _ $4,923.14
Disbursements
Advertising, circular letter and cards _ _ _ $ 10.00
Postage _ 73.50
Salary of Secretary _ 200.00
Express _ 2.16
Printing _ 2,613.73
Total disbursements _ $2,899.39
Balance in State Treasury April 1, 1929 _ $2,023.75
Securities and cash on hand April 1 , 1929
City of Madison bonds _ _ _ $2,000.00
Mortgage bonds on Chapman Block, Madison _ _ _ 400.00
Trust agreement (Cent. Wis. Trust Co.) _ 1,000.00
Bonds of Commonwealth Telephone Co. _ _ _ 200.00
Cash _ 16.23
$3,616.23
Chancey Juday,
Secretary.
Audited and found correct.
R. N. Buckstaff,
George Wagner,
Auditors.
610 Wisconsin Academy of Sciences, Arts, and Letters .
LIST OF OFFICERS AND MEMBERS
Corrected to May 1, 1929
Officers
President, Samuel A. Barrett, Milwaukee.
Vice-President, Sciences, Storrs B. Barrett, Williams Bay.
Vice-President, Arts, Arnold Dresden, Madison.
Vice-President, Letters , E. K. J. H. Voss, Madison.
Secretary-Treasurer, Chancey Juday, Madison.
Curator , C. E. Brown, Madison.
Librarian, Walter M. Smith, Madison.
Committee on Publication
The President, ex-officio,
The Secretary, ex-officio,
W. E. Tottingham, Madison.
Council
The President, Vice-Presidents, Secretary, Treasurer, Librarian
and Past Presidents retaining their residence in Wisconsin.
Committee on Library
The Librarian, ex-officio,
W. H. Barber, Ripon,
A. E. Culver, Stevens Point,
George Wagner, Madison.
Committee on Membership
The Secretary, ex-officio,
P. W. Boutwell, Beloit,
R. N. Buckstaff, Oshkosh,
Huron H. Smith, Milwaukee.
Past Presidents
Honorable John W. Hoyt,1 1870-1875.
Doctor P. R. Hoy,1 1876-1878.
President A. L. Chapin,1 1879-1881.
Professor Ronald D. Irving,1 1882-1884.
Professor Thomas C. Chamberlin,1 1885-1887.
Professor William F. Allen,1 1888-1889.
1 Deceased.
List of Officers and Members.
611
Professor Edward A. Birge, 1889-1890.
Librarian George W. Peckham,1 1891-1893.
President Charles R. Van Hise,* * 1894-1896.
Professor C. Dwight Marsh, 1897-1899.
Professor Charles S. Slighter, 1900-1902.
Doctor John J. Davis, 1903-1905.
Professor Louis Kahlenberg, 1906-1909.
President Samuel Plantz,1 1910-1912.
Professor Dana C. Munro, 1913-1915.
Director Henry L. Ward, 1916-1918.
President Edward A. Birge, 1919-1921.
President Melvin A. Brannon, 1922-1924.
Professor Leon J. Cole, 1925-1927.
HONORARY MEMBERS
Garland, Hamlin, New York, N. Y.
Jordan, David Starr, Stanford University, Calif.
Merrick, George B., 350 W. Washington Ave., Madison.
Trelease, William, University of Illinois, Urbana, Ill.
Wheeler, William Morton, Forest Hills, Boston, Mass.
LIFE MEMBERS
Beatty, Arthur, 1824 Vilas Street, Madison.
*Birge, Edward A., 2011 Van Hise Ave., Madison
♦Davis, John J., 803 State Street, Madison.
Hixon, Frank P., La Crosse
Hobbs, William H., Ann Arbor, Michigan
Horlick, A. J., Racine
* Leith, Charles K., Moraine, Old Sauk Road, Madison
Logan, Frank A., Chicago, Illinois
♦Marsh, C. Dwight, 1882 Monroe St., Washington, D. C.
Norris, Mrs. Fanny, 1906 Grand Ave., Milwaukee
Peckham, Mrs. Elizabeth G., 46 Marshall St., Milwaukee
Sharp, Frank C., University of Wisconsin, Madison
♦Skinner, Ernest Brown, 210 Lathrop St., Madison
♦Slichter, Charles Sumner, 636 North Frances St., Madison
CORRESPONDING MEMBERS
Crooker, Joseph Henry, Orlando, Florida
Eckels, William Alexander, La Fayette College, Easton, Pa.
Hendrickson, Geo. L., Yale University, New Haven, Conn.
Hoskins, Leander Miller, Palo Alto, California
1 Deceased.
* Member of American Association for Advancement of Science.
612 Wisconsin Academy of Sciences , Arts , and Letters .
Kinley, David, University of Illinois, Urbana, Illinois
Leverett, Frank, University of Michigan, Ann Arbor, Mich.
Libby, Orin Grant, Grand Forks, North Dakota
Marx, Charles David, Palo Alto, California
McClumpha, Charles Flint, 56 Church St., Amsterdam, N. Y.
Munro, Dana C., Princeton University, Princeton, N. J.
Olive, Edgar W., Brooklyn Botanic Garden, Brooklyn, N. Y.
Stone, Ormond, Clifton Station, Virginia
Tolman, Albert H., University of Chicago, Chicago, Ill.
Townley, Sidney Dean, Stanford University, California
Turner, Frederick Jackson, 2214 Van Hise Ave., Madison
ACTIVE MEMBERS
♦Addleman, Irving M., 511 Third Street, Wausau
♦Addoms, Ruth M., Biology Building, U. W., Madison
Aitken, Robert, 1058 Fourteenth Ave., Honolulu, Hawaii
Alderman, William E., Beloit College, Beloit
♦Aldrich, H. R., 1705 Jefferson Street, Madison
♦Allen, Charles E., 2014 Chamberlin Ave., Madison
♦Arzberger, Emil G., Bureau of Plant Industry, Washington, D. C.
♦Bachmann, Freda M., Stout Institute, Menomonie
♦Bagg, Rufus M., 16 Brokaw Place, Appleton
♦Baird, Edgar A., University, Grand Forks, N. D.
♦Banfield, Walter M., 1220 Mound St., Madison
Barber, Wm. H., Ripon College, Ripon
♦Bardeen, Charles R., University of Wisconsin, Madison
Barrett, Samuel A., Public Museum, Milwaukee
♦Barrett, Storrs B., Yerkes Observatory, Williams Bay
Barth, George P., 3006 Juneau Ave., Milwaukee
Bascom, Lelia, 419 Sterling Place, Madison
♦Beams, H. W., Biology Bldg., U. W., Madison
♦Bean, E. F., Science Hall, U. W., Madison
♦Bean, Elizabeth Smith, Dalhousie University, Halifax, N. S.
♦Beckwith, Ethelwynn R., Milwaukee-Downer College, Milwaukee
Behncke, Nile J., 785 Algoma Blvd., Oshkosh
♦Bennett, Edward, 1919 Jefferson Street, Madison
♦Blakslee, George C., Yerkes Observatory, Williams Bay
♦Blank, Lester M., 1717 Summit Ave., Madison
♦Bobb, Newton, 1008 Second Ave., West, Ashland
♦Boutwell, Paul W., Beloit College, Beloit
♦Bradley, Harold C., Memorial Institute, U. W., Madison
Braun, Adolph R., 832 38th Street, Milwaukee
♦Brink, R. Alex., University of Wisconsin, Madison
Brown, Charles E., 2011 Chadbourne Ave., Madison
♦Browne, Frederick L., 119 Ash Street, Madison
* Member of American Association for Advancement of Science.
List of Officers and Members.
613
♦Browning, Harold W., State College, Kingston, R. I.
♦Bryan, George S., Biology Building, U. W., Madison
♦Buckstaff, Ralph N., 1122 South Main Street, Oshkosh
Buehler, Henry A., Bureau of Geology, Rolla, Missouri
♦Bunting, Charles H., Memorial Institute, U. W., Madison
Bussewitz, M. A., 435 Kenwood Boulevard, Milwaukee
Butterfield, Ellen F., 713 Milwaukee Ave., South Milwaukee
Cahn, A. R., University of Illinois, Urbana, Ill.
Cairns, Wm. B., University of Wisconsin, Madison
♦Calderwood, H. N., Chemistry Building, U. W., Madison
♦Caldwell, Ralph M., 803 State Street, Madison
♦Calvert, Mary R., Yerkes Observatory, Williams Bay
Carter, Sylvester J., Public Library, Milwaukee
♦Carter, Thomas C., 931 West Johnson St., Madison
♦Chandler, Elwyn F., University, Grand Forks, N. D.
Chase, Wayland J., University of Wisconsin, Madison
♦Churchill, B. P., 679 Sixth Ave., Milwaukee
♦Clapp, Grace L., Milwaukee-Downer College, Milwaukee
♦Clark, Paul F., Memorial Institute, U. W., Madison
Clas, Alfred C., 445 Milwaukee Street, Milwaukee
Clawson, Arthur B., 1884 Monroe St., N. W., Washington, D. C.
♦Closs, John O’Neill, Chemistry Bldg., U. W., Madison
Clowes, Herbert, Public Museum, Milwaukee
♦Cole, Leon J., University of Wisconsin, Madison
♦Coleman, William W., 575 Marshall Street, Milwaukee
♦Collie, George L., Beloit College, Beloit
Commons, John R., University of Wisconsin, Madison
♦Comstock, George C., 635 College Street, Beloit
Conklin, George H., 2011 John Avenue, Superior
♦Conwell, H. H., Beloit College, Beloit
♦Cooper, Delmer C., Biology Bldg., U. W., Madison
♦Cooper, George Olds, Biology Bldg., U. W., Madison
Costello, David F., Marquette University, Milwaukee
♦Cromwell, Hobart W., Memorial Institute, U. W., Madison
♦Culver, Garry E., State Teachers College, Stevens Point
♦Daniels, Farrington, Chemistry Building, U. W., Madison
♦Davis, Carl Henry, 141 Wisconsin Street, Milwaukee
♦Dawson, Percy M., Memorial Institute, U. W., Madison
Dean, Alletta F., 87 West Street, Mansfield, Mass.
♦De Cleene, Rev. L. A. V., St. Norbert College, West De Pere
♦Delwiche, Edmond J., R. F. D. 3, Green Bay
Demetrio di Demetrio, Via Malcanton II/II, Trieste, Italy
♦Denniston, Rollin H., Biology Building, U. W., Madison
♦Densmore, H. D., Beloit College, Beloit
Dodge, B. O., Botanical Garden, New York, N. Y.
Dodge, R. E. N., Bascom Hall, U. W., Madison
♦Domogalla, Bernhard P., 803 State Street, Madison
* Member of American Association for Advancement of Science.
614 Wisconsin Academy of Sciences , Arts , and Letters .
♦Dove, William F., University of Maine, Orono, Maine
Downes, Robert H., 574 Algoma Boulevard, Oshkosh
♦Drechsler, Charles, Bureau of Plant Industry, Washington, D. C.
♦Dudley, John E., Jr., 1532 University Avenue, Madison
♦Duggar, B. M., Biology Bldg., U. W., Madison
Du Mez, Andrew G., University of Maryland, Baltimore, Md.
Edwards, Ira, Public Museum, Milwaukee
Elias, Marcel, Boulevard de la Constitution 25, Liege, Belgium
Ely, Richard T., Northwestern University, Chicago, Ill.
♦Elvey, C. T., Yerkes Observatory, Williams Bay
♦Ericson, Henry, 3414 Juneau Avenue, Milwaukee
♦Evans, Clarence T., 367 Sixth Avenue, Wauwatosa
♦Evans, Edward, 121 South 13th Street, La Crosse
Evans, R. I., Biology Bldg., U. W., Madison
♦Fajardo, T. G., State College, Corvallis, Oregon
♦Fanselow, J. R., '921 East Alton Street, Appleton
♦Farley, John H., Lawrence College, Appleton
♦Fassett, Norman C., Biology Bldg., U. W., Madison
♦Fiebiger, Geo. J., Waterloo
Finch, V. C., Science Hall, U. W., Madison
Finkler, Adolph, 612 Commerce Street, Milwaukee
♦Fischer, Richard, Chemistry Bldg., U. W., Madison
♦Fisk, Emma L., Biology Bldg., U. W., Madison
♦Fitzpatrick, Edward A., Marquette University, Milwaukee
Fluke, Charles L., University of Wisconsin, Madison
♦Fracker, Stanley B., 3716 Ingomar St., N. W., Washington, D. C.
♦Fred, Edwin B., Agricultural Hall, U. W., Madison
Frey, Charles N., 103 West 183rd St., New York, N. Y.
Frizzi, Oscar, Via Gatteri, 9/1, Trieste, Italy
Fuller, Albert M., Public Museum, Milwaukee
Gay, Lucy M., Bascom Hall, U. W., Madison
♦Gerry, Eloise, Forest Products Laboratory, Madison
♦Gilbert, Edward M., Biology Bldg., U. W., Madison
Glicksman, Harry, South Hall, U. W., Madison
Gloyer, Walter 0., Agric. Experiment Station, Geneva, N. Y.
♦Graber, L. F., Agronomy Bldg., U. W., Madison
♦Granovsky, Alexander A., University of Wisconsin, Madison
Greene, Howard, 559 Marshall Street, Milwaukee
♦Groves, James F., Ripon College, Ripon
♦Gunther, William H., Sheboygan Clinic, Sheboygan
Gutsch, Milton R., University of Texas, Austin, Texas
♦Guyer, M. F., Biology Building, U. W., Madison
Haase, Ewald, 182 East Wisconsin Ave., Milwaukee
♦Hamilton, J. M., Horticultural Bldg., U. W., Madison
♦Hampton, Harold C., Hawkesbury, Ontario, Canada
Hanley, Miles L., Bascom Hall, U. W., Madison
♦Hansell, J. M., Science Hall, U. W., Madison
* Member of American Association for Advancement of Science.
List of Officer's and Members.
615
Harper, Robert A., Columbia University, New York, N. Y.
Harper, Mrs. Robert A., R. R. No. 1, Ridgewood, N. J.
♦Hastings, E. G., Agricultural Hall, U. W., Madison
*Heineman, Paul G., 5707 Kenwood Ave., Chicago, Ill.
* Heller, Edmund, 4600 Vliet Street, Milwaukee
*Henmon, V. A. C., Bascom Hall, U. W., Madison
♦Henshaw, Paul S., Biology Building, U. W., Madison
♦Herrick, C. A., Biology Bldg., U. W., Madison
♦Hiestand, William A., Biology Bldg., U. W., Madison
♦Higgs, Charles D., Fontana
♦Hilton, George, 375 Washington Blvd., Oshkosh
♦Hisaw, F. L., Biology Bldg., U. W., Madison
♦Hodgson, A. J., 1008 E. Broadway, Waukesha
Hohlfeld, A. R., Bascom Hall, U. W., Madison
Holmes, Samuel J., University of California, Berkeley, Calif.
Honey, E. E., 314 Burr Oak St., Albion, Mich.
Hotchkiss, W. O., Michigan College of Mines, Houghton, Mich.
Hotton, J. Sidney, Williams Bay
♦Horlick, William, Jr., Racine
♦Humphrey, Clarence J., Bureau of Science, Manila, P. I.
♦Ingersoll, Leonard R., Sterling Hall, U. W., Madison
♦Ingraham, Mark Hoyt, North Hall, U. W., Madison
* Jackson, H. H. T., Biological Survey, Washington, D. C.
Jahr, Charles A., Elkhorn
* Jansky, Cyril M., University of Wisconsin, Madison
* Jasper, Thomas McL., A. O. Smith Corp., Milwaukee
♦Jastrow, Joseph, 143 E. 49th St., New York, N. Y.
♦Johnson, Aaron G., Dept. Agric., Washington, D. C.
♦Johnson, James, Horticultural Bldg., U. W., Madison
♦Jones, Fred R., Horticultural Bldg., U. W., Madison
Jones, James A., High School, Fond du Lac
♦Jones, Leon K., Agric. Exp. Sta., Geneva, N. Y.
♦Jones, Lewis R., Horticultural Bldg., U. W., Madison
♦Jones, Sarah Van H., Rochester, Mich.
♦Juday, Chancey, Biology Bldg., U. W., Madison
♦Kahlenberg, Louis, Chemistry Bldg., U. W., Madison
Keefe, Anselm M., St. Norbert College, West De Pere
♦Keitt, George W., Horticultural Bldg., U. W., Madison
♦Kenyon, W. A., Hamline Univ., St. Paul, Minn.
♦Ketcham, H. H., 2011 Chadbourne Ave., Madison
Kiekhofer, W. H., Sterling Hall, U. W., Madison
♦Koehler, Arthur, Forest Products Lab., Madison
♦Kowalke, O. L., Chemical Engr. Bldg., U. W., Madison
Kraus, Ezra J., University of Chicago, Chicago, Ill.
♦Kremers, Edward, Chemistry Bldg., U. W., Madison
♦Kutchin, Harriet L., Green Lake
♦Lee, Oliver J., Dearborn Observatory, Evanston, Ill.
Member of American Association for Advancement of Science.
616 Wisconsin Academy of Sciences , Arts , and Letters.
* Leonard, Clifford S., Yale University, New Haven, Conn.
Lewis, F. F., 115 Jackman St., Janesville
*Lindsay, Ruth H., Nashotah
*Lobeck, Armin K., Science Hall, U. W., Madison
*Lounsbury, James A., Marquette University, Milwaukee
Lowe, John N., State Normal, Marquette, Mich.
^McAllister, Fred, University of Texas, Austin, Tex.
*McCoy, Elizabeth F., Agriculutral Hall, U. W., Madison
*McElvain, S. M., Chemistry Bldg., U. W., Madison
*MacLean, J. D., Forest Products Lab., Madison
McLeod, Andrew F., Crane Junior College, Chicago, Ill.
*McMillan, Mary B., 1341 Third St., South, Wisconsin Rapids
*McMinn, Amelia, State Normal School, Milwaukee
Main, Angie K., Fort Atkinson
Majerowski, Stephen J., Public Museum, Milwaukee
*Manresa, Miguel, Agricultural College, Los Banos, P. I.
*March, Herman W., North Hall, U. W., Madison
*Marlatt, Abby L., Home Ec. Bldg., U. W., Madison
*Marquette, William G., Pleasantville, N. Y.
"Marschall, A. J., 14 Proudfit St., Madison
Marshall, Ruth, Rockford College, Rockford, Ill.
^Marshall, William S., Biology Bldg., U. W., Madison
* Martin, Stephen J., Biology Bldg., U. W., Madison
*Mason, Carol Y., Milwaukee-Downer College, Milwaukee
*Mason, Max, Rockefeller Foundation, New York, N. Y.
*Mavor, James W., Union College, Schenectady, N. Y.
*Mathews, Joseph H., Chemistry Bldg., U. W., Madison
*Maurer, Edward R., Engr. Bldg., U. W., Madison
Meachem, John G., 745 College Ave., Racine
*Mead, Daniel W., Engr. Bldg., U. W., Madison
*Mead, Warren J., Science Hall, U. W., Madison
*Meloche, V. W., Chemistry Bldg., U. W., Madison
*Mendenhall, Charles E., Sterling Hall, U. W., Madison
*Metzner, Albertine E., Plymouth
Meyer, B. H., 3327 P St., Washington, D. C.
*Miller, Eric R., North Hall, U. W., Madison
*Miller, William Snow, Memorial Inst. Bldg., U. W., Madison
*Miloslavich, Edward L., 415 Eastwood PI., Milwaukee
*Moe, Carl, International Falls, Minn.
Moffitt, George W., Williams Bay
Monroe, Charles E., 1400 Wisconsin Ave., Milwaukee
*Morris, Harold H., 624 High St., Newark, N. J.
*Mossman, Harland W., Science Hall, U. W., Madison
*Muehlberger, Clarence W., Memorial Institute, U. W., Madison
*Mullenix, Rollin C., Lawrence College, Appleton
*Musbach, Fred L., 409 South Cherry St., Marshfield
Muttkowski, Richard A., University of Detroit, Detroit, Mich.
* Member of American Association for Advancement of Science.
List of Officers and Members.
617
*Nagler, Ellen T., 1913 Kendall Ave., Madison
*Nardin, F. Louise, Lathrop Hall, U. W., Madison
Naylor, Wilson S., Lawrence College, Appleton
Nettles, Curtis P., Bascom Hall, U. W., Madison
*Nevins, Beatrice I., Biology Bldg., U. W., Madison
*Noland, Lowell E., Biology Bldg., U. W., Madison
Oberholser, Harry C., National Museum, Washington, D. C.
* Ogden, Henry V., 436 Summit Ave., Milwaukee
*0’Shea, M. Vincent, Bascom Hall, U. W., Madison
Otto, Max C., Bascom Hall, U. W., Madison
^Overton, James B., Biology Bldg., U. W., Madison
Owen, E. T., 624 State St., Madison
Pammel, L. H., Iowa State College, Ames, la.
*Parsons, Helen T., Home Ec. Bldg., U. W., Madison
Paxson, Frederick L., Bascom Hall, U. W., Madison
Pearse, A. S., Duke University, Durham, N. C.
Peter, George, Public Museum, Milwaukee
*Peterson, William H., Agric. Chem. Bldg., U. W., Madison
*Petzke, Ernest A., Hixton
^Phillips, James D., Administration Bldg., U. W., Madison
Pitman, Annie M., Extension Bldg., U. W., Madison
*Poetker, A. H., Marquette University, Milwaukee
*Pollitz, Louise, 65% Otter St., Oshkosh
Pope, Thomas B., Public Museum, Milwaukee
Potter, George F., University of New Hampshire, Durham, N. H.
* Public Museum, Milwaukee
Rakette, Maximilian, Unterlindau 66, Frankfurt am Main, Germany
*Reed, George M., Botanic Garden, Brooklyn, N. Y.
*Reith, Allan F., A. O. Smith Corp., Milwaukee
^Richards, C. Audrey, Forest Products Lab., Madison
*Richtmann, W. D., Chemistry Bldg., U. W., Madison
*Rickett, Harold W., University of Missouri, Columbia, Mo.
*Riker, Regina S., Horticultural Bldg., U. W., Madison
Rimsky-Korsakoff, Vladimir N., Biology Bldg., U. W., Madison
Ripley, Caryl, Williams Bay
*Roark, Raymond J., Engr. Bldg., U. W., Madison
* Roebuck, John R., Sterling Hall, U. W., Madison
Roedder, E. C. L. C., Bascom Hall, U. W., Madison
* Rogers, Walter E., Lawrence College, Appleton
*Rohde, H. W., 1275 Stowell Ave., Shorewood, Milwaukee
Ross, Frank E., Yerkes Observatory, Williams Bay
*Russell, Fred G., 141 E. Wisconsin Ave., Milwaukee
*Ruth, Elbert B., Biology Bldg., U. W., Madison
*Sammis, J. L., Hiram Smith Hall, U. W., Madison
Schaefer, David A., Lake Geneva
*Schlundt, Herman, University of Missouri, Columbia, Mo.
*Schmidt, Albert H., Palatine, Cook Co., Ill.
Member of American Association for Advancement of Science.
618 Wisconsin Academy of Sciences , Arts, and Letters .
♦Schmidt, Erwin R., Wis. Gen. Hosp., U. W., Madison
Schorger, A. W., 2021 Kendall Ave., Madison
Schroeder, Laura J., Beloit College, Beloit
Schubring, E. J. B., 415 N. Park St., Madison
Schubring, Selma L., 415 N. Park St., Madison
♦Schuette, H. A., Chem. Bldg., U. W., Madison
♦Schultz, Alfred R., 800 Vine St., Hudson
*Schuster, B. L., Central State Hosp., Waupun
Seaman, Gilbert E., 312 Goldsmith Bldg., Milwaukee
Showerman, Grant, Bascom Hall, U. W., Madison
Sieker, William C., 1542 Prospect Ave., Milwaukee
*Simon, Arthur, 3216 Chestnut St., Milwaukee
♦Skinner, Charles H., Marquette University, Milwaukee
Smith, Elmer A., 735-9th St., Secaucus, N. J.
♦Smith, Gilbert B. L., 99 Livingston St., Brooklyn, N. Y.
♦Smith, Harvey M., Wis. Gen. Hosp., U. W., Madison
*Smith, H. H., Public Museum, Milwaukee
Smith, Walter McM., Library, U. W., Madison
♦Snell, Walter H., Brown University, Providence, R. I.
♦Stebbins, Joel, Washburn Observatory, U. W., Madison
♦Steenbock, Harry, Agr. Chem. Bldg., U. W., Madison
*Steil, William N., Marquette University, Milwaukee
* Steiner, Gotthold, Bureau of Plant Industry, Washington, D. C.
♦Stickney, Gardner P., 864 Summit Ave., Milwaukee
Stickney, M. E., Denison University, Granville, O.
* Stock, Kurt, Fish Creek
* Stout, Arlow B., Botanical Garden, Bronx Park, New York, N. Y.
♦Sullivan, Walter E., Science Hall, U. W., Madison
*Taylor, James H., North Hall, U. W., Madison
* Taylor, L. W., University of Kentucky, Lexington, Ky.
Taylor, Warner, Bascom Hall, U. W., Madison
*Thorkelson, H. J. B., Kohler
Thwaites, Frederick T., Science Hall, U. W., Madison
*Tisdale, William B., Agric. Exp. Sta., Gainesville, Fla.
Tomlinson, Charles W., 1610 Bixby Ave., Ardmore, Okla.
♦Toole, Eben II., Bureau of Plant Industry, Washington, D. C.
*Tottingham, W. E., Agric. Chem. Bldg., U. W., Madison
*Trautmann, William J., Beloit College, Beloit
♦Tressler, Willis L., Biology Bldg., U. W., Madison
Triebel, Robert, Beuthen, O. S., Germany
♦Truog, Emil, New Soils Bldg., U. W., Madison
♦Turneaure, F. E., Engr. Bldg., U. W., Madison
Tutunjian, K. H., Ripon College, Ripon
♦Twenhofel, William H., Science Hall, U. W., Madison
Tyrrell, Edward R., Public Museum, Milwaukee
♦Uehling, Edward A., 1468 Fourth St., Milwaukee
♦Ullrich, Fred T., State Teachers College, Platteville
* Member of American Association for Advancement of Science.
List of Officers and Members.
*Van Biesbroeck, George, Yerkes Observatory, Williams Bay
Van Ostrand, Mrs. E. H., 214 W. Gilman St, Madison
-Van Vleck, Edward B., North Hall, U. W., Madison
*Van Vleck, John H., Sterling Hall, U. W., Madison
Vaughan, R. E., Horticultural Bldg., U. W., Madison
Voss, E. K. J. H., Bascom Hall, U. W., Madison
Wadmond, Samuel C., 305 Washington St., Delavan
*Wagner, George, Biology Bldg., U. W., Madison
*Wakeman, Nellie A., Chemistry Bldg., U. W., Madison
* Walker, J. Charles, Horticultural Bldg., U. W., Madison
* Walker, Ruth L, Biology Bldg., U. W., Madison
*' Warfield, Louis M., 193 Prospect Ave., Milwaukee
* Waters, Ralph M„ Wis, Gen. Hosp., U. W., Madison
* Weber, George F., Agric. Exp. Sta., Gainesville, Fla.
* Wellman, F. L., United Fruit Co., Tela, Honduras
*Wenstrand, D. E. W., 210 E. Wisconsin Ave., Milwaukee
* Werner, Fred W., 99 1-1 6th St, Milwaukee
West, George A., 2828 Highland Blvd., Milwaukee
*Whealdon, Albert D., 817 N. 16th St., Superior
nVhitbeck, R. H., Science Hall, U. W., Madison
Wiebe, Abraham H., Biological Station, Fairport, la.
*Wilson, H. F., Econ, Ent. Bldg., U. W., Madison
Wimmer, E. J., Agricultural College, Manhattan, Kan.
*Winchell, A. N., Science Hall, U. W., Madison
*Windesheim, Gustave, 453 Durkee Ave., Kenosha
*Witzemann, Edgar J., Science Hall, U. W., Madison
Woodmansee, Wilson R., Ripon College, Ripon
Wright, Stillman, University of Michigan, Ann Arbor, Mich.
Young, Karl, Yale University, New Haven, Conn.
Zdanowicz, Casimir, Bascom Hall, U. W., Madison
Zimmerman, Oliver B., 324 S. Ashland Ave., La Grange, III.
*Zoch, Richmond T., U. S. Weather Bureau, Washington, D. C.
* Member of American Association for Advancement of Science.
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